Relation of water transport to leaf gas exchange properties in three mangrove species

Mangrove species more tolerant to salinity may function with less efficient water transport, which may be related to more conservative water use. To test the hypothesis, we investigate the gas exchange and hydraulic properties of three mangrove species: Rhizophora mangle L., Laguncularia racemosa Gaert and Avicennia germinans (L.)L. Experiments were performed with adult plants growing naturally in the field under a salinity of 35‰. Gas exchange parameters showed that A. germinans had significantly higher photosynthetic rates, and lower stomatal conductance and transpiration rates, compared to the other two mangroves. In concert with this, instantaneous water use efficiency was significantly high in A. germinans, intermediate in L. racemosa and lowest in R. mangle. The hydraulic parameters of the three mangrove species were in the lowest end of the range reported for tropical trees. However, the three mangrove species exhibited measurable differences in hydraulic parameters related to the control of water requirements for maintenance of carbon gain. L. racemosa and A. germinans showed less efficient water transport at shoot level but were the more efficient species in water use at the leaf level in comparison to R. mangle. Received: 7 April 1999 / Accepted: 25 July 1999     

Increasing fluctuations of soil salinity affect seedling growth performances and physiology in three Neotropical mangrove species

Background

Micro-tidal wetlands are subject to strong seasonal variations of soil salinity that are likely to increase in amplitude according to climate model predictions for the Caribbean. Whereas the effects of constant salinity levels on the physiology of mangrove species have been widely tested, little is known about acclimation to fluctuations in salinity.

Aims and methods

The aim of this experiment was to characterize the consequences of the rate of increase in salinity (slow versus fast) and salinity fluctuations over time versus constant salt level. Seedling mortality, growth, and leaf gas exchange of three mangrove species, Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle were investigated in semicontrolled conditions at different salt levels (0, 685, 1025, and 1370 mM NaCl).

Results

Slow salinity increase up to 685 mM induced acclimation, improving the salt tolerance of A. germinans and L. racemosa, but had no effect on R. mangle. During fluctuations between 0 and 685 mM, A. germinans and R. mangle were not affected by a salinity drop to zero, whereas L. racemosa took advantage of the brief freshwater episode as shown by the durable improvement of photosynthesis and biomass production.

Conclusions

This study provides new insights into physiological resistance and acclimation to salt stress. We show that seasonal variations of salinity may affect mangrove seedlings’ morphology and physiology as much as annual mean salinity. Moreover, more severe dry seasons due to climate change may impact tree stature and species composition in mangroves through higher mortality rates and physiological disturbance at the seedling stage.     

Distribution of mangrove vegetation along inundation, phosphorus, and salinity gradients on the Bragança Peninsula in Northern Brazil

Background and aims

The Bragança Peninsula, in northern Brazil is characterized by macrotides (4 m) and specific edaphic conditions, which determine the local mangrove forest’s development. This study, conducted during the dry season evaluated the spatial patterns of Rhizophora mangle and Avicennia germinans species across an inundation gradient.

Methods

Along a transect of 700 m, measurements of structure forest, soil moisture, porewater salinity, extractable phosphorus (extr.-P) in sediments, and phosphorus in the leaves (leaf-P) were conducted.

Result

The A. germinans (100 %) occurred in high intertidal (HI) zone. A. germinans (59 %) and R. mangle (41 %) co-occurred in mid intertidal (MI) zone, while R. mangle (58 %) predominated in low intertidal (LI) zone, followed by A. germinans (37 %) and Laguncularia racemosa (5 %). Covariance analysis (ANCOVA) indicated that salinity and soil moisture means are significantly different between the mangrove forests, but do not correlate with inundation frequency (IF). The means of extr.-P were significantly different in mangrove forests and correlated with IF and leaf-P.

Conclusion

The inundation frequency, the availability of P in the sediments, phosphorus in the leaves and interstitial salinity are all important factors contributing to the distribution of the mangrove tree species A. germinans and R. mangle on the Bragança Peninsula.     

