|Clerodendrum trichotomum fruit|
Clerodendrum is a genus of flowering plants in the family Lamiaceae. Its common names include glorybower, bagflower and bleeding-heart. It is currently classified in the subfamily Ajugoideae, being one of several genera transferred from Verbenaceae to Lamiaceae in the 1990s, based on phylogenetic analysis of morphological and molecular data.
Estimates of the number of species in Clerodendrum vary widely, from about 150 to about 450. This is partly because about 30 species have been transferred to Rotheca, about 30 more to Volkameria, and 1 to Ovieda. The type species for the genus is Clerodendrum infortunatum. It is native to Sri Lanka and the Andaman Islands.
The genus is native to tropical and warm temperate regions of the world, with most of the species occurring in tropical Africa and southern Asia, but with a few in the tropical Americas and northern Australasia, and a few extending north into the temperate zone in eastern Asia.
They are shrubs, lianas, and small trees, usually growing to 1–12 m (3 ft 3 in–39 ft 4 in) tall, with opposite or whorled leaves. C. floribundum can grow to 30 m (98 ft) tall. Clerodendrum fistulosum and Clerodendrum myrmecophila have hollow stems that are inhabited by ants. Clerodendrum trichotomum is a common ornamental in warmer parts of the world. Eight other species are also grown in the tropics for their abundant and attractive flowers. One of these, Clerodendrum macrostegium, suckers abundantly from the roots, often producing a thicket within a few years. A few other species are also found, somewhat rarely, in cultivation.
Clerodendrum species are used as food plants by the larvae of some Lepidoptera species including Endoclita malabaricus and Endoclita sericeus. Both butterflies and hummingbirds are often attracted to blooming clerodendrum.
Formerly placed here
- Pseudocaryopteris foetida (D.Don) P.D.Cantino (as C. foetidum D.Don)
- Rotheca incisa (Klotzsch) Steane & Mabb. (as C. incisum Klotzsch or C. macrosiphon Hook.f.)
- Rotheca myricoides (Hochst.) Steane & Mabb. (as C. myricoides (Hochst.) Vatke or C. ugandense Prain)
- Rotheca serrata (L.) Steane & Mabb. (as C. serratum (L.) Moon)
- Volkameria aculeata L. (as C. aculeatum (L.) Schltdl.)
- Volkameria glabra (E.Mey.) Mabb. & Y.W.Yuan (as C. glabrum E.Mey.)
- Volkameria inermis L. (as C. inerme (L.) Gaertn.)
- Volkameria ligustrina Jacq. (as C. ligustrinum (Jacq.) R.Br.)
Stamens 4 (rarely 5), usually in 2 pairs of unequal length and projecting well beyond the mouth of the corolla.
The flowers are protandrous. When the flower opens, the stamens stand erect, parallel to the central axis of the flower, while the style bends over, holding the stigma beyond the rim of the corolla. After the pollen is shed, the stamens curl up or bend over, and the style straightens out, bringing the stigma to the center of the flower. Except for Aegiphila, which is heterostylous, this breeding strategy is shared by all members of the clade consisting of Kalaharia, Clerodendrum, Volkameria, Aegiphila, Ovieda, Tetraclea, and Amasonia.
Clerodendrum was named by Linnaeus in Species Plantarum in 1753. The name is derived from two Greek words, kleros, meaning "chance or fate" and dendron, "a tree". It refers to the considerable variation in reports of the usefulness of Clerodendrum in medicine.
Regional revisions of Clerodendrum have been done for local floras, but the last monograph of the entire genus was by John Isaac Briquet in 1895. He recognized about 90 species, defining the genus broadly to include species that others had placed in Rotheca, Volkameria, and Ovieda. His circumscription was followed by most authors for the next 100 years, even though it was widely believed to be problematic.
In 1997, phylogenetic analysis of DNA data showed that Clerodendrum, as then understood, was polyphyletic. This situation was remedied in 1998 with the revival of Rotheca. This taxonomic change was based on previous work and on a molecular phylogenetic study that was not published until the following year.
In 2004, a study of DNA sequences showed that the monospecific Australian genus Huxleya was embedded in a clade of Clerodendrum species that had formerly been placed in Volkameria. Huxleya was then sunk into synonymy with Clerodendrum. The 2004 study sampled Aegiphila, Tetraclea, and Amasonia, three New World genera of Ajugoideae that had not previously been sampled for DNA. The results of this study cast doubt, once again, upon the monophyly of Clerodendrum.
In 2010, a study of four chloroplast DNA intergenic spacers showed that part of Clerodendrum was closer to the New world genera than to other Clerodendrum, and that one species of Clerodendrum was nested within the clade of New World genera. The authors of this study resurrected the genus Volkameria and assigned to it about 30 species that had been in Clerodendrum. They also resurrected Ovieda as a monotypic genus consisting of Ovieda spinosa. Volkameria and Ovieda had been erected by Linnaeus in 1753. Modern cladistic analysis has largely vindicated his concepts of Clerodendrum and its relatives.
