Anales del Jardín Botánico de Madrid 80 (1)
July-December 2023, e134
ISSN: 0211-1322, eISSN: 1988-3196
https://doi.org/10.3989/ajbm.565

Checklist of the vascular plants of Río Muni (Equatorial Guinea): floristic analysis, diversity, endemicity, and threatened status

Catálogo de las plantas vasculares de Río Muni (Guinea Ecuatorial): análisis florístico, diversidad, endemicidad y estado de amenaza

Guillermo VELAYOS

Universidad Complutense de Madrid, Departamento de Biodiversidad, Ecología y Evolución, José Antonio Novais 12, 28040 Madrid, Spain

https://orcid.org/0000-0001-9722-1042

Patricia BARBERÁ

Missouri Botanical Garden, Africa and Madagascar Department, 4344 Shaw Blvd., St. Louis, Missouri 63110, USA

https://orcid.org/0000-0002-4110-8419

Francisco J. CABEZAS

Universidad Complutense de Madrid, Departamento de Biodiversidad, Ecología y Evolución, José Antonio Novais 12, 28040 Madrid, Spain

https://orcid.org/0000-0003-0614-0445

Maximiliano FERO

Universidad Nacional de Guinea Ecuatorial. Av. de Hassan II s.n., Malabo, Equatorial Guinea

https://orcid.org/0000-0002-5169-9355

Mauricio VELAYOS

Real Jardín Botánico (RJB), CSIC, Departamento de Biodiversidad y Conservación, Plaza de Murillo 2, 28014 Madrid, Spain

https://orcid.org/0000-0002-0919-3545

Abstract

An updated checklist of the vascular plants from the continental region of Equatorial Guinea (Río Muni) is presented. The catalogue (Appendix 1) is the result of the compilation of herbarium specimens (6850), species records (7985) from the botanical literature and online databases (10109 GBIF records and 8897 RAINBIO records). A database of 23517 georeferenced records was prepared by updating the nomenclatural and standardizing the locality names from all these sources. The catalogue comprises 2707 taxa (2598 species, 81 subspecies, and 28 varieties) included in 1020 genera and 178 families. About 90.6% of the taxa are considered native, 1.17% introduced and 5.96% naturalized. The 10 most diverse families are Rubiaceae (294 species), Fabaceae (290), Orchidaceae (168), Poaceae (105), Euphorbiaceae (87), Apocynaceae (85), Cyperaceae (79), Annonaceae (68), Acanthaceae (65) and Melastomataceae (61), which comprise 49.22% of the species in Río Muni. Only 11 species can be considered endemic to Río Muni; this low number reflects the absence of natural barriers in the territory. The number of threatened taxa (VU, EN and CR) is 134 (5.02% of the total evaluated), of which 43 are at risk of extinction, being within the categories of Endangered or Critically Endangered. Five species restricted to the Gulf of Guinea are considered threatened: three Endangered (Grossera angustifolia, Polyscias aequatoguineensis and Rhipidoglossum montealenense), and two Critically Endangered (Asplenium carvalhoanum and Macropodiella uoroensis), thus they should be considered as priority in management plans development and conservation strategies.

Keywords: 
Biodiversity; endemism; flora; Equatorial Guinea; Río Muni.
Resumen

Se presenta un catálogo actualizado de las plantas vasculares de la región continental de Guinea Ecuatorial (Río Muni). El catálogo es fruto de la compilación de especímenes de herbario (6,850), registros de especies de la literatura botánica (7,985) y bases de datos en línea (10,109 registros de GBIF y 8,897 de RAINBIO). Se elaboró una base de datos de 23,517 registros georreferenciados realizando la actualización nomenclatural y estandarización de los nombres de localidades de todas estas fuentes. El catálogo comprende 2707 taxones (2598 especies, 81 subespecies y 28 variedades) incluidos en 1,020 géneros y 178 familias. El 90.6% de los taxones se consideran nativos, el 1.17% introducidos y el 5.96% naturalizados. Las 10 familias más diversas son Rubiaceae (294 especies), Fabaceae (290), Orchidaceae (168), Poaceae (105), Euphorbiaceae (87), Apocynaceae (85), Cyperaceae (79), Annonaceae (68), Acanthaceae (65) y Melastomataceae (61), que comprenden el 49.22% de las especies de Río Muni. Solo 11 especies pueden considerarse endémicas de Río Muni; este bajo número refleja la ausencia de barreras naturales en el territorio. El número de taxones amenazados (VU, EN y CR) es de 134 (5.02% del total evaluado) de los cuales 43 se encuentran en riesgo de extinción, al estar dentro de las categorías de En Peligro o En Peligro Crítico. Cinco especies restringidas al Golfo de Guinea se consideran amenazadas: tres En Peligro (Grossera angustifolia, Polyscias aequatoguineensis y Rhipidoglossum montealenense), y dos En Peligro Crítico (Asplenium carvalhoanum y Macropodiella uoroensis), por lo que deberían considerarse prioritarias en los planes de gestión y conservación.

Palabras clave: 
Biodiversidad; endemismo; flora; Guinea Ecuatorial; Río Muni.

Received: 16 December 2022; accepted: 23 February 2023; published online: 14 June 2023

Associate editor: Carlos Aedo.

How to cite this article: Velayos G., Barberá P., Cabezas F.J., Fero M., Velayos M. 2023. Checklist of the vascular plants of Río Muni (Equatorial Guinea): floristic analysis, diversity, endemicity, and threatened status. Anales del Jardín Botánico de Madrid 80: e134. https://doi.org/10.3989/ajbm.565

CONTENT

INTRODUCTION

 

The Republic of Equatorial Guinea is formed by an oceanic island called Annobón (17 km²), four continental islands: Corisco (15 km2), Elobey Grande (2.27 km²) and Elobey Chico (0.19 km²), and Bioko (2,017 km²), and a continental region, Río Muni. Politically, it is divided into seven provinces: Bioko Norte and Bioko Sur, Annobón, Litoral, Wele-Nzas, Kie-Ntem, and Centro Sur. The last four are on the mainland. Equatorial Guinea has maintained a close relationship with Europe. The island of Bioko was discovered in 1471, becoming part of the Portuguese crown. Four years later the same happened with the island of Annobon. In 1777, with the signing of the Treaty of San Ildefonso between Spain and Portugal, Equatorial Guinea passed to the Spanish crown, and would remain so until 1968, when it achieved its independence.

