Review Article, J Mar Biol Oceanogr Vol: 15 Issue: 2
Marine Biodiversity of São Tomé and PrÃncipe: Geographic, Climatic, Oceanographic, and Biogeochemical settings
Maria Emilia Cunha1*, Armando J Almeida2, Wadmilton Fernandes3 and Hugulay Maia3
1Centro de Ciências do Mar do Algarve (CCMAR/CIMAR LA), Universidade do Algarve, 8005-139 Faro, Portugal
2Centro de Ciências do Mar e doAmbiente (MARE), Portugal e Departamento de Biologia (Bloco C2), Faculdade de Ciências, Universidade de Lisboa, 1749-016 Lisboa, Portugal
3Faculdade de Ciências e Tecnologias da Universidade de São Tomé e PrÃncipe (FCT-USTP), Quinta de Santo António. C.P. 546 São Tomé, São Tomé e PrÃncipe
- *Corresponding Author:
- Maria Emilia Cunha
Centro de Ciências do Mar do Algarve (CCMAR/CIMAR LA), Universidade do Algarve, 8005-139 Faro, Portugal
E-mail: mecunha@ualg.pt
Received: 23-March-2026; Manuscript No: JMBO-26-186751; Editor Assigned: 26-March-2026; PreQC Id: JMBO-26-186751(PQ); Reviewed: 13-April-2026; QC No: JMBO-26-186751(Q); Revised: 21-April-2026; Manuscript No: JMBO26-186751(R); Published: 30-April-2026, DOI: 10.4172/jmbo.1000343
Citation: Cunha ME, Almeida AJ, Fernandes W, Maia H (2026) Marine Biodiversity of São Tomé and PrÃncipe: Geographic, Climatic, Oceanographic and Biogeochemical settings. J Mar Biol Oceanogr 15: 343
Abstract
Aim: To understand and describe the oceanic conditions that influence marine biodiversity in the São Tomé and Principe archipelago.
Methods: Review of published and unpublished research publications on marine biodiversity, ocean productivity and oceanography in São Tomé and Principe and the nearby regions.
Results: The geographic location of the oceanic islands of São Tomé and Principe, in the Gulf of Guinea, strongly determine its climate, oceanic circulation and ultimately marine biodiversity. The climate is controlled by the seasonal displacement of the Intertropical Convergence Zone (ITCZ) that also controls the year-round oceanic biological production with two seasons: a cold upwelling season from July to September marked by cooling and nitrate enrichment of the surface waters and a predominant warm rainy season (from October to June) characterized by nutrients input by runoff. Despite being geographically isolated, these islands have a relatively low level of marine endemism although high species richness. Many shallow-water marine fish and invertebrate species and genera found here are also present in Cape Verde Islands and Tropical Western Atlantic. This similarity is largely attributed by the authors to the strong oceanographic connections with the coast of African and the Tropical Western Atlantic, facilitated by zonal equatorial currents like the Guinea Current and the subsurface Equatorial Undercurrent.
Main conclusion: Marine biodiversity in São Tomé is strongly influenced by the ITCZ and the oceanographic connectivity within the Tropical Eastern and Western Atlantic.
Keywords: Marine Biodiversity; São Tomé and Principe; Eastern Tropical Atlantic; Gulf of Guinea; Bay of Biafra; Biological Productivity; Oceanography; Climate; ITCZ; Biogeography
Introduction
Global change is significantly impacting Earth’s biodiversity [1]. As species population’s decline and extinction risks increase worldwide [2], it is crucial to identify sites critical for the conservation of biodiversity. São Tomé and Príncipe, situated in the tropical eastern Atlantic along the Cameroon Line, possess geographic isolation and diverse environmental conditions that have fostered distinctive marine life with tropical origins and high species richness. The Gulf of Guinea has been identified as one of the world’s marine biodiversity hotspots, ranking within the upper two-thirds in terms of risk from human activities [3]. Overfishing and other threats to species populations continue to deplete resources and alter ecosystems. Despite recent advances, including the recognition of São Tomé and Príncipe as a UNESCO World Biosphere Reserve in 2025, a gap persists between available scientific knowledge and its integration into public policies. To make informed decisions that align with global biodiversity commitments, such as those outlined in the Global Biodiversity Framework (GBF), there is a need to establish a robust monitoring system that tracks progress towards conservation goals and enhance accountability in biodiversity management based on knowing what, where, and when species are present. In addition, the data need to be placed, and/or mapped, into an environmental context.
A comprehensive synthesis of existing scientific knowledge is both timely and essential, not only to consolidate fragmented data but also to inform evidence-based conservation and foster future research agendas. Through this synthesis, we seek to position São Tomé and Príncipe within the broader context of the Tropical Atlantic Island biogeography and to highlight its role as a keystone system for understanding marine diversity in tropical oceanic environments. Extensive information exists regarding species biodiversity, yet there is limited understanding of the environmental conditions that support their thriving populations.
A review of published and unpublished research related to the regional marine biodiversity was used in the study [Annex I]. Five hundred and sixty-eight (568) entries in total were recovered and the documents are now available for consultation at the Fisheries and Aquaculture Directorate (Direcção das Pescas e Aquacultura) of São Tomé and Principe. Of these, 69% refer to the presence of taxonomic groups or species. Most examined specimens belonged to Phylum Chordata (56%), but Mollusca (17%) and Arthropoda (13%) (mainly Crustacea) also attracted considerable interest. Among the Chordata, particular emphasis was placed on birds (45%) and fish (37%).
The present paper provides a summary of research conducted in São Tomé and Principe concerning marine biodiversity, regional marine biogeography, and the oceanographic factors that have influenced the development of its biodiversity.
Geographical setting of São Tomé and Principe
São Tomé and Príncipe are two volcanic islands located right north of the equator in the eastern part of the Gulf of Guinea [Figure 1]. It is a small island developing nation consisting of two principal islands and several smaller islets, all of which are part of the Cameroon Volcanic Line. This geological line is aligned in a southwest direction extending from Cameroon on the African continent through the continental island of Bioko into the ocean. It includes the oceanic Principe Island, situated approximately 220 km southwest of Bioko and formed around 31 million years ago; São Tomé, located 146 km from Príncipe is younger with an estimated age of 13 million years old; and Annobón, the youngest island in the chain with 4.8 million years, and is positioned 180 km from São Tomé [4]. Located at the east inner most part of the Guinea Gulf, the two islands are true oceanic formations separated among them and from the African mainland by depths exceeding 3000 meters. The coast is rocky and very steep, with a shelf break on both islands around 80 – 100 m depth and bottom depths typically deeper than 1000 m off the shelf [5]. The emerged portion of the island of São Tomé with its southern Rola islet (836 km²) is much larger than Príncipe (130 km²), but if measured from the shelf break at 100 m depth, Príncipe is larger. São Tomé shelf covers an area of 432 km² that in the north-western quarter of the island is very narrow reaching up to 1.2 km from the coastline, whereas in the north-eastern part, the shelf extends as far as 5.4 km. Príncipe’s shelf area covers 1,097 km², with the 100 m isobaths extending 7 km north and 27 km south to the Tinhosas islets. The shapes of the two islands, considering the limit of the 100 m isobaths, are remarkably similar [Figure 1].
São Tomé and Príncipe feature a few large river basins with dense networks [Figure 2]. The watersheds are numerous and small. Their areas range from 4 to 111 km², developing around watercourses that have a length between 5 and 27 km and an elevation difference between highest points and sea level of 1000 to 1500 meters [70].
Climatological and Oceanographic overview
Climatology
The tropical Atlantic (30° N and 30° S), exhibits northeasterly and southeasterly trade winds converging into an Intertropical Convergence Zone (ITCZ) north of the equator, all of which display a seasonal cycle that dominates the ocean-atmosphere signal in the tropical Atlantic [7] [Figure 3a]. The ITCZ is a narrow band marked by warm temperatures above 27°C and upward movement of moist air. As air ascends and undergoes cooling, water vapor condenses, resulting in a global belt of substantial precipitation. The cooled, elevated air subsequently moves toward the poles from the ITCZ, descends, and produces drier clear weather conditions on the opposite side resulting in atmospheric cells. This is referred to as Hadley cells, one in the North and the other in the South of the Equator [Figure 3b]. In the Atlantic Ocean, the ITCZ is always found several degrees north of the geographical equator, slightly tilting northward toward the eastern basin. The ITCZ exhibits seasonal variation in its location, typically trailing the sun by approximately one month. During the northern hemisphere summer, it shifts northward, while in winter, it moves southward. The northward migration results in a variable wind forcing field across the tropical Atlantic region because the sea- surface air pressure difference between the north (warm) and south (cooler) drives an unusually strong wind from the southeast [8] trade.