Reproductive phenology and physiological traits in the red mangrove hybrid complex (Rhizophora mangle and R. racemosa) across a natural gradient of nutrients and salinity

Factors modulating introgressive hybridization between the red mangrove species Rhizophora mangle and R. racemosa in spatially defined sites are poorly understood. To investigate this, we evaluated the reproductive phenology and the nutrient and physiological traits in those two species and their F₁ hybrids genotyped with microsatellite data across a natural hybrid zone from the Pacific coast of Panama. We found no evidence that reproductive phenology represents a barrier to gene flow, because R. mangle and the F₁ hybrids produced flowers and propagules throughout the annual cycle, while R. racemosa flowered only in the dry season. Soil nutrient concentrations decreased landward, while soil salinity varied only slightly. Foliar nutrients and δ¹⁵N signatures varied according to the soil nutrient gradient, but only foliar phosphorus and carbon varied among species. In contrast, two structural variables (height and trunk diameter) and leaf variables related to salinity tolerance (Na, Cl:Na, K:Na, cation:anion) and water-use efficiency (i.e., δ¹³C) differed among species, suggesting higher salinity tolerance for R. mangle and F₁ hybrids compared with R. racemosa. We conclude that parental species and F₁ hybrids differ in salinity tolerance and water-use efficiency, which could be associated with adaptive evolution of the red mangrove hybrid complex.     

Herbivore feeding preferences as measured by leaf damage and stomatal ingestion: a mangrove crab example

The diet of the mangrove crab, Aratus pisonii, was assessed by determining the percent of damaged leaves at selected mangrove communities and by examining herbivore gut contents. This study compared the utility of both methods and tested if comparable levels of damage and dietary preference occurred using the methods. Percent of damaged leaves was determined for the three species of mangroves within Tampa Bay, FL, USA, including: the red, black, and white mangroves (Rhizophora mangle, Avicennia germinans, and Laguncularia racemosa, respectively) in four 5×10-m quadrats during summer 2001. For each species, in each of the quadrats, 200 leaves per tree were assessed for the presence or absence of crab damage. A. pisonii were sampled from the same quadrats from which leaf damage data were collected. Stomach contents were dissected and food items were classified into a number of categories.Species damaged and preferred were determined by comparing relative numbers of mangrove leaf stomata from the three mangrove species in gut contents. Results suggested that both methods provide similar estimates of preference. R. mangle leaves were preferred over those of A. germinans and L. racemosa. The percent of R. mangle leaves with damage was about 20-30 times greater than the other species, and R. mangle leaf stomata were 3 to 20 times more abundant in crab guts compared to leaf stomata of the other species. Gut contents indicated that A. pisonii is omnivorous, that average-sized adult crabs (1.4-1.7-cm width) prefer R. mangle, and that the diet of males is more varied than of females. While use of both percent leaf damage and crab gut contents reliably indicates preference, gut contents may describe better the actual diet and elucidate trends for different size or sex classes within a population.     

Are mangroves in the tropical Atlantic ripe for invasion? Exotic mangrove trees in the forests of South Florida

Two species of mangrove trees of Indo-Pacific origin have naturalized in tropical Atlantic mangrove forests in South Florida after they were planted and nurtured in botanic gardens. Two Bruguiera gymnorrhiza trees that were planted in the intertidal zone in 1940 have given rise to a population of at least 86 trees growing interspersed with native mangrove species Rhizophora mangle, Avicennia germinans and Laguncularia racemosa along 100 m of shoreline; the population is expanding at a rate of 5.6% year⁻¹. Molecular genetic analyses confirm very low genetic diversity, as expected from a population founded by two individuals. The maximum number of alleles at any locus was three, and we measured reduced heterozygosity compared to native-range populations. Lumnitzera racemosa was introduced multiple times during the 1960s and 1970s, it has spread rapidly into a forest composed of native R. mangle, A. germinans, Laguncularia racemosa and Conocarpus erectus and now occupies 60,500 m² of mangrove forest with stem densities of 24,735 ha⁻¹. We estimate the population growth rate of Lumnitzera racemosa to be between 17 and 23% year⁻¹. Populations of both species of naturalized mangroves are dominated by young individuals. Given the long life and water-dispersed nature of propagules of the two exotic species, it is likely that they have spread beyond our survey area. We argue that the species-depauperate nature of tropical Atlantic mangrove forests and close taxonomic relatives in the more species-rich Indo-Pacific region result in the susceptibility of tropical Atlantic mangrove forests to invasion by Indo-Pacific mangrove species.     

