The Eragrostiella Bsr (, Eragrostideae)

M. Lazarides

Herbarium Australiense, CSIRO, P.O. Box 1600, Canberra City, A.C.T. 2601.

Abstract Lazarides, M. The genus Eragrostiella Bor (Poaceae, Eragrostideae). Contrib. Herb. Aust. 22: 1-7 (1976). The taxa of Eragrostiella Bor are reviewed. As a result, the genus is recorded in and for the fist time, and E. brachyphylla (Stapf) Bor for the first time in (Ceylon). Two species are reduced to varietal rank, and one varietal name to synonymy. The genus comprises five species, one with three varieties. A key to their identification is provided. The Australian is considered to have resurrection properties.

Eragrostiella Bor, Indian For. 66: 269 (1940). Type species: E. leioptera (Stapf) Bor (Eragrostis leioptera Stapf). Eragrostiella comprises a small group of closely allied taxa originally described in Eragrostis, which are segregated primarily on the basis of their spicate inflorescence and their compact habit of basal leaves and persistent leaf sheaths. Further details of the history of the genus are available in Bor (loc. cit.). On review, the genus consists of seven taxa, the majority confined to the Indian and Indo-Chinese region, but one species having a disjunct distribution in the Old World tropics.

DIAGNOSTIC CHARACTERS OF THE GENUS Inflorescence a solitary terminal spike of subsecund biseriate mainly imbricate spikelets, the rhachis scabrous-pubescent on the upper surface. Spikelets laterally compressed, with many to numerous florets, the rhachilla usually tough. Florets mainly bisexual, closely imbricate. Glumes deciduous, equal or slightly unequal; lower 1-nerved, with an acute scabrous keel; upper 1-sub 3-nerved, thickened and rounded on the back, usually acutely keeled near the apex. Lemmas deciduous, 3-nerved, usually glabrous, the lateral nerves usually somewhat closer to the margins than to the midnerve. Paleas subequal in length to their lemmas, per- sistent, usually glabrous on the back and flaps, with narrowly winged usually ciliolate keels. Grain obtusely trigonous to almost terete, obtuse, plump, reticulately striate, the embryo about half as long as the grain itself. Slender perennials, com- pactly tufted; culms simple; foliage mainly basal, with ribbed nerves; leaf sheaths persistent, the basal ones becoming fibrous; ligule a short fringed membrane; auricles usually bearded; blades convolute or complicate or flat, often filiform, usually scabrous-hispid and pilose on the upper surface. 2 M. Lazarides

KEY TO THE TAXA OF ERAGROSTIELLA

1. Keels of palea with relatively broad (c. 0.3 mm), glabrous wings; collar represented by a thin groove; rigid plant with thin, rolled, subpungent blades 20-30 cm long ...... 1. leioptera 1. Keels of palea with narrower, ciliolate wings; collar absent; rather delicate; blades thin or thick, rolled or flat, subacute or obtuse, usually shorter than 20 cm (except bijaria var. walkeri) 2. Palea scabrous-pubescent on the back and flaps; spikelets purplish ...... 2. collettii 2. Palea glabrous on back and flaps (rarely puberulous or scaberulous on the back near the apex); spikelets leaden grey, olive green or straw coloured 3. Leaf blades glabrous and smooth on the upper surface, flat or complicate, obtuse, coriaceous, 1.5-3 mm wide (flattened) ...... 3. brachyphylla 3. Leaf blades densely scabrous and usually pilose on the upper surface, usually convolute, subacute, not coriaceous, and much narrower 4. Leaf sheaths densely scabrous, the hairs extending upwards on the lower surface of the blades; blades rather densely pilose on upper surface; spikes 2.5-5 mm wide; spikelets 8-9 X 2-2.5 rnm, c. 13-flowered; lemmas lanceolate ...... 4. nardoides 4. Leaf sheaths and lower surface of blades glabrous and usually smooth; blades glabrous or sparsely pilose on upper surface; spikes 5-10 mrn wide; spikelets 6-20 X 1.5-4 mm, 9-34-flowered; lemmas ovate ...... 5. bifaria 5. Spikelets 10- 14 X 2-4 mm, 12-23-flowered, narrow-elliptic, olive green; lemmas c. 3 mm long, slightly longer than wide, acute ...... 5b. var. secunda 5. Spikelets 6-20 X 1.5-2 mm, 9-34-flowered, narrow-oblong, leaden grey or straw coloured; lemmas 2-2.5 mrn long, slightly wider than long, subacute or obtuse 6. Plant 30-45 cm high; culms 1-2-noded; leaf blades mainly basal, usually con- volute filiform and less than 20 cm long ...... 5a. var. bijaria 6. Plant 45-60 cm; culrns usually 2-4-noded; leaf blades evenly distributed, flat or complicate, usually more than 25 cm long ...... 5c. var. walkeri

