Upper and lower mesophotic coral reef fish communities evaluated by underwater visual censuses in two Caribbean locations

Despite more than 60 yr of coral reef research using scuba diving, mesophotic coral ecosystems (MCEs) between 30 and 150 m depth remain largely unknown. This study represents the first underwater visual census of reef fish communities in the Greater Caribbean on MCEs at depths up to 80 m in Bermuda and 130 m in Curaçao. Sampling was performed using mixed-gas closed-circuit rebreathers. Quantitative data on reef fish communities were obtained for four habitats: coral reefs (45–80 m), rhodolith beds (45–80 m), ledges (85–130 m) and walls (85–130 m). A total of 38 species were recorded in Bermuda and 66 in Curaçao. Mesophotic reef fish communities varied significantly between the two localities. MCEs in Bermuda had lower richness and abundance, but higher biomass than those in Curaçao. Richness, abundance and biomass increased with depth in Bermuda, but decreased in Curaçao. A high turnover of species was found among depth strata and between Bermuda and other Caribbean upper MCEs (45–80 m), indicating that depth was an important driver of community structure at all localities. However, local and evolutionary factors (habitat and endemism) are likely the main factors shaping communities in isolated locations such as Bermuda. High fishing pressure is evident in both localities, as total biomass of apex predators was generally low, and thus may be driving a “refugia” scenario in Bermuda, as the abundance and biomass of macro-carnivores increased with depth and distance from the coast.     

Intraspecific variability in the life histories of endemic coral-reef fishes between photic and mesophotic depths across the Central Pacific Ocean

Mesophotic coral ecosystems (MCEs) represent the lowest depth distribution inhabited by many coral reef-associated organisms. Research on fishes associated with MCEs is sparse, leading to a critical lack of knowledge of how reef fish found at mesophotic depths may vary from their shallow reef conspecifics. We investigated intraspecific variability in body condition and growth of three Hawaiian endemics collected from shallow, photic reefs (5–33 m deep) and MCEs (40–75 m) throughout the Hawaiian Archipelago and Johnston Atoll: the detritivorous goldring surgeonfish, Ctenochaetus strigosus, and the planktivorous threespot chromis, Chromis verater, and Hawaiian dascyllus, Dascyllus albisella. Estimates of body condition and size-at-age varied between shallow and mesophotic depths; however, these demographic differences were outweighed by the magnitude of variability found across the latitudinal gradient of locations sampled within the Central Pacific. Body condition and maximum body size were lowest in samples collected from shallow and mesophotic Johnston Atoll sites, with no difference occurring between depths. Samples from the Northwestern Hawaiian Islands tended to have the highest body condition and reached the largest body sizes, with differences between shallow and mesophotic sites highly variable among species. The findings of this study support newly emerging research demonstrating intraspecific variability in the life history of coral-reef fish species whose distributions span shallow and mesophotic reefs. This suggests not only that the conservation and fisheries management should take into consideration differences in the life histories of reef-fish populations across spatial scales, but also that information derived from studies of shallow fishes be applied with caution to conspecific populations in mesophotic coral environments.

     

Mesophotic depths as refuge areas for fishery-targeted species on coral reefs

Coral reefs are subjected to unprecedented levels of disturbance with population growth and climate change combining to reduce standing coral cover and stocks of reef fishes. Most of the damage is concentrated in shallow waters (<30 m deep) where humans can comfortably operate and where physical disturbances are most disruptive to marine organisms. Yet coral reefs can extend to depths exceeding 100 m, potentially offering refuge from the threats facing shallower reefs. We deployed baited remote underwater stereo-video systems (stereo-BRUVs) at depths of 10–90 m around the southern Mariana Islands to investigate whether fish species targeted by fishing in the shallows may be accruing benefits from being at depth. We show that biomass, abundance and species richness of fishery-targeted species increased from shallow reef areas to a depth of 60 m, whereas at greater depths, a lack of live coral habitat corresponded to lower numbers of fish. The majority of targeted species were found to have distributions that ranged from shallow depths (10 m) to depths of at least 70 m, emphasising that habitat, not depth, is the limiting factor in their vertical distribution. While the gradient of abundance and biomass versus depth was steepest for predatory species, the first species usually targeted by fishing, we also found that fishery-targeted herbivores prevailed in similar biomass and species richness to 60 m. Compared to shallow marine protected areas, there was clearly greater biomass of fishery-targeted species accrued in mesophotic depths. Particularly some species typically harvested by depth-limited fishing methods (e.g., spearfishing), such as the endangered humphead wrasse Cheilinus undulatus, were found in greater abundance on deeper reefs. We conclude that mesophotic depths provide essential fish habitat and refuge for fishery-targeted species, representing crucial zones for fishery management and research into the resilience of disturbed coral reef ecosystems.     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

