Helictotrichon devesae, a new endemic grass species from Castilla-La Mancha (Central Spain)

Helic o ichon de esae, a new endemic g ass species om Cas illa-La Mancha (Cen al Spain) by Ca los Rome o Za co Depa amen o de Biología Vege al y Ecología, Uni e sidad de Se illa. Apa ado 1095, 41080-Se illa, Spain. [email p o ec ed] Abs ac Rome o Za co, C. 2007. Helic o ichon de esae, a new endemic g ass species om Cas illa-La Mancha (cen al Spain). Anales Ja d. Bo . Mad id 64(2): 205-211. Helic o ichon de esae Rome o Za co (A eneae; Poaceae), a new g ass species is desc ibed and i s ana omical and mo pho- logical cha ac e s a e compa ed in ela ion o o he species in he genus. Keywo ds: Helic o ichon, A eneae, G amineae, axonomy, lea ana omy, Spain. Resumen Rome o Za co, C. 2007. Helic o ichon de esae, una nue a es- pecie de g amínea endémica de Cas illa-La Mancha (cen o de España). Anales Ja d. Bo . Mad id 64(2): 205-211 (en inglés). Se desc ibe Helic o ichon de esae Rome o Za co (A eneae; Poaceae), una nue a especie de g amínea y se compa an sus ca- ac e es ana ómicos y mo ológicos con los de o as especies del géne o. Palab as cla e: Helic o ichon, A eneae, G amineae, axono- mía, ana omía olia , España. In oduc ion The genus Helic o ichon Besse (A eneae; Poa- ceae), sensu s ic o (Helic o ichon subgen. Helic o i- chon sensu Röse , 1989), was he subjec o se e al axonomic su eys in he las cen u y (Ge ais, 1973; Rome o Za co, 1984, 1985a; Röse , 1989, 1996). Those s udies p o ided a la ge da a se o Wes e n Eu opean and No h A ican axa. A p esen , up o 7 Helic o ichon axa, belonging o 3 (Rome o Za co, 1984) o 4 (Röse , 1989) species, ha e been iden i ied o he Ibe ian Peninsula. The Medi e anean egion is he likely scena io o ecen specia ion e en s in ol ing ances o s o he p e-Medi e anean lo a, p obably simila o now ex- is ing species in Sou h-eas e n Asia o opical Eas A ica (Röse , 1996: 256). The genus’ geog aphic ange in Spain is condi ioned by he o ophilic o somewha s eppe cha ac e and mainly basiphilic a ini y o i s species. Two isola ed a eas a e known o include Helic o i- chon species: one in he No h and ano he in he Sou h-Eas (Fig. 1). The o me comp ises a small po - ion o he Ibe ian Sys em, Cen al and Wes e n Py e- nees and he Can ab ian Range, whe e he endemic H. can ab icum (Lag.) Ge ais g ows in a g ea a ie y o plan communi ies om sea le el o 2200 m. The meso-xe ophilic Alpine-A lasic H. sedenense (DC.) J. Holub occu s in alpine meadows, subalpine and o o- medi e anean dwa -sh ub communi ies, and open subalpine coni e ous woodlands (Röse , 1996). In he second a ea, sou h-eas e n Ibe ia, H. sede- nense g ows in open hemic yp ophy ic o ma ions a ec ed by c yo u ba ion (c yo omedi e anean) and a lowe al i udes in xe ophilic spiny sh ubs (o o- medi e anean), om 2000 o 3300 m abo e sea le el. The H. ili olium (Lag.) Hen a d complex occu s in mos o SE Spain, om he Sie a de G azalema (Cádiz p o ince, S Spain) o Sego be (Cas ellón p o ince, E Spain), including Sie a de Cazo la and he Júca alley. In his in e es ing complex, low ploidy le els (2x, 4x, 8x) a e ound in he cen al pa o he Be ic Ranges (S Spain: Sie a de Ru e, Sie a Tejeda, sou h o Jaén p o ince and se e al places in G anada p o ince, specially in Sie a Ne ada). In a p e ious pape Rome o Za co (1984) iden i ied hese popula ions as H. ili olium subsp. elu inum (Boiss.) Rome o Za co. Howe e , Röse (1989, 1996, 2006), Anales del Ja dín Bo ánico de Mad id Vol. 64(2): 205-211 julio-diciemb e 2007 ISSN: 0211-1322 conside ed hese plan s o be e y well delimi ed om he es o he complex and wo h o speci ic ecogni- ion as H. sa aceno um (Gand.) J. Holub. Awai ing o u u e phylogene ic e idences, we accep Röse ’s p oposal, mainly because i seems mo e consis en wi h he axonomic ea men o he genus. I has been sugges ed ha plan s wi h 2n = 70 (10x) om he wes e n Be ic anges (coexis ing wi h Abies pin- sapo Boiss.) co espond o H. ili olium subsp. a un- danum Rome o Za co, a axon accep ed by Röse in his axonomic (Röse , 1989) and ecogeog aphic (Rös- e , 1996) s udies. The mo e ex ended H. ili olium subsp. ili olium includes wo a ie ies: a . cazo lense Rome o Za co (2n = 98) dis ibu ed in Sie a de Ca- zo la, Sie a del Pozo and pe haps in o he cen al Be ic moun ains, and a . ili olium (2n = 84) some- wha smalle and wi h ela i ely na owe lea es, being he single Helic o ichon ep esen a i e ocu- C. Rome o Za co206 Anales del Ja dín Bo ánico de Mad id 64(2): 205-211, julio-diciemb e 2007. ISSN: 0211-1322 ing in Cen al Spain (Albace e and Ciudad Real p o inces, and Se anía de Cuenca). In 1989, we ound a he ba ium specimen