Systematics of the Guadua Angustifolia Complex (Poaceae: Bambusoideae) Author(S): Stephen M
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Systematics of the Guadua angustifolia Complex (Poaceae: Bambusoideae) Author(s): Stephen M. Young and Walter S. Judd Source: Annals of the Missouri Botanical Garden, Vol. 79, No. 4 (1992), pp. 737-769 Published by: Missouri Botanical Garden Press Stable URL: http://www.jstor.org/stable/2399719 Accessed: 08-08-2014 17:43 UTC Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Missouri Botanical Garden Press is collaborating with JSTOR to digitize, preserve and extend access to Annals of the Missouri Botanical Garden. http://www.jstor.org This content downloaded from 158.135.136.72 on Fri, 08 Aug 2014 17:43:50 UTC All use subject to JSTOR Terms and Conditions Volume79 Annals Number4 ofthe 1992 Missouri Botanical Garden SYSTEMATICS OF THE StephenM. Young2and GUADUA ANGUSTIFOLIA WalterS. Judd3 COMPLEX (POACEAE: BAMBUSOIDEAE)' ABSTRACT Guadua angustifolia Kunth is the largest and most economically important bamboo in the Western Hemisphere. It is the primary source of building material for urban and rural dwellings in many areas within its geographical range, especially in Colombia and Ecuador. Guadua aculeata Rupr. ex Fourn. from Central America and Bambusa (Guadua) chacoensis N. Rojas from Argentina and Paraguay are two closely related taxa that were often confused with or identified as G. angustifolia. These three taxa comprise the Guadua angustifolia complex. A detailed morphological analysis was performed on all three species using traditional taxonomic methods as well as principal components analysis. The Guadua angustifolia complex was also compared with other sympatric species of the genus. The results indicate that G. angustifolia should be regarded as one polymorphic species consisting of two subspecies, subspecies angustifolia and subspecies chacoensis, which are differentiatedby a mosaic of vegetative and reproductive characters. Subspecies angustifJlia consists of populations from Central America (formally G. aculeata) and north- western South America that exhibit variability in certain vegetative and reproductive characters. The woody bamboos of the grass subfamilyBam- species that vary from slender, thornless plants busoideae are widespread throughout the tropics, with scandent culms to large, thorny plants with and bamboos of the genus Guadua Kunth com- erect culms. mony occur in the hot lowland regions of Latin The largest and one of the most beautiful of America. The genus contains approximately 30 these bamboos is Guadua angustifolia.It was first ' This studyis dedicatedto the memoryof Thomas R. Soderstrom,whose encouragement, support, and enthusiasm forthe studyof bamboosmade thiswork possible. We thankour friendsand the staffof the Universityof Florida herbarium(FLAS) and the Departmentof Botanyfor help in completingthe research.Additional thanks go to the HerbariumServices Unit of the United States National Herbarium (US) forhelp in obtainingspecimens and information. We thankthe curatorsof the followingherbaria for allowing access to materialunder their care: B, COL, CTES, ENCB, F, G, LE, MEXU, MO, NY, P, S, UC, US, and VEN. Special thanksto LynnClark of Iowa State University, Camilo Quarin of Argentina,and Ximena Londofioand Oscar Hidalgo of Colombia for providingmuch useful information.Photos were taken by the firstauthor except whereotherwise indicated. 2 MercerArboretum and BotanicGardens, 22306 AldineWestfield Road, Humble,Texas 77338, U.S.A.; present address: New York Natural HeritageProgram, New York Departmentof EnvironmentalConservation, 700 Troy- SchenectadyRoad, Latham,New York 12110-2400, U.S.A. 3 Departmentof Botany,220 BartramHall, Universityof Florida,Gainesville, Florida 32611, U.S.A. ANN. MISSOURI BOT. GARD. 79: 737-769. 1992. This content downloaded from 158.135.136.72 on Fri, 08 Aug 2014 17:43:50 UTC All use subject to JSTOR Terms and Conditions 738 Annals of the Missouri Botanical Garden discovered by Humboldt and Bonpland during their single variable (polymorphic) species consisting of expedition to South America and was formallyde- two subspecies (i.e., subsp. angustifolia and cha- scribed by them in 1806. Its distributionranges coensis), with the former showing several slightly from southern Mexico to northern Argentina, and divergent geographic races. its economic importance is unsurpassed by any Guadua angustifoliasubsp. chacoensis is found other native bamboo in the Western Hemisphere. along rivers in northeastern Argentina and eastern Where temperature ranges are high enough