Río Muni is the continental region of Equatorial Guinea. It is in the Atlantic sector of Central Africa. It occupies 26017 km² between latitudes 1°01’N and 2°21’N, and longitudes 9°00 ‘E (the Atlantic coast) and 11°20’E. It limits to the north with Cameroon, along the Ntem River, and to the east and south with Gabon. Río Muni is part of a large granite plateau that runs from the Atlantic coast to the Adamawa Massif on the north, to the Angola Plateau on the south and to the Congo River Basin on the west. This granite block comes into contact with sandstones and marls of the Tertiary by the west and with sandstones and slates of the Secondary by the southwest.

From a biogeographical point of view, Río Muni is included entirely in the strip of lowland and plain African rainforest, which barely exceeds 1000 m in altitude. It is integrated within the Guineo-Congolian Region, in the Nigerian-Cameroonian Province (Takhtajan 1986: 199Takhtajan A. 1986. Floristic regions of the world. University of California, Berkeley.; Linder & al. 2012Linder H.P., de Klerk H.M., Born J., Burgess N.D., Fjeldså J. & Rahbek C. 2012. The partitioning of Africa: Statistically defined biogeographical regions in sub-Saharan Africa. Journal of Biogeography 39: 1189-1205.; Droissart & al. 2018Droissart V., Dauby G., Hardy O.J., Deblauwe V., Harris D.J., Janssens S., Mackinder B.A., Blach-Overgaard A., Sonké B., Sosef M.S.M., Stévart T., Svenning J.C., Wieringa J.J. & Couvreur T.L.P. 2018. Beyond trees: Biogeographical regionalization of tropical Africa. Journal of Biogeography 45: 1153-1167.). This biogeographical province contains several endemic genera and many endemic species, and is currently covered by dense equatorial rainforests.

The soils of Río Muni are derived from the decomposition of granite and gneiss. They are lateritic soils with a concentration of iron oxide, aluminum, titanium, and magnesium hydrates higher than 30% (De Castro & De la Calle 1985De Castro M.L. & De la Calle M.L. 1985. Geografía de Guinea Ecuatorial. Secretaría General Técnica. Ministerio de Educación y Ciencia, Madrid.; Fa 1991: 19Fa J.E. 1991. Conservación de los Ecosistemas Forestales de Guinea Ecuatorial. IUCN, Gland, Cambridge.).

The climate is equatorial with two dry seasons: one from July to September, the most important, and another from December to mid-February, and two rainy seasons: one from September to November and another from March to June. The average temperature is 25ºC with an oscillation not exceeding 5ºC. The average relative humidity is 90%, dropping slightly, to 85%, in the dry season. Annual precipitation ranges from 1800 to 3800 mm mainly between the months of September to December, with a considerable decrease in precipitation from March to May (De Castro & De la Calle 1985: 15-16De Castro M.L. & De la Calle M.L. 1985. Geografía de Guinea Ecuatorial. Secretaría General Técnica. Ministerio de Educación y Ciencia, Madrid.; Fa 1991: 20Fa J.E. 1991. Conservación de los Ecosistemas Forestales de Guinea Ecuatorial. IUCN, Gland, Cambridge.). However, these figures vary greatly from one year to another, with an estimated mean precipitation of 2500 mm/year (Wilks & Issembé 2000: 3Wilks C. & Issembé Y. 2000. Les arbres de la Guinée Equatoriale: Guide pratique d’identification: région continentale. Projet CUREF, Bata.). Rainfall decreases from west to east and from south to north. Mountainous areas, such as Mount Mitra, and the basin drained by the Mitemele river (Utamboni), on the southern border with Gabon, are the most humid areas.

Botanical collectors in Río Muni

 

The mainland of Equatorial Guinea has been much less explored than the island territories. The first known collections were made by the German Gustaf Mann (1836-1916), expeditionary of the Baikie’s Niger Expedition. Although he focused his work on Bioko, he also collected fifty specimens in Litoral province (Corisco, Elobey, and in the Kongui river [Congüe]) in 1862. These specimens are preserved mainly in the K herbarium.

It took almost half a century for the explorations in this area to continue, carried out by another German explorer, Günther Tessmann (1884-1969), who in 1908 collected 100 specimens in Litoral province and especially in Wele Nzas (hosted at K), at the Nkolentangan (Alén), Uelleburg and Bebai stations. In 1928, the Spanish forestry engineer Fernando Nájera, commissioned by the “Dirección General de Marruecos y Colonias”, visited the continental area to study the forest richness of Spanish Guinea. During the four months that he stayed in Río Muni, he studied and identified around 60 types of wood. These collections were probably lost from the Madrid “Escuela de Ingenieros de Montes” due to a fire. The result of Nájera’s study was published by the “Instituto Forestal de Investigaciones y Experiencias” (Nájera 1930Nájera F. 1930. La Guinea española y su riqueza forestal. Instituto forestal de investigaciones y experiencias, Madrid.).

