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Catabrosa Versus Colpodium (Poaceae: Poeae) in Southern Africa, with a Key to These Genera and Their Species in Africa

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Catabrosa Versus Colpodium (Poaceae: Poeae) in Southern Africa, with a Key to These Genera and Their Species in Africa KEW BULLETIN VOL. 66: 1 – 10 (2011) Catabrosa versus Colpodium (Poaceae: Poeae) in southern Africa, with a key to these genera and their species in Africa Robert J. Soreng1 & Lyn Fish2 Summary. Differences between the genera Colpodium Trin. and Catabrosa P. Beauv. are discussed. Colpodium drak- ensbergense Hedberg & I. Hedberg is transferred to Catabrosa drakensbergense (Hedberg & I. Hedberg) Soreng & Fish and a lectotype is chosen for the species. This leaves only two Colpodium species in Africa (Ethiopia, Kenya and Tanzania), and extends the range of Catabrosa from northwestern Africa to southern Africa. Key Words. Africa, Catabrosa, Colpodium, Miliinae, Poaceae, Poeae, Puccinelliinae. Introduction chionogeiton (Pilg.) Tzvelev (Kenya and Tanzania), C. Colpodium drakensbergense Hedberg & I. Hedberg was drakensbergense (Lesotho), and C. hedbergii (Ethiopia, described by Olov Hedberg and Inga Hedberg (1994), Kenya, Tanzania). from specimens collected in high altitude bogs in On receiving two specimens of Colpodium drakensber- Lesotho. Collections from Lesotho were initially iden- gense from Lesotho as a gift to US from PRE it was tified as Catabrosa aquatica (L.) P. Beauv (Anderson noticed that the glumes were unequal in length and 1969), then, beginning with annotations as early as some samples had numerous 2-flowered spikelets. Thus 1972 (on Werger 1614), as Colpodium hedbergii (Melderis) it looked as if these specimens belonged to Catabrosa P. Tzvelev (van Zinderen Bakker & Werger 1974;Gibbs Beauv. rather than to Colpodium. However, in Africa Russell et al. 1990), a species otherwise known from Mt Catabrosa was known only from northwest Africa (Maire Kilimanjaro to Ethiopia. The Hedbergs provided a 1955; Clayton et al. 2006), and a study was therefore discussion of the taxonomic history of the genus undertaken to examine the variation in this species, Colpodium Trin. s.l. (sensu Clayton & Renvoize 1986), including the type material, in order to resolve the along with a detailed analysis and key to the species of generic position of the southern African material. Africa. Tzvelev & Bolchovskich (1965)dividedColpo- The genus Colpodium was merged with Catabrosa by dium into four genera: Catabrosella (Tzvelev) Tzvelev, Boissier (1884)asCatabrosa sect. Colpodium (Trin.) Colpodium, Hyalopoa (Tzvelev) Tzvelev, and Paracolpo- Boiss. but nearly all other authors since Bentham & dium (Tzvelev) Tzvelev. The three segregate genera are Hooker (1883) have kept the genera, s.s. or s.l., apart. sometimes accepted (Alexeev 1980, 1988;Alexeev& Several characteristics differentiate Catabrosa from Tzvelev 1981; Cope 1982;Tzvelev1976;Mill1985), and Colpodium (see Discussion and Table 1). Catabrosa has sometimes not (Bor 1970; Clayton & Renvoize 1986; two widely accepted species: the polymorphic C. Hedberg & Hedberg 1994;Dicoré1995; Wu & Phillips aquatica (circumboreal, with a few stations in southern 2006). Colpodium s.s., including Keniochloa Melderis Patagonia and Tierra del Fuego in Argentina and (Tzvelev 1976; Hedberg & Hedberg 1994), sometimes Chile) and C. werdermannii (Pilg.) Nicora & Rúgolo recognised as Colpodium sect. Keniochloa (Melderis) E. (Andes of central Argentina and Chile to southern B. Alexeev (Alexeev 1980)primarilybasedonthe Bolivia). Sometimes, C. aquatica subspecies (Tzvelev higher chromosome number Hedberg & Hedberg 1976) are treated as species: C. capusii Franch. (China, (1977) found in the two African species known at that Iran, Iraq, Kyrgyz Republic, Tajikistan, Turkey, Uzbeki- time (2n =8versus2n = 4), is the only one of these stan) and C. pseudairoides (J. C. G. Herrm.) Tzvelev genera found in Africa. At present, in Colpodium s.s. (Azerbaijan, Iran, Iraq, Israel or Palestine, Russia there are two species in southeast Europe and south- (around the Caspian and Aral Seas), Syria, Turkey) west Asia, C. versicolor (Steven) Schmalh. (Georgia, Iraq, (Czerepanov 1995; Mill 1985; Wu & Phillips 2006). Iran, Russia [Caucasus], and Turkey) and C. violaceum Tzvelev (1976), in the GrassesofSovietUnion, (Boiss.) Griseb. (Iran and Iraq), and three in Africa, C. arranged these genera in the subtribe Poinae, in the Accepted for publication April 2011. 