Stronger winds from the south increasingly cool the southern water and help to push the ITCZ northward bringing clear skies and dryer weather to the south.
The climate in São Tomé and Príncipe typically reflects the location of the ITCZ. There is a warm rainy season from November to May, and a cold season from June to October. The northernmost position of the ITCZ is reached in July-August, corresponding to the peak of cooler and dryer season. During April-May, which corresponds to the heavy rainy season, the ITCZ is at its southernmost position and just north of the equator.
Oceanographic settings
The upper-level geostrophic circulation in the Atlantic Ocean, south of the 10°N, is dominated by a system of gyres and by the Equatorial and Circumpolar Current Systems. [Figure 4] provides a schematic representation of the large-scale, upper-level geostrophic circulation in the region. The Subtropical gyre, from which the Benguela Current integrates the eastern limb of the South Equatorial Current moving westwards, is located mostly south of latitude 20°S. The Benguela Current moves away from the African territory near 30°S and heads north-westwards towards the Brazilian coast where the most part turns south integrating the Brazilian Current and some form the north Brazil Current that moves north along the coast [9]. Centred at around 10°S and 9°E there is a subsurface prominent feature during the southern hemisphere summer (January-April), the Angola dome. It is a cold-water dome off Angola with a cyclonic circulation. Some of the water from South Equatorial Current retroflex close to the Brazilian tip at around 5°S giving origin to the South Equatorial Counter Current that moves eastwards. The Antarctic Circumpolar Current flows from west to east around Antarctica.
Details of the system of equatorial currents, located between 5°S and 5°N is shown in [Figure 5]. This is the main system influencing the region of São Tomé and Principe and is marked by variability in time and space. The equatorial ocean responds strongly to changes in the overlying wind fields because the Coriolis parameter vanishes in the presence of density stratification. Fluctuations of current velocity in response to seasonal changes in equatorial wind stress fields are therefore mainly zonal along the equatorial wave guide. The hydrographic regime of the waters of the São Tomé and Príncipe region is primarily influenced by three major zonal currents. Two are surface currents: the Guinea Current (GC) and the northern branch of the South Equatorial Current (nSEC), and the third is subsurface: the Equatorial Undercurrent (EUC) [Figure 5]. The nSEC is separated from the GC by the northern tropical convergence along 3°N. The Guinea Current is mainly supplied by the North Equatorial Countercurrent (NECC) in July, and a southern branch of the Canary Current (sCC) in January [11]. The current moves eastward along the African coast from Senegal spilling into the Bay of Biafra, where the STP archipelago is located. The Guinea Current is not very deep on average, extending from the surface to 15 m near the coast and to 25 m offshore. The GC flows east at depths of around 20−50 m at about 3° − 5°N (São Tomé and Príncipe are located around 0° and 2°N) and exhibits seasonally varying velocities ranging from 0.2 to 0.6 m s−1. At the Bay of Biafra, and due to the barrier presented by the African continental mass, a part of the water retroflects. The Equatorial Undercurrent (EUC) is a strong subsurface current that crosses west to east the entire Atlantic at the equator level and slopes progressively upward to the east. The EUC is a permanent, stable current, originated in the western part of the Atlantic Ocean, in northern Brazil. The Coriolis force (the Coriolis parameter becomes very small, going to zero at the equator) keeps the EUC centred close to the equator. The EUC has a width of around 200 km, a thickness of 150 m and maximum speed within the core of up to 0.5−1.25 m s−1. This current is present at a depth of 50 to 125 m along the entire equator and feeds surface currents off the African coast. It is clearly visible with a core at about 100 m depth associated with maxima in salinity and oxygen and temperatures around 20°C, which is about 5°C cooler than the surface waters in the equatorial region. During its way to the east the flow shallows and weakens. A numerical study by [13] indicate that in the Gulf of Guinea the remainder of the EUC splits into two poleward branches after impinging upon the African continent at São Tomé Island thereby impacting the surface water characteristics of the southern Gulf of Guinea.
Figure 5: Schematic map showing the horizontal distribution of the major eastern tropical currents. Shown are the North Equatorial Counter Current (NECC), the Guinea Current (GC), the South Equatorial Current (SEC) with the northern (nSEC) and equatorial branches (eSEC), the Equatorial Undercurrent (EUC), the Gabon-Congo Undercurrent (GCUC), the Angola Current (AC), the South Equatorial Undercurrent and the South Equatorial Counter Current (SECC) [Adapted from [8]). Darker blue arrows indicate to sub-surface currents; lighter blue arrows are for surface currents].
Biogeochemical settings
The Bay of Biafra is characterized by thermal stability and a strong pycnocline, with warm and low salinity waters (25°-29°C of temperature and 33-34 of salinity), the Tropical Surface Water, strongly influenced by rainfall, drainage and runoff of the Niger river [14,15] overlying high salinity Subtropical Underwater (19°-28°C, 35-36.5 salinity) [8]. The thermocline is close to the surface (20-50 m).
The year-round production is driven by two kinds of enrichment linked to the seasonal displacement of the ITCZ: 1. a cold upwelling season from July to September marked by cooling and nitrate enrichment of the surface of the water supply and 2. a predominant warm rainy season (from October to June) characterized by the existence, of a thin surface layer of warm, and low salinity water from runoff. During July to September and due to the intensification of the southeasterly trade winds [16,17] that push the ITCZ northward [18,19], the equatorial near surface mixing is intensified and there is upwelling of cooler waters and nitrate enrichment of the surface of the water along the equator [20,21]. The surface extension of the upwelled waters is limited to the north by a thermal front where they plunge under the less salty waters of the Gulf of Guinea creating the Equatorial Upwelling Front [22]. The intensification of the equatorial upwelling is the result of the dynamic uplifting of the thermocline (and the nutricline and associated subsurface chlorophyll maximum) in the eastern tropical Atlantic in response to the increasing easterly winds in the western Atlantic [23, 24]. During the rainy season (October to June), the thermal ridge has two layers divided by the nitracline. The upper layer’s primary production is depleted of nutrients, while the deeper layer is limited by light. Production and chlorophyll are maximum at the top of the nitracline whose level is determined by the level of the maximum vertical gradient of density that screens the diffusion of nutrient salts. During periods of upwelling, the density barrier destroys the two-layer system and most of the production is located at the surface.
Another significant oceanic consequence of the ITCZ in enhancing the regional primary productivity is the development of the Atlantic Cold Tongue (ACT) [16, 25, 26] by the action of southeasterly Trade Winds. The ACT is a large pool of locally cooler ocean surface water (down to 23°C), appearing every year in boreal spring and summer, on and slightly south of the equator and extending to approximately 20°W. This phenomenon is primarily caused by the upwelling of deep cold waters that enhance biological production by nitrate supply [27, 28]. Because it is centred slightly south of the equator it affects the ecosystem of the region, from primary production to high trophic levels [27–31].
The Tropical Atlantic Marine Biogeography
Marine bio geographers divide the biosphere into major provinces or regions by identifying unique patterns of endemism [32–34]. Each province features a particular group of species found only in certain geographic areas called areas of endemism. Provinces are defined by natural barriers that separate them [33,35–38], as well as by considering their origins and how they are interconnected [34]. According to these researchers, the tropical Atlantic reef fauna consists of four primary provinces: Brazil, the Greater Caribbean, the Mid-Atlantic Ridge, and the Eastern Atlantic. Five major barriers distinguish these provinces [33]: the first is the Atlantic Ocean, which stretches over 3,500 km; the second is the Red Sea land bridge, separating Atlantic and Indian Ocean tropical faunas; the third is the freshwater discharge from the Orinoco and Amazon rivers, spanning 2,300 km along South America’s northeast coast between Brazilian and Caribbean reefs; the fourth is the Isthmus of Panama, which splits the Caribbean and Tropical Eastern Pacific; and the fifth is the cold Benguela Current that keeps the Atlantic’s tropical reef fauna apart from the southern Indian Ocean. Those geographic and oceanographic factors act as essential barriers or filters responsible for divergence and speciation among the Atlantic’s regional faunas.
