Ethnobotany, Phytochemistry and Pharmacology of Podocarpus Sensu Latissimo (S.L.) ⁎ H.S
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Available online at www.sciencedirect.com South African Journal of Botany 76 (2010) 1–24 www.elsevier.com/locate/sajb Review Ethnobotany, phytochemistry and pharmacology of Podocarpus sensu latissimo (s.l.) ⁎ H.S. Abdillahi, G.I. Stafford, J.F. Finnie, J. Van Staden Research Centre for Plant Growth and Development, School of Biological and Conservation Sciences, University of KwaZulu-Natal Pietermaritzburg, Private Bag X01, Scottsville 3209, South Africa Received 26 August 2009; accepted 2 September 2009 Abstract The genus Podocarpus sensu latissimo (s.l.) was initially subdivided into eight sections. However, based on new information from different morphological and anatomical studies, these sections were recognised as new genera. This change in nomenclature sometimes is problematic when consulting ethnobotanical data especially when selecting plants for pharmacological screening, thus there is a need to clear any ambiguity with the nomenclature. Species of Podocarpus s.l. are important timber trees in their native areas. They have been used by many communities in traditional medicine and as a source of income. Podocarpus s.l. is used in the treatment of fevers, asthma, coughs, cholera, distemper, chest complaints and venereal diseases. Other uses include timber, food, wax, tannin and as ornamental trees. Although extensive research has been carried out on species of Podocarpus s.l over the last decade, relatively little is known about the African species compared to those of New Zealand, Australia, China and Japan. Phytochemical studies have led to the isolation and elucidation of various terpenoids and nor- and bis- norditerpenoid dilactones. Biflavonoids of the amentoflavone and hinokiflavone types have also been isolated. Nor- and bis-norditerpenes are said to be taxonomic markers for this genus. Recent in vitro and in vivo studies have shown antitumor, antimicrobial, anti-inflammatory, antioxidant, larvicidal, plant and insect growth regulation activities. Various studies have yielded important natural bioactive products and two of them are worth mentioning. Taxol, a significant anticancer agent has been isolated from Podocarpus gracilior and totarol, a diterpenoid isolated from various species and now commercially produced as a potent antibacterial and antioxidant agent. Findings from this review supports the use of an ethnobotanical and chemotaxonomical approach in selecting plants for pharmacological screening since most of the species in the different morphological groups have similar uses. Also the isolated compounds have chemotaxonomic value amongst the groups. Some of the biological activities identified from extracts and compounds isolated from Podocarpus s.l. support the rationale behind the medicinal uses of these species. © 2009 SAAB. Published by Elsevier B.V. All rights reserved. Keywords: Chemotaxonomy; Ethnobotany; Pharmacology; Podocarpus; Traditional uses Contents 1. Introduction ................................................................2 1.1. Morphology ............................................................2 1.2. Taxonomy .............................................................2 1.3. Distribution ............................................................2 1.4. Chemistry .............................................................2 2. Ethnobotanical uses of Podocarpus....................................................3 2.1. Non-medicinal uses ........................................................3 2.2. Medicinal uses...........................................................3 ⁎ Corresponding author. E-mail address: [email protected] (J. Van Staden). 0254-6299/$ - see front matter © 2009 SAAB. Published by Elsevier B.V. All rights reserved. doi:10.1016/j.sajb.2009.09.002 2 H.S. Abdillahi et al. / South African Journal of Botany 76 (2010) 1–24 3. Phytochemistry ............................................................ 11 4. Biological activity ........................................................... 15 4.1. Anticancer/cytotoxic activities ................................................. 15 4.2. Antimicrobial activity ..................................................... 16 4.3. Plant growth inhibitory activity ................................................ 17 4.4. Insect growth inhibitory activity ................................................ 18 4.5. Gastroprotective activity .................................................... 18 4.6. Other activities ......................................................... 19 5. Conclusions .............................................................. 20 Acknowledgements ............................................................. 