Anthoxanthum odoratum L., Sp. Pl. 28 (1753).

T: habitat in Europae pratis, n.v.

Vegetative form. Perennial, caespitose. Leaves scattered along the culms. Culms slender or robust (9.5-)30-60(-105) cm high, unbranched above, 2-3 noded. Mid-culm nodes glabrous, exposed, pigmented, constricted. Mid-culm internodes hollow, often pigmented below the nodes, usually glabrous or puberulous (rarely, the hairs sparse below the node), terete. Young shoots extravaginal. Shoots aromatic (coumarin scented). Leaves non-auriculate (but with long setae at the blade-sheath junction). Basal leaf sheaths not keeled, terete, firmly encircling the culm, usually glabrous or pilose, the hairs variable in density and distribution but usually restricted to the upper part of the sheath, usually the same colour as the lamina or purple (often at the base of upper sheaths, the pigmentation more extensive and conspicuous in the basal sheaths), with the veins equally striate, with margins free, hyaline to membranous, occasionally pigmented, smooth. Ligule (1-)2-6 mm long, hyaline, occasionally pigmented, smooth or ciliolate (minutely), obtuse, entire. Collar glabrous. Leaf blades flat, linear, (20-)30-310 mm long, (1.5-)2-9 mm wide; rarely adaxially glabrous or adaxially scabrous or adaxially pilose (the hairs variable in density), shallowly grooved to not grooved; rarely abaxially glabrous or abaxially scabrous or abaxially pilose (the hairs variable in density), with midrib prominent (in the lower 1/3); with margins smooth or with margins scabrous and with margins ciliate, apices acute, flat or hooded.

Reproductive organization. Plants bisexual. Rudimentary spikelets usually at the base of the inflorescence (and at the base of primary branches in the lower half of the inflorescence). Incomplete spikelets absent. Hermaphrodite florets protogynous. Plants self incompatible. Inflorescence of chasmogamous spikelets.

Inflorescence. Inflorescence a panicle, straw coloured, green or purple, erect, contracted, 5-18 mm wide, narrowly oblong to ovate or interrupted (rarely), symmetrical, fully exserted. Main inflorescence axis (60-)100-490 mm long. Peduncles (45-)80-420 mm long, sometimes pigmented in the upper part, glabrous, smooth. Rachides 15-75(-120) mm long, terete, glabrous. Primary inflorescence branches glabrous; solitary or paired, branching at the base, spreading, spiral. Spikelets on first order branches or on second order branches, erect, spreading.

Hermaphrodite (`perfect') spikelets. Hermaphrodite spikelets subsessile and pedicellate, 6-10.2 mm long, strongly laterally compressed, narrowly ovate, not disarticulating as a separate unit, disarticulating above the glumes. Pedicels short, 0.4-2 mm long, erect, occasionally pigmented, slender, rarely glabrous or pubescent, usually straight. Glumes two per spikelet, similar, unequal in length, equalling the proximal lemma and longer than the proximal lemma (G1 equal and G2 longer). Lower glume c. 1/2 the length of the upper glume, (dorsally) ovate, 3.5-5.1 mm long, 0.67-0.85 mm wide, hyaline to membranous, keeled (the keel green), acute, entire, apiculate, margin hyaline to margin membranous, margin smooth or margin ciliolate (rarely); 1 veined, veins prominent, rarely the midvein glabrous or midvein scabrous to midvein pilose (toward the base); rarely intercostal regions glabrous, or intercostal regions pubescent (the hairs varying in density and usually only on one half of the glume). Upper glume large, completely enclosing the florets, (dorsally) narrowly ovate, 6.6-10.2 mm long, 0.67-0.83 mm wide, membranous, keeled (the keel green), acute, entire, apiculate, margin membranous, margin smooth; 3 veined, mid veins prominent, midvein glabrous or midvein scabrous to midvein pilose (towards the base); usually intercostal regions glabrous, or intercostal regions papillose, or intercostal regions puberulous (the hairs variable in distribution and sparse). Rudimentary florets absent. Incomplete florets present, proximal to the hermaphrodite florets. Proximal incomplete florets 2 per spikelet, neuter (reduced to well developed lemmas), awned, epaleate. Incomplete lemmas becoming decidedly firmer than the glumes, clearly longer than the hermaphrodite lemma, about equal in length, (dorsally) narrowly oblong or narrowly elliptic, 2.8-3.5 mm long, 0.33-0.67 mm wide, becoming chartaceous (and golden brown at maturity), not keeled, apices rounded, shortly bifid (the two lobes colourless, glabrous), awned, with margins ciliate (the hairs long, c. 1.6mm, and silky golden brown); 5 veined. Incomplete lemma veins not confluent apically, obscure; pilose (the hairs golden brown), with hairs on all veins; intercostal regions pilose (the hairs golden brown), the hairs over most of the dorsal surface (but not the lobes). Incomplete lemma awns 1, median. Incomplete lemma median awn dorsal (arising from the midpoint or upper half of the first lemma and from near the base of the second lemma), much longer than the body of the lemmas, markedly unequal in length (the awn of the first lemma 2.25-3.4 mm long, relatively slender, straight; the awn of the second lemma 6.6-8 mm long, stout, geniculate with a twisted column pigmented at maturity), 2.25-8 mm long. Hermaphrodite florets 1 per spikelet, symmetrical (?), becoming shiny and golden-brown at maturity. Rachilla not disarticulating between the florets, glabrous, not obviously elongated between glumes or florets, not apically prolonged. Callus absent. Lemma completely enclosing the palea prior to anthesis and becoming decidedly firmer than the glumes, (dorsally) narrowly elliptic (but rotund when flat), 1.7-2.55 mm long, 0.4-0.6 mm wide, becoming indurate, not keeled, obtuse to truncate, emarginate, muticous. Lemma margins smooth. Lemma veins obscure, glabrous; intercostal regions glabrous. Palea fully developed, equalling the lemmas, thinner than the lemmas, tightly clasped by the lemmas, narrowly elliptic, 1.2-2 mm long, 0.25-0.42 mm wide, hyaline, keel-less, obtuse, entire, margin smooth; 1 veined; intercostal regions glabrous. Lodicules absent. Stamens 2. Anthers 3-5.6 mm long, basally 2-lobed. Ovary glabrous. Styles 2, apical, free to their bases.

