Gradients in embolism resistance within stems driven by secondary growth in herbs.

The stems of some herbaceous species can undergo basal secondary growth, leading to a continuum in the degree of woodiness along the stem. Whether the formation of secondary growth in the stem base results in differences in embolism resistance between the base and the upper portions of stems is unknown. We assessed the embolism resistance of leaves and the basal and upper portions of stems simultaneously within the same individuals of two divergent herbaceous species that undergo secondary growth in the mature stem bases. The species were Solanum lycopersicum (tomato) and Senecio minimus (fireweed). Basal stem in mature plants of both species displayed advanced secondary growth and greater resistance to embolism than the upper stem. This also resulted in significant vulnerability segmentation between the basal stem and the leaves in both species. Greater embolism resistance in the woodier stem base was found alongside decreases in the pith-to-xylem ratio, increases in the proportion of secondary xylem, and increases in lignin content. We show that there can be considerable variation in embolism resistance across the stem in herbs and that this variation is linked to the degree of secondary growth present. A gradient in embolism resistance across the stem in herbaceous plants could be an adaptation to ensure reproduction or basal resprouting during episodes of drought late in the lifecycle.

An assessment of commonly employed satellite-based remote sensors for mapping mangrove species in Mexico using an NDVI-based classification scheme.

Optimizing the classification accuracy of a mangrove forest is of utmost importance for conservation practitioners. Mangrove forest mapping using satellite-based remote sensing techniques is by far the most common method of classification currently used given the logistical difficulties of field endeavors in these forested wetlands. However, there is now an abundance of options from which to choose in regards to satellite sensors, which has led to substantially different estimations of mangrove forest location and extent with particular concern for degraded systems. The objective of this study was to assess the accuracy of mangrove forest classification using different remotely sensed data sources (i.e., Landsat-8, SPOT-5, Sentinel-2, and WorldView-2) for a system located along the Pacific coast of Mexico. Specifically, we examined a stressed semiarid mangrove forest which offers a variety of conditions such as dead areas, degraded stands, healthy mangroves, and very dense mangrove island formations. The results indicated that Landsat-8 (30 m per pixel) had  the lowest overall accuracy at 64% and that WorldView-2 (1.6 m per pixel) had the highest at 93%. Moreover, the SPOT-5 and the Sentinel-2 classifications (10 m per pixel) were very similar having accuracies of 75 and 78%, respectively. In comparison to WorldView-2, the other sensors overestimated the extent of Laguncularia racemosa and underestimated the extent of Rhizophora mangle. When considering such type of sensors, the higher spatial resolution can be particularly important in mapping small mangrove islands that often occur in degraded mangrove systems.

Diagnosing the level of stress on a mangrove species (Laguncularia racemosa) contaminated with oil: A necessary step for monitoring mangrove ecosystems.

Monitoring the effects of pollution on mangrove vegetation is a challenge. A specific study using an oil spill simulation on mangrove species was conducted to address this challenge. We tested the effectiveness of the chlorophyll a fluorescence kinetics as a fast and robust method to diagnose the vitality of Laguncularia racemosa. We used L. racemosa plants contaminated with marine fuel oil in mangrove microcosm models. Several parameters of the JIP-test were capable of detecting the impairment of the photosynthetic function prior to the visual manifestation of symptoms in response to oil contamination. The results support the use of the chlorophyll fluorescence transient as a reliable, fast and easy to apply diagnostic method for evaluating oil-impacted mangroves. To the best of our knowledge, it is the first time that consistent data showing photosynthetic impairment in response to oil contamination is shown for a mangrove tree species.

Environmental tolerances of rare and common mangroves along light and salinity gradients.

Although mangroves possess a variety of morphological and physiological adaptations for life in a stressful habitat, interspecific differences in survival and growth under different environmental conditions can shape their local and geographic distributions. Soil salinity and light are known to affect mangrove performance, often in an interactive fashion. It has also been hypothesized that mangroves are intrinsically shade intolerant due to the high physiological cost of coping with saline flooded soils. To evaluate the relationship between stress tolerance and species distributions, we compared responses of seedlings of three widespread mangrove species and one narrow endemic mangrove species in a factorial array of light levels and soil salinities in an outdoor laboratory experiment. The more narrowly distributed species was expected to exhibit a lower tolerance of potentially stressful conditions. Two of the widespread species, Avicennia germinans and Lumnitzera racemosa, survived and grew well at low-medium salinity, regardless of light level, but performed poorly at high salinity, particularly under high light. The third widespread species, Rhizophora mangle, responded less to variation in light and salinity. However, at high salinity, its relative growth rate was low at every light level and none of these plants flushed leaves. As predicted, the rare species, Pelliciera rhizophorae, was the most sensitive to environmental stressors, suffering especially high mortality and reduced growth and quantum yield under the combined conditions of high light and medium-high salinity. That it only thrives under shaded conditions represents an important exception to the prevailing belief that halophytes are intrinsically constrained to be shade intolerant.

