End of the chain? Rugosity and fine-scale bathymetry from existing underwater digital imagery using structure-from-motion (SfM) technology

The rugosity or complexity of the seafloor has been shown to be an important ecological parameter for fish, algae, and corals. Historically, rugosity has been measured either using simple and subjective manual methods such as ‘chain-and-tape’ or complicated and expensive geophysical methods. Here, we demonstrate the application of structure-from-motion (SfM) photogrammetry to generate high-resolution, three-dimensional bathymetric models of a fringing reef from existing underwater video collected to characterize the seafloor. SfM techniques are capable of achieving spatial resolution that can be orders of magnitude greater than large-scale lidar and sonar mapping of coral reef ecosystems. The resulting data provide finer-scale measurements of bathymetry and rugosity that are more applicable to ecological studies of coral reefs than provided by the more expensive and time-consuming geophysical methods. Utilizing SfM techniques for characterizing the benthic habitat proved to be more effective and quantitatively powerful than conventional methods and thus might portend the end of the ‘chain-and-tape’ method for measuring benthic complexity.     

Do coral-reef fish faunas have a distinctive taxonomic structure?

Do the highly diverse fish faunas that associate with coral reefs have distinguishing taxonomic and ecological characteristics, as proposed by Choat and Bellwood (1991) and Bellwood (1996)? Does a 50?my old (Eocene) fossil fish fauna from Italy represent a coral-reef fish assemblage that provides unique information about the evolution of such assemblages, as claimed by Bellwood (1996)? I compared the structure of the reef fish faunas of adjacent tropical regions rich and poor in coral reefs, in both America and Polynesia, and found that they exhibit no substantive differences in relative species richness among families of typical “coral-reef” fishes. While coral-rich regions have larger reef fish faunas, a variety of factors probably contribute to such differences. Thus coral-reef fish faunas may lack a distinctive taxonomic structure. A similar comparative approach would be useful for assessing whether assemblages of fishes on coral reefs have distinctive ecological characteristics. Based on patterns of habitat use by modern tropical shorefishes, the Italian Eocene fauna includes few definite reef fishes, and may well consist primarily of non-reef fishes preserved in a non-reef habitat. Until we know more about the environment in which those fossils were preserved, that fauna can contribute little to understanding how coral reef fish assemblages have evolved.     

Error,Power, and Blind Sentinels: The Statistics of Seagrass Monitoring

We derive statistical properties of standard methods for monitoring of habitat cover worldwide, and criticize them in the context of mandated seagrass monitoring programs, as exemplified by Posidonia oceanica in the Mediterranean Sea. We report the novel result that cartographic methods with non-trivial classification errors are generally incapable of reliably detecting habitat cover losses less than about 30 to 50%, and the field labor required to increase their precision can be orders of magnitude higher than that required to estimate habitat loss directly in a field campaign. We derive a universal utility threshold of classification error in habitat maps that represents the minimum habitat map accuracy above which direct methods are superior. Widespread government reliance on blind-sentinel methods for monitoring seafloor can obscure the gradual and currently ongoing losses of benthic resources until the time has long passed for meaningful management intervention. We find two classes of methods with very high statistical power for detecting small habitat cover losses: 1) fixed-plot direct methods, which are over 100 times as efficient as direct random-plot methods in a variable habitat mosaic; and 2) remote methods with very low classification error such as geospatial underwater videography, which is an emerging, low-cost, non-destructive method for documenting small changes at millimeter visual resolution. General adoption of these methods and their further development will require a fundamental cultural change in conservation and management bodies towards the recognition and promotion of requirements of minimal statistical power and precision in the development of international goals for monitoring these valuable resources and the ecological services they provide.     