1. Eragrostiella leioptera (Stapq Bor, Indian For. 66: 270 (1940), et F1. 5: 106 (1940); Bor, Grasses of Burma, Ceylon, and Pakistan 495 (1960). Eragrostis leioptera Stapf in Hook. f., F1. Brit. Ind. 7: 325 (1896). Type: India, East Bengal, Khasia Hills, Hrbr. Griffith (K, CAL). Rigid perennial; culms wiry, 1-2-noded; blades 20-30 cm long, complicate (appearing terete), subpungent, somewhat sinuate, possibly pruinose when young; Zigule a ciliolate membrane c. 3 mm long; auricles bearded though glabrous with age; collar represented by a thin groove. Spike 13-21.5 cm long, interrupted in the lower part. Spikelets c. 10 X 3-4 mm, 13-19-flowered, oblong-elliptic, olive green or leaden grey. Lower glume 1.5 mm long. Upper glume 1.75-2 mm long. Lemmas 2.25-2.5 mm long, rather longer than broad, acute, cartilaginous, inflated. Palea as long as its lemma and sometimes slightly longer, with glabrous wings c. 0.3 mrn wide on the keels, often the winged keels exserted beyond the margins of the lemma. Grain 0.75-1 mm long, very plump and almost terete, with very prominent embryo. Distribution Endemic in India. Specimens examined Only the type specimens (cited above) seen. Eragrosiielk Bor 3

2. Eragrostiella collettii (Stapf) Bor, Indian For. 66: 270 (1940), et F1. Assam 5: 107 (1940); Bor, Grasses of Burma, Ceylon, India and Pakistan 494 (1960). Eragrostis collettii Stapf in Hook. f., F1. Brit. Ind. 7: 326 (1896). Syntypes: Burma: Meiktalla, Collett 19, x.1888 (K); Pinmona, Abdul Huk s.n., 20.viii. 1890 (CAL). Diagnostic characters Very slender perennial; culms extremely thin, 1-2-noded; blades up to 10 cm long, convolute-complicate, rather densely pilose on the upper surface; leaf sheaths with broad scarious margins. Spike up to 17 cm long. Spikelets 10-12 X 2-2.5 mm, 12-15-flowered, linear-oblong, purple (though glumes pallid), usually crowded. Glumes subequal, c. 2 mm long, scaberulous near apex, otherwise glabrous and smooth. Lemmas c. 2 mm long, slightly wider than long, very broadly ovate, obtuse. Palea scabrous-pubescent on the back and flaps, pallid except for purplish keels. Grain c. 0.5 mm long and almost as wide, the embryo slightly less than half as long. Distribution Endemic in Burma. Specimens examined Only the syntypes (cited above) seen.