The history of Spanish botanical collections in the continental region begins with the pharmaceutical inspector Lope del Val Cordón (1896-1967), who visited the area between June and July 1939, collecting 650 specimens in the four continental provinces. The first botanist to take a decided interest in the flora of Equatorial Guinea was Emilio Guinea López (1907-1985). He made only one trip to the continent between June and September 1945. Guinea indicates that he collected 3,000 specimens. This number is difficult to confirm due to the complicated numbering system that he used. What we know for sure is that more than 1000 vouchers are preserved in MA. A part of these specimens is found in Lisbon (LISC), where Emilio Guinea traveled to identify this material, and perhaps in London (K). The itinerary of his trip shows that he traveled through most of the continental region, with long walking journeys through the most intricate of the tropical forest. Although he collected in the four provinces, it is noteworthy his systematic and intense tours to the Wele Nzas and Kie Ntem provinces. A first advance of his work was published in the Anales del Jardín Botánico de Madrid (Guinea 1945Guinea E. 1945. Novedades sistemáticas de mi primer viaje a la Guinea Continental Española (verano de 1945). Anales del Jardín Botánico de Madrid 6: 463-486.). The following year he published what is still today the main work on the flora of Río Muni: Ensayo Geobotánico de la Guinea Continental española (Guinea 1946Guinea E. 1946. Ensayo geobotánico de la Guinea Continental Española. Dirección General de Marruecos y Colonias, Madrid.). Guinea was occasionally accompanied by J. Gómez Moreno. It will be almost fifty years after (1946Guinea E. 1946. Ensayo geobotánico de la Guinea Continental Española. Dirección General de Marruecos y Colonias, Madrid.), before floristic studies were resumed in the continental region starting in 1990. The Spanish-Portuguese Manuel Fidalgo do Carvalho, who had already collected in Bioko, moved his residence from Malabo (Bioko) to Bata on April 10, 1991, although he had already made some collections in Río Muni in December 1990. Carvalho remained on the continent until September of 1997. During his time in Río Muni, he made 1896 botanical collections. The abundant material that he collected from each specimen is very remarkable, always from fertile plants, and perfectly prepared and preserved. A complete series of his collections is held in MA, with numerous duplicates in major European and North American herbaria (AAU, B, BM, BR, COI, FI, G, H, K, LG, LISC, MO, NY, P, S, UPS, US, WAG, etc.) as well as in the National Herbarium of Equatorial Guinea (BATA).

At the same time of Guinea’s collecting work, in the 90s, the CUREF (“Conservación y Utilización Racional de los Ecosistemas Forestales de Guinea Ecuatorial”) project, funded by the European Union, started, with special emphasis on the study of plants. The botanical leader of the project was Jean Lejoly (BRLU) with the invaluable help of Stanislaw Lisowski (POZG). Several European botanists collected with the support of this project: Tariq Stévart (BRLU, MO), Ingrid Parmentier (BRLU), Bruno Senterre (BRLU) and Stephan Porembski (ROST) who collaborated in the training of a group of Guinean botanists from the INDEFOR (“Instituto de Desarrollo Forestal”), under the newly created BATA herbarium: Crisantos Obama, Pablo Esono, Norberto Nguema, Ildefonso Ndong, Domingo Ngomo, Francisca Eneme, and Diosdado Obiang Mbomio. In addition, there were six additional plant collectors who sometimes accompanied the CUREF-INDEFOR collecting team: Pierre van Asbroeck (BRLU) who collected in 1998 with Lejoly; N. Biedinger (BRLU) in 1998 with Porembski and N. Nguema, and Elad Maurice Epah (BRLU) in 1998 with Lejoly.

In 1997, with the financial support, of the “Agencia Española de Cooperación Internacional y Desarrollo” first, and later the Spanish Ministerio de Ciencia (in their different denominations), the Real Jardín Botánico, CSIC, began a project to study the flora of Equatorial Guinea, including Río Muni, Bioko and Annobón. It was directed by Mauricio Velayos (1955-) from the Real Jardín Botánico, CSIC. Since then, apart from M. Velayos, a team of researchers participated and directed the exploration and collection trips through the four continental provinces: C. Aedo (MA), F.J. Cabezas (MA, currently at MACB), M. de la Estrella (MA, currently at COA), P. Barberá (MA, currently at MO) and M. Fero Meñe (MA, currently at UNGE). In total, 6,800 specimens were collected under the Flora of Equatorial Guinea project.

Other people also participated in plant collecting campaigns organized by the Real Jardín Botánico: Raquel Pérez-Viso, who lived in Bata between 1998 and 2000 and collected intensively in Río Muni (4,500 specimens, especially from the Centro Sur Province). Myriam López Serrano, a pharmacist from Granada, focused on the study of the traditional use of medicinal plants of the Centro Sur Province. Encarnación Gómez Marín and Laureano Merino Cristóbal collected 160 plant specimens for the publication of the book Plantas Medicinales de Guinea Ecuatorial. Up to 48 more botanists visited Equatorial Guinea sporadically and collected in the country, but all together collected less than 100 specimens.

Floristic knowledge

 

Most recent published data from Río Muni are those of Sosef & al. (2017)Sosef M.S., Dauby G., Blach-Overgaard A., van der Burgt X., Catarino L., Damen T., Deblauwe V., Dessein S., Dransfield J., Droissart V., Duarte M.C., Engledow H., Fadeur G., Figueira R., Gereau R.E., Hardy O.J., Harris D.J., de Heij J., Janssens S., Klomberg Y., Klomberg Y., Ley A.C., Mackinder B.A., Meerts P., de Poel J.L. van, Sonke B., Stevart T., Stoffelen P., Svenning J.C., Sepulchre P., Zaiss R., Wieringa J.J. & Couvreur T.L. 2017. Exploring the floristic diversity of tropical Africa. BMC biology 15: 15.. In this study, the reported plant species richness for the whole of Equatorial Guinea is 3,049 species, increasing the estimated figure up to 3,821 species. This approach was based on the study of 15,341 records reflecting a Collection Density Index (CDI) for the country (number of collections/100 km2) of 54.69 (Campbell 1989Campbell D.G. 1989. The Importance of Floristic Inventory in the Tropics. In Campbell D.G. & Hammond D. (eds.), Floristic inventory of tropical countries: The status of plant systematics, collections, and vegetation, plus recommendations for the future: 6-29. New York Botanical Garden, Nueva York.). When CDI is calculated, the figures reveal a moderate level of knowledge of the region and more precisely from the continental region. These authors suggest a minimum of a CDI of 100 before one can state that the botanical inventory of a country or a region is of an acceptable level. If we calculate the CDI of the four provinces of Río Muni, only the Litoral Province is near the level of the CDI of 100, being Kie Ntem and Wele Nzas the provinces with the lowest botanical knowledge (Table 1).