Acoustic mapping using a multibeam echosounder reveals cold-water coral reefs and surrounding habitats

Multibeam surveys were carried out in four areas to the west of Scotland where the coral Lophelia pertusa had previously been recorded. Distinctive seabed mounds were found in one area; video images from the mounds showed coral reef formation, and grab samples recovered L. pertusa reef framework and rubble. Skeleton samples were dated to 3,800 years BP. Grab samples contained 123 species of fauna. The reef structures, termed the Mingulay Reef Complex, were identified as topographic mound-like structures from the bathymetric data and were also visible on the backscatter images. The location of the reefs coincides with Atlantic bottom waters, close to a primary productivity centre and mixing zone, in an area where currents are likely to be accelerated by rocky seafloor ridges. This study shows that multibeam echosounders are powerful tools to locate and map deep-water coral reefs irrespective of water depth. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Reef fish assemblages in north-western Sri Lanka: distribution patterns and influences of fishing practises

The distribution and abundance of reef fishes in relation to habitat structure were studied within Bar Reef Marine Sanctuary (BRMS) and on an adjacent reef, disturbed by destructive fishing techniques, in north-western Sri Lanka, by visually censusing 135 species groups using fifty metre belt-transects. Two types of continental shelf patch-reefs are found in the study area: coral reefs and sandstone reefs, which are divided into distinct habitats, four for the coral reef (shallow reef flat, shallow patch reef, deep reef flat and Porites domes) and two for the sandstone reef (structured sandstone-reef and flat sandstone-reef). Fish assemblages varied in structure between reef types and among habitats within reef types. Functional aspects of habitat structure and composition, such as available food and shelter, seemed to be important factors influencing distribution patterns. The strongest separation in the organisation of fish assemblages in BRMS was between reef types: 19% of all species were confined to the coral-reef patches while 22% were restricted to the sandstone reef patches and 59% were represented on both reef types. In terms of distribution among habitats, 21% of all species were restricted to one habitat while only 1.5% were present in all. The highest density of fish was in the coral reef habitats while highest species diversity was found in the most structurally complex habitat: the structured sandstone-reef. This habitat also had the highest proportion of species with restricted distribution. Planktivores were the most abundant trophic group in BRMS, and the species composition of the group varied among habitats. The comparison of the disturbed reef with BRMS suggested that habitat alteration caused by destructive fishing methods has strongly influenced the fish community. Within the fished area the structure of the fish assemblages was more heterogeneous, fish abundance was lower by an order of magnitude and species numbers were lower than in BRMS.     

Predicting spatially explicit coral reef fish abundance, richness and Shannon–Weaver index from habitat characteristics

The assessment of biodiversity in coral reefs requires the application of geographic information systems (GIS), remote sensing and analytical tools in order to make cost-effective spatially explicit predictions of biodiversity over large geographic areas. Here we present a spatially explicit prediction for coral reef fish diversity index, as well as habitat classification according to reef fish diversity index values in Chinchorro Bank Biosphere Reserve, one of the most important plain/atoll type reef systems in the Caribbean. We have used extensive ecological data on depth, fish and habitat characteristics to perform such prediction. Fish species assemblages and different biotic variables of benthic organisms were characterized using visual censuses and video-transects, respectively at 119 sampling stations. The information was integrated in a GIS, along with satellite imagery (LANSDAT 7 ETM+) and a digital bathymetric model. From the recorded data and a hierarchical classification procedure, we obtained nine different classes of habitats. We used a generalized regression analysis and spatial prediction methodology to create predictive maps (GIS layers) of the different reef benthic components, and a second modeling run produced predictive maps of coral reef fish diversity index. Predictive accuracy of the diversity index map presented a good correlation coefficient (r = 0.87), with maximum diversity index values en reefscapes composed of aggregation of coral colonies with seagrass beds. The implementation of our application was successful for the prediction of fish diversity hot spots and surrogate habitats.     

