A taxonomic revision of Rhizophora L. (Rhizophoraceae) in Thailand.

A taxonomic revision of Rhizophora L. (Rhizophoraceae) in Thailand is presented. Two species, R. apiculata Blume and R. mucronata Poir., are enumerated with updated morphological descriptions, illustrations and a taxonomic identification key, together with notes on distributions, habitats and ecology, phenology, conservation assessments, etymology, vernacular names, uses, and specimens examined. Three names in Rhizophora, are lectotypified: R. apiculata and two associated synonyms of R. mucronata (i.e., R. latifolia Miq. and R. macrorrhiza Griff.). R. longissima Blanco, a synonym of R. mucronata, is neotypified. All two Rhizophora species have a conservation assessment of Least Concern (LC). Based on the morphological identification, these two species can be distinguished from one another by the shape and width of the leaf laminae and the length of a terminal stiff point of the leaf laminae; the type and position of the inflorescences and the number of flowers per inflorescence; the character and color of the bracteoles; the presence or absence of the flower pedicels; the shape of the mature flower buds; the shape, color, and texture of the sepals; the shape, character, and the presence or absence of hairs of the petals; the number of stamens per flower; the size of the fruits; the color and size of the hypocotyls; the color and diameter of the cotyledonous cylindrical tubes; and the color of the colleters and exudate. The thick cuticles, sunken stomata, large hypodermal cells, and cork warts are adaptive anatomical features of leaves in Rhizophora that live in the mangrove environments. The pollen grains of Thai Rhizophora species are tricolporate, prolate spheroidal or oblate spheroidal shapes, small-sized, and reticulate exine sculpturing.

Hydrodynamics of mangrove-type root models: the effect of porosity, spacing ratio and flexibility.

Mangrove trees play a prominent role in coastal tropic and subtropical regions, providing habitats for many organisms and protecting shorelines against high energy flows. In particular, the species Rhizophora mangle (red mangrove) exhibits complex cluster roots interacting with different hydrological flow conditions. To better understand the resilience of mangrove trees, we modeled the roots as a collection of cylinders with a circular pattern subject to unidirectional flow. We investigated the effect of porosity and spacing ratio between roots by varying both the diameter of the patch, D, and inset cylinders, d. In addition, we modeled hanging roots of red mangroves as cantilevered rigid cylinders on a hinge. Force and velocity measurements were performed in a water tunnel (Reynolds numbers from 2200 to 11 000). Concurrently, we performed 2D flow visualization using a flowing soap film. We found that the frequency of the vortex shedding increases as the diameter of the small cylinders decreases while the patch diameter is constant, therefore increasing the Strouhal number, [Formula: see text]. By comparing the change of Strouhal numbers with a single solid cylinder, we introduced a new length scale, the effective diameter. The effective diameter of the patch decreases as the porosity increases. In addition, we found that patch drag scales linearly with the patch diameter but decreases linearly as the spacing ratio increases. After a spacing ratio of ([Formula: see text]), the force scales linearly with the free stream velocity, and the mean velocity behind the patch is independent of the Reynolds number and the patch effect disappears. For flexible cylinders, we found that a decrease in stiffness increases both patch drag and the wake deficit behind the patch in a similar fashion as increasing the blockage of the patch. This information has the potential to help in the development of methods to design resilient bio-inspired coastline structures.

Pb uptake and tolerance in the two selected mangroves with different root lignification and suberization.

Metal pollution has been widely reported in mangrove wetlands; however, the mechanisms involved in metal detoxification by mangroves are still poorly understood. This study aimed to investigate the possible function of root lignification/suberization on Pb uptake and tolerance in mangroves. Two mangroves, Acanthus ilicifolius and Rhizophora stylosa with different root lignification/suberization were selected as plant materials; the former exhibits a thin exodermis and low lignification/suberization, while the latter possesses a thick exodermis and high lignification/suberization. A pot trial with addition of Pb was conducted to investigate the differences in Pb uptake and tolerance between the two mangroves. The experiment of rhizobox was designed to explore Pb dynamics and availabilities in the rhizosphere soils, besides, the ability of Pb uptake by the excised roots and X-ray analysis for Pb distribution within roots were also detected. The results revealed that R. stylosa exhibited relatively higher Pb tolerance together with less Pb accumulations when compared to A. ilicifolius. For both species, lower proportion of exchangeable and Carbonate Pb and higher higher Fe-Mn oxides Pb were observed in the rhizosphere zone when compared to the respective non-rhizosphere zone. The results from metal uptake by the excised roots and X-ray analysis clearly showed that the thick lignified/suberized exodermis of R. stylosa could more efficiently delay Pb entering into the roots, leading to less Pb accumulation. In summary, the present study proposes a barrier property of the lignified/suberized exodermis in dealing with the stresses of Pb.

