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Caryopsis Micromorphological Survey of the Genus Themeda (Poaceae) and Allied Spathaceous Genera in the Andropogoneae

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Caryopsis Micromorphological Survey of the Genus Themeda (Poaceae) and Allied Spathaceous Genera in the Andropogoneae Turkish Journal of Botany Turk J Bot (2014) 38: 665-676 http://journals.tubitak.gov.tr/botany/ © TÜBİTAK Research Article doi:10.3906/bot-1308-27 Caryopsis micromorphological survey of the genus Themeda (Poaceae) and allied spathaceous genera in the Andropogoneae 1,2 1 1 1, Yu ZHANG , Xiaoying HU , Yunxiao LIU , Qing LIU * 1 Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, P.R. China 2 Graduate University of Chinese Academy of Sciences, Beijing, P.R. China Received: 16.08.2013 Accepted: 19.02.2014 Published Online: 20.05.2014 Printed: 19.06.2014 Abstract: A caryopsis micromorphological survey of 15 species of the genus Themeda Forssk. (Poaceae) and 4 species of allied spathaceous genera in tribe Andropogoneae allowed the recognition of 2 types of caryopses based on the ventral face and embryo shape for the first time. One kind of caryopsis had a concave ventral face with a grooved embryo, and the other kind of caryopsis had a flat ventral face with a keel-shaped embryo. Although the ventral face and embryo shapes had limited taxonomic value at the infrageneric level, the caryopsis shapes, embryo proportion, and sculpturing patterns were found to be valuable taxonomic characters at the interspecific level in Themeda. Consistent micromorphological characters of style base persistence and dorsiventral compression were observed for the first time in the caryopses of representative species of spathaceous genera in the Andropogoneae. Key words: Micromorphology, spathaceous genera, taxonomy 1. Introduction (Sendulsky et al., 1986; Clayton, 1990; Long, 1999). The genus Themeda Forssk. (Andropogoneae, Poaceae) Caryopses are significant raceme components enclosed comprises 27 drought-tolerant species exhibiting by spatheoles; thus, caryopsis micromorphology surveys considerable ecological and morphological diversity in might provide clues to unravel the mystery of the tropical and subtropical arid areas (Chen and Phillips, ecological function of spatheoles in Andropogoneae. 2006; Zhang and Liu, 2012). Northern India and Southwest Caryopsis micromorphology, as a reliable approach for China (Figure 1) were inferred as the center of diversity assessing phylogenetic relationship, had been definitely for the genus with 20 species (Zhang and Liu, 2012). recognized in Poaceae (Bogdan, 1966; Barthlott, 1981; Two sections of Themeda were recognized based on the Wang et al., 1986; Liu et al., 2005b). For example, an characters of raceme: section Primothemedae S.L.Chen & examination revealed that 7 subfamilies corresponded T.D.Zhuang with homogamous spikelet pairs arranged at to 7 caryopsis subtypes for Chinese grasses (Wang et al., different levels, and section Themeda with homogamous 1986). In another example, the caryopsis ventral face and spikelet pairs arranged at the same level in the raceme base embryo shape were valuable characters for recognizing (Chen and Zhuang, 1989; Chen and Phillips, 2006). 3 suprageneric groups in the gramineous subfamily The spathaceous genera were characterized by Chloridoideae (Liu et al., 2005b). However, almost compound panicles, with spatheoles intervening at the nothing was known about the caryopsis micromorphology base of racemes and paraclades (repeating branches; of Themeda and allied spathaceous genera in the Liu et al., 2005a) in the tribe Andropogoneae (Figure Andropogoneae. 