Pollen fertility and intraspecific and interspecific compatibility in mangroves of Fiji

 Pollen fertility and compatibility status were investigated in three species of mangroves: Bruguiera gymnorhiza, Rhizophora samoensis, R. stylosa and a hybrid between the two Rhizophora species. Pollen fertility was greater than 80% in the three species but less than 10% in the hybrid. Also, all three species were intraspecifically compatible, while the hybrid showed very low compatibility which may be due to its low pollen fertility. The genera Bruguiera and Rhizophora were found to be completely cross incompatible, while a low reciprocal cross compatibility was observed in interspecific crosses between Rhizophora stylosa and R. samoensis. Received: 5 June 1997 / Revision accepted: 10 October 1997     

Molecular phylogeny of mangroves VIII: Analysis of mitochondrial DNA variation for species identification and relationships in Indian mangrove Rhizophoraceae

Ten species belonging to four genera of the mangrove tribe Rhizophoreaefound in the Indian subcontinent were analysed for species identification andgenetic relationship using nine mitochondrial gene probes. RFLP patternobservedwith 27 probe enzyme combinations the genera Rhizophora,Bruguiera, Ceriops andKandelia differentiated these species into three classesofmitotypes with further resolution within them. Clustering of these mitotypesindicated that Rhizophora was more closely related toCeriops-Kandelia than to theBruguiera. Though the component species of each genusclustered together, a high degree of heterogeneity was observed among fourspecies of the genus Rhizophora and three species of genusBruguiera. The variation between two species ofCeriops was minimal. Species-specific profiles wereobserved for all the species in some probe-enzyme combination. Though themonotypic genus Kandelia shared a number of loci withgenusCeriops, it remained distinct. The putative parents of thenaturally occuring interspecific hybrid in Pichavaram were reconfirmed to beR. apiculata and R. mucronata. Theresults are discussed with regard to the taxonomic and phylogeneticrelationships between different species and genera of the tribe Rhizophoreae.     

Litter production and seasonality of mangroves in Papua New Guinea

Productivity, as measured by litter trapping, of a mangrove dominated by Rhizophora stylosa Griff. at Motupore Island (lat. 9° 31′34′?S; long. 147° 17$\text{\$}\text{?}$02‘ E) was 1430 g dry weight m^?2 year^?1. This is a high figure when compared with other studied mangrove sites. Phenological data were obtained for R. stylosa, R. apiculata Blume and Sonneratia alba Smith. All species have a major leaf fall in the wet season. Flowering in both Rhizophora species is greatest during the wet season, but S. alba peaks during the dry season. Leaf emergence data were only available for R. stylosa and these appeared to be trimodal. Correlations were calculated for each component with rainfall, wind run, maximum temperature and minimum temperature. Some significant correlations were obtained with rainfall and maximum temperature. These correlations are discussed and it is suggested that the seasonality of mangrove phenologies needs to be considered in terms of whole tree physiology rather than isolated environmental factors.     

Applications of Multiple Nuclear Genes to the Molecular Phylogeny,Population Genetics and Hybrid Identification in the Mangrove Genus Rhizophora

The genus Rhizophora is one of the most important components of mangrove forests. It is an ideal system for studying biogeography, molecular evolution, population genetics, hybridization and conservation genetics of mangroves. However, there are no sufficient molecular markers to address these topics. Here, we developed 77 pairs of nuclear gene primers, which showed successful PCR amplifications across all five Rhizophora species and sequencing in R. apiculata. Here, we present three tentative applications using a subset of the developed nuclear genes to (I) reconstruct the phylogeny, (II) examine the genetic structure and (III) identify natural hybridization in Rhizophora. Phylogenetic analyses support the hypothesis that Rhizophora had disappeared in the Atlantic-East Pacific (AEP) region and was re-colonized from the IWP region approximately 12.7 Mya. Population genetics analyses in four natural populations of R. apiculata in Hainan, China, revealed extremely low genetic diversity, strong population differentiation and extensive admixture, suggesting that the Pleistocene glaciations, particularly the last glacial maximum, greatly influenced the population dynamics of R. apiculata in Hainan. We also verified the hybrid status of a morphologically intermediate individual between R. apiculata and R. stylosa in Hainan. Based on the sequences of five nuclear genes and one chloroplast intergenic spacer, this individual is likely to be an F1 hybrid, with R. stylosa as its maternal parent. The nuclear gene markers developed in this study should be of great value for characterizing the hybridization and introgression patterns in other cases of this genus and testing the role of natural selection using population genomics approaches.     