Root biomechanics in Rhizophora mangle: anatomy, morphology and ecology of mangrove's flying buttresses.

BACKGROUND AND AIMS: Rhizophora species of mangroves have a conspicuous system of stilt-like roots (rhizophores) that grow from the main stem and resemble flying buttresses. As such, the development of rhizophores can be predicted to be important for the effective transmission of dynamic loads from the top of the tree to the ground, especially where the substrate is unstable, as is often the case in the habitats where Rhizophora species typically grow. This study tests the hypothesis that rhizophore architecture in R. mangle co-varies with their proximity to the main stem, and with stem size and crown position. METHODS: The allometry and wood mechanical properties of R. mangle (red mangrove) trees growing in a mangrove basin forest within a coastal lagoon in Mexico were compared with those of coexisting, non-buttressed mangrove trees of Avicennia germinans. The anatomy of rhizophores was related to mechanical stress due to crown orientation (static load) and to prevailing winds (dynamic load) at the study site. KEY RESULTS: Rhizophores buttressed between 10 and 33 % of tree height. There were significant and direct scaling relationships between the number, height and length of rhizophores vs. basal area, tree height and crown area. Wood mechanical resistance was significantly higher in the buttressed R. mangle (modulus of elasticity, MOE = 18·1 ± 2 GPa) than in A. germinans (MOE = 12·1 ± 0·5 GPa). Slenderness ratios (total height/stem diameter) were higher in R. mangle, but there were no interspecies differences in critical buckling height. When in proximity to the main stem, rhizophores had a lower length/height ratio, higher eccentricity and higher xylem/bark and pith proportions. However, there were no directional trends with regard to prevailing winds or tree leaning. CONCLUSIONS: In comparison with A. germinans, a tree species with wide girth and flare at the base, R. mangle supports a thinner stem of higher mechanical resistance that is stabilized by rhizophores resembling flying buttresses. This provides a unique strategy to increase tree slenderness and height in the typically unstable substrate on which the trees grow, at a site that is subject to frequent storms.

Metal (Pb, Zn and Cu) uptake and tolerance by mangroves in relation to root anatomy and lignification/suberization.

Metal pollution has been widely reported in mangrove wetlands; however, the mechanisms involved in metal detoxification by mangroves are still poorly understood. This study aimed to investigate the possible function of root anatomy and lignification/suberization on metal uptake and tolerance in seedlings of six species of mangroves. The results revealed that the three rhizophoraceous species (Bruguiera gymnorrhiza (L.) Poir, Kandelia obovata Sheue, Liu & Yong and Rhizophora stylosa Griff) consistently exhibited higher metal tolerances than the three pioneer species (Aegiceras corniculatum (Linn.) Blanco, Acanthus ilicifolius L. and Avicennia marina (Forsk.) Viern.). Moreover, metal-tolerant species often exhibited a thick exodermis with high lignification and suberization. The tolerance indices of the mangroves were found to be positively correlated with the amounts of lignin and suberin deposition within the exodermal cell walls. The observed metal uptake by the excised roots further illustrated that a lignified/suberized exodermis directly delayed the entry of metals into the roots, and thereby contributed to a higher tolerance to heavy metals. In summary, the present study proposes a barrier property of the lignified/suberized exodermis in dealing with the stresses of heavy metals, such that the mangroves which possessed more extensive lignification/suberization within the exodermis appeared to exhibit higher metal tolerance.

Modelling reveals endogenous osmotic adaptation of storage tissue water potential as an important driver determining different stem diameter variation patterns in the mangrove species Avicennia marina and Rhizophora stylosa.

