Functional Traits of Terrestrial Plants in the Intertidal: A Review on Mangrove Trees.

AbstractTrue mangroves are vascular plants (Tracheophyta) that evolved into inhabiting the mid and upper intertidal zone of tropical and subtropical soft-sediment coasts around the world. While several dozens of species are known from the Indo-West Pacific region, the Atlantic-East Pacific region is home to only a mere dozen of true mangrove species, most of which are rare. Mangrove trees can form dense monospecific or multispecies stands that provide numerous ecosystem services. Despite their eminent socioecological and socioeconomic relevance and the plethora of studies on mangroves, many details of the ecology of mangrove ecosystems remain unknown; and our knowledge about general ecological principles in mangrove ecosystems is scarce. For instance, the functional trait concept has hardly been applied to mangroves. Here we provide an inventory of 28 quantitative and 8 qualitative functional traits of true mangrove species and stipulate some insight into how these traits may drive ecosystem structure and processes. The differentiation between true mangroves and mangrove associates, which can dwell inside as well as outside mangrove forests, is reflected by a number of leaf traits. Thus, true mangroves exhibit lower specific leaf area, lower leaf N content, and lower K∶Na ratio, and higher leaf succulence, higher Na and Cl content, and higher osmolality than mangrove associates. True mangrove species that form pure stands produce larger leaves and exhibit higher N content per leaf area, higher leaf K and Ca content, greater maximum plant height, longer propagules, and lower root porosity than more sporadic species. The species-specific expression of most traits does not reflect the species' position along intertidal gradients, suggesting that adaptation to tidal inundation does not explain these traits. Rather, many of the traits studied herein exhibit strong phylogenetic signals in true mangroves. Thus, wood density is high in most species of the Rhizophoraceae, irrespective of their habitat or maximum height. On the other hand, species of the genus Sonneratia exhibit low wood density and do not grow taller than 20 m. Some leaf traits of true mangroves are more like those of plants from drier environments, reflecting the perception that a saline environment creates physiological drought stress. Along the same line, most true mangrove species exhibit sclerophyllous leaf traits. The few major mangrove tree species of the Atlantic-East Pacific are as distinct from each other, with regard to some traits, as are the many mangrove species of the Indo-West Pacific. We hypothesize that this phenomenon explains the similarly high biomass of mangrove forests in both the species-rich Indo-West Pacific and the species-poor Atlantic-East Pacific.

Nutrient History Affects the Response and Resilience of the Tropical Seagrass Halophila stipulacea to Further Enrichment in Its Native Habitat.

Eutrophication is one of the main threats to seagrass meadows, but there is limited knowledge on the interactive effects of nutrients under a changing climate, particularly for tropical seagrass species. This study aimed to detect the onset of stress in the tropical seagrass, Halophila stipulacea, by investigating the effect of in situ nutrient addition during an unusually warm summer over a 6-month period. We measured a suite of different morphological and biochemical community metrics and individual plant traits from two different sites with contrasting levels of eutrophication history before and after in situ fertilization in the Gulf of Aqaba. Nutrient stress combined with summer temperatures that surpassed the threshold for optimal growth negatively affected seagrass plants from South Beach (SB), an oligotrophic marine protected area, while H. stipulacea populations from North Beach (NB), a eutrophic and anthropogenically impacted area, benefited from the additional nutrient input. Lower aboveground (AG) and belowground (BG) biomass, reduced Leaf Area Index (LAI), smaller internodal distances, high sexual reproductive effort and the increasing occurrence of apical shoots in seagrasses from SB sites indicated that the plants were under stress and not growing under optimal conditions. Moreover, AG and BG biomass and internodal distances decreased further with the addition of fertilizer in SB sites. Results presented here highlight the fact that H. stipulacea is one of the most tolerant and plastic seagrass species. Our study further demonstrates that the effects of fertilization differ significantly between meadows that are growing exposed to different levels of anthropogenic pressures. Thus, the meadow's "history" affects it resilience and response to further stress. Our results suggest that monitoring efforts on H. stipulacea populations in its native range should focus especially on carbohydrate reserves in leaves and rhizomes, LAI, internodal length and percentage of apical shoots as suitable warning indicators for nutrient stress in this seagrass species to minimize future impacts on these valuable ecosystems.

Unusually Warm Summer Temperatures Exacerbate Population and Plant Level Response of Posidonia oceanica to Anthropogenic Nutrient Stress.

