An underwater imagery identification guide for shallow, mesophotic and deep-sea benthos in Maldives.

Background: During the 2022 Nekton Maldives Mission, we deployed a variety of platforms (snorkelling, remotely-operated vehicles and manned submersibles) to conduct video surveys of the biodiversity and composition of shallow (< 30 m), mesophotic (30-150 m) and deep-sea (> 150 m) benthos found in the Maldives' central and southern atolls. In total, ~ 80 hrs of stereo-video footage were collected during the benthic transect surveys, which were subsequently processed using annotation software in order to evaluate benthic biodiversity and community composition. Here, we present a photographic guide for the visual, in situ identification of reef benthos encountered, including corals, sponges and other invertebrates that inhabit Maldives' nearshore habitats. We hope that this identification guide will aid future imagery-based surveys or observations of organisms during fieldwork. New information: A total of 283 morphotypes were identified, including those belonging to Octocorallia (61), Scleractinia (57), Porifera (38), Asteroidea (22), Antipatharia (15), Decapoda (13), Hydrozoa (12), Holothuroidea (10), Actiniaria (9), Echinoidea (8), Annelida (6), Chlorophyta (5), Gastropoda (4), Bivalvia (4), Ascidiacea (3), Crinoidea (3), Bryozoa (2), Cyanobacteria (2), Zoantharia (2), Cephalopoda (1), Ceriantharia (1), Corallimorpharia (1), Ctenophora (1), Ophiuroidea (1), Rhodophyta (1) and to an unknown category (1). Out of these, we identified 40 to species level, 120 to genus, 47 to family, 14 to order and suborder, 58 to class and subclass, two to phylum and one was of unknown phylum. This represents the first attempt to catalogue the mesophotic and deep-sea benthic megafaunal diversity in the Maldives using underwater imagery.

Mesophotic.org: a repository for scientific information on mesophotic ecosystems.

Mesophotic coral ecosystems (MCEs) and temperate mesophotic ecosystems (TMEs) occur at depths of roughly 30-150 m depth and are characterized by the presence of photosynthetic organisms despite reduced light availability. Exploration of these ecosystems dates back several decades, but our knowledge remained extremely limited until about a decade ago, when a renewed interest resulted in the establishment of a rapidly growing research community. Here, we present the 'mesophotic.org' database, a comprehensive and curated repository of scientific literature on mesophotic ecosystems. Through both manually curated and automatically extracted metadata, the repository facilitates rapid retrieval of available information about particular topics (e.g. taxa or geographic regions), exploration of spatial/temporal trends in research and identification of knowledge gaps. The repository can be queried to comprehensively obtain available data to address large-scale questions and guide future research directions. Overall, the 'mesophotic.org' repository provides an independent and open-source platform for the ever-growing research community working on MCEs and TMEs to collate and expedite our understanding of the occurrence, composition and functioning of these ecosystems. Database URL: http://mesophotic.org/.

Assessing population changes of historically overexploited black corals (Order: Antipatharia) in Cozumel, Mexico.

Black corals (Antipatharians) are crucial structural and ecological components of many mesophotic coral ecosystems (MCEs; reefs 30-150 m depth). In Mexico, black corals are harvested for the jewellery industry, which has historically led to population depletion. Harvesting began in the early 1960s and was concentrated around Cozumel Island until 1995. Since then, harvesting permits have been granted only for the mainland coast. Here we compare Cozumel populations between 1998 and 2016 for the two black coral species targeted by the jewellery industry. We found that densities of Plumapathes pennacea in 2016 were substantially lower than in 1998. However, the 2016 P. pennacea population has shifted to be dominated by larger colonies, suggesting disproportionate juvenile mortality or recruitment failure. Low numbers of Antipathes caribbeana were recorded, and no change in population density or colony size was detected between 1998 and 2016. Despite harvesting occurring for almost 70 years in the Mexican Caribbean, no information on reproduction, recruitment and other dynamics of the targeted species is available. We advocate for harvesting permits to be based on scientific evidence, and for implementation of future restrictions to prevent total depletion of black corals in the area.