3. Eragrostiella brachyphylla (Stapf) Bor, Indian For. 66: 270 (1940), et F1. Assam 5: 107 (1940); Bor, Grasses of Burma, Ceylon, India and Pakistan 494 (1960). Eragrostis brachyphylla Stapf in Hook. f., F1. Brit. Ind. 7: 327 (1896); Fischer in Gamble, F1. Madras 3: 1265 (1956). Syntypes: India: Hrbr. Wight, no. 1773b (K). Wallich Gzt. no. 382 7A (CAL). Diagnostic characters Perennial, extremely compact at the base; culms c. 2-noded; blades mostly 6-12.5 cm long and 1.5-3 mm wide, coriaceous, flat or complicate, obtuse, glabrous and usually glaucous on the upper surface, scabrous on the thickened margins and usually also below on the ribbed midnerve, otherwise smooth; ligule a densely ciliate membrane, 0.7-1 rnm long; auricles glabrous or bearded; leaf sheaths with broad scarious margins. Spike usually 8-18 cm long and 5-7.5 mm wide, usually crowded with spikelets. Spikelets 6-8 mm long, 1.5-2.75 mm wide, 10-1 5-flowered, linear-oblong, leaden grey to straw coloured. Glumes subequal, 1.5-2 mm long; lower ovate, acute or acuminate; upper lanceolate-ovate, obtuse. Lemmas 1.75-2.5 mm long and wide, broadly ovate-elliptic, obtuse. Palea with very narrow ciliolate wings on keels. Grain c. 0.75 X 0.5 mm, very plump and almost terete, reddish brown. Distribution Previously known only from India (Bengal, Bihar, Bombay, Madras; fide Bor, loc. cit.). Clayton's collection (cited below) represents a new record for Sri Lanka. Specimens examined INDIA: Syntypes (cited above). CEYLON: North Central Province, Polonnaruwa District, 13 miles E. of Polomaruwa on Trincomalee road, Clayton 5125 (CANB). 4 M. Lazarides 4. Eragrostiella nardoides (Trin.) Bor, Indian For. 66: 270 (1940), et F1. Assam 5: 107 (1940); Bor, Grasses of Burma, Ceylon, India and Pakistan 495 (1960). Eragrostis nardoides Trin. in Mem. Acad. Sci. Petersb. ser. 6, 1: 415 (1830); Stapf in Hook. F., F1. Brit. Ind. 7: 326 (1896); Collett, F1. Simlensis 625 (1921). Type: , Wallich Cat. no. 3827F (holotype in LE, not seen; isotype in K). Diagnostic characters Extremely slender perennial; culrns c. 2-noded; blades convolute, up to 22 cm long and reaching the lower part of the spike, rather densely pilose on the upper surface; ligule c 0.3 mm long; auricles sparsely bearded or glabrous; leaf sheaths scabrous, the hairs extending upwards on the lower surface of the blades. Spike 14-19 cm long, 2.5-5 mm wide, interrupted in the lower part, crowded with splke- lets upwards. Spikelets 8-9 X 2-2.5 nun, c 13-flowered, dark leaden grey. Glumes subequal; lower c. 1.5 mm long, acute; upper c. 2 mm long, acuniinate, scaberulous on the keel near the apex. Lemmas c. 2 mm long, longer than wide, lanceolate, acuminate, scaberulous or puberulous near the apex. Palea slightly shorter than its lemma; rhachilla glabrous, flexuose. Grain not seen. Distribution Nepal (type, cited above); India (Himalaya, Parasnath; fide Collett 1921, Bor 1960). Specimens examined Only the type (cited above) seen

5. Eragrostiella bifaria (Vahl) Bor, Indian For. 66: 270 (1940), et F1. Assam 5: 107 (1940); Senaratna, Grasses of Ceylon 77 (1956); Abeywickrama, Ceyl. J. Sci. (Bio. Sci.) 2(2): 124 (1959); Bor, Grasses of Burma, Ceylon, India and Pakistan 494 (1960); Maheshwari, F1. Delhi 383 (1963); Napper, Grasses of Tanganyika 27, fig. 74 (1965); Maheshwari, Illusts F1. Delhi, fig. 248 (1966); Santapau, F1. Khandala 310 (1967); Phillips in F1. Trop. East Afr., Gramineae part 2, 244, fig. 67 (1974). Poa buaria Vahl, Symb. Bot. 2: 19 (1791); Roxb., F1. Ind. 1: 331 (1820). Eragrostis bifaria (Vahl) Wight, Cat. Ind. PI. 105, no. 1773 (1834) nom. nud., ex Steud., Nom. Bot. ed 2, 1: 562 (1840) nom. nud., et Syn. P1. Glum. 1: 264 (1854); Stapf in Hook.f., F1. Brit. Ind. 7: 325 (1896); Cooke, F1. Bombay 2(5): 550 (1908); Fischer in Gamble, F1. Madras 3: 1205 (1956); Bogdan, Revised List of Grasses 18 (1968). Tjjpe: India, without collector (holotype in C). Eragrostis coroma~ldeliana(Koen. ex Rottl.) Trin., Mem. Acad. Sci. Petersb. ser. 6, 1: 415 (1830) (sphalm. 'coromandelina'); Steud., Syn. P1. Glum. 1: 264 (1854); Stapf in Hook. f., F1. Brit. Ind. 7: 326 (1896); Stapf in Trimen, Handb. F1. Ceyl. 5: 298 (1900). Poa corornandeliana Koen. ex Rottl., Ges Naturf. Freunde Berlin, Neue Schrift 4: 191 (1803). Type: India, 'Trankenbar nach Madras' (not seen). Eragrostielh Bor 5

