Nordic Journal of Botany 31: 161–165, 2013 doi: 10.1111/j.1756-1051.2012.01207.x, © 2013 Th e Authors. Nordic Journal of Botany © 2013 Nordic Society Oikos Subject Editor: Henrik Ærenlund Pedersen. Accepted 13 March 2012

Nanooravia gen. nov., subtribe Dimeriinae (– – ) from

M. S. Kiran Raj , M. Sivadasan , J. F. Veldkamp , A. H. Alfarhan and Jacob Thomas

M. S. Kiran Raj, Dept of Botany, Univ. of Calicut,Calicut University PO, 673 635, Kerala, India. MSK also at: Dept of Botany, Sree Narayana College, Cherthala, S. N. Puram PO, 688 582, Alappuzha, Kerala, India. – M. Sivadasan (drmsivadasan@rediff mail.com) , A. H. Alfarhan and Jacob Th omas, Dept of Botany and Microbiology, College of Science, King Saud Univ., PO Box 2455, 11451 Riyadh, Kingdom of Saudi Arabia. – J. F. Veldkamp, Netherlands Centre of Biodiversity Naturalis, Leiden Univ., PO Box 9514, NL-2300 RA Leiden, the Netherlands .

Th e new Nanooravia Kiran Raj & Sivad. (Poaceae – Andropogoneae – Dimeriinae) from the southwestern Ghats in India is described and illustrated, and the new combination N. santapaui (M. R. Almeida) Kiran Raj & Sivad. is made. Th e genus is characterized by its usually unequal and intertwined racemes, triquetrous rachis, extremely oblique and glabrous pedicel tip, distantly arranged spikelets, long trigonous callus with oblique tip and densely covered with golden – yellow or yellowish– brown hairs along one angle, keel-less glumes with a dorsally echinate apex and apically auricled margins, and an upper lemma with a stout awn having a long column. Th e new genus is distinct from Dimeria R. Br. in which the species was originally described, but is similar to the monotypic Indian genus Pogonachne Bor currently placed in the subtribe Ischaeminae. It occurs in Peninsular India, a region considered as the centre of diversity of the subtribe with more than 50% of the known Dimeria species, including numerous endemics.

Dimeriinae Hack., a little known palaeotropical subtribe of distinguished from D. santapaui by having binate racemes Poaceae, belongs to the tribe Andropogoneae of the subfam- with intertwined rachises, smaller spikelets, and a paleate ily Panicoideae and was hitherto represented by the single upper fl oret. Apart from the type collections, no specimens genus Dimeria R. Br. It is considered as an enigmatic mem- of these taxa are known to have been collected. ber of the tribe (Clayton and Renvoize 1986). Dimeria is Our investigations revealed that the description of mainly distributed from India to the west Pacifi c, with D. santapaui given in the protologue does not perfectly some species occurring in North Australia and Madagascar. match the accompaning illustration. Th e stout rachis with More than 50% of its species occur in Peninsular India a single raceme and spikelets with the glumes having an api- (Bor 1960, Kiran Raj and Sivadasan 2008), the area obvi- cal winged margin are the important diagnostic features of ously considered as the centre of diversity of the subtribe D. santapaui as given in the protologue. But the “ apical and abode of several recently described species of the tribe winged margin ” of both glumes as mentioned in the descrip- Andropogoneae (Ravi et al. 1998, 2000a, 2000b, 2001a, tion of D. santapaui was not shown in the drawings of 2001b, 2004, Kiran Raj et al. 2003, Sunil and Pradeep both the glumes; instead, a ‘ keel-wing ’ , a feature which is 2005). During the post monsoon season (Nov – Dec), they characteristic of the genus Dimeria alone was shown. Th is form a prominent element of the vegetation of the Ghat main and unique characteristic feature of the glumes of regions of Peninsular India, especially along lateritic hilly D. santapaui was neither given in the table comparing the slopes (Kiran Raj 2008). characters of D. keralae and D. santapaui provided by Nair et al. (1984) or in the protologue of D. keralae . Instead, Identity of Dimeria santapaui and D. keralae and they mistakenly pointed out features like “ wingless lower recognition of a new genus glume ” in D. santapaui which actually means the absence of keel on the lower glume and “ keel-wing of the upper Dimeria santapaui was described by Almeida (1970) based glume ” which indicated the presence of distinct keel with on specimens collected by L. J. Sedgwick and T. R. D. Bell a wing. in 1919 from ‘ North Kanara ’ (Uttar Kannada Dist.) of Th e present authors have examined the type specimens Karnataka State. Later, Nair et al. (1984) described another of D. santapaui (isotype: BSI) and D. keralae (isotype: species, viz. Dimeria keralae, based on collections made MH) and found that in both species, glumes of the spike- from Paramba and Periya of the Kannur District in north- lets are dorsally keel-less with an echinate apex and have an ern Kerala. According to the protologue of D. keralae, it is apical auricled (not winged) margin. Th e study of recent

