Frontiers | First evidence of live mesophotic coral reefs on the Benin continental shelf

First evidence of live mesophotic coral reefs on the Benin continental shelf

• Coffi Gérard Franck Zinzindohoué ^1*

• Houangninan Midinoudéwa ¹

• Bôla-Nlé Christelle Akondé ¹

• Zacharie Sohou ¹

• Ursula Schauer ²

• Björn Fiedler ³

• Kanna Rajan ⁴

• Jonatha Giddens ⁵

• 1. Institut de Recherches Halieutiques et Océanologiques du Bénin (IRHOB), Cotonou, Benin

• 2. Alfred Wegener Institute Helmholtz Center for Polar and Marine Research, Bremerhaven, Germany

• 3. GEOMAR Helmholtz Centre for Ocean Research, Kiel, Germany

• 4. Faculty of Engineering, University of Porto, Porto, Portugal

• 5. Exploration Technology Lab, National Geographic Society, Washington, DC, United States

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Abstract

Mesophotic coral ecosystems (MCE) remain poorly documented along the West African coastline, particularly on the continental shelf of the Gulf of Guinea. Coral reef barriers were reported off the coast of Benin during studies conducted in 1963-1964, but their ecological status remained unknown and they were presumed dead. Between April and December 2025, exploratory marine surveys combining side-scan sonar mapping and underwater video exploration were conducted to re-evaluate these historical data. Over a surveyed west-east extent of approximately 7 km, side-scan sonar data revealed two spatially distinct zones of elevated acoustic backscatter, interpreted as consolidated hard-bottom structures. Visual observations conducted at selected sonar targets documented living coral communities at depths of 54 m on rocky substrate at discrete locations within the surveyed area along the Benin continental shelf. The assemblage was dominated by octocorals, forming a mesophotic coral garden rather than a structuring scleractinian reef. At least six octocoral morphotypes and two black coral taxa, and eight associated reef fish species were observed from imagery. These observations provide the first confirmed visual evidence of living mesophotic coral communities off Benin and, to our knowledge, the first documented occurrence on the Gulf of Guinea continental shelf. More broadly, this study contributes baseline information for a region where mesophotic ecosystems remain largely unexplored and highlights the need for broader acoustic mapping, quantitative surveys, and physical sampling to better understand the extent and ecological significance of these habitats.

Introduction

Mesophotic coral ecosystems (MCEs) are light-dependent communities typically occurring at depths beyond those commonly accessible through standard recreational diving and extend to the lower limits of reef-building corals constrained by light availability and temperature (). This depth, commonly reaching approximately 150 m in clear tropical waters, defines the lower photic zone where reef growth remains possible (; ). Ecologically, MCEs function as transitional systems between shallow reefs and deeper benthic habitats, often harboring distinct species assemblages and environmental conditions compared to shallow-water counterparts (; ).

Despite increasing recognition of their ecological and climatic significance, mesophotic reefs remain poorly documented and constitute one of the least explored components of tropical and subtropical marine biodiversity (). Particularly in the Gulf of Guinea, coral reef research across both shallow and deep habitats is extremely limited, and mesophotic coral communities remain largely undocumented. This region therefore represents a major knowledge gap in global coral reef science, despite its location within the tropical Atlantic.

Coral reefs along the Benin – Togo continental shelf were first identified during surveys conducted in 1963–1964 by the Office de la Recherche Scientifique et Technique des Territoires Outre-Mer (ORSTOM, now Institut de Recherche pour le Développement). These surveys, carried out at the request of the governments of Dahomey (now Benin) and Togo, aimed primarily to map trawlable seabed and document fish diversity. During these surveys, a coral reef barrier was observed at depths of 52–56 m (), but the reefs were assumed dead and their ecological status remained unknown. Since then, no research targeting corals has been conducted on the Benin continental shelf, leaving the status, distribution, composition, and ecological significance unknown.

