Characterization and management of the commercial sector of the Pohnpei coral reef fishery,Micronesia

Commercial coral reef fisheries in Pohnpei (Micronesia) extract approximately 1,521 kg of reef fish daily (∼500 MT year⁻¹) from 152 km² of surrounding reef. More than 153 species were represented during surveys, with 25 species very common or common within combined-gear catch. Acanthurids contributed the greatest to catch volume, with bluespine unicornfish, Naso unicornis, and orangespine unicornfish, Naso lituratus, among the most frequently observed herbivores. Nighttime spearfishing was the dominant fishing method and inner lagoon areas were primarily targeted. A seasonal sales ban (March–April), intended to reduce pressure on reproductively active serranids, significantly increased the capture volume of other families. Catch was significantly greater during periods of low lunar illumination, suggesting higher fishing success or greater effort, or both. The marketed catch was dominated by juveniles and small adults, based on fishes of known size at sexual maturity. Artificially depressed market prices appear to be catalyzing (potential or realized) overfishing by increasing the volume of fish needed to offset rising fuel prices. These results support the need for comprehensive fisheries management that produces sustainable fishing and marketing practices and promotes shared management and enforced responsibilities between communities and the state. To be effective, management should prohibit nighttime spearfishing. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.

Phase shifts and the role of herbivory in the resilience of coral reefs

Cousin Island marine reserve (Seychelles) has been an effectively protected no-take marine protected area (MPA) since 1968 and was shown in 1994 to support a healthy herbivorous fish assemblage. In 1998 Cousin Island reefs suffered extensive coral mortality following a coral bleaching event, and a phase shift from coral to algal dominance ensued. By 2005 mean coral cover was <1%, structural complexity had fallen and there had been a substantial increase in macroalgal cover, up to 40% in some areas. No clear trends were apparent in the overall numerical abundance and biomass of herbivorous fishes between 1994 and 2005, although smaller individuals became relatively scarce, most likely due to the loss of reef structure. Analysis of the feeding habits of six abundant and representative herbivorous fish species around Cousin Island in 2006 demonstrated that epilithic algae were the preferred food resource of all species and that macroalgae were avoided. Given the current dominance of macroalgae and the apparent absence of macroalgal consumers, it is suggested that the increasing abundance of macroalgae is reducing the probability of the system reverting to a coral dominated state.     

Experimental biology of coral reef ecosystems

Coral reef ecosystems are at the crossroads. While significant gaps still exist in our understanding of how “normal” reefs work, unprecedented changes in coral reef systems have forced the research community to change its focus from basic research to understand how one of the most diverse ecosystems in the world works to basic research with strong applied implications to alleviate damage, save, or restore coral reef ecosystems. A wide range of stressors on local, regional, and global spatial scales including over fishing, diseases, large-scale disturbance events, global climate change (e.g., ozone depletion, global warming), and over population have all contributed to declines in coral cover or phase shifts in community structure on time scales never observed before. Many of these changes are directly or indirectly related to anthropogenically induced changes in the global support network that affects all ecosystems. This review focuses on some recent advances in the experimental biology of coral reef ecosystems, and in particular scleractinian corals, at all levels of biological organization. Many of the areas of interest and techniques discussed reflect a progression of technological advances in biology and ecology but have found unique and timely application in the field of experimental coral reef biology. The review, by nature, will not be exhaustive and reflects the author's interests to a large degree. Because of the voluminous literature available, an attempt has been made to capture the essential elements and references for each topic discussed.     

Larval retention and connectivity among populations of corals and reef fishes: history,advances and challenges

The extent of larval dispersal on coral reefs has important implications for the persistence of coral reef metapopulations, their resilience and recovery from an increasing array of threats, and the success of protective measures. This article highlights a recent dramatic increase in research effort and a growing diversity of approaches to the study of larval retention within (self-recruitment) and dispersal among (connectivity) isolated coral reef populations. Historically, researchers were motivated by alternative hypotheses concerning the processes limiting populations and structuring coral reef assemblages, whereas the recent impetus has come largely from the need to incorporate dispersal information into the design of no-take marine protected area (MPA) networks. Although the majority of studies continue to rely on population genetic approaches to make inferences about dispersal, a wide range of techniques are now being employed, from small-scale larval tagging and paternity analyses, to large-scale biophysical circulation models. Multiple approaches are increasingly being applied to cross-validate and provide more realistic estimates of larval dispersal. The vast majority of empirical studies have focused on corals and fishes, where evidence for both extremely local scale patterns of self-recruitment and ecologically significant connectivity among reefs at scales of tens of kilometers (and in some cases hundreds of kilometers) is accumulating. Levels of larval retention and the spatial extent of connectivity in both corals and fishes appear to be largely independent of larval duration or reef size, but may be strongly influenced by geographic setting. It is argued that high levels of both self-recruitment and larval import can contribute to the resilience of reef populations and MPA networks, but these benefits will erode in degrading reef environments.     