Reproductive phenology of a northeast Brazilian mangrove community: Environmental and biotic constraints

Brazil has the third largest area of mangrove in the world, which is widely threatened by anthropogenic pressures. We carried out the first long-term phenological study investigating whether environment and competition for pollinators shape the reproduction of a western mangrove community in Brazil, and provide new information for mangrove conservation. We monitored monthly the flowering and fruiting of Avicennia schaueriana, Conocarpus erectus, Laguncularia racemosa and Rhizophora mangle, the only species composing this mangrove community. We applied circular statistics to detect seasonal trends, null models to test for aggregated, staggered or random flowering patterns, performed correlations between phenophases and climate, and calculated intra-specific phenological synchrony. Each species presented a different flowering pattern, from brief annual to continuous and from regular to irregular, resulting in a bimodal pattern at community level. Fruiting was annual or continuous and seasonally unimodal at community level. Precipitation showed the strongest correlation with reproduction for all species, except L. racemosa. Flowering was randomly distributed among species sharing pollinators and each species presented high intra-specific synchrony. The studied mangrove showed a diversity of flowering patterns despite the low number of species. Annual to sub-annual sequential flowering were prevalent, sustaining the pollinators of species all the year long, while the wind-pollinated species flowered continuously. We provide strong evidence that daylength, rainfall and temperature are driving the flowering and fruiting rhythm of these mangrove species.     

Small Gap Creation in Belizean Mangrove Forests by a Wood–Boring Insect1

We investigated the role of wood-boring insects in the creation of light gaps within mangrove forests. We compared the frequency of gaps caused by wood borers to other gap-forming processes and characterized the physical attributes of light gaps in mangrove forests on small islands in Belize. Methods of quantifying light gaps included aerial photography, ground surveys, and experimental plots. Small light gaps (≤12 m²) were very common in Rhizophom mangle fringe, comprising almost 22 percent of these forests. Rhizophora mangle gaps were smaller than gaps in Avicennia germinans forests. In R. mangle forests, gaps were caused by branch death, and in A. germinans forests, gaps were caused primarily by downed trees. More than 91 percent of the gap-forming branches and boles in the R. mangle fringe were killed by a wood-boring cerambycid beetle, Elaphidion mimeticum, indicating that it is the major cause of small-scale disturbances in these forests. No trees or branches in the A. germinans forest were attacked by this beetle. In R. mangle forests, small gaps had significantly higher light levels and soil temperatures than areas under the closed canopy; however, soil conditions for sulfide concentrations, porewater salinity, and redox potentials were similar in small gaps and under the closed canopy. Survival of R. mangle, A. germinans, and Laguncularia racemosa seedlings was also higher inside these small gaps, indicating their importance in regeneration of mangrove forests. Feeding by wood borers is thus an important type of indirect herbivory in mangrove forests, with a critical role in ecological processes such as gap dynamics.     

Long-Term Growth and Succession in Restored and Natural Mangrove Forests in Southwestern Florida