Between April and December 2025, the Coral Reefs Rediscovering and Exploration project in Benin (COREB) was launched to address this knowledge gap. This effort aimed to assess whether living mesophotic coral communities occur on the Benin continental shelf and to provide first ecological characterization of these communities

Methods

Study area

The study was conducted on the Benin continental shelf in the Gulf of Guinea (Figure 1). Benin’s coastline extends approximately 125 km, and the continental shelf has a predominantly sandy bottom covering roughly 2,800 km² between the 10 and 100 m isobaths with an average width of approximately 27 km ()

Survey design and data acquisition

Our study aimed at recapturing examples of coral structures which were reported by to exist at depths between 52 and 56 m on the continental shelf off Benin. The objective was to confirm the occurrence of the mesophotic coral communities to find out whether they are dead, as presumed by , or whether they are alive, and if alive, to achieve a first overview of the composition of the ecosystem. The framework of the study did not allow to conduct a quantitative assessment of benthic cover or habitat prevalence across the entire continental shelf. Our survey consisted of side-scan sonar mapping to identify potential hard-bottom features, which were subsequently investigated using underwater drone and stationary video systems for visual verification. The surveys were conducted between April and December 2025.

Side-scan sonar mapping

Side-scan sonar surveys were conducted using a DeepVision DeepEye 680 DSSP towfish side-scan sonar equipped with a dual transducer operating at 680 kHz, providing a range resolution of approximately 1 cm and a lateral range of 10–100 m on each side. The system is rated for a maximum operating depth of 100 m and was deployed from small motorboats using a 200 m lightweight tow cable. The survey area was chosen using maps of reef structures published by . Survey track lines were run parallel to the coastline to ensure consistent coverage of the continental shelf. Data acquisition was conducted at a vessel speed of approximately 1.5 knots. Georeferencing of sonar data was achieved using the integrated DeepVision GPS system based on a u-blox positioning chip, which synchronized geographic coordinates with acoustic records to accurately locate seabed features identified during the survey.

A total of 35 side-scan sonar transects were conducted between April and May 2025. Individual transect lengths ranged from 4 to 1,260 m, resulting in a cumulative survey distance of 11.5 km. Surveys were conducted using 150 m and 200 m swath widths, with the majority of high-quality acoustic returns obtained using the 150 m range setting, which provided higher spatial resolution. Sonar data were processed to identify potential hard-bottom features based on elevated acoustic backscatter, structural relief, and seabed texture.

Underwater drone surveys

Targets interpreted as potential hard-bottom features were investigated using a Qysea Fifish V-EVO underwater drone, equipped with a 4K (60 fps) camera, dual LED lights producing 5000 lumens, and an ultra-wide 166°field of view. The drone was tethered from the surface and is rated for a maximum operating depth of 100 m. The underwater drone provides real-time depth readings based on an internal pressure sensor displayed on the operator interface during deployment. Two underwater drone deployments were conducted at the first sonar-identified target area, with survey durations of 1 h 45 s and 41 min 47 s, respectively. Due to logistical and budgetary constraints, visual inspections were limited to the first identified reef complex. The system was used to conduct targeted visual and close-range observations of hard-bottom features interpreted from side-scan sonar data.

Stationary video surveys

Benthic observations were also obtained using the National Geographic Deep Sea Camera System, a stationary baited video platform designed for deep-sea biodiversity surveys (). The system consists of a Sony Handycam FDR-AX33 in a 33 cm borosilicate glass sphere, positioned at a 45-degree downward declination for optimized benthic imaging. Illumination was provided by two Cree XLamp CMT1930 LED modules. Following the baited survey period, an internal timer activated a burn-wire release mechanism to detach a 10 kg ballast, allowing the system to ascend autonomously to the surface for recovery using VHF and Iridium satellite localization. To quantify the vertical profile and benthic environment, depth and high-resolution environmental data were logged using two integrated systems. An external sensor (RBR Ltd. RBRcoda³ T.ODO) recorded temperature and dissolved oxygen at 1-second intervals. Depth was gauged using an internal pressure sensor (Keller 7LY HP).

A total of five stationary video deployments were conducted in August 2025 at sonar-identified target locations within the study area. One deployment lasted approximately three hours, whereas the remaining four deployments lasted approximately two hours each. As the system used bait to attract mobile fauna, observations of fish assemblages may have been influenced by attraction bias toward scavenging and predatory species.