Advances on the use of remote sensing methods to estimate oceanic primary production [39-44] made possible to focus on pelagic environments. The oceans were then classified into biomes and biogeochemical provinces based on oceanographic data and global chlorophyll profiles [45-46]. This system is regarded as one of the most thorough ways to categorize pelagic life, although its usefulness is limited when applied to the complex dynamics of coastal waters. According to the biogeochemical classification, São Tomé and Príncipe islands lie at the intersection of the Eastern Tropical Atlantic Province (ETRA) within the Atlantic Trade Wind Biome and the Guinea Current Coastal Province (GUIN) in the Atlantic Coastal Biome [46]. The author of this classification specifies that the series of smaller islands stretching from Annobón to Fernando Pó, which include São Tomé and Príncipe, in the Bay of Biafra are considered part of the ETRA province [46].
An ecosystem-based approach to managing marine fisheries has been recommended by various researchers [47–53]. This approach stresses the importance of using well-defined, place-specific ecosystem units for evaluating and managing marine environments. One practical example is the Large Marine Ecosystem (LME) strategy, developed over several years by experts and presented in several books and chapters [54–56]. LMEs are defined as large areas - typically at least 200,000 km² - that have distinct bathymetry, hydrography, productivity, and interconnected populations (www.lme.noaa.gov/Portal/). Those LMEs provide a framework for addressing transboundary management challenges related to fisheries, pollution, habitat restoration, productivity, socioeconomic concerns, and governance. The system mainly focuses on coastal and shelf seas, as well as adjacent ocean regions, with an emphasis on biological productivity and oceanographic processes. Within the framework referred in [54-56], São Tomé and Príncipe integrates the Guinea Current Large Marine Ecosystem (GCLME), one of the Large Marine Ecosystem (LME) of the world’s oceans [57] and is in the subsystem of the Eastern Gulf of Guinea. However, this is a classification used to distinguish regions of ocean space encompassing coastal areas out to the seaward boundary of continental shelves, and the outer margins of coastal current systems and the islands of São Tomé and Príncipe are mostly oceanic.
From a biogeographical standpoint, São Tomé and Príncipe is part of the Gulf of Guinea Islands Ecoregion, within the Gulf of Guinea Province according to the Marine Ecoregion of the World (MEOW) system [58]. This region has a relatively uniform species mix that stands out from neighbouring systems. The types of species found here are influenced by just a few ecosystems or unique oceanographic and topographic features. Although these islands cover a small area, they offer a remarkable variety of habitats, including subtidal rocky reefs, rhodolite beds, marginal coral groups, and soft-bottom communities. Extensive information exists regarding species biodiversity, yet there is limited understanding of the environmental conditions that support their thriving populations.
Marine Biological Diversity of São Tomé and Principe
Research into marine biodiversity on the islands of the the Gulf of Guinea began in the late nineteenth and early twentieth centuries [Figure 6], with naturalists Balthazar Osório and António Nobre concentrating on fish, crustaceans, and gastropods, while José Vicente Barbosa du Bocage focused on birds. After almost forty years of very low activity, research resumed in the 1960s but it was only in the nineties that interest exploded. Since then, many studies and expeditions have taken place on São Tomé and Príncipe.
Numerous monographs and field guides have been published that explore marine biodiversity on the islands of São Tomé and Príncipe. Publicaly available information on marine biodiversity, highlighting its value for ongoing research and conservation initiatives is presented in [59]. This document acts as a catalogue of marine species present on São Tomé and Príncipe but does not cover their ecology or biology. Such works are crucial for systematically understanding and analysing the diversity and distribution of marine life at a national level.
A phylogeographer review analyzing warm-water reef-fish genera in the Atlantic, providing insights into patterns of species distribution and relationships within the Tropical Atlantic fauna is found in [60]. The authors conducted analysis utilizing a presence/absence species distribution database to elucidate the relationships among the West Atlantic and the East Atlantic Provinces. Based on their analysis, São Tomé and Príncipe belong to the Tropical Eastern Atlantic Province (TEA) and within TEA the archipelago belongs to a subgroup that involves also the islands of Cape Verde [Figure 7]. According to their results there was 92 reef fish species shared between the Cape Verde islands and São Tomé making up roughly 95% of the shallow-water TEA fish population. The other TEA subgroup corresponded to the coastal regions of tropical West Africa (North and Tropical) that shared 90% of the population.
Figure 7: Relationships of 25 areas in the Atlantic Ocean based on maximum-parsimony analysis using the presence/absence distribution data base of 2605 reef fish species. Values at nodes represent bootstrap support (1000 replicates). [Adapted from [60]].
The TEA province is separated from the western side provinces of the Atlantic from a barrier that is the > 3500-km wide Atlantic itself and from the southern Indian Ocean by the cold Benguela Current [Figure 8]. These two geographic and oceanographic features are regarded as the primary barriers or filters producing divergences and speciation among the regional faunas of the Atlantic. There were 105 reef fish species shared between the Caribbean at NWA and Tropical Eastern Atlantic (TEA)-making up roughly 27% of the shallow-water TEA fish population. Similarly, 107 reef fish species from the Brazilian (SWA) province are found in the TEA while only 38 are shared with eastern South Africa. Most trans-Atlantic species inhabit both CA and BR areas, amounting to 112 species [60]. This distribution signifies continuing connections among the Atlantic’s warm regions
Figure 8: Biogeographical barriers and relationships among areas based on the number of reef fish species common to each pair of areas. Soft barriers: (III) Amazon discharge; (IV) Mid-Atlantic Barrier; (V) Benguela Barrier. Areas/provinces: NWA, Northwestern Atlantic (Caribbean + Bermuda + Carolinian); SWA, Southwestern Atlantic (Brazil + Brazilian oceanic islands + Argentina); MAR, Mid-Atlantic Ridge (Ascension + St Helena); TEA, Tropical Eastern Atlantic (Cape Verde and São Tomé oceanic islands+ eastern tropical coastal Africa); SWIO, Southwest Indian Ocean. Arrow widths are sized proportionally to the number of species common to the areas. The proportion of the number of species common to a pair of areas relative to its entire reef fish fauna is given at each end of the arrows. [Adapted from [60]].
[61] or suggests recent migrations within the Atlantic basin [62]. Phylogenetic evidence primarily points to these migratory events occurring from west to east [60, 63, 64]. Although the Caribbean has played a major role in shaping warm-region biodiversity, the tropical Atlantic continues to acquire new species from other regions.
A more recent study offered an updated taxonomic checklist of marine fish-including coastal, offshore, and deep-water species-in São Tomé and Príncipe (and Annobón), along with a summary of marine ecosystems, biogeography, fish evolution, and conservation [65]. These authors refer that most fish species documented from São Tomé and Príncipe are also found throughout other regions of the Gulf of Guinea, with only a limited number of species being endemic to the islands. The low level of marine endemism - just 3% for fishes [66], has been attributed to the high degree of oceanographic connectivity with the African coast. However, the studies cited in support of this statement referred to connectivity with the Tropical Western Atlantic rather than the eastern Atlantic [67].
Conclusions
The geographic location and topography of shelf of the oceanic islands of São Tomé and Principe, in the Gulf of Biafra, strongly determine its climate, oceanic circulation and ultimately its marine biodiversity. The climate is controlled by the seasonal displacement of the ITCZ that also controls the year-round oceanic biological production with two seasons: a cold upwelling season from July to September marked by cooling and nitrate enrichment of the surface waters and a predominant warm rainy season (from October to June) characterized by nutrients input by runoff. Another significant oceanic consequence of the Intertropical Convergence Zone (ITCZ) in the enhancement of the regional primary productivity, is the development of the Atlantic Cold Tongue (ACT), a large pool of locally cooler ocean surface water (down to 23°C), appearing every year in boreal spring and summer, on and south of the equator.