Historical and contemporary evidence of a mutton snapper (<Emphasis Type="Italic">Lutjanus analis</Emphasis> Cuvier, 1828) spawning aggregation fishery in decline

Scientific information on reef fish spawning aggregation fisheries is sparse in light of numerous regional declines and extirpations from overexploitation. Fisher interviews of the small-scale commercial mutton snapper (Lutjanus analis) spawning aggregation fishery at Gladden Spit, Belize, suggests a historic decadal decline. The reported trend is supported by analysis of inter-seasonal catch and effort and yield (2000–2002) that reveals a 59% decline in catch per unit effort (CPUE) and a 22% decrease in mean landings per boat. Declining population-level trends are also supported by a significant decrease in inter-annual median lengths of mutton snappers (2000–2006). These findings demonstrate the need for additional life history information that includes length-associated age and details on growth to provide clearer support of the effects on, and responses by, populations following fishing. In view of the historical changes to mutton snapper CPUE and landings at Gladden Spit and the fishery-associated declines in fish spawning aggregations observed globally, a precautionary approach to spawning aggregation management is warranted that provides full protection from fishing to enhance population persistence. The findings also highlight the need for substantially greater enforcement and long-term fisheries monitoring under a comprehensive regional management strategy.     

White-band disease and the changing face of Caribbean coral reefs

In recent decades, the cover of fleshy macroalgae has increased and coral cover has decreased on most Caribbean reefs. Coral mortality precipitated this transition, and the accumulation of macroalgal biomass has been enhanced by decreased herbivory and increased nutrient input. Populations of Acropora palmata (elkhorn coral) and A. cervicornis (staghorn coral), two of the most important framework-building species, have died throughout the Caribbean, substantially reducing coral cover and providing substratum for algal growth. Hurricanes have devastated local populations of Acropora spp. over the past 20–25 years, but white-band disease, a putative bacterial syndrome specific to the genus Acropora, has been a more significant source of mortality over large areas of the Caribbean region.Paleontological data suggest that the regional Acropora kill is without precedent in the late Holocene. In Belize, A. cervicornis was the primary ecological and geological constituent of reefs in the central shelf lagoon until the mid-1980s. After constructing reef framework for thousands of years, A. cervicornis was virtually eliminated from the area over a ten-year period. Evidence from other parts of the Caribbean supports the hypothesis of continuous Holocene accumulation and recent mass mortality of Acropora spp. Prospects are poor for the rapid recovery of A. cervicornis, because its reproductive strategy emphasizes asexual fragmentation at the expense of dispersive sexual reproduction. A. palmata also relies on fragmentation, but this species has a higher rate of sexual recruitment than A. cervicornis. If the Acropora spp. do not recover, macroalgae will continue to dominate Caribbean reefs, accompanied by increased abundances of brooding corals, particularly Agaricia spp. and Porites spp. The outbreak of white-band disease has been coincident with increased human activity, and the possibility of a causal connection should be further investigated.     

