Multiple impact pathways of the 2015-2016 El Niño in coastal Kenya.

The 2015-2016 El Niño had large impacts globally. The effects were not as great as anticipated in Kenya, however, leading some commentators to call it a 'non-event'. Our study uses a novel combination of participatory Climate Vulnerability and Capacity Analysis tools, and new and existing social and biophysical data, to analyse vulnerability to, and the multidimensional impacts of, the 2015-2016 El Niño episode in southern coastal Kenya. Using a social-ecological systems lens and a unique dataset, our study reveals impacts overlooked by conventional analysis. We show how El Niño stressors interact with and amplify existing vulnerabilities to differentially impact local ecosystems and people. The policy significance of this finding is that the development of specific national capacities to deal with El Niño events is insufficient; it will be necessary to also address local vulnerabilities to everyday and recurrent stressors and shocks to build resilience to the effects of El Niño and other extremes in climate and weather.

Channel network structure determines genetic connectivity of landward-seaward Avicennia marina populations in a tropical bay.

Mangrove ecosystems along the East African coast are often characterized by a disjunct zonation pattern of seaward and landward Avicennia marina trees. This disjunct zonation may be maintained through different positions in the tidal frame, yielding different dispersal settings. The spatial configuration of the landscape and coastal processes such as tides and waves is expected to largely influence the extent of propagule transport and subsequent regeneration. We hypothesized that landward sites would keep a stronger genetic structure over short distances in comparison with enhanced gene flow among regularly flooded seaward fringes. We tested this hypothesis from densely vegetated A. marina transects of a well-documented mangrove system (Gazi Bay, Kenya) and estimated local gene flow and kinship-based fine-scale genetic structure. Ten polymorphic microsatellite markers in 457 A. marina trees revealed no overall significant difference in levels of allele or gene diversities between sites that differ in hydrological proximity. Genetic structure and connectivity of A. marina populations however indicated an overall effect of geographic distance and revealed a pronounced distinction between channels and topographic setting. Migration models allowed to infer gene flow directionality among channels, and indicated a bidirectional steppingstone between seaward and nearest located landward stands. Admixed gene pools without any fine-scale structure were found within the wider and more exposed Kidogoweni channel, suggesting open systems. Elevated kinship values and structure over 5 to 20 m distance were only detected in two distant landward and seaward transects near the mouth of the Mkurumuji River, indicating local retention and establishment. Overall, our findings show that patterns of A. marina connectivity are explained by hydrological proximity, channel network structure, and hydrokinetic energy, rather than just their positioning as disjunct landward or seaward zones.

Carbon storage in the seagrass meadows of Gazi Bay, Kenya.

Vegetated marine habitats are globally important carbon sinks, making a significant contribution towards mitigating climate change, and they provide a wide range of other ecosystem services. However, large gaps in knowledge remain, particularly for seagrass meadows in Africa. The present study estimated biomass and sediment organic carbon (Corg) stocks of four dominant seagrass species in Gazi Bay, Kenya. It compared sediment Corg between seagrass areas in vegetated and un-vegetated 'controls', using the naturally patchy occurence of seagrass at this site to test the impacts of seagrass growth on sediment Corg. It also explored relationships between the sediment and above-ground Corg, as well as between the total biomass and above-ground parameters. Sediment Corg was significantly different between species, range: 160.7-233.8 Mg C ha-1 (compared to the global range of 115.3 to 829.2 Mg C ha-1). Vegetated areas in all species had significantly higher sediment Corg compared with un-vegetated controls; the presence of seagrass increased Corg by 4-6 times. Biomass carbon differed significantly between species with means ranging between 4.8-7.1 Mg C ha-1 compared to the global range of 2.5-7.3 Mg C ha-1. To our knowledge, these are among the first results on seagrass sediment Corg to be reported from African seagrass beds; and contribute towards our understanding of the role of seagrass in global carbon dynamics.

Evaluating, predicting and mapping belowground carbon stores in Kenyan mangroves.

