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Addressing the Vepris Verdoorniana Complex (Rutaceae) in West Africa, with Two New Species

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Addressing the Vepris Verdoorniana Complex (Rutaceae) in West Africa, with Two New Species KEW BULLETIN (2019) 74:53 ISSN: 0075-5974 (print) DOI 10.1007/S12225-019-9837-Y ISSN: 1874-933X (electronic) Addressing the Vepris verdoorniana complex (Rutaceae) in West Africa, with two new species Martin Cheek1 , Jean-Michel Onana2, Shigeo Yasuda1, Poppy Lawrence3, Gabriel Ameka4 & Greta Buinovskaja5 Summary. Vepris verdoorniana (Rutaceae) has long been regarded as a widespread and variable species occurring from Guinea to Gabon. Recent evidence has shown that the original material from the type locality in Cameroon consists of two different species, V. verdoorniana in the strict sense, endemic to Lower Guinea (Cameroon, Gabon and C.A.R. in W-C Africa) and V. letouzeyi Onana, which was thought to occur from Guinea to Cameroon. In this paper we show that the West African (Upper Guinea) material previously referred to as V. letouzeyi/V. verdoorniana is separate morphologically from that in Cameroon (Lower Guinea). In fact the West African material referred to comprises two distinct species, here described as V. occidentalis Cheek & Onana (Guinea to Ghana, but probably extending to Benin) a tree of lowland marginal dry evergreen and semi-deciduous forest, and V. fer Cheek, a species restricted to four mainly lower submontane forest locations on iron substrates in Liberia and Sierra Leone (also Upper Guinea). Vepris occidentalis is here assessed as Least Concern due to its wide range and high frequency, whilst V. fer is assessed as Endangered due to the low number of locations and threats from iron ore mining projects. Both new species are illustrated and mapped. Key Words. Arcelor Mittal, conservation assessments, endangered, Gola, iron substrate, mining natural products, Teclea. Introduction evergreen shrubs and trees, predominantly of tropical Vepris Comm. ex A.Juss. (Rutaceae-Toddalieae), is a lowland evergreen forest, but with some species genus with 86 accepted species, 22 in Madagascar and extending into submontane forests and some into 65 in Continental Africa with one species extending to drier forests and woodland. Vepris species are often Arabia and another endemic to India (Plants of the indicators of good quality, relatively undisturbed World Online, downloaded 25 May 2018). Four new evergreen forest since they are not pioneers (Cheek, species were recently described from Cameroon Onana pers. obs. 1992 – 2016). (Onana & Chevillotte 2015; Cheek et al. 2018a) and Species of Vepris in Africa extend from South one more is in press (Onana et al. 2019), taking the Africa, e.g. Vepris natalensis (Sond.) Mziray, to the total in Cameroon to 22 species, the joint highest Guinean woodland in the fringes of the Sahara desert number for any country. Many of these 22 species are (V. heterophylla (Engl.) Letouzey). Mziray (1992) sub- endemic to western Cameroon (SouthWest Region sumed the genera Araliopsis Engl., Diphasia Pierre, and NorthWest Regions of Cameroon) and several are Diphasiopsis Mendonça, Oricia Pierre, Teclea Delile, and threatened (Onana & Cheek 2011). Western Camer- Toddaliopsis Engl. into Vepris, although several species oon is the most species-diverse area for vascular plants were only formally transferred subsequently (e.g. per degree square in tropical Africa and contains Cheek et al. 2009; Onana & Chevillotte 2015). Mziray’s several refugia areas (Barthlott et al. 1996; Cheek et al. conclusions were confirmed by the molecular phylo- 2001). genetic studies of Morton (2017) but sampling was In continental Africa, Vepris are easily recognised. limited, identifications appeared problematic (several They differ from all other Rutaceae because they have species appear simultaneously in different parts of the digitately (1 –)3(– 5)-foliolate (not pinnate) leaves, phylogenetic trees) and more molecular work would and unarmed (not spiny) stems. The genus consists of be desirable. Morton studied about 14 taxa of Vepris, Accepted for publication 7 August 2019. 1 Science, Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AE, UK. e-mail: [email protected] 2 Faculté des Sciences de l’Université de Yaoundé I, BP 812, Yaoundé, Cameroon. 3 Biosciences, College of Life and Environmental Sciences, University of Exeter, Exeter, EX4 4QD, UK. 4 Herbarium, Botany Department, University of Ghana, P.O. Box 55, Legon, Ghana. 