Is the WBE model appropriate for semi-arid shrubs subjected to clear cutting?

It is crucial to understand the adaptive mechanisms of woody plants facing periodic drought to assess their vulnerability to the increasing climate variability predicted in the Sahel. Guiera senegalensis J.F.Gmel is a semi-evergreen Combretaceae commonly found in Sahelian rangelands, fallows and crop fields because of its value as an agroforestry species. We compared canopy leafing, and allometric measurements of leaf area, stem area and stem length and their relationships with leaf water potential, stomatal conductance (gs) and soil-to-leaf hydraulic conductance (KS-L), in mature and current-year resprouts of G. senegalensis in Sahelian Niger. In mature shrubs, seasonal drought reduced the ratio of leaf area to cross-sectional stem area (AL : AS), mainly due to leaf shedding. The canopy of the current-year resprouts remained permanently leafed as the shrubs produced leaves and stems continuously, and their AL : AS ratio increased throughout the dry season. Their KS-L increased, whereas gs decreased. West, Brown and Enquist's (WBE) model can thus describe allometric trends in the seasonal life cycle of undisturbed mature shrubs, but not that of resprouts. Annual clear cutting drives allometric scaling relationships away from theoretical WBE predictions in the current-year resprouts, with scaling exponents 2.5 times greater than those of mature shrubs. High KS-L (twice that of mature shrubs) supports this intensive regeneration process. The adaptive strategy described here is probably common to many woody species that have to cope with both severe seasonal drought and regular disturbance over the long term.

Poleward expansion of mangroves is a threshold response to decreased frequency of extreme cold events.

Regional warming associated with climate change is linked with altered range and abundance of species and ecosystems worldwide. However, the ecological impacts of changes in the frequency of extreme events have not been as well documented, especially for coastal and marine environments. We used 28 y of satellite imagery to demonstrate that the area of mangrove forests has doubled at the northern end of their historic range on the east coast of Florida. This expansion is associated with a reduction in the frequency of "extreme" cold events (days colder than -4 °C), but uncorrelated with changes in mean annual temperature, mean annual precipitation, and land use. Our analyses provide evidence for a threshold response, with declining frequency of severe cold winter events allowing for poleward expansion of mangroves. Future warming may result in increases in mangrove cover beyond current latitudinal limits of mangrove forests, thereby altering the structure and function of these important coastal ecosystems.

Cyanobacterial diversity in the phyllosphere of a mangrove forest.

The cyanobacterial community colonizing phyllosphere in a well-preserved Brazilian mangrove ecosystem was assessed using cultivation-independent molecular approaches. Leaves of trees that occupy this environment (Rhizophora mangle,Avicennia schaueriana and Laguncularia racemosa) were collected along a transect beginning at the margin of the bay and extending upland. The results demonstrated that the phyllosphere of R. mangle and L. racemosa harbor similar assemblages of cyanobacteria at each point along the transect. A. schaueriana, found only in the coastal portions of the transect, was colonized by assemblages with lower richness than the other trees. However, the results indicated that spatial location was a stronger driver of cyanobacterial community composition than plant species. Distinct cyanobacterial communities were observed at each location along the coast-to-upland transect. Clone library analysis allowed identification of 19 genera of cyanobacteria and demonstrated the presence of several uncultivated taxa. A predominance of sequences affiliated with the orders Nostocales and Oscillatoriales was observed, with a remarkable number of sequences similar to genera Symphyonemopsis/Brasilonema (order Nostocales). The results demonstrated that phyllosphere cyanobacteria in this mangrove forest ecosystem are influenced by environmental conditions as the primary driver at the ecosystem scale, with tree species exerting some effect on community structure at the local scale.

Secondary metabolites and nutrients of woody plants in relation to browsing intensity in African savannas.