Despite being geographically isolated, these islands have a low level of marine endemism. Many shallow-water marine fish and invertebrates found here are also present in Cape Verde Islands and the Tropical Western Atlantic. A credible explanation for this phenomenon involves species migration/colonization facilitated by the zonal easterly and westerly flowing Equatorial currents. The Guinea Current is mainly supplied by the North Equatorial Counter Current (NECC) in July, and a southern branch of the Canary Current (sCC) in February [68] which easily explains the faunistic similarities with Cape Verde. The Guinea Current moves eastward along the African coast from Senegal pilling into the Gulf of Biafra, where the São Tomé and Príncipe archipelago is located. The pattern of oceanic circulation provides also substantial support for the trans- Atlantic transport of planktonic larvae, particularly via the Equatorial Undercurrent. Furthermore, planktonic productivity is enhanced by Equatorial upwelling, which mitigates larval starvation during transit.
The described pattern of colonization supports the Marine Centres of Speciation model proposed by [69], where biodiversity hotspots, such as the Caribbean Sea, function both as sources and recipients of new species, facilitating a mutual exchange referred to as biodiversity feedback. Due to the geographic position the Principe Island, closer to the African continent, is probable that the faunistic and floristic composition will be slightly different from São Tomé with the Principe Island hosting more coastal species than São Tomé [70].
From a marine biogeographic perspective, the review indicates that the marine biodiversity of São Tomé and Príncipe is part of the Eastern Tropical Atlantic Islands sub-province within the Eastern Tropical Atlantic province.
Acknowledgements
The authors acknowledge Projeto de Apoio à Comercialização, Marketing, Produtividade Agrícola e Nutrição (COMPRAN), co- funded by the International Fund for Agricultural Development and the Ministério do Plano, Finanças e Economia Azul da República Democrática de São Tomé e Príncipe. They also acknowledge António Jorge da Silva for critical reviews of previous drafts of the manuscript that help to shape the actual version.
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ANNEX I
RESEARCH ON MARINE BIODIVERSITY AROUND SÃO TOMÉ AND PRÍNCIPE
This list contains documents that help deepen understanding of marine biodiversity and habitats in São Tomé and Príncipe, spanning from 1551 to today. It includes references to scientific articles, brief notes in journals, reports published by national and international organisations, books, species identification guides, conference presentations, as well as expedition and research cruise reports conducted locally. Alongside these resources, a database has been created to compile all available information, aiming to make it accessible to individuals and groups interested in furthering work related to these subjects.
GENERAL
- Afonso, P., Morato, T., Santos, R.S., 2016. Marine biodiversity of oceanic islands: The São Tomé and Príncipe case study. Marine Ecology, 37(2): 173–186
- Albuquerque C., Cesarini D., Tagliabue L.C., 2009. Plano de Manejo do Parque Nacional Obô de São Tomé. ECOFAC IV, São Tomé, São Tomé e Príncipe, 117 pp
- Albuquerque, C., Cesarini, D., 2014 Plano de Manejo do Parque Natural Príncipe. 2009-2014. ECOFAC IV, São Tomé, São Tomé e Príncipe 146 pp.
- Barns, A., 1928. In Portuguese West Africa: Angola and the isles of the Guinea Gulf. Geographical Journal, 72:18-37
- Bento, M., Niza, H., Cartaxana, A., Bandeira, S., Paula, J., Correia, A.M. 2023. A Dataset of Marine Macroinvertebrate Diversity from Mozambique and São Tomé and Príncipe. Data, 8:76. https://doi.org/10.3390/data8050076
- Bento, M., Niza, H., Cartaxana, A., Bandeira, S., Paula, J., Correia, A.M., 2023. Mind the Gaps: Taxonomic, Geographic and Temporal Data of Marine Invertebrate Databases from Mozambique and São Tomé and Príncipe. Diversity 2023, 15, 70. https://doi.org/10.3390/d15010070
- Board, E., 2021. World Register of Marine Species.
- Bocage, V.B., 1889. Breves considerações sobre a fauna de São Thomé. Jornal de Sciências Mathematicas, Physicas e Naturaes, 1(1):33-36
- Bocage, J.V.B., 1903. Contribution à la faune des quatre îles du Golfe de Guinée. Jornal de Sciências Mathematicas, Physicas e Naturaes, (2)7:25–59
- Bocage, V.B., 1903. Contributions à la faune des quatre îles du Golfe de Guinée. I. Ile de Anno Bom. Jornal de Sciências Mathematicas, Physicas e Naturaes,7: 54-59
- Bocage, J.V.B., 1903. Contributions à la faune des quatre îles du Golfe de Guinée. IV. Ile de St. Thomé. Jornal de Sciências Mathematicas, Physicas e Naturaes, 7: 65-96
- Bowen, W., Rocha, L.A., Toonen, R.J., Karl, S.A., 2013. The origins of tropical marine biodiversity. Trends in Ecology & Evolution, 28 (6), 359–366. doi:10.1016/j.tree.2013.01.018
- Briggs, C., 2003. Marine centres of origin as evolutionary engines. Journal of Biogeography, 30: 1–18
- Briggs, C., Bowen, B.W., 2012. Marine biogeographic provinces: a realignment with reference to fish distributions.
Journal of Biogeography, 39: 12–30
- Briggs, J.C., Bowen, B.W., 2013. Marine shelf habitat: biogeography and evolution. Journal of Biogeography, 40 (6):1023-1035
- Brito, B.R., 2021. Conserving marine life in São Tomé and Príncipe: Concerted actions with Agenda 2030. In: W. Leal Filho et al. (eds.), Life Below Water, Encyclopedia of the UN Sustainable Development Goals, 1-16 pp. Springer Nature Switzerland AG 2021
- Buhl-Mortensen, L., Buhl-Mortensen, P., Dolan, M.J.F., Gonzalez-Mirelis, G., 2015. Habitat mapping as a tool for conservation and sustainable use of marine resources: Some perspectives from the MAREANO Programme, Norway. Journal of Sea Research, 100: 46-61
- Buhl-Mortensen, L., Myhr, A.I., Wellin, S., 2005. Carbon sequestration, the precautionary approach and the responsibility of scientists. Water Science and Technology, 6: 205-212
- Buhl-Mortensen, L., Vanreusel, A., Gooday, A.J., Levin, L., G. Priede, I.G., Buhl-Mortensen, P., Gheerardyn, H., King, N.J., Raes, M., 2010. Biological structures as a source of habitat heterogeneity and biodiversity on the deep ocean margins. 2009 Marine Ecology, 31: 21-50
- Carpenter, K.E., De Angelis, N., (Eds),2014. The living marine resources of the Eastern Central Atlantic. Volume 1: Introduction, crustaceans, chitons, and cephalopods. FAO Species Identification Guide for Fishery Purposes, Rome, FAO. pp. 1–663
- Carpenter, K.E., De Angelis, N., (Eds), 2016. The living marine resources of the Eastern Central Atlantic. Volume 3: Bony Fishes Part 1 (Elopiformes to Scorpaeniformes). FAO Species Identification Guide for Fishery Purposes, Rome, FAO. pp. 1511–2350, 2343–3124.
- Carpenter, K.E., De Angelis, N., (Eds), 2016. The living marine resources of the Eastern Central Atlantic. Volume 4: Bony fishes part 2 (Perciformes to Tetradontiformes) and Sea turtles. FAO Species Identification Guide for Fishery Purposes, Rome, FAO. pp. 2343–3124
- Carpenter, K.E., De Angelis, N., (Eds), 2016. The living marine resources of the Eastern Central Atlantic. Volume 2: Bivalves, gastropods, hagfishes, sharks, batoid fishes, and chimaeras. FAO Species Identification Guide for Fishery Purposes, Rome, FAO. pp. 665–1509
- Coelho, M.T., Barreto, E., Rangel, T.F., Diniz-Filho, F.A.F., Wüest, R.O., Bach, W., Skeels, A., McFadden, I.R., Roberts, D.W., Pellissier, L., Zimmermann, N.E., Graham, C.H., 2023. The geography of climate and the global patterns of species diversity. Nature, 622: 537-544
- Fea, 1902. S. Thomé. Ricordi ed impressioni. Bolletino della Società Geografica Italiana fasc. I, 22 pp.