Despite covering only approximately 138 000 km2 , mangroves are globally important carbon sinks with carbon density values three to four times that of terrestrial forests. A key challenge in evaluating the carbon benefits from mangrove forest conservation is the lack of rigorous spatially resolved estimates of mangrove sediment carbon stocks; most mangrove carbon is stored belowground. Previous work has focused on detailed estimations of carbon stores over relatively small areas, which has obvious limitations in terms of generality and scope of application. Most studies have focused only on quantifying the top 1 m of belowground carbon (BGC). Carbon stored at depths beyond 1 m, and the effects of mangrove species, location and environmental context on these stores, are poorly studied. This study investigated these variables at two sites (Gazi and Vanga in the south of Kenya) and used the data to produce a country-specific BGC predictive model for Kenya and map BGC store estimates throughout Kenya at spatial scales relevant for climate change research, forest management and REDD+ (reduced emissions from deforestation and degradation). The results revealed that mangrove species was the most reliable predictor of BGC; Rhizophora muronata had the highest mean BGC with 1485.5 t C ha-1 . Applying the species-based predictive model to a base map of species distribution in Kenya for the year 2010 with a 2.5 m2 resolution produced an estimate of 69.41 Mt C [±9.15 95% confidence interval (C.I.)] for BGC in Kenyan mangroves. When applied to a 1992 mangrove distribution map, the BGC estimate was 75.65 Mt C (±12.21 95% C.I.), an 8.3% loss in BGC stores between 1992 and 2010 in Kenya. The country-level mangrove map provides a valuable tool for assessing carbon stocks and visualizing the distribution of BGC. Estimates at the 2.5 m2 resolution provide sufficient details for highlighting and prioritizing areas for mangrove conservation and restoration.

Applying Climate Compatible Development and economic valuation to coastal management: A case study of Kenya's mangrove forests.

Mangrove forests are under global pressure. Habitat destruction and degradation persist despite longstanding recognition of the important ecological functions of mangroves. Hence new approaches are needed to help stakeholders and policy-makers achieve sound management that is informed by the best science. Here we explore how the new policy concept of Climate Compatible Development (CCD) can be applied to achieve better outcomes. We use economic valuation approaches to combine socio-economic data, projections of forest cover based on quantitative risk mapping and storyline scenario building exercises to articulate the economic consequences of plausible alternative future scenarios for the mangrove forests of the South Kenya coast, as a case study of relevance to many other areas. Using data from 645 household surveys, 10 focus groups and 74 interviews conducted across four mangrove sites, and combining these with information on fish catches taken at three landing sites, a mangrove carbon trading project and published data allowed us to make a thorough (although still partial) economic valuation of the forests. This gave a current value of the South Coast mangroves of USD 6.5 million, or USD 1166 ha(-1), with 59% of this value on average derived from regulating services. Quantitative risk mapping, projecting recent trends over the next twenty years, suggests a 43% loss of forest cover over that time with 100% loss at the most vulnerable sites. Much of the forest lost between 1992 and 2012 has not been replaced by high value alternative land uses hence restoration of these areas is feasible and may not involve large opportunity costs. We invited thirty eight stakeholders to develop plausible storyline scenarios reflecting Business as Usual (BAU) and CCD - which emphasises sustainable forest conservation and management - in twenty years time, drawing on local and regional expert knowledge of relevant policy, social trends and cultures. Combining these scenarios with the quantitative projections and economic baseline allowed the modelling of likely value added and costs avoided under the CCD scenario. This suggests a net present value of more than US$20 million of adoption of CCD rather than BAU. This work adds to the economic evidence for mangrove conservation and helps to underline the importance of new real and emerging markets, such as for REDD + projects, in making this case for carbon-rich coastal habitats. It demonstrates a policy tool - CCD - that can be used to engage stakeholders and help to co-ordinate policy across different sectors towards mangrove conservation.

Effects of experimental sedimentation on the phenological dynamics and leaf traits of replanted mangroves at Gazi bay, Kenya.

Sedimentation results in the creation of new mudflats for mangroves to colonize among other benefits. However, large sediment input in mangrove areas may be detrimental to these forests. The dynamics of phenological events of three mangrove tree species (Avicennia marina, Ceriops tagal, and Rhizophora mucronata) were evaluated under experimental sediment burial simulating sedimentation levels of 15, 30, and 45 cm.While there was generally no shift in timing of phenological events with sedimentation, the three mangrove tree species each responded differently to the treatments.Partially buried A. marina trees produced more leaves than the controls during the wet season and less during the dry season. Ceriops tagal on the other hand had higher leaf loss and low replacement rates in the partially buried trees during the first 6 months of the experiment but adapted with time, resulting in either equal or higher leaf emergence rates than the controls.Rhizophora mucronata maintained leaf emergence and loss patterns as the unaffected controls but had a higher fecundity and productivity in the 15-cm sedimentation level.The results suggest that under incidences of large sedimentation events (which could be witnessed as a result of climate change impacts coupled with anthropogenic disturbances), mangrove trees may capitalize on "advantages" associated with terrestrial sediment brought into the biotope, thus maintaining the pattern of phenological events.