Traditional medicinal use
Clerodendrum is strongly supported as monophyletic in molecular phylogenetic analyses. It consists of two clades, each of which receives strong bootstrap support. One clade contains mostly African species. The other is mostly Asian. The African and Asian groups can not confidently be divided into sections without more extensive sampling of taxa in phylogenetic studies. The Madagascan species, in particular, are poorly studied.
- "Genus: Clerodendrum L.". Germplasm Resources Information Network. United States Department of Agriculture. 2010-05-27. Retrieved 2011-02-17.
- Raymond M. Harley, Sandy Atkins, Andrey L. Budantsev, Philip D. Cantino, Barry J. Conn, Renée J. Grayer, Madeline M. Harley, Rogier P.J. de Kok, Tatyana V. Krestovskaja, Ramón Morales, Alan J. Paton, and P. Olof Ryding. 2004. "Labiatae" pages 167-275. In: Klaus Kubitzki (editor) and Joachim W. Kadereit (volume editor). The Families and Genera of Vascular Plants volume VII. Springer-Verlag: Berlin; Heidelberg, Germany. ISBN 978-3-540-40593-1
- Steven J. Wagstaff, Laura Hickerson, Russ Spangler, Patrick A. Reeves, and Richard G. Olmstead. 1998. "Phylogeny in Labiatae s.l., inferred from cpDNA sequences". Plant Systematics and Evolution 209(3-4):265-274.
- Yao-Wu Yuan, David J. Mabberley, Dorothy A. Steane, and Richard G. Olmstead. 2010. "Further disintegration and redefinition of Clerodendrum (Lamiaceae): Implications for the understanding of the evolution of an intriguing breeding strategy". Taxon 59(1):125-133.
- Dorothy A. Steane and David J. Mabberley. 1998. "Rotheca (Lamiaceae) Revived". Novon 8(2):204-206.
- Rosette B. Fernandes and Bernard Verdcourt. 2000. "Rotheca (Labiatae) revived - more new combinations". Kew Bulletin 55(1):147-154.
- Clerodendrum In: Index Nominum Genericorum. In: Regnum Vegetabile (see External links below).
- Anthony J. Huxley, Mark Griffiths, and Margot Levy (editors). 1992. The New Royal Horticultural Society Dictionary of Gardening. The Macmillan Press Limited, London; The Stockton Press, New York.
- David J. Mabberley. 2008. Mabberley's Plant-Book third edition (2008). Cambridge University Press: UK. ISBN 978-0-521-82071-4
- Floyd, A.G., Australian Rainforests in New South Wales Volume 2 - 1990 ISBN 0-949324-32-9 page 179
- George W. Staples and Derral R. Herbst "A Tropical Garden Flora" Bishop Museum Press: Honolulu (2005)
- "GRIN Species Records of Clerodendrum". Germplasm Resources Information Network. United States Department of Agriculture. Retrieved 2011-02-17.
- Clerodendrum page 637. In: Carolus Linnaeus. 1753. Species Plantarum volume 2. Laurentii Salvii. (see External Links below).
- Umberto Quattrocchi. 2000. CRC World Dictionary of Plant Names volume I, page 559. CRC Press: Boca Raton; New York; Washington,DC;, US. London, UK. ISBN 978-0-8493-2673-8 (set).
- John Isaac Briquet. 1895. "Clerodendrum" pages 174-176. In: "Verbenaceae" pages 132-182. In: Die Natürlichen Pflanzenfamilien volume IV, part 3a. Verlag von Wilhelm Engelmann: Leipzig, Germany.
- Dorothy A. Steane, Robert W. Scotland, and David J. Mabberley (1997). "Phylogenetic Relationships of Clerodendrum s.l. (Lamiaceae) Inferred from Chloroplast DNA". Systematic Botany 22 (2): 229–243. doi:10.2307/2419455.
- Dorothy A. Steane, Robert W. Scotland, David J. Mabberley, and Richard G. Olmstead (1999). "Molecular systematics of Clerodendrum (Lamiaceae): ITS sequences and total evidence". American Journal of Botany 86 (1): 98–107. doi:10.2307/2656958. PMID 21680349.
- Dorothy A. Steane, Rogier P.J. de Kok, and Richard G. Olmstead (2004). "Phylogenetic relationships between Clerodendrum (Lamiaceae) and other Ajugoid genera inferred from nuclear and chloroplast DNA sequence data". Molecular Phylogenetics and Evolution 32 (1): 39–45. doi:10.1016/j.ympev.2003.11.011. PMID 15186795.
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