Mesophotic coral ecosystems in the Hawaiian Archipelago

Efforts to map coral reef ecosystems in the Hawaiian Archipelago using optical imagery have revealed the presence of numerous scleractinian, zoothanthellate coral reefs at depths of 30–130+ m, most of which were previously undiscovered. Such coral reefs and their associated communities have been recently defined as mesophotic coral ecosystems (MCEs). Several types of MCEs are found in Hawai‘i, each of which dominates a different depth range and is characterized by a unique pattern of coral community structure and colony morphology. Although MCEs are documented near both ends of the archipelago and on many of the islands in between, the maximum depth and prevalence of MCEs in Hawai‘i were found to decline with increasing latitude. The Main Hawaiian Islands (MHI) had significantly deeper and greater percentages of scleractinian coral, and peaks in cover of both scleractinian corals and macroalgae occurred within depth bins 20 m deeper than in the Northwestern Hawaiian Islands (NWHI). Across the archipelago, as depth increased the combined percentage of living cover of mega benthic taxa declined sharply with increasing depth below 70 m, despite the widespread availability of hard substrate.     

Effects of habitat,wave exposure,and marine protected area status on coral reef fish assemblages in the Hawaiian archipelago

The relationships between fish assemblages, their associated habitat, and degree of protection from fishing were evaluated over a broad spatial scale throughout the main Hawaiian islands. Most fish assemblage characteristics showed positive responses to protection whether it was physical (e.g. habitat complexity), biological (e.g. coral cover growth forms), or human-induced (e.g. marine reserves). Fish biomass was lowest in areas of direct wave exposure and highest in areas partially sheltered from swells. Higher values for fish species richness, number of individuals, biomass, and diversity were observed in locations with higher substrate complexity. Areas completely protected from fishing had distinct fish assemblages with higher standing stock and diversity than areas where fishing was permitted or areas that were partially protected from fishing. Locations influenced by customary stewardship harbored fish biomass that was equal to or greater than that of no-take protected areas. Marine protected areas in the main Hawaiian islands with high habitat complexity, moderate wave disturbance, a high percentage of branching and/or lobate coral coupled with legal protection from fishing pressure had higher values for most fish assemblage characteristics.     

Characterization of the mesophotic reef fish community in south Florida,USA

The southeastern coast of Florida, USA supports a substantial recreational fishery, yet little is known of the coral reef ecosystem or fisheries resources past 50 m depth. Fish assemblages associated with low‐relief substrate and three vessel reefs between 50 and 120 m depth off southeast Florida were surveyed by remotely operated vehicles providing the first characterization of the mesophotic fish assemblages in the region. Two distinct assemblages were observed on low‐relief substrate and high‐relief vessel reefs. A total of 560 fishes of 42 species was recorded on 27 dives on low‐relief substrate, and 50 152 fishes of 65 species were recorded on 24 dives on three vessel reefs. Small planktivorous Anthiinae fishes and several economically valuable species were common on vessel reefs but rare on low‐relief substrate. Fish assemblages on vessel reefs more closely resembled those found at similar depths in high‐relief natural areas off east‐central Florida and the Gulf of Mexico than those associated with adjacent low‐relief habitat or nearby coral reef tracts. From a fisheries perspective, these results provide limited support to the hypothesis that in deep‐water regions with limited relief, vessel reefs may provide an opportunity to increase fish diversity and abundance by creating high‐relief habitat without compromising adjacent fish assemblages.     

Reef habitats and associated sessile-benthic and fish assemblages across a euphotic–mesophotic depth gradient in Isla Desecheo, Puerto Rico

Quantitative surveys of sessile benthos and fish populations associated with reef habitats across a 15–50 m depth gradient were performed by direct diver observations using rebreathers at Isla Desecheo, Puerto Rico. Statistically significant differences between depths were found for total live coral, total coral species, total benthic algae, total sponges and abiotic cover. Live coral cover was higher at the mid-shelf (20 m) and shelf-edge (25 m) stations, whereas benthic algae and sponges were the dominant sessile-benthic assemblage at mesophotic stations below 25 m. Marked shifts in the community structure of corals and benthic algae were observed across the depth gradient. A total of 119 diurnal, non-cryptic fish species were observed across the depth gradient, including 80 species distributed among 7,841 individuals counted within belt-transects. Fish species richness was positively correlated with live coral cover. However, the relationship between total fish abundance and live coral was weak. Abundance of several numerically dominant fish species varied independently from live coral cover and appeared to be more influenced by depth and/or habitat type. Statistically significant differences in the rank order of abundance of fish species at euphotic vs mesophotic stations were detected. A small assemblage of reef fishes that included the cherubfish, Centropyge argi, sunshine chromis, Chromis insolata, greenblotch parrotfish, Sparisoma atomarium, yellowcheek wrasse, Halichoeres cyanocephalus, sargassum triggerfish, Xanthichthys ringens, and the longsnout butterflyfish, Chaetodon aculeatus was most abundant or only present from stations deeper than 30 m, and thus appear to be indicator species of mesophotic habitats.     

