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Journal of Ethnopharmacology 103 (2006) 1–24 Review Plectranthus: A review of ethnobotanical uses Catherine W. Lukhoba a , Monique S.J. Simmonds b,∗ , Alan J. Paton b a University of Nairobi, Department of Botany, P.O. Box 30197, Nairobi, Kenya b Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, UK Received 23 May 2005; received in revised form 7 September 2005; accepted 8 September 2005 Abstract Plectranthus is a large and widespread genus with a diversity of ethnobotanical uses. The genus is plagued with numerous nomenclatural disharmonies that make it difficult to collate accurate data on the uses. The aim of this review is to gather together all ethnobotanical information on Plectranthus and to map the data onto the most up-to-date phylogenetic classification in order to see if there are similar uses among related species and hence provide a framework for the prediction and exploration of new uses of species. The uses of 62 species of Plectranthus were mapped onto a current phylogeny based on DNA sequence data. The phylogeny reveals two major Clades, 1 and 2. The members of Clade 1 (corresponding to the formally recognized genus Coleus) were richer in number and diversity of uses than members of Clade 2 (comprising the remaining species of Plectranthus). The high incidence of synonymy can lead to problems in uncovering a species’ ethnobotanical profile. About 30% of all citations of Plectranthus use a synonym and most of the synonyms are attributed to 10 of the most used species, 9 of which are in Clade 1. Members of the ‘Coleus’ Clade are the most studied group both taxonomically and economically. The higher incidence of study may be as a result of the higher diversity of uses and the fact that species in Clade 1, such as Plectranthus barbatus, Plectranthus amboinicus and Plectranthus mollis, are geographically more widespread than those in Clade 2. Plectranthus species in Clade 1 are frequently used as medicines and are used to treat a range of ailments, particularly digestive, skin, infective and respiratory problems. Plectranthus used as foods, flavours, fodder and materials are also mostly found in Clade 1. Monoterpenoids, sesquiterpenoids, diterpenoids and phenolics have been reported in species of Plectranthus. The abietane diterpenoids are the most diverse of the diterpenoids isolated from species of Plectranthus. The labdane diterpenoid, forskolin, occurs in Plectranthus barbatus and could explain some of the traditional uses of this species. This review highlights the fact that not enough is known about the chemistry of other species of Plectranthus to explain their traditional uses. © 2005 Elsevier Ireland Ltd. All rights reserved. Keywords: Plectranthus; Ethnobotanical uses; Coleus Contents 1. 2. 3. ∗ Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Synonymy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Ethnobotanical uses. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3.1. Medicine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3.1.1. Digestive conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3.1.2. Skin conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3.1.3. Respiratory conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3.1.4. Infections and fever . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 3.1.5. Genito-urinary conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 3.1.6. Pain . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Corresponding author. Tel.: +44 208332 5328; fax: +44 208332 5340. E-mail address: m.simmonds@kew.org (M.S.J. Simmonds). 0378-8741/$ – see front matter © 2005 Elsevier Ireland Ltd. All rights reserved. doi:10.1016/j.jep.2005.09.011 2 C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 4. 5. 6. 7. 3.1.7. Muscular–skeletal conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.1.8. Other medical conditions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.2. Horticultural uses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.3. Food . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.4. Food additives . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.5. Fodder . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3.6. Materials . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Geographical distribution . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chemistry of Plectranthus species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ethnobotanical uses and phylogeny . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Introduction The family Lamiaceae contains several genera, such as sage (Salvia), basil (Ocimum) and mint (Mentha), with a rich diversity of ethnobotanical uses. Another important genus is Plectranthus, a large genus containing about 300 species found in Tropical Africa, Asia and Australia. Some species of Plectranthus are difficult to identify because of a lack of clear-cut morphological criteria to discriminate not only among species within the genus but also among the closely related genera. This has resulted in numerous taxonomic problems in the naming of species with the result that species have often been placed in several closely related genera like Coleus, Solenostemon and Englerastrum. In addition, some species formally placed in Plectranthus, are now recognized as the more distantly related Isodon (Paton et al., 2004). Because of these taxonomic issues, different names have been used for the same species of Plectranthus and thus it has been difficult to collate information about the ethnobotanical uses of this genus. Furthermore, the most commonly used medicinal species of Plectranthus have a high degree of synonymy. This review aims to collate data about the different uses of species of Plectranthus, held in disparate sources and frequently under several synonymous Latin names. Information about the uses of the different species was gathered from searching the published literature using the databases NAPRALERT (Pharmacological Sciences (PCRPS), College of Pharmacy, University of Illinois), Web of Science (http://isiknowledge.com), CAB-direct (http://www. cabdirect.org), SEPASAL (http://www.kew.org/ceb/sepasal), Ingenta connect (http://www.ingentaconnect.com/;jsessionid= fq5k5kae0rgi9.victoria), Medline (http://medline.cos.com), Kew Library Catalogue, Elsevier ScienceDirect (www. sciencedirect.com), Dr. Duke’s database (http://www.arsgrin.gov/duke/ethnobot.html) as well as looking through the Floras of countries in Africa, South America, Asia and Australia. Terms used in the searches included ‘ethnobotany, traditional uses, chemicals, phytochemistry, of Plectranthus, Coleus, Solenostemon, Anisochilus and Tetradenia’. All voucher specimens of Plectranthus and Coleus as well as relevant ones of in Solenostemon in herbaria at Kew (K), British Museum (BR), East Africa (EA) and Nairobi (NAI) were also 12 12 13 13 13 13 14 14 14 16 17 18 18 examined to verify the different synonyms used to describe the species and to examine them for notes about their uses. Collectors often made notes either on their vouchers or in their field notebooks about the local use made of species they collect. (Herbarium vouchers cited in the text and/or tables are given at the end of the reference section.) Finally, the ethnobotanical data were mapped onto the most recent phylogeny of the genera (Paton et al., 2004) to see if there is any relationship among species of Plectranthus with similar ethnobotanical uses. 2. Synonymy This review covers the ethnobotanical uses of 62 species of Plectranthus and about 30% of the literature citations covered by this review use synonyms to describe these species. The 10 species with the most uses also have the greatest number of synonyms. For example, Plectranthus barbatus Andr. has commonly been referred to as Plectranthus forskohlii Briq, Plectranthus forskalaei Willd., Plectranthus kilimandschari (Gürke) H.L. Maass., Plectranthus grandis (Cramer) R.H. Willemse, Coleus forskohlii Briq., Coleus kilimandschari Gürke ex Engl., Coleus coerulescens Gürke and Coleus barbatus (Andr.) Benth. Plectranthus amboinicus synonyms include Plectranthus aromaticus Roxb., Coleus aromaticus Benth. and Coleus amboinicus Lour. Those of Plectranthus esculentus N.E.Br are Coleus esculentus (N.E.Br.) G. Tayl., Coleus dazo A. Chev. and Coleus floribundus N.E.Br. and that of Plectranthus vettiveroides (K.C. Jacob) H.I. Maass is Coleus vettiveroides K.C. Jacob. Those of Plectranthus laxiflorus Benth. and Plectranthus mollis (Aiton) Spreng. are Plectranthus albus Gürke and Plectranthus incanus Link, respectively. Synonyms of Plectranthus rotundifolius (Poir.) Spreng. are Coleus rotundifolius (Poir.) A. Chev. & E. Perrot, Coleus dysentericus Bak., Solenostemon rotundifolius (Poir.) J.K. Morton and Plectranthus tuberosus Blume. Plectranthus alpinus (Vatke) O. Ryding has been called Coleus assurgens Bak. and Plectranthus assurgens (Bak.) J.K. Morton. Plectranthus aegyptiacus (Forssk.) C. Chr. is called Plectranthus tenuiflorus (Vatke) Agnew, whereas Plectranthus hadiensis (Forssk.) Schweinf. ex Spreng. is refered to as Plectranthus zeylanicus Benth., Plectranthus cyaneus Gürke ex Engler and Plectranthus zatarhendi (Benth.) E.A. Bruce. The percentage of synonymy would rise if the species Plectranthus vettiveroides (K.C. Jacob) C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 H.I. Maass, which is thought to be conspecific with Plectranthus rotundifolius (Poir.) Spreng., is reduced to synonymy under the latter name, as it is the earliest name. The lack of an up-to-date, authoritative synonomised checklist is a significant barrier to synthesizing knowledge on plantuse from the diverse and fragmented data sources available. The need for such checklists has been identified in other disciplines: for example, in the Global Strategy for Plant Conservation (UNEP, 2004). The fact that most taxonomic revisions of Plectranthus have tended to be regional rather than global could have contributed to the confusion that has ensured the use of multiple taxonomic names to describe the same species. Another issue is the range of common names that can be used for different species of Plectranthus, but collating this information is beyond the remit of this review. 3. Ethnobotanical uses The ethnobotanical data collated for this review have been grouped using the categories in Economic Botany Data Collection Standard (Cook, 1995). A summary of the different ethnobotanical uses of the 62 best documented species of Plectranthus is presented in Tables 1 and 2. 