Assessing mesophotic coral ecosystems inside and outside a Caribbean marine protected area.

Widespread shallow coral reef loss has led to calls for more holistic approaches to coral reef management, requiring inclusion of ecosystems interacting with shallow coral reefs in management plans. Yet, almost all current reef management is biased towards shallow reefs, and overlooks that coral reefs extend beyond shallow waters to mesophotic coral ecosystems (MCEs; 30-150 m). We present the first detailed quantitative characterization of MCEs off Cozumel, Mexico, on the northern Mesoamerican Reef in the Mexican Caribbean, and provide insights into their general state. We documented MCE biodiversity, and assessed whether MCEs adjacent to a major town and port, where coastal development has caused shallow reef damage, have similar benthic and fish communities to MCEs within a National Park. Our results show that overall MCE communities are similar regardless of protection, though some taxa-specific differences exist in benthic communities between sites within the MPA and areas outside. Regardless of protection and location, and in contrast to shallow reefs, all observed Cozumel MCEs were continuous reefs with the main structural habitat complexity provided by calcareous macroalgae, sponges, gorgonians and black corals. Hard corals were present on MCEs, although at low abundance. We found that 42.5% of fish species recorded on Cozumel could be found on both shallow reefs and MCEs, including 39.6% of commercially valuable fish species. These results suggest that MCEs could play an important role in supporting fish populations. However, regardless of protection and depth, we found few large-body fishes (greater than 500 mm), which were nearly absent at all studied sites. Cozumel MCEs contain diverse benthic and fish assemblages, including commercially valuable fisheries species and ecosystem engineers, such as black corals. Because of their inherent biodiversity and identified threats, MCEs should be incorporated into shallow-reef-focused Cozumel National Park management plan.

Lionfish (Pterois spp.) invade the upper-bathyal zone in the western Atlantic.

Non-native lionfish have been recorded throughout the western Atlantic on both shallow and mesophotic reefs, where they have been linked to declines in reef health. In this study we report the first lionfish observations from the deep sea (>200 m) in Bermuda and Roatan, Honduras, with lionfish observed to a maximum depth of 304 m off the Bermuda platform, and 250 m off West End, Roatan. Placed in the context of other deeper lionfish observations and records, our results imply that lionfish may be present in the 200-300 m depth range of the upper-bathyal zone across many locations in the western Atlantic, but currently are under-sampled compared to shallow habitats. We highlight the need for considering deep-sea lionfish populations in future invasive lionfish management.

Identifying zooplankton community changes between shallow and upper-mesophotic reefs on the Mesoamerican Barrier Reef, Caribbean.

Mesophotic coral ecosystems (MCEs, reefs 30-150 m) are understudied, yet the limited research conducted has been biased towards large sessile taxa, such as scleractinian corals and sponges, or mobile taxa such as fishes. Here we investigate zooplankton communities on shallow reefs and MCEs around Utila on the southern Mesoamerican Barrier Reef using planktonic light traps. Zooplankton samples were sorted into broad taxonomic groups. Our results indicate similar taxonomic zooplankton richness and overall biomass between shallow reefs and MCEs. However, the abundance of larger bodied (>2 mm) zooplanktonic groups, including decapod crab zoea, mysid shrimps and peracarid crustaceans, was higher on MCEs than shallow reefs. Our findings highlight the importance of considering zooplankton when identifying broader reef community shifts across the shallow reef to MCE depth gradient.

Depth-dependent effects of culling-do mesophotic lionfish populations undermine current management?

