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Elionurus purpureus E.J.Thomps. (Panicoideae: Andropogoneae: Tripsacinae), a new species for Queensland: circumscription and breeding system PDF

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Elionurus purpureus E.J.Thomps. (Panicoideae: Andropogoneae: Tripsacinae), a new species for Queensland: circumscription and breeding system E. J. Thompson Summary Thompson, E.J. (2017). Elionurus purpureus E.J.Thomps. (Panicoideae: Andropogoneae. Tripsacinae), a new species for Queensland: circumscription and breeding system. Austrobaileya 10(1): 139-162. A new species of Elionurus Willd. endemic to northeast Queensland is described and illustrated. It is distinguishable from E. citreus by the annual growth habit and spikelet morphology. The leaf anatomy of the two Australian species of Elionurus is illustrated and compared. The spikelet morphology of the two Australian species is contextualised within Elionurus, Tripsacinae and Rottboellinae. A new distinguishing morphological character for Elionurus is presented, viz. the proximal beak on the sessile spikelets, which is shared by the African genus Urelytrum Hackel (Tripsacinae), and some Australian genera in other subtribes of Andropogoneae. The breeding system that involves cleistogamy is discussed for the two Australian species of Elionurus. Key Words: Poaceae, Andropogoneae, Rottboelliinae, Tripsacinae, Elionurus, Elionurus citreus, Elionurus purpureus, Queensland flora, taxonomy, new species, cleistogamy, morphological characters, leaf anatomy, stem anatomy, cultivated plants E.J. Thompson, Queensland Herbarium, Department of Science, Information Technology and Innovation, Brisbane Botanic Gardens, Mt Coot-tha Road, Toowong, Queensland 4066, Australia. Email: [email protected] Introduction Elionurus Willd. is a genus of 15 species of spikelets (sessile and pedicellate) and a tropical and subtropical caespitose grasses, rachilla (Stapf 1934; Renvoize 1978; Watson 13 perennial and two annual. Nine (six & Dallwitz 1992). These spikelet pairs have endemic) species occur in Africa, five (two phylogenetic significance and are considered endemic) from southern North America, by Zanotti et al. (2010) to be the evolutionary four (three endemic) from South America, origin of the solitary spikelets in the sister one from India, and one species recorded for tribe, Paniceae. Australia and New Guinea (Clayton 1973; Tripsacinae is currently represented Clayton & Renvoize 1986; Watson & Dallwitz worldwide by a further six genera, none of 1992). Elionurus has been placed in the tribe which are native to Australia and includes the Andropogoneae subtribe Rottboelliinae significant food plant maize {Zea L.) (Soreng on the basis of morphology (Watson & et al. 2017). In the updated phylogeny of these Dallwitz 1992) and a molecular phylogeny grasses, Soreng et al. (2017) transferred five by Soreng et al. (2015), although Kellogg genera from Rottboelliinae to Tripsacinae that (2015) placed it in Andropogoneae, incertae had previously comprised only Tripsacum sedis based on earlier molecular phylogenies. L. and Zea, thus leaving Rottboelliinae Soreng et al. (2017) in an updated phylogeny with 16 genera. Species of Tripsacinae and placed Elionurus in subtribe Triplacinae. Rottboelliinae have unbranched inflorescences Members of the tribe typically have racemose and lack the conspicuous geniculate spiralled inflorescences composed of disarticulating awn on the sessile spikelets present in most segments made up of a pair of differentiated other subtribes of Andropogoneae (Clayton 1973; Clayton & Renvoize 1986; Watson & Dallwitz 1992; Kellogg 2015), although awnless spikelets also occur in subtribes Chionachninae and Coicinae (Soreng et al. Accepted for publication 21 July 2017 2017). Clayton (1973) used morphological 140 Austrobaileya 10(1): 139-162(2017) characters in a numerical analysis of species of male and female inflorescences in