3.1. Medicine The most frequently cited use of species of Plectranthus is for their medicinal properties, which accounts for over 85% of all uses. Plectranthus amboinicus and Plectranthus barbatus are used to treat a wide range of diseases (13 categories) and accounts for about 68% of all traditional uses of the genus (Tables 1 and 2). Plectranthus laxiflorus is used in 10 categories, Plectranthus mollis in 8 categories, both Plectranthus vettiveroides and Plectranthus aegyptiacus in 6, Plectranthus montanus (syn. Plectranthus cylindraceus) in 5 and both Plectranthus sylvestris and Plectranthus bojeri in 4. The rest of the species are used in one to three categories (Table 1). 3.1.1. Digestive conditions Disorders of the digestive system are treated using 21 species of Plectranthus (Table 1). Species are used to treat stomach pain, nausea, vomiting, diarrhoea, mouth and throat infections and are used as purgatives, carminatives and as antihelmintics. Plectranthus barbatus, Plectranthus amboinicus, Plectranthus laxiflorus, Plectranthus esculentus and Plectranthus vettiveroides are the most frequently used species and Plectranthus barbatus and Plectranthus amboinicus are used to treat a wide variety of digestive problems. For instance, Plectranthus barbatus is used for the treatment of stomachache and as a purgative (Rounce, 1933; Jarrett, 1950; Johns et al., 1990; Kokwaro, 1993; Simon and Mollel, 1999), for nausea (Hamill et al., 2003) and for gastritis and intestinal spasms in Brazil (Camara et al., 2003). Plectranthus amboinicus is popular in the treatment of dyspepsia, indigestion, diarrhoea and as a carminative in India and Africa (Morton, 1992; Gurib-Fakim et al., 1996; Jain and Lata, 1996; Ong and Nordiana, 1999). Plectranthus caninus, Plectranthus laxiflorus and Plectranthus barbatus are used in the treatment of teeth and gum disorders (Hulme, 3 1954; Kokwaro, 1993; Meyerhoff, 1978b). Plectranthus laxiflorus is also used as a purgative (Kokwaro, 1993), while Plectranthus defoliatus is used to treat diarrhoea (Schlage et al., 2000). Plectranthus vettiveroides is used in India to treat stomachaches, dyspepsia, nausea and vomiting (Dash and Kashyap, 1987; Yoganarasimhan, 2000). Plectranthus esculentus and the leaves of Plectranthus aegyptiacus are also used to treat stomachache (Morris, 1996; Parkia and Cooke, 2003). In Eastern and Southern Africa, both Plectranthus elegans and Plectranthus esculentus are reported to be used as anthelmintics (Kokwaro, 1993; Burkill, 1995; Allemann et al., 2004). 3.1.2. Skin conditions A total of 20 species are recorded as being used for skin conditions (Table 1). Plectranthus barbatus and Plectranthus amboinicus are the most frequently cited species for the treatment of burns, wounds, sores, insect bites and allergies. In Kenya and the Democratic Republic of Congo, Plectranthus barbatus is used in the treatment of wounds and ringworms (Githinji and Kokwaro, 1993; Chifundera, 2001), to reduce swelling on bruises (Gentry, 1970) and for bathing babies suffering from measles (Kokwaro, 1993). This species has anti-ageing effects in combination with tocotrienol (Adachi et al., 1996). The whole plant contains a forskolin-like compound that is used in hair dyeing (Sugiyama et al., 1988). In addition, Plectranthus barbatus contains an essential oil that exhibits anti-allergic activities through passive cutaneous anaphylaxis inhibition (Gupta et al., 1993a). Plectranthus amboinicus is used in Brazil for the treatment of skin ulcerations caused by Leishmania braziliensis (França et al., 1996). It is also used to treat burns and as a poultice for centipedes and scorpion bites in Malay (Morton, 1992). In India, the juice of the leaves is used to treat skin allergies (Harsha et al., 2003). Other species used to treat skin allergies include Plectranthus amboinicus (Harsha et al., 2003) and Plectranthus asirensis which is used in Saudi Arabia for diaper rash and itching, as well as an antiseptic dressing (Abulfatih, 1987a,b). The leaves of Plectranthus congestus are used in Papua New Guinea as an antiseptic dressing for wounds (Holdsworth, 1977; Woodley, 1991) and in the treatment of scabies (Holdsworth and Rali, 1989). Plectranthus madagascariensis is also used in the treatment of scabies and small wounds (Watt and Breyer-Brandwijk, 1962; Roberts, 1990; Rabe and van Staden, 1998; Neuwinger, 2000), the finely ground leaves of Plectranthus bojeri are applied as dressing on wounds and abscesses (Neuwinger, 2000), Plectranthus laxiflorus is rubbed onto the skin to treat leprosy (Chifundera, 2001), Plectranthus vettiveroides is a hair tonic (Yoganarasimhan, 2000) and Plectranthus fruticosus was used in traditional Romanian medicine for its healing properties, especially for treating burns (Pages et al., 1991). Plectranthus ecklonii is used in Zimbabwe to treat skin infections (Nyanyiwa and Gundidza, 1999) while in Rwanda, the essential oil of Plectranthus sylvestris is used to treat skin diseases (Chalchat et al., 1996). 3.1.3. Respiratory conditions A total of 15 species are recorded for the treatment of respiratory conditions. Plectranthus barbatus is the most fre- 4 Table 1 Medicinal uses of species of Plectranthus categories are defined in Cook (1995) Species Geographical distribution Plectranthus aegypticus (Forssk.) C. Chr. Plectranthus tenuiflorus (Vatke) Agnew Arabia, East and Northeast Tropical Africa Plectranthus alpinus (Vatke) O. Ryding Plectranthus ambiguus (Bolus) Codd Plectranthus assurgens (Baker) J.K. Morton East and Central Africa Plectranthus amboinicus (Lour.) Spreng. Coleus amboinicus Lour., Coleus aromaticus Benth., Plectranthus aromaticus Roxb. Plectranthus asirensis J.R.I. Wood Plectranthus barbatus Andr. Plectranthus beddomei Raiz. Plectranthus bojeri (Benth.) Hedge Plectranthus caninus Roth Medicinal categories Digestive Skin Parkia and Cooke (2003) Meyerhoff (1978c), Bos et al. (1983), de Padua (1988), Morton (1992), Gurib-Fakim et al. (1996), Jain and Lata (1996), Rabe and van Staden (1998), Ong and Nordiana (1999), Rajendran et al. (1999), Yoganarasimhan (2000), Hadi and Bremner (2001) Arabia Coleus kilimandschari Gurke ex Engl., Coleus barbatus (Andr.) Benth., Coleus forskohlii Briq., Coleus comosus A. Rich., Plectranthus kilimandschari (Gürke) H.L. Maas, Plectranthus forskohlii auct Briq. North, East, Central Africa, Asia, South America de Padua (1988), Morton (1992), França et al. (1996), Neuwinger (2000), Harsha et al. (2003) Genito-urinary Abulfatih (1987b), Othman and Shoman (2004) Parkia and Cooke (2003) Pain Muscular–skeletal Hedberg et al. (1983), Neuwinger (2000) Hulme (1954), Hutchings et al. (1996), Rabe and van Staden (1998), Neuwinger (2000) Ayensu (1978), de Padua (1988), Morton (1992), Jain and Lata (1996), Ruiz et al. (1996), Castillo and González (1999), Yoganarasimhan (2000), Cano and Volpato (2004) Morton (1992), Gurib-Fakim et al. (1996), Jain and Lata (1996), Harsha et al. (2002) Morton (1992), Jain and Lata (1996), Ayensu (1998), Neuwinger (2000), Yoganarasimhan (2000) Meyerhoff (1978c), de Padua (1988) Meyerhoff (1978c) Williams (1975), Holdsworth (1977), Milliken (1977), Meyerhoff (1978b), Baerts and Lehmann (1989), Githinji and Kokwaro (1993), Gupta et al. (1993a,b), Rwangabo (1993), Rajendran et al. (1999), Neuwinger (2000), Matu and van Staden (2003), Vigneron et al. (2005) Watt and Breyer-Brandwijk (1962), Innamorati (1973), Boily and van Puyvelde (1986), Rwangabo (1993), Neuwinger (2000), Chifundera (2001), Cos et al. (2002) Neuwinger (2000), Yoganarasimhan (2000), Chifundera (2001) Baerts and Lehmann (1989), Githinji and Mwangangi (1989), Githinji and Kokwaro (1993), Kokwaro (1993), Rwangabo (1993), Neuwinger (2000) Abulfatih (1987a,b) Bally (1934, 1937), Jarrett (1950), Hulme (1954), Meyerhoff (1978a,b,c), Baerts and Lehmann (1989), Johns et al. (1990), Githinji and Kokwaro (1993), Gupta et al. (1993a,b), Kokwaro (1993), Rwangabo (1993), Johns et al. (1995), Rounce (1933), Rodrı́guez et al. (1995), Simon and Mollel (1999), Neuwinger (2000), Yoganarasimhan (2000), Chifundera (2001), Geissler et al. (2002), Camara et al. (2003), Costa and Nascimento (2003), Hamill et al. (2003) Baerts and Lehmann (1989), Gachathi (1989), Kokwaro (1993), Rwangabo (1993), Neuwinger (2000), Chifundera (2001) India Rajendran et al. (1999) Madagascar Neuwinger (2000) East Tropical Africa Infections/fevers Al-Yahya et al. (1985) Burkill (1995) South Africa North, East, Central Africa, Asia, South America, Caribbean, Pacific Respiratory Kokwaro (1993), Neuwinger (2000) Bouquet (1969), Boily and van Puyvelde (1986), Baerts and Lehmann (1989), van Puyvelde et al. (1994), Rajendran et al. (1999), Yoganarasimhan (2000), Chifundera (2001), Cos et al. (2002), Benerjee (2003), Schanberg and Ikan (2003) Neuwinger (2000) Githinji (1990), Githinji and Kokwaro (1993) C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Synomyms encountered in the study Plectranthus coeruleus (Gürke) Agnew Plectranthus congestus R.Br. East Tropical Africa Australia Plectranthus variifolius De Wild Plectranthus esculentus N.E.Br. Coleus dazo A. Chiev., Coleus esculentus G. Tayl., Plectranthus floribundus N.E.Br. Plectranthus hereroensis Engl. Plectranthus igniarius (Schweinf.) Agnew Plectranthus insignis Hook.f. Plectranthus kamerunensis (Gürke) Plectranthus lactiflorus (Vatke) Agnew. Plectranthus lanceolatus Bojer ex Benth. Plectranthus lanuginosus (Benth.) Agnew. Plectranthus laxiflorus Benth. Plectranthus longipes Baker Africa-DRC Neuwinger (2000) Asia Tropical Schlage et al. (2000) South Africa East Tropical Africa East and Southern Africa Southern Africa, West-Central Africa, Asia Tropical Nyanyiwa and Gundidza (1999) Githinji and Kokwaro (1993) Kokwaro (1993), Neuwinger (2000) West-Central Africa–Cameroon South Africa Plectranthus zeylanicus Benth., Plectranthus cyaneus Gürke ex Engler, Plectranthus zatarhendi (Benth.) E.A. Bruce Plectranthus myrianthus Briq. Northeast and East Tropical Africa to India Mehrotra et al. (1989) Sothern Africa Batista et al. (1995), Ferraeira et al. (1997) Tanner (1959) Hutchings et al. (1996) Timberlake (1987) West-Central Africa–Cameroon West and East Tropical Africa Cheek et al. (2000) East Tropical Africa–Tanzania Africa–Malgaches Tanner (1961b) East Tropical Africa Kokwaro (1993), Wirtu et al. (1997), Neuwinger (2000) Roberts (1990), Kokwaro (1993), Hutchings et al. (1996), Rabe and van Staden (1998), Neuwinger (2000) Neuwinger (2000) East Tropical Africa–Kenya Morris (1996), Neuwinger (2000) Ngassoum (2001) East Tropical Africa Plectranthus amaniensis Gürke Morris (1996) Bally (1937), Githens (1949), Neuwinger (2000) Pages et al. (1991) Plectranthus urticoides Baker West-Central Africa, East and Southern Africa Neuwinger (2000) Burkill (1995), Neuwinger (2000) South Africa Plectranthus albus Gürke Woodley (1991) Holdsworth (1977), Holdsworth and Rali (1989), Woodley (1991) C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Plectranthus decurrens (Gürke) J.K. Morton Plectranthus defoliatus Hochst. ex Benth. Plectranthus ecklonii Benth. Plectranthus edulis (Vatke) Agnew Plectranthus elegans Britten Plectranthus fruticosus L’Her. Plectranthus glandulosus Hook.f. Plectranthus