Fish assemblages on sunken vessels and natural reefs in southeast Florida,USA

Derelict ships are commonly deployed as artificial reefs in the United States, mainly for recreational fishers and divers. Despite their popularity, few studies have rigorously examined fish assemblages on these structures and compared them to natural reefs. Six vessel-reefs off the coast of southeast Florida were censused quarterly (two ships per month) to characterize their associated fish assemblages. SCUBA divers used a non-destructive point-count method to visually assess the fish assemblages over 13- and 12-month intervals (March 2000 to March 2001 and March 2002 to February 2003). During the same intervals, fish assemblages at neighboring natural reefs were also censused. A total of 114,252 fishes of 177 species was counted on natural and vessel-reefs combined. Mean fish abundance and biomass were significantly greater on vessel-reefs in comparison to surrounding natural reef areas. Haemulidae was the most abundant family on vessel-reefs, where it represented 46% of total fish abundance. The most abundant family on natural reefs was Labridae, where it accounted for 24% of total fish abundance. Mean species richness was significantly greater on vessel-reefs than neighboring natural reefs and also differed among vessel-reefs. Both mean fish abundance and mean species richness were not significantly different between natural reefs neighboring vessel-reefs and natural reefs with no artificial structures nearby. This suggests the vessel-reefs are not, in general, attracting fish away from neighboring natural reefs in our area. Additionally, economically important fish species seem to prefer vessel-reefs, as there was a greater abundance of these species on vessel-reefs than surrounding natural reef areas. Fish assemblage structure on natural versus artificial reefs exhibited a low similarity (25.8%). Although no one species was responsible for more than 6% of the total dissimilarity, fish assemblage trophic structure differed strikingly between the two reef types. Planktivores dominated on vessel-reefs, accounting for 54% of the total abundance. Conversely, planktivores only made up 27% of total abundance on natural reefs. The results of this study indicate vessel-reef fish assemblages are unique and that these fishes may be utilizing food resources and habitat characteristics not accessible from or found at natural reefs in our area. Production may also be occurring at vessel-reefs as the attraction of fish species from nearby natural reefs seems to be minimal. Electronic supplementary material Supplementary material is available for this article at and accessible for authorised users     

Designing marine reserves to reflect local socioeconomic conditions: lessons from long-enduring customary management systems

Coral reef conservation strategies such as marine protected areas have met limited success in many developing countries. Some researchers attribute part of these shortcomings to inadequate attention to the social context of conserving marine resources. To gain insights into applying Western conservation theory more successfully in the socioeconomic context of developing countries, this study examines how long-enduring, customary reef closures appear to reflect local socioeconomic conditions in two Papua New Guinean communities. Attributes of the customary management (including size, shape, permanence, and gear restrictions) are examined in relation to prevailing socioeconomic conditions (including resource users’ ability to switch gears, fishing grounds, and occupations). Customary closures in the two communities appear to reflect local socioeconomic circumstances in three ways. First, in situations where people can readily switch between occupations, full closures are acceptable with periodic harvests to benefit from the closure. In comparison, communities with high dependence on the marine resources are more conducive to employing strategies that restrict certain gear types while still allowing others. Second, where there is multiple clan and family spatial ownership of resources, the communities have one closure per clan/family; one large no-take area would have disproportionate affect on those compared to the rest of the community. In contrast, communities that have joint ownership can establish one large closure as long as there are other areas available to harvest. Third, historical and trade relationships with neighboring communities can influence regulations by creating the need for occasional harvests to provide fish for feasts. This study further demonstrates the importance of understanding the socioeconomic context of factors such as community governance and levels of dependence for the conservation of marine resources.     

Diving behavior and movements of juvenile hawksbill turtles Eretmochelys imbricata on a Caribbean coral reef

As historically abundant spongivores, hawksbill turtles Eretmochelys imbricata likely played a key ecological role on coral reefs. However, coral reefs are now experiencing global declines and many hawksbill populations are critically reduced. For endangered species, tracking movement has been recognized as fundamental to management. Since movements in marine vertebrates encompass three dimensions, evaluation of diving behavior and range is required to characterize marine turtle habitat. In this study, habitat use of hawksbill turtles on a Caribbean coral reef was elucidated by quantifying diel depth utilization and movements in relation to the boundaries of marine protected areas. Time depth recorders (TDRs) and ultrasonic tags were deployed on 21 Cayman Islands hawksbills, ranging in size from 26.4 to 58.4 cm straight carapace length. Study animals displayed pronounced diel patterns of diurnal activity and nocturnal resting, where diurnal dives were significantly shorter, deeper, and more active. Mean diurnal dive depth (±SD) was 8 ± 5 m, range 2–20 m, mean nocturnal dive depth was 5 ± 5 m, range 1–14 m, and maximum diurnal dive depth was 43 ± 27 m, range 7–91 m. Larger individuals performed significantly longer dives. Body mass was significantly correlated with mean dive depth for nocturnal but not diurnal dives. However, maximum diurnal dive depth was significantly correlated with body mass, suggesting partitioning of vertical habitat by size. Thus, variable dive capacity may reduce intraspecific competition and provide resistance to degradation in shallow habitats. Larger hawksbills may also represent important predators on deep reefs, creating a broad ecological footprint over a range of depths. Communicated by Biology Editor Dr Mark McCormick     

