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Morphometric Analysis of the Rattan Calamus Javensis Complex (Arecaceae: Calamoideae) Author(S): Mega Atria, Harald Van Mil, William J

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Morphometric Analysis of the Rattan Calamus Javensis Complex (Arecaceae: Calamoideae) Author(S): Mega Atria, Harald Van Mil, William J Morphometric Analysis of the Rattan Calamus javensis Complex (Arecaceae: Calamoideae) Author(s): Mega Atria, Harald van Mil, William J. Baker, John Dransfield, and Peter van Welzen Source: Systematic Botany, 42(3):494-506. Published By: The American Society of Plant Taxonomists URL: http://www.bioone.org/doi/full/10.1600/036364417X696168 BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder. BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Systematic Botany (2017), 42(3): pp. 494–506 © Copyright 2017 by the American Society of Plant Taxonomists DOI 10.1600/036364417X696168 Date of publication August 25, 2017 Morphometric Analysis of the Rattan Calamus javensis Complex (Arecaceae: Calamoideae) Mega Atria,1,2,6 Harald van Mil,3 William J. Baker,4 John Dransfield,4 and Peter van Welzen1,5 1Naturalis Biodiversity Center, Research Group Biodiversity Discovery, P. O. Box 9517, 2300 RA Leiden, The Netherlands 2Department of Biology, E building, Faculty of Mathematics and Natural Sciences, University of Indonesia, Depok 16424, Indonesia 3Department of Mathematics, University of Leiden, P. O. Box 9512, 2300 RA Leiden, The Netherlands 4Royal Botanic Gardens, Kew, Richmond, Surrey, TW9 3AB, U.K. 5Institute of Biology Leiden, University of Leiden, P.O. Box 9505, 2300 RA Leiden, The Netherlands 6Author for correspondence ([email protected]) Communicating Editor: Martin Wojciechowski Abstract—Calamus javensis (Calamoideae) is a slender rattan common in tropical rainforests. The species is very polymorphic and forms a species complex together with some undetermined forms and the species C. acuminatus, C. amplijugus, C. congestiflorus, C. corrugatus, C. elopurensis, C. hypertrichosus, C. impar,andC. tenompokensis. Within the complex, the morphological variation and similarities among the entities are too difficult to solve with traditional morphological observation. Therefore, two multivariate analyses, MDS (multidimensional scaling) and hierarchical cluster analysis (HCA) with UPGMA (unweighted pair group method with arithmetic mean) were used for two datasets, quantitative data only and in combination with qualitative data. The inclusion of qualitative characters did not make significant differences in the cluster results. Of all nine taxa included in the complex, only two clusters resulted from the analysis next to two forms based on few specimens. Only C. tenompokensis was recognized as a separate cluster, while all other taxa were combined into one typical C. javensis cluster. More material may result in the recognition of the two forms as distinct species. Keywords—Calamus tenompokensis, hierarchical cluster analysis, multivariate analysis, species complex, West Malesia. Calamus L. is the largest genus within the palms and belongs alternative rattan source for industrial needs. However, the fact to the rattans or climbing palms (Dransfield 1974). The genus is that C. javensis is a polymorphic species complex with variable characterized by the presence of fruit scales. Most Calamus quality of the cane makes it less valuable unless the complex is species are climbing, and may reach 150 m in length; however, taxonomically resolved. erect or stemless species/forms are also known. They climb Calamus javensis was first described by Blume (1847). He with a flagella or a cirrus. Other typical characters are scattered transferred his own Calamus equestris into the synonymy of C. leaf sheath spines, unisexual inflorescences, and solitary or javensis. He also noted morphological variation within C. javensis clumped stems (Furtado 1956). Within the genus Calamus, C. and distinguished three varieties based mainly on the variation javensis Blume is known as an extremely polymorphic species in the leaves and spines on the leaf sheaths (Table 1; var. a is (Dransfield 1992). Cronk (1988) stated that polymorphic considered to be the typical variety, and should according to species (or ochlospecies) are species with chaotic infraspecific modern nomenclature be called var. javensis). Beccari (1884, variation intractable to formal taxonomic treatment. Calamus 1908) recognized Blume’s varieties and made modifications, javensis is an understory rattan belonging