grallatus Briq. Plectranthus hadiensis (Forssk.) Schweinf. ex Spreng. Neuwinger (2000) Githinji and Kokwaro (1993) Lusigi et al. (1984) Githinji and Kokwaro (1993) Kakudidi et al. (1996), Neuwinger (2000), Chifundera (2001) Neuwinger (2000) Neuwinger (2000) Hutchings et al. (1996), Rabe and van Staden (1998), Neuwinger (2000) Watt and Breyer-Brandwijk (1962), Roberts (1990), Hutchings et al. (1996), Rabe and van Staden (1998) Lindsay and Hepper (1978), Kokwaro (1985), Neuwinger (2000) Kokwaro (1993), Neuwinger (2000) Kokwaro (1993), Neuwinger (2000) 5 6 Table 1 (Continued ) Species Geographical distribution Plectranthus hirtus Benth. Southern Africa Plectranthus luteus Gürke East Africa Madagascar Plectranthus incanus Link India Plectranthus montanus Benth. Plectranthus cylindraceus Hochst. ex Benth. East Tropical Africa Plectranthus parviflorus (Poir.) Henckel Plectranthus prostratus Gürke Plectranthus pseudomarrubioides Willemse Plectranthus pubescens Baker Plectranthus punctatus L’Her. Plectranthus stachyoides Oliv. Plectranthus stolzii Gilli Plectranthus sylvestris Gürke Plectranthus tetensis (Bak.) Agnew Plectranthus vettiveroides (K.C. Jacob) H.I. Maass Plectranthus austalis R.Br., Coleus parviflorus Benth. Australia, Pacific Plectranthus madagascarensis Benth. Plectranthus mandalensis Baker Plectranthus melleri Baker Plectranthus mollis (Aiton) Spreng. Respiratory Holdsworth (1977), Roberts (1990), Rabe and van Staden (1998), Neuwinger (2000) Githens (1949), Roberts (1990), Rabe and van Staden (1998), Neuwinger (2000) Infections/fevers Genito-urinary Pain Morris (1996) Jain et al. (1994) Timberlake (1987), Githinji and Kokwaro (1993), Kokwaro (1993) Yoganarasimhan (2000) Yoganarasimhan (2000) Rahman et al. (2004) Kokwaro (1993) Sebastian and Bhandari (1984), Yoganarasimhan (2000) Kokwaro (1993) Tewtrakul et al. (2003) Mugisha (2002) Luke (2004) Morris (1996) Malawi Northeast and East Tropical Africa–Ethiopia East Tropical Africa–Tanzania East Tropical Africa–Tanzania East and Central Africa Hedberg (1979) Tanner (1961a) Gaetan (1952) Githinji and Kokwaro (1993) Chalchat et al. (1996), Neuwinger (2000) Kokwaro (1993), Neuwinger (2000) Meyerhoff (1978a) Dash and Kashyap (1987), Yoganarasimhan (2000) Yoganarasimhan (2000) Yoganarasimhan (2000) East Tropical Africa Coleus vettiveroides K.C. Jacob Asia Tropical–India Asia Tropical–Papua New Guinea East Tropical Africa–Tanzania Synomyms encountered in the study Geographical distribution Plectranthus tenuiflorus (Vatke) Agnew Plectranthus assurgens (Baker) J.K. Morton Coleus amboinicus Lour., Coleus aromaticus Benth., Plectranthus aromaticus Roxb. Arabia, East and Northeast Tropical Africa East and Central Africa Kokwaro (1993), Neuwinger (2000) Yoganarasimhan (2000) Holdsworth (1977) Tanner (1961c) Kokwaro (1993) Medicinal categories Circulatory and blood North, East, Central Africa, Asia, South America, Caribbean, Pacific Muscular–skeletal Yamada (1999) East Tropical Africa, Uganda East Tropical Africa, Asia temperate Plectranthus sp. Plectranthus aegypticus (Forssk.) C. Chr. Plectranthus alpinus (Vatke) O. Ryding Plectranthus amboinicus (Lour.) Spreng. Skin Africa–Malawi Plectranthus sp. Species Medicinal categories Digestive Nervous Sensory Poisons treatment Abulfatih (1987b) Unspecific Inflammation Othman and Shoman (2004) Hedberg et al. (1983) Morton (1992), Neuwinger (2000) Ayensu (1978), Morton (1992), Jain and Lata (1996), Ruiz et al. (1996), Salmán et al. (1996), Neuwinger (2000), Camara et al. (2003) Zepernick (1972), Morton (1992) Yoganarasimhan (2000) Kuebel and Tucker (1988), Prudent et al. (1995), França et al. (1996), Hutchings et al. (1996), Bennett and Prance (2000) Morton (1992) C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Synomyms encountered in the study Plectranthus barbatus Andr. Coleus kilimandschari Gurke ex Engl., Coleus barbatus (Andr.) Benth., Coleus forskohlii Briq., Coleus comosus A. Rich., Plectranthus kilimandschari (Gürke) H.L. Maass, Plectranthus forskohlii auct Briq. North, East, Central Africa, Asia, South America Madagascar Plectranthus fruticosus L’Her. Plectranthus glandulosus Hook.f. Plectranthus grallatus Briq. Plectranthus grandidentatus Gürke Plectranthus hadiensis (Forssk.) Schweinf. ex Spreng. South Africa Plectranthus igniarius (Schweinf.) Agnew Plectranthus kivuensis (Lebrun & Touss.) R.H. Willemse Plectranthus lanceolatus Bojer ex Benth. Plectranthus lanuginosus (Benth.) Agnew. Plectranthus laxiflorus Benth. Plectranthus longipes Baker Plectranthus madagascarensis Benth. Plectranthus mandalensis Baker Plectranthus mollis (Aiton) Spreng. Plectranthus montanus Benth. Plectranthus occidentalis B.J. Pollard Plectranthus urticoides Baker Neuwinger (2000), Chifundera (2001) Baerts and Lehmann (1989), Neuwinger (2000), Geissler et al. (2002), Benerjee (2003) Greenway (1946), Tanner (1946), Akendenué and Louis (1994) Neuwinger (2000) Neuwinger (2000) Fratkin (1975), Caufield (1978), Githinji and Kokwaro (1993), Svoboda (1996), Barnett (2000), Bennett and Prance (2000), Byavu et al. (2000) Australia Maiden (1899) East Tropical Africa Githinji (1990), Lukhoba and Mathenge (1993) Pages et al. (1991) West-Central Africa–Cameroon South Africa Yoganarasimhan (2000), Chifundera (2001) Cheek et al. (2000) Hutchings et al. (1996) Cerqueira et al. (2004) Plectranthus zeylanicus Benth., Plectranthus cyaneus Gürke ex Engler, Plectranthus zatarhendi (Benth.) E.A. Bruce Neuwinger (2000) Northeast and East Tropical Africa to India East Tropical Africa Timberlake (1987) East Tropical Africa Napier (1933) Africa–Malgaches East Tropical Africa Kakudidi et al. (1996) Kokwaro (1993), Neuwinger (2000) Lindsay and Hepper (1978), Neuwinger (2000) Plectranthus albus Gürke West-Central Africa, East and Southern Africa Plectranthus amaniensis Gürke Plectranthus hirtus Benth. East Tropical Africa–Kenya Neuwinger (2000) Southern Africa Fleurentin et al. (1983) Africa–Malawi Plectranthus incanus Link India Plectranthus cylindraceus Hochst. ex Benth. Solenostemon mannii (Hook. f.) Baker East Tropical Africa West Africa C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Plectranthus bojeri (Benth.) Hedge Plectranthus congestus R.Br. Plectranthus edulis (Vatke) Agnew Neuwinger (2000), Yoganarasimhan (2000), Chifundera (2001) Morris (1996) Yoganarasimhan (2000) Singh and Ali (1992) Jain et al. (1994) Sebastian and Bhandari (1984) Fleurentin et al. (1983), Barnett (2000) Harvey et al. (2004) 7 C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Gaetan (1952), Cunningham and Barigyira (1992) Shankar and Majumdar (1995), Tandon (1996) Lovett and Kayombo (1989a,b) Yoganarasimhan (2000) Morris (1996) Plectranthus pubescens Baker Plectranthus punctatus L’Her. Plectranthus sp. aff. occidentalis Plectranthus stolzii Gilli East Tropical Africa–Tanzania Asia Tropical–India Plectranthus vettiveroides (K.C. Jacob) H.I. Maass Plectranthus viphyensis Brummitt & J.H. Seyani Coleus vettiveroides K.C. Jacob East Tropical Africa–Tanzania Northeast and East Tropical Africa–Ethiopia West Tropical Africa Malawi Medicinal categories Species Table 1 (Continued ) Synomyms encountered in the study Geographical distribution Circulatory and blood Nervous Sensory Hedberg (1979) Poisons treatment Unspecific Leeuwenberg (1972) Inflammation 8 quently cited species used to relieve colds (Rajendran et al., 1999), cough (Baerts and Lehmann, 1989; Neuwinger, 2000; Yoganarasimhan, 2000; Chifundera, 2001), bronchitis (Boily and van Puyvelde, 1986; Cos et al., 2002), pneumonia (Bouquet, 1969) and for general respiratory ailments (van Puyvelde et al., 1994). Studies have shown that it reverses allergic bronchospasm (Kreutner et al., 1985). Plectranthus amboinicus is also frequently cited in the treatment of chronic coughs, asthma, bronchitis and sore throat in India and the Carribean (Morton, 1992; Jain and Lata, 1996; Ruiz et al., 1996), and in Cuba it is used to treat catarrhal infections and asthma (Castillo and González, 1999; Cano and Volpato, 2004). The leaves of Plectranthus amboinicus have been found to have bronchodilator activity in guinea pig and anti-Mycobacterium tuberculosis activity (Carbajal et al., 1991; Frame et al., 1998). Plectranthus madagascariensis is used for colds (Githens, 1949), cough and chest complaints (Watt and Breyer-Brandwijk, 1962; Rabe and van Staden, 1998; Neuwinger, 2000) and asthma (Roberts, 1990). A decoction of the whole plant of Plectranthus bojeri, together with Ocimum gratissimum, is drunk to treat pneumonia (Neuwinger, 2000). Plectranthus aegyptiacus is used in Saudi Arabia, for the treatment of sore throats and laryngitis (Al-Yahya et al., 1985) and in Kenya, Plectranthus elegans is used to alleviate sore throats (Bally, 1937). Plectranthus montanus (Rahman et al., 2004) and Plectranthus tetensis (Meyerhoff, 1978a) are used to treat sore throats. Plectranthus mollis is used in India as a respiratory stimulant and vasoconstrictor (Yoganarasimhan, 2000), while extracts from the roots of Plectranthus caninus are used in Kenya for the treatment of coughs (Githinji, 1990). In South Africa, tea made from the leaves of Plectranthus laxiflorus is taken for coughs and colds (Hutchings et al., 1996; Rabe and van Staden, 1998) and an infusion of the crushed leaves of Plectranthus ambiguus is mixed with a little hot water and sipped for coughs (Hulme, 1954; Hutchings et al., 1996). The roots of Plectranthus stolzii are eaten in Tanzania for coughs (Gaetan, 1959). 