Consequences of pre-dispersal damage by insects for the dispersal and recruitment of mangroves

Herbivores may enhance plant recruitment, but such positive interactions may be overlooked in favour of obvious negative effects of herbivory on propagules. My objective was to determine whether larval insects that feed and develop within fruit of the mangrove Avicennia marina act as mutualist herbivores by increasing the dispersal of propagules without affecting their viability and emerging successfully as adults following dispersal of the propagule by water. Surveys revealed that frugivory is common throughout the mangrove forest, and fruit had up to six exit holes where larvae had emerged as adults. Larval insects did not affect the flotation of propagules with pericarps, a thin structure that provides buoyancy for dispersal by water. In contrast, after simulating germination by removing the pericarp, the majority of propagules with three exit holes floated on average for 20 h longer than those without exit holes, which sank immediately. Based on this evidence that frugivory could increase the dispersal potential of propagules, I predicted that propagules consumed by larval insects should disperse farther than undamaged propagules, and this was tested by quantifying the potential viability of propagules stranded on beaches at increasing distances (up to 20 km) from mangrove forests. Flies and moths emerged as adults after being transported tens of kilometres within mangrove propagules, revealing a novel mode of dispersal. Proportionally fewer potentially viable propagules were supplied to beaches at increasing distances from mangrove forests, however, indicating that larval insects negatively affect recruitment and are thus not acting as mutualist herbivores. Nevertheless, when transported back to the mangrove forest, seedlings established from propagules damaged by larval insects and stranded on beaches. Therefore, although frugivory does not preclude mangrove recruitment, its negative effects in the pre-dispersal environment may be intensified with increasing dispersal distance, thus limiting the long-distance supply of propagules and recruitment of mangroves.     

The current status of mangroves along the Kenyan coast: a case study of Mida Creek mangroves based on remote sensing

Mangroves form a unique ecosystem of limited extent covering an area of about 53 000 ha along the Kenyan coast which need protection from overexploitation. Background information on the areal extent and status of these mangroves is limited and makes their protection and management difficult.A model study has been carried out on the Mida Creek mangroves based on a double sampling approach starting with SPOT multispectral satellite imagery followed by ground checks to provide information on species composition, density and distribution of mangroves. The utilization of mangroves and uses around and within the mangroves were determined.During this study, seven mangrove species were identified in Mida Creek, namely:Avicennia marina, Bruguiera gymnorrhiza, Ceriops tagal, Rhizophora mucronata, Sonneratia alba, Xylocarpus granatum andLumnitzera racemosa. This report briefly outlines the present utilisation of mangrove species. It is recommended that a multidisciplinary management plan be developed in order to conserve and manage the mangroves of Kenya on a sustainable yield basis.     

福建漳江口红树林区秋茄幼苗生长动态

通过福建漳江口红树林自然保护区内8个样方24个小样方人工种植600个秋茄胚轴,在3a时间内对秋茄胚轴建立、幼苗生长以及环境因子进行定期观测。研究结果表明：林缘空地的秋茄生长状况良好,白骨壤林内最不利于秋茄幼苗的生长。潮位、盐度、底质土壤理化因子不是造成该样地各样方间秋茄幼苗生长差异的主要原因。动物取食、光照状况以及种间竞争是限制秋茄生长的主要环境因子。秋茄胚轴在长根前易于随潮水漂走,底质土壤中自骨壤致密的根系抑制了秋茄胚轴的定植,导致白骨壤林内秋茄幼苗漂走的数量最多。昆虫和螃蟹等动物的取食是导致林内已经固着生长的秋茄幼苗大量死亡的最主要原因,而林外被取食的幼苗个体极少。此后秋茄幼苗能否继续成长,主要取决于幼苗所接受到的光照条件。3a后,在荫蔽的树冠下,秋茄幼苗无法存活;而在林外,秋茄幼苗已经长成幼树。在林外滩涂上迅速生长的互花米草,也将影响秋茄幼苗的更新和生长。     

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.     

Early growth responses of mangroves to different rates of nitrogen and phosphorus supply

Experiments were conducted in an outdoor facility to quantify growth responses of six mangrove species to rates of dissolved inorganic nitrogen and phosphorus supply mimicking the range of N and P mineralization rates in natural soils. Growth of all six species on nitrogen was nonlinear. Stem extension rates of Rhizophora apiculata and Xylocarpus granatum were enhanced to the highest rate of N supply (50 mmol m^− 2 d^− 1); Bruguiera gymnorrhiza, Avicennia marina, and Xylocarpus moluccensis stem growth leveled off by 10 mmol m^− 2 d^− 1. Stem growth of Ceriops tagal peaked at 24-26 mmol N m^− 2 d^− 1. Except for A. marina and C. tagal, rates of biomass increase declined at the highest supply rate, indicating NH₄⁺ toxicity. At different rates of P supply, stem extension rates and rates of biomass increase of R. apiculata and C. tagal best-fit Gaussian curves and B. gymnorrhiza stem growth and biomass increase best-fit sigmoidal and Gaussian curves, respectively; X. moluccensis stem and biomass growth increased linearly, but stem and biomass growth rates of A. marina did not vary in relation to P supply. Stem growth of X. granatum was Gaussian but rates of biomass increase best-fit a quadratic equation. Changes in leaf and root N and P content mirrored the growth responses. As rates of N and P mineralization in natural mangrove soils overlap with the lowest rates of N and P supplied in these experiments, the growth responses imply that mangroves are intrinsically nutrient-limited at mineralization rates often encountered in nature. Such species specificity may have significant implications for recruitment success and the establishment of species gradients within mangrove forests.     