Table 1.  Collection Density Index (CDI) of Río Muni and its provinces.
Territory Nº collections Area km² CDI
Centro Sur 4,065 9,931 40
Kie Ntem 1,019 3,943 27
Litoral 4,953 6,665 74
Wele Nzas 1,295 5,478 23
Unknown territory 217 - -
Total Río Muni 11,549 26,017 44

If we plot the collection localities from the 23,517 specimen records (Fig. 1) we observed that plant-collecting efforts are geographically biased and some inaccessible places remain poorly known, thus botanical exploration in Río Muni is still one of the main tasks for future biodiversity studies.

medium/medium-AJBM-80-01-e134-gf1.png
Fig. 1.  Locations of the plant collections (white circles) from Río Muni across its four provinces, based on 23,517 georeferenced specimen records. Protected areas are highlighted in green (see Fig. 3).

MATERIAL AND METHODS

 

We built a checklist (Appendix 1) based primarily on both herbarium specimens and bibliographic references. We studied 11,549 specimens collected in Río Muni, mainly from the herbaria BATA, BRLU, HBG, K, LBV, MA, and WAG; although to a lesser extent, also from BM, BR, G, MO, NY, P, and S. For comparative purposes specimens from Cameroon, Gabon, São Tomé and Príncipe, and other territories of Equatorial Guinea kept in those herbaria were also studied.

Of the 11,549 specimens compiled, only 6,850 could be identified to species, this is, assigned to an accepted name. Bibliographic references for all families of vascular plants were also checked. In total 7,985 species records from bibliographic sources were compiled from Alston (1956Alston A.H.G. 1956. New African ferns. Boletim da Sociedade Broteriana, sér. 2, 2: 30: 5-27., 1959)Alston A.H.G. 1959. The ferns and fern-allies of West Tropical Africa. A Supplement of the second edition of the Flora of West Tropical Africa. Crown Agents, London., Arends (1992)Arends J.C. 1992. Studies in Begoniaceae IV. Biosystematics of Begonia squamulosa Hook.f. and affiliated species in section Tetraphila A.DC. Wageningen Agricultural University Papers 91(6): 1-222., Aubréville (1961)Aubréville A. 1961. Sapotacées. In Aubréville A. (ed.), Flore du Gabon 1: 1-162. Muséum National d´Histoire Naturelle, Paris., Aubriot (2020)Aubriot X. 2020. Solanaceae. In Sosef S.M. & al. (eds.), Flore du Gabon 55: 89-137. Naturalis Biodiversity Center & Botanic Garden Meise, Robdorf. , Bamps (1970Bamps P. 1970. Notes sur les Guttiferae d’Afrique tropicale. VI et VII. Bulletin du Jardin Botanique National de Belgique 40: 281-290., 1974)Bamps P. 1974. Contribution a l’etude des Araliacees africaines. Bulletin du Jardin Botanique National de Belgique 44: 101-139., Barberá & al. (2013Barberá P., Aedo C. & Velayos M. 2013. Annotated checklist and identification keys of the Acalyphoideae (Euphorbiaceae) of Equatorial Guinea (Annobón, Bioko and Río Muni). Phytotaxa 140: 1-25., 2014Barberá P., Velayos M. & Aedo C. 2014. Taxonomic Revision of Grossera (Crotonoideae, Euphorbiaceae): A Central African Genus. Systematic Botany 39: 490-509. , 2015)Barbera P., Riina R. & Velayos M. 2015. A new species of Grossera (Euphorbiaceae) from Equatorial Guinea. Phytotaxa 221: 175-180., Bauters & al. (2018)Bauters K., Goetghebeur P., Asselman P., Meganck K. & Larridon I. 2018. 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(2019)Veranso-Libalah M.C., Lachenaud O., Stone R.G. & Kadereit G. 2019. Nothodissotis (Melastomataceae), a new genus from Atlantic Central Africa, including the new species N. alenensis from Equatorial Guinea. PhytoKeys 118: 89-103. , Verdcourt (1996)Verdcourt B. 1996. Piperaceae. In Polhill R.M. (ed.), Flora of Tropical East Africa: 1-24. A.A. Balkema, Rotterdam and Brookfield., Vermeulen (1987)Vermeulen J.J. 1987. A taxonomic revision of the continental African Bulbophyllinae. Orchids Monographs 2: 1-298., Verstraete & al. (2013)Verstraete B., Lachenaud O., Smets E., Dessein S. & Sonké B. 2013. Taxonomy and phylogenetics of Cuviera (Rubiaceae-Vanguerieae) and reinstatement of Globulostylis with the description of three new species. Botanical Journal of the Linnean Society 173: 407-441., Villiers (1990)Villiers J.F. 1990. Contribution à l’étude du genre Newtonia Baillon (Leguminosae-Mimosoideae) en Afrique. Bulletin du Jardin Botanique National de Belgique 60: 119-138, White (1963White F. 1963. Notes on the Ebenaceae V. Some new names and new species in Diospyros. Bulletin du Jardin Botanique de l’État à Bruxelles 33: 345-367., 1976White F. 1976. The taxonomy, ecology and chorology of African Chrysobalanaceae (excluding Acioa). Bulletin du Jardin Botanique National de Belgique 46: 265-350., 1978)White F. 1978. The taxonomy, ecology and chorology of African Ebenaceae I. The Guineo-Congolian species. Bulletin du Jardin Botanique National de Belgique 48: 245-358., Whitmore (2008)Whitmore T.C. 2008. The genus Macaranga. A prodromus. Royal Botanic Gardens, Kew., Wieringa (1999)Wieringa J.J. 1999. Monopetalanthus exit. A systematic study of Aphanocalyx, Bikinia, Icuria, Michelsonia and Tetraberlinia (Leguminosae, Caesalpinioideae). Wageningen Agricultural University Papers 99 (4): 1-320., Wilde (1968Wilde J.J.F.E. de 1968. A revision of the species of Trichilia P. Browne (Meliaceae) on the African continent. Mededelingen Landbouwhogeschool Wageningen 68(2): 1-207., 1971)Wilde W.J.J.O. de 1971. A monograph of the genus Adenia Forsk. (Passifloraceae). Mededelingen Landbouwhogeschool Wageningen 71(18): 1-281., Wilks & Issembé (2000)Wilks C. & Issembé Y. 2000. Les arbres de la Guinée Equatoriale: Guide pratique d’identification: région continentale. Projet CUREF, Bata., Zemagho & al. (2017Zemagho L., Liede-Schumann S., Lachenaud O., Dessein S. & Sonké B. 2017. Taxonomic revision of Sabicea subgenus Anisophyllae (Ixoroideae, Rubiaceae) from Tropical Africa, with four new species. Phytotaxa 293: 1-68., 2018)Zemagho L., Lachenaud O. & Sonké B. 2018. Four new species of Sabicea (Rubiaceae) from tropical Africa, with additional notes on the genus. Candollea 73: 277-293.. Names were updated according to the African Plant Database (https://africanplantdatabase.ch/en), or in the cases where supporting specimens were mentioned, the corresponding accepted names were assigned. Names of localities were also updated and standardized. Complementarily, 10,109 records from GBIF (2022)GBIF.org. 2022. GBIF Occurrence Download (16 Dec. 2022), https://doi.org/10.15468/dl.gpnzaa and 8,897 records from RAINBIO (Dauby & al. 2016Dauby G., Zaiss R., Blach-Overgaard A., Catarino L., Damen T., Deblauwe V., Dessin S., Dransfield J., Droissart V., Duarte M.C., Engledow H., Fadeur G., Figueira R., Gereau R.E., Hardy O.J., Harris D.J., de Heij J., Janssens S.B., Klomberg Y., Ley A.C., Mackinder B.A., Meerts P., van de Poel J.L., Sonké B., Sosef M.S.M., Stévart T., Stoffelen P., Svenning J.-C., Sepulchre P., van der Burgt X.M., Wieringa J.J. & Couvreur T.L.P. 2016. RAINBIO: a mega-database of tropical African vascular plant distributions. PhytoKeys 74: 1-18.) databases were also checked and updated, building a dataset of 36,984 species occurrence records from Río Muni, of which 23,517 were georeferenced.