Determining the Extent and Characterizing Coral Reef Habitats of the Northern Latitudes of the Florida Reef Tract (Martin County)

Climate change has recently been implicated in poleward shifts of many tropical species including corals; thus attention focused on higher-latitude coral communities is warranted to investigate possible range expansions and ecosystem shifts due to global warming. As the northern extension of the Florida Reef Tract (FRT), the third-largest barrier reef ecosystem in the world, southeast Florida (25–27° N latitude) is a prime region to study such effects. Most of the shallow-water FRT benthic habitats have been mapped, however minimal data and limited knowledge exist about the coral reef communities of its northernmost reaches off Martin County. First benthic habitat mapping was conducted using newly acquired high resolution LIDAR bathymetry and aerial photography where possible to map the spatial extent of coral reef habitats. Quantitative data were collected to characterize benthic cover and stony coral demographics and a comprehensive accuracy assessment was performed. The data were then analyzed in a habitat biogeography context to determine if a new coral reef ecosystem region designation was warranted. Of the 374 km² seafloor mapped, 95.2% was Sand, 4.1% was Coral Reef and Colonized Pavement, and 0.7% was Other Delineations. Map accuracy assessment yielded an overall accuracy of 94.9% once adjusted for known map marginal proportions. Cluster analysis of cross-shelf habitat type and widths indicated that the benthic habitats were different than those further south and warranted designation of a new coral reef ecosystem region. Unlike the FRT further south, coral communities were dominated by cold-water tolerant species and LIDAR morphology indicated no evidence of historic reef growth during warmer climates. Present-day hydrographic conditions may be inhibiting poleward expansion of coral communities along Florida. This study provides new information on the benthic community composition of the northern FRT, serving as a baseline for future community shift and range expansion investigations.     

The importance of physical habitat assessment for evaluating river health

1. Physical habitat is the living space of instream biota; it is a spatially and temporally dynamic entity determined by the interaction of the structural features of the channel and the hydrological regime. 2. This paper reviews the need for physical habitat assessment and the range of physical habitat assessment methods that have been developed in recent years. These methods are needed for assessing improvements made by fishery enhancement and river restoration procedures, and as an intrinsic element of setting environmental flows using instream flow methods. Consequently, the assessment methods must be able to evaluate physical habitat over a range of scales varying from the broad river segment scale (up to hundreds of kilometres) down to the microhabitat level (a few centimetres). 3. Rapid assessment methods involve reconnaissance level surveys (such as the habitat mapping approach) identifying, mapping and measuring key habitat features over long stretches of river in a relatively short space of time. More complex appraisals, such as the Physical Habitat Simulation System (PHABSIM), require more detailed information on microhabitat variations with flow. 4. Key research issues relating to physical habitat evaluation lie in deciding which levels of detail are appropriate for worthwhile yet cost-effective assessment, and in determining those features that are biologically important and hence can be considered habitat features rather than simple geomorphic features. 5. The development of new technologies particularly relating to survey methods should help improve the speed and level of detail attainable by physical habitat assessments. These methods will provide the necessary information required for the development of the two-and three-dimensional physical and hydraulic habitat models. 6. A better understanding of the ways in which the spatial and temporal dynamics of physical habitat determine stream health, and how these elements can be incorporated into assessment methods, remains a key research goal.     

Mapping Cold-Water Coral Habitats at Different Scales within the Northern Ionian Sea (Central Mediterranean): An Assessment of Coral Coverage and Associated Vulnerability

In this study, we mapped the distribution of Cold-Water Coral (CWC) habitats on the northern Ionian Margin (Mediterranean Sea), with an emphasis on assessing coral coverage at various spatial scales over an area of 2,000 km² between 120 and 1,400 m of water depth. Our work made use of a set of data obtained from ship-based research surveys. Multi-scale seafloor mapping data, video inspections, and previous results from sediment samples were integrated and analyzed using Geographic Information System (GIS)-based tools. Results obtained from the application of spatial and textural analytical techniques to acoustic meso-scale maps (i.e. a Digital Terrain Model (DTM) of the seafloor at a 40 m grid cell size and associated terrain parameters) and large-scale maps (i.e. Side-Scan Sonar (SSS) mosaics of 1 m in resolution ground-truthed using underwater video observations) were integrated and revealed that, at the meso-scale level, the main morphological pattern (i.e. the aggregation of mound-like features) associated with CWC habitat occurrences was widespread over a total area of 600 km². Single coral mounds were isolated from the DTM and represented the geomorphic proxies used to model coral distributions within the investigated area. Coral mounds spanned a total area of 68 km² where different coral facies (characterized using video analyses and mapped on SSS mosaics) represent the dominant macro-habitat. We also mapped and classified anthropogenic threats that were identifiable within the examined videos, and, here, discuss their relationship to the mapped distribution of coral habitats and mounds. The combined results (from multi-scale habitat mapping and observations of the distribution of anthropogenic threats) provide the first quantitative assessment of CWC coverage for a Mediterranean province and document the relevant role of seafloor geomorphology in influencing habitat vulnerability to different types of human pressures.     