- Floeter, R., Rocha, L. A., Robertson, D. R., Joyeux, J. C., Smith-Vaniz, W. F., Wirtz, P., Edwards, A. J., Barreiros,
- P., Ferreira, C. E. L., Gasparini, J. L., Brito, A., Falcón, J. M., Bowen, B. W., Bernardi G., 2008. Atlantic reef fish biogeography and evolution. Journal of Biogeography, 35: 22–47
- Floeter, S.R., Ferreira, C.E. L., Rocha, , Solé-Cava, A., Wirtz, P., Sampaio C., Gasparini, J.L.,Gascoigne, A., 2017. The evolution and ecology of a poorly known Atlantic biodiversity hotspot. Research Report Grant #7937-05, National Geographic Committee for Research and Exploration, 12 pp.
- Freitas, R., Menezes, G.M., Prieto, R., Silva, M.A. (2019). Insular marine biodiversity: Patterns and conservation challenges. Frontiers in Marine Science, 6: 124
- Froese, , Pauly., D. 2022. FishBase. World Wide Web electronic publication www.fishbase.org
- Garcia, F.J., Bertcsch, H., 2009 Diversity and distribution of the Gastropoda Opisthobranchia from the Atlantic Ocean: A global biogeographic approach. Scientia Marina, 73(1): 153-160
- Girard, A. 1895. Les explorations zoologiques des Portugais dans les iles du Golfe de Guiné. Portugal em Africa, 2 (19): 809-824
- Greathead, G., Magni, P., Vanaverbeke, J., Buhl-Mortensen, L., Janas, U., Blomqvist, M., Craeymeersch, J., Dannheim, J., Darr, A., Degraer, S., Desroy, N., Donnay, A., Griffiths, Y., Guala, I., Guerin, L., Hinchen, H., Labrune, , Reiss, H., Van Hoey, G. Birchenough, S.N.R. 2020. –– A generic framework to assess the representation and protection of benthic ecosystems in European marine protected areas. Aquatic conservation: Marine and Freshwater Ecosystems, 30(7):1253-1275
- Hachich, N. F., Bonsall, M.B., Arraut, E.M., Barneche, D.R., Leqinsohn, T.M., Floeter, S.R., 2015. Island biogeography: Patterns of marine shallow-water organisms in the Atlantic Ocean. Journal of Biogeography, 42: 1871–1882
- Henning Reiss, Silvana Birchenough, Angel Borja, Lene Buhl-Mortensen, Johan Craeymeersch, Jennifer Dannheim, Alexander Darr, Ibon Galparsoro, Mayya Gogina, Hermann Neumann, Jacques Populus, Anna M. Rengstorf, Mireia Valle, Gert van Hoey, Michael L. Zettler, Steven Degraer, 2015. Benthos distribution modelling and its relevance for marine ecosystem management. ICES Journal of Marine Science, 72: 297-315
- Henriques, J.A. 1917. A ilha de São Thomé sob o ponto da vista histórico-natural e agrícola. Boletim da Sociedade Broteriana, 27: 1-197
- Hofer, , 1991. Die Tierwelt von São Tomé. Privately published, Berne, 24 pp.
- Jones, J., 1994. Biodiversity in the Gulf of Guinea: an overview. Biodiversity and Conservation, 3(9): 772-784.
- Juste, J., Fa, J.E., 1994. Biodiversity conservation in the Gulf of Guinea: taking stock and preparing action.
Biodiversity and Conservation, 3(9): 759-771
- Karstadt, J.-O., Bagøien, E., 2016. Recruitment studies in the coastal waters of Gabon, Republic of Congo, and Angola in 2014 and 2016.
- Katsanevakis, S., Smit, W., Pipitone, C., Leopold, M., Cronin, M., Scheidat, M., Doyle, T. K., Buhl-Mortensen, L., Buhl-Mortensen, P., D'Anna, G., Boois, I. de, Dalpadado, P., Damalas, D., Fiorentino, F., Garofalo, G., Giacalone, V.M, Hawley, K., Issaris, Y., Jansen, J., Rosten, C., Knittweis, L, Kröncke, I., Mirto, S., Muxika, I., Reiss, H, Skjoldal, H.R., Vöge, S., 2012. Monitoring marine populations and communities: methods dealing with imperfect detectability. Aquatic Biology, 16: 31-52
- Krakstad, J-O., Bagøien, E., Ensrud, T.M., Lewmbe, J.deD. Samba, J., Andre, A.M., 2014. Recruitment studies on Sardinella aurita and maderensis in the coastal waters of Gabon, Congo and Northern Angola 24. May - 14. June 2014
- Krakstad, J-O., Skagseth, Ø., Bagøien, E., Ensrud, T.M., Lewembe, J.deD., Samba, J., Nsaku, D., 2016. Recruitment studies on Sardinella, Sardinella aurita and maderensis, in the coastal waters of Gabon and Republic of the Congo
- Longhurst, , 1998. Ecological Geography of the Sea. Academic Press, San Diego, CA.
- Maia, A., Morais, R.A., Siqueira, A.C., Hanazaki, N., Floeter, S.R., Bender, M.G., 2018. Shifting baselines among traditional fishers in São Tomé and Príncipe islands, Gulf of Guinea. Ocean Coastal Management, 154: 133–142
- Niza, H., Bento, M., Lopes, L.F., Cartaxana, A., Correia, A.M., 2021. A picture is worth a thousand words: using digital tools to visualise marine invertebrate diversity data along the coasts of Mozambique and São Tomé & Príncipe. Biodiversity Data Journal, 9: e68817, doi: 10.3897/BDJ.9.e68817
- Pina, A.A., Prazeres, M.J., Leal, A., Ramos, A., Bird, T., Benitez, L., Costa, E., Matos, L., 2022, Parque Natural do Príncipe (PNP) - Plano de Manejo 2022-2026. 104 pp. DOI: 10.13140/RG.2.2.11.20322
- Polidoro, B., Ralph, G.M., Strongin, K., Harvey, M., Carpenter, K.E., Adeofe, T.E., Arnold, R., Bannerman, P., Bi Nguema, J.N.B., Buchanan, J.R., Camara, K.M.A., Camara, Y.H., Cissoko, K., Collette, B.B., Comeros-Raynal, T., De Bruyne, G., Diouf, M., Djiman, R., Ducrocq, M., Gon, O. Harold, A.S., Harwell, H., Hilton-Taylor, C., Hines, A., Hulley, P.A., Iwamoto, T., Knudsen, S., Lewembe, J.D.D., Linardich, C., Lindeman, K., Mbye, E.M., Mikolo, J.E., Monteiro, V., Mougoussi, J.B., Munroe, T., Beh, J.H.M., Nunoo, F.K.E., Pollock, C., Poss, S., Quartey, R., Russell, B., Sagna, A., Sayer, C., Sidibe, A., Smith-Vaniz, W., Stump, E., Sylla, M., Tito De Morais, L., Vié, J.- C., Williams, A., 2016. Red List of Marine Bony Fishes of the Eastern Central Atlantic. Gland, Switzerland: IUCN. viii + 80 pp.
- República Democrática de São Tomé e Príncipe, 2022. Plano de Manejo do Parque Natural Obô de São Tomé, 2021-
- República Democrática de São Tomé e Príncipe, Plano de Manejo do Parque Natural do Príncipe, 2022-2026
- Rupert, E. E., Fox, R. S., Barnes, R. D., 2004. Invertebrate Zoology: A Functional Evolutionary Approach. . Thomson Brooks/Cole, 1008 pp.
- Sequeira , 1907. Adolfo Frederico Moller. Portugal em Africa, 14(167):528-529
- Silva, J.D., 1940. Francisco Newton. Explorador naturalista. "Coleção pelo Império" nº.68, Agência Geral das Colónias, Ministério das Colónias, Lisboa, 41 pp.
- Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N., Ferdaña, Z.A., Finlayson, M., Halpern, B.S., Jorge, M.A., Lombana, A., Lourie, S. A., Martin, K.D., Mcmanus, E., Molnar, J., Recchia, C. A., Robertson, J., 2007. Marine Ecoregions of the World: A Bioregionalization of Coastal and Shelf Areas. BioScience, 57(7) : 573-583
- Tams, H.T., 1933-4. A visit to the islands in the Gulf of Guinea. Natural History Magazine, 4(28-30): 126-218.