Posidonia oceanica is a key foundation species in the Mediterranean providing valuable ecosystem services. However, this species is particularly vulnerable towards high coastal nutrient inputs and the rising frequency of intense summer heat waves, but their combined effect in situ has received little attention so far. Here, we investigated the effects of in situ nutrient addition during an unusually warm summer over a 4-month period, comparing different morphological, physiological and biochemical population metrics of seagrass meadows growing in protected areas (Ischia) with meadows already exposed to significant anthropogenic pressure (Baia - Gulf of Pozzuoli). Our study highlights that the effects of warmer than usual summer temperatures on the population level of seagrass meadows can be exacerbated if the plants are already exposed to higher anthropogenic pressures. Morphological and population level indicators mainly changed over time, possibly impacted by season and the warmer temperatures, and displayed more pronounced reductions in seagrasses from impacted sites. The additional nutrient supply had even more deleterious effects, as shown by a decrease in approximately 67% in cover in fertilized plots at high impacted sites and 33% at low impacted sites. Moreover, while rhizome starch concentration showed a seasonal increase in plants from low impacted sites it displayed a trend of a 27% decrease in fertilized plots of the high impacted sites. Epiphyte biomass was approximately four-fold higher on leaves of plants growing in impacted sites and even doubled with the additional nutrient input. Predicting and anticipating stress in P. oceanica is of crucial importance for conservation and management efforts, given the limited colonizing and reproductive ability and extremely slow growth of this ecosystem engineer. Our results suggest that monitoring efforts should focus especially on leaf area index (LAI), carbohydrate concentrations in the rhizomes, and epiphyte cover on leaves as indicators of the onset of stress in Posidonia oceanica, which can be used by decision makers to take appropriate measures before damage to the ecosystem becomes irreversible, minimize future human interference and strengthen the resilience of these important ecosystems.

Projected Rapid Habitat Expansion of Tropical Seagrass Species in the Mediterranean Sea as Climate Change Progresses.

During the last 150 years, the tropical seagrass species Halophila stipulacea has established itself in the southern and eastern parts of the Mediterranean Sea. More recently (2018), Halophila decipiens was observed for the first time in the eastern Mediterranean, and was described as the second non-native seagrass species in the Mediterranean Sea. We implemented a species distribution model (SDM) approach to (1) hindcast the habitat suitability of H. stipulacea over the last 100 years in the Mediterranean basin, and (2) to model the increase in the potential habitat suitability of H. stipulacea and H. decipiens during the current century under two very different climate scenarios, RCP 2.6 (lowest carbon emission scenario) and RCP 8.5 (highest carbon emission scenario). In addition, a principal component analysis (PCA) and k-means cluster based on temperature and salinity drivers were applied to visualize the distance and relatedness between the native and invasive H. stipulacea and H. decipiens populations. Results from this PCA suggest that the H. stipulacea populations of the Mediterranean and Red Sea are likely to be similar. In contrast, H. decipiens from the Mediterranean is more related to the Atlantic populations rather than to the Red Sea populations. The hindcast model suggests that the expansion of H. stipulacea was related to the increases in seawater temperatures in the Mediterranean over the last 100 years. The SDMs predict that more suitable habitat will become available for both tropical species during this century. The habitat suitability for H. stipulacea will keep expanding westward and northward as the Mediterranean continues to become saltier and warmer. In comparison, the SDMs built for H. decipiens forecast a restricted habitat suitability in the south-eastern Mediterranean Sea at the present environmental conditions and predicts a progressive expansion with a potential increase in habitat suitability along 85% of the Mediterranean coastline. The predicted rapid expansion of non-native seagrass species could alter the Mediterranean's seagrass community and may entail massive impacts on associated ecosystem functions and services, impacts that have severe socio-economic consequences.

Do differences in mating behaviour lead to differences in connectivity patterns of reef fishes? Insights from two sympatric surgeonfish species in the Indian Ocean.

Disentangling the contribution of biotic and abiotic factors in the structuring of the genetic diversity of reef species is critical to illuminate the diversification of evolutionary lineages in marine environment. Howevr, previous studies have mainly focused on determining the influence of pelagic larval duration on the connectivity and demographic history of reef fishes, whereas few studies have examined the effects of other biotic factors, such as mating behaviour and habitat preference. Here, we use mitochondrial DNA (ATPase 6/8) and ten microsatellite loci to compare the population genetic structure and demographic history of the spawning aggregating Acanthurus triostegus with the monogamous spawning Acanthurus leucosternon. Pairwise comparisons and discriminant analysis of principal components showed that the genetic structuring patterns of the two species are not consistent with the influence of mating behaviour, suggesting the possible role of other biotic and abiotic factors. However, demographic history estimates revealed that these species may have responded differently to sea level fluctuations during the glacial maxima.