different Andropogoneae with awnless sessile spikelets parts of the same plant (Watson & Dallwitz covering 22 of the 23 genera from Tripsacinae 1992). All of the species of Tripsacinae and (except Zed) and Rottboelliinae and presented Rottboelliinae have bisexual flowers and similarity diagrams for the genera. Clayton most are solely chasmogamous (CH), where (1973) considered Rottboelliinae to be a anthers and stigmas emerge from florets with recent evolutionary branch disjunct from the ability to outcross and thereby provide other members of the tribe. gene flow (Watson & Dallwitz 1992; Simon & Alfonso 2011). The one known exception The previous single Australian species, until now is the presence of cleistogamy (CL) Elionurus citreus (R.Br.) Munro ex Benth., (self-pollination within a closed flower), in lemon-scented grass, occurs along the Rottboellia exaltata L.f. reported by Heslop- northern tropical and eastern subtropical Harrison (1959). Various types of CL have coast of Australia and New Guinea (Simon & been reported for species in other subtribes of Alfonso 2011). This species is described and Andropogoneae (Connor 1979; Campbell et illustrated by Tothill & Hacker (1983) and al. 1983; Watson & Dallwitz 1992; Culley & Jacobs et al. (2008). Klooster 2007). Lemon-scented foliage also occurs in This paper provides a taxonomic account other species of Andropogoneae, where of a new species of Elionurus with lemon- fragrance of leaves has been used sometimes scented foliage from north Queensland. The as a taxonomic character in keys to genera breeding systems of the two Australian species by Stapf (1934) and Simon (2002). Lemon- of Elionurus are presented and categorised scented foliage is found in the Australian according to current CL classifications grass Cymbopogon ambiguus A.Camus from the literature (Table 1). Morphological (Australian lemon-scented grass), several affinities of Elionurus to other genera of species and cultivars of Cymbopogon Rottboelliinae are also discussed. Spreng. (lemon grass, citronella grass) from Asia, and Elionurus muticus (Spreng.) Materials and methods Kunth (lemon grass) from South America Field survey at the site of the type collection and Africa. The Asian and South African of Elionurus purpureus was conducted in grasses are commercial sources of lemon- three successive years, April 2015, April 2016 scented essential oils (e.g., citral, citronellol, and May 2017. geraniol) (Soenarko 1977; Watson & Dallwitz 1992; Sangwan et al. 2001; Nakaharar et al. A comparative study of gross morphology 2003; Fuller et al. 2014; Kellogg 2015). Other and anatomy was made for E. purpureus essential oils (predominantly campherenone) and E. citreus. Morphological data for have been reported in leaves and roots of E. purpureus were obtained from four Elionurus elegans Kunth, an annual from accessions at BRI which includes the type Africa (Mevy et al. 2002). (Fig. 1). Spikelets from terminal and axillary inflorescences were dissected to examine their The breeding systems in grasses are highly characters. Six topotypes from Queensland diverse (Connor 1979) and in Panicoideae this Herbarium (BRI) collections of E. citreus is particularly so for the genera comprising were selected for sampling (Appendix 1). Tripsacinae as represented by Soreng et al. These specimens were collected in the vicinity (2017). Tripsacum L. and Zea provide unusual of the type and are considered representative diversity in breeding systems for subtribes substitutes for the type, a collection by Robert in Andropogoneae. Tripsacum has male and Brown {Bennett no. 6176) made in 1802 from female flowers in separate portions of the same Northumberland Island, Queensland. Online inflorescences, as is found in Chionachne images of the isotype {Andropogon citreus R.Br. (subtribe Chionachninae) and Zea has R.Br.) at E, K and W were examined. Thompson, Elionurus purpureus 141 Table 1. Morphological character differences between the Australian