3.1.4. Infections and fever Fifteen species have been recorded for the treatment of infections and fever. Plectranthus barbatus is used to treat a range of infections including throat and mouth infections (Meyerhoff, 1978b), tonsillitis (Neuwinger, 2000), gastro-intestinal infections (Baerts and Lehmann, 1989; Gupta et al., 1993b), genitourinary infections (Rwangabo, 1993; Neuwinger, 2000) as well as ear and eye infections (Baerts and Lehmann, 1989). It is used to treat malaria in Brazil and to break fevers in East Africa and India (Holdsworth, 1977; Rwangabo, 1993; Githinji and Kokwaro, 1993; Milliken, 1977; Rajendran et al., 1999; Neuwinger, 2000; Schlage et al., 2000; Steele et al., 2002), whereas in Kenya it is used for the treatment of East Coast Fever in cattle (Ole-Marion, 2003). The whole plant is said to have antibacterial, antiviral and antifungal activity (Boily and van Puyvelde, 1986; van Puyvelde et al., 1994; Vlietinck et al., 1995; Alasbahi et al., 1999; Cos et al., 2002; Matu and van Staden, 2003) and against protozoa, such as Giardia (Johns et al., 1995). Plectranthus amboinicus is important in Asia and South America for the treatment of fevers (Morton, 1992; Harsha et Table 2 Non-medicinal uses of species of Plectranthus Species Synomyms encountered Geographical in the study distribution Non-medicinal uses Poisons Plectranthus caninus East Tropical Africa Roth Plectranthus ciliatus E. Plectranthus natalensis South Africa Mey. Bak. Plectranthus crassus N.E.Br. Plectranthus decurrens (Gürke) J.K. Morton Plectranthus ecklonii Benth. Soil improvers Food Schoelhorn (2002) Dymoc (1885) Plectranthus edulis (Vatke) Agnew East Tropical Africa Plectranthus elegans Britten Plectranthus esculentus Coleus dazo A. Chiev., N.E.Br. Coleus esculentus G. Tayl., Plectranthus floribundus N.E.Br. East and Southern Africa Southern Africa, West-Central Africa, Asia Tropical Animal feeds Materials Social uses Bees Yuncker (1953), Morton (1992), Morton (1992), Prudent et al. (1995) Prudent et al. (1995) Epling (1981), Purseglove (1987), Craig and Mayenda (1990), Mayenda (1991), Morton (1992), Prudent et al. (1995), Brown (1997) Schoelhorn (2002) Owili (1977), Mitchell and Rook (1979) Grimshaw (1993), Bennett and Prance (2000) Maher (1935), Hendy (1977), Caufield (1978) Fleurentin et al. (1983), Drummond and Battiscombe Wiersema and Leon Hemsley (1953), (1912), Rounce Mwangangi (1982) (1999) (1933), Bush (1943), Templer (1960), Rwaburindore (1969), Mathew (1970), Mwangangi (1982), Riley and Brokensha (1988), Gachathi (1989) Githinji (1990) Bally (1934), Gentry (1969, 1970), Mwangangi (1971, 1982), Brokensha and Riley (1978), Githinji (1988) Tabuti et al. (2003) Githinji (1990) Githinji (1990) Roberts (1990), Rabe and van Staden (1998) Malawi Eest-Central Africa-DRC South Africa Food additives Cheek et al. (2000) Australia North, East, Central Africa, Asia, South America Hedges Spicy hu (2004) Yamada (1999) Auchland Regional Council (2002), Schoelhorn (2002) Fleurentin et al. (1983), Lukhoba and Mathenge (1993) Githinji (1990) Napier (1933) C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Plectranthus argentatus S.T. Blake Plectranthus barbatus Coleus kilim & schari Andr. Gurke ex Engl., Coleus barbatus (&r.) Benth., Coleus forskohlii Briq., Coleus comosus A. Rich., Plectranthus kilim & schari (Gürke) H.L. Maass., Plectranthus forskohlii auct Briq. Ornmamentals Rahman et al. (2004) Plectranthus aegypticus Plectranthus tenuiflorus Arabia, East and (Forssk.) C. Chr. (Vatke) Agnew Northeast Tropical Africa Plectranthus alpinus Plectranthus assurgens East and Central (Vatke) O. Ryding (Baker) J.K.Morton Africa North, East, Central Plectranthus amboinicus Coleus amboinicus Africa, Asia, South (Lour.) Spreng. Lour., Coleus America, Caribbean, aromaticus Benth., Plectranthus aromaticus Pacific Roxb. Jaarsveld (1991) Allemann and Ward (1935), Msyla Hammes (1998) (1944), Semsei (1956a,b), Fleurentin et al. (1983), Purseglove (1987), Burkill (1995), Morris (1996), Mooi et al. (1999), Cheek et al. (2000), Reinten and Cootzee (2002), Allemann and Hammes (2003), Allemann et al. (2004) 9 10 Table 2 (Continued ) Species Synomyms encountered Geographical in the study distribution Non-medicinal uses Poisons Plectranthus fruticosus L’Her. Plectranthus glandulosus Hook.f. Plectranthus graveolens R.Br. Plectranthus hadiensis (Forssk.) Schweinf. Ex Spreng. South Africa Plectranthus urticoides Baker West-Central Africa–Cameroon Australia Hedges Soil improvers Food Harvey et al. (2004) Plectranthus longipes Baker Plectranthus amaniensis East Tropical Gürke Africa–Kenya Plectranthus madagascarensis Benth. Plectranthus melleri Baker Plectranthus mollis (Aiton) Spreng. Plectranthus hirtus Benth. Southern Africa Plectranthus luteus Gürke Plectranthus incanus Link East Africa Madagascar India Tanzania Bees Roberts (1990) Hutchings et al. NMK Expd. (1996) (1974), Timberlake (1987) Sato (1976), Timberlake (1987) Cheek et al. (2000) Glover et al. (1961a) Glover et al. (1961a) Cheek et al. (2000) Githinji and Kokwaro (1993) Parkia and Cooke (2003) Hepper and Jaeger (1978) Neuwinger (2000) Koritschoner (1935a,b), Kakudidi et al. (1996) Glover et al. (1961b), Glover and Samuel (1962) Schoelhorn (2002) Yamada (1999) Maikhuri and Gangwar (1993) Jain et al. (1994), Yoganarasimhan (2000) Jain et al. (1994) Tanner (1952b) Wiersema and Leon (1999), Welsh and Cotner (2000) Tourle (2003), Schmidt (2005) Australia, Pacific East Tropical Africa, Ug&a East Tropical Africa, Asia temperate Social uses Roberts (1990) Tanner (1952a) East Tropical Africa Plectranthus cylindraceus Hochst. ex Benth. Brazil, South Africa Plectranthus oertendahlii T.C.E.Fr. Plectranthus parviflorus Plectranthus austalis (Poir.) Henckel R.Br., Coleus parviflorus Benth. Plectranthus prostratus Gürke Plectranthus Plectranthus pseudomarrubioides marrubioides Hochst. Willemse Plectranthus radiatus A.J. Paton in ed. Materials Anon. (1874) East Tropical Africa–Tanzania East Tropical Africa West-Central Africa, East and Southern Africa Animal feeds Ramachandran and Nair (1981), Purseglove (1987) McNew (2002) Omolo et al. (2004) Meyer (1964) Zaroug (1981) Singh et al. (2003) C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 West-Central Africa–Cameroon West and East Tropical Africa Food additives Schoelhorn (2002) Plectranthus zeylanicus Northeast and East Tropical Africa to Benth., Plectranthus India cyaneus Gürke ex Engler, Plectranthus zatarhendi Benth.) E.A. Bruce East Tropical Africa Plectranthus igniarius (Schweinf.) Agnew Plectranthus insignis Hook.f. Plectranthus kamerunensis (Gürke) Plectranthus lactiflorus (Vatke) Agnew. Plectranthus lanuginosus (Benth.) Agnew. Plectranthus laxiflorus Plectranthus albus Benth. Gürke Plectranthus montanus Benth. Ornmamentals C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Lusigi et al. (1984) Riley and Brokensha (1988) Winter and Leistner (1957) Githinji and Kokwaro (1993) al., 2002) and as a cure of cholera in Rodrigues (Gurib-Fakim et al., 1996). It also has antimicrobial activity (Bos et al., 1983; Castillo and González, 1999) and is reported to have antiviral activity against Herpes simplex virus-1 (Hattori et al., 1995) and anti-HIV inhibition activity (Kusumoto et al., 1995). Plectranthus aegyptiacus and Plectranthus punctatus are used to treat ear infections in Ethiopia (Hedberg, 1979) and Saudia Arabia (Abulfatih, 1987b), respectively. Plectranthus glandulosus is used to treat influenza in Cameroon (Ngassoum, 2001) and Plectranthus sylvestris is given to patients with malaria in East Africa (Kokwaro, 1993). A decoction of the whole plant of Plectranthus bojeri is used in Madagascar, to bathe children in order to reduce fever (Neuwinger, 2000). Plectranthus mollis is a recognized febrifuge (Varma and Sharma, 1963; Yoganarasimhan, 2000), whereas Plectranthus laxiflorus is used by the Pokot in Kenya as an enema for the treatment of influenza (Watt and BreyerBrandwijk, 1962). Plectranthus fruticosus is also antimicrobial and this activity may be attributed, in part, to the presence of kaurane diterpenoids (Gaspar-Marques et al., 2003). Both Plectranthus ecklonii and Plectranthus montanus have antibacterial and antifungal activity (Chadya and Gundidza, 1999; Nyanyiwa and Gundidza, 1999). Recently, Plectranthus parvifolius was shown to contain compounds that inhibit HIV-1 integrase (Tewtrakul et al., 2003). Coe (2001) South Africa Europe–Belgium Plectranthus nummularius Briq. Southern Africa Plectranthus ungentarius Codd Plectranthus verticillatus Druce Plectranthus zuluensis T. Cooke Plectranthus sp. Plectranthus dissitiflorus West Tropical Africa (Gürke) J.K. Morton East and Central Africa Plectranthus sp. aff. occidentalis Plectranthus sylvestris Gürke Plectranthus saccatus Benth. Plectranthus rotundifolius (Poir.) Spreng. Coleus rotundifolius (Poir.) A. Chev. & E. Perrot.; Solenostemon rotundifolius (Poir.) J.K. Morton South Africa Motolese et al. (1991) Buitendag (1971) Southern Africa Joffe (1993, 2001), Kahurananga and Kiilu (1976) Leeuwenberg (1972) Riley and Brokensha (1988) Fleurentin et al. (1983) Mooney (1954), Tadesse (1967), Scott (1979) Northeast and East Tropical Africa–Ethiopia Tropical Asia, Southern Africa Plectranthus punctatus L’Her. 11 3.1.5. Genito-urinary conditions Eight species are used to treat genito-urinary conditions. This includes Plectranthus barbatus that is used in the treatment of syphilis in Central Africa (Boily and van Puyvelde, 1986; Cos et al., 2002), as an emmenagogue and oral abortifacient or contraceptive (Watt and Breyer-Brandwijk, 1962; Almeida and Lemonica, 2000) and as an aphrodisiac in Somalia (Innamorati, 1973). The contraceptive property of this species is corroborated by the fact that the leaves were found to have an anti-implantation effect in rats (Almeida and Lemonica, 2000). The leaves of Plectranthus amboinicus are frequently utilized in the treatment of urinary diseases in the Amazon and India (Jain and Lata, 1996; Yoganarasimhan, 2000). This species is also reported to relieve kidney troubles, treat vaginal discharges and is drunk after childbirth (Morton, 1992). The Giriama of Kenya use Plectranthus aegypticiacus as a medicine to treat ailments associated with sexual misconduct (Parkia and Cooke, 2003), while in Marakwet, Kenya, Plectranthus laxiflorus is used as an abortifacient (Lindsay and Hepper, 1978; Kokwaro, 1985). Plectranthus vettiveroides is used as an emmenagogue in India (Yoganarasimhan, 2000). Plectranthus prostratus is used by women in Uganda to increase sexual stimulation and vaginal fluids (Mugisha, 2002), while Plectranthus elegans is a potency medicine in Malawi (Morris, 1996). In laboratory experiments, Plectranthus caninus has been found to have diuretic activity (Painuly and Tandon, 1983; Aswal et al., 1984), although we could not find any ethnobotanical information to support this activity. 3.1.6. Pain Plectranthus barbatus is used for the treatment of gastric and spasmodic pain in Africa and Asia (Dubey et al., 1981; 12 C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Yoganarasimhan, 2000; Chifundera, 2001), abdominal pain (Baerts and Lehmann, 1989; Neuwinger, 2000), painful micturition (Dubey et al., 1981), muscular pains (Githinji and Kokwaro, 1993) and for generalized pain (Elisabetsky and Castilhos, 1990; Woodley, 1991). In Kenya, an extract of Plectranthus sylvestris is inserted into two holes cut in the chest to relieve chest pain. When the leaf sap of Plectranthus sylvestris is mixed with decoctions of Senna didymototrya and Clerodendron johnsonii it is drunk for headache and abdominal pain and Plectranthus montanus mixed with Microglossa oblongifolia is a remedy for fever and severe headache (Kokwaro, 1993). In Africa, Plectranthus amboinicus is a remedy for headaches (Meyerhoff, 1978c; de Padua, 1988) as is Plectranthus esculentus (Morris, 1996) and Plectranthus laxiflorus is a treatment for abdominal pain (Kokwaro, 1993; Neuwinger, 2000). 