The checklist was arranged in four main plant groups: Pteridophytes, Magnoliids, Monocotyledons and Eudicotyledons. Within each group, families, genera, and species were arranged alphabetically. Family classification follows PPG I (Schuettpelz & al. 2016Schuettpelz E., Schneider H., Smith A.R., Hovenkamp P., Prado J., Rouhan G., Salino A., Sundue M., Almeida T.E. , Parris B., Sessa E.B., Field A.R., Gasper A.L., Rothfels C.J., Windham M.D., Lehnert M., Dauphin B., Ebihara A., Lehtonen S., Schwartsburd P.B., Metzgar J., Zhang L.B., Kuo L.Y., Brownsey P.J., Kato M. & Arana M.D. 2016. A community-derived classification for extant lycophytes and ferns. Journal of Systematics and Evolution 54: 563-603.) for Pteridophytes and APG IV (Byng & al. 2016Byng J.W., Chase M.W., Christenhusz M.J.M., Fay M.F., Judd W.S., Mabberley D.J., Sennikov A.N., Soltis D.E., Soltis P.S. & Stevens P.F. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181: 1-20.) for Angiosperms. When a discrepancy with the classification of Flora de Guinea Ecuatorial (Velayos & al. 2008Velayos M., Aedo C., Cabezas F. & Estrella M. 2008. Flora de Guinea Ecuatorial. Claves de plantas vasculares de Annobón, Bioko y Río Muni vol. I. Psilotaceae-Vittariaceae. Real Jardín Botánico (CSIC), Madrid. ) was found we indicated it in brackets. Authors of plant names were abbreviated according to Brummit & Powell (1992)Brummit P.K. & Powell C.E. 1992. Authors of plant names. Royal Botanic Gardens, Kew. Londres., and to the updated version available at http://www.ipni.org/ipni/query_author.html.

Under each accepted name, we included the following information: i) the updated IUCN category (https://www.iucnredlist.org/ accessed September 2022) (NE indicates the species has not been evaluated); ii) distribution information, indicating if the species is native, endemic, subendemic (considering subendemic plants with a reduced distribution area in which most of the localities are in Río Muni), introduced, naturalized, or cultivated and also the complete distribution range with the biogeographical divisions of Takhtajan (1986)Takhtajan A. 1986. Floristic regions of the world. University of California, Berkeley.; and iii) a list of the studied specimens arranged by provinces. References from the most relevant literature were also included, but these were not considered in the floristic analysis. Taxa marked with a question mark are doubtful because of imprecise information and must be corroborated in the future.

A map (Fig. 1) and an updated and standardized list of collection localities of Río Muni is also provided (Appendix 2). A list of 665 localities was gathered, checked, and correctly spelled, and 450 localities were revised and georeferenced.