Fruit. Fruit free from both lemma and palea, elliptical, 1.4-1.75 mm long, 0.5-0.7 mm wide, not grooved, glabrous. Embryo 0.6 mm long, not waisted.

Abaxial leaf blade epidermis. Leaf anatomical data recorded.

Costal/intercostal zonation conspicuous; the intercostal zones bordering the midrib 12-15 cells wide; epidermis differentiated into long- and short-cells; long-cells different in shape costally and intercostally (the costal long-cells narrower); long-cells of similar wall thickness costally and intercostally (fairly thin-walled).

Microhairs absent.

Crown cells absent. Prickles present; costal and marginal (and also present intercostally in RSBS 33, Black Mtn. ACT); antrorse and retrorse; variable in size and form to fairly uniform in size and form; of two types (as hooks and macrohairs). Prickle bases not paired with a short-cell. Costal prickles along all zones; frequent in their files. Bases of the costal prickles about as long as the width of an intercostal long-cell to longer than the width of an intercostal long-cell; barbs of the costal prickles about as long as the bases, or more than twice as long as the bases. Macrohairs absent, or present; when present, costal; intergraded with long prickles; unicellular; lax; with thickened walls; sparse but frequent, or infrequent and irregularly dispersed; more than twice as long as an intercostal long-cell. Macrohair bases one-celled; attached at cuticle only; uniform in mode of insertion.

Intercostal long-cells fairly constant in shape to variable in shape; varying in shape within a single intercostal zone (being fusiform centrally and rectangular laterally). Mid-intercostal long-cells markedly elongated; fusiform; hexagonal to inflated; long-cell walls straight to tessellate; the undulations regular; end walls vertical, or angled; the intercostal long-cell walls associated with conspicuous pitting; outer surfaces of intercostal long-cells not pitted. Papillae absent.

The costal zones all histologically similar; costal short-cells solitary. Costal silica bodies present but imperfectly developed; throughout the costal zones; horizontal-smooth, horizontal-crenate, and cuboid.

Intercostal short-cells absent; silica bodies absent.

Stomata infrequent; absent from some intercostal zones; arranged in definite rows; 43.5-49-57 m long; with guard-cells overlapped by the interstomatals. Subsidiaries parallel-sided.

Photosynthetic pathway and related features. C3. XyMS+.

Transverse section of the leaf blade. Lamina mid-zone in transverse section open; more or less flat. Width of lamina across a primary vascular bundle 135-156-183 m. Lamina mid-zone in transverse section exhibiting adaxial ribs only. Adaxial furrows slight; wide. Adaxial furrows between all the vascular bundles. Adaxial ribs more or less constant in size; round topped; opposite all vascular bundles. Vascular bundles in the mid-lamina 14-20; 1 per rib. Abaxial furrows absent.

Midrib pronounced in outline; adaxially raised and rounded; skightly prominent abaxially; tissue layout similar to that of other primary vascular bundles. Vascular bundles in the mid-lamina region of the midrib 1. The median vascular bundle with a protoxylem cavity; with an enlarged protoxylem vessel; without sclerosed phloem. Midrib without colourless tissue adaxially; with thin-walled sclerenchymatous cells adaxially; without lacunae. The lamina symmetrical on either side of the midrib.

Mesophyll chlorenchyma non-radiate; loosely packed. Mesophyll without lacunae; without any obvious adaxial palisade; without `circular cells'; not traversed by columns of colourless cells; without arm cells; without fusoids. Bulliforms present in discrete groups. The bulliform groups situated between vascular bundles. The bulliform groups without contiguous colourless mesophyll cells; small to large; simple, fan-shaped. Abaxial epidermis of bulliform-like epidermal cells. Abaxial epidermal cell walls not thickened. The cells regular in shape, rounded.

