The functioning of coral reef communities along environmental gradients

One of the primary challenges in ecology is to understand how environmental disturbance affects diversity and community structure, and what are the subsequent consequences on ecosystem functioning. Coral reefs are some of the most diverse ecosystems on the planet resulting in complex sets of interactions between benthic, habitat-forming constituents and mobile fish consumers. However, scleractinian corals, the primary habitat engineers, are dependent on high-light, low-nutrient water conditions and thus are highly responsive when the environment varies from this status. In Southeast Asia, an increase in human coastal populations centred around urban areas has resulted in extensive changes to the coastal environment such as degraded water quality and removal of fish consumers. This has resulted in highly varied abiotic and biotic conditions in relation with distance from the shore. Often, coral reefs closer to shore are much lower in benthic and fish diversity than those further from anthropogenic influences, with direct impacts on ecosystem functioning. Therefore the aim of this thesis was to explore coral reef ecosystem functioning with respect to changes in benthic community structure and fish diversity in relation to varying environmental conditions in the Spermonde Archipelago, Indonesia. A combination of observational, experimental and theoretical analyses were conducted on the functioning of coral reefs using eight islands on a transect of increasing distance from the mainland, varying from 1 to 55 km. At these eight sites, benthic and pelagic surveys identified variation in the status of coral reef communities, while recruitment and feeding assays identified variation in important ecological processes. Lastly, experimental observations were further qualified with stable isotope analysis and the application of contemporary indices of functional diversity. It was found that indeed, the coral reefs varied along a continuum of structure, assemblage and processes. Increasing distance from shore was associated with greater live coral cover and structural complexity, while sites closer to shore were dominated by turf algae and rubble. Furthermore, turf algae was observed as playing a particularly important role, as this group was dominant during recruitment and subsequent development of open benthic space as supplied by terracotta tiles. Fish diversity, along with redundancy in the important herbivore group, also increased with distance from shore, resulting in an increasingly diverse response to Sargassum and Padina assays. The functional composition of the fish assemblages became increasingly variable with loss in coral cover and structural complexity, suggesting communities become destabilised under habitat degradation. Furthermore, stable isotope analysis indicated that the trophic niche of a fish species can increase at sites with more degradation suggesting varying functional utility. However, functioning is not determined only by exposure to chronic, abiotic conditions. Outbreaks of the crown-of-thorns starfish (Acanthaster planci) and mechanical destruction (bomb fishing) resulted in extreme loss of live coral. At these sites, biological and functional diversity displayed some of the lowest values among all sites. Coral reefs can exist in systems with altered water condition if physiological and ecological capacity of the organisms allow for their continuation. Nevertheless, degraded water condition will select against many species, resulting not only in the observed lower biological diversity, but also in less species taking part in functional roles as reflected in higher functional variability. Combined, these results show that the functioning of coral reefs does not exist in discrete states; rather, their functioning is a result of abiotic stressors and biological feedbacks. It is becoming increasingly clear that pristine coral reefs are not a reality in many cases around the world. Thus understanding coral reef functioning at all stages of degradation will help with future management. This thesis adds to the ever-growing knowledge about disturbed coral reefs, but more importantly, it describes the changing relationship between diversity and functioning of coral reefs in relation to disturbance

Trophic ecology of parrotfish in Zanzibar : applications of stable isotope analysis