- Tams, W.H.T., 1934. The Percy Sladen and Godman Trusts Expedition to the Islands in the Gulf of Guinea, October 1932-March 1933. - I. Introduction. Annals and Magazine of Natural History, series 10 14(80):213-219.
- Tenreiro, , 1961. A ilha de São Tomé. Memórias da Junta de Investigações do Ultramar, 2ª Série 24:1–289
- Tittensor, P., Mora, C., Jetz, W., Lotze, H. K., Ricard, D., Berghe, E. V., Worm, B., 2010. Global patterns and predictors of marine biodiversity across taxa. Nature, 466: 1098–1101
- Vaz, , 2017. O enigma de São Tomé e Príncipe, ou um segredo por revelar. National Geographic Portugal https://nationalgeographic.pt/natureza/grandes-reportagens/1267-o-enigma-de-sao-tome-e-principe-ou-um-segredo-por-revelar
- Wirtz, , 2001. New records of marine invertebrates from the Cape Verde Islands. Arquipélago Life and Marine Sciences, 18A: 81-84
- Wirtz, , 2003. New records of marine invertebrates from São Tomé Island. Journal of the Marine Biological Association of the UK, 83: 735–736
- Wirtz, , 2009. Expedition to São Tomé. Journal of the Marine Biological Association of the UK, Global Marine Environment: 26-28
- Wirtz, , 2018. New records of marine invertebrates from São Tomé and Príncipe, eastern Atlantic Ocean.
Arquipélago Life and Marine Sciences, 35:41–46
- Worms, J., 1996. From subsistence to Commerce: Fisheries in São Tomé e Príncipe. Gulf of Guinea Conservation Newsletter, 6:3-4
MISCELLANEOUS REPORTS
- Airuad, , 2021. Relatório da 9ª missão de Assistência técnica em cogestão das pescas e AMP em São Tomé e Príncipe.
- Albuquerque C., Cesarini D., Tagliabue L.C., 2009. Plano de Gestão do Parque Natural Obô de São Tomé. ECOFAC IV, São Tomé, São Tomé e Príncipe, 109 pp.
- Barrencua, , 2016. Coordinator, S. & Haroun, R. BIO-Príncipe Scientific EXPEDITION – Final Report.
- Bollen, A., 2017. Compilation marine work, marine and coastal species and ecosystems of Príncipe. Fauna & Flora International, Fundação Príncipe, Santo António, 26 pp.
- Carneiro, G., 2011. ‘They come, they fish, and they go’: EC fisheries agreements with Cape Verde and São Tomé e Príncipe. Marine Fisheries Review, 73: 1–25
- De Labra, , Vilela López, B., Prieto Porriños, G., Blanc, P.P., Vasta, A., Anibal, O., 2023. The coastal pelagic value chain in São Tomé and Príncipe: Summary report. Rome, FAO.
- Direção das Pescas, 2015. Relatório do Estudo de Seguimento de Espécies Demersais da Zona Costeira de S. Tomé e Príncipe.
- Direção Geral do Ambiente, 2014(?). Estratégia Nacional e Plano de Acção para a Biodiversidade 2015-2020 (ENPAB II). 119 pp.
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- Ferreira-Airaud, B., Vieira, S., Henriques, N.S., Airaud, F., 2021. Relatório de realização do mapeamento submarino do distrito de Lobata, São Tomé. São Tomé e Príncipe, Programa Tatô and Marapa, São Tomé, 7 pp.
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- Omali Vida Nón, Sumário de atividades de projeto e resultados preliminares
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RESEARCH CRUISE REPORTS
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CLIMATE AND OCEANOGRAPHY
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Deep-Sea Research, 39 (7/8): 1273-1298
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- Wauthy, , 1983. Introduction à la climatologie du golfe de Guinée. Océanographie tropicale, 18(2):103-38 BIOLOGICAL PRODUCTIVITY
- Belkin, M., Cornillon, P.C., 2007. Fronts in the World Ocean’s Large Marine Ecosystems. ICES CM 2007/D:21
- Dessier, A, 1979. Ecologie dynamique des peuplements zooplanctoniques cotiers, et plus particulièrement des copepodes, du sud du golfe de Guinee (cotes du Congo, du Gabon et de l'Angola). Tese de Doctorat d'Etat ES- Sciences Naturelles. L'Universite Pierre et Marie Curie. 389 pp.
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MARINE HABITATS
- Afonso, F. M. G. 2019. A importância dos mangais de São Tomé: perceções e serviços ecossistémicos. (Faculty of Sciences of the University of Lisbon, MSc Thesis).
- Afonso, F., Félix, P.M., Chainho, P., Heumüller, J.A., de Lima, R.F., Ribeiro F., Brito, A.C. 2021. Assessing Ecosystem Services in Mangroves: Insights from São Tomé Island (Central Africa). Frontiers in Environmental Science, https://doi.org/10.3389/fenvs.2021.501673
- Airaud, F., Vieira, S., Henriques, N.S., Ferreira-Airaud, B. 2020. Relatório de realização do mapeamento submarino do distrito de Caué, São Tomé, São Tomé e Príncipe. Programa Tatô, Oikos-Cooperação e Fauna & Flora International, São Tomé, 13 pp.
- Airuad, F., 2021.Relatório da 9a missão de Assistência técnica em cogestão das pescas e AMP em São Tomé e Príncipe.
- Azevedo e Silva, J., 2021. Ecological characterization of the rocky shores of Príncipe Island, Gulf of Guinea. (Faculdade de Ciências, Universidade de Lisboa, MSc Thesis).
- Borja, , Andersen, J.H., D. Arvanitidis, C.D., Basset, A., Buhl-Mortensen, L., Carvalho, S., Dafforn, K.A., Devlin, M.J., Escobar-Briones, E.G., Grenz, C., Harder, T., Katsanevakis, S., Liu, D., Metaxas, A., Moran, X.A.G., Newton, A., Piroddi, C., Pochon, X., Queiros, A.M., Snelgrove, P.V.R., Solidoro, C., St. John, M.A., 2020. Past and Future
Grand Challenges in Marine Ecosystem Ecology. Frontiers in Environmental Science, 7 https://doi.org/10.3389/fmars.2020.00362
- Brito, C., Silva, T., Beltrán, C., Chainho, P., Lima, R. F. de, 2017. Phytoplankton in two tropical mangroves of São Tomé Island (Gulf of Guinea): A contribution towards sustainable management strategies. Regional Studies in Marine Science, 9: 89–96
- Buhl-Mortensen, L., Serigstad, B., Buhl-Mortensen, P., Olsen, M, Ostrowski, M., Blazewicz-Paszkowycz, M., Appoh, E., 2017. First observations of the structure and megafaunal community of a large Lophelia reef on the Ghanaian shelf (the Gulf of Guinea). Deep-Sea Research Part II: Tropical Studies in Oceanography, 137: 148-156
- Buhl-Mortensen, L., Serigstad, B., Buhl-Mortensen, P., Olsen, M., Ensrud, T.M., Ostrowski, M., Blazewicz- Paszkowycz, M., Jozwiak, P., Appoh. E., 2015. 6. First observations of the structure and megafaunal community of a large Lophelia reef on the Ivorian-Ghanaian margin (the Gulf of Guinea). Deep-Sea Research Part II: Tropical Studies in Oceanography, 137: 148-156
- Canterle, A. M., 2017. A influência da composição bentônica nas interações e diversidade de peixes recifais na ilha de Príncipe, Golfo da Guiné, África. (Universidade Federal de Santa Catarina), MSc Thesis
- Canterle, A.M., Nunes, L.T., Fontoura, L., Maia, H.A., Floeter, S.R., 2020. Reef microhabitats mediate fish feeding intensity and agonistic interactions at Príncipe Island Biosphere Reserve, Tropical Eastern Atlantic. Marine Ecology, 41(5):e12609
- Carpine, C., 1959. Campagne de la "Calypso" dans le golfe de Guinée el aux îles Príncipe, São Tomé, Annobon (1956). 11. Aperçu sur les peuplements littoraux. Annales d’Institut Océanographique, Paris NS, 37(4): 75-90
- Cowburn, B., 2018. Marine Habitats of Príncipe, Eastern Tropical Atlantic—description and map. Príncipe Trust, Santo António, 18 pp Available via the Omali Vida Nón project. https://omaliprincipeen.weebly.com/resources.html.