The conservation status of threatened species was assessed using the IUCN criteria (IUCN 2019). A preliminary risk of extinction assessment using the IUCN Red List categories and criteria (IUCN 2019) was carried out for the endemics and subendemic species that have not yet been published in the IUCN. Area of Occupancy (AOO) and Extent of Occurrence (EOO), estimated using a 2 × 2 km grid, were calculated using GeoCAT (2020)GeoCAT. 2020. Geospatial Conservation Assessment Tool. Royal Botanic Gardens, Kew. Website: http://geocat.kew.org [accessed 5 Nov. 2022].

RESULTS AND DISCUSSION

 

Floristic Analysis

 

Due to their biological importance, floristic inventories of tropical African underexplored areas are an urgent need. Despite of the huge development of communication infrastructures in the continental region of Equatorial Guinea, a large percentage of its territory is still inaccessible (Fig. 1). Besides, some of the published data are not supported by herbarium specimens, thus so much of the information available is not reliable. Many of the analyses, conservation plans, or management strategies that have been developed in African countries have been made with outdated bibliographic data and assuming complete biodiversity inventories, which are far from being so. Thus, it is quite common in these regions (tropical Africa) that when new collections are added, species numbers change drastically. Similar results are usually observed with new taxonomic treatments or systematic analyses. Endemic or threatened species are usually known from a few collections, which require new data to properly evaluate their real taxonomic status or threat level. The Río Muni Vascular Plants catalogue is a compendium of numerous cases of the above-mentioned problems.

The checklist of Vascular Plants from Río Muni comprises 178 families, 1,020 genera and 2,707 species and infraspecific taxa (Appendix 1). Pteridophytes represent the 5% of the species (142) and the 12.35% of the families of Río Muni (22). With these figures, the floristic knowledge of Río Muni, in terms of number of species, increases in 26.14% from the most recent studies (Table 2). The number of taxa considered as native is 2,411 (89.96%), 32 species (1.2%) are introduced, 36 cultivated (1.35%), and 158 (5.92%) should be considered as naturalized since they are able to establish new populations.

Table 2.  Number of Rio Muni vascular plants species compared with the main previous works.
Guinea 1946Guinea E. 1946. Ensayo geobotánico de la Guinea Continental Española. Dirección General de Marruecos y Colonias, Madrid. Lejoly & Lisowski 1998Lejoly J. & Lisowski S. 1998. Catalogue des Plantes Vasculaires de la Region Continentale de Guinee Equatorial. Ministerio de Bosques y Medio Ambiente República de Guinea Ecuatorial. CUREF, Bata. Sosef & al. 2017Sosef M.S., Dauby G., Blach-Overgaard A., van der Burgt X., Catarino L., Damen T., Deblauwe V., Dessein S., Dransfield J., Droissart V., Duarte M.C., Engledow H., Fadeur G., Figueira R., Gereau R.E., Hardy O.J., Harris D.J., de Heij J., Janssens S., Klomberg Y., Klomberg Y., Ley A.C., Mackinder B.A., Meerts P., de Poel J.L. van, Sonke B., Stevart T., Stoffelen P., Svenning J.C., Sepulchre P., Zaiss R., Wieringa J.J. & Couvreur T.L. 2017. Exploring the floristic diversity of tropical Africa. BMC biology 15: 15. This work
Pteridophyta 52 53 133 142
Spermatophyta 1,064 1,141 2,013 2,565
Total 1,116 1,194 2,146 2,707

The most diverse families are Rubiaceae (294 species), Fabaceae (290), Orchidaceae (168) and Poaceae (105). The 10 most species-rich families are completed by Euphorbiaceae (87), Apocynaceae (85), Cyperaceae (79), Annonaceae (68), Acanthaceae (65) and Melastomataceae (61). These 10 families alone comprise about half of the species (49.22%) of Río Muni (Fig. 2).

medium/medium-AJBM-80-01-e134-gf2.png
Fig. 2.  Diversity of species per family in Río Muni. The eleven most specious families (≥ 60 spp.) are highlighted. The “Others” category includes 167 families with fewer than 60 species.

This floristic pattern of family diversity is similar to the one found in Annobon island (Velayos & al. 2014Velayos M., Barberá P., Cabezas F., Estrella M., Fero M. & Aedo C. 2014. Checklist of the Vascular Plants of Annobón (Equatorial Guinea). Phytotaxa 171: 1-078. ). Such pattern is observed even when total species-richness figures are 10-times bigger (Annobon 24 species of Fabaceae, Río Muni 290) and it highlights the importance of the flow of species across the territories of the Gulf of Guinea. This idea is supported by the low number of endemic species observed in these territories.

Although the botanical exploratory level of Río Muni is relatively high when compared to the Central African countries, the number of species recorded so far is still far from the observed in neighboring Cameroon or Gabon, with 6,883 and 5,236 species respectively (Sosef & al. 2017Sosef M.S., Dauby G., Blach-Overgaard A., van der Burgt X., Catarino L., Damen T., Deblauwe V., Dessein S., Dransfield J., Droissart V., Duarte M.C., Engledow H., Fadeur G., Figueira R., Gereau R.E., Hardy O.J., Harris D.J., de Heij J., Janssens S., Klomberg Y., Klomberg Y., Ley A.C., Mackinder B.A., Meerts P., de Poel J.L. van, Sonke B., Stevart T., Stoffelen P., Svenning J.C., Sepulchre P., Zaiss R., Wieringa J.J. & Couvreur T.L. 2017. Exploring the floristic diversity of tropical Africa. BMC biology 15: 15.). Besides the differences in area among the three of them, these figures are pointing out the need of continuing with the collection effort in Río Muni. The number of species in the continental region and the entire Equatorial Guinea would likely increase if plant collecting efforts were extended to the poorly explored areas in the near future.

