Otolith growth of Springer's demoiselle, Chrysiptera springeri (Pomacentridae, Allen & Lubbock), on a protected and non-protected coral reef

The structural complexity of coral reefs is important for their function as shelter and feeding habitats for coral reef fishes, but physical disturbance by human activities often reduce complexity of the reefs by selectively destroying fragile and more complex coral species. The damselfish Springer's demoiselle Chrysiptera springeri primarily utilize complex coral heads for shelter and are hence vulnerable to human disturbance. In order to evaluate the potential effect of habitat degradation on juvenile fish growth, coral reef cover, fish age at settling and otolith growth, juvenile Springer's demoiselle was investigated on a protected and non‐protected coral reef in Darvel Bay, Borneo. The protected reef had higher coverage of complex branching corals and exhibited a more complex 3‐dimensional structure than the non‐protected reef. Springer's demoiselle settled at the same age on non‐protected and protected reefs. The growth rates of the otoliths from Springer's demoiselle were similar during the pre‐settlement period on the two reefs (manova , P > 0.05), but from age 20 to 48 days (post‐settlement period) the otolith growth rate of juveniles on the non‐protected reef was reduced compared to those from the protected reef (manova , P = 0.017). However, the differences in the otolith size, and by inference, fish size, after 48 days were small. The small effect of habitat degradation on growth is likely related to the fact that the Springer's demoiselles collected on the non‐protected reef were associated with the few remaining complex coral heads. Increased foraging‐predation tradeoffs on the non‐protected reef may decrease food intake and growth of juvenile Springer's demoiselle, but the main effect of habitat degradation on their abundance is likely to be related to lack of suitable shelter, and consequently reduced carrying capacity, on disturbed reefs.     

Relationships between habitat structure and fish communities on coral

The influence of habitat structure on reef-fish communities at Bar Reef Marine Sanctuary, Sri Lanka, was investigated. The relationship between habitat characteristics and the distribution and abundance of 135 species of fishes was examined on two reef types: coral and sandstone reefs. Results suggested that the reef-fish communities were strongly influenced by various aspects of reef structure. However, relationships between habitat variables and fish communities structure, varied between the two reef types. Fish species diversity was correlated with a number of habitat variables on the sandstone reefs, although structural complexity seemed to play the dominant role. There were no correlations between habitat structure and fish diversity on the coral reefs. Total abundance was not related to any one habitat parameter on either reef type. However, abundances of some species, families and trophic groups were correlated with habitat features. These specific correlations were commonly related to food or shelter availability. For example, coral feeders were correlated with live coral cover, and pomacentrid species, which used branching corals for protection, showed a significant relationship with the density of Acropora colonies. This shows that a summary statistic such as total abundance may hide important information. Effects of habitat structure on the distribution patterns of the fish communities was further investigated using multi-dimensional scaling ordination (MDS) and the RELATE-procedure. With the MDS ordinations for both habitat and fish-community composition it was possible to show that the multivariate pattern between the two ecological components was clearly correlated.     

The effect of coral morphology on shelter selection by coral reef fishes

While the loss of structural complexity causes declines in coral reef fish diversity, the processes leading to this decline are largely unexplained. To explore the role of coral morphology in providing shelter for fishes, tabular, branching and massive corals were filmed with video cameras and their usage by large reef fishes compared. Tabular corals were utilised more than the other two morphologies, with at least triple the abundance, biomass and residence times of large fishes. The preference of coral reef fishes for specific structural traits of tabular corals was also examined using artificial structural units. This experimental component showed that large reef fishes preferred opaque rather than translucent canopies. It appears that large fishes cue to tabular corals because of the concealment and/or shade provided. It is suggested that a loss of tabular corals as a result of climate change would have significant ecological impacts for the coral reef fishes that use these structures for shelter.     