161 collections including those from the type localities of Racemes 1 or 2, usually unequal and intertwined; rachis both the species and the type specimens suggested that triquetrous; pedicel-lip extremely oblique and glabrous. D. santapaui and D. keralae are conspecifi c and showed Spikelets distantly arranged along the tough rachis, 8 – 10 that there is variation only in spikelet size which is a vari- on each raceme; callus trigonous, extremely acute at base, able character with no delimitating value. Th is study of the oblique at the tip, densely covered with golden– yellow or species further showed that it diff ered in several features yellowish– brown coloured hairs along one angle. Lower from the characteristics of the genus Dimeria. A morpho- glume sub-crustaceous, glabrous, thickened and rounded logical analysis warranted the establishment of a new genus on back with echinate apex, auricled along both sides of to accommodate it, and we propose the name Nanooravia the margin towards the tip. Upper glume subcrustaceous, for the new genus. Our analysis was based on herbarium rounded on back with echinate apex, auricled along the specimens available at CAL, BSI, MH and K (all types) margin on both sides at the apical portion. Upper lemma and several fresh collections from diff erent localities of the with a long stout awn from the sinus; column chestnut southwestern Ghats. brown coloured, bristles pale and scabrid.

Nanooravia Kiran Raj & Sivad. gen. nov. Similar taxa Dimeriae similis, pedicello longo labello obliquo, callo longo Nanooravia santapaui is distinct from the species of pilis fl avide aureis secus unum latus dense obtecto, uterque Dimeria by having a very long callus with dense golden glumis crustaceis dorsaliter rotundatis apice echinato alato et yellow hairs, keel-less glumes with a dorsally echinate apex apice margine auriculato, lemmatis superioris arista valida and apically auricled margins, an upper glume with long columna longa diff ert. hairs apically, and an upper lemma with a stout awn with a long column. Th e close resemblance of infl orescences in Type : Nanooravia santapaui (M. R. Almeida) Kiran Raj & their general morphology and the occurrence of species Sivad. comb. nov. ( Dimeria santapaui M. R. Almeida). of the two genera, viz. Dimeria and Nanooravia in the same geographical region indicate a close relationship of Etymology the two despite the diff erences in their fl oral characters. Th e new genus is named in honour of Prof. N. Ravi Th e spikelets of Nanooravia show certain morphological (Nanoo Ravi), a dedicated teacher of taxonomy and similarities with Pogonachne Bor, (Andropogoneae – subtribe a well-known agrostologist of Kerala, India. His silent, Ischaeminae), an endemic monotypic genus of the north but serious eff orts were instrumental in kindling interest in Western Ghats. A comparison of the signifi cant morpho- plant taxonomy among the young students which produced logical features of the genera is given in Table 1. good plant taxonomists. In the tribe Andropogoneae, the three genera, viz. Dimeria, Nanooravia and Pogonachne have solitary pedi- Description celled spikelets instead of the usual ‘ paired ’ (pedicelled Annuals. Culms slender; leaves mostly confi ned to the base, and sessile) spikelets characteristic for most of the gen- sheaths loosely arranged, keeled all along the back; ligule era (except for some species of Arthraxon P. Beauv. and membranous, fi mbriate at apex; leaf blades linear-subulate. Ophiuros C. F. Gaertn. where they are solitary and sessile).