Mangrove community structure and regeneration potential on a rapidly expanding,river delta in Java

Key message

Mangroves in rapidly expanding Southeast Asian river deltas form floristically simple zones dominated by a few highly regenerative species adaptable or tolerant to rapid sedimentation and extensive river flooding.

Abstract

The size class distribution, community composition and spatial structure of five representative mangrove forests in the rapidly expanding Cimanuk river delta on Java were determined. These deltaic forests are species-poor (eight true mangrove species) and spatially segregated into three distinct floristic zones: (1) a fringing, low intertidal zone co-dominated by Avicennia marina and A. officinalis, with less abundant Bruguiera parviflora, Rhizophora apiculata, and R. mucronata; (2) a zone transitional between the low and mid intertidal in which Avicennia and Rhizophora spp. co-dominate; and (3) a mid intertidal zone dominated by R. mucronata and R. apiculata. Numerically dominated by seedlings (52,500–73,500 seedlings ha^?1) and saplings (5,268–5,660 saplings ha^?1), all five forests are relatively young and actively regenerating. Positive correlations of tree stem diameter and tree height with soil organic matter and P concentrations, salinity, the soil C/N ratio, pH, and silt/clay composition highlight the importance of soil factors in sustaining forest growth. The low diversity and relative structural simplicity of these rapidly growing and regenerating forests may be attributed to adaptation or tolerance to flooding and the rapid sedimentation and seaward expansion of the delta.     

Effects of light and intertidal position on seedling survival and growth in tropical tidal forests

Reciprocal transplant experiments were used to study the effects of tidal inundation and light level on growth and survival of four species of mangroves in Australia: Avicennia marina (Forsk.) Vierh., Bruguiera gymnorrhiza (L.) Lam., Ceriops tagal C.T. White, and Rhizophora stylosa Griff. Seedlings were planted in the high or low intertidal and in light gaps or under the shade of a closed canopy. Survival and growth of the seedlings were monitored for 30–36 months. Significant differences in survival were found among species, between intertidal zones and due to light level. Averaged across intertidal zones and light level, survival was greatest for Rhizophora and decreased in the order Ceriops, Avicennia, and Bruguiera. For all species survival was greater (P ⩽ 0.001) in the high than in the low intertidal treatment, regardless of light level. Within the high intertidal all species survived better in light gaps than under the canopy. Relative growth of Rhizophora stylosa, Avicennia marina, and Ceriops tagal were greater in the high versus low intertidal and in gaps versus under the canopy. For Bruguiera gymnorrhiza growth was not significantly different between gap and canopy or high and low intertidal. Although Avicennia marina, Bruguiera gymnorrhiza, and Rhizophora stylosa survived and/or grew best in the high intertidal they reach maximum abundance in the low intertidal. Ceriops tagal, however, performed best in the region where it is most abundant, yet even there, it was out-performed by the former three species. These results indicate that the species zonation patterns often observed across the intertidal cannot be explained by physiological adaptation alone. Factors such as propagule dispersal, competition and predation on propagules may also be important.     

Pattern and process in mangrove forests of the Andaman Islands

Abatract The structural and functional aspects of mangrove forests of the Andaman Islands were described. The mangrove forests of Andaman are highly diversified and rich in species composition. Twenty-six species were collected, Rhizophora lamarckii and R. stylosa were reported for the first time from these islands. The mangroves of Andaman are very productive in comparison to other forest types. Maximum production of biomass was found in the undisturbed Oralkatcha forest. Maximum litter fall was also found at Oralkatcha.     

Fungal endophyte communities in four Rhizophoraceae mangrove species on the south coast of China

This study investigated the taxonomic identities and diversity of fungal endophytes isolated from four Rhizophoraceae mangrove plant species, Ceriops tagal, Rhizophora apiculata, R. stylosa and Bruguiera sexangula var. rhynchopetala, using a combination of morphological and molecular approaches. Two hundred ninety-five isolates were classified into 38 taxa by morphological characteristics. The representative 38 isolates from each taxa were selected for further molecular identification using nuclear ribosomal DNA sequences, including both the internal transcribed spacers (ITS1 and ITS2) and the 5.8S gene region. The 38 representative endophytes were identified to various taxonomic levels. These results suggest that Pestalotiopsis and Phomopsis were the most frequent endophytes in the four host species. Some of the endophytes exhibit host and tissue specificity. The colonization frequencies of endophytic fungi in the stems of the four host plants are evidently higher than in the roots. The four Rhizophoraceae mangrove species have low similarities of endophyte communities.     