The influence of coral reef benthic condition on associated fish assemblages

Accumulative disturbances can erode a coral reef's resilience, often leading to replacement of scleractinian corals by macroalgae or other non-coral organisms. These degraded reef systems have been mostly described based on changes in the composition of the reef benthos, and there is little understanding of how such changes are influenced by, and in turn influence, other components of the reef ecosystem. This study investigated the spatial variation in benthic communities on fringing reefs around the inner Seychelles islands. Specifically, relationships between benthic composition and the underlying substrata, as well as the associated fish assemblages were assessed. High variability in benthic composition was found among reefs, with a gradient from high coral cover (up to 58%) and high structural complexity to high macroalgae cover (up to 95%) and low structural complexity at the extremes. This gradient was associated with declining species richness of fishes, reduced diversity of fish functional groups, and lower abundance of corallivorous fishes. There were no reciprocal increases in herbivorous fish abundances, and relationships with other fish functional groups and total fish abundance were weak. Reefs grouping at the extremes of complex coral habitats or low-complexity macroalgal habitats displayed markedly different fish communities, with only two species of benthic invertebrate feeding fishes in greater abundance in the macroalgal habitat. These results have negative implications for the continuation of many coral reef ecosystem processes and services if more reefs shift to extreme degraded conditions dominated by macroalgae.     

Non‐reef habitats in a tropical seascape affect density and biomass of fishes on coral reefs

Nonreef habitats such as mangroves, seagrass, and macroalgal beds are important for foraging, spawning, and as nursery habitat for some coral reef fishes. The spatial configuration of nonreef habitats adjacent to coral reefs can therefore have a substantial influence on the distribution and composition of reef fish. We investigate how different habitats in a tropical seascape in the Philippines influence the presence, density, and biomass of coral reef fishes to understand the relative importance of different habitats across various spatial scales. A detailed seascape map generated from satellite imagery was combined with field surveys of fish and benthic habitat on coral reefs. We then compared the relative importance of local reef (within coral reef) and adjacent habitat (habitats in the surrounding seascape) variables for coral reef fishes. Overall, adjacent habitat variables were as important as local reef variables in explaining reef fish density and biomass, despite being fewer in number in final models. For adult and juvenile wrasses (Labridae), and juveniles of some parrotfish taxa (Chlorurus), adjacent habitat was more important in explaining fish density and biomass. Notably, wrasses were positively influenced by the amount of sand and macroalgae in the adjacent seascape. Adjacent habitat metrics with the highest relative importance were sand (positive), macroalgae (positive), and mangrove habitats (negative), and fish responses to these metrics were consistent across fish groups evaluated. The 500‐m spatial scale was selected most often in models for seascape variables. Local coral reef variables with the greatest importance were percent cover of live coral (positive), sand (negative), and macroalgae (mixed). Incorporating spatial metrics that describe the surrounding seascape will capture more holistic patterns of fish–habitat relationships on reefs. This is important in regions where protection of reef fish habitat is an integral part of fisheries management but where protection of nonreef habitats is often overlooked.     

Maintenance of fish diversity on disturbed coral reefs

Habitat perturbations play a major role in shaping community structure; however, the elements of disturbance-related habitat change that affect diversity are not always apparent. This study examined the effects of habitat disturbances on species richness of coral reef fish assemblages using annual surveys of habitat and 210 fish species from 10 reefs on the Great Barrier Reef (GBR). Over a period of 11 years, major disturbances, including localised outbreaks of crown-of-thorns sea star (Acanthaster planci), severe storms or coral bleaching, resulted in coral decline of 46–96% in all the 10 reefs. Despite declines in coral cover, structural complexity of the reef framework was retained on five and species richness of coral reef fishes maintained on nine of the disturbed reefs. Extensive loss of coral resulted in localised declines of highly specialised coral-dependent species, but this loss of diversity was more than compensated for by increases in the number of species that feed on the epilithic algal matrix (EAM). A unimodal relationship between areal coral cover and species richness indicated species richness was greatest at approximately 20% coral cover declining by 3–4 species (6–8% of average richness) at higher and lower coral cover. Results revealed that declines in coral cover on reefs may have limited short-term impact on the diversity of coral reef fishes, though there may be fundamental changes in the community structure of fishes.     