BACKGROUND: Stem diameter variations are mainly determined by the radial water transport between xylem and storage tissues. This radial transport results from the water potential difference between these tissues, which is influenced by both hydraulic and carbon related processes. Measurements have shown that when subjected to the same environmental conditions, the co-occurring mangrove species Avicennia marina and Rhizophora stylosa unexpectedly show a totally different pattern in daily stem diameter variation. METHODS: Using in situ measurements of stem diameter variation, stem water potential and sap flow, a mechanistic flow and storage model based on the cohesion-tension theory was applied to assess the differences in osmotic storage water potential between Avicennia marina and Rhizophora stylosa. KEY RESULTS: Both species, subjected to the same environmental conditions, showed a resembling daily pattern in simulated osmotic storage water potential. However, the osmotic storage water potential of R. stylosa started to decrease slightly after that of A. marina in the morning and increased again slightly later in the evening. This small shift in osmotic storage water potential likely underlaid the marked differences in daily stem diameter variation pattern between the two species. CONCLUSIONS: The results show that in addition to environmental dynamics, endogenous changes in the osmotic storage water potential must be taken into account in order to accurately predict stem diameter variations, and hence growth.

Influence of N deficiency and salinity on metal (Pb, Zn and Cu) accumulation and tolerance by Rhizophora stylosa in relation to root anatomy and permeability.

Effects of N deficiency and salinity on root anatomy, permeability and metal (Pb, Zn and Cu) translocation and tolerance were investigated using mangrove seedlings of Rhizophora stylosa. The results showed that salt could directly reduce radial oxygen loss (ROL) by stimulation of lignification within exodermis. N deficiency, oppositely, would reduce lignification. Such an alteration in root permeability may also influence metal tolerance by plants. The data indicated that a moderate salinity could stimulate a lignified exodermis that delayed the entry of metals into the roots and thereby contributed to a higher metal tolerance, while N deficiency would aggravate metal toxicity. The results from sand pot trail further confirmed this issue. This study provides a barrier property of the exodermis in dealing with environments. The plasticity of root anatomy is likely an adaptive strategy to regulate the fluxes of gases, nutrients and toxins at root-soil interface.

Interactions among Fe2+, S2-, and Zn2+ tolerance, root anatomy, and radial oxygen loss in mangrove plants.

Root anatomy, radial oxygen loss (ROL), and tolerances to ferrous (Fe(2+)), sulphide (S(2-)), and zinc (Zn(2+)) ions were investigated in seedlings of eight species of mangrove, including three pioneer species, three rhizophoraceous and two landward semi-mangrove species. The results showed an interesting co-tolerance to Fe(2+), S(2-), and Zn(2+). The three rhizophoraceous species (Bruguiera gymnorrhiza, Kandelia obovata and Rhizophora stylosa), which possessed the thickest lignified exodermis and the 'tightest barrier' in ROL spatial pattern, consistently exhibited the highest tolerance to Fe(2+), S(2-), and Zn(2+). B. gymnorrhiza could directly reduce ROL by increasing lignification within the exodermis. Such an induced barrier to ROL is a probable defence response to prevent further invasion and spread of toxins within plants. The data also indicated that, in B. gymnorrhiza, Fe(2+) or S(2-), or both, induced a lignified exodermis that delayed the entry of Zn(2+) into the roots and thereby contributed to a higher tolerance to Zn(2+). This study provides new evidence of exclusive strategies of mangrove seedling roots in dealing with contaminations. The information is also important in the selection and cultivation of tolerant species for the bioremediation of contaminated waters or soils.

Effect of wastewater discharge on root anatomy and radial oxygen loss (ROL) patterns of three mangrove species in southern China.

The effects of wastewater discharge on radial oxygen loss (ROL) and root anatomy varied among mangrove species. ROL of Bruguiera gymnorrhiza (L) increased from 22.44 ng cm(-2) min(-1) in the control (just fresh water) to 31.09 ng cm(-2) min(-1) when received normal wastewater (NW) and to 44.22 ng cm(-2) min(-1) when treated with strong wastewater (10NW). However, discharge of both NW and 10NW caused 28% decreases of ROL in the root tip of Excoecaria agallocha L., and the decreases in Acanthus ilicifolius L were even more significant, with 45% when treated by 10NW The changes of ROL were related to the root anatomy. Among three species, A. ilicifolius had the highest proportional cross-sectional area of aerenchyma air spaces, suggesting that the internal oxygen transfer to root tip was the fastest. However, the area of aerenchyma air spaces in the root tip of 10NW treated A. ilicifolius was significantly reduced while area of epidermis and hypodermis (E + H) increased leading to less oxygen supply to root tip. Compared to B. gymnorrhiza and E. agallocha, the (E + H) layer of A. ilicifolius was the thinnest, and the cells without suberized walls were loosely packed in all three treatments. These results suggested that the root anatomy and ROL of B. gymnorrhiza was least affected by wastewater discharge, followed by E. agallocha, and A. ilicifolius was the most susceptible species thus was not suitable for treating strong wastewater.