Human hydrographical changes interact with propagule predation behaviour in Sri Lankan mangrove forests

For mangroves with almost no tidal influence and a subsequent mosaic vegetation structure we describe the relation between propagule predators and both vegetation structure and environmental factors on a vegetation assemblage or ‘forest patch’ level. We then report the interaction between natural propagule predation and human influence. In different forest patches dominated by Avicennia officinalis, Excoecaria agallocha, Lumnitzera racemosa, Rhizophora apiculata or R. mucronata or in open or mixed areas identified from aerial photographs, the predation on propagules of A. officinalis, B. gymnorrhiza, R. apiculata and R. mucronata was monitored in a total of 24 experimental plots (3 per forest patch). Relationships to environmental factors (topography, water level, rainfall and season) on the level of predation were investigated. The highest predation intensity by crabs was on A. officinalis propagules, and in E. agallocha forest patches. Predation by crabs, snails, insects and mammals was observed, and the significant differences in predation intensity across species and across forest patches were explained directly by propagule predator identity and abundance, and by the interaction between rainfall, water level and microtopography. We illustrated how hydrography changes alter the behaviour of propagule predators and play a role in the shaping of vegetation structure by reconstructing the lagoon water level of a site over a period of 50 years based on rainfall data. This time frame covered both a condition without and with human impacts, and past and present propagule predation settings were confronted with the simulated hydrology. The importance of spatial and temporal microhabitat variations in opening multiple successional pathways in vegetation dynamics is illustrated, and is highly relevant for ecosystems with unpredictable or short-lived (< 20 years) patchy vegetation structures and microhabitats.     

Distinct responses of growth and respiration to growth temperatures in two mangrove species

Background and AimsMangrove plants are mostly found in tropical and sub-tropical tidal flats, and their limited distribution may be related to their responses to growth temperatures. However, the mechanisms underlying these responses have not been clarified. Here, we measured the dependencies of the growth parameters and respiration rates of leaves and roots on growth temperatures in typical mangrove species.MethodsWe grew two typical species of Indo-Pacific mangroves, Bruguiera gymnorrhiza and Rhizophora stylosa, at four different temperatures (15, 20, 25 and 30 °C) by irrigating with fresh water containing nutrients, and we measured growth parameters, chemical composition, and leaf and root O₂ respiration rates. We then estimated the construction costs of leaves and roots and the respiration rates required for maintenance and growth.Key ResultsThe relative growth rates of both species increased with growth temperature due to changes in physiological parameters such as net assimilation rate and respiration rate rather than to changes in structural parameters such as leaf area ratio. Both species required a threshold temperature for growth (12.2 °C in B. gymnorrhiza and 18.1 °C in R. stylosa). At the low growth temperature, root nitrogen uptake rate was lower in R. stylosa than in B. gymnorrhiza, leading to a slower growth rate in R. stylosa. This indicates that R. stylosa is more sensitive than B. gymnorrhiza to low temperature.ConclusionsOur results suggest that the mangrove species require a certain warm temperature to ensure respiration rates sufficient for maintenance and growth, particularly in roots. The underground temperature probably limits their growth under the low-temperature condition. The lower sensitivity of B. gymnorrhiza to low temperature shows its potential to adapt to a wider habitat temperature range than R. stylosa. These growth and respiratory features may explain the distribution patterns of the two mangrove species.     

5种红树科植物叶片的比较解剖及其生态适应研究

用石蜡切片法对红树（Rhizophora apiculata Bl.）、红海榄（R.stylosa Griff.）、秋茄（Kandelia candel(Linn.) Druce）、木榄（Bruguiera gymnorrhiza(Linn.) Savigny）和海莲（B.sexangula(Lour.) Poir.）等5种红树科植物的叶片进行解剖学的观察和研究,结果表明：5种红树科植物叶片的表皮均有厚的角质膜和下皮层,表皮和下皮层的细胞常含有单宁;上表皮无气孔器;栅栏组织多层;木质部非常发达。说明了红树科植物的叶片具有很强的耐旱能力;同属植物的叶片在解剖结构上有着相似的特征。     

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.     