Relationships Between Reef Fish Communities and Remotely Sensed Rugosity Measurements in Biscayne National Park,Florida, USA

The realization that coral reef ecosystem management must occur across multiple spatial scales and habitat types has led scientists and resource managers to seek variables that are easily measured over large areas and correlate well with reef resources. Here we investigate the utility of new technology in airborne laser surveying (NASA Experimental Advanced Airborne Research Lidar (EAARL)) in assessing topographical complexity (rugosity) to predict reef fish community structure on shallow (<10 m deep) patch reefs. Marine portions of Biscayne National Park, Florida, USA, were surveyed remotely using the EAARL, and reef fish populations were visually surveyed on 10 patch reefs at independent, randomly selected stations (n = 10–13 per reef). Rugosity at each station was assessed in situ by divers using the traditional chain-transect method (10-m scale), and remotely using the EAARL submarine topography data at multiple spatial scales (2, 5, and 10 m). The rugosity and biological datasets were analyzed together to elucidate the predictive power of EAARL rugosity in describing the variance in reef fish community variables and to assess the correlation between chain-transect and EAARL rugosity. EAARL rugosity was not well correlated with chain-transect rugosity, or with species richness of fishes (although statistically significant, the amount of variance explained by the model was very low). Variance in reef fish community attributes was better explained in reef-by-reef variability than by physical variables. However, once the reef-by-reef variability was taken into account in a two-way analysis of variance, the importance of rugosity could be seen on individual reefs. Fish species richness and abundance were statistically higher at high rugosity stations compared to medium and low rugosity stations, as predicted by prior ecological research. The EAARL shows promise as an important mapping tool for reef resource managers as they strive to inventory and protect coral reef resources.     

Spatial variation in sea urchins, fish predators, and bioerosion rates on coral reefs of Belize

Although sea urchins are critical for controlling macroalgae on heavily fished coral reefs, high densities threaten reefs, as urchins are also prodigous bioeroders. This study examined urchin population characteristics, bioerosion rates, their fish predators (Labridae), and potential competitors (Scaridae) on unprotected reefs and a reef within a marine protected area (MPA) in the lagoonal regions off Belize. Urchin density (<1 m⁻²) and bioerosion rates (∼0.2 kg CaCO₃ m⁻² year⁻¹) were lowest and members of the Labridae were the highest (∼20 fish 200 m⁻³) within the MPA, while several unprotected reefs had higher (∼18–40 m⁻²) urchin densities, lower Labridae abundances (1–3 fish 200 m⁻³), and bioerosion rates ranging from ∼0.3–2.6 kg CaCO₃ m⁻² year⁻¹. Urchin abundances were inversely related to Labridae (wrasses and hogfish) densities; however, on reef ridges, low algal cover (∼15%), small urchin size (∼14 mm), and low proportion of organic material in urchin guts suggested food limitation. Both top–down (predation) and bottom–up factors (food limitation) likely contribute to the control of urchins, predominantly Echinometra viridis, off Belize, thereby potentially diminishing the negative impacts of bioerosion activities by urchins.     

Reef fish assemblage structure affected by small-scale spacing and size variations of artificial patch reefs

To examine how varying the distance between patch reefs affects reef fish assemblage structure, replicate concrete reef modules (∼ 1 m³ each) were deployed on sand bottom at 8 m depth off Ft. Lauderdale, Florida, USA (26°07N, 80°05W). Modules were positioned at the apices of one of four differently sized equilateral triangles. Triangular configurations had side lengths of: 25 m, 15 m, 5 m, and 0.33 m; each treatment with two replicates. Two additional configurations: (1) a solitary module (Single) and (2) two modules side by side (Double), also with two replicates, were deployed in order to examine the interaction of reef size with fish assemblages. SCUBA divers censused fishes monthly, for 2 years, recording the species present, their abundance and sizes (TL). Fishes were assigned to one of five length categories: < 2 cm, > 2-5 cm, > 5-10 cm, > 10-20 cm, and > 20 cm. In general and excluding the smallest three-module spacing treatment (0.33 m treatment), which may have provided unique treatment-specific refuge, total fish abundance and richness were shown to increase when isolation distance increased. However, there were also species-specific and size class differences in response to isolation distance. The second part of this study indicated varying reef size, by doubling and tripling the number of reef modules, increased total fish abundance and species richness. Nevertheless, fish abundance and species richness did not change by an identical multiplier (e.g., doubling modules ≠ double abundance). These results suggest that scientists and marine managers alike should consider reef size and isolation as habitat attributes capable of altering the structure and dynamics of reef fish assemblages.     