The growth of Kandelia candel seedlings in mangrove habitats of the Zhangjiang estuary in Fujian, China

Propagules of Kandelia candel collected from the Zhangjiang estuary were planted in mangrove habitats along the intertidal gradient. The rooting rates of K. candel propagules varied spatially. The lowest rate occurred in Avicennia marina forest (69.7%). The rates were higher in K. candel forest (90.0%), at the fringe of the mangrove forest (89.3%) and on the bare tidal flat outside the mangrove forest (82.7%). After one year, the survival rates of seedlings planted under A. marina forest, K. candel forest, at the fringe of the mangrove forest, and on the bare tidal flat were 13.7%, 54.7%, 76.0%, and 34.7%, respectively. Among the surviving K. candel seedlings, those at the fringe of the mangrove forest and on the bare tidal flat had greater height, stem diameter, leaf number, leaf area, and biomass than those under A. marina and K. candel forests. These results demonstrated that establishment and growth of K. candel seedlings occurred successfully at the fringe of the mangrove forest, but were the worst under A. marina forest. The performance of K. candel seedlings was independent of physico-chemical characters of sediment. However, interspecies competition, propagule predation by insects and crabs, and the incident light had significant effects on seedling survival and growth.     

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.     

Stable soil pH of thai mangroves in dry and rainy seasons and its relation to zonal distribution of mangroves

Soil characteristics of mangrove forests at Amphur Laemngob, Trat Province, east Thailand were investigated in both dry andrainy seasons. Sonneratia alba, Avicennia alba andRhizophora mucronata mainly grew as seaward mangroves.Rhizophora apiculata, Ceriops tagal, Excoecaria agallocha andBruguiera gymnorrhiza grew as meso mangroves in most cases.Lumnitzera racemosa was found as a landward mangrove. The salinity of the soil in which these 8 species grew was significantly higher in the dry season than in the rainy season. Soil pH did not change irrespective of the season. Each species seemed to grow at inherent soil pH. Bruguiera gymnorrhiza, Sonneratia alba andAvicennia marina were found both in Japan and Thailand. The soil pH of each mangrove species was similar in Japan and Thailand. The soil salinity of each mangrove species in Japan was similar to that in Thailand in the dry season. It was suggested that soil pH and salinity in the dry season are the important factors governing the zonal distribution of mangroves.     

Soil salinity and pH in Japanese mangrove forests and growth of cultivated mangrove plants in different soil conditions

Soil conditions of mangrove forests in southern Japan were found to correlate largely with zonal distributions of the species.Kandelia candel grew in soils with low salinity and low pH,Avicennia marina, Rhizophora stylosa andSonneratia alba in soils with high salinity and high pH, andBruguiera gymnorrhiza in soil with a wide range of pH but limited range of salinity.Lumnitzera racemosa colonized soil with a wide range of pH and medium salinity. Seedlings ofKandelia candel, Bruguiera gymnorrhiza andRhizophora stylosa were planted in soils with differing salinity and pH. Optimum seedling growth ofKandelia, Bruguiera andRhizophora occurred when plants were cultivated in soils similar to those of their natural habitats, suggesting that growth of mangrove species and their zonal distributions were regulated by salinity and soil pH.     

The growth of Kandelia candel seedlings in mangrove habitats of the Zhangjiang estuary in Fujian, China

Propagules of Kandelia candel collected from the Zhangjiang estuary were planted in mangrove habitats along the intertidal gradient. The rooting rates of K. candel propagules varied spatially. The lowest rate occurred in Avicennia marina forest (69.7%). The rates were higher in K. candel forest (90.0%), at the fringe of the mangrove forest (89.3%) and on the bare tidal flat outside the mangrove forest (82.7%). After one year, the survival rates of seedlings planted under A. marina forest, K. candel forest, at the fringe of the mangrove forest, and on the bare tidal flat were 13.7%, 54.7%, 76.0%, and 34.7%, respectively. Among the surviving K. candel seedlings, those at the fringe of the mangrove forest and on the bare tidal flat had greater height, stem diameter, leaf number, leaf area, and biomass than those under A. marina and K. candel forests. These results demonstrated that establishment and growth of K. candel seedlings occurred successfully at the fringe of the mangrove forest, but were the worst under A. marina forest. The performance of K. candel seedlings was independent of physico-chemical characters of sediment. However, interspecies competition, propagule predation by insects and crabs, and the incident light had significant effects on seedling survival and growth.