Distributions and habitat associations of the bridled monocle bream Scolopsis bilineatus (Nemipteridae): a demographic approach

This study focussed on the demography and ecology of Scolopsis bilineatus at three locations on the Great Barrier Reef: the Lizard Island Group, Orpheus Island and One Tree Island. Scolopsis bilineatus lived for up to 16 years and had four distinct life‐history stages, which varied in their distribution patterns, habitat use and reproductive behaviour. Pre‐maturational sex change occurred whereby all males were derived from immature females, and males grew faster and larger than females. Small females and larger males generally formed pairs, which influenced their spatial distributions at small scales. Distributions of S. bilineatus were influenced by depth and exposure within reefs, particularly for juveniles, and most fish were found in shallow, sheltered habitats. Abundance was influenced by benthic cover, and was higher in areas of high coral cover and low where algae were abundant. Habitat associations were stronger at the microhabitat scale, and shelter sites were important for adults. Ontogenetic changes in microhabitat associations were found: juveniles occupied sand and rubble, and adults occupied shelters such as caves and overhangs. Adults showed site fidelity for shelter sites over a period of 4 days and returned to specific shelter sites repeatedly. These findings illustrate the importance of understanding the spatial ecology and habitat use of coral reef fishes, particularly with reference to size‐based changes within species.     

The importance of structural complexity in coral reef ecosystems

The importance of structural complexity in coral reefs has come to the fore with the global degradation of reef condition; however, the limited scale and replication of many studies have restricted our understanding of the role of complexity in the ecosystem. We qualitatively and quantitatively (where sufficient standardised data were available) assess the literature regarding the role of structural complexity in coral reef ecosystems. A rapidly increasing number of publications have studied the role of complexity in reef ecosystems over the past four decades, with a concomitant increase in the diversity of methods used to quantify structure. Quantitative analyses of existing data indicate a strong negative relationship between structural complexity and algal cover, which may reflect the important role complexity plays in enhancing herbivory by reef fishes. The cover of total live coral and branching coral was positively correlated with structural complexity. These habitat attributes may be creating much of the structure, resulting in a collinear relationship; however, there is also evidence of enhanced coral recovery from disturbances where structural complexity is high. Urchin densities were negatively correlated with structural complexity; a relationship that may be driven by urchins eroding reef structure or by their gregarious behaviour when in open space. There was a strong positive relationship between structural complexity and fish density and biomass, likely mediated through density-dependent competition and refuge from predation. More variable responses were found when assessing individual fish families, with all families examined displaying a positive relationship to structural complexity, but only half of these relationships were significant. Although only corroborated with qualitative data, structural complexity also seems to have a positive effect on two ecosystem services: tourism and shoreline protection. Clearly, structural complexity is an integral component of coral reef ecosystems, and it should be incorporated into monitoring programs and management objectives.     

Ecological condition of coral reef assemblages in the Cuban Archipelago

The condition of coral reefs in the Cuban Archipelago is poorly known. We aimed to analyse coral assemblages across 199 reef sites belonging to 12 localities. Crest and fore reefs were assessed using six metrics: species richness, density, coral cover, mortality, coral size and reef complexity. The condition of reefs varied across the archipelago from healthy to depleted reefs. The localities with best scores were Cienfuegos, Bahía de Cochinos and Cazones. These reefs have values of living coral cover (>20%) and complexity (>50?cm) similar to the best preserved Caribbean reefs. However, the majority of crest biotopes suffered important deterioration with old mortality of Acropora palmata populations and moderate coral cover (15%); although crest reefs still maintained their structural complexity. Despite moderate levels of coral cover in fore reefs (18%), their condition was alarming because 25% of the sites had cover below the recovery threshold of 10%, accumulated mortality and structural flattening. Compared with the 1980s, the species richness was roughly the same (42) for crest and fore reefs, although dominance has changed to widespread tolerant species. Coral reef assemblages varied at local and regional scales in similar magnitude, suggesting the combined effects of natural and anthropogenic drivers.     

Predation Risk,Resource Quality,and Reef Structural Complexity Shape Territoriality in a Coral Reef Herbivore

For many species securing territories is important for feeding and reproduction. Factors such as competition, habitat availability, and male characteristics can influence an individual’s ability to establish and maintain a territory. The risk of predation can have an important influence on feeding and reproduction; however, few have studied its effect on territoriality. We investigated territoriality in a haremic, polygynous species of coral reef herbivore, Sparisoma aurofrenatum (redband parrotfish), across eight reefs in the Florida Keys National Marine Sanctuary that were either protected or unprotected from fishing of piscivorous fishes. We examined how territory size and quality varied with reef protection status, competition, predation risk, and male size. We then determined how territory size and quality influenced harem size and female size to understand the effect of territoriality on reproductive potential. We found that protected reefs trended towards having more large predatory fishes and that territories there were smaller but had greater algal nutritional quality relative to unprotected reefs. Our data suggest that even though males in protected sites have smaller territories, which support fewer females, they may improve their reproductive potential by choosing nutritionally rich areas, which support larger females. Thus, reef protection appears to shape the trade-off that herbivorous fishes make between territory size and quality. Furthermore, we provide evidence that males in unprotected sites, which are generally less complex than protected sites, choose territories with higher structural complexity, suggesting the importance of this type of habitat for feeding and reproduction in S. aurofrenatum. Our work argues that the loss of corals and the resulting decline in structural complexity, as well as management efforts to protect reefs, could alter the territory dynamics and reproductive potential of important herbivorous fish species.     