Coral reef crisis in deep and shallow reefs: 30 years of constancy and change in reefs of Curacao and Bonaire

Coral reefs are thought to be in worldwide decline but available data are practically limited to reefs shallower than 25 m. Zooxanthellate coral communities in deep reefs (30–40 m) are relatively unstudied. Our question is: what is happening in deep reefs in terms of coral cover and coral mortality? We compare changes in species composition, coral mortality, and coral cover at Caribbean (Curacao and Bonaire) deep (30–40 m) and shallow reefs (10–20 m) using long-term (1973–2002) data from permanent photo quadrats. About 20 zooxanthellate coral species are common in the deep-reef communities, dominated by Agaricia sp., with coral cover up to 60%. In contrast with shallow reefs, there is no decrease in coral cover or number of coral colonies in deep reefs over the last 30 years. In deep reefs, non-agaricid species are decreasing but agaricid domination will be interrupted by natural catastrophic mortality such as deep coral bleaching and storms. Temperature is a vastly fluctuating variable in the deep-reef environment with extremely low temperatures possibly related to deep-reef bleaching. An erratum to this article can be found at     

Could some coral reefs become sponge reefs as our climate changes?

Coral reefs across the world have been seriously degraded and have a bleak future in response to predicted global warming and ocean acidification (OA). However, this is not the first time that biocalcifying organisms, including corals, have faced the threat of extinction. The end‐Triassic mass extinction (200 million years ago) was the most severe biotic crisis experienced by modern marine invertebrates, which selected against biocalcifiers; this was followed by the proliferation of another invertebrate group, sponges. The duration of this sponge‐dominated period far surpasses that of alternative stable‐ecosystem or phase‐shift states reported on modern day coral reefs and, as such, a shift to sponge‐dominated reefs warrants serious consideration as one future trajectory of coral reefs. We hypothesise that some coral reefs of today may become sponge reefs in the future, as sponges and corals respond differently to changing ocean chemistry and environmental conditions. To support this hypothesis, we discuss: (i) the presence of sponge reefs in the geological record; (ii) reported shifts from coral‐ to sponge‐dominated systems; and (iii) direct and indirect responses of the sponge holobiont and its constituent parts (host and symbionts) to changes in temperature and pH. Based on this evidence, we propose that sponges may be one group to benefit from projected climate change and ocean acidification scenarios, and that increased sponge abundance represents a possible future trajectory for some coral reefs, which would have important implications for overall reef functioning.     

Conservation,management, and restoration of coral reefs

The 8th International Conference on Coelenterate Biology (ICCB 8) was held in Eilat, Israel from December 1st to 5th 2013. The conference included 15 sessions, one of which discussed the latest information on the conservation, management, and restoration of Coelenterata in different parts of the world. A total of 16 oral presentations and 5 posters were presented in this session. Of these 21 papers, 11 were related to conservation issues, 7 described management, and 3 discussed restoration. This session provided insights on the current conservation, management, and restoration of coelenterates in different parts of the world.     

Connectivity,biodiversity conservation and the design of marine reserve networks for coral reefs