3.1.7. Muscular–skeletal conditions Six species of Plectranthus are used to treat musculo-skeletal conditions. Plectranthus amboinicus and Plectranthus barbatus are used for treating stiff neck and backache (Meyerhoff, 1978b; Githinji, 1990; Githinji and Kokwaro, 1993). The leaves of Plectranthus barbatus when burned a little and placed on the skin of the neck relieve stiffness (Githinji and Mwangangi, 1989). Plectranthus barbatus is also used in the treatment of bone dislocations (Baerts and Lehmann, 1989). In Africa, Plectranthus barbatus (Rwangabo, 1993), Plectranthus laxiflorus (Koritschoner, 1935b; Kokwaro, 1993; Neuwinger, 2000), Plectranthus mandalensis and Plectranthus pubescens (Morris, 1996) are used to treat rheumatism, whereas in India the species used is Plectranthus mollis (Sharma and Sharma, 1981; Sebastian and Bhandari, 1984). Plectranthus mollis is reported to exhibit relaxant activity on smooth and skeletal muscles (Yoganarasimhan, 2000). 3.1.8. Other medical conditions Plectranthus barbatus is used to treat heart, blood and circulation conditions, such as myalgia, angina, haemorrage and hypertension in Africa and Asia (Dubey et al., 1981; Valdes et al., 1987; Yoganarasimhan, 2000; Chifundera, 2001). Plectranthus amboinicus is used in the Caribbean, to treat congestive heart failure (Morton, 1992) and Plectranthus mollis is used in India as a cardiac depressant (Yoganarasimhan, 2000). Species of Plectranthus used to treat various blood conditions include Plectranthus mollis as a cure for haemorrahage (Yoganarasimhan, 2000) and Plectranthus grandidentatus which has antiproliferation activity on human lymphocytes (Cerqueira et al., 2004). Species of Plectranthus used in the treatment of nervous disorders include Plectranthus amboinicus. It is prescribed in cases of epilepsy and convulsions (Morton, 1992; Ruiz et al., 1996; Castillo and González, 1999) and meningitis (Neuwinger, 2000). Plectranthus barbatus is used to treat psychiatric problems in Tanzania (Chhabra et al., 1993; Neuwinger, 2000) as well as treatment of insomnia and convulsions in Asia (Schanberg and Ikan, 2003). Plectranthus alpinus is used in Tanzania as an anticonvulsant (Hedberg et al., 1983). Plectranthus pubescens is used in Malawi for depression (Morris, 1996) and in Asia Plec- tranthus mollis is used for the treatment of mental retardation (Singh and Ali, 1992). In Uganda, Plectranthus laxiflorus is used to treat psychological problems including madness (Kakudidi et al., 1996). Plectranthus mandalensis is used in Malawi to treat depression (Morris, 1996) and in India Plectranthus vettiveroides is used as a stimulant (Yoganarasimhan, 2000). Species of Plectranthus are also used to treat sensory disorders associated with ear and eye problems. For example, Plectranthus amboinicus seed oil is a treatment for acute edematous otitis acuta in Polynesia (Zepernick, 1972), whereas in India its leaves are rubbed onto the eyes to alleviate conjunctivitis (Morton, 1992). Plectranthus barbatus is a recognized source of useful antiglaucoma agents (Caprioli et al., 1984; Svoboda, 1996; Schanberg and Ikan, 2003). The leaves are used to treat conjunctivitis in the Democratic Republic of Congo (Baerts and Lehmann, 1989; Neuwinger, 2000) and earache in Kenya (Geissler et al., 2002). In Malgaches, the finely ground leaves of Plectranthus bojeri are used for the treatment of conjunctivitis (Neuwinger, 2000). In Kenya, the leaves of Plectranthus igniarius and Plectranthus laxiflorus are used to treat inflamed eyes and ophthalmia, respectively (Lindsay and Hepper, 1978). A number of species including Plectranthus amboinicus (Bhakuni et al., 1969), Plectranthus barbatus (Zelnik et al., 1977; Schanberg and Ikan, 2003), Plectranthus caninus (Painuly and Tandon, 1983), Plectranthus esculentus (Mooi et al., 1999; Ali et al., 2000), Plectranthus mollis (Bhakuni et al., 1971) and Plectranthus ternifolius (Aswal et al., 1984) have cytotoxic and anti-tumour promoting activity and can be used in the treatment of cancer. Plectranthus barbatus is reported to stimulate the immune system (Lasure et al., 1995). Plectranthus amboinicus (Yoganarasimhan, 2000), Plectranthus barbatus (Greenway, 1946), Plectranthus bojeri (Neuwinger, 2000) and Plectranthus mollis (Jain et al., 1994) have been used against snakebites in India, Gabon and Kenya. In Uganda and Kenya, Plectranthus barbatus and Plectranthus aegyptiacus, respectively, are used to treat spiritual ailments (Parkia and Cooke, 2003; Tabuti et al., 2003). Plectranthus barbatus and Plectranthus amboinicus are used to prevent or alleviate inflammation (Morton, 1992; Chifundera, 2001). Plectranthus barbatus is also used in ethnoveterinary medicine in Africa (Holdsworth, 1977; Byavu et al., 2000; Ole-Marion, 2003). A number of Plectranthus species were recorded as being used in medicine but no specific condition was stated. For example, Plectranthus barbatus was cited as being used as a traditional medicine in Machakos (Barnett, 2000), a children’s tonic (Caufield, 1978), an emetic used by Samburu of Kenya for strength (Fratkin, 1975) and as an Ayurvedic medicinal plant (Svoboda, 1996). Plectranthus amboinicus is used medicinally in South East Asia and Martinique (Kuebel and Tucker, 1988; Prudent et al., 1995). Others include Plectranthus aegyptiacus used medicinally in Egypt (Othman and Shoman, 2004), Plectranthus congestus used in Australia for internal complaints (Maiden, 1899; Lassak and McCarthy, 1983), Plectranthus edulis is medicinal in Kenya (Lukhoba and Mathenge, 1993), Plectranthus gradulosus and Plectranthus occidentalis in Cameroon (Cheek et al., 2000; Harvey et al., 2004) and Plectranthus grallatus in South Africa (Cunningham, 1995). C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Plectranthus vettiveroides is medicinal (Shankar and Majumdar, 1995) and Plectranthus mollis is a tonic in India (Sebastian and Bhandari, 1984). Some species of Plectranthus are reported to have caused some adverse responses to people and domestic animals. Plectranthus barbatus causes perianal dermatitis (Owili, 1977) and contact with unnamed species of Plectranthus has been reported to cause skin irritations and dermatitis in susceptible persons (Mitchell and Rook, 1979; Motolese et al., 1991). Plectranthus caninus and Plectranthus longipes are poisonous to livestock (Glover and Samuel, 1962; Githinji, 1990). Plectranthus barbatus (Kemtoff et al., 2002) and Plectranthus fruticosus (Chamorro et al., 1991; Pages et al., 1998) have embryo- and fetotoxic activities on rodents. 3.2. Horticultural uses Twenty species were reported as having horticultural uses. The majority are ornamentals either planted for their coloured and attractive foliage or for their beautiful flowers. Many species of Plectranthus grown as ornaments are resistant to diseases, they are usually succulent and can survive in dry conditions. Some species of Plectranthus planted as ornamentals in Africa, Asia, Northern and Southern America and Australia include Plectranthus oertendahlii, Plectranthus parvifolius (syn Plectranthus australis) (Wiersema and Leon, 1999; Tourle, 2003; Schmidt, 2005), Plectranthus argentatus, Plectranthus amboinicus, Plectranthus fruticosus, Plectranthus madagascariensis, Plectranthus ciliatus (Garden Plants List, 2004), Plectranthus barbatus, (Grimshaw, 1993; Bennett and Prance, 2000), Plectranthus ecklonii (Auchland Regional Council, 2002), Plectranthus zuluensis (Coe, 2001) and Plectranthus saccatus (Kahurananga and Kiilu, 1976; Joffe, 1993, 2001). Plectranthus lanuginosus makes a splendid rock garden plant (Hepper and Jaeger, 1978), Plectranthus elegans (Jaarsveld, 1991) and Plectranthus verticillatus (Buitendag, 1971; Garden Plants List, 2004) are planted in gardens in South Africa while Plectranthus parvifolius is grown as a house plant (Schmidt, 2005). Plectranthus prostratus has decorative, fuzzy leaves that develop red markings in high light and is ideal for hanging baskets (McNew, 2002) and likewise for Plectranthus purpatus and Plectranthus oertendahlii (Welsh and Cotner, 2000). In East Africa, Plectranthus barbatus (Rounce, 1933; Greenway, 1946; Battiscombe, 1956; Templer, 1960; Mathew, 1970; Gachathi, 1989; Grimshaw, 1993) and Plectranthus sylvestris are planted as a hedge, fence or boundary marker. Cowpeas, green grams and maize are cultivated in areas where Plectranthus barbatus has been cleared as the plant is said to be a good indicator of fertile soil (Riley and Brokensha, 1988). The plant is used for making manure (Mwangangi, 1982) and is planted on the hillsides to prevent soil erosion (Drummond and Hemsley, 1953). 3.3. Food Nine species of Plectranthus are reported to be edible. For example, the tubers of Plectranthus esculentus are rich in 13 carbohydrates, Vitamin A and minerals and are usually boiled or roasted and eaten as a substitute for sweet potato in most parts of Africa (Dudgeon, 1909; Holland, 1915; Perrot, 1944; Purseglove, 1944; Semsei, 1956a,b; Roberts, 1990; Morris, 1996; Blench, 1997; Allemann and Hammes, 2003; Allemann et al., 2003). In Malaysia, the tuber is eaten as a vegetable (Mooi et al., 1999). Those of Plectranthus punctatus (Mooney, 1954; Tadesse, 1967; Scott, 1979) and Plectranthus edulis (Lukhoba and Mathenge, 1993) are eaten in Ethiopia and Kenya, respectively. Tubers of Plectranthus rotundifolius are a popular food in South Africa and research is currently underway to see if they can be cultivated for use as a domestic crop (Venter et al., 2000), whereas in Tropical Asia, the tubers of Plectranthus parviflorus are popular (Ramachandran and Nair, 1981; Purseglove, 1987). In India, the fruits of Plectranthus parvifolius are also eaten (Ramachandran and Nair, 1981). The leaves of Plectranthus mollis (Maikhuri and Gangwar, 1993) and Plectranthus barbatus (Fleurentin et al., 1983; Wiersema and Leon, 1999) are cooked as a vegetable. Leaves of Plectranthus amboinicus are chopped, made into flour balls and fried in oil or butter (Dymoc, 1885). 3.4. Food additives The three species of Plectanthus reported as food additives are Plectranthus amboinicus, Plectranthus esculentus and Plectranthus crassus (Table 2). The leaves of Plectranthus amboinicus are used in food stuffings (Purseglove, 1987), for flavouring and marinating beef and chicken (Epling, 1981; Kuebel and Tucker, 1988; Bodner and Gereau, 1988; Craig and Mayenda, 1990; Brown, 1997), to mask odor of strong smells associated with goat, fish and shellfish (Morton, 1992) and to spice dishes containing tomato sauces (Mayenda, 1991). The leaves are sometimes eaten raw with bread and butter and in India, they may be added to beer and wine (Morton, 1992). The stems of Plectranthus esculentus are used in Embu, Kenya, to sweeten gruel (porridge) (Allemann and Hammes, 1998) and in Hungary, the leaves of Plectranthus crassus are used for culinary purposes (Spicy hu, 2004). 