Climate warming, marine protected areas and the ocean-scale integrity of coral reef ecosystems

Coral reefs have emerged as one of the ecosystems most vulnerable to climate variation and change. While the contribution of a warming climate to the loss of live coral cover has been well documented across large spatial and temporal scales, the associated effects on fish have not. Here, we respond to recent and repeated calls to assess the importance of local management in conserving coral reefs in the context of global climate change. Such information is important, as coral reef fish assemblages are the most species dense vertebrate communities on earth, contributing critical ecosystem functions and providing crucial ecosystem services to human societies in tropical countries. Our assessment of the impacts of the 1998 mass bleaching event on coral cover, reef structural complexity, and reef associated fishes spans 7 countries, 66 sites and 26 degrees of latitude in the Indian Ocean. Using Bayesian meta-analysis we show that changes in the size structure, diversity and trophic composition of the reef fish community have followed coral declines. Although the ocean scale integrity of these coral reef ecosystems has been lost, it is positive to see the effects are spatially variable at multiple scales, with impacts and vulnerability affected by geography but not management regime. Existing no-take marine protected areas still support high biomass of fish, however they had no positive affect on the ecosystem response to large-scale disturbance. This suggests a need for future conservation and management efforts to identify and protect regional refugia, which should be integrated into existing management frameworks and combined with policies to improve system-wide resilience to climate variation and change.     

Competition for shelter in a high-diversity system: structure use by large reef fishes

Competition among large reef fishes for shelter beneath tabular structures provides a rare opportunity to study competition in a species-rich environment. The system permits a detailed study of localised competition with major implications for coral reefs with respect to human impacts including climate change. Using underwater video cameras, this study examined competition among 30 species of large reef fishes (from nine families) for access to shelter provided by 26 tabular structures, which may be the highest reported diversity of vertebrates competing for a single resource. Mean concentrations of fishes under tabular structures were also among the highest biomass recorded on reefs (4.71 kg m⁻²). A generated dominance hierarchy for the occupation of shelter appeared to be primarily driven by the size of fishes. In contrast to previous studies, fishes higher in the hierarchy tended to exhibit the lowest levels of aggression. However, size difference between fishes was found to be strongly negatively correlated with the proportion of aggressive interactions (R ² = 0.971, P < 0.0001). The strong competition for the shade provided by these corals highlights concerns about future shifts in the structure of large reef fish communities as corals are lost. This is particularly concerning given the critical functional roles played by certain species of large reef fishes that utilise tabular structure for shelter and which occupy the lower ranks of the dominance hierarchy.

     

Multiscale habitat associations of detrivorous blennies (Blenniidae: Salariini)

The distribution and habitat associations of detrivorous blennies on a tropical coral reef were investigated at several spatial scales and compared with other fish that feed on the epilithic algal matrix to assess density and biomass contributions of small detrivorous fishes to these assemblages. At broad spatial scales total blenny abundance and biomass were highest on the tops of reefs exposed to prevailing winds. On the finer scale of microhabitat use, all species showed a preference for non-living corals, although the type of coral utilised differed between species. The high abundance of blennies on reef tops and non-living corals may be partially related to the quality and availability of detritus in these habitats. Comparisons of total blenny abundance and biomass with other territorial detrivores found that blennies accounted for approximately 60% of this functional group's density and 21% of their biomass on exposed reef tops. Overall, territorial detrivores were found to constitute approximately 37% of the density and 26% of the biomass of the detrivorous/herbivorous fish assemblage on exposed reef tops. Small detrivorous fish therefore represent a substantial proportion of fish assemblages that feed on epilithic algae and associated detritus on coral reefs.     