- Denechaud, C., Buhl-Mortensen, L., Joanny, T., Dovlo, E., Staby, A., 2022. Spatial environmental variability shapes bottom ecosystems diversity in the western Gulf of Guinea. African Journal of Marine Science, 47 (2): 191–205
- Ecologically or Biologically Significant Marine Areas (EBSAs). https://www.cbd.int/ebsa/repository
- Ferreira-Airaud, B., Vieira, S., Henriques, N. S., Airaud, F., 2021. Relatório de realização do Mapeamento Submarino do Distrito de Lobata.
- Haroun, R., Herrero-Barrencua, A., Abreu, A.D., 2018. Mangrove habitats in São Tomé and Príncipe (Gulf of Guinea, Africa): conservation and management status. In: Makowski C, Finkl C (eds) Threats to mangrove forests. Coastal Research Library, vol 25. Springer, Cham, pp. 589–605
- Herrero, H., Haroun, R., Abreu, A., 2017. Caracterización Preliminar de los Manglares de la Isla de Príncipe (São Tomé y Príncipe). IU-ECOAQUA, Universidad de Las Palmas de Gran Canaria. ecoaqua.eu
- Heumuller, J. A. et al., 2017. Diversidade e Fenologia dos Peixes como Ferramentas para Determinar a Diversidade Funcional nos Mangais da Ilha de São Tomé (São Tomé e Príncipe). In: 16º Encontro Nacional de Ecologia
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- Maia, H. A., 2018. Padrões Espaciais e Temporais das Comunidades Recifais nas Ilhas De São Tomé e Príncipe, Golfo de Guiné, África. PhD thesis (Universidade Federal de Santa Catarina)
- Maia, H. A., Morais,A., Quimbayo, J.P., Dias, M.S., Sampaio,C.S.S., Horta,P.A., Ferreira, C.E.L., Floeter, S.R., 2019. Spatial patterns and drivers of fish and benthic reef communities at São Tomé Island, Tropical Eastern Atlantic. Marine Ecology, 39(6):e12520
- McGlade, J.M., Cury, P., Koranteng, K., Hardman-Mountford, N.J., (ed), 2002. The Gulf of Guinea Large Marine Ecosystem. Environmental Forcing Sustainable Development of Marine Resources. 392 p. Sherman, K. (ed) Large Marine Ecosystems Series, Elsevier
- Morais, R. A., Maia, H. A., 2016. Reef Sites: Lush underwater forests in mesophotic reefs of the Gulf of Guinea. In: Coral Reefs 1 (Springer, 2016). doi:10.1007/s00338–016–1523-z.
- Otero-Ferrer, F., Tuya, F., Bosch Guerra, N.E., Herrero-Barrencua, A., Abreu, A.D., Haroun, R., 2020. Composition, structure and diversity of fish assemblages across seascape types at Príncipe, an understudied tropical island in the Gulf of Guinea (Eastern Atlantic Ocean). African Journal of Marine Science, 42(4):381–391
- Pinheiro, H. T., Ferreira, C. E. L., Joyeux, J. C., Santos, R. G., Horta, P. A., 2011. Reef fish structure and distribution in a south-western Atlantic Ocean tropical island. Journal of Fish Biology, 79: 1984–2006
- São Tomé e Príncipe, National report on the status of biodiversity in S. Tomé and Príncipe. 108 pp.
- Secrétariat de la Convention sur la diversité biologique, 2020. Aires marines d’importance écologique ou biologique (AIEB). Endroits spéciaux des océans du monde. Volume 6: L’océan Atlantique Sud-Est. Montréal : 108 pp.
- Spalding, M.D., Fox, H.E., Allen, G.R., Davidson, N., Ferdaña, Z.A., Finlayson, M., Halpern, B.S., Jorge, M.A., Lombana, A., Lourie, S. A., Martin, K.D., Mcmanus, E., Molnar, J., Recchia, C. A., Robertson, J., 2007. Marine Ecoregions of the World: A Bioregionalization of Coastal and Shelf Areas. BioScience, 57(7) : 573-583
- Tilot, V.. King, A., 1993. A Review of the Subsystems of the Canary Current and Gulf of Guinea Large Marine Ecosystems. IUCN Marine Programme
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- Ukwe, C.N., Ibe, C.A., Alo, B.I., Yumkella, K.K., 2003. Achieving a paradigm shift in environmental and living resources management in the Gulf of Guinea: the large marine ecosystem approach. Marine Pollution Bulletin, 47(1– 6):219–225
- Watson R., Pauly, D., Christensen, V., Froese, R., Longhurst, A., Platt, T. Sathyendranath, S., Sherman, K., O'Reilly, J., Celone, , 2003. Mapping Fisheries onto Marine Ecosystems for Regional, Oceanic and Global Integrations. In G. Hempel and K. Sherman (Ed) Large Marine Ecosystems of the World. 376-395 pp.
TAXA STUDIES
MACROALGAE AND SEAGRASSES
- Alexandre, A., Silva, J., Ferreira, R., Paulo, D., Serrão, E.A., Santos, R., 2017. First description of seagrass distribution and abundance in São Tomé and Príncipe. Aquatic Botany, 142:48–52
- John, D. M., Prud’homme van Reine, W. F., Lawson, G. W., Kostermans, T. B. & Price, J. H., 2004. A taxonomic and geographical catalogue of the seaweeds of the western coast of Africa and adjacent islands. Beihefte zur Nova Hedwigia 127
- Steentoft, B. Y. M., 1967. A revision of the marine algae of São Tomé and Principe (Gulf of Guinea). Linnean Society of London Proceedings Botany Journal, 60: 99–146
PROTOZOA
- Mangin, , 1959. Campagne de la Calypso dans le golfe de Guinée et aux îles Príncipe, São Tomé et Annobón (1956). IV. Foraminifères. Annales de l'lnstitut Oceanographique. Paris NS, 37(4): 91-114
- Reis Moura, , 1961. Contribuição para o conhecimento dos foraminíferos das praias levantadas de São Tomé e Príncipe. Garcia de Orta, 9(4):751-758
- Rocha, A.T., Pissara, J.B., 1963. Contribuição para o estudo mineralógico e da microfauna dos rgossolos psamiticos calcarios de São Tomé. Garcia de Orta, 11(1): 171-178
- Serralheiro, R., 1957. Novos elementos para o conhecimento da fauna fóssil do Miocenico da ilha do Príncipe .
Garcia da Orta, 5: 287-296
- Silva, G.H., 1956. La faune Miocene de l’Ile du Prince. Memórias e Notícias, Museu Mineralógico e Geológico da Universidade de Coimbra, 42: 29-53
- Silva, H., 1956. O Miocenico da ilha do Príncipe. Conferência International dos Africanistas Ocidentais 6ª sessão,
- Tomé. Comunicações, 2: 231-269
- Silva, H., 1958. Contribuição para o conhecimento da microfauna do Miocénico marinho da Ilha do Príncipe.
Garcia de Orta, 6(3):507-510
- Silva, G.H., 1958. Nota sobre a microfauna do Miocenio marinho da ilha do Príncipe. Memórias e Noticias. Museu Mineralógico e Geológico da Universidade de Coimbra, 45:56-60
PORIFERA
- C., 1959. Campagne 1956 dans le Golfe de Guinée el aux Iles Principe, São Tomé et Annobon. 5. Spongiaires.
