Studies of prop-root damage from wood-boring animals are primarily focussed upon sphaeromid isopods (Perry, 1988). In Clifton, J., et al. CAS  frequency of herbivore attack and chemical defence investment (Brooks & Bell, 2002). However, significant differences were found with tissue regrowth among the root treatments (GLM, F Parenchyma cell differentiation may change from generation of parenchyma to generation of cork and sclerenchyma cells (Bloch, 1952; Wier et al., 1996). Rhizophora stylosa. Sal-voza), and together they characterize most stands of IWP stilt mangroves. This was also apparent with naturally occurring severely damaged roots, as evidenced by the many teredinid tunnels. Stilt roots arises from the trunk or branches of the mangrove and grows toward the soil where the stilt root will develop an underground root system. Digital images were taken of each treated root over 12 months. Schimp. 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. Wound healing in higher plants II. Proceedings of the. A colour with red set to 255 will appear bright red, or fully saturated with colour. Crow, B. These data demonstrate that R. stylosa prop roots are able to defend against teredinid larval settlement when exposed to superficial damage, and prop roots are able to produce an over compensatory level of tissue regrowth with moderate damage increasing root fitness. The objective of this study is to investigate the spatial structure of Rhizophora stylosa prop roots in different community types in terms of 3 parameters of the uniform angle index, neighborhood comparison and nearest neighbor by setting the location of R. stylosa prop roots. 2,932 = 79.8, P  ≤ 0.001), and time (Fig. The adult root system. The percentage of red pixel intensity used as a proxy for tannin production from each damaged root (mean ± SE, n = 81). Google Scholar. Biotropica 3: 63–77. Uses of R. stylosa in traditional medicine have not been reported. Oecologia 118: 316–323. Wound periderm development in red mangrove, Rhizophora mangle L. International Journal of Plant Science 157: 63–70. Suh, S. S., J. Hwang, M. Park, H. S. Park & T. K. Lee, 2014. 2017) Climate change mitigation • Root biomass is a major The sites were chosen because damaged roots were frequent and teredinid activity was also common. Rhizophora "rhizo" meaning root and "phora" meaning bear or carry in reference to the numerous prop roots growing from the trunk and branches of the mangrove. The growth of the new vascular tissues may provide some structural rigidity for the new cortex. Rhizophora stylosa prop roots in six plots measuring 10 m × 10 m were accessed for damage to determine the level required for teredinid recruitment. Differences of numbers of teredinid tunnels in sections exposed to the surgical treatments among the three mangrove localities were examined using PERMANOVA in Bray-Curtis matrices and post hoc pairwise tests. Australian Journal of Ecology 16: 433–443. 2C). & S. S. Bell, 2002. Rhizophora stylosa develops little brown fruits which are about 3-5cm small. Rhizophora stylosa grows naturally in Japan, China, Taiwan, Cambodia, Vietnam, Malesia and Australia (New South Wales and Queensland). Brooks, R. A. As a result, mangrove soil tends to have a low nitrogen content (Alongi et al., 1992; Feller et al., 2003a, b; Lovelock et al., 2004; Reef et al., 2010). The damaged area produces an increase of parenchyma cells, and the new outgrowth of non-radial vascular tissue may offer structural rigidity for the increased mass of cortex tissues. 4). Tomascik, T., Mah, A. J., Nontji, A., & Moosa, M. K., 1997. For each of the levels of damage, eight roots were used. stylosa (Griff.) Many roots exposed to moderate and severe levels of damage had necrotic cell damage. Rhizophora stylosa was an arbor with well-developed prop roots,strong power of wind-resistance and wave absorption.The characteristics of support structure of prop roots of R.stylosa were discussed in this paper.The results showed that in the pure forests of R.stylosa,mixed forests of R.stylosa and Avicennia marina,mixed forests of R.stylosa and Bruguiera gymnorhiza,its prop roots … Aerial roots growing from the tree´s limbs also help the plant breathe. Yet, when the loss of tissues penetrates the tannin-free vascular cylinder normally protected by the cortex layer, teredinid larvae are able to settle and tunnel into the prop root. Isolation of Salt Stress Tolerance Genes from Roots of Mangrove Plant, Rhizophora stylosa Griff., Using PCR-Based Suppression Subtractive Hybridization Shedding prevents possible risk of decay