We compared colonization, growth and succession from 1989 to 2000 in a restored mangrove site and in gap and closed canopy sites in a natural mangrove forest. The restored site was created in 1982 and planted with Rhizophora mangle (≈2 m⁻²) propagules. By 1989, Laguncularia racemosa, with densities up to 12.9 tree m⁻², was a dominant in all plots, although densities were greater at edge plots relative to inner plots, and near open water (west plots) relative to further inland (east plots), and in tall mangrove plots relative to scrub plots. Rhizophora mangle (1989 tree densities about 2 m⁻²) was a codominant in inner and scrub plots, while Avicennia germinans had the lowest densities (<1 tree m⁻²) in all plots. From 1989 to 2000 L. racemosa experienced reduced recruitment and apparent density-dependent mortality of canopy individuals in plots with high initial densities. Scrub plots experienced high rates of colonization by R. mangle and L. racemosa, rapid growth in height of all species (1989–1996), followed by a dieoff of L. racemosa in later years (1997–2000) as the canopy came to resemble that of tall mangrove plots. Colonization and growth rates were lower in gap and closed canopy regions of the natural forest relative to rates in the restored site. After 11 years, densities of L. racemosa were 10–20× lower and R. mangle slightly less in the gap relative to densities in tall mangrove plots in the restored site at the same age. Although the restored stand had converged with the natural forest by 2000 in terms of some factors such as species richness, vegetation cover, litterfall, and light penetration, trees were still much smaller and stem densities much higher. Full development of mature structure and ecological function will likely require decades more development.     

Leaf Longevity,Construction, and Maintenance Costs of Three Mangrove Species Under Field Conditions

This study assessed the effect of leaf age on construction cost (CC) in the mangrove species Avicennia germinans, Laguncularia racemosa, and Rhizophora mangle growing in their natural habitat. Leaf osmolality values were species-specific, the highest in A. germinans (1 693 mmol kg^–1) and the lowest in L. racemosa (1 270 mmol kg^–1). In the three species, contents of chlorophyll (a+b) (Chl_a+b) and nitrogen (N) per unit of leaf area were maximal in adult leaves and tended to decline with age. Leaf mass to leaf area ratio (LMA) and ash content increased during leaf ageing. Similarly, as leaves aged, a significant increase in leaf construction cost per leaf area (CC_a) was observed, while per leaf mass (CC_m) it remained almost constant, suggesting a sustained production of leaf compounds as leaves became older. CC was positively correlated with LMA and heat of combustion (Hc) per leaf area, suggesting differences among species in the quantity and composition of expensive compounds. Leaf half lifetime (t_0.5) showed contrasting values in the three mangrove species (60, 111, and 160 d in L. racemosa, R. mangle, and A. germinans, respectively). Overall, L. racemosa was the species with less expensive leaves to construct while leaves of A. germinans and R. mangle had the highest CC_m and CC_a, respectively. Leaf longevity was positively correlated with the ratio between CC and maximum photosynthetic rate (P _max), clearly showing the existence of a balance between leaf costs and benefits.     

Seedling recruitment patterns in a Belizean mangrove forest: effects of establishment ability and physico-chemical factors

A field study was conducted to evaluate the relative importance of factors affecting seedling establishment and survival on a mangrove-dominated island in Belize. An examination of spatial patterns of seedling relative densities in relation to reproductive adults and physico-chemical conditions provided correlative information on factors affecting mangrove regeneration patterns. Distance from reproductive adults explained 89–94% of the variation in relative density of Rhizophora mangle seedlings, whereas availability of resources (light and NH₄) explained 73–80% of variation in Avicennia germinans seedling relative density. Just after dispersal (December), 89% of the variation in Laguncularia racemosa seedling relative density was attributable to distance from reproductive adults, but 7 months later (July) 74% of the variation was explained by intensity of flooding- and salinity-related stresses. Survivorship (after 2.5 years) of propagules and seedlings of R. mangle and A. germinans transplanted to zones of contrasting physico-chemical conditions demonstrated that: (1) mortality was highest during the establishment phase and major causes were failure to strand before viability was lost, consumption by predators and desiccation; and (2) after establishment, differences in sensitivity to physicochemical stress factors such as flooding (A. germinans) and initial orientation of the seedling axis (R. mangle) exerted a further influence on seedling survival. The results indicate that seedling recruitment in these neotropical forests is strongly influenced by dispersal patterns, differential establishment abilities and effects of physico-chemical factors that vary with elevation and distance from the shoreline.     