species of Elionurus Character Elionurus purpureus Elionurus citreus Growth habit annual perennial Inflorescence type multiple racemes at nodes at least single racemes at nodes on on cultivated plants cultivated plant and topotypes Culms Transverse section shape broadly concavo-convex narrowly concavo-convex Anatomy sclerenchyma along convex edge mostly 3 or 4 cells wide mostly 6-8 cells wide Leaf blade and sheath Distribution cauline mostly basal Margin prickle hairs absent medium (60-70 p) Abaxial surface length of stomata (pm) 45-60 41-51 length of silica bodies (pm) 15-20 8-15 Margins and mid-vein indumentum smooth usually scabrid Mid-vein obtusely keeled acutely keeled Anatomy of vascular bundles adaxial sclerenchyma absent present Sessile spikelet (mid raceme*) Total length (beak, body and lobes) 8.1-9.2, purple 10.6-13.1, pallid to pale pink. (mm), colour at maturity rarely purplish Lower glume lobe length (mm) 2.6-3.2 5.5-7 body width x length (mm), texture 1.7-2 x 3.5-4.2, chartaceous 1.5-1.7 x 3.6-4.3, cartilaginous indumentum type, orientation pubescent, appressed glabrous to pilose, ascending hair length (mm) c. 0.2 c. 2 keeled margin of lower glume not winged narrowly winged Upper lemma margin hyaline, pilose, hairs c. 0.3 mm long hyaline, ciliate apically or glabrous apex acute attenuate Callus length (mm) 1-2.3 1.8-2.7 * apical spikelets are smaller and basal larger Pedicellate spikelet Lower glume width x length (mm), colour at 0.8-1.3 x 5.2-57, purple; c. 0.9 x 7.2-8.5, pallid to pale pink. maturity, body, margins lanceolate, asymmetrical, 5-veined, rarely purplish, linear to narrow 1-keeled, both margins with oil lanceolate, asymmetrically. streak 3-veined, 1-keeled, one margin with oil streak Upper glume subequal to lower glume, 5-veined, unequal to lower glume, 3-veined, back rounded back laterally compressed Anther length (mm) Chasmogamous 0.9-2 1.3-2.7 Cleistogamous c. 0.8 0.7-1.8 Rachilla Width x length (mm), shape 1.2-1.4 x 4.3-4.4, distinctly clavate 0.9-1.3 x 3.3-5, clavate distinctly winged, unequally usually narrowly winged and bilobed unequally bilobed, or not winged Apex rim Sub-apical beard length of longest hairs (mm) 2.8-2.9 3.6-4.9 Pedicel Width x length (mm), shape c. 0.8 x 2.5-3.5 0.5-0.6 x 2.9-4 142 Austrobaileya 10(1): 139-162(2017) . Fig. 1 Holotype specimen of Elionuruspurpureus (McDonaldKRM16860 & Thompson, BRI). Thompson, Elionurus purpureus 143 Herbarium specimens and/or images scutellum. Germination was at ambient of type specimens of all the other species temperature on damp tissue paper in a covered of Elionurus and species in other genera transparent container in November 2016. of Tripsacinae and Rottboelliinae were Plants were cultivated in pots in a well- examined in detail (Appendices 1 & 2). drained potting medium in full sun in Leaf and culm transverse sections were Brisbane, Australia (lat. 27° 26’ 37”S). Plants prepared using a modified freehand sectioning were watered daily, unless there had been version of the method described by Frohlich sufficient rain, to maintain continuously (1984). Samples of herbarium material were moist potting medium. Watering rate was at rehydrated by initial immersion in hot water about 7 mm per day, measured using a rain and left to soak for up to several days. Leaves gauge, equating to about 1000 mm for the from upper culms were used and sections wet season (December - March). Plants were taken from near the middle of each leaf. Leaf occasionally fertilised with a commercial samples were placed on a glass slide covered pelletised chicken manure. with a glass slide cover that served as a cutting Rainfall data from Climate Data Online guide. Culm sections were cut without the use (BoM) for weather stations near the collection of the slide cover. Thin sections were cut using localities of BRI accessions of E. purpureus a razor blade while viewing under a binocular were examined to investigate the potential microscope at *40 