Rapid losses of surface elevation following tree girdling and cutting in tropical mangroves.

The importance of mangrove forests in carbon sequestration and coastal protection has been widely acknowledged. Large-scale damage of these forests, caused by hurricanes or clear felling, can enhance vulnerability to erosion, subsidence and rapid carbon losses. However, it is unclear how small-scale logging might impact on mangrove functions and services. We experimentally investigated the impact of small-scale tree removal on surface elevation and carbon dynamics in a mangrove forest at Gazi bay, Kenya. The trees in five plots of a Rhizophora mucronata (Lam.) forest were first girdled and then cut. Another set of five plots at the same site served as controls. Treatment induced significant, rapid subsidence (-32.1±8.4 mm yr-1 compared with surface elevation changes of +4.2±1.4 mm yr-1 in controls). Subsidence in treated plots was likely due to collapse and decomposition of dying roots and sediment compaction as evidenced from increased sediment bulk density. Sediment effluxes of CO2 and CH4 increased significantly, especially their heterotrophic component, suggesting enhanced organic matter decomposition. Estimates of total excess fluxes from treated compared with control plots were 25.3±7.4 tCO2 ha-1 yr-1 (using surface carbon efflux) and 35.6±76.9 tCO2 ha-1 yr-1 (using surface elevation losses and sediment properties). Whilst such losses might not be permanent (provided cut areas recover), observed rapid subsidence and enhanced decomposition of soil sediment organic matter caused by small-scale harvesting offers important lessons for mangrove management. In particular mangrove managers need to carefully consider the trade-offs between extracting mangrove wood and losing other mangrove services, particularly shoreline stabilization, coastal protection and carbon storage.

Turning the tide: how blue carbon and payments for ecosystem services (PES) might help save mangrove forests.

In this review paper, we aim to describe the potential for, and the key challenges to, applying PES projects to mangroves. By adopting a "carbocentric approach," we show that mangrove forests are strong candidates for PES projects. They are particularly well suited to the generation of carbon credits because of their unrivaled potential as carbon sinks, their resistance and resilience to natural hazards, and their extensive provision of Ecosystem Services other than carbon sequestration, primarily nursery areas for fish, water purification and coastal protection, to the benefit of local communities as well as to the global population. The voluntary carbon market provides opportunities for the development of appropriate protocols and good practice case studies for mangroves at a small scale, and these may influence larger compliance schemes in the future. Mangrove habitats are mostly located in developing countries on communally or state-owned land. This means that issues of national and local governance, land ownership and management, and environmental justice are the main challenges that require careful planning at the early stages of mangrove PES projects to ensure successful outcomes and equitable benefit sharing within local communities.

How to catch the patch? A dendrometer study of the radial increment through successive cambia in the mangrove Avicennia.

BACKGROUND AND AIMS: Successive vascular cambia are involved in the secondary growth of at least 200 woody species from >30 plant families. In the mangrove Avicennia these successive cambia are organized in patches, creating stems with non-concentric xylem tissue surrounded by internal phloem tissue. Little is known about radial growth and tree stem dynamics in trees with this type of anatomy. This study aims to (1) clarify the process of secondary growth of Avicennia trees by studying its patchiness; and (2) study the radial increment of Avicennia stems, both temporary and permanent, in relation to local climatic and environmental conditions. A test is made of the hypothesis that patchy radial growth and stem dynamics enable Avicennia trees to better survive conditions of extreme physiological drought. Methods Stem variations were monitored by automatic point dendrometers at four different positions around and along the stem of two Avicennia marina trees in the mangrove forest of Gazi Bay (Kenya) during 1 year. KEY RESULTS: Patchiness was found in the radial growth and shrinkage and swelling patterns of Avicennia stems. It was, however, potentially rather than systematically present, i.e. stems reacted either concentrically or patchily to environment triggers, and it was fresh water availability and not tidal inundation that affected radial increment. CONCLUSIONS: It is concluded that the ability to develop successive cambia in a patchy way enables Avicennia trees to adapt to changes in the prevailing environmental conditions, enhancing its survival in the highly dynamic mangrove environment. Limited water could be used in a more directive way, investing all the attainable resources in only some locations of the tree stem so that at least at these locations there is enough water to, for example, overcome vessel embolisms or create new cells. As these locations change with time, the overall functioning of the tree can be maintained.

An Evaluation of Plotless Sampling Using Vegetation Simulations and Field Data from a Mangrove Forest.