21 References ................................................................. 21 1. Introduction ceae using the single most parsimonious tree based on 54 characters showed the genera Nageia, Retrophyllum and Afro- 1.1. Morphology carpus which initially were included in Podocarpus s.l. to belong to the tropical clade and very closely related. Dacrycarpus s.l., Until the 1970s, the Podocarpaceae family was composed of though a tropical clade, was separated from Podocarpus s.l. seven genera; Podocarpus L' Her. ex Pers., Dacrydium Sol. ex because of being a tetragonal-leaved clade (Kelch, 1997). A recent Forst., Phyllocladus Rich. Ex Mirb., Acmopyle Pilg., Micro- analysis of the family based on rbcL sequences retrieved three cachrys Hook.f., Saxegothaea Lindl. and Pherosphaera W. main clades (Podocarpoid, Dacrydioid and Prumnopityoid) and Archer bis (= Microstrobos J. Garden & L.A.S. Johnson, Afrocarpus, Nageia and Retrophyllum were grouped just as nom. inval.: Brummitt et al., 2004). Based on the leaf anatomy, Kelch's analysis indicated (Conran et al., 2000). This study also Podocarpus was initially subdivided into eight sections; Afro- placed Sundacarpus amarus within Prumnopitys, suggesting it is carpus J. Buchholz & N. E. Gray, Dacrycarpus Endl., Eupodo- just a specialised member of that genus and is not itself a good carpus Endl., Microcarpus Pilg., Nageia (Gaertn.) Endl., genus (Quinn and Price, 2003). Polypodiopsis C. E. Bertrand, Stachycarpus Endl., and Sunda- carpus J. Buchholz & N. E. Gray (Buchholz and Gray, 1948). The 1.3. Distribution AfricantaxawereplacedinPodocarpus (Leistner, 1966). In the past 25 years, research, based on new information from studies of In the early 1940s and 1950s, the biogeographical information wood anatomy, embryology and chemistry, have proposed new on Podocarpaceae was published (Florin, 1940; Buchholz and genera and endorsed recognition of the 19th century segregates Gray, 1948; Li, 1953). However, due to great changes in the from Podocarpus sensu latissimo (s.l.) and Dacrydium (s.l.) (De taxonomy of Podocarpaceae (De Laubenfels, 1969; Quinn, 1982; Laubenfels, 1969, 1972, 1985; Quinn, 1982; Page, 1989). Molloy, 1995; Kelch, 1997, 1998), the earlier biogeographical However, herbaria and authors of floristic works have been statement is rendered invalid. For example the genus Dacrydium slow to accept these changes, probably because the broadly is not present in both western and eastern hemispheres (Sinclair defined genera are unnatural. et al., 2002). The conventional view was that the Podocarpaceae had a Gondwanan origin and migrated northwards to reach the 1.2. Taxonomy present northerly limits in the Caribbean, Ethiopia and eastern Asia. The only limitation to this view is that it does not consider Subsequent studies based on morphological and molecular the numerous Laurasian fossils that have been assigned to (DNA sequence) data showed that Podocarpus section Afrocar- Podocarpaceae (Mill, 2003). The greatest generic diversity of pus is related to Podocarpus section Nageia,andtheother Podocarpaceae is in Malesia and Australasia where 17 of the 19 sections were regarded as paraphyletic (Kelch, 1997). More recent living genera are found. Malesia, New Caledonia and New studies by Conran et al. (2000) and Sinclair et al. (2002), Zealand each have eight genera, seven genera in Australia, four concludes that there is considerable molecular evidence favouring genera in South America and Africa, and Asia two genera each. the generic level recognition of Afrocarpus and the other genera as With respect to living endemism, Australia has three endemic proposed by Page (1989). Thus Podocarpaceae is now represented genera (Lagarostrobos, Microcachrys and Pherosphaera), two by two genera in Africa: Podocarpus and Afrocarpus,andthe for New Zealand (Halocarpus and Manoao), New Caledonia, species Podocarpus falcatus (Thunb.) R. Br. ex Mirb. and Po- South America and Africa each have one (Parasitaxus, Saxe- docarpus gracilior Pilg. are now Afrocarpus falcatus (Thunb.) gothaea and Afrocarpus,respectively)(Mill, 2003). C. N. Page and Afrocarpus gracilior (Pilg.) C. N. Page (Barkera et al., 2004). Cladistic analysis of morphological, anatomical and 1.4. Chemistry embryological characteristics indicated strongly the segregation of the new genera and the relationships between closely related Presence or absence of different compounds can be used to genera. For instance, the morphological analysis of Podocarpa- relate the new and the old taxa. For example, flavonoids can be H.S. Abdillahi et al. / South African Journal of Botany 76 (2010) 1–24 3 a useful chemotaxonomic tool in this group of plants, since Pitman, 1972). For example, fine timber known as Yellowwood biflavonoids of the amentoflavone and hinokiflavone groups is obtained from the South African species P. falcatus (Thunb.)