magnification. Several effect of soil moisture levels on plant growth sections were made for both E. purpureus and flowering. Frequency of rainy days of and E. citreus. Three accessions of E. citreus more than 5 mm and more than 10 mm are were examined to capture variation and recorded in Table 2 as a comparison with the because some leaf blades retained distorted watering regime of cultivated plants. parenchyma following rehydration. Leaf blades and culms of near equivalent width Spikelets from the cultivated plants were were used for both species. examined in detail at flowering and fruiting in April - June 2017. Spikelets from the upper Leaf, spatheole and culm surface replicas and lower halves of racemes were harvested on were prepared using the method described by a daily basis as they matured. The most distal Hilu & Randall (1984). spikelet of racemes disarticulated readily at Micromorphology was studied from maturity. Ripe spikelets were gathered by images taken using a compound binocular gently touching. The proportion of CH:CL microscope at 5x, 10x? 20 x and 40 x for was examined by randomly selecting 20 transverse sections, and at 20x and 40x for spikelets for two batches, each batch from the surface replicas. upper halves and lower halves of racemes. Plants of E. citreus and E. purpureus were Botanical terminology follows Beentje cultivated for three main purposes: to study (2010) and Harris & Harris (1994) for general any impact of environment on growth habit, usage. An exception is the use of the term effect of environment on the breeding system, convexo-concave to differentiate the shape of and provide a supply of viable caryopses for the culm transverse section of E. purpureus. further studies. Caryopses were taken from Anatomy descriptions and terminology follow the type of E. purpureus and BRI accession, Metcalfe (1960), Ellis (1976, 1979), Renvoize McDonald KRM11354 for E. citreus. The (1982) and Watson & Dallwitz (1992). caryopses were scarified by scraping off a small portion of pericarp just above the 144 Austrobaileya 10(1): 139-162(2017) Taxonomy Key to the Australian species of Elionurus 1 Perennial; spikelets pallid to pale pink; sessile spikelet with lobes longer than body; body of lower glume pubescent; upper glume glabrous on margin; NE NSW, EQld,N NT, NWA.E. citreus 1. Annual; spikelets purple at maturity; sessile spikelets with lobes shorter than body; body of lower glume pilose or glabrous; upper glume pilose on margin; N Qld.E. purpureus Elionurus purpureus E.J.Thomps., sp. nov. long. Glumes dissimilar; lower glume 6.1-7.5 similar to E. citreus (R.Br.) Munro ex Benth. mm long; body 1.7-2 x 3.5-4.2 mm, broadest in differing by the annual growth habit and middle, 9-veined, symmetrical, chartaceous; purple spikelets 8.1-9.1 mm long with body of back flattened, pubescent with appressed pale lower glume longer than the lobes and margin hairs c. 0.2 mm long or sometimes glabrous; of upper glume pilose. Typus: Queensland. margins keeled, keels not winged, with two Cook District: 0.8 km along Mt Spurgeon rows of pectinate white bristles to 1.3 mm road from Mulligan Highway, Mt Carbine, long at 45° to margin decreasing in length 10 April 2015, K.R.McDonald KRM16860 & towards apex, sub-margins with broad dark E.J.Thompson (holo: BRI). purple glabrous oil streak continuous with lobes; apex 2-lobed, 2.6-3.2 mm long, obtuse, Erect caespitose annual to 1.3 m high. Culms lobes fused in lower third to half; upper glume < 4 mm wide, shortly branched, nodes 4.2-4.5 mm long, symmetrical, lanceolate, 3-9; internodes purplish, smooth, hairless, membranous, 3-veined, carinate, midvein furrowed; nodes glabrous often purplish, half with short erect proximal hairs; margins to two-thirds of total length exserted from winged, wing c. 0.3 mm wide, membranous, sheath. Basal leaf sheaths purplish; culm pilose; short appressed to ascending hairs sheaths with scattered white hairs mostly near on back, glabrous in upper 