Table 1. Comparison of morphological characters of Dimeria R. Br., Nanooravia Kiran Raj & Sivad. and Pogonachne Bor.

Dimeria Nanooravia Pogonachne Callus of spikelets short (0.5– 1.0 mm Callus of spikelets long (1.5– 2.0 mm), trigonous, Callus of spikelets long (ca 1.0 – 1.5 mm) long), cylindric, apex truncate, shortly narrowed at base and oblique at apex, densely cylindric, apex truncate, densely bearded or glabrous covered with golden – yellow hairs along the bearded dorsal ridge Racemes equal, usually divergent (rarely Racemes unequal, non-divergent (sometimes Racemes unequal, non-divergent with coiled or rolled at maturity) with many intertwined) with few spikelets (8 – 10) in each few spikelets (8– 10) in each raceme spikelets (15 – 50) in each raceme raceme Raceme-rachis tough, non-articulated Raceme-rachis tough, non-articulated Raceme-rachis fragile, articulated Spikelets small (2.5– 6.0 mm), laterally Spikelets small (5 – 6 mm), never laterally Spikelets large (8 – 10 mm), never compressed compressed laterally compressed Glumes laterally compressed, completely Glumes neither compressed nor keeled, dorsally Glumes neither compressed nor keeled or upper half keeled along back, rounded, dorsally sparsely hairy, echinate (except near apex), dorsally rounded, winged or wingless along the keel, and towards apex smooth towards apex smooth or hairy towards apex Glume-margins not auricled at apex Glume-margins auricled at apex Glume-margins not auricled at apex Upper glume glabrous or hairy with long Upper glume sparsely hairy with a few long hairs Upper glume glabrous, and with a hairs all along dorsal side along dorsal side at the tip only median ‘ tuft of long hairs ’ along dorsal side Awn of upper lemma 10 – 15 mm long with Awn of upper lemma 22 – 30 mm long with a Awn of upper lemma 30 – 40 mm long a column 2 – 4 1 mm column 10 – 12 2 mm with a column 12 – 15 3 mm Lower fl oret reduced with a lower palea Lower fl oret reduced without a lower palea Lower fl oret reduced with a lower palea only (sometimes absent) Upper fl oret with two stamens Upper fl oret with two stamens Upper fl oret with three stamens