Allelopathic effects of fruits of the Brazilian pepper Schinus terebinthifolius on growth, leaf production and biomass of seedlings of the red mangrove Rhizophora mangle and the black mangrove Avicennia germinans

Exotic plant species can negatively affect native flora and fauna by changing the diversity and productivity of the system and altering successional processes. The novel weapons hypothesis has been suggested as one mechanism for invasion and spread of exotic plant species. It states that exotic species with “new weapons”, such as allelopathic chemicals, may have greater impacts on species in the invaded range without co-evolved defense strategies against the exotic chemicals. In Florida, one successful exotic species, Schinus terebinthifolius (Brazilian pepper), has invaded nearly every habitat throughout the state. Past studies have documented allelopathic properties of S. terebinthifolius negatively impacting the growth of native terrestrial flora. Here we document the effect of S. terebinthifolius for the first time on mangrove systems. We examined growth and survival of Rhizophora mangle (red mangrove) and Avicennia germinans (black mangrove) seedlings when exposed to a range of densities of intact and crushed S. terebinthifolius fruits (0, 25, 50) at two different salinities (15, 30 ppt). We also documented the natural density of S. terebinthifolius fruits beneath the tree canopy after fruit production in mangrove habitat. Growth and biomass were significantly reduced in A. germinans when exposed to the highest density of intact S. terebinthifolius fruits growing in 30 ppt saltwater. Trials with intact fruits had no significant effects on R. mangle; however, crushed fruits significantly decreased growth, and leaf production. The ecological importance of the different effects of crushed and intact fruits is supported by field observations which found equal or greater numbers of crushed fruits compared to intact fruits beneath the tree canopy of female S. terebinthifolius. In addition, abiotic factors, such as salinity, may interact with allelopathic chemicals in natural systems and needs to be addressed when planning future allelopathic studies.     

Selective feeding by larval dytiscids (Coleoptera:Dytiscidae) and effects of fish predation on upper littoral zone macroinvertebrate communities of arctic lakes

Carbon and nitrogen stable isotopic data from the primary producers in mangrove ecosystems are needed to investigate trophic links and biogeochemical cycling. Compared with other mangrove species (e.g. Rhizophora mangle) very few measurements have been conducted on the white mangrove, Laguncularia racemosa. The carbon and nitrogen stable isotopic and elemental compositions of L. racemosa were analyzed and compared from Florida and Belize. ¹³C values of L. racemosa from Florida (mean = –26.4) were slightly higher than those from Twin Cays, Belize (mean = -27.4), which may be due to higher salinity in some parts of the Florida site. There was no difference between the ¹⁵N values from L. racemosa from these two sites (Florida mean = 0.6; Belize mean = 0.3), which are indicative of nitrogen derived from nitrogen fixation in a planktonic marine system. However, higher ¹⁵N values from L. racemosa at Man of War Cay in Belize (11.4 and 12.3), which is fertilized by roosting marine birds (14.0), illustrate that L. racemosa can sensitively reflect alternative nitrogen sources. Although the isotopic data could not distinguish between Avicennia germinans, R. mangle and L. racemosa in Belize the L. racemosa had considerably higher C/N ratios (46.5 – 116.1) compared with the Florida samples (42.2 – 76.0) or the other mangrove species. Unlike some previous findings from R. mangle, substrate characteristics (e.g. salinity, NH₄ ⁺, and H₂S) were not related to the isotopic or elemental composition of L. racemosa. ¹³C, ¹⁵N and C/N were analyzed for ecosystem components from L. racemosa habitats at Twin Cays, including other plants (e.g. R. mangle, A. germinans and seagrass), detritus, microbial mats and sediments. Results from mass-balance calculations show that mangrove detritus composes very little of the sediment, which is principally composed of microbial biomass (80 – 90%). Detritus at some sites is also influenced by sources other than that from L. racemosa, including seagrass leaves.     

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.     