Carbon-based secondary metabolites (CBSMs) are assumed to function as defences that contribute to herbivore-avoidance strategies of woody plants. Severe browsing has been reported to reduce concentrations of CBSMs and increase N concentrations in individual plants, causing heavily browsed plants to be characterised by N-rich/C-poor tissues. We hypothesised that concentrations of condensed tannins (CT) and total polyphenols (TP) should decrease, or N increase, in relation to increasing intensity of browsing, rendering severely browsed plants potentially more palatable (increased N:CT) and less N-limited (increased N:P) than lightly browsed ones. We sampled naturally browsed trees (taller than 2 m) of four abundant species in southern Kruger National Park, South Africa. Species-specific relationships between N:CT, CT, TP and P concentrations and increasing browsing intensity were detected, but N and N:P were consistently invariable. We developed a conceptual post-hoc model to explain diverse species-specific CBSM responses on the basis of relative allocation of C to total C-based defence traits (e.g. spines/thorns, tough/evergreen leaves, phenolic compounds). The model suggests that species with low allocation of C to C-based defence traits become C-limited (potentially more palatable) at higher browsing intensity than species with high allocation of C to C-based defences. The model also suggests that when N availability is high, plants become C-limited at higher browsing intensity than when N availability is low.

The effect of disturbance on an ant-plant mutualism.

Protective ant-plant mutualisms-where plants provide food or shelter to ants and ants protect the plants from herbivores-are a common feature in many ecological communities, but few studies have examined the effect of disturbance on these interactions. Disturbance may affect the relationship between plants and their associated ant mutualists by increasing the plants' susceptibility to herbivores, changing the amount of reward provided for the ants, and altering the abundance of ants and other predators. Pruning was used to simulate the damage to buttonwood mangrove (Conocarpus erectus) caused by hurricanes. Pruned plants grew faster than unpruned plants, produced lower levels of physical anti-herbivore defenses (trichomes, toughness), and higher levels of chemical defenses (tannins) and extrafloral nectaries. Thus, simulated hurricane damage increased plant growth and the amount of reward provided to ant mutualists, but did not have consistent effects on other anti-herbivore defenses. Both herbivores and ants increased in abundance on pruned plants, indicating that the effects of simulated hurricane damage on plant traits were propagated to higher trophic levels. Ant-exclusion led to higher leaf damage on both pruned and upruned plants. The effect of ant-exclusion did not differ between pruned and unpruned plants, despite the fact that pruned plants had higher ant and herbivore densities, produced more extrafloral nectaries, and had fewer physical defenses. Another common predator, clubionid spiders, increased in abundance on pruned plants from which ants had been excluded. I suggest that compensatory predation by these spiders diminished the effect of ant-exclusion on pruned plants.

Relationship of water transport to anatomical features in the mangrove Laguncularia racemosa grown under contrasting salinities.

The purpose of this study was to investigate the xylem anatomy and hydraulic characteristics of the mangrove Laguncularia racemosa grown under contrasting salinities. The study addressed the hypothesis that, at high salinity, water transport capacity may decrease in association with higher water use efficiency. Plants were grown in media to which 0, 15 and 30 NaCl was added. Vessel density and diameter were determined in transverse sections of stem and midrib leaves in terminal shoots, and hydraulic parameters were measured. In stems, the vessel density increased with salinity, while the anatomical diameter (d(a)) and hydraulic diameter (d(h)) declined; in leaves, these parameters remained unchanged with salinity. Huber value and hydraulic and specific conductivities decreased with salinity. Leaf blade resistance increased with salinity and represented the largest fraction of twig resistance. Xylem anatomy and leaf tissue of L. racemosa appeared to be modulated by salinity, which led to a coordinated decline in hydraulic properties as salinity increased. Therefore, these structural changes would reflect functional water use characteristics of leaves under salinity.

Seed shadow, seedling recruitment, and spatial distribution of Buchenavia capitata (Combretaceae) in a fragment of the Brazilian Atlantic forest.