5a. var. bifaria Eragrostis bifaria var. australiana F. M. Bail., Qd F1. 6: 1908 (1902), syn. nov. Type: Australia, Queensland, Cook District, Walsh River, Burton 2 (holotype in BRI). Diagnostic char(ccters Slender perennial usually 30-45 cm high; culms few-noded; blades usually 5-15 cm long (often up to 25 cm in Australian specimens), complicate-convolute (rarely flat); ligule 0.3-0.6 mm long; leaf sheaths scaberulous on the nerves or smooth. Spikes usually 10-16 cm long and 0.5-1 cm wide. Spikelets usually c. 1 cm long, up to 2 mm wide and 10-34-flowered, narrow-oblong, leaden grey or straw coloured, often curved. Glumes subequal; lower 1.7-2.7 mm long, upper 2-2.8 mm long. Lemmas 2-2.5 mm long, rather wider than long, broadly ovate, subacuie or obtuse, the apex often splitting into 2 small lobes in mature spikelets. Palea slightly shorter than its lemma, sometimes puberulous or scaberulous near the apex, with relatively wide ciliolate wings on the keels and broad flaps. Grain c. 0.5 mm long and almost as wide. A detailed description and excellent illustration of this plant is available in Phillips (loc. cit.). However, not any of the specimens examined in this study have inverse glumes as recorded by Phillips. In diagnostic features, Australian plants agree with specimens seen from tropical . Consequently, there is little justification for upholding Bailey's variety, which is hereby considered to be a taxonomic synonym. Distribution The typical variety of this species occurs over virtually the entire distribution range of the genus in India, Ceylon, Burma, eastern and north-eastern Australia. Furthermore, the species almost certainly occurs in Thailand (see collec- tion cited below, the determinstion of which requires confirmation), and probably in Indonesia. In Australia, plants are known from low altitudes, growing in sandy soils developed over granite and laterite or in inhospitable sites on granite outcrops. All the specimens seen were collected in June-July in a somewhat overmature condition, so active growth probably takes place in the summer wet season from December to March. Associated species include Eucalyptus alba Reinw. ex Bl., Chrysopogon fallax S. T. Blake and Eriachne spp. Corlservation category On present records, Australian plants of the species are known to occur sparsely in a few localities in a relatively small area in north-eastern Queensland. Consequently, in relation to the conservation categories laid down by Specht et al. (1974)*, the taxon might be allocated to category (3) and considered as a rare species requiring constant monitoring, or to category (6) and designated a species of geographical importance (with a disjunct or isolated distribution).

* Specht, R. L., Roe, E. M., and Boughton, V. H. (1974). Conservation of major plant communities in Australia and Papua New Guinea. Aust. J. Bot. Suppl. Ser. 1974, No. 7, 1-667. 6 M. Lazarides

Resurrection features On request Dr D. F. Gaff, Monash University, Victoria, kindly provided the following remarks in relation to his studies on the two collections cited below and now located in CANB. 'Plants of Eragrostiella, found in an airdry condition in Northern Queensland in June, were rehydrated in water. Tests to establish the semipermeability of the cell membranes indicated that the leaves were alive, and this was confirmed by the fact that the plants re-established active growth in the greenhouses at Monash University. The water content of the dry plants was comparable to that for the desiccation tolerant plants of Southern Africa (Gaff 1971, Desiccation-Tolerant Flowering Plants in Southern Africa, Science 174: 1033-4). Consequently Eragrostiella must also be considered a 'resurrection plant'.' Specimens examined AUSTRALIA: Queensland: Cook District: Newcastle Range, Blackman s.n., vii.1906 (BRI, K); between 'Carpentaria Downs' and Einasleigh (c. 10 miles from 'Carpentaria Downs' and 17 miles SE. from Einasleigh), Blake 19502 (BRI); Walsh Range, Burton 2 (BRI); 10.9 miles S. of 'The Lynd' on the Hughenden to Mt Garnet road, Gafy s.n. (CANB 49881, 55602); 10 miles W. of Carpentaria Downs Station, Lazarides 4679 (CANB). North Kennedy District: 11 miles NW. of Wairuna Station, Lazarides 3769 (CANB); 16 miles NW. of Wairuna Station, Lazarides 3779 (CANB). CEYLON: Uva Province: Monaragala District, near Mullagama, Lazarides 7260 (CANB, K, US, PDA), Lazarides 7264 (CANB, K, US, PDA). Northerr Province: Vavuniya District, near Kokkavil, Claytorz 5288 (CANB). North Central Province: Polonnaruwa District, Polonnaruwa, Clayton 5110 (CANB); Anuradhapura District, Galpitigala, a few miles from Ritigala, Ballard I460 (K). Southern Province: Hambantota District, Ruhuna National Park, Clayton 5950 (CANB). INDIA: Holotype (cited above) (C). Belgaum, Kitchie 851 (K). Hrbr. Rottlerianum, Gingee, xi.1732 and Wandawasi, 6.i.1800, without collectors (K); Bombay, No. 1, comm. Lisboa s.n., xii.1891 (K). Without loc., Walliclz Cat. no. 3827C (P). BURMA: Upper Burma, in 1890, Abdul Huk s.12. (K). THAILAND: Ban Na, Phumipol Dam, Tak 623 (Hrbr. No. 32694), v.1959 (BK). This specimen represents the first generic record for Thailand, but its specific determination requires confirmation. EASTERN AFRICA: Fide Phillips (1974).