extending into the healthy tissues (Mattheck & Breloer, 1994). Herbivorous attack from sphaeromids and teredinids is different within mangroves. The greatest amount of tissue regrowth was found in the moderately damaged roots (PERMANOVA, F The levels of damage upon the prop roots were replicated experimentally on non-damaged prop roots to determine which of the prop-root surfaces are colonisable by teredinids. Mangrove wood herbivores range from insects such as beetles (Perry, 1988; Feller & Mathis, 1997) to wood-boring aquatic molluscs (Teredinidae) (Robertson & Daniel, 1989). Hendy, I.W., Cragg, S.M. A colour with red set to 0 specifies the absence of colour and emits no red light. face roots (Excoecaria agallocha), prop roots (Rhizophora apiculata ), stilt roots ( R . Bakau pasir (Rhizophora stylosa) Ng, Peter K. L. & N. Sivasothi, 1999. This means that wood borers may benefit mangrove ecosystems by breaking down dead wood, even though they do cause damage to some living tissues (Barkati & Tirmizi, 1991). Rhizophora mucronata and R. stylosa are sibling species (i.e., possibly R. stylosa =R. This study benefited from discussions with Mike J Swift, and an unpublished study of MJS and SMC. The line within the box marks the median. Feller, I. C., 2002. To add, many undamaged roots showed signs of full wound healing and recovery. Compartmentalization of decay in trees. This study highlights the resilience and ability of mangroves to heal damaged roots and defend against woodborers. Revista Brasileira de Ciência Avícola 14: 57–62. No roots died when exposed to this level of damage (Fig. Rhizophora stylosa. Google Scholar. Oikos 97: 167–176. This study aimed to determine why Rhizophora stylosa Griff trees were not toppled by wood boring teredinids by assessing the level of damage required to expose roots to teredinid colonisation. Buy Rhizophora stylosa Stilted Mangrove, select amount and size and put your mangrove to the shopping basket. Within each plot, each treatment was conducted on three individual roots totalling nine roots per plot and 27 roots per site. Superficially damaged roots did not succumb to teredinid attack. Applied Science, Barkin, UK, pp. Gill, A. M. & P. B. Tomlinson, 1971. The lenticels are air-filled spaces that connect with underground root structures. Growth and differentiation of aerial roots. Induced plant responses to herbivory. The role of herbivory by wood-boring insects in mangrove ecosystems in Belize. Ecology 69: 1064–1075. Marine Ecology Progress Series 516: 177–185. Rhizophora stylosa prop roots even when damaged prevent wood-boring teredinids from toppling the trees. Zero percent, 15 and 8% of the roots exposed to superficial, moderate and severe levels of damage (respectively) died. Within all the sites, a total of 81 roots were used. Bloch, R., 1952. The most prevalent level of damage was severe damage, which is a measure of un-healable damage (PERMANOVA, F The vascular cylinder is tannin-free (Gill & Tomlinson, 1971), making teredinid settlement possible (Turner, 1976). 409–416. Damage to prop roots may also occur from physical actions such as abrasion (Gill & Tomlinson, 1977), or by falling branches from the canopy. PubMed  Outliers are marked as asterisks. Filho, C. S., C. H. Tagliaro & C. R. Beasley, 2008. No differences were found with the number of teredinid tunnels in sections among the three mangrove localities (PERMANOVA, F Ecological role and services of tropical mangrove ecosystems: a reassessment. Plant Biology 10: 252–259. However, significant differences were found with the number of teredinid tunnels in sections among different surgical treatments (PERMANOVA, F Part of Springer Nature. Nowadays Rhizophora stylosa grows between latitude 20 north and 25 south from the equator. Rhizophora community is represented by three species, namely, R. apiculata, R. mucronata and R. stylosa, and two hybrids. Relationship between color and tannin content in sorghum grain: application of image analysis and artificial neural network. Root tissue loss was also great, with −31 ± 13% loss of the original circumference (Fig. Annual Review of Ecology and Systematics 20: 331–348. Teredinids are the major detritivores in mangrove forests that have high levels of dead wood, and they mechanically break down the wood (Robertson, 1990; Kohlmeyer et al., 1995). Losses of root tissue (percentage of cm2) were greatest in sections cut from severely damaged roots. 