1/3, apex acute. apical margins; collar margins with hairs to Florets 2; lower floret neuter, lemma c. 3.5 3.3 mm. Ligule a fringed membrane, c. 0.9 mm long, lanceolate, hyaline, veins indistinct, mm. Leaves cauline, blades up to 45 cm long margin pilose, apex acute. Lower palea absent. and 0.8-2 mm wide, discolorous, cauline, Upper floret hermaphrodite, upper lemma, keeled, pilose adaxially, hairs to 0.2 mm, 2.5-2.8 mm long, elliptic, hyaline, veins margins smooth, inrolled on drying, lemon- indistinct, margin pilose, apex obtuse. Upper scented when crushed. Inflorescences of single palea absent. Lodicules 2, c. 0.5 mm long. spatheolate one-sided racemes, 4 to 13 cm Anthers chasmogamous 3, 0.9-2 mm long, long. Peduncle absent. Up to 9 nodes with 1-3 cream when fresh, or cleistogamous 3 and c. one-noded branches each bearing a raceme; 0.8 mm long. Caryopses c. 1 x 0.6 x 2.5-27 axillary racemes exserted, partly exserted mm, dorsiventrally compressed, broadest in or enclosed with spatheole; spatheoles c. the middle, bowed longitudinally, buff to light 4.5 mm wide and up to 13 cm long, pilose; brown; scutellum c. 1.4 mm long; hilum c. 0.5 culms extending to 15 cm beyond spatheole. mm long. Pedicellate spikelet neuter; glumes Spikelets paired (sessile and pedicellate), dissimilar, subequal, 5-veined, chartaceous, dimorphic, purple at maturity, closely lanceolate, lobes orientation parallel to body; overlapping in two opposing staggered rows lower glume 0.8-1.3 x 5.2-57 mm, dorsally along disarticulating jointed axis; diaspore compressed, back glabrous, asymmetrical, composed of paired spikelets and rachilla. broadest at the base, acute apex; 1-lobed, Callus elliptical, c. 0.4 x 1-2.3 mm. Sessile lobe orientation horizontal; margin 1-keeled, spikelet fertile, dor si-ventral ly compressed, with a single row of pectinate white bristles 1.4-1.6 x 8.1-9.2 mm , gradational (larger c. 0.6 mm long decreasing in length apically, from base to apex), proximal beak 2-2.2 mm submargins with oil streak,; upper glume, c. Thompson, Elionurus purpureus 145 0.8 x 4.5-5.3 mm, glabrous, symmetrical, key character annual, specimens key to E. laterally compressed; back convex, carinate hensii K.Schum from the African continent, in upper half, keel pilose with stiff parallel although examination of the online image hairs c. 0.2 mm long; apex acute; 1-lobed, of the isosyntype provided confirmation lobe orientation vertical; margins flat. Pedicel of the dissimilarity with E. hensii. If the c. 0.8 x 2.5-3.5 mm, free from internode, growth habit is assumed to be perennial then clavate, hollow, plano-convex in cross- specimens key to E. citreus but do not have section, adaxial wall membranous and abaxial the lower glume with lobes longer than the wall chartaceous; abaxial edge pubescent with body. Table 3 lists the character differences white hairs, short and appressed along most between E. purpureus and E. citreus and Fig. of length and with a longer sub-apically tuft 4 provides images of the respective spikelet to 2.4 mm. Rachilla 1.2-1.4 x 4.3-4.4 mm, pairs for comparison. dissimilar to pedicel, free from internode, Diaspore dispersal in the Australian clavate, hollow, plano-convex in cross- species of Elionurus occurs via two modes. section, adaxial wall membranous and abaxial Most of the diaspores readily disarticulate wall chartaceous, abaxial edge pubescent from the racemes at maturity and can disperse with short white appressed hairs; apex c. 1.2 x by various vectors in similar ways to other 2.3 mm, oblique, adaxially flanged, unequally grasses. Secondly, the basal spikelet pair bilobed, subapical ring of white hairs to 2.8- that is CH or CF of each raceme is resistant 2.9 mm long and longest on abaxial margin; to disarticulation. This delayed release of disarticulation scar c. 0.7 mm x 1.5 mm, flat, the spikelet may have some advantage with elliptical. Figs. 1, 2, 3 & 4. dispersal allowing the spikelet to fall further Additional specimens