Multi-scale approach for predicting fish species distributions across coral reef seascapes

Two of the major limitations to effective management of coral reef ecosystems are a lack of information on the spatial distribution of marine species and a paucity of data on the interacting environmental variables that drive distributional patterns. Advances in marine remote sensing, together with the novel integration of landscape ecology and advanced niche modelling techniques provide an unprecedented opportunity to reliably model and map marine species distributions across many kilometres of coral reef ecosystems. We developed a multi-scale approach using three-dimensional seafloor morphology and across-shelf location to predict spatial distributions for five common Caribbean fish species. Seascape topography was quantified from high resolution bathymetry at five spatial scales (5–300 m radii) surrounding fish survey sites. Model performance and map accuracy was assessed for two high performing machine-learning algorithms: Boosted Regression Trees (BRT) and Maximum Entropy Species Distribution Modelling (MaxEnt). The three most important predictors were geographical location across the shelf, followed by a measure of topographic complexity. Predictor contribution differed among species, yet rarely changed across spatial scales. BRT provided ‘outstanding’ model predictions (AUC = >0.9) for three of five fish species. MaxEnt provided ‘outstanding’ model predictions for two of five species, with the remaining three models considered ‘excellent’ (AUC = 0.8–0.9). In contrast, MaxEnt spatial predictions were markedly more accurate (92% map accuracy) than BRT (68% map accuracy). We demonstrate that reliable spatial predictions for a range of key fish species can be achieved by modelling the interaction between the geographical location across the shelf and the topographic heterogeneity of seafloor structure. This multi-scale, analytic approach is an important new cost-effective tool to accurately delineate essential fish habitat and support conservation prioritization in marine protected area design, zoning in marine spatial planning, and ecosystem-based fisheries management.     

Volcanic ash supports a diverse bacterial community in a marine mesocosm

Shallow‐water coral reef ecosystems, particularly those already impaired by anthropogenic pressures, may be highly sensitive to disturbances from natural catastrophic events, such as volcanic eruptions. Explosive volcanic eruptions expel large quantities of silicate ash particles into the atmosphere, which can disperse across millions of square kilometres and deposit into coral reef ecosystems. Following heavy ash deposition, mass mortality of reef biota is expected, but little is known about the recovery of post‐burial reef ecosystems. Reef regeneration depends partly upon the capacity of the ash deposit to be colonised by waterborne bacterial communities and may be influenced to an unknown extent by the physiochemical properties of the ash substrate itself. To determine the potential for volcanic ash to support pioneer bacterial colonisation, we exposed five well‐characterised volcanic and coral reef substrates to a marine aquarium under low light conditions for 3 months: volcanic ash, synthetic volcanic glass, carbonate reef sand, calcite sand and quartz sand. Multivariate statistical analysis of Automated Ribosomal Intergenic Spacer Analysis (ARISA) fingerprinting data demonstrates clear segregation of volcanic substrates from the quartz and coral reef substrates over 3 months of bacterial colonisation. Overall bacterial diversity showed shared and substrate‐specific bacterial communities; however, the volcanic ash substrate supported the most diverse bacterial community. These data suggest a significant influence of substrate properties (composition, granulometry and colour) on bacterial settlement. Our findings provide first insights into physicochemical controls on pioneer bacterial colonisation of volcanic ash and highlight the potential for volcanic ash deposits to support bacterial diversity in the aftermath of reef burial, on timescales that could permit cascading effects on larval settlement.     