The role of radial oxygen loss and root anatomy on zinc uptake and tolerance in mangrove seedlings.

Root anatomy, radial oxygen loss (ROL) and zinc (Zn) uptake and tolerance in mangrove plants were investigated using seedlings of Aegiceras corniculatum, Bruguiera gymnorrhiza and Rhizophora stylosa. The results revealed that B. gymnorrhiza, which possessed the 'tightest barrier' in ROL spatial patterns among the three species studied, took up the least Zn and showed the highest Zn tolerance. Furthermore, zinc significantly decreased the ROL of all three plants by inhibition of root permeability, which included an obvious thickening of outer cortex and significant increases of lignification in cell walls. The results of SEM X-ray microanalysis further confirmed that such an inducible, low permeability of roots was likely an adaptive strategy to metal stress by direct prevention of excessive Zn entering into the root. The present study proposes new evidence of structural adaptive strategy on metal tolerance by mangrove seedlings.

[Costa Rica mangroves: the north Pacific]. / Los manglares de Costa Rica: el Pacífico norte.

Costa Rica has mangrove forests on both the Caribbean and Pacific coasts. The Pacific side has 99% of the mangrove area of the country. In this review we compile available information on the mangroves of the north Pacific coast of Costa Rica, from Bahía Salinas, on the border with Nicaragua, to the tip of the Peninsula de Nicoya at Cabo Blanco. We provide information on the location of the mangroves and all available information for each mangrove forest. These mangrove communities are smaller in extension and tree sizes, and have lower diversity compared to the mangroves on the southern section of the Pacific coast of Costa Rica. The dominant species are Rhizophora mangle and Rhizophora racemosa along the canal edges, backed by Avicennia germinans, and farther inland Avicennia bicolor, Laguncularia racemosa and Conocarpus erectus. At Potrero Grande a healthy population of Pelliciera rhizophorae, a rare species, has been reported. We recognized 38 mangrove communities in this part of the country, based on the National Wetland Inventory, published papers, field observations, theses, technical reports, and the national topographic maps (1:50,000, Instituto Geográfico Nacional). Relatively detailed information could be found for only five mangrove forests, for 14 more only prelimary and incomplete lists of plants and in some cases of animal species are available, for nine there is even less information, and for nine more only their location is known, which in some cases was not correct. Detail mapping, characterization of the vegetation and fauna, physiological studies, analyses of biogeochemical and physical processes, economic valuations, and determination of the health status of the mangrove of the northern Pacific coast, as well as for the rest of Costa Rica, are neccesary and urgent.

Los manglares de Costa Rica: el Pacífico norte

Abstract: Costa Rica mangroves: the north Pacific. Costa Rica has mangrove forests on both the Caribbean and Pacific coasts. The Pacific side has 99% of the mangrove area of the country. In this review we compile available information on the mangroves of the north Pacific coast of Costa Rica, from Bahía Salinas, on the border with Nicaragua, to the tip of the Peninsula de Nicoya at Cabo Blanco. We provide information on the location of the mangroves and all available information for each mangrove forest. These mangrove communities are smaller in extension and tree sizes, and have lower diversity compared to the mangroves on the southern section of the Pacific coast of Costa Rica. The dominant species are Rhizophora mangle and Rhizophora racemosa along the canal edges, backed by Avicennia germinans, and farther inland Avicennia bicolor, Laguncularia racemosa and Conocarpus erectus. At Potrero Grande a healthy population of Pelliciera rhizophorae, a rare species, has been reported. We recognized 38 mangrove communities in this part of the country, based on the National Wetland Inventory, published papers, field observations, theses, technical reports, and the national topographic maps (1:50 000, Instituto Geográfico Nacional). Relatively detailed information could be found for only five mangrove forests, for 14 more only prelimary and incomplete lists of plants and in some cases of animal species are available, for nine there is even less information, and for nine more only their location is known, which in some cases was not correct. Detail mapping, characterization of the vegetation and fauna, physiological studies, analyses of biogeochemical and physical processes, economic valuations, and determination of the health status of the mangrove of the northern Pacific coast, as well as for the rest of Costa Rica, are neccesary and urgent. Rev. Biol. Trop. 57 (3): 473-488. Epub 2009 September 30.