Standing crop and aboveground biomass partitioning of a dwarf mangrove forest in Taylor River Slough,Florida

The structure and standing crop biomass of a dwarf mangrove forest, located in the salinity transition zone ofTaylor River Slough in the Everglades National Park, were studied. Although the four mangrove species reported for Florida occurred at the study site, dwarf Rhizophora mangle trees dominated the forest. The structural characteristics of the mangrove forest were relatively simple: tree height varied from 0.9 to 1.2 meters, and tree density ranged from 7062 to 23 778 stems ha^–1. An allometric relationship was developed to estimate leaf, branch, prop root, and total aboveground biomass of dwarf Rhizophora mangle trees. Total aboveground biomass and their components were best estimated as a power function of the crown area times number of prop roots as an independent variable (Y = B × X^–0.5083). The allometric equation for each tree component was highly significant (p<0.0001), with all r² values greater than 0.90. The allometric relationship was used to estimate total aboveground biomass that ranged from 7.9 to 23.2 ton ha^–1. Rhizophora mangle contributed 85% of total standing crop biomass. Conocarpus erectus, Laguncularia racemosa, and Avicennia germinans contributed the remaining biomass. Average aboveground biomass allocation was 69% for prop roots, 25% for stem and branches, and 6% for leaves. This aboveground biomass partitioning pattern, which gives a major role to prop roots that have the potential to produce an extensive root system, may be an important biological strategy in response to low phosphorus availability and relatively reduced soils that characterize mangrove forests in South Florida.     

Fungal diversity and plant disease in mangrove forests: salt excretion as a possible defense mechanism

Keywords. Salt excretion in leaves of some mangrove species may serve as an important defense against fungal attack, reducing the vulnerability of typically high-density, monospecific forest stands to severe disease pressure. In field surveys of a Caribbean mangrove forest in Panama, Avicennia germinans suffered much less damage from foliar diseases than did Laguncularia racemosa or Rhizophora mangle. Similarly, Avicennia leaves supported the least superficial fungal growth, endophytic colonization, and diversity, followed by Laguncularia and Rhizophora. Host specificity of leaf-colonizing fungi was greater than expected at random. We hypothesize that the different salt tolerance mechanisms in the three mangrove species may differentially regulate fungal colonization. The mangroves differ in their salt tolerance mechanisms such that Avicennia (which excretes salt through leaf glands) has the highest salinity of residual rain water on leaves, Laguncularia (which accumulates salt in the leaves) has the greatest bulk salt concentration, and Rhizophora (which excludes salt at the roots) has little salt associated with leaves. The high salt concentrations associated with leaves of Avicennia and Laguncularia, but not the low salinity of Rhizophora, were sufficient to inhibit the germination of many fungi associated with mangrove forests.     

Investigating the role of impoundment and forest structure on the resistance and resilience of mangrove forests to hurricanes

The present study evaluates how a prolonged artificial flooding regime in impounded mangrove forests influences the regeneration after medium-sized forest destruction created by two hurricanes in 2004 in the Indian River Lagoon, Florida, USA. Disturbance patterns, forest structure, and regeneration were investigated. We found disturbed areas, characterized by uprooted and snapped trees, and intact forests in close proximity. The canopy turnover was greater in forests in higher succession stages with median tree diameter at breast height of 7.6 ± 5.7 cm compared to the intact forest with 3.7 ± 1.2 cm. Larger trees with lower densities were more susceptible to hurricane damage. We observed that regeneration of the open patches was dominated by the flood-tolerant species Rhizophora mangle (89.9%) instead of the faster-growing pioneer species Laguncularia racemosa (7.0%). Some of the disturbed areas created by the hurricanes were not recolonized. The regeneration rate of the disturbed areas expressed by vegetation closure >50% differed between almost zero to 100% in three different impoundments. We concluded that the artificial flooding regime imposed by impoundment is the predominant selective force in the successive process according to the species composition. However, we were not able to detect the driving factors that prevented mangrove establishment in disturbed areas.     