In vegetation science and forest management, tree density is often used as a variable. To determine the value of this variable, reliable field methods are necessary. When vegetation is sparse or not easily accessible, the use of sample plots is not feasible in the field. Therefore, plotless methods, like the Point Centred Quarter Method, are often used as an alternative. In this study we investigate the accuracy of different plotless sampling methods. To this end, tree densities of a mangrove forest were determined and compared with estimates provided by several plotless methods. None of these methods proved accurate across all field sites with mean underestimations up to 97% and mean overestimations up to 53% in the field. Applying the methods to different vegetation patterns shows that when random spatial distributions were used the true density was included within the 95% confidence limits of all the plotless methods tested. It was also found that, besides aggregation and regularity, density trends often found in mangroves contribute to the unreliability. This outcome raises questions about the use of plotless sampling in forest monitoring and management, as well as for estimates of density-based carbon sequestration. We give recommendations to minimize errors in vegetation surveys and recommendations for further in-depth research.

Species mixing boosts root yield in mangrove trees.

Enhanced species richness can stimulate the productivity of plant communities; however, its effect on the belowground production of forests has scarcely been tested, despite the role of tree roots in carbon storage and ecosystem processes. Therefore, we tested for the effects of tree species richness on mangrove root biomass: thirty-two 6 m by 6 m plots were planted with zero (control), one, two or three species treatments of six-month-old Avicennia marina (A), Bruguiera gymnorrhiza (B) and Ceriops tagal (C). A monoculture of each species and the four possible combinations of the three species were used, with four replicate plots per treatment. Above- and belowground biomass was measured after three and four years' growth. In both years, the all-species mix (ABC) had significant overyielding of roots, suggesting complementarity mediated by differences in rhizosphere use amongst species. In year four, there was higher belowground than aboveground biomass in all but one treatment. Belowground biomass was strongly influenced by the presence of the most vigorously growing species, A. marina. These results demonstrate the potential for complementarity between fast- and slow-growing species to enhance belowground growth in mangrove forests, with implications for forest productivity and the potential for belowground carbon sequestration.

Intra- and interspecific facilitation in mangroves may increase resilience to climate change threats.

Mangroves are intertidal ecosystems that are particularly vulnerable to climate change. At the low tidal limits of their range, they face swamping by rising sea levels; at the high tidal limits, they face increasing stress from desiccation and high salinity. Facilitation theory may help guide mangrove management and restoration in the face of these threats by suggesting how and when positive intra- and interspecific effects may occur: such effects are predicted in stressed environments such as the intertidal, but have yet to be shown among mangroves. Here, we report the results of a series of experiments at low and high tidal sites examining the effects of mangrove density and species mix on seedling survival and recruitment, and on the ability of mangroves to trap sediment and cause surface elevation change. Increasing density significantly increased the survival of seedlings of two different species at both high and low tidal sites, and enhanced sediment accretion and elevation at the low tidal site. Including Avicennia marina in species mixes enhanced total biomass at a degraded high tidal site. Increasing biomass led to changed microenvironments that allowed the recruitment and survival of different mangrove species, particularly Ceriops tagal.

A patchy growth via successive and simultaneous cambia: key to success of the most widespread mangrove species Avicennia marina?

BACKGROUND AND AIMS: Secondary growth via successive cambia has been intriguing researchers for decades. Insight into the mechanism of growth layer formation is, however, limited to the cellular level. The present study aims to clarify secondary growth via successive cambia in the mangrove species Avicennia marina on a macroscopic level, addressing the formation of the growth layer network as a whole. In addition, previously suggested effects of salinity on growth layer formation were reconsidered. METHODS: A 1-year cambial marking experiment was performed on 80 trees from eight sites in two mangrove forests in Kenya. Environmental (soil water salinity and nutrients, soil texture, inundation frequency) and tree characteristics (diameter, height, leaf area index) were recorded for each site. Both groups of variables were analysed in relation to annual number of growth layers, annual radial increment and average growth layer width of stem discs. KEY RESULTS: Between trees of the same site, the number of growth layers formed during the 1-year study period varied from only part of a growth layer up to four growth layers, and was highly correlated to the corresponding radial increment (0-5 mm year(-1)), even along the different sides of asymmetric stem discs. The radial increment was unrelated to salinity, but the growth layer width decreased with increasing salinity and decreasing tree height. CONCLUSIONS: A patchy growth mechanism was proposed, with an optimal growth at distinct moments in time at different positions around the stem circumference. This strategy creates the opportunity to form several growth layers simultaneously, as observed in 14 % of the studied trees, which may optimize tree growth under favourable conditions. Strong evidence was provided for a mainly endogenous trigger controlling cambium differentiation, with an additional influence of current environmental conditions in a trade-off between hydraulic efficiency and mechanical stability.