4, PERMANOVA pairwise, P ≤ 0.05 and ≤0.001, respectively). Induced responses to herbivory in wild radish: effects on several herbivore and plant fitness. The wood of Rhizophora stylosa has very high density and therefore very heavy wood which makes it very attractive as timber for boats, houses, fences and also fire wood. Tissue regeneration reduces the risk from potential infection (Wier et al., 1996). Dense soils increased total root biomass and primary root diameter, while the primary root length decreased. The benefits of induced defences against herbivores. Tannins in mangrove tree roots and their role in the root environment. The experiments took place in three intertidal R. stylosa-dominated mangrove forests in the Tukang Besi archipelago, Wakatobi Marine Park, East Sulawesi, Indonesia (see Cragg & Hendy, 2010 for site details). The Stilted Mangrove was als spread by human for coastal protection and aquaculture. Examples are Rhizophora x lamarckii, a cross between R. stylosa and R. apiculata (Chan, 1996; Ragavan et al., 2011; Ng & Chan, 2012b), and Rhizophora x mohanii, a cross between R. stylosa and R. mucronata (Ragavan et al., 2015). PubMed  Coupled with prolonged immersion, with greater root damage, the level of teredinid tunnelling increased. & Stout, M. J. Rhizophora stylosa is commonly known as the Red Mangrove. volume 803, pages333–344(2017)Cite this article. The influence of mangrove-derived tannins on intertidal meiobenthos in tropical estuaries. Marine Ecology 3: 13–19. A The removal of the periderm layer (superficial damage), B removal of the cortex tissue (moderate damage) exposing the vascular cylinder, and C removal of the vascular cylinder, (severe damage). The blackening of the wounds in the roots is due to the formation of a tannin-ferric iron complex, which counteracts potential toxicity to the roots via oxidation (Kimura & Wada, 1989). The bark is dark brown to black. Postharvest Biology and Technology 19: 73–83. Studies on the growth of red mangrove (Rhizophora mangle L.) 2. The greatest percentage of tissue regrowth was 21% greater in circumference compared to its original size. As renewed cortex grows over the damaged area, a new formation of the vascular cylinder is produced that appears to grow into the new cortex tissue. A, Strauss, S. Y. All statistical analyses were performed using MINITAB (MINITAB Inc, version 13.20) and PRIMER 6.1 (PrimerE Ltd: Plymouth Routines in Multivariate Ecological Research). It is not uncommon for trees to shed dead or damaged areas. There are few reports of teredinids attacking live Rhizophora prop roots (Roonwal, 1954, see Fig. Asterisks denote when intensity was significantly less in superficially damaged roots (GLM ANOVA with time period and severity of treatments as fixed factors, with Tukey’s post hoc pairwise comparisons). The wound response of a mangrove is increased by the level of injury. The degree of tolerance can be expressed as compensation to the plants ability to tolerate herbivore attack (Strauss & Agrawal, 1999), or stress. Tissue regrowth from roots exposed to moderate levels of damage was significantly more compared to the regrowth measured from superficially or severely damaged roots (Fig. Under these conditions Rhizophora stylosa is able to reach a height of up to 30 meters but usually stops height between 5 to 20 meters. Laboratory screening of tropical hardwoods for natural resistance to the marine borer Limnoria quadripunctata: the role of leachable and non-leachable factors. The number of teredinid tunnels within sections exposed to superficial, moderate and severely damaged roots. Percentage data were normalised using arcsine transformation. Doorn, W. G. V. & P. Cruz, 2000. 2,108 = 4.5, P ≤ 0.001). Shigo, A. L., 1985. Treatments on 24 of the experimentally damaged roots were cut transversely into sections after 12 months. The roots are then open to infection, and cell necrosis will ensue. The variation of tissue loss and regrowth between the different surgeries may be due to the degree of stress tolerance of each root. Six plots measuring 10 m × 10 m were designated to estimate the number of naturally occurring damaged prop roots in the mangrove forests., DOI:, Over 10 million scientific documents at your fingertips, Not logged in To combat attack, some plants release tannins (Bloch, 1952; Alongi, 1987). Debris brought in by the tides (Lee et al., 2014) and extreme storm events may also damage mangrove roots (McIvor et al., 2013; Jusoff, 2013), which are predicted to increase (Bhatt & Kathiresan, 2012). Wound-initiated tissue regrowth, also known as wound periderm, compartmentalises wounds and may prevent the spread of potential pathogens to healthy plant tissues (Wier et al., 1996). Herbivory