Isolation and characterization of RARE-1, a Ty1/copia-like retrotransposon domesticated in the genome of Rhizophora apiculata

Long terminal repeat (LTR) retrotransposons are predominant mobile elements that play important roles in plant genome evolution. Here, we isolated the first putative complete Ty1/copia-like retrotransposon of 6303 bp in mangrove Rhizophora apiculata, named RARE-1. RARE-1 was homologous to the soybean retroelement 1 (SORE-1) and exhibited abundant cis-regulatory motifs involved in various stress responses in its LTRs. Using the sequence-specific amplification polymorphism (S-SAP) technique, we obtained a total of 112 bands for two R. apiculata populations from Hainan, China and Ranong, Thailand. The Hainan population showed slightly higher S-SAP polymorphism but fewer unique bands than the Ranong population. Moreover, the Hainan population also had significantly more copies of RARE-1 than the Ranong population as revealed by quantitative real-time PCR (qPCR). Our results suggested that RARE-1 might have been domesticated in the R. apiculata genome, as a result of the long-term evolution of mangroves under the extreme environment.     

Isolation and characterization of microsatellite loci in the red mangrove Rhizophora mangle (Rhizophoraceae) and its related species

Fourteen microsatellite markers were isolated from the red mangrove Rhizophora mangle (Rhizophoraceae), a widely distributed mangrove plant in the New World and West Africa. The range of expected heterozygosity of these markers was 0.000–0.672 in the two populations of R. mangle. Cross-species testing was examined for five other species of Rhizophora, and Kandelia candel and Bruguiera gymnorrhiza. All 14 markers could be amplified in R. samoensis, thirteen in R. racemosa, and six markers in all other species of Rhizophora. Our findings greatly increase the utility of these markers.     

Chloroplast microsatellite markers for the mangrove tree species Bruguiera gymnorrhiza, Kandelia candel, and Rhizophora stylosa, and cross-amplification in other mangrove species

Chloroplast microsatellite (cpSSR) markers were developed for three ecologically and economically important tree species in the mangrove family, Rhizophoraceae: Bruguiera gymnorrhiza, Kandelia candel, and Rhizophora stylosa. Noncoding regions of chloroplast DNA (cpDNA) from each species were separately amplified using universal chloroplast primers. Six, two, and three polymorphic cpSSR loci in B. gymnorrhiza, K. candel, and R. stylosa, respectively, were developed from amplified noncoding cpDNA regions. Characterization of 216, 156, and 253 individuals of B. gymnorrhiza, K. candel, and R. stylosa, respectively, collected from different natural mangrove populations (B. gymnorrhiza, 9; K. candel, 7; R. stylosa, 9) on Iriomote Island in Japan showed that these loci provide cpSSR markers with polymorphisms ranging from two to four alleles per locus and gene diversity between 0.027 and 0.480. These cpSSR markers will be useful for analyzing the maternal lineage distributions and population genetic structures of the three species. Several of these markers may also be useful in similar studies of other mangrove species.     

Effects of artificial grassland establishment on soil nutrients and carbon properties in a black-soil-type degraded grassland

Despite a growing knowledge of nutrient limitation for mangrove species and how mangroves adapt to low nutrients, there is scant information about the relative importance of N:P ratio and leaf phenolics variability in determining nutrient conservation. In this study, we evaluated possible nutrient conservation strategies of a mangrove Rhizophora stylosa under nutrient limitation. 1. The leaf nutrient concentrations of R. stylosa changed with season, with the highest N concentration in winter and the highest P concentration in spring for both mature and senescent leaves. Leaf N and P concentrations decreased significantly during leaf senescence. Based on N:P ratios R. stylosa forest was N-limited. Accordingly, the nitrogen resorption efficiency (NRE) was significantly higher than phosphorus resorption efficiency (PRE) for the R. stylosa leaves during leaf senescence. The NRE and PRE both reached the highest in the autumn. Average N and P concentrations in the senescent leaves were 0.15% and 0.06% for R. stylosa, respectively, indicating a complete resorption of N and an incomplete resorption of P. There was a significant negative correlation between nitrogen resorption proficiency (NRP) and NRE, meanwhile phosphorus resorption proficiency (PRP) and PRE correlation was also highly significantly. 2. R. stylosa leaves contained relatively high tannin level. Total phenolics, extractable condensed tannins and total condensed tannins contents increased during leaf senescence, and changed between seasons. The lowest concentrations of total phenolics, extractable condensed tannins and total condensed tannins occurred in summer, total phenolics concentrations were inversely related to nitrogen or phosphorus concentrations. 3. Our results confirmed that resorption efficiency during leaf senescence depends on the type of nutrient limitation, and NRE was much higher than PRE under N-limited conditions. R. stylosa forest developed several nutrient conservation strategies in the intertidal coastline surroundings, including high nitrogen resorption efficiency, low nutrient losses and high tannins level.     