Towards an understanding of resilience in isolated coral reefs

In 1998, seawater temperature anomalies led to unprecedented levels of coral bleaching on reefs worldwide. We studied the direct effects of this thermal event on benthic communities and its indirect effects on their associated coral reef fish communities at a group of remote reefs off NW Australia. Long‐term monitoring of benthic and fish assemblages on these reefs allowed us to compare the responses of these communities to coral bleaching using a data series that included 4 years before, and 6 years following, this bleaching event. While bleaching mortality was evident to >30 m depth, it was patchy among the shallower survey sites with decreases in live coral cover ranging from 30% to 90% across seven surveyed locations Within 2 years of the bleaching, hard coral recovery had begun at all sites and by 2003 reef‐wide coral cover had increased to ～39% of its preimpact levels. We exploited this pattern of differential survival of corals among sites, the associated changes in these benthic communities, and their patterns of recovery, to better understand links between benthic community dynamics and their associated fish communities. Temporal changes in the resident fish communities strongly reflected the differential shifts in the benthic communities, but were lagged by 12–18 months. Five years after the bleaching event, the fish communities on five of the seven surveyed locations showed evidence of recovery, however, none had regained their preimpact structures. Analyses of these communities by taxonomic family revealed a range of responses to the disturbance reflective of their life‐histories and trophic and habitat affiliations. The slow but recognizable recovery of this isolated reef system has parallels with other relatively isolated systems that displayed resilience to the 1998 bleaching event, e.g. the Chagos archipelago, but it also contrasts sharply with low levels of resilience documented in other isolated reef systems subject to the same disturbance, e.g. the Seychelles. In this context, our results highlight the significant knowledge gaps remaining in understanding the resilience of these ecosystems to disturbance.     

Disturbance and the structure of coral reef fish communities on the reef slope

Three levels of physical disturbance were applied to corals in permanent 10x10 m quadrats along a section of fringing reef at Lizard Island on the Great Barrier Reef to investigate the response of fish assemblages. Tabular and corymbose corals were overturned and left in situ, reducing total hard coral cover from ˜55% to ˜47%, ˜43%, and ˜34%. Despite pre-existing associations with benthic cover, all fish groups examined (pomacentrids, labrids, chaetodontids, and acanthurids) were resistent to benthic disturbances at the level and scale at which they were applied. Partial Mantel's tests, in combination with partial Canonical Correspondence Analysis enabled spatial and temporal variation to be factored out from experimental effects. Most of the variation in the fish community could be assigned to spatio-temporal variables, indicating that spatial structure over the reef landscape may moderate localised disturbance effects. This study indicates that coral reef fish assemblages may be more resistant to disturbance than many correlative studies would suggest, and highlights a need for further information on levels and scales of natural habitat disturbance in order to apply a structured approach to the experimental investigation of the importance of habitat in structuring coral-reef fish assemblages.     

High macroalgal cover and low coral recruitment undermines the potential resilience of the world's southernmost coral reef assemblages

Coral reefs are under increasing pressure from anthropogenic and climate-induced stressors. The ability of reefs to reassemble and regenerate after disturbances (i.e., resilience) is largely dependent on the capacity of herbivores to prevent macroalgal expansion, and the replenishment of coral populations through larval recruitment. Currently there is a paucity of this information for higher latitude, subtropical reefs. To assess the potential resilience of the benthic reef assemblages of Lord Howe Island (31°32'S, 159°04'E), the worlds' southernmost coral reef, we quantified the benthic composition, densities of juvenile corals (as a proxy for coral recruitment), and herbivorous fish communities. Despite some variation among habitats and sites, benthic communities were dominated by live scleractinian corals (mean cover 37.4%) and fleshy macroalgae (20.9%). Live coral cover was higher than in most other subtropical reefs and directly comparable to lower latitude tropical reefs. Juvenile coral densities (0.8 ind.m(-2)), however, were 5-200 times lower than those reported for tropical reefs. Overall, macroalgal cover was negatively related to the cover of live coral and the density of juvenile corals, but displayed no relationship with herbivorous fish biomass. The biomass of herbivorous fishes was relatively low (204 kg.ha(-1)), and in marked contrast to tropical reefs was dominated by macroalgal browsing species (84.1%) with relatively few grazing species. Despite their extremely low biomass, grazing fishes were positively related to both the density of juvenile corals and the cover of bare substrata, suggesting that they may enhance the recruitment of corals through the provision of suitable settlement sites. Although Lord Howe Islands' reefs are currently coral-dominated, the high macroalgal cover, coupled with limited coral recruitment and low coral growth rates suggest these reefs may be extremely susceptible to future disturbances.     