on plants elicits a physiological response that can create an over compensation (excess) regrowth of tissue (Lennartsson et al., 1998). The experimental damage consisted of one of three surgical treatments with increasing severity upon individual roots. This may be due to the variation and frequency of teredinid attack coupled with the severity of damage imposed upon each root. Yet, few roots with severe damage demonstrated the same level of excessive tissue regrowth, and many roots lost tissue to necrosis and teredinid attack. The purpose of this research was to: (1) document the path of internal airflow, (2) describe and quantify tissues of internal airflow, and (3) determine if the amounts of aerenchyma in leaves, stems, and roots are relatively constant among plants of Rhizophora stylosa.Cork warts of leaves (average of 10.1 cork warts per mm 2 of leaf surface) are sites of air uptake. Indeed, there were a large number of naturally damaged roots in the mangrove forests in this study, and many of those roots showed signs of complete recovery. Red mangroves in North Queensland may grow to 20 m high, though trees of 4 to 5 m are more common elsewhere. In addition, exposure of the woody tissues may induce settlement behaviour, while the spongy cortex layer may not provide the necessary settlement cues. The cortex of the root took on a strong red colouration after being experimentally damaged. Rhizophora stylosa roots of seedling Evolutionary Ecology 14: 551–562. This work tested the prediction that after 1 year R. stylosa roots with: (1) superficial levels of damage (removal of the outer bark, the periderm) will not be attacked by teredinids and will make a full recovery; (2) moderate levels of damage (removal of the cortex) will elicit an over compensatory level of tissue regrowth; and (3) severe levels of damage (removal of the inner radial wall of the vascular cylinder) will die due to a high level of stress. A Guide to the Mangroves of Singapore I (Plant Diversity). Sixty-one percent of those roots had exceeded their initial circumference with an increase of 9 ± 2% before surgery. However, those roots with teredinid tunnels were either necrotic or dead in a similar way to that observed in the experimentally severely damaged roots. Therefore, tissue loss after one year within the severely damaged roots went beyond the original scaring. 1). When tissue patterns arise that differ from the norm, or when a lack of pattern is encountered, the cellular regrowth can be classified as atypical (Bloch, 1952). Responses of 81 R. stylosa roots to three levels of experimental damage were investigated: superficial, moderate and severe. The energy required to produce tannins can be costly (Agrawal et al., 1999; Karban et al., 1997). Robertson, A. I. 168 pp. Typically Rhizophora stylosa develops a one columnar stem which often soon starts to develop a few main branches. 2,41 = 0.6, P ≥ 0.05). The first surgical treatment (superficial) consisted of removal of the outermost layer of the bark, the periderm (Fig. Sections were inspected for evidence of necrotic cellular damage and then measured to calculate the area of cellular regrowth or loss using ImageTool Version 3.00 (The University of Texas Health Science Centre at San Antonio). Stilt roots arises from the trunk or branches of the mangrove and grows toward the soil where the stilt root will develop an underground root system. 2,17 = 93, P ≤ 0.001). Tissue regrowth was minimal with an increase of 3% ± 0.3% of the original circumference before surgery. Conversely, herbivory may be beneficial (Paige & Whitham, 1987; Paige, 1992). This study demonstrates the remarkable ability of mangroves to combat herbivorous activity by the bark barrier, by the production of tannins in damaged cortex tissue and/or by an over compensatory regrowth of tissue. An increase of suberin helps with parasite resistance and provides immunity to further infection in the damaged tissues (Karban & Myers, 1989; Franke & Schreiber, 2007). Sites were chosen because damaged roots and teredinid activity were frequent. Primary herbivory by wood-boring insects along an architectural gradient of Rhizophora mangle. The Ecology of the Indonesian Seas Part Two. I'm adding this plant onto the blog due to its ability to be able to survive in saline environments. The effect of moisture content and drying temperature on the colour of two poplars and robinia wood. A Chi square test was used to examine differences between sections and the number of teredinid tunnels after 12 months. However, it is beneficial for the plant to produce