3.5. Fodder Plectranthus spp. are mostly used as dry season fodder. Plectranthus barbatus is the most frequently cited of the five species used (Table 2). In Kenya and Yemen, it is fed to sheep, goats and cattle (Maher, 1935; Hendy, 1977; Caufield, 1978). Plectranthus igniarius is an important succulent fodder for camels, goats and cattle (Sato, 1976; Lusigi et al., 1984), Plectranthus sylvestris is recognized as fodder for domestic animals (Lusigi et al., 1984) and in Oman, Plectranthus pseudomarrubioides is sometimes given to cattle (Zaroug, 1981) while wild growing Plectranthus kamerunensis is reported to be eaten by domestic animals in the Rift Valley of Kenya (Glover et al., 1961a). Plectranthus hadiensis is eaten by rock rabbits in Tanzania (Tanner, 1952a) and Plectranthus edulis is eaten by elephants in the Aberdare National Park of Kenya (Githinji, 1990). 14 C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 3.6. Materials Plectranthus insignis is the only species whose wood is used to build huts and temporary houses and for firewood (Cheek et al., 2000; Table 2). In Kenya, the soft velvety leaves of Plectranthus barbatus are used as sanitary tissue (Githinji, 1988) to clean milk guards (Brokensha and Riley, 1978) and both the leaves and stems are used to hasten the ripening of bananas (Mwangangi, 1971, 1982). Plectranthus amboinicus has scented leaves and these are often rubbed into the hair and body after bathing (Morton, 1992). In the Amazon, the leaves are mixed with sugar and used as an intoxicant (Jain and Lata, 1996), while in Tonga and Martinique the leaves are used in the cleaning of textiles to perfume them (Yuncker, 1953; Prudent et al., 1995). A decoction of Plectranthus kamerunensis is used in bathing newborn babies in Kenya (Githinji and Kokwaro, 1993). Plectranthus ciliatus and Plectranthus montanus are used in personal hygiene and to wash clothes and animal skins (Tanner, 1952b; Watt and Breyer-Brandwijk, 1962). In Namibia, the roots of Plectranthus unguentarius are used as an ingredient of aromatic pomade for use by ladies (Winter and Leistner, 1957). In Kenya, old men use the leaves of Plectranthus caninus as snuff, a habit that can become addictive (Gentry, 1969). The mothers in Marakwet are reported to use the succulent leaves of this species to clean their breasts before suckling a baby after arriving from a journey (Githinji, 1990). A number of species are used for spiritual or religious purposes (see social use in Table 2). For instance, Plectranthus amboinicus is offered to the spirits when a house is being built (Morton, 1992). Plectranthus mollis, Plectranthus sylvestris and the roots of Plectranthus laxiflorus are used to drive away evil spirits in India, Kenya and Tanzania (Koritschoner, 1935a; Githinji and Kokwaro, 1993; Jain et al., 1994). The tubers of Plectranthus viphyensis are given to children in Tanzania to stop them grinding their teeth in their sleep (Lovett and Kayombo, 1989a,b). The seeds of Plectranthus mollis are fried in mustard oil and then massaged all over the body as an insect repellant (Jain et al., 1994). Plectranthus fruticosus, Plectranthus amboinicus and Plectranthus pseudomarrubioides are also used as insect repellants (Roberts, 1990; Prudent et al., 1995; Omolo et al., 2004). 4. Geographical distribution A survey of the ethnobotanical information by geographical areas shows that 45 species are used in the African continent (Watt and Breyer-Brandwijk, 1962; Roberts, 1990; Kokwaro, 1993; Githinji and Kokwaro, 1993; Morris, 1996; Neuwinger, 2000), 14 species in Asia (Morton, 1992; Yoganarasimhan, 2000), 10 species in America (Prudent et al., 1995; Ruiz et al., 1996), five species in Australasia (Morton, 1992) and one species in the pacific (Mayenda, 1991) (Tables 1 and 2). In Africa, the most frequently used species are Plectranthus barbatus and Plectranthus laxiflorus, whereas in Asia Plectranthus amboinicus and Plectranthus mollis are the most frequently cited species. Overall, Plectranthus amboinicus and Plectranthus barbatus have the widest geographical range occurring beyond Africa and Asia continents into the Americas, whereas Plectranthus laxiflorus is typically African in distribution occurring in Ethiopia through Eastern Africa to Southern Africa and Plectranthus mollis is mainly Asiatic. Most of the species in Africa and Asia are wild harvested, although some, such as Plectranthus esculentus, are now being commercially cultivated for use as a food source (Reinten and Cootzee, 2002). Overall, there is very little information about the harvesting methods used to supply these species and whether these methods are sustainable. 5. Chemistry of Plectranthus species To date, the majority of phytochemical studies on species of Plectranthus have focused on the isolation of a range of diterpenoids. Although, some of these studies report the biological activity of the diterpenoids isolated (Abdel-Mogib et al., 2002), very few have used activity-guided fractionation to isolate the compounds associated with a specific ethnobotanical use. Of the different types of diterpenoids found in the genus, the abietane diterpenoids are the most diverse group. Some of the abietane diterpenoids isolated include 6␤-formyloxy-7␣hydroxyroyleanone, coleon U and V in Plectranthus argentatus (Adler et al., 1984), coleon F, C, coleonol C, cariocal and plectrin in Plectranthus barbatus (Rüedi and Eugster, 1973; Grob et al., 1978; Tandon et al., 1978; Kelecom and Dos Santos, 1985; Kreutner et al., 1985; Jin et al., 1990), coleon M, N, O, P, Q and R in Plectranthus caninus (Arihara et al., 1975; Grob et al., 1978), edulone A and 16-O-acetylcoleon D in Plectranthus edulis (Buchbauer et al., 1978; Kunzle et al., 1987), 7␣,11-dihydroxy-12-methoxy-8,11,13-abietatriene in Plectranthus elegans (Dellar et al., 1996), grandidone A and grandidone B in Plectranthus grandidentatus (Uchida et al., 1981), 6␤,7␤-dihydroxyroyleanone in Plectranthus hadiensis (syn Plectranthus zeylanicus) (Mehrotra et al., 1989), horminone and 7␣,12-dihydroxy-17(15 → 16)-abeo-abieta-8,12,16-triene11,14-dione in Plectranthus hereroensis (Batista et al., 1994, 1995, 1996; Ferraeira et al., 1997) and lanugon M, N, S, 15epilanugin F, coleon J and H in Plectranthus lanuginosus (Moir et al., 1973a,b; Schmid et al., 1982; Matloubi-Moghadam et al., 1984). A few of these compounds have been tested for antimicrobial activity and this could explain some of medicinal uses, especially for the treatment of infections, fever and inflammation (Table 1). For example, the abietane diterpenes 7-epigrandidone D and grandidone D isolated from Plectranthus grandidentatus have antimicrobial activity (Teixeira et al., 1997). 11-Hydroxy12-oxo-7,9(11),13-abietatriene, isolated from Plectranthus elegans inhibited spore germination of the fungus Cladosporium cucumerinum and growth of Gram-positive bacteria, particularly Bacillus subtilis (Dellar et al., 1996; Gibbons, 2004). Similarly, Plectranthus hereroensis produces an acetylated abietane quinone related to horminone that has potent antimicrobial activity, especially against Staphylococcus (Batista et al., 1994, 1995; Htwe et al., 2001; Gibbons, 2004). Horminone isolated from the roots of Plectranthus hereroensis can cause liver damage, an adverse response that justifies further investigation as extracts of Plectranthus hereroensis are used to treat digestive C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 disorders (Ferraeira et al., 1997). Other diterpenoids including derivatives of barbatusol, cariocal and coleonol derivatives have hypotensive activity (Kelecom, 1983; Tandon et al., 1984, 1992; Kelecom and Dos Santos, 1985). Labdane diterpenoids found in Plectranthus barbatus, include forskolin, forskolin E and F (Valdes et al., 1987; Gabetta et al., 1989; Jin and He, 1998), ent-labda-8(17),12Z,14-trien-2␣ol and ent-3␤-acetoxylabda-8(17),12Z,14-trien-2␣-ol in Plectranthus fruticosus (Gaspar-Marques et al., 2003) and plectrornatin C and D in Plectranthus ornatus (Rijo et al., 2002). The fact that forskolin directly activates adenyl cyclase and thus the modulation of cAMP could underlie the diversity of different traditional uses of forskolin-containing species of Plectranthus, such as Plectranthus barbatus. For example, activation of adenyl cyclase can stimulate gastric secretions and the presence of forskolin in Plectranthus barbatus could explain why extracts from the plant are used for the treatment of digestive disorders (Bhat et al., 1979, 1980; Hersey et al., 1983; Mukherjee et al., 2000). Forskolin is known to directly activate the catalytic subunit of adenylate cyclases in smooth muscle cells, an action that is made possible by enhanced calcium intake (Metzger and Lindner, 1981; Lindner and Metzger, 1983). Forskolin is a known cardiotonic agent that mediates contractility by increasing levels of intracellular cAMP or by inhibiting its metabolism (Ghisalberti, 1977). It is also effective in the treatment of hypertension (Lindner et al., 1978; Valdes et al., 1987). Thus, forskolin could explain why extracts of Plectranthus barbatus are reported to lower blood pressure of anaesthetized rat due to relaxation of vascular smooth muscle (Dubey et al., 1981). The fact that forskolin can activate the cAMP-dependent protein kinase A signalling pathway (Huang et al., 2005) indicates that it could have beneficial anti-cancer properties. In fact, early studies showed that forskolin strongly inhibits melanoma cell-induced human platelet aggregation (Agarwal and Parks, 1983). Forskolin could also contribute to the therapeutic antidepressive potential of Plectranthus barbatus (Ozawa et al., 1997). Recently, products containing forskolin have been sold in the United States of America as sliming treatments as extracts of Plectranthus barbatus are reported to stimulate the breakdown of fat. However, there are concerns about the use of these products by patients taking medicinal drugs as forskolin can influence the drug-metabolizing enzymes in the liver and thus could be implicated in adverse herb–drug interactions (Ding and Staudinger, 2005). As yet there are little data on the biological properties of kaurane diterpenoids found in Plectranthus fruticosus (GasparMarques et al., 2003) and Plectranthus purpuratus (Katti et al., 1985), the rare phyllocladane diterpenoids from Plectranthus ambiguus (Liu and Rüedi, 1996; Liu et al., 2003), the neoclerodanes from Plectranthus ornatus (Rijo et al., 2002) and the methylenequinones from Plectranthus barbatus (Rüedi, 1986; Yoganarasimhan, 2000). Biologically active mono- and sesquiterpenoids are frequently found in many species of Plectranthus but there are little published data that directly link the presence of specific compounds in a species with the traditional uses of that species. For example, Plectranthus barbatus contains a 15 wide range of mono- and sesquiterpenoids including humulene and ␤-caryophyllene that are known for their antimicrobial properties (Ascensão et al., 1998). Plectranthus sylvestris also contains ␤-caryophyllene that could explain its use in treating infections (Chalchat et al., 1996). Plectranthus mollis contains fenchone, limonene, piperitenone, ␤-bisabolene, ␤-cubebene