A community and functional comparison of coral and reef fish assemblages between four decades of coastal urbanisation and thermal stress

1. Urbanized coral reefs experience anthropogenic disturbances caused by coastal development, pollution, and nutrient runoff, resulting in turbid, marginal conditions in which only certain species can persist. Mortality effects are exacerbated by increasingly regular thermal stress events, leading to shifts towards novel communities dominated by habitat generalists and species with low structural complexity.
2. There is limited data on the turnover processes that occur due to this convergence of anthropogenic stressors, and how novel urban ecosystems are structured both at the community and functional levels. As such, it is unclear how they will respond to future disturbance events.
3. Here, we examine the patterns of coral reef community change and determine whether ecosystem functions provided by specialist species are lost post‐disturbance. We present a comparison of community and functional trait‐based changes for scleractinian coral genera and reef fish species assemblages subject to coastal development, coastal modification, and mass bleaching between two time periods, 1975–1976 and 2018, in Nakagusuku Bay, Okinawa, Japan.
4. We observed an increase in fish habitat generalists, a dominance shift from branching to massive/sub‐massive corals and increasing site‐based coral genera richness between years. Fish and coral communities significantly reassembled, but functional trait‐based multivariate space remained constant, indicating a turnover of species with similar traits. A compression of coral habitat occurred, with shallow (<5 m) and deep (>8 m) coral genera shifting towards the mid‐depths (5–8 m).
5. We show that although reef species assemblages altered post disturbance, new communities retained similar ecosystem functions. This result could be linked to the stressors experienced by urban reefs, which reflect those that will occur at an increasing frequency globally in the near future. Yet, even after shifts to disturbed communities, these fully functioning reef systems may maintain high conservation value.

     

Differential Response of Fish Assemblages to Coral Reef-Based Seaweed Farming

As the global demand for seaweed-derived products drives the expansion of seaweed farming onto shallow coral ecosystems, the effects of farms on fish assemblages remain largely unexplored. Shallow coral reefs provide food and shelter for highly diverse fish assemblages but are increasingly modified by anthropogenic activities. We hypothesized that the introduction of seaweed farms into degraded shallow coral reefs had potential to generate ecological benefits for fish by adding structural complexity and a possible food source. We conducted 210 transects at 14 locations, with sampling stratified across seaweed farms and sites adjacent to and distant from farms. At a seascape scale, locations were classified by their level of exposure to human disturbance. We compared sites where (1) marine protected areas (MPAs) were established, (2) neither MPAs nor blast fishing was present (hence “unprotected”), and (3) blast fishing occurred. We observed 80,186 fish representing 148 species from 38 families. The negative effects of seaweed farms on fish assemblages appeared stronger in the absence of blast fishing and were strongest when MPAs were present, likely reflecting the positive influence of the MPAs on fish within them. Species differentiating fish assemblages with respect to seaweed farming and disturbance were typically small but also included two key target species. The propensity for seaweed farms to increase fish diversity, abundance, and biomass is limited and may reduce MPA benefits. We suggest that careful consideration be given to the placement of seaweed farms relative to MPAs.     

Quantifying the direct and indirect effects of flow‐related disturbance on stream fish assemblages

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Mesophotic reef fish assemblages of the remote St. Peter and St. Paul’s Archipelago,Mid-Atlantic Ridge,Brazil