Here we describe the seed shadow, seedling recruitment, ontogenetic structure and spatial distribution of Buchenavia capitata (an emergent canopy tree) in a 380-ha fragment of the Atlantic forest in northeast Brazil. In particular, we examine seed distribution around 10 parental trees and both seedling recruitment and mortality, during an 18 month period beneath and around parental trees. Moreover, we describe: (1) B. capitata occurrence within treefall gaps; (2) population structure in terms of ontogenetic stages for the whole site; and (3) spatial distribution of adults within an area of 51 hectares. 99% of seeds were found beneath parent crowns (n = 4,236) and seed density reached 14.6 +/- 29.9 seeds/m2 (0-140 seeds/m2). 49% of all seeds germinated but seedling mortality reached 100% after an 18 month period. In addition, saplings of B. capitata were not found in forest understory and within 30 treefall gaps (94-2,350 m2). The adults showed an average DBH of 69.3 +/- 22.1 cm, were 19.2 +/- 2.9 m tall and presented a clumped spatial distribution. B. capitata matched some of the features presented by shade intolerant trees or large-gap specialists, and we hypothesize that low rates or even lack of long distance seed dispersal events may be reducing the probability of B. capitata seeds reaching suitable habitats for successful seedling recruitment and growth. Because of that (1) seedlings face high levels of early mortality; (2) there is no sapling recruitment at the study site; and (3) local population faces senility and it is threatened by local extinction.

Seed shadow, seedling recruitment, and spatial distribution of Buchenavia capitata (Combretaceae) in a fragment of the Brazilian Atlantic Forest / Sombra de sementes, recrutamento de plântulas e distribuição espacial de Buchenavia capitata em um fragmento da Floresta Atlântica Brasileira

Here we describe the seed shadow, seedling recruitment, ontogenetic structure and spatial distribution of Buchenavia capitata (an emergent canopy tree) in a 380-ha fragment of the Atlantic forest in northeast Brazil. In particular, we examine seed distribution around 10 parental trees and both seedling recruitment and mortality, during an 18 month period beneath and around parental trees. Moreover, we describe: (1) B. capitata occurrence within treefall gaps; (2) population structure in terms of ontogenetic stages for the whole site; and (3) spatial distribution of adults within an area of 51 hectares. 99 percent of seeds were found beneath parent crowns (n = 4,236) and seed density reached 14.6 ± 29.9 seeds/m² (0-140 seeds/m²). 49 percent of all seeds germinated but seedling mortality reached 100 percent after an 18 month period. In addition, saplings of B. capitata were not found in forest understory and within 30 treefall gaps (94-2,350 m²). The adults showed an average DBH of 69.3 ± 22.1 cm, were 19.2 ± 2.9 m tall and presented a clumped spatial distribution. B. capitata matched some of the features presented by shade intolerant trees or large-gap specialists, and we hypothesize that low rates or even lack of long distance seed dispersal events may be reducing the probability of B. capitata seeds reaching suitable habitats for successful seedling recruitment and growth. Because of that (1) seedlings face high levels of early mortality; (2) there is no sapling recruitment at the study site; and (3) local population faces senility and it is threatened by local extinction.

Neste estudo, são descritas a sombra de sementes, o recrutamento de plântulas, a estrutura ontogenética e a distribuição espacial de Buchenavia capitata (árvore emergente) em um fragmento de 380 ha da floresta Atlântica do Nordeste do Brasil. Particularmente, são examinadas a distribuição de sementes de 10 árvores e o recrutamento e a mortalidade de plântulas durante 18 meses, tanto embaixo quanto em volta dos adultos. Além disso, descrevem-se: (1) a ocorrência de B. capitata em 30 clareiras; (2) a estrutura populacional em termos de estágios ontogenéticos para toda a área; e (3) a distribuição espacial dos adultos dentro de uma área de 51 hectares. Noventa e nove por cento das sementes foram encontradas embaixo das copas das matrizes (n = 4236) e a densidade de sementes alcançou 14,6 ± 29,9 sementes/m² (0-140 sementes/m²). Quarenta e nove por cento de todas as sementes germinaram, mas a mortalidade de plântulas atingiu 100 por cento após 18 meses. Além disso, jovens de B. capitata não foram encontrados no sub-bosque e em 30 clareiras (94-2350 m²). Os adultos mostraram um DAP médio de 69,3 ± 22,1 cm, 19,2 ± 2,9 m de altura e apresentaram uma distribuição espacial agregada. B. capitata apresenta características comuns às árvores intolerantes à sombra ou especialistas de grandes clareiras. Baixas taxas, ou mesmo a ausência, de dispersão de sementes a longa distância podem estar reduzindo a probabilidade de sementes de B. capitata alcançarem hábitats adequados ao recrutamento e ao desenvolvimento das plântulas. Por causa disto (1) as plântulas enfrentam altos níveis de mortalidade; (2) não tem ocorrido recrutamento de jovens; e (3) a população é senil e está ameaçada de extinção local.

Effect of root feeding by Diaprepes abbreviatus (Coleoptera: Curculionidae) larvae on leaf gas exchange and growth of three ornamental tree species.