Mangrove sesarmid crab feeding experiments in Peninsular Malaysia

The feeding ecology of mangrove sesarmid crabs in Peninsular Malaysia was investigated by field and laboratory experiments using four mangrove leaf species (Avicennia officinalis, Bruguiera gymnorrhiza, B. parviflora and Rhizophora apiculata) and leaves of different condition (fresh and senescent). Leaves tethered on strings at high (Bruguiera zone) and low (Rhizophora zone) intertidal positions, both upstream (Sungai Pasir) and downstream (Lower Merbok) showed significant amounts of leaf litter removal in 24 h (mean 79±3% initial dry mass). Significantly more B. gymnorrhiza was consumed in Bruguiera zones and significantly less senescent A. officinalis in the upstream Rhizophora zone. In Bruguiera zones, significant numbers of leaves were taken down burrows but there were no preferences for leaf species or condition of leaf taken down burrows at all sites. In 24 h, under laboratory conditions, the sesarmid crabs Sesarma (Perisesarma) eumolpe and S. (Perisesarma) onychophorum were offered with a mangrove species choice of either fresh or senescent leaves. There was no difference in mangrove species taken when the leaves were senescent for both crab species, but when the leaves were fresh, significantly more A. officinalis leaves were consumed by both sesarmid crab species. S. onychophorum ate significantly more B. parviflora than did S. eumolpe. The crab distribution in the field was related to the preferred tree species dominance, indicating that tree species may be important for crab species distribution, or vice versa. The mean rate of leaf consumption was not significantly different between the crab species; S. eumolpe was 29.9±5.9 and S. onychophorum was 35.3±7.2 mg dry mass per wet mass gram of crab in 24 h. Rhizophora spp. were the least preferred species in all feeding experiments, a finding which may have implications for ecosystem functioning in monoculture rehabilitation projects.     

天然红海榄支柱根空间结构研究

通过对红海榄及其支柱根的定位,利用角尺度、大小比数和最小距离3个参数对特呈岛不同群落的红海榄支柱根空间结构进行研究。结果表明,红海榄纯林、红海榄+白骨壤、红海榄+木榄混交林3种不同群落中红海榄支柱根的平均角尺度分别是0.772、0.692和0.636,大于均匀分布的取值范围［0.475,0.517］,属于聚集分布格局;支柱根的直径大小比数和高度大小比数均接近0.5,从直径大小比数来看,在参照支柱根与相邻支柱根的结构单元中,径级分布较均匀;高度大小比数和直径大小比数在描述红海榄各参照支柱根与相邻根之间的生长优势方面具有较强的一致性;3种群落中红海榄支柱根的最小距离分别是2.83、1.00和2.24 cm,平均距离分别是10.54、9.07和8.63 cm。3个结构参数量化了参照支柱根与其相邻根之间的空间关系,对红海榄的有效保护及合理利用具有一定的实际指导意义。     

The abundance and growth of Avicennia marina and Rhizophora stylosa in the low shore zone of Darwin Harbour,Northern Territory

Abstract The growth and distribution of the two mangrove seedling species, Avicennia marina (Forsk.) Vierh. and Rhizophora stylosa Griff., were studied in the low-shore zone at three locations along two estuaries of Darwin Harbour to determine how growth varied along the estuary and within the habitats of the low shore zone. The low shore environment was heterogenous for factors such as salinity and light, with substantial variation within and between locations. Seedlings of Avicennia were in higher densities in light gaps than in the adjacent forest, and density in both habitats decreased towards the mouth of the estuary. Growth of Avicennia seedlings, as measured through in situ growth trials and plant harvests, increased towards the mouth of the estuary. Results for Rhizophora seedlings were more complex although some similar trends were evident. This may have been due to the greater early survival of Rhizophora seedlings compared to Avicennia seedlings. The importance of propagule origin was investigated, for Rhizophora, by transplanting propagules between locations along one estuary. Increased seedling survival was observed at the propagule source site; however, this effect was subtle and probably not the most important factor affecting seedling survival at a site.     

Salinity tolerance in some mangrove species from Pakistan

Growth, ionic and water relations of three mangrove species viz. Avicennia marina, Ceriops tagal and Rhizophora mucronata werestudied in different seawater concentrations (0, 25, 50, 75 and 100%).All mangrove species showed optimal growth at 50% seawater. Relativelymore biomass was accumulated by R. mucronata while C. tagalhad the tallest individuals. Tissue water potential became more negativewith the increase in salinity and stomatal conductance was decreased in allplants. Higher stomatal conductance was noted in R. mucronata,followed by A. marina and C. tagal. Sodium and chloride ionsincreased with the increase in salinity and this accumulation was muchhigher in A. marina.