Annales de l'lnstitut Oceanographique, Paris NS, 37(4): 115-141
- Moller, E., 1894. Esponjas de S.Thomé. Annaes das Sciencias Naturaes, 1:202-203 CNIDARIA
- Billard, A.. 1907. Hydroides recoltes par M. Ch. Gravier a l’île de San Thomé. Bulletin du Muséum National d'Histoire Naturelle. Paris, 13: 274-275
- Boschma, H. 1961: Resultats scientifiques des campagnes de la Calypso. XVII. Campagne de le Calypso dans le golfe de Guinee et aus lies Principe, Sao Tome, Annobon (1956). 12. Stylasterina. Annales de l'lnstitut Oceanographique Paris NS, 395: 193-225
- Carrison-Stone, D., Van Syoc, R., Williams, G., Brian Simison, W., 2013. Two new species of the gorgonian inhabiting barnacle, Conopea (Crustacea, Cirripedia, Thoracica), from the Gulf of Guinea. Zookeys, 270: 1–20
- Cordeiro, , McFadden, C., van Ofwegen, L., Williams, G., 2021. World List of Octocorallia. Acanthogorgia hirsuta Gray, 1857. World Register of Marine Species https://www.marinespecies.org/aphia.php?p=taxdetails&id=125349
- Creed, J. C., Fenner, D., Sammarco, P.W., Cairns, S.D., Capel, K., Junqueira, A.O.R., Cruz, I.C.S., Miranda, R.J., Carlos Junior, L.A., Mantelatto, M.C., Oigman-Pszczol, S,S., 2017. The invasion of the azooxanthellate coral Tubastraea (Scleractinia: Dendrophylliidae) throughout the world: history, pathways and vectors. Biological Invasions, 19: 283–305
- Forest, J., 1959. Campagne de la "Calypso" dans le Golfe de Guinéc ct aux i1es Príncipe, São Tomé et Annobon ( 1956). Resultats scientifiques de la campagne de la Calypso, 5: 36 pp.
- Gravier, C., 1906. Sur les formations corallienes de l'île San Thomé (golfe de Guinée). Bulletin du Muséum National d'Histoire Naturelle. Paris, 12(7):543-547
- Grasshoff, M., 1988. The Genus Leptogorgia (Octocorallia: Gorgoniidae) in West Africa. Scientific Results of the Danish Expedition to the Coasts of Tropical West Africa, 14.
- Gravier, C., 1906. Sur les formations corallienes de l'ile San Thomé (golfe de Guinée). Bulletin du Muséum National d’Histoire Naturelle, Paris, 12(7): 543-547
- Gravier, , 1909. Sur l'habitat el le polymorphisme du Siderastrea radians. Bulletin de Muséum Nationale d'Histoire Naturelle, Paris, 15:365-368
- Gravier, C., 1909. Sur l`habitat e la polymorphisme du Siderastrea radians. Bulletin de Muséum Nationale d'Histoire Naturelle, Paris, 15: 365-368
- Gravier, C., 1910. Madreporaires des Îles São Tomé et du Príncipe. Annales de l’Institut Océanographique, Monaco, 1: 1-27
- Greef, R., 1886. Über Westafrikanische Stylasteridea. Sitzungsberichte der Gesellschaft zur Beförderung der gesamten Naturwissenschaften zu Marburg.
- Laborel, J., 1974. West African reef corals: an hypothesis on their origin. Proceedings of the 2nd International Coral Reef Symposium, 1:425-443
- Repelin, R., 1965. Quelques méduses de l’ile Anno Bon (Golfe de Guinée). Cahiers d'O.R.S.T.O.M. Océanographiques, 3:73-79
- Rodríguez, E., Fautin, D., Daly, , 2023. World List of Actiniaria. Actinostella flosculifera (Le Sueur, 1817). World Register of Marine Species https://www.marinespecies.org/aphia.php?p=taxdetails&id=289355
- Tuder-Durivault, 1961: Resultats scientifiques des Campagnes de la Calypso. XVII. Campagne de la Calypso dans le golfe de Guinee et aux iles de Principe, Sao Tome, Annobon (1956). 14. Les octocoralliaires du Golfe de Guinee et des iles du Cap-Vert (Alcyonacea, Pennatulacea). Annales de l'lnstitut Oceanographique Paris NS, 39: 237-262
- Von Koch, W., Uber die von Herrn Prof. Dr. Greef in Gulf von Guinea gesammelten Anthozoen. Bonn. 34 pp.
- Wirtz, P., Berenger, L., 2017. Range extension of Pseudocorynactis caribbeorum den Hartog, 1980 in the eastern Atlantic. Revista de la Academia Canaria de Ciencias, 24: 203–206
- Wirtz, 2024. A pictorial catalogue of the Cnidaria Malacacyonacea of São Tomé and Príncipe. ResearchGate.
- Wirtz, 2024. Revised pictorial catalogue of the shallow water Scleractinia of São Tomé and Príncipe Islands. ResearchGate.
- Wirtz, P., Santos, M. E. A. 2025. A pictorial catalogue of the Zoantharia (Cnidaria) of São Tomé and Príncipe (STP).
- Wirtz, 2023. A pictorial catalogue of the shallow water Corallimorpharia (Cnidaria) of São Tomé and Príncipe. ResearchGate.
“WORMS”: NEMATODA, ANNELIDA, and CHAETOGNATA
- Britayev, T. A., Gil, J., Altuna, Á., Calvo, M. & Martín, D., 2014. New symbiotic associations involving polynoids (Polychaeta, Polynoidae) from Atlantic waters, with redescriptions of Parahololepidella greeffi (Augener, 1918) and Gorgoniapolynoe caeciliae (Fauvel, 1913). Memoirs of Museum Victoria, 71: 27–43
- Brown, J., Nishi, E. & Wirtz, P., 2019. The polychaete Lygdamis wirtzi at Ascension and St Helena Islands (Annelida, Polychaeta, Sabellariidae). Arquipélago Life and Marine Sciences, 1: 19–22
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MOLLUSCA
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- Nobre, A., 1887. Remarques sur la faune malacologique de marine des possessions portugaises de l' Afrique Occidentale. Jornal de Sciencias mathematicas physicas e naturaes, 12(46): 107·-120
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- Rolan, E., Fernandes, F., 1990. Coralliophila adansoni (Kosuge and Fernandes, 1989): new name for Ocinebrina adansoni. Boletino Malacologico 26(5-9): 143-144
- Rolan, E., Fernandes, F., 1991. Muricopsis (Risomurex) (Gastropoda, Muricidae) de las islas de São Tomé y Principe (Golfo de Guinea, Africa Occidental). Apex, Brussels, 6(1):11-20
- Rolan, E., Fernandes, F., 1992. Aportaciones aI conocimiento de la familia Turridae Swainson, 1840 (Mollusca, Gastropoda) en las islas de São Tomé y Príncipe (Golfo de Guinea). Nova Acta Cientifica Compostelana (Bioloxia), 3: 135-143
- Rolan, E., Fernandes, F., 1995. Aprovechamiento del carbonato calcico por una especie endemica de la isla de Príncipe. Noticiario SEM, 23: 30-31
- Rolán, E., Swinnen, F., 2004. A new Barleeia (Mollusca, Barleeidae) from Principe Island (Gulf of Guinea, West Africa). Gloria Maris, 43, 1–6
- Rolan, , Templado, J., 1993. Una especie nueva del genero Tropidorissoia (Mollusca: Gastropoda: Barleeidae) para la costa oesteafricana. Boletino Malacologico, 29: 237-242
- Rubio, F., Rolan, E., 1990. Aportaciones a 1os conocimientos sobre micromoluscos de Africa occidental. 2. Archaeogastropoda de São Tomé y Príncipe. Iberus, 9(1-2): 209-219
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Journal of Conchology 13:319-320
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ARTHROPODA CHELICERATA
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- Fage, , Stock, J.H., 1966. Campagne de la Calypso aux iles du Cap Vert (1959). 6. Pycnogonides. Annales de l'Institut Oceanographique, 44: 315-327
CRUSTACEA
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- Azevedo e Silva, J., Almeida, A.J., Cravo, M., Pais, M.P., Santos, Y., Paula, J., Patterns of fish distribution in tropical rock pools at Príncipe Island, Gulf of Guinea. African Journal of Marine Science, 44: 353-366
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- Crosse, H., Fischer, P., 1888. Observations sur le Bulimus exaratus Müller. Journal de Conchyliologie. 3 ser. 36: 11- 12
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- Osório, B., 1906. Uma nova lista de Crustaceos africanos. Jornal de Sciências Mathematicas, Physicas e Naturaes Lisboa, ser. 2, 7: 149,150
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