Parrotfish are a critical component of the herbivore functional group on tropical coral reefs around the world because they mediate competition that occurs between algae and scleractinian corals. Also, because of their feeding technique, which consists of rasping at the substratum with their beak-like teeth, they play an important role in carbonate turnover and the clearing of reef surface area for the settlement of new sessile organisms. Because of these roles, parrotfishes are an important structuring component of coral reef communities. However, individual species can play different roles depending on their physiology, behaviour and ecology. Despite the possible ecological differences that may exist amongst species, specific roles of the fishes remain unclear as the group is most often studied at higher community levels. This thesis applied stable isotope analysis to differing levels of organisation within a parrotfish community to help elucidate their trophic ecology on coral reefs in Zanzibar. Firstly, blood and muscle tissues were compared to identify differences in their isotope signatures. In other organisms, blood turns over faster than muscle tissue so that muscle tissue represents the diet as integrated over a longer period of time. In most species of parrotfish the blood and muscle δ¹³C signatures were not found to be significantly different, but the δ¹⁵N signatures were significantly different between tissues. This indicated that the δ¹³C signature of both tissues would reveal similar dietary information. Conversely, differences in the δ¹⁵N signature indicated that the nitrogen relationship between tissues was more complicated. Secondly, spatial variability in parrotfish, coral, detritus and macroalgae isotope signatures was assessed at different scales. In macroalgae and coral tissues (zooxanthellae and polyp treated separately), the δ¹³C signatures were shown to differ with depth, presumably because of changes in photosynthetic processes related to depth-associated changes in light. While δ¹⁵N signatures were not affected by depth, all organisms showed enrichment at the Nyange reef, the closest reef to the capital of Zanzibar, Stone Town, presumably reflecting the effects of sewage outfall. These results show that processes that impact the δ¹⁵N signatures of primary producers (macroalgae and zooxanthellae) can be traced to higher trophic levels (coral polyps and fish). Lastly, δ¹³C and δ¹⁵N signatures were used to identify ontogenetic dietary changes in multiple species of parrotfish. Four of the species showed stages that varied from the diets that are normally assumed on the basis of their dentition and feeding technique. This indicates that functional roles based on taxonomy or morphology may fail to include possible ontogenetic dietary changes, and may also fail to elucidate the full impact a species could have on coral reef communities. The conclusions from these studies indicate that the species-specific ecological role of parrotfish in coral reef communities can be complex within and between species, and may differ amongst reefs. In light of the natural and anthropogenic pressures that affect coral reef systems, management decisions based on a more complete understanding of the role of these fish in coral reef communities will help decisions that maintain resilience in these fragile systems

Contrasting responses in the niches of two coral reef herbivores along a gradient of habitat disturbance in the Spermonde Archipelago, Indonesia

Habitat modification of coral reefs is becoming increasingly common due to increases in coastal urban populations. Coral reef fish are highly dependent on benthic habitat; however, information on species-specific responses to habitat change, in particular with regard to trophic strategies, remains scarce. This study identifies variation in the trophic niches of two herbivorous coral reef fishes with contrasting trophic strategies, using Stable Isotopes Bayesian Ellipses in R, along a spatial gradient of changing coral reef habitats. In the parrotfish Chlorurus bleekeri, a roving consumer, the range of δ15N and δ13C and their niche area displayed significant relationships with the amount of rubble in the habitat. In contrast, the farming damselfish, Dischistodus prosopotaenia, showed a narrow range of both δ15N and δ13C, displaying little change in niche parameters among sites. This may indicate that parrotfish vary their feeding according to habitat, while the damselfish continue to maintain their turf and invertebrate resources. Assessing isotopic niches may help to better understand the specific trophic responses to change in the environment. Furthermore, the use of isotopic niches underlines the utility of stable isotopes in studying the potential impacts of environmental change on feeding ecology

Spatio-temporal patterns in the coral reef communities of the Spermonde Archipelago, 2012–2014, II: Fish assemblages display structured variation related to benthic condition