162 In Nanooravia and Dimeria, the spikelets are arranged sin- Key to the genera of subtribe Dimeriinae gly along a non-articulated rachis on pedicels. Th e pedicel never detaches from the rachis. In Pogonachne , the rachis 1. Racemes usually divergent; callus short (0.5 – 1.0 mm is articulated and the spikelets are borne at each node and long); glumes keeled with or without wing, margin not the pedicel gets detached from the point of articulation auricled ……………………………………… Dimeria with the rachis after the spikelet falls off from the pedicel. – Racemes never divergent; callus long (1.5 – 2.0 mm According to Clayton and Renvoize (1986), the subtribe long); glumes keel-less and wingless, margin auricled at Dimeriinae is derived from Ischaeminae by suppression the apex ………………………………… Nanooravia of the sessile spikelet. Th e spikelets of Nanooravia santapaui (Fig. 1B) Ravia santapaui (M. R. Almeida) Kiran Raj & resemble that of Pogonachne racemosa Bor (Fig. 1A) in hav- Sivad. comb. nov . (Fig. 2) ing dorsally rounded glumes and the upper lemma with Type : ‘ North Kanara, Nirjan Flat’, 300 ft, Oct 1919, a stout awn, but diff ers in having smaller spikelets with a L. J. Sedgwick and T. R. D. Bell 6875 (holotype: BLAT, long callus, absence of a median tuft of hairs along the not found; isotypes: BSI!, K, cibachrome photograph!). dorsal side of the upper glume, but with apical hairs. In Dimeria, the spikelets are laterally compressed and have Taxonomic synonym: Dimeria keralae N. C. Nair, Sreek. & dorsally keeled glumes. In all species of Dimeria and V. J. Nair (1984, p. 626). Nanooravia santapaui the spikelets are borne on a tough, Type : India, Kerala State, Kannur Dist.: Paramba, 150 m non-articulated rachis. Th e spikelets of the species in a.s.l., 16 Nov 1981, P. V. Sreekumar 71717 (holotype: Dimeria sect. Capillares , e.g. D. hohenackeri Hochst. ex CAL!; isotypes: MH!, K, cibachrome photograph!). Miq. (Fig. 1C) diff er from those of sect. Loriformes , e.g. D. kurumthotticalana Jacob (Fig. 1D) by having the upper Description glumes dorsally keeled above the middle. In Nanooravia up to 30 cm tall. Culms slender, erect or some- santapaui the upper glumes are dorsally rounded through- times bent at a right angle to the axis at the extreme apical out, but not keeled. However, in D. hohenackeri the node; nodes bearded. Leaves mostly confi ned to the base; upper glumes are dorsally rounded only at the basal half sheaths 3 – 5 cm long, shorter than the internodes, loosely and keeled at the upper half. In D. kurumthotticalana arranged, keeled all along the back, glabrous; blades linear- they are not dorsally rounded, but keeled throughout the subulate, 2– 5 cm 1 – 2 mm, keeled on the midrib below, length. Th e spikelet morphology of Pogonachne, Nanooravia continuous with that of the sheath, acuminate at apex, and Dimeria is schematically shown in Fig. 1A – D. glabrous or margins with a few tubercle-based hairs. With the recognition of Nanooravia gen. nov., the subtribe Ligule up to 1 mm long, ovate, fi mbriate at apex. Racemes Dimeriinae now comprises two genera, viz. Dimeria and usually unequal, sometimes intertwined, 2– 5 cm long; Nanooravia and they can be identifi ed as in the below key. rachis triquetrous, ca 0.25 mm broad, glabrous; pedicels 1.0 – 1.5 mm long, oblong, with extremely oblique lip, gla- brous. Spikelets linear oblong-oblanceolate, 5 – 6 mm long, cuneate at base, distantly arranged along the rachis. Callus 1.5 – 2.0 mm long, extremely acute at base, oblique at apex, covered with golden-yellow or yellowish – brown coloured hairs along one angle, hairs up to 1.5 mm long. Lower glume subcrustaceous, more or less linear-oblong, 4.0 – 4.5 mm long, glabrous, thickened and rounded on the back, echinate at apex, with margin coriaceous, distally auricled on both sides towards the tip. Upper glume sub- crustaceous, oblong-lanceolate, 5.0 – 5.5 mm long, glabrous, echinate at apex, with auricled margins and a few long hairs at the tip. Lower fl oret empty; lemma hyaline, oblan- ceolate, 3 – 4 mm long, 1-nerved, sparsely ciliate along the upper two-thirds of the margin; palea absent. Upper fl oret bisexual; lemma hyaline, oblanceolate, subcoria- ceous, 1-nerved, inconspicuous in the upper half, 3 – 4 mm long, apex bifi d with acute lobes; awn stout from the Figure 1. Spikelet morphology of Nanooravia Kiran Raj & Sivad. sinus, simple, scaberulous, 22 – 30 mm long, geniculate; and allied taxa. (A) spikelet of Pogonachne racemosa Bor, (B) spikelet column 10 – 12 mm long, twisted, chestnut brown, with of Nanooravia santapaui (M. R. Almeida) Kiran Raj & Sivad., (B1) pale bristles; palea minute, ca 0.5 mm long, hyaline, ovate- cross section of the upper glume of P. racemosa & N. santapaui lanceolate, without nerves, shortly ciliate along margins. (schematic), (C) spikelet of Dimeria hohenackeri Hochst. ex Miq. Lodicules 2, ca 0.4 0.2 mm, 2- or 3-lobed at apex. ( Dimeria sect. Cappillares ), (C1) cross section of the upper glume Stamens 2; anthers 1.0– 1.5 mm long, pale yellow to red- of D. hohenackeri showing keel (schematic), (D) spikelet of D. kurumthotticalana Jacob (Dimeria sect. Loriformes ), (D1) cross dish. Stigma pinkish. Grain linear-oblong, 2.0 – 2.5 section of the upper glume of D. kurumthotticalana showing 0.4 mm, subcompressed, acute – apiculate at apex. Flower- keel-wing (schematic). Scale bar 1 mm. ing and fruiting occurs between Sep and Dec.