Green sturgeon physical habitat use in the coastal Pacific Ocean

The green sturgeon (Acipenser medirostris) is a highly migratory, oceanic, anadromous species with a complex life history that makes it vulnerable to species-wide threats in both freshwater and at sea. Green sturgeon population declines have preceded legal protection and curtailment of activities in marine environments deemed to increase its extinction risk. Yet, its marine habitat is poorly understood. We built a statistical model to characterize green sturgeon marine habitat using data from a coastal tracking array located along the Siletz Reef near Newport, Oregon, USA that recorded the passage of 37 acoustically tagged green sturgeon. We classified seafloor physical habitat features with high-resolution bathymetric and backscatter data. We then described the distribution of habitat components and their relationship to green sturgeon presence using ordination and subsequently used generalized linear model selection to identify important habitat components. Finally, we summarized depth and temperature recordings from seven green sturgeon present off the Oregon coast that were fitted with pop-off archival geolocation tags. Our analyses indicated that green sturgeon, on average, spent a longer duration in areas with high seafloor complexity, especially where a greater proportion of the substrate consists of boulders. Green sturgeon in marine habitats are primarily found at depths of 20-60 meters and from 9.5-16.0°C. Many sturgeon in this study were likely migrating in a northward direction, moving deeper, and may have been using complex seafloor habitat because it coincides with the distribution of benthic prey taxa or provides refuge from predators. Identifying important green sturgeon marine habitat is an essential step towards accurately defining the conditions that are necessary for its survival and will eventually yield range-wide, spatially explicit predictions of green sturgeon distribution.     

The Influence of Data Resolution on Predicted Distribution and Estimates of Extent of Current Protection of Three ‘Listed’ Deep-Sea Habitats

Modelling approaches have the potential to significantly contribute to the spatial management of the deep-sea ecosystem in a cost effective manner. However, we currently have little understanding of the accuracy of such models, developed using limited data, of varying resolution. The aim of this study was to investigate the performance of predictive models constructed using non-simulated (real world) data of different resolution. Predicted distribution maps for three deep-sea habitats were constructed using MaxEnt modelling methods using high resolution multibeam bathymetric data and associated terrain derived variables as predictors. Model performance was evaluated using repeated 75/25 training/test data partitions using AUC and threshold-dependent assessment methods. The overall extent and distribution of each habitat, and the percentage contained within an existing MPA network were quantified and compared to results from low resolution GEBCO models. Predicted spatial extent for scleractinian coral reef and Syringammina fragilissima aggregations decreased with an increase in model resolution, whereas Pheronema carpenteri total suitable area increased. Distinct differences in predicted habitat distribution were observed for all three habitats. Estimates of habitat extent contained within the MPA network all increased when modelled at fine scale. High resolution models performed better than low resolution models according to threshold-dependent evaluation. We recommend the use of high resolution multibeam bathymetry data over low resolution bathymetry data for use in modelling approaches. We do not recommend the use of predictive models to produce absolute values of habitat extent, but likely areas of suitable habitat. Assessments of MPA network effectiveness based on calculations of percentage area protection (policy driven conservation targets) from low resolution models are likely to be fit for purpose.     

Bathymetry,water optical properties,and benthic classification of coral reefs using hyperspectral remote sensing imagery

The complexity and heterogeneity of shallow coastal waters over small spatial scales provides a challenging environment for mapping and monitoring benthic habitats using remote sensing imagery. Additionally, changes in coral reef community structure are occurring on unprecedented temporal scales that require large-scale synoptic coverage and monitoring of coral reefs. A variety of sensors and analyses have been employed for monitoring coral reefs: this study applied a spectrum-matching and look-up-table methodology to the analysis of hyperspectral imagery of a shallow coral reef in the Bahamas. In unconstrained retrievals the retrieved bathymetry was on average within 5% of that measured acoustically, and 92% of pixels had retrieved depths within 25% of the acoustic depth. Retrieved absorption coefficients had less than 20% errors observed at blue wavelengths. The reef scale benthic classification derived by analysis of the imagery was consistent with the percent cover of specific coral reef habitat classes obtained by conventional line transects over the reef, and the inversions were robust as the results were similar when the benthic classification retrieval was constrained by measurements of bathymetry or water column optical properties. These results support the use of calibrated hyperspectral imagery for the rapid determination of bathymetry, water optical properties, and the classification of important habitat classes common to coral reefs.     