Identification of potential coral reef structures from side-scan sonar

Acoustic backscatter signatures were visually inspected for features indicative of rocky or consolidated substrates, such as high-relief structures, angular blocks, and areas of heterogeneous texture. Survey track lines were examined for consistent patterns of elevated backscatter that could correspond to coral-bearing substrates. Targets identified in this manner were subsequently prioritized for visual verification using the underwater drone and stationary video surveys.

Features identified from sonar data were subsequently ground-truthed using underwater drone and stationary video observations to confirm substrate characteristics and the presence of benthic biological communities

Coral and fish identification

Coral taxa were provisionally identified from high-resolution video imagery using a combination of morphological characteristics, including colony growth form, branching pattern, colony architecture, apparent skeletal rigidity, and polyp arrangement visible in the footage. Morphotypes were defined as visually distinct colony forms that were consistently recognizable across multiple video frames and observations. No physical specimens were collected. Identification at the species level was considered preliminary due to the absence of skeletal examination and genetic analysis; therefore, identifications are reported at the lowest reliable taxonomic level, based on visual assessment alone. Fish species were also identified from high-resolution videos and still images, based on external morphological characteristics and color patterns.

Results

Environmental conditions during visual surveys

To analyze the bottom-water conditions of the coral habitat, environmental data from five Deep Sea Cam deployments were used when the system was deployed near the seabed between 50 and 60 m depth. The total of the bottom-water temperatures recorded ranged from 18.9 to 25.9 °C. The dissolved oxygen saturation values ranged from 44.5 to 92.5%, and the dissolved oxygen concentrations varied between 104.4 and 197.7 µmol L⁻¹.

Comparing the temperature and oxygen measurements of the bottom-mounted Deep Sea Cam with one of the rare CTD profiles off Benin taken on a French cruise in 2011 () reveals that the corals grow within the thermocline which explains the large variation of the Deep Sea Cam data. Also, the oxygen profiles show a steep decline with depth. Our lowest temperatures recorded by the Deep Sea Cam correspond to profile values at 80 m, while the shallowest values compare to profile CTD data values at a depth at 40 m.

Side-scan sonar observations

Over a west–east extent of approximately 7 km, side-scan sonar analysis revealed two spatially distinct areas of elevated acoustic backscatter (Figure 2). These features appeared as irregular high-reflectivity patches contrasting with the surrounding low-backscatter sandy seabed. The identified acoustic signatures were characterized by heterogeneous texture and localized high-amplitude backscatter, consistent with consolidated or hard-bottom substrates. These two areas represent discrete zones of elevated backscatter within an otherwise predominantly low-reflectivity sandy shelf environment.

These two areas were interpreted as consolidated hard-bottom structures potentially associated with mesophotic coral habitats. The Area 2 located approximately 5.4 km west of Area 1, suggesting that hard-bottom habitats within the study area may occur as spatially discrete patches. At Area 1, some of the detected hard-bottom structures measured 208 m in length, with elevations reaching up to 7.4 m in certain areas. At Area 2, one structure measured 275 m in length and reached a maximum elevation of 10.3 m. Measurements were obtained using the distance and height measurement tools integrated into the DeepView software, which is used for the processing and analysis of DeepVision sonar data. The detected features occurred at depths between 52 and 56 m, based on depth readings obtained from sensors embedded in the Deep Sea Cam and the underwater drone, which closely matched the depths reported during the surveys conducted in the 1960s (). Acoustic targets identified from side-scan sonar data were subsequently investigated using underwater drone and the stationary baited Deep Sea Cam.

Coral community composition

Visual surveys revealed living coral communities at depths of 54 m on consolidated rocky substrate along the Benin continental shelf. The assemblage was dominated by octocorals (soft corals), forming a mesophotic coral garden (Figure 3A) rather than a framework-building scleractinian reef. At least six distinct octocoral morphotypes were observed (Figures 3C, D), including monospecific, fan-shaped, and planar growth forms.

In addition, at least two morphologically distinct black coral (Antipatharia) taxa were recorded. Gorgonian sea fans (Figure 3C) were locally abundant and structurally prominent within the surveyed area. Framework-building scleractinian corals appeared to be rare and limited to small colonial forms with pale (whitish to violet) tissue. No substantial carbonate framework indicative of active scleractinian reef accretion was evident in the imagery. Coral colonies were attached to large, sub-angular rocky blocks and consolidated hard substrate, which formed the physical foundation of the coral garden (Figure 3B).