Invasive lionfish (Pterois volitans and P. miles) have spread widely across the western Atlantic and are recognized as a major threat to native marine biodiversity. Although lionfish inhabit both shallow reefs and mesophotic coral ecosystems (MCEs; reefs from 30 to 150 m depth), the primary management response implemented by many countries has been diver-led culling limited to reefs less than 30 m. However, many reef fish undergo ontogenetic migrations, with the largest and therefore most fecund individuals found at greatest depths. Here, we study lionfish density, body size, maturity and dietary patterns across the depth gradient from the surface down to 85 m on heavily culled reefs around Utila, Honduras. We found lionfish at increased densities, body size and weight on MCEs compared with shallow reefs, with MCEs also containing the greatest proportion of actively spawning females, while shallow reefs contained the greatest proportion of immature lionfish. We then compared lionfish behaviour in response to divers on shallow culled and mesophotic unculled Utilan reefs, and on shallow unculled reefs in Tela Bay, on the Honduran mainland. We found that mesophotic lionfish exhibited high alert distances, consistent with individuals previously exposed to culling despite being below the depth limits of removal. In addition, when examining stomach content, we found that fish were the major component of lionfish diets across the depth gradient. Importantly, our results suggest that despite adjacent shallow culling, MCEs retain substantial lionfish populations that may be disproportionately contributing towards continued lionfish recruitment onto the shallow reefs of Utila, potentially undermining current culling-based management.

Assessing Caribbean Shallow and Mesophotic Reef Fish Communities Using Baited-Remote Underwater Video (BRUV) and Diver-Operated Video (DOV) Survey Techniques.

Fish surveys form the backbone of reef monitoring and management initiatives throughout the tropics, and understanding patterns in biases between techniques is crucial if outputs are to address key objectives optimally. Often biases are not consistent across natural environmental gradients such as depth, leading to uncertainty in interpretation of results. Recently there has been much interest in mesophotic reefs (reefs from 30-150 m depth) as refuge habitats from fishing pressure, leading to many comparisons of reef fish communities over depth gradients. Here we compare fish communities using stereo-video footage recorded via baited remote underwater video (BRUV) and diver-operated video (DOV) systems on shallow and mesophotic reefs in the Mesoamerican Barrier Reef, Caribbean. We show inconsistent responses across families, species and trophic groups between methods across the depth gradient. Fish species and family richness were higher using BRUV at both depth ranges, suggesting that BRUV is more appropriate for recording all components of the fish community. Fish length distributions were not different between methods on shallow reefs, yet BRUV recorded more small fish on mesophotic reefs. However, DOV consistently recorded greater relative fish community biomass of herbivores, suggesting that studies focusing on herbivores should consider using DOV. Our results highlight the importance of considering what component of reef fish community researchers and managers are most interested in surveying when deciding which survey technique to use across natural gradients such as depth.

Reef Fish Community Biomass and Trophic Structure Changes across Shallow to Upper-Mesophotic Reefs in the Mesoamerican Barrier Reef, Caribbean.

Mesophotic coral ecosystems (MCEs; reefs 30-150m depth) are of increased research interest because of their potential role as depth refuges from many shallow reef threats. Yet few studies have identified patterns in fish species composition and trophic group structure between MCEs and their shallow counterparts. Here we explore reef fish species and biomass distributions across shallow to upper-MCE Caribbean reef gradients (5-40m) around Utila, Honduras, using a diver-operated stereo-video system. Broadly, we found reef fish species richness, abundance and biomass declining with depth. At the trophic group level we identified declines in herbivores (both total and relative community biomass) with depth, mostly driven by declines in parrotfish (Scaridae). Piscivores increased as a proportion of the community with increased depth while, in contrast to previous studies, we found no change in relative planktivorous reef fish biomass across the depth gradient. In addition, we also found evidence of ontogenetic migrations in the blue tang (Acanthurus coeruleus), striped parrotfish (Scarus iserti), blue chromis (Chromis cyanea), creole wrasse (Clepticus parrae), bluehead wrasse (Thalassoma bifasciatum) and yellowtail snapper (Ocyurus chrysurus), with a higher proportion of larger individuals at mesophotic and near-mesophotic depths than on shallow reefs. Our results highlight the importance of using biomass measures when considering fish community changes across depth gradients, with biomass generating different results to simple abundance counts.