and ␣-humulene (Chalchat et al., 1996). These compounds have antimicrobial activity and could explain the use of Plectranthus mollis as a febrifuge (Varma and Sharma, 1963; Yoganarasimhan, 2000). The use of extracts of leaves of Plectranthus barbatus in birth control could be associated with the presence of sabinyl acetate as this compound has been found to be embryotoxic in rodents (Chamorro et al., 1991; Pages et al., 1991, 1998; Kemtoff et al., 2002). Sabinyl acetate has also been reported from Plectranthus fruticosus (Pages et al., 1998). Other monoterpenes and sesquiterpenes have been reported from different species and many have antimicrobial activity including ␤-caryophyllene, p-cymene and thymol from Plectranthus aegyptiacus (Smith et al., 1996), limonene, linalool, myrcene and thymol from Plectranthus amboinicus (Baslas and Kumar, 1981; Prudent et al., 1995), borneol, camphene, ␤-phellandrene and ␣-thujene from Plectranthus barbatus (Mathela et al., 1986; Maia et al., 1988), fenchone, nerol and terpinolene from Plectranthus glandulosus (Ngassoum, 2001), ␤-caryophyllene, fenchone and ␣-fenchyl acetate from Plectranthus madagascariensis (Ascensão et al., 1998), menthone and thymol from Plectranthus montanus (Chadya and Gundidza, 1999). ␣-Amorphene and ␤-cubebene occur in Plectranthus amboinicus (Prudent et al., 1995; Pino et al., 1996), ␦-selinene, ␣-ionone and humulene are found in Plectranthus barbatus (Mathela et al., 1986; Maia et al., 1988; Camara et al., 2003), caryophyllene oxide and 15-hydroxyspathulenol in Plectranthus fruticosus (Gaspar-Marques et al., 2003) and germacrene D occurs in Plectranthus sylvestris (Chalchat et al., 1996). The role of phenolic compounds in the medicinal properties of species of Plectranthus has not been well studied. An antioxidant activity-guided fractionation of Plectranthus ambiguus resulted in the isolation of 5,6-dihydroxy-7,4′ -dimethoxyflavone (ladanein) (Liu and Rüedi, 1996). Phenolics have also been reported from Plectranthus amboinicus (Brieskorn and Riedel, 1977), Plectranthus caninus (Painuly and Tandon, 1983) and Plectranthus sylvestris (Juch and Rüedi, 1997). As part of a chemotaxonomic study, the flavonoids of 42 species of Plectranthus were studied and found to contain two non-flavonoid phenolics, the caffeic acid derivatives, nepetoidin A and B (Grayer et al., 2003). Grayer et al. (2003) showed that nepetoidin B had potent free radical scavenging activity and was more active than rosmarinic acid and gallic acid. Nepetoidin A was not tested due to lack of material. Both compounds had antifungal activity. Nepetoidin A and B were found in the water extracts of leaves from 13 of the species of Plectranthus reported in this review to have medicinal activity: Plectranthus ambiguus, Plectranthus asirensis, Plectranthus barbatus, Plectranthus coeruleus, Plectranthus ecklonii, Plectranthus elegans, Plectranthus hadiensis, Plectranthus igniarius, Plectranthus kivuensis, Plectranthus lanuginosus, Plectranthus madagascarensis, Plectranthus parviflorus and Plectranthus pseudomarrubioid. Whether these 16 C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 two compounds contribute to the medicinal properties of these species has not been studied. In addition, lipids have been reported from Plectranthus mollis (Yoganarasimhan, 2000). Lipids, resinous gum and compounds of nutritional value like calcium, iron, proteins and carbohydrates have been recorded from Plectranthus esculentus (Allemann and Hammes, 2003; Allemann et al., 2003). Alcohols occur in Plectranthus aegyptiacus (Smith et al., 1996), Plectranthus amboinicus (Prudent et al., 1995) and Plectranthus glandulosus (Ngassoum, 2001). 6. Ethnobotanical uses and phylogeny In order to see if there are any relationships among the species of Plectranthus with ethnobotanical uses, the ethnobotanical data were mapped onto a phylogeny of the genus. The most recent traditional classification of the genus proposed by Codd (1975) is problematic because it does not include all species encountered in the literature and it groups species by superficial morphological similarity based on one or two characters leading to groupings which are incongruent with the phylogeny proposed by Paton et al. (2004). The phylogeny by Paton et al. (2004), based on DNA sequence data and augmented by morphological data, provides an informal classification that divides the species into two main groups or clades (Fig. 1). Clade 1, the ‘Coleus’ Clade of Paton et al. (2004), broadly corresponding to the formally recognized genus Coleus, is divided into two subclades, Clades 1a and b. Clade 2 is recognized as the ‘Plectranthus’ Clade (Fig. 1). Not all species encountered in this ethnobotanical survey are covered in the phylogeny. Species not covered are grouped together with those included in the phylogeny that are morphologically similar. In this way it is possible to assign most species to groups within one of the Clades 1a, b or 2 (Fig. 1). Species that form one of these groups were given numbers within the clade, for example, Clade 1a group 8 contains species morphologically similar to Plectranthus amboinicus. Occasionally, it was not possible to assign a species cited in the literature to numbered groups in Clade 1 or 2 because they did not share morphological characters with species in those clades. Such species were placed in groups annotated by letters. For example, Plectranthus edulis belongs in Clade 1 as its corolla, calyx and nutlet morphology are similar to species in that Clade. However, Plectranthus edulis is not very similar to Fig. 1. Figure illustrating the placement of utilized Plectranthus species on the phlyogeny following Paton et al. (2004). The thick unbroken lines represent groups with species cited as useful; the broken lines indicate groups with no/or few cited uses; groups shown in bold have recorded medicinal uses. C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 any of the numbered groups represented in the phylogeny. Thus, Plectranthus edulis is placed in Clade 1, group B along with another species, Plectranthus punctatus that is morphologically similar. When the ethnobotanic data were mapped onto the phylogenetic cladogram for Plectranthus it shows that 70% of all species of Plectranthus with medicinal uses are found in Clade 1 (Fig. 1). In fact, the majority is found in groups in Subclade 1b, except those in group 6. It is of interest that species in group 6 have no recorded medical uses. Subclade 1a has two medicinally important groups, groups 2 and 8. The unplaced groups of species in Clade 1 are all used medicinally except group D (Fig. 1). In Clade 2, the only medicinally used species fall into groups 3, 6, B and E. All species used as food or in flavouring food are found in Clade 1 with the exception of Plectranthus glandulosus that belongs to Clade 2. Within Clade 1, species used as food occur within groups A, B and E and in Subclade 1b group 5. The majority of species utilized as fodders also occur within Clade 1 (particularly Subclade 1a groups 2 and 7 and Subclade 1b groups 2 and 8), except Plectranthus kamerunensis that belongs to Clade 2 group 6. All species used in apiculture are also found in Clade 1. The majority of species used in the materials and social use categories are also from Clade 1 (Table 2; Fig. 1) as are the species used for magic-religious activities. The same trend is seen in the horticultural category, where Clade 1 has a greater number of species than in Clade 2. In addition, species of Clade 1 are used in a wide range of agricultural and horticultural procedures, such as preparing manure, preventing erosion and as hedges (Bush, 1943; Drummond and Hemsley, 1953; Mwangangi, 1982). The species recorded as having horticultural uses that are placed in Clade 2 all occur in group 3 and are ornamentals. 7. Conclusion Of the 300 species of Plectranthus, 62 species are reported to be used as medicines, ornamentals, foods, flavours, fodder and/or material. It could be that the number of species used is greater as there were citations to unconfirmed species that were only cited at the level of genus, but because it was unclear as to the species used very few of these references have been cited in this review. There are some interesting trends that emerge from this review. Species with ethnobotanical uses are not randomly distributed thoughout the genus but are related. The majority of species with ethnobotanical uses belong to Clade 1, a group of Plectranthus formally known as Coleus. Most species with medicinal uses occur in Clade 1 Subclade 1b and Subclade 1a groups 2 and 8. Whether other species in the other groups within Clade 1 or those in Clade 2 have potential to be used as medicinal plants justifies further study as the compounds, especially the diterpenoids, occur in species from both clades. To date the majority of the phytochemical studies have concentrated on the isolation of diterpenoids and although these compounds have been shown to have potent antimicrobial activity few have been tested in bioassays that are directly related to the traditional uses of the species the compounds were isolated from. Further 17 research is needed to identify the active compounds, especially in those species that are most frequently cited. One of the most studied Plectranthus-derived compounds is the labdane forskolin isolated from Plectranthus barbatus. It has a range of pharmalogical propoperties and could explain many of the diverse medicinal uses of Plectranthus barbatus. Despite the interest in the activity of this compound our knowledge about its distribution in the genus is poor. Natural product chemists have mostly reported on novel diterpenoids in species of Plectranthus and maybe not always reported on the occurrence of known compounds. Thus, there is a scientific case for more research to be undertaken on the activity and distribution of diterpenoids in this genus. The majority of the 300 species of Plectranthus occur in Africa and around half are distributed within sub-Saharan Africa of these most, about 110, species belong to Clade 1 and 40 species belong to Clade 2 (Paton et al., 2004). Madagascar and Southern Africa both have around 50 species, although in these areas around 75% of species belong in Clade 2. There are around 70 species in Asia and Australia together and these species can be placed in Clade 1 (Paton et al., 2004). The genus is not native in the New World, the species which have been introduced there largely belong to Clade 1. In this review, we have collected information about the uses of 21% of the 300 species. The lack of documentation for the other species could reflect the fact that it is not easy to differentiate among some of the species and species with a limited distributed could have been confused with the more common widespread distributed species. In this study, we