these chemical compounds only when they are required as herbivore attack is random and variable (Hol et al., 2004). Rimmer, M. A., S. L. Battaglene & P. L. Dostine, 1983. To estimate tannin content, we focussed on red light intensity in each image. Mattheck, K. & H. Breloer, 1994. Ribi, G., 1982. Zoological Society 7: 91–103. Each treatment was conducted no more than 20 cm from the substratum. Biotropica 29: 440–451. Nova Science Publishers, Hauppauge, NY. However, it is thought that sphaeromids are unable to burrow into older developed roots that reach the substratum due to the development of woody tissue in the older roots (Perry, 1988). It takes some experience to be able to distinguish between the Rhizophora propagules and to identify propagules of Rhizophora stylosa undoubtedly. The blossoms of Rhizophora stylosa are small and inconspicuous as all blossoms of all Rhizophora species., Decomposition and the annual flux of detritus from fallen timber in tropical mangrove forests. A Fisher’s exact test with a post hoc pairwise test was used to test for differences between sections with or without lateral outgrowth of new vascular tissue after 12 months. Rhizophora stylosa prop roots to heal damagedtissues and defend against herbivorous attack from teredinids in three mangrove forests. Damage and alteration of mangroves inhabited by a marine wood-borer. Lateral view and transverse sections of three treatments of roots characterised by different levels of damage. Novel in-growth containers were used to assess the effect of soil bulk density (BD: 0.4, 0.8 and 1.2 g cm-3) on morphological, anatomical and chemical traits of the below-ground fraction of aerial roots of the mangrove Rhizophora stylosa. This study examines the ability of live Rhizophora stylosa prop roots to heal damaged tissues and defend against herbivorous attack from teredinids in three mangrove forests. Mangrove plants may likely use the tolerance strategy against damage or stress (Brooks & Bell, 2002), as many other angiosperms have evolved this response to combat herbivore attack (Strauss & Agrawal, 1999). Sphaeromids may benefit the plant by inducing increases of lateral out-branching of new aerial roots (Ribi, 1982). Not all roots were tolerant to the removal of tissues. Overcompensation in response to mammalian herbivory: the advantage of being eaten. Decomposition of mangrove wood by marine fungi and teredinids in Belize. Induction of overcompensation in the field gentian, Gentianella campestris. Barkati, S. & N. M. Tirmizi, 1991. Sections cut from superficially damaged roots had almost no measurable changes. The boundaries of the boxes indicate the 1st and 3rd quartiles. Wier et al. The halophytic Rhizophora stylosa is useful for the study of the molecular mechanisms behind salinity tolerance in mangrove trees. PubMed  Turner, R. D., 1976. In Costa Rica, the destructive effect of the sphaeromid, Sphaeroma peruvianum Richardson, 1910 on live mangrove root tissues can reduce the growth rates of Rhizophora mangle L. aerial prop roots by 50% (Perry, 1988). Guest editors: K. W. Krauss, I. C. Feller, D. A. Friess & R. R. Lewis III / Causes and Consequences of Mangrove Ecosystem Responses to an Ever-Changing Climate. Plant Life of the Great Barrier Reef... (1985) p 230-32 Parts Shown: Flower, Habit Photo. Rhizophora stylosa. This study examines the ability of live Rhizophora stylosa prop roots to heal damaged tissues and defend against herbivorous attack from teredinids in three mangrove forests. 2,41 = 12.6, P ≤ 0.001). The extent of infestation by teredinids was quantified. Twelve percent of the severely damaged roots were attacked by teredinids, and sections cut from severely damaged roots had between three and 10 teredinid tunnels. Alongi, D. M., 1987. Thus, particular mangrove trees may express resistance only when it is needed to reduce energetic cost into producing the tannins. The first one is the adaption to low oxygen. Mangrove Forests of the Wakatobi National Park. mucronata var. Sites were chosen because damaged roots and teredinid activity were frequent. The third treatment (severe) consisted of additionally removing the inner radial wall of the vascular cylinder, exposing the inner pith (Fig. Some trees are protected from herbivorous attack by their chemical and structural anti-herbivore defence mechanisms (Turner, 1976; Brooks & Bell, 2002). Australian Journal of Marine and Freshwater Research 34: 355–357. Rhizophora stylosa grows along the coast and sometimes directly in the ocean near the coast, often in areas where rivers flow into the ocean, soils