Size-structural shifts reveal intensity of exploitation in coral reef fisheries

Fisheries exploitation represents a considerable threat to coral reef fish resources because even modest levels of extraction can alter ecological dynamics via shifts of stock size, species composition, and size-structure of the fish assemblage. Although species occupying higher trophic groups are known to suffer the majority of exploitative effects, changes in composition among lower trophic groups may be major, though are not frequently explored. Using size-based biomass spectrum analysis, we investigate the effects of fishing on the size-structure of coral reef fish assemblages spanning four geopolitical regions and determine if patterns of exploitation vary across trophic groups. Our analyses reveal striking evidence for the variety of effects fisheries exploitation can have on coral reef fish assemblages. When examining biomass spectra across the entire fish assemblage we found consistent evidence of size-specific exploitation, in which large-bodied individuals experience disproportionate reductions. The pattern was paralleled by and likely driven by, strongly size-specific reductions among top predators. In contrast, evidence of exploitation patterns was variable among lower trophic groups, in many cases including evidence of reductions across all size classes. The breadth of size classes and trophic groups that showed evidence of exploitation related positively to local human population density and diversity of fishing methods employed. Our findings highlight the complexity of coral reef fisheries and that the effects of exploitation on coral reefs can be realized throughout the entire fish assemblage, across multiple trophic groups and not solely restricted to large-bodied top-predators. Size-specific changes among fishes of lower trophic groups likely lead to altered ecological functioning of heavily exploited coral reefs. Together these findings reinforce the value of taking a multi-trophic group approach to monitoring and managing coral reef fisheries.     

The effect of natural variation in substrate architecture on the survival of juvenile bicolor damselfish

Coral reefs offer settling fish larvae a spatial mosaic of microhabitats that differ not only in structural complexity but also in the abundance and diversity of predators. This paper provides evidence that interactions between predators and prey are causally linked to variation in relevant architectural characteristics of natural substrates. Juvenile bicolor damselfish, Stegastes partitus, experienced greater mortality on Montastrea annularis boulder coral than on piles of Porites porites coral rubble. This pattern was consistent on both back and fore reef habitats. Architectural differences, variable encounter rates with predators, and access to different food sources all contributed to higher mortality. Spatial and structural differences among refuges provided by the two substrates were most important in affecting survival. Porites rubble contained almost three times the number of crevices (39.9 vs. 14.3 crevices m^–2AM²), had a much smaller mean crevice size (15.9 vs. 48.5 cm²), and had a more complex internal structure than Montastrea coral. Survival of juvenile fish living on Montastrea coral was positively correlated with crevice density (r² = 0.41) whereas survival of fish living on Porites rubble was negatively correlated with crevice size (r² = 0.43). These patterns were evident on fore reef habitats only, whereas on back reef habitats no clear patterns emerged. The effect of these natural differences in architecture on mortality rates of juvenile S. partitus was experimentally tested in the field using a combination of natural and artificial substrates. As the number of large shelters was increased I found that the density of potential predators increased but the survival rates of juvenile bicolor damselfish declined. These results highlight the importance of structural architecture among common reef substrates in affecting predator-prey relationships and in determining the survivorship and small-scale distribution patterns of juvenile reef fish.     

The effect of microhabitat patch size on settlement differs among co-occurring coral reef fishes

Over small spatial scales, coral reefs represent a mosaic of suitable settlement microhabitat patches of varying size for late-stage larval reef fishes. Few studies have specifically examined how variation in patch size influences density of recently settled coral reef fishes (recruits). Using standardized units of coral rubble settlement substrate deployed on sandy bottom, we monitored the concurrent settlement of three reef fish taxa onto differently sized patches (0.28–1.68 m²) at 5-d intervals during a lunar settlement peak. We found marked differences among taxa in how recruit density scaled with patch size. Recruit density of a damselfish and a parrotfish decreased and increased, respectively, with the increase in patch size, while that of a wrasse was similar among patch sizes. Our results highlight the importance of the interaction between taxon-specific settlement behaviour and patch size in establishing initial spatial differences in density within and among coral reef fish taxa in a heterogeneous landscape.