Coral and sandstone reef-habitats in north-western Sri Lanka: patterns in the distribution of coral communities

Coral and sandstone reefs cover a significant part ofSri Lanka‘s continental shelf and contain incomparison unique reef structures. Despite this, reefsin this region of the northern Indian Ocean havereceived little research attention. In an attempt tobetter understand these ecosystems and their innatecharacter, this study describes the variety of reeftypes and habitats that are found in this area. Thestudy concentrated on four major reef areas: the BarReef Marine Sanctuary (BRMS), Kandakuliya Reefs,Talawila Reef, and Mampuri Reef. These reefs showedapparent differences in habitat structure in terms ofthe proportion coral cover, coral species compositionand structural complexity. Two reef types were presentwithin the continental shelf of BRMS: coral andsandstone patch-reefs. Acropora was the mostdominant coral genera however in total 118madreporarian species and 50 coral genera wererecorded in the sanctuary. Distinct habitats wereidentified within reef types including shallow reefflat, shallow patch reef, deep reef flat and Porites dome habitats for the coral-reef patcheswhile the sandstone-reef patches were divided intostructured and flat sandstone reef habitats.Kandakuliya Reef south of BRMS was by large dominatedby coral rubble. Talawila Reef and Mampuri Reef showedunique structures with the former being dominated bymassive corals and the latter mainly containingsandstone structures. In addition to factors such asbio-erosion, sedimentation, hydrodynamics, andrecruitment or colonisation processes, some reefs wereclearly under significant direct human impact whichappeared to play a dominant role in habitatstructuring. However, type and degree of humandisturbance varied among the reefs. Habitat alterationat Kandakuliya Reef was the result of intense fishingusing destructive fishing methods. Talawila Reef andMampuri Reef was also influenced by fishing activitiesthough reef structure seemed less affected. This revised version was published online in July 2006 with corrections to the Cover Date.     

Predictability of reef fish diversity and abundance using remote sensing data in Diego Garcia (Chagos Archipelago)

The diversity, abundance and distribution of reef fish are related to heterogeneity and physical complexity of benthic habitat. However, the field effort required to evaluate these aspects of the benthos in situ, at the scale of entire reefscapes, is greatly constrained by logistical and resource limitations. With moderate ground truthing, both substratum type and seabed topography are amenable to monitoring using satellite data. Here, remote sensing imagery was used to resolve the bathymetry and benthic character of a reef system in Diego Garcia (British Indian Ocean Territory). Replicate fish counts were made at seven measurement stations across the study area using visual census. Monte Carlo simulation revealed that species richness and abundance of several guilds and size groupings of reef fish appraised in situ were correlated with the satellite-derived seabed parameters over areas of seafloor as large as 5,030 m². The study suggests that satellite remote sensing is capable of predicting habitat complexity at a scale relevant to fish. Furthermore, as larger size classes of fish were better predicted with the satellite habitat complexity data, this technique could be used to predict fish stocks and identify potential sites for marine protected areas where intensive field surveys are not practical.     