The Spermonde Archipelago is a complex of ~70 mostly populated islands off Southwest Sulawesi, Indonesia, in the center of the Coral Triangle. The reefs in this area are exposed to a high level of anthropogenic disturbances. Previous studies have shown that variation in the benthos is strongly linked to water quality and distance from the mainland. However, little is known about the fish assemblages of the region and if their community structure also follows a relationship with benthic structure and distance from shore. In this study, we used eight islands of the archipelago, varying in distance from 1 to 55 km relative to the mainland, and 3 years of surveys, to describe benthic and fish assemblages and to examine the spatial and temporal influence of benthic composition on the structure of the fish assemblages. Cluster analysis indicated that distinct groups of fish were associated with distance, while few species were present across the entire range of sites. Relating fish communities to benthic composition using a multivariate generalized linear model confirmed that fish groups relate to structural complexity (rugosity) or differing benthic groups; either algae, reef builders (coral and crustose coralline algae) or invertebrates and rubble. From these relationships we can identify sets of fish species that may be lost given continued degradation of the Spermonde reefs. Lastly, the incorporation of water quality, benthic and fish indices indicates that local coral reefs responded positively after an acute disturbance in 2013 with increases in reef builders and fish diversity over relatively short (1 year) time frames. This study contributes an important, missing component (fish community structure) to the growing literature on the Spermonde Archipelago, a system that features environmental pressures common in the greater Southeast Asian region

The Tropospheric Distribution and Budget of Ethane

From about 1500 measurements of ethane in the remote troposphere the longitudinally and vertically averaged latitudinal and seasonal variability of ethane was derived. To improve the data coverage, several data sets from literature were included. There are only very few data sets available for the southern hemisphere. Nevertheless, the uncertainty of the average seasonal/latitudinal ethane profile is estimated to less than 30%. The global annually averaged ethane mixing ratio is 860 ppt. There is a strong interhemispheric gradient with an average north/south ratio of 3.5. Within the northern hemisphere there is an average gradient from the highest annual mean value of 2500 ppt around 65°N to about 600 ppt at the equator. In the southern hemisphere there is only a small gradient at low latitudes and at middle and high southern latitudes no significant gradient can be seen. In both hemispheres a significant seasonal cycle with highest mixing ratios in late winter is observed. The ethane source strength needed to balance the atmospheric budget of ethane is estimated to 15.5 Tg/yr, with most of the emissions in the northern hemisphere. An independent estimate of the sources indicate that most of the emissions are due to natural gas losses (6 Tg/yr) and biomass burning (6.4 Tg/yr). This is also compatible with the latitudinal and seasonal variation of the atmospheric ethane removal rates. However, these estimates have substantial uncertainties and it should be noted that the role of the biosphere for the atmospheric budget of ethane is presently not understood

Spatio-temporal patterns in coral reef communities of the Spermonde Archipelago, 2012-2014, I: Comprehensive reef monitoring of water and benthic indicators reflect changes in reef health

Pollution, fishing, and outbreaks of predators can heavily impact coastal coral reef ecosystems, leading to decreased water quality and benthic community shifts. To determine the main environmental drivers of coral reef status in the Spermonde Archipelago, Indonesia, we monitored environmental variables and coral reef benthic community structure along an on-to-offshore gradient annually from 2012 to 2014. Findings revealed that concentrations of phosphate, chlorophyll a-like fluorescence, suspended particulate matter, and light attenuation significantly decreased from on-to-offshore, while concentrations of dissolved O2 and values of water pH significantly increased on-to-offshore. Nitrogen stable isotope signatures of sediment and an exemplary common brown alga were significantly enriched nearshore, identifying wastewater input from the city of Makassar as primary N source. In contrast to the high temporal variability in water quality, coral reef benthic community cover did not show strong temporal, but rather, spatial patterns. Turf algae was the dominant group next to live coral, and was negatively correlated to live coral, crustose coralline algae (CCA), rubble and hard substrate. Variation in benthic cover along the gradient was explained by water quality variables linked to trophic status and physico-chemical variables. As an integrated measure of reef status and structural complexity, the benthic index, based on the ratio of relative cover of live coral and CCA to other coral reef organisms, and reef rugosity were determined. The benthic index was consistently low nearshore and increased offshore, with high variability in the midshelf sites across years. Reef rugosity was also lowest nearshore and increased further offshore. Both indices dropped in 2013, increasing again in 2014, indicating a period of acute disturbance and recovery within the study and suggesting that the mid-shelf reefs are more resilient to disturbance than nearshore reefs. We thus recommend using these two indices with a selected number of environmental variables as an integral part of future reef monitoring