Mangrove litter dynamics in La Mancha Lagoon,Veracruz, Mexico

In the mangrove surrounding the coastal lagoon of La Mancha, Veracruz, Mexico, we studied litter fall, litter standing crop, and turnover rates in four different mangrove settings, based on the ecological classification of Lugo and Snedaker (1974). We studied those three prominent ecological processes at the basin, fringe and riverine mangrove settings, being the last one a relict riverine stand. The aim was to describe and compare litter dynamics among mangrove types in a lagoon with an ephemeral inlet, as a way of understanding functional heterogeneity within this coastal ecosystem. The daily average values of litter fall were different (P < 0.01) among mangrove site basin I, formed by Avicennia germinans and Rhizophora mangle (2.35 g/m²/day); basin II, formed by Laguncularia racemosa, Avicennia germinans, and Rhizophora mangle (2.93 g/m²/day); fringe with Rhizophora mangle (2.13 g/m²/day); and relic riverine, also with R. mangle (4.70 g/m²/day). The amount of litter standing crop was different among sites (P < 0.001), and also between the dry and rainy season, for each mangrove type (P < 0.001). Turnover ratios were higher in basin I and basin II sites (6.34 and 7.44 times per year) than in relic riverine and fringe mangroves (1.49 and 2.39 times per year). Interstitial salinity and sediment nutrients varied among mangrove types and could influence litter production. Since this lagoon has an ephemeral inlet, continuous inundation throughout 7–8 months per year has an important effect on litter dynamics.     

Pollen morphology of the shrub and arboreal flora of mangroves of Northeastern Brazil

The Brazilian coast, especially in the Northeastern one, possesses a vast area of mangrove which acts as the interface that links terrestrial ecosystems and estuaries. After an inventory of the shrub and tree species present in the mangroves of Northeastern Brazil, polliniferous material was collected for morphological study of the pollen grains, which were acetolysed, measured and analyzed using light and scanning electron microscopy. Fourteen species belonging to seven families were grouped as restricted and peripheral, according to their position into the mangrove vegetation. The restricted species are six: Avicennia germinans and A. schaueriana (Avicenniaceae), Rhizophora harrisonii, R. mangle and R. racemosa (Rhizophoraceae), Laguncularia racemosa (Combretaceae). The peripheral species are eight: Annona glabra (Annonaceae), Conocarpus erectus (Combretaceae), Dalbergia ecastophyllum, Inga subnuda, Sophora tomentosa (Leguminosae), Hibiscus pernambucensis, H. tiliaceus (Malvaceae) and Suriana maritima (Surianaceae). All species had their pollen grains described and illustrated in that first atlas of the pollen flora of the Brazilian mangroves.     

Mangrove habitat partitioning by Ucides cordatus (Ucididae): effects of the degree of tidal flooding and tree-species composition during its life cycle

Environmental factors strongly affect mangrove crabs, and some factors modulate population structure and habitat partitioning during the crabs’ life cycle. However, the effect of these environmental factors on habitat selection by mangrove crabs is still unknown. We evaluated habitat selection by the mangrove crab Ucides cordatus in mangrove forests with different degrees of predominance of Rhizophora mangle, Laguncularia racemosa or Avicennia schaueriana, two tidal flooding levels (less- and more-flooded), and two biological periods (breeding and non-breeding seasons). Sampling was conducted in four mangrove forests with different influences of these biotic and abiotic parameters. We used the data for sex ratio to explain environmental partitioning by this species. Females predominated in R. mangle mangroves, independently of the biological period (breeding or non-breeding seasons), and males predominated only in the less-flooded L. racemosa mangroves. The flooding level affected the sex ratio of U. cordatus, with a predominance of males in less-flooded mangroves, independently of the biological period; and a gender balance in the more-flooded mangroves only during the breeding season. Outside the breeding season, the largest specimens were recorded in the R. mangle mangroves, but in the breeding season, the largest crabs were recorded in the L. racemosa mangroves with a higher level of flooding. These results suggest that tree-species composition and tidal flooding level can have a significant effect on the habitat partitioning of sexes and sizes of the mangrove crab U. cordatus both during and outside the breeding season.