163 Figure 2. Nanooravia santapaui comb. nov. (A) habit, (B) spikelet, (C) a portion of rachis with a pedicel, (D) callus, (E) lower glume, lateral view, (E1) apical portion of lower glume, dorsal view, (F) upper glume, lateral view, (F1) apical portion of upper glume, dorsal view, (G) lower lemma, (H) upper lemma, dorsal view, (H1) upper lemma, lateral view, (I) palea, (J) lodicules, stamens and pistil. Drawing by V. B. Sajeev after M. S. Kiran Raj CU 92838 (CALI).

164 Distribution for the award of a senior research fellowship, and to the Endemic to south Western Ghats; northern region of ‘ International Association for Plant Taxonomy ’ (IAPT), Vienna Kerala, especially in the Kannur and Kasaragode Districts for the award of a plant systematics research grant in 2007. Th e and the Dakshin Kannada and Uttar Kannada Districts of second, fourth and fi fth authors acknowledge the support from the ‘ National Plan for Science, Technology and Innovation’ Karnataka. (NPST), King Abdulaziz City for Science and Technology, Kingdom of Saudi Arabia through the research project Habitat and ecology no. 11-ENV1753-02. More or less sub-gregarious during the post monsoon season; usually found in dry lateritic rocky slopes, some- times on wet laterite hillocks along with other endemic References grasses like Bhidea burnsiana Bor, Danthonidium gammiei (Bhide) C. E. Hubb., Dimeria bialata C. E. C. Fisch., Almeida, M. R. 1970. Th ree new grasses from the former Bombay Glyphochloa acuminata (Hack.) Clayton and Arundinella Presidency. – J. Bombay Nat. Hist. Soc. 66: 510 – 513. spp. at 50 – 120 m a.s.l. Bor, N. L. 1960. Th e grasses of Burma, Ceylon, India and Pakistan (excluding Bambuseae). – Pergamon Press. Clayton, W. D. and Renvoize, S. A. 1986. Genera Graminum. Notes Grass genera of the world. – Kew. Bull. Add. Ser. 12, R. Bot. Th e isotype of N. santapaui at Kew is bearing a label Gard. ‘ ‘ Dimeria kanarensis ’ ’ written by M. R. Almeida in 29 Jul Kiran Raj, M. S. 2008. Taxonomic revision of the subtribe 1968. Th is specimen was later designated as the isotype of Dimeriinae Hack. of Andropogoneae (Poaceae– Panicoideae) D. santapaui by Almeida (1970). in Peninsular India. – PhD thesis, Univ. of Calcutta. Kiran Raj, M. S. and Sivadasan, M. 2008. A new species of Dimeria R. Br. (Poaceae, Panicoideae, Andropogoneae) from Additional specimens examined Goa, India. – Novon 18: 183 – 186. India: Kerala State: Kannur Dist., Paramba, 150 m a.s.l., Kiran Raj, M. S. et al. 2003. Grass diversity of Kerala – endemism 17 Nov 1981, P. V. Sreekumar 71727 (MH); ibid, 30 Oct and its phytogeographical signifi cance. – In: Janardhanam, 1999, Ravi TBGT 41566 (TBGT, CALI); ibid, Periya, M. K. and