Diaprepes abbreviatus L. (Coleoptera: Curculionidae), feeds on a variety of ornamental plants grown in southern Florida. Studies were conducted to evaluate the effects of root feeding by D. abbreviatus larvae on leaf gas exchange and growth of three ornamental tree species commonly grown in southern Florida that are known hosts of this weevil: green buttonwood, Conocarpus erectus L.; live oak, Quercus virginiana Mill.; and pygmy date palm, Phoenix roebelenii O'Brien. These hosts were grown in containers and infested with weevil larvae. Net CO2 assimilation, transpiration, and stomatal conductance of CO, were measured monthly. Leaf, stem, and root fresh and dry weights of each species also were determined. In one of two tests, larval root feeding significantly reduced net CO2 assimilation, transpiration, and stomatal conductance of CO2 of infested green buttonwood trees. Leaf gas exchange of live oak was not affected by larval infestation. In addition to testing cumulative effects of multiple infestations of larvae, the effects of incremental infestations on leaf gas exchange and fresh and dry weights also were tested for each plant species. Net CO2 assimilation, transpiration, stomatal conductance of CO2, and dry weights of green buttonwood were reduced as a result of larval root feeding, whereas there was no effect of incremental larval infestations on leaf gas exchange of live oak or pygmy date palm within the experimental time frame. There was no effect of incremental larval infestations on dry weights of live oak, but leaf, stem, and dry root weight of pygmy date palm were lower for infested plants than for noninfested plants. Overall, green buttonwood was more susceptible to larval root feeding damage than either live oak or pygmy date palm.

Analysis of the plant architecture via tree-structured statistical models: the hidden Markov tree models.

Plant architecture is the result of repetitions that occur through growth and branching processes. During plant ontogeny, changes in the morphological characteristics of plant entities are interpreted as the indirect translation of different physiological states of the meristems. Thus connected entities can exhibit either similar or very contrasted characteristics. We propose a statistical model to reveal and characterize homogeneous zones and transitions between zones within tree-structured data: the hidden Markov tree (HMT) model. This model leads to a clustering of the entities into classes sharing the same 'hidden state'. The application of the HMT model to two plant sets (apple trees and bush willows), measured at annual shoot scale, highlights ordered states defined by different morphological characteristics. The model provides a synthetic overview of state locations, pointing out homogeneous zones or ruptures. It also illustrates where within branching structures, and when during plant ontogeny, morphological changes occur. However, the labelling exhibits some patterns that cannot be described by the model parameters. Some of these limitations are addressed by two alternative HMT families.

Propagule size and predispersal damage by insects affect establishment and early growth of mangrove seedlings.

Variation in rates of seedling recruitment, growth, and survival can strongly influence the rate and course of forest regeneration following disturbance. Using a combination of field sampling and shadehouse experiments, we investigated the influence of propagule size and predispersal insect damage on the establishment and early growth of the three common mangrove species on the Caribbean coast of Panama: Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle. In our field samples, all three species exhibited considerable intraspecific variation in mature propagule size, and suffered moderate to high levels of predispersal attack by larval insects. Rates of insect attack were largely independent of propagule size both within and among trees. Our experimental studies using undamaged mature propagules showed that, for all three species, seedlings established at high rates regardless of propagule size. However, propagule size did have a marked effect on early seedling growth: seedlings that developed from larger propagules grew more rapidly. Predispersal insect infestations that had destroyed or removed a substantial amount of tissue, particularly if that tissue was meristematic or conductive, reduced the establishment of propagules of all three species. The effect of sublethal tissue damage or loss on the subsequent growth of established seedlings varied among the three mangrove species. For Avicennia, the growth response was graded: for a propagule of a given size, the more tissue lost, the slower the growth of the seedling. For Laguncularia, the response to insect attack appeared to be all-or-none. If the boring insect penetrated the outer spongy seed coat and reached the developing embryo, it usually caused sufficient damage to prevent a seedling from developing. On the other hand, if the insect damaged but did not penetrate the seed coat, a completely healthy seedling developed and its growth rate was indistinguishable from a seedling developing from an undamaged propagule of the same size. Similar to Avicennia, if an infestation did not completely girdle a Rhizophora seedling, it survived, but grew at a reduced rate. In summary, our experiments demonstrated that natural levels of variation in propagule size and predispersal damage by insects translate into significant differences in seedling performance in terms of establishment and/or early growth. Such differences are sufficiently large that they could influence the intensity and outcome of competitive interactions during forest regeneration.