2) (Clayton and Renvoize, 1986). Themeda and allied The present study aims to: 1) give detailed descriptions spathaceous genera, the members of a specialized of caryopses in 15 Themeda species and 4 species of allied lineage in the tribe (Skendzic et al., 2007; Soreng et al., spathaceous genera in the tribe Andropogoneae; 2) discuss 2009; Teerawatananon et al., 2011), were predominantly the taxonomic value of caryopsis micromorphological distributed in extreme habitats of arid rocky mountain characters at the suprageneric and interspecific level; and slopes (authors’ field observations). The ecological 3) evaluate the evolutionary tendency of the caryopsis function of spatheoles remained mysterious for years micromorphological character in Themeda. * Correspondence: [email protected] 665 ZHANG et al. / Turk J Bot 80° E 90° E 100° E 110° E T. anathera T. tremula T. arundinacea T. trichiata T. caudata T. villosa T. cymbaria T. yunnanensis T. hookeri T. helferi T. huttonensis T. laxa T. intermedia T. quadrivalvis T. mooneyi T. sabarimalayan T. pseudotremula T. strigosa 30° N T. saxicola T. triandra 20° N N 4000 km Figure 1. Distribution center of Themeda (yellow regions). Generic distribution regions shown by closed red lines in the lower right corner (color on the web only). A B Figure 2. Spathaceous inflorescence (A) and diagram structure (B) of Themeda caudata. MF = main florescence, Pa = paraclade, Pr = prophyll, Sp = spatheole, hollow circle = raceme, hollow oblong = sterile spikelet, solid oblong = fertile spikelet (color on the web only). 2. Materials and methods After shape observation and drawing with a Carl Zeiss Mature caryopses of 15 species in Themeda and 4 species SM, the caryopses were cleaned in a series of ethanol of allied spathaceous genera [5–16 caryopses for each solutions (70%, 90%, and 100%) 3 times and then sputter- species, excluding T. unica S.L.Chen & T.D.Zhuang and coated with gold in a JFC-1100 sputter coater (JEOL, Elymandra lithophila (Trin.) Pilg.] were examined either Tokyo, Japan). Micromorphological characters were from field collections or from herbarium specimens (Table examined and photographed with a JEOL JSM-6360LV 1). The length measurements of spatheole, lemma, and scanning electron microscope (SEM) at 10 kV. Caryopses caryopsis were conducted under a Carl Zeiss Stemi SV 11 were orientated with the stylopodium pointing up and the stereomicroscope (SM) (Carl Zeiss AG, Jena, Germany) embryo pointing down. The dorsal face with embryo was and data are available upon request. Statistical analyses orientated towards the lemma and the ventral face with were conducted using Microsoft Excel 2007. sculptures was orientated towards the palea (Wang et al., 666 ZHANG et al. / Turk J Bot Table 1. Material of Themeda [following the classifications of Chen and Phillips (2006) and The Plant List (2010)] and allied spathaceous genera examined in this study. Species Voucher Location No. Figures Themeda Section Primothemedae S.L.Chen & T.D.Zhuang T. anathera (Nees ex Steud.) Hack. Walter K 22194 (US); PI 215612 India: Madhes 5 3A, 4B, 7A, 9A, 10A Rock JF 825 (US) Burma: Chinwin District T. arundinacea (Roxb.) A.Camus 20 4D, 6B, 6C, 9B, 9C, 10C Mao PY 6793 (IBSC) China : Yunnan T. avenacea (F.Muell.) Maiden & Betche White CT 8736 (US) Australia: Queensland 5 4E, 6D, 8I, 10D T. caudata (Nees) A.Camus Qing Liu 154 (IBSC) China: Guangdong 5 3G, 7B, 9D, 10E Clayton 5789 (US) Ceylon: Nuwara Eliya District T. cymbaria Hack. 11 4F, 6E, 8J, 10F V. Narayana Surasny s.n. (IBSC) India Anonymous 16764 (US) ; PI 247413 Philippines: Luzon T. gigantea (Cav.) Hack. 20 3B, 3H, 6F, 9E, 10G Ye HG 891 (IBSC) China: Guangdong T. hookeri (Griseb.) A.Camus Qingzang Exped. 