Threatened status and endemicity

 

The number of threatened taxa (VU, EN and CR) is 144 (5.31% of the total taxa) of which 48 are under extinction risk, i.e., considered Endangered or Critically Endangered (Table 3). Two endemic species, Polyscias aequatoguineensis and Scleria mongomoensis are categorized as Endangered and should be regarded as prioritary species for Conservation plans and strategies. Nonetheless, a revaluation of the real situation of these species should be even more important to do since preliminary data indicate that the real threatened status of Vascular Plants in Equatorial Guinea is far from well known. As an example, we focus our discussion below on endemic or subendemic plants from Río Muni.

Table 3.  Number of taxa per conservation status for each major plant group in Río Muni. Percentages were calculated considering the total number of taxa in each plant group.
NE-DD LC NT VU EN CR
% % % % % %
Pteridophyta 132 92.96 9 6.34 0 0 1 0.7 0 0 0 0
Gymnospermae 1 50 0 0 1 50 0 0 0 0 0 0
Magnoliods 42 43.75 43 44.79 3 3.12 6 6.25 1 1.04 1 1.04
Monocots 389 65.79 177 31.21 1 0.17 8 1.41 11 1.94 1 0.17
Eudicots 1,082 57.55 652 34.68 31 1.65 81 4.3 30 1.6 4 0.21
Total 1,646 881 36 96 42 6

Of all threatened taxa, only three are non-native to Río Muni namely Terminalia neotaliala, Gossypium hirsutum (both naturalized) and Memecylon erythranthum (introduced); and only 11 species could be considered as strictly endemic to Río Muni (Table 4). This low number of endemic taxa is reflecting the absence of natural barriers in the territory.

Table 4.  Endemic and subendemic species from Río Muni. Accepted Name, threatened category according to the IUCN, or preliminary assessments already published or preliminary assessed here. Preliminary risk of extinction assessments using the IUCN Red List categories and criteria (IUCN 2022) are provided for the endemic and subendemic species that have not yet been published in the IUCN.
Family Species IUCN category Preliminary assessment
Araliaceae Polyscias aequatoguineensis Lejoly & Lisowski EN (2021) -
Aspleniaceae Asplenium carvalhoanum Herrero, Aedo, Velayos & Viane NE CR (here)
Begoniaceae Begonia aequatoguineensis Sosef & Nguema NE VU D2 (here)
Begoniaceae Begonia monte-alenensis Sosef NE VU D2 (Sosef 2014Sosef M. 2014. Novitates Gabonenses 86. The Begonia clypeifolia complex (Begoniaceae) unravelled. Plant Ecology and Evolution 147(2): 224-236)
Cyperaceae Scleria mongomoensis Bauters EN (2020) -
Euphorbiaceae Grossera angustifolia Barberá & Riina EN (2022) -
Malvaceae Scaphopetalum obiangianum M.E. Leal NE VU D2 (here)
Melastomataceae Guyonia obamae (Lejoly & Lisowski) Veranso-Libalah & R. D. Stone (=G. arenaria (Jacq.-Fél.) Verano-Libalah & R.D. Stone NE NT (Lachenaud & al. 2018Lachenaud O., Stévart T., Boupoya A., Texier N., Dauby G. & Bidault E. 2018. Novitates Gabunensis 88: additions to the flora of Gabon and new records of little-known species. Plant Ecology and Evolution 151: 393-422.)
Melastomataceae Nothodissotis alenensis Veranso-Libalah & O. Lachenaud NE VU D2 (Veranso-Libalah & al. 2019Veranso-Libalah M.C., Lachenaud O., Stone R.G. & Kadereit G. 2019. Nothodissotis (Melastomataceae), a new genus from Atlantic Central Africa, including the new species N. alenensis from Equatorial Guinea. PhytoKeys 118: 89-103. )
Orchidaceae Polystachya riomuniensis Stévart & Nguema NE LC/EN
Orchidaceae Rhipidoglossum montealenense Descourvières, Stévart & P.J. Cribb - EN (Descourvières & al. 2013)
Orchidaceae Vanilla chalotii Finet NE VU/EN
Podostemaceae Macropodiella uoroensis Rial NE CR (here)

From the 13 endemic/subendemic species, three of them (Scleria mongomoensis, Polyscias aequatoguineensis, and Grossera angustifolia) have been recently evaluated by the IUCN as EN (Larridon & Bauters 2020Larridon I. & Bauters K. 2020. Scleria mongomoensis. The IUCN Red List of Threatened Species 2020: e.T172032833A172033445. ; Paradis & al. 2021Paradis A.H., Ikabanga D.U., Bidault E., Texier N. & Stévart T. 2021. Polyscias aequatoguineensis. The IUCN Red List of Threatened Species 2021: e.T173360456A174512880. ; Ruiz de Diego & al. 2022Ruiz de Diego P., Riina R. & Barberá P. 2022. Grossera angustifolia. The IUCN Red List of Threatened Species 2022: e.T176584058A176601312.). Guyonia obamae records from Río Muni have been identified as G. arenaria, and the species was preliminary assessed as NT (Lachenaud & al. 2018Lachenaud O., Stévart T., Boupoya A., Texier N., Dauby G. & Bidault E. 2018. Novitates Gabunensis 88: additions to the flora of Gabon and new records of little-known species. Plant Ecology and Evolution 151: 393-422.), nonetheless its taxonomic situation is still unclear.

We confirm that 12 out 13 of the endemic species evaluated should be considered as threatened (Table 4). Only for the subendemic Polystachya riomuniensis, since the distribution range of the species is quite large, the Extent of Occurrence (EOO) criteria states it as Least Concern while the Area of Occupancy (AOO) indicates the taxon as EN. We consider a more realistic conservative approach until taxonomic studies confirm the identity of these 12 species or new collecting campaigns confirm the presence of these taxa beyond the current distribution ranges.