Discovery of a “Living Dinosaur”: Globally unique modern hexactinellid sponge reefs off British Columbia, Canada

Summary Globally unique hexactinellid sponge reefs occur on the continental shelf off British Columbia, Canada. They cover about 425 km² of seafloor on the continental shelf off British Columbia (Canada) in water depths between 165 and 240 metres and occur on a low-angle deep shelf, iceberg scoured seafloor, characterized by very low sedimentation rates and very stable environmental conditions. The sponge bioherms are up to 19 metres high with steep flanks, whereas the biostromes are 2–10 metres thick and many kilometres wide. They all consist of dense populations of only seven hexactinellid species. Three of them, all hexactinosan species (Aphrocallistes vastus, Heterochone calyx, Farrea occa) are the main frambuilders, composing a true rigid framework of sponge skeletons encased in a organic rich matrix of modern clay baffled by the sponges. Growth rates of hexactinosan sponges range in the order of 0–7 centimetres per year. The base of the oldest sponge reefs date from approximately 9000 years b.p. Different invertebrate and fish faunas occupy the reefs than occur on adjacent seafloor areas and some species appear to use the sponge reef complex structures as refugia where they can hide. Sidescan sonar data and direct observation by manned submersible clearly show that large areas of sponge reefs have been heavily damaged by seafloor trawling in the past decade. These unique extant siliceous sponge reefs can be used as a modern analogue for a better understanding and interpretation of fossil siliceous sponge reefs, known from many ages and many locations world wide.     

Detached kelps from distant sources are a food subsidy for sea urchins

Trophic subsidies link habitats and can determine community structure in the subsidised habitats. Knowledge of the spatial extents of trophic interactions is important for understanding food webs, and for making spatial management practices more efficient. We demonstrate trophic linkages between detached (drift) fragments of the kelp Ecklonia radiata and the purple sea urchin Heliocidaris erythrogramma among discrete rocky reefs separated by kilometres. Sea urchins were abundant at one inshore reef, where the biomass of drift was usually high. There, sea urchins trapped detached kelp at high rates, although local kelp abundance was low. Most detached kelp present on the reef was retained by sea urchins. Detached seagrass, which was abundant on the reef, was not retained by sea urchins in large quantities. Experiments with tethered pieces of kelp showed that sea urchins only consumed detached fragments, and did not consume attached kelps. Comparisons of the morphology of detached fragments of kelp collected from the inshore reef to attached kelps from reefs further offshore showed that a large proportion (30-95%, varying among dates) of the fragments originated at distant reefs (>/=2 km away). At the inshore reef, the sea urchin H. erythrogramma is subsidised by detached kelps, and detached kelp fragments have been transported across landscapes. Cross-habitat resource subsidies therefore link discrete reef habitats separated by kilometres of non-reef habitat.     

Fitness consequences of habitat variability,trophic position,and energy allocation across the depth distribution of a coral-reef fish

Environmental clines such as latitude and depth that limit species’ distributions may be associated with gradients in habitat suitability that can affect the fitness of an organism. With the global loss of shallow-water photosynthetic coral reefs, mesophotic coral ecosystems (~30–150 m) may be buffered from some environmental stressors, thereby serving as refuges for a range of organisms including mobile obligate reef dwellers. Yet habitat suitability may be diminished at the depth boundary of photosynthetic coral reefs. We assessed the suitability of coral-reef habitats across the majority of the depth distribution of a common demersal reef fish (Stegastes partitus) ranging from shallow shelf (SS, <10 m) and deep shelf (DS, 20–30 m) habitats in the Florida Keys to mesophotic depths (MP, 60–70 m) at Pulley Ridge on the west Florida Shelf. Diet, behavior, and potential energetic trade-offs differed across study sites, but did not always have a monotonic relationship with depth, suggesting that some drivers of habitat suitability are decoupled from depth and may be linked with geographic location or the local environment. Feeding and diet composition differed among depths with the highest consumption of annelids, lowest ingestion of appendicularians, and the lowest gut fullness in DS habitats where predator densities were highest and fish exhibited risk-averse behavior that may restrict foraging. Fish in MP environments had a broader diet niche, higher trophic position, and higher muscle C:N ratios compared to shallower environments. High C:N ratios suggest increased tissue lipid content in fish in MP habitats that coincided with higher investment in reproduction based on gonado-somatic index. These results suggest that peripheral MP reefs are suitable habitats for demersal reef fish and may be important refuges for organisms common on declining shallow coral reefs.