3168 (KIB) China: Sichuan 6 3D, 7D, 8B, 10I T. intermedia (Hack.) Bor Waterhouse JH 952 (US) New Guinea: Bismarck Archipelago 5 3I, 6G, 9F, 10J T. unica S.L.Chen & T.D.Zhuang Courtois P 12259 (NAS) China: Anhui 1 7E, 8C, 11C T. villosa (Poir.) A.Camus Liborio EE 854 (US) Philippines: Silangan Mt. 5 4G, 7F, 9J, 11D T. yunnanensis S.L.Chen & T.D.Zhuang Wu SK 362 (IBSC) China: Yunnan 5 4H, 6J, 9I, 11E, 11F Section Themeda T. arguens (L.) Hack. Frank WG 13027 (US) Ceylon: Peradeniya 5 4C, 6A, 8H, 10B T. helferi Munro ex Hack. Ban NL s.n. (US) India: Madzae 5 3C, 7C, 8A, 10H T. quadrivalvis (L.) Kuntze Li YL et Ma WJ YDDXSB-158 (KIB) China: Yunnan 8 3E, 6H, 9G, 11A Yang GH 69627 (KIB); ILCA 18381 China: Sichuan T. triandra Forssk. 20 3J, 6I, 9H, 11B Jiang B 1 (IBSC) China: Shandong Allied spathaceous genera Elymandra lithophila (Trin.) Clayton Anonymous 11.763 (US) Angola: Lunda 1 4I, 7G, 8D, 11G Shauty HJ 227 (US) Africa: Nyasaland Heteropogon contortus (L.) P.Beauv. ex Roem. & Schult. 20 4A, 7H, 8E, 11H Zhang ZQ 10113 (IBSC) China : Guangxi Anonymous 4022 (KIB) China: Yunnan Hyparrhenia filipendula (Hochst.) Stapf 20 3F, 7I, 8F, 11I Davidse G 8915 (US) Sri Lanka, Uva Hyperthelia dissoluta (Nees ex Steud.) Clayton Hitchcock AS 25002 (US) Kenya: Eldoret 5 4J, 7J, 8G, 11J Abbreviations: IBSC = Herbarium of South China Botanical Garden, Chinese Academy of Sciences (CAS); ILCA = International Livestock Centre for Africa, Addis Ababa, Ethiopia; KIB = Herbarium of 667 Kunming Institute of Botany, CAS; NAS = Herbarium of Institute of Botany, Jiangsu Province and CAS; PI = Germplasm Resources Information Network of United States Department of Agriculture at Beltsville; US = United States National Herbarium. ZHANG et al. / Turk J Bot 1986; Terrell and Peterson, 1993). SEM photos were taken of section Themeda, namely T. helferi; and 4 species of in the middle area of the ventral face for each sample. allied spathaceous genera including Elymandra lithophila, Terminology following Sendulsky et al. (1986), Stearn Hyparrhenia filipendula, Heteropogon contortus, and (1992), Terrell and Peterson (1993), Koul et al. (2000), and Hyperthelia dissoluta. Bona (2013) was used. Flat ventral face: Caryopses had relatively flat ventral faces, rounded dorsal faces, and keel-shaped embryos. 3. Results They were abbreviated as the FK type (Figure 3B). According to the corresponding relationship between Taxa with the FK type of caryopses: 6 species of section ventral face and embryo morphology, 2 types of caryopses Primothemedae, including Themeda arundinacea, T. were observed in the examined taxa. A detailed description avenacea, T. cymbaria, T. gigantea, T. intermedia, and T. is given below. yunnanensis, and 3 species of section Themeda, including Concave ventral face: Caryopses had concave ventral T. arguens, T. quadrivalvis, and T. triandra. faces, convex dorsal faces, and grooved embryos. They In addition to the ventral face, 5 diagnostic characters, were abbreviated as the CG type (Figure 3A).
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