have made use of the notes on herbarium sheets and on collector notes for herbarium vouchers lodged in herbaria in parts of Africa and the United Kingdom. If this review was expanded to cover samples of Plectranthus from herbaria in other countries as well as unpublished literature, such as locally produced ethnobotanical reports from African and Asian universities or research institute, our knowledge about the diversity of species used and how they are used would most likely increase. Whatever the source of information, it is important that the authors refer to voucher samples lodged in a herbarium. These vouchers can be used to support the identification of the species. The value and importance of vouchers and the notes associated with the vouchers can be illustrated by reference to Plectranthus stolzii. Plectranthus stolzii is restricted in its natural distribution to parts of Southern Tanzania and Northern Malawi, and there is very little information about its uses in the published literature. A survey of different herbaria found vouchers deposited at Kew that had been made during general collecting trips (Gaetan, 1952) and projects aimed to record the importance of medicinal plants to the people of Africa to support their conservation (Cunningham and Barigyira, 1992). The vouchers of P. stolzii record the occurance of the species and its ethnobotanical uses (Table 1). These data have not yet been documented in published papers. The species is reported to treat a range of medicinal conditions including respiratory conditions (Table 1). This species is very similar in appearance to the more widespread Plectranthus laxiflorus that occurs in the same areas as Plectranthus stolzii 18 C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 and is used to treat some of the sample medicinal conditions (Table 1). However, because voucher specimens of Plectranthus stolzii were lodged in the herbarium at Kew, along with notes about its traditional uses, the identification of the species can be confirmed. Species that have the greatest number of different medicinal uses, such as Plectranthus barbatus and Plectranthus amboinicus, appear to be distributed widely in both the Old and New World. The wide distribution of these frequently used species reflects the fact that they have been semi-cultivated as “herbs” and taken by people from one place to another as they have migrated across the world. Generally, the uses in the New World are similar to those in the Old World, although there are some differences. In Africa and Asia, Plectranthus species were mostly used to treat digestive, skin and infections, whereas Plectranthus amboinicus is used in the treatment of epilepsy and convulsions in the Caribbean (Morton, 1992; Ruiz et al., 1996) and as an intoxicant in the Amazon (Jain and Lata, 1996). These species are also used in the treatment of pain in Africa but not so frequently in Asia. This could reflect the fact that in Asia other more active species were available, whereas this was not the case in Africa. Plectranthus includes a few species that are often found in dry open cleared areas near villages and could be classed as a “weed” using the definition proposed by Baker (1965). Baker (1965) defines a weed as a species that can successfully grow in “any specified geographical area, its population grow entirely or predominantly in situations markedly disturbed by man, without, of course, being deliberately cultivated”. Species classed as weeds are often medicinal plants (Stepp, 2004). The natural habitat of species, such as Plectranthus laxiflorus, can be associated with the behaviour of village people as it is found at the margin of forests and near paths. Nevertheless, there are many other species of Plectranthus found in similar habitats but they are not recorded as having medicinal or other economic uses. This could be because some are rare and thus not frequently used or their economic uses have not been documented. Nevertheless, within Plectranthus the link between being classed as a weed and having medicinal properties is at the moment weak, as it is only supported by a few species. The phenomenon of synonymy within the genera has also made it difficult to collate together the uses of specific species. Synonymy is most prevalent for species in Clade 1 and particularly in Subclade 1b, which contains both of the popularly used species, Plectranthus barbatus and Plectranthus amboinicus. The diversity of names used to describe the same species could in part reflect the widespread use of these species along with the difficulties in differentiating among closely related species. In this review, we have shown how a cladogram produced from the new molecular phylogeny of the genus (Paton et al., 2004) has provided a framework to superimpose our knowledge about the uses of Plectranthus. Furthermore, the cladogram can also serve as a guide to direct the search for information on the uses of other species, especially those in Clade 2. Within Clade 1b it is highly likely that those species closely related to those with traditional uses could also share these uses. Acknowledgements The authors are grateful to the BAT Biodiversity Partnership with RBG Kew for the grant that enabled this work to be carried out. We are also grateful to Dr. T. Kokubun, Royal Botanic Gardens, Kew, for designing the database used in this project. 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P. barbatus Andr. Herbarium specimen held at K collected from Kenya (Greenway 7879). Grimshaw, 1993. P. barbatus Andr. Herbarium specimen held at K collected from Tanzania (Grimshaw 9339). Hedberg, 1979. P. punctatus L’Her. Herbarium specimen held at K collected from Ethiopia (Hedberg 6942B). Hendy, 1977. P. barbatus Andr. Herbarium specimen held at K collected from Yemen (Hendy 8). Hepper, Jaeger, 1978. P. lanuginosus (Benth.) Agnew. Herbarium specimen held at K collected from Kenya (Hepper & Jaeger 7032). Jaarsveld, 1991. P. elegans Britten. Herbarium specimen from held at K collected from South Africa (Jaarsveld 11328). Jarrett, 1950. P. barbatus Andr. Herbarium specimen held at K collected from Uganda (Jarrett 16). Kahurananga, Kiilu, 1976. P. saccatus Benth. Herbarium specimen held at EA collected from Kenya (Kahurananga & Kiilu 2953). Koritschoner, 1935a. P. laxiflorus Benth. Herbarium specimen held at K collected from Tanzania (Koritschoner 880). Koritschoner, 1935b. P. laxiflorus Benth. Herbarium specimen held at K collected from Tanzania (Koritschoner 1365). Leeuwenberg, 1972. Plectranthus aff. occidentalis. Herbarium specimen held at K collected from Cameroon (Leeuwenberg 10289). Lovett, Kayombo, 1989a. P. viphyensis Brummit & J.H. Seyani. Herbarium specimen held at K collected from Tanzania (Lovett & Kayombo 3317). Lovett, Kayombo, 1989b. P. viphyensis Brummit & J.H. Seyani. Herbarium specimen held at K collected from Tanzania (Lovett & Kayombo 3508). Lukhoba, Mathenge, 1993. P. edulis (Vatke) Agnew. Herbarium specimen held at NAI collected from Kenya (Lukhoba & Mathenge 220). Maher, 1935. P. barbatus Andr. Herbarium specimen held at K collected from Kenya (Maher 3229). Mathew, 1970. P. barbatus Andr. Herbarium specimen held at K collected from Kenya (Mathew 6163). Mayenda, 1991. P. amboinicus (Lour.) Spreng. Herbarium specimen held at K collected from the Pacific (Mayenda 43). Meyer, 1964. P. pseudomarrubioides Willemse. Herbarium specimen at EA collected from Ethiopia (Meyer 8590). Meyerhoff, 1978a. P. tetensis (Bak.) Agnew. Herbarium specimen held at K collected from Kenya (Meyerhoff 14M). Meyerhoff, 1978b. P. barbatus Andr. Herbarium specimen held at K collected from Kenya (Meyerhoff 28M). Meyerhoff, 1978c. P. amboinicus (Lour.) Spreng. Herbarium specimen held at K collected from Kenya (Meyerhoff 94M). Mooney, 1954. P. punctatus L’Her. Herbarium specimen held at K collected from Ethiopia (Mooney 6225). Msyla, 1944. P. esculentus N.E.Br. Herbarium specimen held at EA collected from Uganda (Msyla 1600). Mwangangi, 1971. P. barbatus Andr. Herbarium specimen held at K collected from Kenya (Mwangangi 1648). Mwangangi, 1982. P. barbatus Andr. Herbarium specimen held at K collected from Kenya (Mwangangi 2241). Napier, 1933. P. kivuensis (Lebrun & Touss.) R.H. Willemse. Herbarium specimen held at K collected from Kenya (Napier 5409). NMK Expd., 1974. P. hadiensis (Forssk.) Schweinf. ex Spreng. Herbarium specimen held at EA collected from Kenya (NMK Expd. 171). Purseglove, 1944. P. esculentus N.E.Br. Herbarium specimen held at K collected from Uganda (Purseglove P1600). Rounce, 1933. P. barbatus Andr. Herbarium specimen held at K collected from Tanzania (Rounce 253). Rwaburindore, 1969. P. barbatus Andr. Herbarium specimen held at K collected from Uganda (Rwaburindore Rwab. 32). Sato, 1976. P. igniarius (Schweinf.) Agnew. Herbarium specimen held at EA collected from Kenya (Sato 22). 24 C.W. Lukhoba et al. / Journal of Ethnopharmacology 103 (2006) 1–24 Scott, 1979. P. punctatus L’Her. Herbarium specimen held at K collected from Ethiopia (Scott 63). Semsei, 1956a. P. esculentus N.E.Br. Herbarium specimen held at EA collected from Tanzania (Semsei 2464). Semsei, 1956b. P. esculentus N.E.Br. Herbarium specimen held at K collected from Tanzania (Semsei 2498). Simon, Mollel, 1999. P. barbatus Andr. Herbarium specimen held at K collected from Tanzania (Simon & Mollel 94). Tadesse, 1967. P. punctatus L’Her. Herbarium specimen held at K collected from Ethiopia (Tadesse 638). Tanner, 1946. P. barbatus Andr. Herbarium specimen held at K collected from Tanzania (Tanner 4877A). Tanner, 1952a. P. hadiensis (Forssk.) Schweinf. ex Spreng. Herbarium specimen held at K collected from Tanzania (Tanner 629). Tanner, 1952b. P. montanus Benth. (P. cylindraceus). Herbarium specimen held at K collected from Tanzania (Tanner 728). Tanner, 1959. P. hadiensis (Forssk.) Schweinf. ex Spreng. Herbarium specimen held at K collected from Tanzania (Tanner 4221). Tanner, 1961a. P. stachyoides Oliv. Herbarium specimen held at K collected from Tanzania (Tanner 5742). Tanner, 1961b. P. lactiflorus (Vatke) Agnew. Herbarium specimen held at K & EA collected from Tanzania (Tanner 5911). Tanner, 1961c. Plectranthus sp. Herbarium specimen held at K collected from Tanzania (Tanner 5907). Templer, 1960. P. barbatus Andr. Herbarium specimen held at K collected from Tanzania (Templer T11). Ward, 1935. P. esculentus N.E.Br. Herbarium specimen held at K & EA collected from Tanzania (Ward H39/35). Williams, 1975. P. barbatus Andr. Herbarium specimen held at EA collected from Tanzania (Williams 60). Winter, Leistner, 1957. Herbarium specimen held at K collected from Namibia (Winter & Leistner 5595). Yuncker, 1953. P. amboinicus (Lour.) Spreng. Herbarium specimen held at BM collected from Tonga (Yuncker 15232). Zaroug, 1981. P. pseudomarrubioides Willemse. Herbarium specimen held at K collected from Oman (Zaroug RMD 23).