Narasimhan, D. (eds), Plant diversity, human 50 m a.s.l., 18 Nov 1981. P. V. Sreekumar 71755 (MH); welfare and conservation. Goa Univ., pp. 8 – 30. McNeill, J. et al. (eds) 2006. International code of botanical ibid, 18 Nov 1999, N. Ravi and M. S. Kiran Raj TBGT nomenclature, Vienna code. Adopted by the 17th Int. Bot. 41541, 41542 (TBGT, CALI); Kasaragode Dist., on the Congr. Vienna, Austria, Jul 2005. Reg. Veg. Vol. 146. – ARG way to Bandaduka, 110 m a.s.l., 6 Oct 2002, M. S. Kiran Gantner Verlag. Raj CU 92838 (CALI); Karnataka State: Dakshin Kannada Nair, N. C. et al. 1984. Dimeria keralae (Poaceae) – a novelty Dist., Mangalore, 100 m a.s.l., 18 Dec 2001, M. S. Kiran from Kerala, India. – J. Bombay Nat. Hist. Soc. 80: 626 –629. Raj CU 81019 (CALI); Uttar Kannada Dist., Mirjan, Nees von Esenbeck, C. G. D. and Martius, C. F. P. S. 1823. Rauia . 120 m a.s.l., 7 Oct 2003, M. S. Kiran Raj CU 92989 – Nova Acta Phys. Med. Acad. Caes. Leop. Carol. Nat. Cur. 11: 151, 167. (CALI); ‘ north Kanara, Mirjan Flat’, 300 ft, Oct 1919, Ravi, N. et al. 1998. Th ree new species of Ischaemum L. (Poaceae) L. J. Sedgwick and T. R. D. Bell 6875 (BSI). from Kerala, India. – Rheedea 8: 149 – 158. Ravi, N. et al. 2000a. Another new species of Ischaemum from Kerala, India. – Rheedea 10: 49 – 53. Acknowledgements – Th e authors are grateful to Dr M. Sanjappa, Ravi, N. et al. 2000b. Two new species of Poaceae from Kerala, former director, Botanical Survey of India, to the joint directors India. – Rheedea 10: 91 – 98. and curators of BLAT, BSI, CAL, K, MH, and TBGT for permit- Ravi, N. et al. 2001a. Th ree new species of Poaceae from south ting to examine the specimens for the present study. Th anks also India. – Rheedea 11: 87 – 96. to Dr V. J. Nair, retired joint director, ‘ Southern Circle of Ravi, N. et al. 2001b. Th ree more new species of Ischaemum L. the Botanical Survey of India’ and emeritus scientist at MH, (Poaceae) from Kerala, India. – Bot. Bull. Acad. Sin. 42: Dr P. Lakshminarasimhan, former Indian liaison offi cer at K 223 – 230. and currently scientist at CAL, Mr Gireesh G. Potdar, Dept of Ravi, N. et al. 2004. Melinis minutifl ora P. Beauv. (Poaceae), an Botany, Shivaji Univ., Maharashtra, and Prof. P. Manimohan, Dept interesting new record for Peninsular India. – J. Econ. Taxon. of Botany, Univ. of Calicut for various help. Th e drawings were Bot. 28: 81 – 83. made by Mr V. B. Sajeev, Ernakulam and he is thankfully remem- Sunil, C. N. and Pradeep, A. K. 2005. Ischaemum glabriglaucum bered. Th e fi rst author expresses his sincere gratitude towards the Sunil (Poaceae), a new species from India. – Candollea 60: ‘ Council of Scientifi c and Industrial Research’ (CSIR), New Delhi 387 – 391.

165