Climate change drives trait-shifts in coral reef communities.

Climate change is expected to have profound, partly unforeseeable effects on the composition of functional traits of complex ecosystems, such as coral reefs, and some ecosystem properties are at risk of disappearing. This study applies a novel spatially explicit, individual-based model to explore three critical life history traits of corals: heat tolerance, competitiveness and growth performance under various environmental settings. Building upon these findings, we test the adaptation potential required by a coral community in order to not only survive but also retain its diversity by the end of this century under different IPCC climate scenarios. Even under the most favourable IPCC scenario (Representative Concentration Pathway, RCP 2.6), model results indicate that shifts in the trait space are likely and coral communities will mainly consist of small numbers of temperature-tolerant and fast-growing species. Species composition of coral communities is likely to be determined by heat tolerance, with competitiveness most likely playing a subordinate role. To sustain ~15% of current coral cover under a 2 °C temperature increase by the end of the century (RCP 4.5), coral systems would have to accommodate temperature increases of 0.1-0.15 °C per decade, assuming that periodic extreme thermal events occurred every 8 years. These required adaptation rates are unprecedented and unlikely, given corals' life-history characteristics.

Genetic population structure of the convict surgeonfish Acanthurus triostegus: a phylogeographic reassessment across its range.

This study investigates the genetic population structure and connectivity of Acanthurus triostegus in five Indo-Pacific biogeographic regions (western and eastern Indian Ocean, western, central and eastern Pacific Ocean), using a mitochondrial DNA marker spanning the ATPase8 and ATPase6 gene regions. In order to assess the phylogeography and genetic population structure of A. triostegus across its range, 35 individuals were sampled from five localities in the western Indian Ocean and complemented with 227 sequences from two previous studies. Results from the overall analysis of molecular variance (AMOVA) without a priori grouping showed evidence of significant differentiation in the Indo-Pacific, with 25 (8.3%) out of 300 pairwise ΦST comparisons being significant. However, the hierarchical AMOVA grouping of Indian and Pacific Ocean populations failed to support the vicariance hypothesis, showing a lack of a genetic break between the two ocean basins. Instead, the correlation between pairwise ΦST values and geographic distance showed that dispersal of A. triostegus in the Indo-Pacific Ocean follows an isolation-by-distance model. Three haplogroups could be deduced from the haplotype network and phylogenetic tree, with haplogroup 1 and 2 dominating the Indian and the Pacific Ocean, respectively, while haplogroup 3 exclusively occurring in the Hawaiian Archipelago of the central Pacific Ocean.

Simulations of long-term community dynamics in coral reefs--how perturbations shape trajectories.

Tropical coral reefs feature extraordinary biodiversity and high productivity rates in oligotrophic waters. Due to increasing frequencies of perturbations--anthropogenic and natural--many reefs are under threat. Such perturbations often have devastating effects on these unique ecosystems and especially if they occur simultaneously and amplify each other's impact, they might trigger a phase shift and create irreversible conditions.We developed a generic, spatially explicit, individual-based model in which competition drives the dynamics of a virtual benthic reef community--comprised of scleractinian corals and algae--under different environmental settings. Higher system properties, like population dynamics or community composition arise through self-organization as emergent properties. The model was parameterized for a typical coral reef site at Zanzibar, Tanzania and features coral bleaching and physical disturbance regimes as major sources of perturbations. Our results show that various types and modes (intensities and frequencies) of perturbations create diverse outcomes and that the switch from high diversity to single species dominance can be evoked by small changes in a key parameter. Here we extend the understanding of coral reef resilience and the identification of key processes, drivers and respective thresholds, responsible for changes in local situations. One future goal is to provide a tool which may aid decision making processes in management of coral reefs.

Status of feral oilseed rape in Europe: its minor role as a GM impurity and its potential as a reservoir of transgene persistence.