Comparative anatomy of intervessel pits in two mangrove species growing along a natural salinity gradient in Gazi bay, Kenya.

BACKGROUND AND AIMS: According to the air-seeding hypothesis, embolism vulnerability in xylem elements is linked directly to bordered pit structure and functioning. To elucidate the adaptive potential of intervessel pits towards fluctuating environmental conditions, two mangrove species with a distinct ecological distribution growing along a natural salinity gradient were investigated. METHODS: Scanning and transmission electron microscopic observations were conducted to obtain qualitative and quantitative characteristics of alternate intervessel pits in A. marina and scalariform intervessel pits in Rhizophora mucronata. Wood samples from three to six trees were collected at seven and five sites for A. marina and R. mucronata, respectively, with considerable differences between sites in soil water salinity. KEY RESULTS: Vestured pits without visible pores in the pit membrane were observed in A. marina, the mangrove species with the widest geographical distribution on global as well as local scale. Their thick pit membranes (on average 370 nm) and minute pit apertures may contribute to reduced vulnerability to cavitation of this highly salt-tolerant species. The smaller ecological distribution of R. mucronata was in accordance with wide pit apertures and a slightly higher pitfield fraction (67 % vs. 60 % in A. marina). Nonetheless, its outer pit apertures were observed to be funnel-shaped shielding non-porous pit membranes. No trends in intervessel pit size were observed with increasing soil water salinity of the site. CONCLUSIONS: The contrasting ecological distribution of two mangrove species was reflected in the geometry and pit membrane characteristics of their intervessel pits. Within species, intervessel pit size seemed to be independent of spatial variations in environmental conditions and was only weakly correlated with vessel diameter. Further research on pit formation and function has to clarify the large variations in intervessel pit size within trees and even within single vessels.

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

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

Growth rings, growth ring formation and age determination in the mangrove Rhizophora mucronata.

BACKGROUND AND AIMS: The mangrove Rhizophora mucronata has previously been reported to lack annual growth rings, thus barring it from dendrochronological studies. In this study the reported absence of the growth rings was reconsidered and the periodic nature of light and dark brown layers visible on polished stem discs investigated. In addition, the formation of these layers in relation to prevailing environmental conditions, as well as their potential for age determination of the trees, was studied. METHODS: Trees of known age were collected and a 2.5-year cambial marking experiment was conducted to determine the periodic nature of the visible growth layers. KEY RESULTS: Annual indistinct growth rings were detected in R. mucronata and are defined by a low vessel density earlywood and a high vessel density latewood. The formation of these growth rings and their periodic nature was independent from site-specific environmental conditions in two forests along the Kenyan coast. However, the periodic nature of the rings was seriously affected by slow growth rates, allowing accurate age determination only in trees with radial growth rates above 0.5 mm year(-1). The onset of the formation of the low vessel density wood coincided with the onset of the long rainy season (April-May) and continues until the end of the short rainy season (November). The high vessel density wood is formed during the dry season (December-March). Age determination of the largest trees collected in the two studied forests revealed the relatively young age of these trees (+/-100 years). CONCLUSIONS: This study reports, for the first time, the presence of annual growth rings in the mangrove R. mucronata, which offers further potential for dendrochronological and silvicultural applications.

Regeneration status of mangrove forests in Mida Creek, Kenya: a compromised or secured future?

The structure and regeneration patterns of Mida Creek mangrove vegetation were studied along belt transects at 2 forest sites of Mida Creek (3 degrees 20'S, 40 degrees 00'E): Uyombo and Kirepwe. Based on the species importance values, the dominant mangrove tree species in Mida were Ceriops tagal (Perr.) C. B. Robinson and Rhizophora mucronata Lamk. Tree density varied from 1197 trees ha(-1) at Kirepwe to 1585 trees ha(-1) at Uyombo and mean tree height was higher at the former site compared to the latter. The size-class structure at both localities of Mida showed the presence of more small trees than large ones. Spatial distribution pattern of adults and juveniles varied greatly between sites and they showed a close to uniform pattern (Morisita's Index I0 << 1) for trees, but a tendency to random distribution (I0 = 1) for juveniles. The present paper shows that unmanaged but exploited mangroves do not necessarily disappear, but change qualitatively from locally preferred R. mucronata to the less preferred C. tagal. Whereas the effects of this change on the ecological function of the mangrove cannot be estimated yet, the economical function of the mangrove has evidently weakened.