and Paraguay and is geographically isolated from G. rainfall or water supply is adequate, these large angustifolia subsp. angustifolia, which is found bamboos provide the raw material for a nearly from northern Peru, through Ecuador, Colombia, endless array of products. It is in Colombia and and central America to southern Mexico, often Ecuador that the uses of plants in the genus Gua- occurring along water courses. The two subspecies dua, especially Guadua angustifolia, reach their can be separated using a combination of vegetative greatest development. Here G. angustifolia can and reproductive characters, but since the mor- be seen in large stands consisting of hundreds of phological gap separating them is smaller than that culms up to 30 m tall and 20 cm diam. Its form separating most species in the genus, the taxa are of growth,strength, and ease of transport,cutting, considered here to be only subspecifically distinct. and working make G. angustifolia the preferred source of constructionmaterial wherever it occurs. TAXONOMICHISTORY Many of the houses of these areas are built entirely THE GENUSGUADUA of the culms of G. angustifolia (Fig. 1). Even though its tremendous size, beauty, economic im- A great deal of controversyhas centered around portance, and broad geographic distribution are the separation of the genus Guadua Kunth from outstanding among the Latin American bamboos, Bambusa Schreber. When Humboldt and Bon- its taxonomy and natural history have not been pland described Bambusa guadua and B. latifolia studied in detail. The last taxonomic treatment of (1806), the firstspecies of large bamboos collected the group was writtenby William Munro in 1868, in South America, they included them in Bambusa. and many new species have been discovered since At that time the genus consisted of six Asian spe- then. cies, and the use of Bambusa Schreber rather than The Guadua angustifolia complex includes G. Bambos Retz., both published in 1789, was still angustifolia and two closely related species, G. controversial. A history of this problem and rec- aculeata and Bambusa chacoensis, oftenconfused ommendations for conserving Bambusa Schreber with or identifiedas G. angustifolia. In order to can be found in McClure (1946) and Holttum delimittaxa withinthe complex, a detailed analysis (1956). Humboldt & Bonpland (1806) chose Bam- of the morphological variabilityof the complex was busa, which they thoughthad been more commonly undertaken. During this study the characters that used and was more pleasing to the ear. The New separate the G. angustifolia complex from other World species of Bambusa are very similar to the species in the genus Guadua also were defined. Asian species in such features as stature, branch The paucity of complete herbarium specimens and complements, thorniness, and spikelet structure, lack of recent revisionary studies have led to a and it was understandable that Humboldt and poor understanding of species delimitation within Bonpland would place them in the same genus. the genus and frequent misidentificationof many A short time later their specimens were closely specimens, including those of G. angustifolia. studied by Carl S. Kunth after he was employed The patterns of variation in over 150 vegetative by Humboldt to help publish the informationgath- and reproductive characters from throughout the ered on Humboldt and Bonpland's journey to the geographic range of Guadua angustifolia, Bam- New World (Stearn, 1956). Kunth (1822a, b) de- busa chacoensis, and G. aculeata were studied cided to place the two New World species in a new using traditionaltaxonomic methods (including her- genus that he called Guadua, but he failed to state barium and field studies) and principal components what characters separated it fromBambusa. Later, analyses. The results of these methods led to the Munro (1868) described a number of character- conclusion that the complex is best treated as a istics that previous authors had used to distinguish FIGURE 1. House constructionusing Guadua angustifolia,Ecuador.-a. House construction,Cayapas River, Esmeraldas.-b. Cayapas Indianhouse, Cayapas River,Esmeraldas. -c. House in Muisne.-d. Experimentalhousing projectin Huaquillas. This content downloaded from 158.135.136.72 on Fri, 08 Aug 2014 17:43:50 UTC All use subject to JSTOR Terms and Conditions Volume 79, Number4 Young & Judd 739 1992 Systematics of Guadua angustifolia t' ' nd.r I f1 EUf ,a S This content downloaded from 158.135.136.72 on Fri, 08 Aug 2014 17:43:50 UTC All use subject to JSTOR Terms and Conditions 740 Annals of the Missouri Botanical Garden the genus. These