Costa Rica tiene bosques de manglar en las costas Caribe y Pacífica. El 99% de los manglares del país se encuentran en la costa Pacífica. En este trabajo recopilamos la información disponible sobre los manglares del Pacífico norte de Costa Rica, desde la frontera con Nicaragua hasta la Reserva Absoluta de Cabo Blanco, en la punta sur de la Península de Nicoya. La ubicación de los manglares y cualquier información disponible para cada manglar es resumida. El clima de esta región es seco con estaciones de lluvia y sequía muy bien definidas. Los manglares del norte son relativamente pequeños en extensión y tamaño de árboles, y de baja diversidad comparados con los del Pacífico sur de Costa Rica. Las principales especies de mangle son: Rhizophora mangle, Rhizophora racemosa a lo largo de los bordes de los canales, seguido por Avicennia germinans, y más tierra adentro Avicennia bicolor, Laguncularia racemosa y Conocarpus erectus. En el manglar de Potrero Grande se informa de una población saludable de la especie de Pelliciera rhizophorae, especies muy poco abundante. Reconocemos 38 comunidades de manglares en el Pacífico norte de Costa Rica basado en el Inventario Nacional de Humedales, artículos publicados, observaciones de campo, tesis, informes técnicos y los mapas 1: 50 000 del Instituto Geográfico Nacional. Se pudo encontrar información relativamente detallada pero fragmentada de únicamente cinco manglares (los descritos en esta publicación), de 14 solamente hay listas preliminares e incompletas de plantas y en algunos casos de animales, de nueve manglares más hay información todavía más limitada, y de otros nueve se conoce solamente su ubicación, que en algunos casos se comprobó que era incorrecta. Trabajos de mapeo detallado, caracterización de la vegetación y la fauna, estudios fisiológicos, análisis de procesos biogeoquímicos, evaluaciones económicas, y determinación del estado de salud de los manglares del Pacífico norte de Costa Rica, y del resto del país son necesarios y urgentes.

Hydraulic redistribution in dwarf Rhizophora mangle trees driven by interstitial soil water salinity gradients: impacts on hydraulic architecture and gas exchange.

Rhizophora mangle L. trees of Biscayne National Park (Florida, USA) have two distinct growth forms: tall trees (5-10 m) growing along the coast and dwarf trees (1 m or less) growing in the adjacent inland zone. Sharp decreases in salinity and thus increases in soil water potential from surface soil to about a depth of 1 m were found at the dwarf mangrove site but not at the tall mangrove site. Consistent with our prediction, hydraulic redistribution detected by reverse sap flow in shallow prop roots was observed during nighttime, early morning and late afternoon in dwarf trees, but not in tall trees. In addition, hydraulic redistribution was observed throughout the 24-h period during a low temperature spell. Dwarf trees had significantly lower sapwood-specific hydraulic conductivity, smaller stem vessel diameter, lower leaf area to sapwood area ratio (LA/SA), smaller leaf size and higher leaf mass per area. Leaves of dwarf trees had lower CO(2) assimilation rate and lower stomatal conductance compared to tall trees. Leaf water potentials at midday were more negative in tall trees that are consistent with their substantially higher stomatal conductance and LA/SA. The substantially lower water transport efficiency and the more conservative water use of dwarf trees may be due to a combination of factors such as high salinity in the surface soil, particularly during dry periods, and substantial reverse sap flow in shallow roots that make upper soil layers with high salinity a competing sink of water to the transpiring leaves. There may also be a benefit for the dwarf trees in having hydraulic redistribution because the reverse flow and the release of water to upper soil layers should lead to dilution of the high salinity in the rhizosphere and thus relieve its potential harm to dwarf R. mangle trees.

Transcriptome profiling of the mangrove plant Bruguiera gymnorhiza and identification of salt tolerance genes by Agrobacterium functional screening.