Dead octocoral colonies were documented in some areas, with visible post-mortem colonization by hydroids and serpulid (Figure 3E). Due to the absence of physical sampling, species-level identifications remain provisional and are reported at the lowest reliable taxonomic level based on visual characteristics

Reef-associated fish community

Video surveys documented the presence of reef-associated fish communities living in and surrounding the mesophotic coral garden. A total of eight fish species were identified from high-resolution videos: Lutjanus fulgens, Myripristis jacobus, Pseudupeneus prayensis, Chromis cadenati, Chromis limbata, Holacanthus africanus, Acanthurus monroviae, and Chaetodon robustus (Figure 4). These taxa were observed in close proximity with coral structures, using the habitat for shelter and feeding. Although no estimates of abundance were made, visual observations suggest that the habitat supports a diverse reef-associated fish assemblage.

Discussion

This study provides the first evidence of living mesophotic coral communities on the Benin continental shelf. More than sixty years after the work of , who reported coral remains along the Benin-Togo shelf and assumed them to be dead, we revisited parts of the same general area using side-scan sonar and targeted visual surveys

The sonar data revealed two main zones of elevated acoustic backscatter within an otherwise soft-sediment dominated shelf. These features correspond to irregular high-reflectivity patches associated with sub-angular rocky blocks and harder substrates. Their origin remains uncertain and they may represent lithified outcrops or remnants of older reefal structures, as previously suggested in the historical surveys. The interpretation of these structures as “dead reefs” in the 1960s likely reflects the limited observational tools available at the time, when analyses relied on dredging records and acoustic returns without direct visual confirmation of benthic communities. Whether the present observations reflect long-term persistence of mesophotic coral communities or changes in ecological conditions since the 1960s cannot be determined from the available data and requires further sampling to clarify their geological origin and ecological history.

Based on the surveyed area, the hard-bottom structures do not form a continuous feature but occur as spatially discrete patches within a sediment-dominated shelf. The idea of a long, uninterrupted coral barrier along the shelf, as initially suggested in earlier reports, is not supported by our acoustic data. That said, our survey only covered a limited portion of the originally described reef area, so we cannot exclude that other structures exist beyond the surveyed tracks. These spatially discrete hard-bottom features provide the physical framework within which benthic communities occur.

Our findings of mesophotic coral communities on isolated hard-bottom structures within the sedimentary shelf off Benin are consistent with observations from other mesophotic ecosystems, including Puerto Rico (), the Abrolhos Shelf (), the Brazilian Equatorial Margin (), and offshore Amazon systems (). Although the Benin shelf differs as a narrow, sediment-dominated system influenced by seasonal upwelling and freshwater inputs from coastal lagoons and river systems, these comparisons suggest that suitable hard substrate is the primary control on mesophotic habitat occurrence. This supports the view that mesophotic hard-bottom habitats may be more widespread across tropical Atlantic shelves than currently documented.

Within this habitat context, bottom-water temperatures ranged from 18.9 to 25.9 °C within the depth interval of coral occurrence. These values are higher than the 16-18 °C range observed at comparable depths (50–60 m) off Cotonou during August, where documented strong seasonal bottom-water temperature variation associated with the development of a pronounced thermocline and the upward displacement of cooler subsurface waters. However, the present dataset is limited temporally and does not allow assessment of seasonal variability or longer-term environmental changes. Nevertheless, these measurements provide the first in situ environmental observations associated with mesophotic coral communities on the Benin continental shelf.

Visual surveys at selected sonar targets confirmed the presence of living mesophotic coral communities at around 54 m depth. These were dominated by octocorals, with relatively few framework-building scleractinians, and are best described as a mesophotic coral garden developing on consolidated rocky substrate. These observations provide additional evidence for structured mesophotic habitats in the eastern tropical Atlantic. They also represent, to our knowledge, the first confirmed living mesophotic coral ecosystem on the Benin continental shelf, and one of the very few records in the wider Gulf of Guinea region, outside São Tomé ().