examined: Queensland. Cook from the mother plant after the culms collapse. District: 9 km from Koolburra on the track S from Koolburra to the Kimba road, Jun 1981, Clarkson 3699 Appendix 2 lists spikelet differences (BRI); Conglomerate Creek, S of the Deighton Road, for all the species of Elionurus. This on Escort Creek Holding, SE of Laura: Lot/Plan 198/ comparison shows that the Australian species SP273726, Apr 2017, Forster PIF45151 & McDonald exhibit morphological similarities between (BRI, MEL); 0.8 km along Mt Spurgeon road from Mulligan Highway, Mt Carbine, May 2017, Thompson themselves but are distinct from the other EJT1083 (BRI). Cultivated. Ashgrove, Apr 2017, species. Only the Australian species have Thompson MOR804 (BRI). the pedicellate spikelets composed of just the Distribution and habitat: Elionurus two glumes and they always neuter. The non- purpureus occurs west of the Great Dividing Australian species have developed pedicellate Range in the range latitude 15°15’ to 16°45’S spikelets with two lemmas and anthers. This and longitude 143°30’ to 145°15’E (Map 1). variation in development of the pedicellate It is found in woodland mostly dominated by spikelet within a genus is uncommon in eucalypts including Corymbia clarksoniana Andropogoneae but has parallels in Arthraxon (D.J.Carr & S.G.M.Carr) K.D.Hill & Beauv. (van Welzen 1981). Furthermore, the L. A. S. Johnson, Eucalyptus leptophleba Australian species have a distinctive character F.Muell. and E. tetrodonta F.Muell. on sandy of the sessile spikelets that they share with alluvial soil adjacent to intermittent creeks or most of the other species. The lower glumes lagoons. have conspicuous submargins that Clayton & Renvoize (1986) referred as a brown oil Phenology: Elionurus purpureus flowers in streak and Watson & Dallwitz (1992) called March to June during the wet season and early glandular. Three species, Elionurus elegans dry season. Kunth, E. hirtifolius Hack, and E. royleanus Notes: The identification key to Elionurus A.Rich. from Africa to NW India, lack this oil species of the world provided by Renvoize streak. These species differ by having tufts of (1978) was used to verify that E. purpureus hairs along the margins of the lower glume is not an introduced species as specimens of the sessile spikelets instead of the row of fail to key out satisfactorily. Using the parallel hairs present in the other species 146 Austrobaileya 10(1): 139-162(2017) E ^ D 3 § OL I §>! § | si "Sg J<UD ‘MEL +O3 B S n in I(cid:9632)oS Q®r a b £o &© .2 T3 w ! 1 5 ^ A2 >!—i kys 2o ho 3 Rj s 1II? s u u ur n of Elio UP>CI > &£- ss'- o b o<touf l -Sog cCIo M n species s iU.ba 3eo t3so §$ 3o©ugo 3-T£©g3 ct3 alia > ucS & E&^ Ea? sa 1PC >b ls str u A e • Su h b b y to t S PC 5cOo?h -QS2 o 5 S) pl E 2 p n P. a at h ose t & £ b b h th tbof i O> -trot 033 wit u§ sHi "32 SB1 O J s OS Qw TO3 OOh ds u u u o r o h to ut > a *“1 s ght s,©a| « o y ei / (cid:9632) — _ d b S 'St« .5©5 S« shSi i e hJ to nt u (cid:9632)3 o © 3 s prese upiic Mug Z a Cca3> TG(U3 £Qa*J 1«."S3C +Eg- 3©W) t3: o a e a e 5^ c a o P S - §D n 1 J § L) i ^ u « C y ( m a *d J £ £ eistog >E§r *©* £3« S0©0h S. of cl £ c g '<5cb s e ori g e at C able 2. bOh UO£h 2bQ^ lUo< £3 s§. t3ug tIt©oo sCoIh T Thompson, Elionurus purpureus 147 (see Fig. 4). However, the three species have have single slender culms to 30 cm high while similarities to the Australian species by the all cultivated plants had multiple culms, one lower glume of the pedicellate spikelets being main robust culm to 1.3 m high and up to 4 asymmetrical with the apex lobed or awned subordinate ones to 50 cm high. Cultivated (Appendix 2). plants had much larger leaves, to 6 mm wide x 42 cm long, compared to the type with Harvesting of ripe spikelets from the leaves 1-2 mm wide x 18 cm long. All of cultivated plants of E. purpureus to collect the cultivated plants closely resembled BRI caryopses revealed a further character accession Clarkson 3699 (Fig. 