To identify key genes in the regulation of salt tolerance in the mangrove plant Bruguiera gymnorhiza, transcriptome profiling in the lateral and main roots under conditions of salt stress was performed. Statistical analysis revealed that 175 and 403 of 11,997 genes shoewd significantly increased high expression in the lateral and main roots respectively. One hundred and sixty genes were up-regulated in both types of roots in the early time period, 1 to 12 h after salt treatment. Expression vectors for 28 selected salt responsive genes were constructed and transformed in Agrobacterium tumefaciens, and then screened for salt tolerance. A. tumefaciens transformed with genes for lipid transfer, zinc finger, and ankyrin repeat proteins showed enhanced salt tolerance. Transgenic Arabidopsis plants expressing these three genes also exhibited increased tolerance to NaCl. These results indicate that Agrobacterium functional screening is an effective supplemental method of pre-screening genes involved in abiotic stress tolerance.

The role of freezing in setting the latitudinal limits of mangrove forests.

Mangrove trees dominate coastal vegetation in tropical regions, but are completely replaced by herbaceous salt marshes at latitudes above 32 degrees N and 40 degrees S. Because water deficit can increase damage caused by freezing, we hypothesized that mangroves, which experience large deficits as a result of saline substrates, would suffer freeze-induced xylem failure. Vulnerability to freeze-induced xylem embolism was examined in the most poleward mangrove species in North America, in an area where freezing is rare but severe, and in Australia, in an area where freezing is frequent but mild. Percentage loss in hydraulic conductivity was measured following manipulations of xylem tension; xylem sap ion concentration was determined using X-ray microanalysis. Species with wider vessels suffered 60-100% loss of hydraulic conductivity after freezing and thawing under tension, while species with narrower vessels lost as little as 13-40% of conductivity. These results indicate that freeze-induced embolism may play a role in setting the latitudinal limits of distribution in mangroves, either through massive embolism following freezing, or through constraints on water transport as a result of vessel size.

Influence of a salinity gradient on the vessel characters of the mangrove species Rhizophora mucronata.

BACKGROUND AND AIMS: Although mangroves have been extensively studied, little is known about their ecological wood anatomy. This investigation examined the potential use of vessel density as a proxy for soil water salinity in the mangrove species Rhizophora mucronata (Rhizophoraceae) from Kenya. METHODS: In a time-standardized approach, 50 wood discs from trees growing in six salinity categories were investigated. Vessel densities, and tangential and radial diameters of rainy and dry season wood of one distinct year, at three positions on the stem discs, were measured. A repeated-measures ANOVA with the prevailing salinity was performed. KEY RESULTS: Vessel density showed a significant increase with salinity, supporting its use as a prospective measure of salinity. Interestingly, the negative salinity response of the radial diameter of vessels was less striking, and tangential diameter was constant under the varying environmental conditions. An effect of age or growth rate or the presence of vessel dimorphism could be excluded as the cause of the absence of any ecological trend. CONCLUSIONS: The clear trend in vessel density with salinity, together with the absence of a growth rate and age effect, validates the potential of vessel density as an environmental proxy. However, it can only be used as a relative measure of salinity given that other environmental variables such as inundation frequency have an additional influence on vessel density. With view to a reliable, absolute proxy, future research should focus on finding wood anatomical features correlated exclusively with soil water salinity or inundation frequency. The plasticity in vessel density with differing salinity suggests a role in the establishment of a safe water transport system. To confirm this hypothesis, the role of inter-vessel pits, their relationship to the rather constant vessel diameter and the underlying physiology and cell biology needs to be examined.

Rhizophores in Rhizophora mangle L: an alternative interpretation of so-called ''aerial roots''

Rhizophora mangle L., uma das mais comuns espécies do mangue, tem um sistema de estruturas aéreas que lhe fornecem estabilidade em solo permanentemente alagado. De fato, essas estruturas, conhecidas por ''raízes aéreas'' ou ''raízes suportes'' demonstraram tratar-se de ramos especiais com geotropismo positivo, que formam grande número de raízes quando em contato com o solo. Esses órgãos apresentam um sistema de ramificação simpodial, medula ampla, córtex pouco espesso, feixes vasculares colaterais, estelo poliarco e protoxilema endarco, como no caule, e uma periderme produzida por um felogênio no ápice, semelhante a uma coifa. Esses ramos apresentam, também, o mesmo tipo de tricoesclereídes que ocorrem no caule com geotropismo negativo, diferente das verdadeiras raízes de Rhizophora, que não formam tricoesclereídes. Por outro lado, esses ramos não formam folhas e nesse aspecto são semelhantes às raízes. Esses ramos especiais são rizóforos, isto é, ramos portadores de raízes, com geotropismo negativo e análogos àqueles encontrados em Lepidodendrales e outras pteridófitas arbóreas do Carbonífero que, usualmente, cresciam em solos alagados.