Networks of no-take reserves are important for protecting coral reef biodiversity from climate change and other human impacts. Ensuring that reserve populations are connected to each other and non-reserve populations by larval dispersal allows for recovery from disturbance and is a key aspect of resilience. In general, connectivity between reserves should increase as the distance between them decreases. However, enhancing connectivity may often tradeoff against a network’s ability to representatively sample the system’s natural variability. This “representation” objective is typically measured in terms of species richness or diversity of habitats, but has other important elements (e.g., minimizing the risk that multiple reserves will be impacted by catastrophic events). Such representation objectives tend to be better achieved as reserves become more widely spaced. Thus, optimizing the location, size and spacing of reserves requires both an understanding of larval dispersal and explicit consideration of how well the network represents the broader system; indeed the lack of an integrated theory for optimizing tradeoffs between connectivity and representation objectives has inhibited the incorporation of connectivity into reserve selection algorithms. This article addresses these issues by (1) updating general recommendations for the location, size and spacing of reserves based on emerging data on larval dispersal in corals and reef fishes, and on considerations for maintaining genetic diversity; (2) using a spatial analysis of the Great Barrier Reef Marine Park to examine potential tradeoffs between connectivity and representation of biodiversity and (3) describing a framework for incorporating environmental fluctuations into the conceptualization of the tradeoff between connectivity and representation, and that expresses both in a common, demographically meaningful currency, thus making optimization possible.     

Ecosystem‐based management of coral reefs under climate change

Coral reefs provide food and livelihoods for hundreds of millions of people as well as harbour some of the highest regions of biodiversity in the ocean. However, overexploitation, land‐use change and other local anthropogenic threats to coral reefs have left many degraded. Additionally, coral reefs are faced with the dual emerging threats of ocean warming and acidification due to rising CO₂ emissions, with dire predictions that they will not survive the century. This review evaluates the impacts of climate change on coral reef organisms, communities and ecosystems, focusing on the interactions between climate change factors and local anthropogenic stressors. It then explores the shortcomings of existing management and the move towards ecosystem‐based management and resilience thinking, before highlighting the need for climate change‐ready marine protected areas (MPAs), reduction in local anthropogenic stressors, novel approaches such as human‐assisted evolution and the importance of sustainable socialecological systems. It concludes that designation of climate change‐ready MPAs, integrated with other management strategies involving stakeholders and participation at multiple scales such as marine spatial planning, will be required to maximise coral reef resilience under climate change. However, efforts to reduce carbon emissions are critical if the long‐term efficacy of local management actions is to be maintained and coral reefs are to survive.     

Spatial and size-frequency distribution of <Emphasis Type="Italic">Acropora</Emphasis> (Cnidaria: Scleractinia) species in Chinchorro Bank,Mexican Caribbean: implications for management

The Mexican Government decreed Chinchorro Bank reef as a Biosphere Reserve in 1996. The aim of this study was to evaluate the spatial and size-frequency distribution of Acropora spp. in order to provide further knowledge and tools to enhance management. A field survey was conducted, within six regions, to locate and measure Acropora patches in the reef lagoon. Density, colony size and living tissue cover of Acropora colonies were evaluated using the line-intercept transect technique, combining direct observations and video transects. The results showed that Acropora spp. was preferentially distributed in the southern regions; where cover and density were high. Based on these results and considering that Acropora spp. produces landscape heterogeneity, which in turn generates shelter for other species, including some of considerable economic importance, then at least the South East region should be considered as a key area for Acropora species conservation, and should be included in the Chinchorro Bank management plan.     

Flattening of Caribbean coral reefs: region-wide declines in architectural complexity

Coral reefs are rich in biodiversity, in large part because their highly complex architecture provides shelter and resources for a wide range of organisms. Recent rapid declines in hard coral cover have occurred across the Caribbean region, but the concomitant consequences for reef architecture have not been quantified on a large scale to date. We provide, to our knowledge, the first region-wide analysis of changes in reef architectural complexity, using nearly 500 surveys across 200 reefs, between 1969 and 2008. The architectural complexity of Caribbean reefs has declined nonlinearly with the near disappearance of the most complex reefs over the last 40 years. The flattening of Caribbean reefs was apparent by the early 1980s, followed by a period of stasis between 1985 and 1998 and then a resumption of the decline in complexity to the present. Rates of loss are similar on shallow (<6 m), mid-water (6–20 m) and deep (>20 m) reefs and are consistent across all five subregions. The temporal pattern of declining architecture coincides with key events in recent Caribbean ecological history: the loss of structurally complex Acropora corals, the mass mortality of the grazing urchin Diadema antillarum and the 1998 El Nino Southern Oscillation-induced worldwide coral bleaching event. The consistently low estimates of current architectural complexity suggest regional-scale degradation and homogenization of reef structure. The widespread loss of architectural complexity is likely to have serious consequences for reef biodiversity, ecosystem functioning and associated environmental services.