to the subtribe creating his own infraspecific taxonomy. Since then (Table 1), Calaminae, tribe Calameae of the palm subfamily Calamoi- many changes in the infraspecific taxa occurred; forms were deae (Dransfield et al. 2008). This slender, generally sparsely united, separated, etc. Also, at the species level many excep- spiny, climbing palm is widely distributed in Southeast Asia, tional forms were described as new species and synonymized ranging from southern Thailand and Peninsular Malaysia, to again, the latter especially by Dransfield (1992, 1999), who Sumatra, Java, Borneo, and Palawan. Characters are: stem mainly recognized one species without infraspecific taxa. As a with leaf sheath removed 2–6 mm in diameter; with sheaths to result, C. javensis is taxonomically the most difficult rattan 10 mm in diameter; internodes up to 30 cm long but usually species complex in the Southeast Asian region (Dransfield 1999). shorter; ocrea distinct, conspicuously deep crimson when Table 1 shows that no subdivision of the complex provided a young; leaves pinnate, ecirrate, up to 40 cm long; terminal satisfactory solution. Forms were described at various levels or leaflets flabellate, lowermost pair often swept back across the moved to different levels, which also shows that the characters stem; flagellum up to 75 cm long; inflorescence long, with red used for recognition were doubtful, too. Madulid (1981) treated crimson rachilla; ripe fruit ovoid (Dransfield 1992; Dransfield C. javensis as a species complex because of the extreme variation and Manokaran 1993). within the species and the relatively subtle differences among People make bindings, rope, and baskets from the cane. The morphologically similar groups of specimens. Cronk (1988) fruits are edible for small mammals and a few birds. The raw introduced the term ochlospecies for this phenomenon, but we cabbage can be eaten and is sometimes used to cure coughs. In prefer to use the term species complex. In species complexes Sarawak the cane is considered excellent because of its length variation is great, often showing complex patterns of disjunc- and strength, but it is only sold locally for baskets, cordage, tions in the ranges of variation (often recognised by separate and handicrafts (Mogea 1994). In Palawan, the cane is deemed names), some of which can be linked to ecological factors only second in quality to Calamus caesius Blume (a well-known or geography, others being apparently random (Cronk 1988, species in the rattan trade: Mogea 1994; Shim and Tan 1994). Dransfield 1999). To solve the problem, Dransfield (1999) According to the FAO, C. javensis is included in the major suggested two options: maintaining several varieties, or commercial species of rattan as identified for Asia by Dransfield blending them all into one species. He also suggested to add and Manokaran (1993). Thus, C. javensis may become a new comprehensive studies in the field along with phenetic and 494 2017] ATRIA ET AL.: MORPHOMETRIC ANALYSIS OF CALAMUS JAVENSIS 495 Table 1. The chronological history of names and taxa involved in the Calamus javensis Blume species complex (adapted from Madulid, 1981). Blume (1847) Beccari and Hook.f. (1892) Ridley (1907) Beccari (1908, 1913) Ridley (1925) Furtado (1956) Dransfield (1984, 1992) C. javensis C. javensis C. penicillatus C. javensis C. javensis C. javensis C. javensis var. a ‘forma typica’ (Dransfield and Manokaran 1993; var. b (firmus) Govaerts and Dransfield 2005) var. inermis var. inermis var. inermis var. laevis var. peninsularis subvar. var. purpurascens var. peninsularis subvar. var. purpurascens var. peninsularis purpurascens purpurascens var. purpurascens subvar. tenuissimus var. tenuissimus var. tenuissimus subvar. pinangianus var. pinangianus var. peninsularis var. pinangianus var. pinangianus var. pinangianus subvar. polyphyllus var. polyphyllus var. polyphyllus var. intermedius var. g (tetrastichus) var. tetrastichus subvar. mollispinus var. sublaevis var. acicularis var. exilis C. kemamanensis C. corrugatus C. corrugatus C. filiformis C. acuminatus C. acuminatus C. hypertrichosus C. hypertrichosus C. impar C. tenompokensis C. tenompokensis (Govaerts and Dransfield 2005) C. amplijugus C. congestiflorus C. elopurensis 496 SYSTEMATIC BOTANY [Volume 42 molecular-phylogenetic techniques to examine the circum- will be used as a preliminary framework for the molecular scription of C. javensis or subdivisions as an alternative. examination of the species complex in order to construct
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