2). uncommonly represented on herbarium specimens. Spikelets on each raceme ripen There were also differences in the and disarticulate successively until the basal inflorescences of cultivated plants. The one that is resistant to detachment from the cultivated plants of E. purpureus had racemes culm but ultimately falls. The specimens of to 13 cm long in fasciculate inflorescences the species of Elionurus observed for this comprised of up to three racemes on branches study frequently had over-mature racemes arising at as many as nine culm nodes. The preventing observation of this phenomenon, holotype and accession Forster PIF45151 although it was present on one specimen of & McDonald have single racemes to 8 cm E. citreus at BRI. The herbarium specimens long on short branches at up to three nodes. usually had complete racemes and sometimes The cultivated plant of E. citreus had single partially disarticulated ones. Permanent racemes at the nodes as was observed for the retention of the basal spikelet is uncommon topotypes listed in Appendix 1. in Australian Andropogoneae but has been The study of the wet season rainfall data reported by Veldkamp et al. (1986) and presented in Table 2 resulted in equivocal observed on herbarium specimens by the conclusions. The sample size is too small for author for Thaumastochloa C.E.Hubb., also any clear trends but there is some evidence Rottboelliinae. to support a hypothesis that December rain Very few of the caryopses obtained from of c. 100 mm may be responsible for plants BRI specimen accessions were viable. Several developing the taller growth habit. This caryopses from the holotype of E. purpureus seems plausible since no intermediate sized germinated and two plants survived to plants were observed despite the presence of maturity. One caryopsis of E. citreus from soil moisture gradients at least for some of McDonald KRM11354 (BRI) germinated and the collection sites (pers. comm. PI. Forster). the plant survived to maturity. The possibility of dimorphic growth habit requires further investigation, particularly Observations from the cultivated progeny from further field survey. of E. purpureus revealed striking differences from the type that initially appeared to be the Breeding system only collection. This promoted a review of In order to categorise the CL found in the 148 accessions of E. citreus at BRI. One Elionurus, the classification schemes of CL of the accessions, Clarkson 3699, matched presented by eight authors were investigated. the cultivated plants. At the point of nearing These schemes are of two broad types in completion of this manuscript another terms of taxonomy, for flowering plants in specimen, Forster PIF45151 & McDonald, general and more specifically for grasses. was incorporated at BRI. This accession The classifications by Campbell et al. (1983) is comprised of plants that are identical to and Hackel (1906) provide more specific the type in growth habit and inflorescence categories relevant to Poaceae. In summary, composition. the schemes assess CL by the presence of The difference in the growth habit of various morphological modifications of the type from its cultivated progeny was the CH morph or whether CL is induced by considerable. The plants comprising the type environmental conditions. The categories 148 Austrobaileya 10(1): 139-162(2017) | QUEENSLAND HERRARIUWli »RlS»AHE~| flora of QUEENSLAND COOK district M R 1 R 1 15-22S 143.54.E(,i,. l coll J.R. Clarkson 3699 25 June 1981 Elionurus citreus (R.Br*) Munro ex Benth, I Fomily 1 HobHa* |*|»|P-|p|fl lyWi |j I ~ IGRAMI -JbWfrfl BRI “"P** *nlfV only- N°.tob» cited in popart. 9km from Koolburra on the track south from Koolburra to the Kimba road. Eucalyptus tetrodonta woodland* An erect grass, Common along the track. QRS, NSW, CANB, PERTH, NT, K, US, PRE dUtENSi.ANb HER BARIUM 294155 brisbTnF Fig. 2. Paratype of Elionurus purpureus (Clarkson 3699, BRI).

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