5b. var. secunda (Nees ex Steud.) Iiizarides, stat. nov. Based on Eragrostis secunda Nees ex Steud., Syn. P1. Glum. 1: 264 (1854); Stapf in Hook. f., F1. Brit. Ind. 7: 326 (1896); Stapf in Trimen, Handb. F1. Ceyl. 5: 298 (1900). Eragrostiella secunda (Nees ex Steud.) Bor, Indian For. 66: 270 (1940), et F1. Assam 5: 107 (1940); Senaratna, Grasses of Ceylon 77 (1956); Abeywickrama, Ceyl. J. Sci. (Bio. Sci.) 2(2): 124 (1959); Bor, Grasses of Burma, Ceylon, India and Pakistan 495 (1960). Type: Ceylon, Sabaragamuwa Province, Galagama, Gardner s.n., ii.1846, (Thwaites C.P. 60, pro parte) (holo- type in ? P, not seen; isotypes in PDA, K, CAL). Diagnostic characters Plants of this taxon closely resemble those of the typical variety in many diagnostic characters, and differ chiefly in the dimensions, shape and colour of the spikelets. Whereas the spikelets of var. bifaria are narrow, pallid and usually very long with numerous florets, those of var. secunda are 10-14 X 2-4 mm, 12-23- flowered, broader and remain olive green in colour. Correspondingly, the lemmas are larger (c. 3 mm long), slightly longer than wide and usually acute. As these differences seem inadequate for distinction at specific level, the taxon is hereby reduced in rank. Eragrostielh Bor

Distribution This variety is endemic in Ceylon, being confined to the montane zone in altitudes above 1200 m and occurring as a component of 'patana' grasslands. Specimens examined CEYLON: Type (cited above) (PDA, K, CAL). Uva Province: Haputale, Clayton 5019 (CANB); about 35 miles south-east of Nuwara Eliya on Ettampitiya-Bandarawela road, Could 13498 (CANB); Bandarawela, Pearson 87 (K), Jowitt s.n., vi.1902 and 19.x.1904 (PDA). Central Province: Hewaheta, Kingdon-Ward 286 (K); Pallekele, Alston 276 (PDA); Pupuressa, Alston I594 (PDA).

5c. var, walkeri (Stapf) Lazarides, stat, nov. Based on Eragrostis walkeri Stapf in Trimen, Handb. F1. Ceyl. 5: 298 (1900); Fischer in Gamble, F1. Madras 3: 1265 (1956). Eragrostiella walkeri (Stapf) Bor, Indian For. 66: 270 (1940), et F1. Assam 5: 107 (1940); Seneratna, Grasses of Ceylon 78 (1956); Abeywickrama, Ceyl. J. Sci. (Bio. Sci.) 2(2): 124 (1959); Bor, Grasses of Burma, Ceylon, India and Pakistan 496 (1 960). Sabaragamuwa Province, Galagama, ii.1846 (Thwaites CP. 60, pro parte) (CAL). Diagnostic characters Undoubtedly, the relationships of this taxon are with E. bifaria and distinguishing features appear to be confined to habit. Plants of var. walkeri are rather more robust and usually 45-60 cm high, with 2-4-noded (not 1-2-noded) culms and longer blades, which are also wider, usually flat or complicate rather than convolute and filiform, and distributed throughout the plant rather than confined chiefly to the base. For these reasons, the taxon is hereby reduced in rank. Distribution Southern India, Ceylon. Specimens examined CEYLON: Syntypes (cited above) (K, CAL). North Central Province: Polonnaruwa District, 13 miles east of Polonnaruwa on Trincomalee road, Clayton 5127 (CANB). North Western Rovince: Kurunegala, Alston 705 (PDA). INDIA: Kurnool District, Cumbum (Cumbuni) Hill, Bourne 4731 (K); Krishna District, Kondapalle, Barber 4399 (K).

ACKNOWLEDGMENTS The author is very grateful to the Director and Staff of Kew for the loan of certain type and authentic material which was essential to this study. Types and other specimens cited from herbaria in Europe and Asia were seen on visits to Copenhagen (C) in 1965-66 during the author's term at Kew as Botanical Liaison Officer, to Peradeniya (PDA) and Calcutta (CAL) in July-October 1970 as a con- tributor to the Flora of Ceylon Project, and to Bangkok (BK) in March-April 1970 on the award of a South-east Asia Treaty Organization Fellowship.