Mesophotic reef fish assemblages (30–90 m depth) of the small and remote St. Peter and St. Paul’s Archipelago (SPSPA), Mid-Atlantic Ridge, Brazil, were characterized using remotely operated vehicles. Ordination analyses identified distinct fish assemblages in the upper (30–50 m) and lower (50–90 m) mesophotic zones, the former characterized by high abundances of species that are also abundant at euphotic reefs (Caranx lugubris, Melichthys niger, Stegastes sanctipauli and Chromis multilineata) and the latter dominated by two mesophotic specialists (Prognathodes obliquus and Chromis enchrysura). Planktivores dominated fish assemblages, particularly in the upper mesophotic zone, possibly due to a greater availability of zooplankton coming from the colder Equatorial Undercurrent in mesophotic depths of the SPSPA. Turf algae, fleshy macroalgae and scleractinian corals dominated benthic assemblages between 30 and 40 m depth, while bryozoans, black corals and sponges dominated between 40 and 90 m depth. Canonical correspondence analysis explained 74 % of the relationship between environmental characteristics (depth, benthic cover and complexity) and structure of fish assemblages, with depth as the most important independent variable. Juveniles of Bodianus insularis and adults of P. obliquus and C. enchrysura were clearly associated with branching black corals (Tanacetipathes spp.), suggesting that black corals play key ecological roles in lower mesophotic reefs of the SPSPA. Results from this study add to the global database about mesophotic reef ecosystems (MREs) and provide a baseline for future evaluations of possible anthropogenic and natural disturbances on MREs of the SPSPA.     

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.     

Theme section on “Mesophotic Coral Ecosystems: Characterization, Ecology, and Management”

Mesophotic coral ecosystems (MCEs) are characterized by the presence of light-dependent corals and associated communities that are typically found at depths ranging from 30 to 40 m and extending to over 150 m in tropical and subtropical regions. The dominant communities providing structural habitat in the mesophotic zone can be comprised of coral, sponge, and algal species. Because working in this depth range is constrained by traditional SCUBA limits, less is known about corals and associated organisms there compared to shallower coral communities. Following the first-ever gathering of international scientists to review and discuss existing knowledge of MCEs, this issue focuses on the ecological characterization, geomorphology, and concept of MCEs as refugia for shallow-water populations. The review and research papers comprising this special issue reflect the current scientific understanding of these ecosystems and the underlying mechanisms that regulate them, as well as potential resource management implications. It is important to understand the value and role of mesophotic coral ecosystems in tropical and subtropical regions as these areas face increasing environmental change and human impacts     

Habitat characteristics can influence fish assemblages in high latitude kelp forests

Kelp forests are patchy fish-associated habitats, which can vary greatly in their size, foundation species, and several physical habitat attributes. The structure of fish assemblages can vary with these characteristics and with the location of the assemblage within the forest, i.e. edges versus interiors. This study quantified the biological and physical heterogeneity within different sized kelp forests and identified which factors are important in structuring the associated fish assemblages. Fish and habitat surveys were conducted at the edge and interiors of ten kelp forests of varying sizes in Kachemak Bay, south-central Alaska. Fish assemblage structure was not correlated with the species composition of surface canopy forming foundation kelps (Nereocystis leutkeana and Eualaria fistulosa) or with kelp forest size. Instead, it correlated with the abundances of two understory foundation kelps (Agarum clathratum and Saccharina latissima), substratum rugosity, and water depth. Together these benthic attributes explained 53.6 % of the fish assemblage variability. Additionally, significantly different fish assemblages were found at edge compared to interior locations with the relative abundance of seven species (Artedius fenestralis, Ammodytes hexapterus, Blepsias cirrhosis, Gadus macrocephalus, Hexagrammos stelleri, Pholis laeta, and Sebastes melanops) explaining 91.4 % of the variability. This study highlights the importance of habitat characteristics such as understory foundation species, substratum rugosity, water depth and location within a patch on the variability of fish assemblages in high latitude kelp forests.     

An assessment of shallow and mesophotic reef brachyuran crab assemblages on the south shore of O‘ahu,Hawai‘i

Shallow coral reefs are extensively studied but, although scleractinian corals have been recorded to 165 m, little is known about other mesophotic coral reef ecosystem (MCE) inhabitants. Brachyuran crabs fill many ecological and trophic niches on reefs, making them ideal candidates for evaluating species composition among depths to ask whether MCEs host the same communities as shallower reef communities that have been well studied. Here we deployed autonomous reef monitoring structures for 2 yr on the south shore of O‘ahu along a depth gradient (12, 30, 60, and 90 m) to sample and assess brachyuran crab communities. A total of 663 brachyuran crabs representing 69 morphospecies (16 families) were found. Community composition was not significantly different within depths, but was highly stratified by depth. Each depth was distinct, but the 30 and 60 m depths were least dissimilar from one another. We show that deeper reefs host significantly different brachyuran communities, and at much lower total abundance, than shallow reefs in Hawai‘i, with 4–27 unique morphospecies per depth and only 3 of 69 morphospecies (~4 %) occurring across the entire depth range sampled.