Preliminary assessments of NE taxa

 

Scaphopetalum obiangianum is a shrub 20-150 cm tall, known from the understory of rain forest, at around 700 m in elevation (Leal 2007Leal M.E. 2007. Novitates Rio Munis 1. A new endemic Scaphopetalum (Malvaceae) from Mount Mitra, Equatorial Guinea. Blumea Journal of Plant Taxonomy and Plant Geography 52: 137-138. ). The species is endemic to Equatorial Guinea and is only known from one collection made by Leal in 2005 in Monte Mitra, which thus represents a single occurrence in the Litoral Province. The AOO is estimated as 4 km2, below the upper threshold for Critically Endangered status under subcriterion B2. The EOO is the same as the AOO. This occurrence therefore represents the entire known population of the species. It is located within a protected area (Monte Alen National Park), and no specific threats have been identified at the present time. Therefore, Scaphopetalum obiangianum is assigned a preliminary status of VU D2.

Begonia aequatoguineensis is an herb up to c. 5 cm high, known from primary forest at c. 450 m altitude (Sosef & Nguema Miyono 2010Sosef M.S.M. & Nguema Miyono N.S. 2010. New species of Begonia section Loasibegonia (Begoniaceae) from the Monte Alen region, Equatorial Guinea. Blumea Journal of Plant Taxonomy and Plant Geography 55: 91-93.). The species is endemic to Equatorial Guinea and is only known from one collection, which thus represents a single occurrence, made by Nguema Miyono at West of Mendung, in the Centro Sur Province. The AOO is estimated as 4 km2, below the upper threshold for Critically Endangered status under subcriterion B2. The EOO is the same as the AOO. This occurrence therefore represents the entire known population of the species. It is located within the protected area Monte Alén National Park and no specific threats have been identified at the present time. Therefore, Begonia aequatoguineensis is assigned a preliminary status of VU D2.

Asplenium carvalhoanum is an epiphyte, known from primary rainforest, endemic to Equatorial Guinea (Herrero & al. 2001Herrero A., Aedo C., Velayos M. & Viane R.L. 2001. A new species of Asplenium (Aspleniaceae, Pteridophyta) from Equatorial Guinea. Annales Botanici Fennici 38: 175-180.). The species is only known from one collection, made by Carvalho in Cogo, between Emangós and Ncó villages, in the Litoral Province, which thus represents a single occurrence. The AOO is estimated as 4 km2, below the upper threshold for Critically Endangered status under subcriterion B2. The EOO is the same as the AOO. This occurrence therefore represents the entire known population of the species. It is not located within a protected area and is threatened by logging and by small-scale harvesting. The most serious plausible threat is the logging. We infer a future decline in the extent and quality of its habitat, therefore Asplenium carvalhoanum is thus assessed as CR B2ab(iii).

Macropodiella uoroensis is endemic to Equatorial Guinea, known from the rapids of the Uoro river, at Centro Sur province. The species is only known from one collection made by Rial & Laso in 1994, representing a single occurrence. The AOO is estimated as 4 km2, below the upper threshold for Critically Endangered status under subcriterion B2. The EOO is the same as the AOO. This occurrence therefore represents the entire known population of the species. It is not located within a protected area and is potentially threatened by water pollution due to increase in urbanization. We infer a future decline in the extent and quality of its habitat hence, Macropodiella uoroensis is assessed as CR B2ab(iii).

Priority conservation areas

 

The geographical analyses of threatened species from Río Muni revealed the value of the National System of Protected Areas of Equatorial Guinea (Fig. 3). A total of 618 records out of the 23,517 compiled correspond to threatened species (CR, EN or VU), 224 of them are outside protected areas, which means that 36.25% of the threatened plants of Río Muni do not have any protection at all. If we consider those Near Threatened taxa, there are 721 total occurrences, 276 outside protected areas (38.28%). Monte Alén National Park is by far the most important Protected Area in Río Muni, hosting 49.23% of the occurrences of threatened species in this continental region. Two protected areas, Río Campo, and Montes Temelón do not include any occurrence from threatened species (Fig. 3).

medium/medium-AJBM-80-01-e134-gf3.png
Fig. 3.  Protected Areas of Río Muni here depicted in different colors. Dots indicate the occurrences of threatened species: CR (red), EN (black), VU (yellow).

Endemicity and areas for conservation

 

The conservation relevance of the system of protected areas of Equatorial Guinea comes to the fore when we consider the endemicity of the flora. Figure 4 shows the spatial distribution of the 13 endemic and subendemic species. Twenty-one of these occurrence records (46.6%) are included in protected areas. In this sense, the situation of Scleria mongomoensis, Macropodiella uoroensis, Polyscias aequatoguineensis, Grossera angustifolia and Asplenium carvalhoanum is particularly delicate since none of their localities has any protection status. As aforementioned, all of them, but Polyscias aequatoguineensis, could be also threatened under the IUCN category of Critically Endangered.

medium/medium-AJBM-80-01-e134-gf4.png
Fig. 4.  Distribution of endemic and subendemic species in Equatorial Guinea. Protected areas of Río Muni (8) are shown in different colors and endemics occurrences are indicated by colored dots and numbers.

Expanding the area of the Monte Alén National Park and establishing a monitoring program for these species to establish corridors between protected areas or new areas of special interest for the flora are necessary strategies to achieve a sustainable management of plant diversity in Equatorial Guinea. Connecting the different protected areas or increasing the surface area of some of them seems to be the best alternative to include the greatest number of threatened species present in this region. Finally, we emphasize that coastal meadows are especially under protected from human impacts.

ACKNOWLEDGMENTS

 

The elaboration of the checklist of vascular plants of Río Muni would have not been possible without the contribution of many people along the last years. We assume that, if we try to mention all of them, we will surely forget some names. We apologize for not even trying. However, it is necessary to mention the main contributors: the authors of the Flora de Guinea project who, over the years, collected, identified and studied most of the specimens on which this checklist is based.

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