are very nutritious and where humidity is between 60 to 80 percent and the air temperature is between 25 to 30°C. Holzforschung 62: 99–111. Roots exposed to moderate and severe levels of damage had lateral out-growths of new vascular tissue extending from the vascular cylinder. Plants also express resistance to herbivory via strategies labelled as tolerance mechanisms (Paige, 1999; Brooks & Bell, 2002), defined as tissue repair and regrowth after exposure from stress. (1996) found proliferations of vascular tissue at the cut ends of the vascular cylinder. The vascular cylinder is the water-carrying component (Gill & Tomlinson, 1971), and the inner limit of the cortex is tannin-free (Gill & Tomlinson, 1971). Svavarsson, J., M. K. Osore & E. Olafsson, 2002. 24–26. Phenol content, antioxidant and tyrosinase inhibitory activity of mangrove plants in Micronesia. Google Scholar. Roonwal, M. L., 1954. Feller, I. C. & W. N. Mathis, 1997. The absence of tannins means there was no chemical defence against teredinid larval settlement. The propagule starts to grow its sprout in the fruit while it is still on the mother tree. This work was carried out in three sites in East Sulawesi, Indonesia. Franke, R. & L. Schreiber, 2007. But, when losses of tissues expose the vascular cylinder, teredinid larvae will settle and tunnel into the root. 1999. gymnorrhiza+A. CAS  A. van Veen & E. van der Meijden, 2004. Estimation of aboveground biomass in a Rhizophora stylosa. A., 1999. Seasonal abundance of the shipworm Neoteredo reynei (Bivalvia, Teredinidae) in mangrove driftwood from a northern Brazilian beach. In addition, a constituent component of the cortex tissue is suberin. In Sharpley, J. M. & A. M. Kaplan (eds), Proceedings of the 3rd International Biodegradation Symposium. Almost 80% of moderately damaged roots had made a full recovery. Mangle is Spanish and means mangrove, ... Rhizophora stylosa was known as Red Mangrove for a long time. Evidence for a wounding-induced xylem occlusion in stems of cut chrysanthemum flowers. - 2A). Title Flora Vitiensis Nova Publication Author Smith. Four types of community (Rhizophora stylosa pure forest, R. stylosa+Bruguiera gymnorrhiza mixed forest, R. stylosa+Avicennia marina mixed forest, and R. stylosa+B. 1854. pronounced: ree-ZOH-for-uh sty-LOH-suh (Rhizophoraceae — the mangrove family)common name: red mangrove. Description. Nitrogen is one of the most essential elements regulating plant growth, and plants have developed source and sink mechanisms for its transport and use (Tegeder and Masclaux-Daubresse, 2018). In the study site, many roots were damaged either by use of tools during firewood collection or by physical abrasion. Compartmentalising sets boundaries that resist the spread of the invading microorganisms (Shigo, 1985). Evolution 53: 1093–1104. However, necrotic regions tend to become isolated from the rest of the tree, so that the stability of the tree itself is not significantly compromised. Cribb, A.B. These do not take root even after … The marine borer, Bactronophorus thoracities (Gould) [Mollusca, Eulamellibranchiata, Teredinidae], as a pest of living trees in the mangrove forests of the Sunderbans, Bengal, India. Fieldwork for IWH was supported by Operation Wallacea and field collections of prop-root measurements were helped by Kungdan. A Fisher’s exact test was used to test for differences between live and dead roots. Rhizophora stylosa roots are able to defend against teredinid larval settlement by production of tannins in damaged cortex tissue and by an over compensatory regrowth. Plants respond to damage and infection by compartmentalising the affected area, reducing the spread of infection to healthy tissues (Bloch, 1952). As most mangrove propagules the propagules of Rhizophora stylosa are viviparous and already develop a sprout on the mother tree. This adaptation may be induced by over compensation of the tolerance mechanism (Haukioja & Koricheva, 2000; Brooks & Bell, 2002). The roots are then open to infection, and cell necrosis. The Nature Conservancy, University of Cambridge, and Wetlands International, Bali, Indonesia, September, pp. All root circumferences and scar depths were measured before and during a 12-month period using a tape measure and callipers. Novel in-growth containers were used to assess the effect of soil bulk density (BD: 0.4, 0.8 and 1.2 g cm −3) on morphological, anatomical and chemical traits of the below-ground fraction of aerial roots of the mangrove Rhizophora stylosa.

rhizophora stylosa roots

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