1 Department of Botany, MRC-166, Smithsonian Institution, Washington, DC 20013–7012, USA. e-mail: [email protected] 2 National Herbarium (PRE), South African National Biodiversity Institute, Private Bag X101, Pretoria 0001, South Africa. © The Board of Trustees of the Royal Botanic Gardens, Kew, 2011 © The Board of Trustees of the Royal Botanic Gardens, Kew, 2011 Table 1. Comparison of Colpodium, Catabrosa and Catabrosa drakensbergense. Cat. aquatica Cat. aquatica Catabrosa (not counting Cat. aquatica subsp. subsp. or var. subsp. Character\Taxon Colpodium Cat. drak.) Cat. drakensbergense aquatica uniflora pseudairoides Habit caespitose stooling (lateral shoots as in Catabrosa sometimes buried, frequently submerged), or cespitose 1 1= À 3= 1= Upper culm sheath near base or up to =5 4 4 2 proportion fused Spikelet length (biflorate) (mm) 2 – 4.2 2.5 – 4.2 2.5 – 4.2 2 – 2.8 (3) Spikelet length (uniflorate) (mm) 2.5 – 6.5 1.5 – 3.5 (1.9) 2.3 – 3.5 2.3 – 3.5 1.7 – 3.5 1.5 – 2.2 Number of florets per 11– 4 (7) 1, 1 – (2), 1 – 21– (2), 1 – 2, (1) 2 11– 3 (4) spikelet (3), 2 – 4 (7) Floret number variable or no yes, within and among panicles yes, within and among panicles highly variable probably variable yes, within not within panicle or and among among panicles panicles Rachilla extension in 1- suppressed suppressed or short suppressed, or up to 0.2 mm long flowered spikelets Glumes (subequal) equal unequal unequal ÀÁ fi 1 1 1 2 1 1 1 1 1 Lower glume relative to rst (subequal) equal or 6 À =2 2 À =3 the proximal lemma 6 À 3 =2 4 À =5 floret exceeding fi Upper glume relative to rst equal or exceeding 1 À 2 ðÞ3= ðÞ2= 3= to nearly equalling or 1 À 2 ðÞ3= 1 À 1= fl 3 3 4 3 4 2 3 4 3 2 oret slightly exceeding the proximal lemma Glume shape lanceolate to broadly broadly lanceolate to broadly ovate broadly lanceolate to oblanceolate, broadly lanceolate to oblanceolate, lanceolate, obtuse to elliptical to oblanceolate to apically obtuse or slightly acute broadly ovate to apically obtuse or acute obovate, apically slightly acute, elliptical to oblanceolate or slightly acute obtuse, sub-flabellate, or truncate to obovate, apically slightly acute, obtuse, sub- flabellate, or truncate Lemma texture relative to thinner than glumes same as glumes or thicker same as glumes or thicker glumes Lemma length (mm) 2.5 – 6.5 1.5 – 3.5 1.9 – 2.5 (2-fld.), 2.2 – 3.5 (1-fld.) 2 – 3 (3.5) 1.7 – 3.5 1.5 – 2.2 Lemma shape lanceolate to ovate, obtuse broadly lanceolate to oblong, broadly lanceolate to oblong, acute truncate to obtuse, acute or slightly to acute truncate to acute or slightly obtuse margin irregular obtuse Lemma no. of veins and vein 3, relatively faint 3, prominent, rarely with faint 3, prominent 3 3 3 prominence laterals (5) Lemma pubescence glabrous or pubescent glabrous or pubescent glabrous or sparsely minutely glabrous or sparsely glabrous or prominently pubescent minutely pubescent sparsely short minutely pubescent KEW BULLETIN VOL. 66(1) pubescent Palea surfaces and keels scarious to hyaline, or faintly subchartaceous, strongly subchartaceous, strongly coloured, green, keels slender, green coloured, keels prominent keels prominent or anthocyanic Lodicules with a well developed hyaline hyaline upper portion reduced hyaline upper portion reduced upper portion Hilum narrowly elliptic to oblong, ovoid, elliptic, oblong, or round, elliptic to oblong, darker than little differentiated in darker than caryopsis body caryopsis body colour from pale body Chromosome number: n= 2 (Asia), 4 (Africa) (5) 10, 15, 21 10 5, 10, 15, 21 5 CATABROSA VERSUS COLPODIUM (POACEAE: POEAE) IN SOUTHERN AFRICA following order: Poa L., Eremopoa Roshev., Catabrosella, chromosome number in Poeae is x =7,andthisis Hyalopoa, Arctophila Rupr., Dupontia R. Br., Catabrosa, common, widespread (only a dozen or so small genera Paracolpodium, Colpodium, Phippsia R. Br., Puccinellia deviate from this) and evidently pleisomorphic in the Parl., Sclerochloa P. Beauv., and Torreyochloa G. L. tribe, as it is also characteristic of the sister tribal lineage Church. Since then other relationships for the above Bromeae plus Triticeae (Clayton & Renvoize 1986). genera have been proposed taking DNA data into The molecular evidence supports the separation of account. Choo et al. (1994) examined chloroplast Colpodium s.s. and Catabrosa. Here, the morphological DNA restriction-site data for the Puccinellia complex and cytogenetic differences between Colpodium s.s. and including Catabrosa, Sclerochloa, Phippsia and Puccinellia. Catabrosa will be evaluated with regard to the disposi- Soreng & Davis (2000) provided additional restriction- tion of Col. drakensbergense. site evidence for this group within the Pooideae. This led to the establishment of a new subtribe for the above genera and the Colpodium complex, namely Materials Puccinelliinae Soreng & Davis (Soreng et al. 2003a), The types and additional material of the African which was originally proposed to include Catabrosa, Colpodium taxa were borrowed from UPS and PRE. All Colpodium, Paracolpodium, Phippsia, Puccinellia and cited specimens have been seen by the first author, the Sclerochloa. Although there are published analyses of second author has seen all PRE specimens. These, and plastid and nuclear DNA sequences with Colpodium in addition material of Catabrosa, Catabrosella, Colpodium, versicolor
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