     

Retention of Habitat Complexity Minimizes Disassembly of Reef Fish Communities following Disturbance: A Large-Scale Natural Experiment

High biodiversity ecosystems are commonly associated with complex habitats. Coral reefs are highly diverse ecosystems, but are under increasing pressure from numerous stressors, many of which reduce live coral cover and habitat complexity with concomitant effects on other organisms such as reef fishes. While previous studies have highlighted the importance of habitat complexity in structuring reef fish communities, they employed gradient or meta-analyses which lacked a controlled experimental design over broad spatial scales to explicitly separate the influence of live coral cover from overall habitat complexity. Here a natural experiment using a long term (20 year), spatially extensive (∼115,000 kms²) dataset from the Great Barrier Reef revealed the fundamental importance of overall habitat complexity for reef fishes. Reductions of both live coral cover and habitat complexity had substantial impacts on fish communities compared to relatively minor impacts after major reductions in coral cover but not habitat complexity. Where habitat complexity was substantially reduced, species abundances broadly declined and a far greater number of fish species were locally extirpated, including economically important fishes. This resulted in decreased species richness and a loss of diversity within functional groups. Our results suggest that the retention of habitat complexity following disturbances can ameliorate the impacts of coral declines on reef fishes, so preserving their capacity to perform important functional roles essential to reef resilience. These results add to a growing body of evidence about the importance of habitat complexity for reef fishes, and represent the first large-scale examination of this question on the Great Barrier Reef.     

Changes in Coral Reef Communities and an Associated Reef Fish Species, <Emphasis Type="Italic">Cephalopholis cruentata</Emphasis> (Lacépède), After 30 years on Curaçao (Netherlands Antilles)

Using the same methodology and identical sites, we repeat a study dating from 1973 and quantify cover of hard coral species, soft corals, sponges, hard substratum and soft substratum, and density of a commercially important reef fish species, the graysby Cephalopholis cruentata, along a depth-gradient of 3–36 m on the coral reefs of Curaçao. The objective was to determine the multi-decade change in benthic coral reef cover and structural complexity, and their effect on densities of an associated reef fish species. Total hard coral cover decreased on average from 52% in 1973 to 22% in 2003, representing a relative decline of 58%. During this time span, the cover of hard substratum increased considerably (from 11 to 58%), as did that of soft corals (from 0.1 to 2.2%), whereas the cover of sponges showed no significant change. Relative decline of hard coral cover and of reef complexity was greatest in shallow waters (near the coast), which is indicative of a combination of anthropogenic influences from shore and recent storm damage. Cover of main reef builder coral species (Agaricia spp., Siderastrea siderea, Montastrea annularis) decreased more than that of other species, and resulted in a significant decrease in reef complexity. Although density of C. cruentata was highly correlated to cover of Montastrea and Agaricia in 1973, the loss of coral cover did not show any effect on the total density of C. cruentata in 2003. However, C. cruentata showed a clear shift in density distribution from shallow water in 1973 to deep water in 2003. It can be concluded that the reefs of Curaçao have degraded considerably in the last three decades, but that this has had no major effect on the population size of one commercially important coral-associated fish species.     

Interactions between coral restoration and fish assemblages: implications for reef management

Corals create complex reef structures that provide both habitat and food for many fish species. Because of numerous natural and anthropogenic threats, many coral reefs are currently being degraded, endangering the fish assemblages they support. Coral reef restoration, an active ecological management tool, may help reverse some of the current trends in reef degradation through the transplantation of stony corals. Although restoration techniques have been extensively reviewed in relation to coral survival, our understanding of the effects of adding live coral cover and complexity on fishes is in its infancy with a lack of scientifically validated research. This study reviews the limited data on reef restoration and fish assemblages, and complements this with the more extensive understanding of complex interactions between natural reefs and fishes and how this might inform restoration efforts. It also discusses which key fish species or functional groups may promote, facilitate or inhibit restoration efforts and, in turn, how restoration efforts can be optimised to enhance coral fish assemblages. By highlighting critical knowledge gaps in relation to fishes and restoration interactions, the study aims to stimulate research into the role of reef fishes in restoration projects. A greater understanding of the functional roles of reef fishes would also help inform whether restoration projects can return fish assemblages to their natural compositions or whether alternative species compositions develop, and over what timeframe. Although alleviation of local and global reef stressors remains a priority, reef restoration is an important tool; an increased understanding of the interactions between replanted corals and the fishes they support is critical for ensuring its success for people and nature.