Ecological effects of full and partial protection in the crowded Mediterranean Sea: a regional meta-analysis

Marine protected areas (MPAs) are a cornerstone of marine conservation. Globally, the number and coverage of MPAs are increasing, but MPA implementation lags in many human-dominated regions. In areas with intense competition for space and resources, evaluation of the effects of MPAs is crucial to inform decisions. In the human-dominated Mediterranean Sea, fully protected areas occupy only 0.04% of its surface. We evaluated the impacts of full and partial protection on biomass and density of fish assemblages, some commercially important fishes, and sea urchins in 24 Mediterranean MPAs. We explored the relationships between the level of protection and MPA size, age, and enforcement. Results revealed significant positive effects of protection for fisheries target species and negative effects for urchins as their predators benefited from protection. Full protection provided stronger effects than partial protection. Benefits of full protection for fish biomass were only correlated with the level of MPA enforcement; fish density was higher in older, better enforced, and - interestingly - smaller MPAs. Our finding that even small, well-enforced, fully protected areas can have significant ecological effects is encouraging for "crowded" marine environments. However, more data are needed to evaluate sufficient MPA sizes for protecting populations of species with varying mobility levels

Five key attributes can increase marine protected areas performance for small-scale fisheries management

Marine protected areas (MPAs) have largely proven to be effective tools for conserving marine ecosystem, while socio-economic benefits generated by MPAs to fisheries are still under debate. Many MPAs embed a no-take zone, aiming to preserve natural populations and ecosystems, within a buffer zone where potentially sustainable activities are allowed. Small-scale fisheries (SSF) within buffer zones can be highly beneficial by promoting local socio-economies. However, guidelines to successfully manage SSFs within MPAs, ensuring both conservation and fisheries goals, and reaching a win-win scenario, are largely unavailable. From the peer-reviewed literature, grey-literature and interviews, we assembled a unique database of ecological, social and economic attributes of SSF in 25 Mediterranean MPAs. Using random forest with Boruta algorithm we identified a set of attributes determining successful SSFs management within MPAs. We show that fish stocks are healthier, fishermen incomes are higher and the social acceptance of management practices is fostered if five attributes are present (i.e. high MPA enforcement, presence of a management plan, fishermen engagement in MPA management, fishermen representative in the MPA board, and promotion of sustainable fishing). These findings are pivotal to Mediterranean coastal communities so they can achieve conservation goals while allowing for profitable exploitation of fisheries resources

Collaboration scripts - a conceptual analysis

This article presents a conceptual analysis of collaboration scripts used in face-to-face and computer-mediated collaborative learning. Collaboration scripts are scaffolds that aim to improve collaboration through structuring the interactive processes between two or more learning partners. Collaboration scripts consist of at least five components: (a) learning objectives, (b) type of activities, (c) sequencing, (d) role distribution, and (e) type of representation. These components serve as a basis for comparing prototypical collaboration script approaches for face-to-face vs. computer-mediated learning. As our analysis reveals, collaboration scripts for face-to-face learning often focus on supporting collaborators in engaging in activities that are specifically related to individual knowledge acquisition. Scripts for computer-mediated collaboration are typically concerned with facilitating communicative-coordinative processes that occur among group members. The two lines of research can be consolidated to facilitate the design of collaboration scripts, which both support participation and coordination, as well as induce learning activities closely related to individual knowledge acquisition and metacognition. In addition, research on collaboration scripts needs to consider the learners’ internal collaboration scripts as a further determinant of collaboration behavior. The article closes with the presentation of a conceptual framework incorporating both external and internal collaboration scripts