PURPOSE: Feral oilseed rape has become widespread in Europe on waysides and waste ground. Its potential as a source of GM impurity in oilseed rape harvests is quantified, for the first time, by a consistent analysis applied over a wide range of study areas in Europe. METHODS: The maximum contribution of feral oilseed rape to impurities in harvested crops was estimated by combining data on feral abundance and crop yield from five established, demographic studies in agricultural habitats in Denmark, Germany (2), France and the UK, constituting over 1,500 ha of land and 16 site-years of observations. Persistence of feral populations over time was compared by visual and molecular methods. RESULTS: Ferals had become established in all regions, forming populations 0.2 to 15 km⁻². The seed they produced was always <0.0001% of the seed on crops of oilseed rape in each region. The contribution of ferals to impurity in crops through accidental harvest of seed and through cross-pollination would be an even smaller percentage. Feral oilseed rape nevertheless showed a widespread capacity to persist in all regions and retain traits from varieties no longer grown. CONCLUSIONS: Feral oilseed rape is not a relevant source of macroscopic impurity at its present density in the landscape but provides opportunity for genetic recombination, stacking of transgenes and the evolution of genotypes that under strong selection pressure could increase and re-occupy fields to constitute an economic weed burden and impurity in future crops.

Information system for monitoring environmental impacts of genetically modified organisms.

BACKGROUND, AIM AND SCOPE: European legislation stipulates that genetically modified organisms (GMO) have to be monitored to identify potential adverse environmental effects. A wealth of different types of monitoring data from various sources including existing environmental monitoring programmes is expected to accumulate. This requires an information system to efficiently structure, process and evaluate the monitoring data. METHODS: A structure for an Information System for Monitoring GMO (ISMO) was developed by a multidisciplinary research team. It is based on the requirement to organise all relevant information in a logical, readily accessible and functional manner. RESULTS: For the ISMO, we present a combination of three interrelated components: Firstly, an ISMO should comprise a knowledge database structured according to information related to the different scale levels of biological organisation relevant to GMO monitoring and scientific hypotheses on cause-effects which should be validated by monitoring data. Secondly, a monitoring database should be part of an ISMO containing GMO-specific monitoring data and meta-data. This monitoring database should be linked with monitoring data from other monitoring programmes which are relevant for GMO-related questions. Thirdly, an ISMO should encompass a database covering administrative and procedural data. Neither national nor international approaches to an ISMO exist yet. CONCLUSIONS: An ISMO as designed in this paper could support competent authorities in both the GMO notification process and in post-market monitoring. This includes evaluating the environmental risks of experimentally releasing GMO and placing them on the market, assessing monitoring plans and evaluating monitoring results. The ISMO should be implemented on both the national and international level, preferably combining different administrative scales. Harmonisation approaches towards GMO monitoring data are at an initial stage, but they are a precondition to coordinated GMO monitoring and to successfully implementing an ISMO. It is recommended to set up a legal basis and to agree on common strategies for the data coordination and harmonisation.

Hazard mitigation or mitigation hazard?

BACKGROUND, AIM AND SCOPE: Transgenic oilseed rape (Brassica napus L.; OSR) is estimated to be environmentally and economically problematic because volunteers and ferals occur frequently and because of its hybridisation potential with several wild and weedy species. A proposed mitigation strategy aims to reduce survival, in particular in conventional OSR crops, by coupling the transgenic target modification with a dwarfing gene to reduce competitive fitness. Our study allowed us to access potential ecological implications of this strategy. MATERIALS AND METHODS: On a large scale (>500 km(2)), we recorded phenological and population parameters of oilseed rape plants for several years in rural and urban areas of Northern Germany (Bremen and surroundings). The characterising parameter were analysed for differences between wild and cultivated plants. RESULTS: In rural areas, occurrences of feral and volunteer OSR together had an average density of 1.19 populations per square kilometre, in contrast to urban areas where we found 1.68 feral populations per square kilometre on average. Throughout the survey, the vegetation cover at the locations with feral OSR ranged from less than 10% to 100%. Our investigations gave clear empirical evidence that feral OSR was, on average, at least 41% smaller than cultivated OSR, independent of phenological state after onset of flowering. DISCUSSION: The findings can be interpreted as phenotypic adaptation of feral OSR plants. Therefore, it must be asked whether dwarfing could be interpreted as an improvement of pre-adaptation to feral environments. In most of the sites where feral plants occurred, germination and establishment were in locations with disturbed vegetation cover, allowing initial growth without competition. Unless feral establishment of genetically modified dwarfed traits are specifically studied, it would not be safe to assume that the mitigation strategy of dwarfing also reduces dispersal in feral environments. CONCLUSIONS AND RECOMMENDATIONS: With respect to OSR, we argue that the proposed mitigation approach could increase escape and persistence of transgene varieties rather than reducing them. We conclude that the development of effective hazard mitigation measures in the risk evaluation of genetically modified organisms requires thorough theoretical and empirical ecological analyses rather than assumptions about abstract fitness categories that apply only in parts of the environment where the plant can occur.