Whether this apparent scarcity reflects real ecological patterns or sampling bias remains an open question. Given how little exploration has been done in this region, both explanations are plausible. This study was exploratory, designed to detect and confirm rather than quantify mesophotic communities cover or to produce spatially exhaustive habitat maps

The combination of multiple octocoral morphotypes, black corals, and associated reef fish species indicates that these habitats are biologically active systems rather than isolated occurrences. The occurrence of reef-associated taxa such as Lutjanus fulgens, previously reported from mesophotic coral habitats near São Tomé and Príncipe (), and Chromis cadenati recognized among fish taxa associated with mesophotic ecosystems in the tropical Atlantic (), further supports the ecological affinity with other eastern tropical Atlantic mesophotic systems All identifications are based on imagery alone and should be considered provisional until confirmed through physical sampling.

Given increasing human pressure on continental shelf environments, including bottom-contact fishing activities, these findings highlight the need to include mesophotic habitats in regional assessments and marine spatial planning. Future work should combine broader acoustic mapping, systematic video transects, and physical sampling to constrain spatial extent, structure, and origin of these coral-associated systems, focusing on (i) geological origin and age of substrates, (ii) connectivity of coral communities across the continental shelf, and (iii) vulnerability to anthropogenic pressures in the Gulf of Guinea.

Statements

Data availability statement

The original contributions presented in the study are included in the article/supplementary material, further inquiries can be directed to the corresponding author/s

Ethics statement

Ethical approval was not required for the study involving animals in accordance with the local legislation and institutional requirements because ethical approval was not required because the study was entirely non-invasive and observational

Funding

The author(s) declared that financial support was received for this work and/or its publication. This research was funded by the National Geographic Society (Grant No. EC-115084R-24). National Geographic Exploration Technology Lab supported this research by providing access to specialized equipment

Acknowledgments

The authors thank the crews of the vessels Piranha, Confiance, and Aurel for their professionalism and invaluable support during offshore operations. We thank Sven Andes, Senior Oceans Specialist at National Geographic for his support and help during this project. We are grateful to André Freiwald and Lydia Beuck (Senckenberg am Meer, Wilhelmshaven, Germany) and Claudia Wienberg and her team (MARUM – Center for Marine Environmental Sciences, Germany) for their expertise in coral image-based identification, and to Rui Freitas (Universidade Técnica do Atlântico, Cabo Verde) and Peter Wirtz (Portugal) for assistance with fish identification. We also thank Ingo Klaucke (GEOMAR Helmholtz Centre for Ocean Research, Germany) for his guidance during side-scan sonar data acquisition and interpretation. We acknowledge the national authorities of Benin, including the Institut de Recherches Halieutiques et Océanologiques du Bénin (IRHOB) and the Autorité Nationale Chargée de l’Action de l’État en Mer (ANCAEM), for their support in facilitating the implementation of offshore research activities. Finally, we are also grateful for the constructive comments from the reviewers, which helped clarify and improve the article.

Conflict of interest

The author(s) declared that this work was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest

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Summary

Keywords

Benin continental shelf, eastern tropical Atlantic, Gulf of Guinea, mesophotic coral ecosystem, octocorals, side-scan sonar, underwater drone

Citation

Zinzindohoué CGF, Midinoudéwa H, Akondé B-NC, Sohou Z, Schauer U, Fiedler B, Rajan K and Giddens J (2026) First evidence of live mesophotic coral reefs on the Benin continental shelf. Front. Mar. Sci. 13:1848226. doi: 10.3389/fmars.2026.1848226

Received

05 April 2026

Revised

09 June 2026

Accepted

10 June 2026

Published

20 July 2026

Volume

13 – 2026

Edited by

Luigi Jovane, University of São Paulo, Brazil

Reviewed by

Nazik Öğretmen, Aarhus University, Denmark

Senem Tarantini, University of Salento, Italy

Updates

Copyright

© 2026 Zinzindohoué, Midinoudéwa, Akondé, Sohou, Schauer, Fiedler, Rajan and Giddens

This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

Disclaimer

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article or claim that may be made by its manufacturer is not guaranteed or endorsed by the publisher

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