Rhizophores in Rhizophora mangle L: an alternative interpretation of so-called ''aerial roots''.

Rhizophora mangle L., one of the most common mangrove species, has an aerial structure system that gives it stability in permanently swampy soils. In fact, these structures, known as "aerial roots" or "stilt roots", have proven to be peculiar branches with positive geotropism, which form a large number of roots when in contact with swampy soils. These organs have a sympodial branching system, wide pith, slightly thickened cortex, collateral vascular bundles, polyarch stele and endarch protoxylem, as in the stem, and a periderm produced by a phellogen at the apex similar to a root cap. They also have the same type of trichosclereid that occurs in the stem, with negative geotropism, unlike true Rhizophora roots, which do not form trichosclereids at all. On the other hand, these branches do not form leaves and in this respect they are similar to roots. These peculiar branches are rhizophores or special root-bearing branches, analogous to those found in Lepidodendrales and other Carboniferous tree ferns that grew in swampy soils.

Leaf gas exchange characteristics of three neotropical mangrove species in response to varying hydroperiod.

We determined how different hydroperiods affected leaf gas exchange characteristics of greenhouse-grown seedlings (2002) and saplings (2003) of the mangrove species Avicennia germinans (L.) Stearn., Laguncularia racemosa (L.) Gaertn. f., and Rhizophora mangle L. Hydroperiod treatments included no flooding (unflooded), intermittent flooding (intermittent), and permanent flooding (flooded). Plants in the intermittent treatment were measured under both flooded and drained states and compared separately. In the greenhouse study, plants of all species maintained different leaf areas in the contrasting hydroperiods during both years. Assimilation-light response curves indicated that the different hydroperiods had little effect on leaf gas exchange characteristics in either seedlings or saplings. However, short-term intermittent flooding for between 6 and 22 days caused a 20% reduction in maximum leaf-level carbon assimilation rate, a 51% lower light requirement to attain 50% of maximum assimilation, and a 38% higher demand from dark respiration. Although interspecific differences were evident for nearly all measured parameters in both years, there was little consistency in ranking of the interspecific responses. Species by hydroperiod interactions were significant only for sapling leaf area. In a field study, R. mangle saplings along the Shark River in the Everglades National Park either demonstrated no significant effect or slight enhancement of carbon assimilation and water-use efficiency while flooded. We obtained little evidence that contrasting hydroperiods affect leaf gas exchange characteristics of mangrove seedlings or saplings over long time intervals; however, intermittent flooding may cause short-term depressions in leaf gas exchange. The resilience of mangrove systems to flooding, as demonstrated in the permanently flooded treatments, will likely promote photosynthetic and morphological adjustment to slight hydroperiod shifts in many settings.

High-resolution time series of vessel density in Kenyan mangrove trees reveal a link with climate.

Tropical trees are often excluded from dendrochronological investigations because of a lack of distinct growth ring boundaries, causing a gap in paleoclimate reconstructions from tropical regions. The potential use of time series of vessel features (density, diameter, surface area and hydraulic conductivity) combined with spectral analysis as a proxy for environmental conditions in the mangrove Rhizophora mucronata was investigated. Intra-annual differences in the vessel features revealed a trade-off between hydraulic efficiency (large vessels) during the rainy season and hydraulic safety (small, more numerous vessels) during the dry season. In addition to the earlywood-latewood variations, a semiannual signal was discovered in the vessel density and diameters after Fourier transformation. The similarity in the Fourier spectra of the vessel features and the climate data, in particular mean relative humidity and precipitation, provides strong evidence for a climatic driving force for the intra-annual variability of the vessel features. The high-resolution approach used in this study, in combination with spectral analysis, may have great potential for the study of climate variability in tropical regions.