The major vascular bundles interspersed with minor bundles; outlines of primary vascular bundles more or less circular to more or less elliptical; primary vascular bundles centrally situated. Primary lateral vascular bundles with adaxial sclerenchyma; with abaxial sclerenchyma; the adaxial sclerenchyma forming girders; the abaxial sclerenchyma forming girders; the combined girders nowhere forming `figures'. Outlines of lower order vascular bundles more or less circular. Lower order vascular bundles centrally situated; with adaxial sclerenchyma; with abaxial sclerenchyma; the adaxial sclerenchyma forming strands, or forming girders; the abaxial sclerenchyma forming strands, or forming girders; the combined girders of the lower order vascular bundles nowhere forming `figures'; the smallest vascular bundles with sclerenchyma. The adaxial sclerenchyma of the mid-lamina all associated with vascular bundles.

Cytology. 2n = 20.

Common name. Sweet Vernal Grass.

Distribution. Introduced. Tasmania, New South Wales, Australian Capital Territory, Victoria, Western Australia, Queensland, and South Australia. World distribution: Western Eurasia, USSR, Mediterranean, Eastern Asia, and North America.

Ecology. Usually in damp sites. Flowers Sept.-Feb. Fruits Nov.-?.

Vouchers. Specimens examined morphologically: W.A.: roadside, Inlet Drive, Denmark, M.McCallum Webster WA/38 (PERTH); Pemberton, M.Koch 2629 (PERTH); Fernhook Falls, Deep River, A.H.Burbidge 3948 (PERTH); Mt Barker, Nov 1927, G.Robertson (PERTH); Albany, Dec 1923, W.Cattan Grasby (PERTH); Harvey, Dec 1924, G.C.Rath (PERTH); Forest Grove, 27 Dec 1979, D.L.Serventy (PERTH). S.A.: margin of Kuitpo Forest on Peters Creek Rd, 14 Nov 1983, D.E. Symon 13452 (AD, CANB, LG, US, MO, BRI). Qld: Springbrook Rd, Springbrook, opposite "Mouses House", dairy farm, 4 Nov. 1989, W.Pieters (BRI 447332, NSW, PERTH, K). N.S.W.: herbaceous swamp 15 km NNE of Guyra, New England Plateau, K.Paijmans 2681 (CANB); Tallaganda State Forest, between Queanbeyan and Braidwood, Southern Tablelands, 1036 m, R.Pullen 4548 (CANB, NSW, BRI, MEL, HO, AD); Tallaganda State Forest, Gourock Ra, SE of Bungendore, Southern Tablelands, 1036 m, R.Pullen 4549 (CANB, PERTH); rd from Fitzroy Falls to Moss Vale, Southern Highlands, R.Pullen 4409 (CANB, NSW, CHR, TNS). A.C.T.: 8.7 km S of Parkers Gap along the fire trail, 18 km by road from Captains Flat, J.G.West 5156 (CANB). Vic.: c. 8 km E of Victorian-S.A. border, Casterton Rd, I.B.Wilson 610 (ADW, CANB). Tas.: Mt Wellington, Hobart, P.J.Darbyshire 1068 (CANB, NSW); Mt Michael, c.2 km N of Poimena, A.Moscal 13235 (HO); Little Split Rock, Central Highlands, 1200 m, A.Moscal 6731 (HO +?); Middlesex Plains, Central Highlands, 760 m, A.Moscal 1018 (HO); Leven R mouth to Somerset, S.J.Berrigan 50 (HO); Pipeclay Beach, George Town, T.E.Burns 799 (HO); Pigstys, Recherche Bay, D.I.Morris 8107 (HO); Safety Cove, Port Arthur, grassy roadside verge, P.Collier 1717 (HO); Gowan Brae Rd Bridge over Nive River, P.Collier 1076 (HO); Flinders Is., Furneaux Group, Lady Barron, J.S.Whinray 1808 (CANB); Munro's, Lower Marshes, Jordan River, 9 Nov 1964, R.Barker (CANB).

Material examined anatomically: ACT: 8.7 km S of Parkers Gap, J.G. West 5156; Black Mtn., RSBS 33. S.A.: Kuito Forest, D.E.Symon 13452. Tas.: Mt Wellington, Hobart, P.J.Darbyshire 1068.

Classification. Pooideae; Poodae; Aveneae.

References. Cytology: Hubbard, 1984. Illustrations: anthox02.gif anthox14.gif anthox15.gif anthox17.gif anthox25.gif

Cite this publication as:
C.M. Weiller, M.J. Henwood, J. Lenz and L. Watson (1995 onwards). `Pooideae (Poaceae) in Australia - Descriptions and Illustrations'. URL http://muse.bio.cornell.edu/delta/
Dallwitz (1980) and Dallwitz, Paine and Zurcher (1993) should also be cited.

References and Acknowledgements