5 Herbarium, Botany Department & Trinity Centre for Biodiversity Research, School of Natural Sciences, Trinity College Dublin, Dublin 2, Ireland. © The Author(s), 2019 53 Page 2 of 16 KEW BULLETIN (2019) 74:53 mainly those from eastern Africa. Nevertheless, charac- Onana & Chevillotte (2015) showed that since teristics of some of the former genera are useful today in Verdoorn (1926), authors had unwittingly included grouping species. The “araliopsoid” species have more than one species in Vepris verdoorniana. Onana & subglobose, 4-locular fruit with 4 external grooves; the Chevillotte (2015)linkedStaudt 493, which has “oricioid” species are apocarpous in fruit; the fruits of winged, not terete petioles, with additional specimens “diphasioid” species are laterally compressed in one from Southwest Region Cameroon and described plane, bilocular and bilobed at the apex; while “tecleoid” them as V. letouzeyi Onana, showing that it is a species are unilocular in fruit and 1-seeded, lacking diphasoid, not a tecleoid species. They gave external lobes or grooves. There is limited support for V. letouzeyi the range formerly attributed to these groupings in Morton’s study, V. verdoorniana, i.e. Guinea to Gabon, that is occurring Due to the essential oils distributed in their leaves, in both Upper Guinea (Guinea to Ghana) and Lower and the alkaloids distributed in their roots, several Guinea (Nigeria to Congo Republic). species of Vepris have traditional medicinal value These two phytogeographic units are separated by (Burkill 1997). Burkill details the uses, essential oils the Dahomey Gap (see e.g. Demenou et al. 2017). and alkaloids known from five species in west Africa: Vepris verdoorniana, differentiated among other char- V. hiernii Gereau (as Diphasia klaineana Pierre), acters by terete petioles and based on the type Staudt V. suaveolens (Engl.) Mziray (as Teclea suaveolens Engl.), 472, was now shown to be restricted to the Cameroon V. afzelii (Engl.) Mziray (as T. afzelii Engl.), coastal plain (Onana & Chevillotte 2015), Bioko, V. heterophylla (Engl.) Letouzey (as T. sudanica A.Chev.) Gabon and C.A.R. and V. verdoorniana (Exell & Mendonça) Mziray (as However, comparison of the West African (that is, T. verdoorniana Exell & Mendonça) (Burkill 1997: 651 Upper Guinea) material attributed to, or attributable – 653). Research into the characterisation and anti- to, Vepris verdoorniana by the definition of Keay (1958) microbial and anti-malarial applications of alkaloid shows that it is morphologically separate from both and limonoid compounds in Vepris is active and V. verdoorniana and V. letouzeyi of Lower Guinea. The ongoing, although sometimes published under gener- Upper Guinean (Guinea to Ghana, possibly Benin) ic names no longer in current use, e.g. Wansi et al. winged-petiole material considered to be V. letouzeyi in (2008). Applications include as synergists for insecti- Onana & Chevillotte (2015) is a different taxon, cides (Langat 2011). Cheplogoi et al. (2008)and named below as V. occidentalis. Further, a second Imbenzi et al. (2014) respectively list 14 and 15 species undescribed taxon is present in Upper Guinea, which of Vepris that have been studied for such compounds. had also been included in Teclea verdoorniana by Keay The most recent revision of the African Toddalieae (1958). The species represented by Barker 1181 (K!) was by Verdoorn (1926), who recognised 49 taxa in has opposite leaves, nearly glabrous inflorescences and the genera now included in Vepris.Herrevision very large, pitted fruits. It occurs in the Gola Forest of underpinned subsequent Flora accounts. In the ac- Sierra Leone near the border with Liberia and in count for Conspectus Florae Angolensis,Exell& Liberia at the southern end of Mt Nimba and in the Mendonça (1951) evidenced that she had mistakenly Putu Hills. This species is here termed V. fer Cheek. attributed the name Teclea grandifolia Engl. (now Vepris Here we present a key and table of characters grandifolia (Engl.) Mziray) for which the type is (Table 1) separating the taxa of the Vepris verdoorniana Angolan, to another, undescribed species diagnosed complex based largely on morphology. We also by pedicellate flowers and axillary inflorescences, for present a key and table of characters (Table 2) to all which they published the new name T. verdoorniana the tecloid (unilocular) West African (Upper Guin- Exell & Mendonça. Verdoorn (1926) had cited three ean) species and present a synopsis of the species with specimens for her “Teclea grandifolia” from Cameroon descriptions, illustrations and conservation assessment (Staudt 472, 493 and Mildbraed 9234) and one from of the two new species to science. Sierra Leone (Barter s.n.). Exell & Mendonça (1951) Approximately 2000 new flowering plant species selected one of these, Staudt 472 (K) as type. In the are described each year (Willis 2017), adding to the second edition of Flora of West Tropical Africa, Keay estimated 369,000 already known to science (Nic (1958) recognised three species of Teclea in West Lughadha et al. 2016) although this total is disputed Africa, separating T. verdoorniana from T. afzelii Engl. (Nic Lughadha et al. 2017). Evidence based conser- and T. sudanica A.Chev. (now V. heterophylla) by the vation assessments exist for about 21 – 26% of these possession of inflorescences both axillary and termi- species, and 30 – 44% of these assessments rate the nal, hairy, to 4 cm long, (vs all axillary, glabrous, to species concerned as threatened (Bachman et al. 2.5 cm long), petioles often winged and leaflets sessile 2018). Widespread species tend to have already been (vs non-winged, leaflets petiolulate). Letouzey follow- discovered, so that many newly discovered species ed this delimitation in Flore du Cameroun (1963) and are those that are range-restricted and so more likely pointed out that the Mildbraed specimen is from to be threatened, e.g. Vepris bali Cheek (Cheek et al.
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