labelled “Helic o ichon ili olium” (MA 447900). The plan was collec ed in 1976 nea Puebla de Almo adiel, in Toledo p o ince, 70 Km away om he nea es locali- ies known o ha species. I was immedia ely e i- den ha i s small spikele s, ex emely na ow lea es and pale basal shea hs did no ma ch well wi h he cha ac e s o any known species o he genus Helic- o ichon. The p esence o hai s on some lea es, a yp- ical cha ac e o young indi iduals o he Helic o i- chon ili olium complex (Rome o Za co, 1984), led o he assump ion ha i was H. sa aceno um, bu he locali y o o igin was ou o he expec ed dis ibu ion ange. Consequen ly, i was se aside as “s udy ma e - ial” pending D . Röse ’s e iew o he genus in 1989. Ne e heless, his ample and excellen e iew did no Fig. 1. Dis ibu ion ange o Spanish Helic o ichon species: 1, ype locali y o H. de esae; 2, H. can ab icum; 3, H. ili olium; 4, H. sa aceno um; 5, H. sedenense. 2-5 acco ding o Rome o Za co (1984) and Röse (1989). 3 including locali ies om Ma ín-Blanco & Ca asco (2005). 1 2 3 4 5 p o ide a sa is ac o y axonomic posi ion o his species. Ma e ial and me hods He ba ium ma e ial om he Uni e si ies o Se ille (SEV) and Ba celona ( o me ly BCF, cu en - ly included in BCN) we e used o illus a e lea ana o- my. Addi ional ma e ial om he Royal Bo anic Ga - den o Mad id has been used o desc ibed he new species. Da a p e iously published by he au ho ha e been included in his s udy (Rome o Za co, 1984, 1985a). C oss-sec ions o lea blades we e made by hand using well de eloped, basal lea es om inno a ions shoo s. Pollen g ains om un- opened an he s whe e obse ed by op ical mi- c oscopy in iso onic wa e solu ion. Resul s Lea ana omy In gene al lea blades o he new axon show a Helic o ichon de esae, a new endemic g ass species om Cas illa-La Mancha s uc u e simila o hose om o he xe ophilic species in he genus. The new axon p esen s sho e and na owe lea blades han o he close species, ac- companied by an o e all educ ion in ascula issue. I p esen s 7 ascula bundles, ins ead o he 9 o 13 which a e common in he es o he species, and he gi de s a e absen o e y educed, a cha ac e ye un- epo ed o his genus (Fig. 2). Pollen g ains Pollen g ains we e obse ed and measu ed. The appea ance was no mal, and he size anged om 29.7 o 33.8 µm. Diagnosis Helic o ichon de esae Rome o Za co, sp. no . Ab Helic o icho ili olio (Lag.) Hen a d p aecipue di e s a u a mino e, aginis s amineis nec ub o inc- is; laminis quidem summe ili o mibus (0,5-0,6 mm diame o), sec ione ans e sali 7-ne iis a que abe- 207 Anales del Ja dín Bo ánico de Mad id 64(2): 205-211, julio-diciemb e 2007. ISSN: 0211-1322 Fig. 2. Idealized c oss-sec ions o lea blades om inno a ion shoo s in 5 axa o Helic o ichon (scle enchyma in black): a, H. sede- nense subsp. sedenense (España, Lé ida, BCF 1481); b, H. sa aceno um (España, Málaga, SEV 86653); c, H. ili olium a . ili olium (Es- paña, Alican e, SEV 86669); d, H. can ab icum (España, Bu gos, SEV 77612); e, H. de esae (España, Toledo, SEV 217062). ab cde 0,5 mm culis des i u is; denique, spiculis mino ibus (c. 10 mm longis). Ab Helic o icho sa aceno um (Gand.) J. Holub p aecipue di e oliis ce e angus issimis, gla- b is el iu enilibus pa ce a que b e i e pilosis (pilis c. 0,1 mm). Typus: in p o incia ole ana Hispaniae, ad i u- lum Cigüela dic um, p ope oppidum La Puebla de Almo adiel, in salsuginosis pascuis appella is a quibusdam «Senecio au iculae-Lygee um e Schoeno nig ican is-Plan agine um ma i imae», legi S. Ci u- jano mense Julio anni 1976; in he ba io Uni e si- a is Hispalensis asse a u sub nume is SEV 217062 (holo ypus) e SEV 217063 (iso ypus) a que in he ba io Ho i Regii Ma i ensis, insupe , sub nu- me o MA 447900 (iso ypus). U pa e , species cla is- simo co dubensi bo anico J.A. De esa obsequen is- sime dica a. Fig. 3. Desc ip ion Pe ennial he b, caespi ose; non- lowe ing shoo s mainly in a aginal; culms 27-35 cm, glab ous; basal lea shea hs subau icula e, s aw-colou ed; ligules 0.5-1 mm, unca e, sho ly cilia e; lea blades 5-16 cm × 0.5-0.6 mm, se aceous, ibbed and scab id abo e, glab ous o sho ly pubescen benea h when young (hai s 0.1-0.2 mm), wi h 7 eins and a con inuous subepide mal laye o scle- enchyma in c oss-sec ion. Panicle 7-12 cm, o a e, lax, wi h 15-25 spikele s and sho ly pubescen b anches; peduncles 3-11 mm, cla a e, sho ly pu- bescen . Spikele s 9.6-10.3 mm, h ee- o ou - low- e ed, he uppe s e ile; hachilla 1.8-2.5 mm, disa - icula ing below he lo e , wi h hai s up o 3 mm; glumes unequal, he lowe 6-6.7 × c. 1,5 mm, 1- ne ed, he uppe 9-9.8 × c. 2.1 mm, 3-ne ed; cal- lus o a e-ellip ic, hi su e wi h hai s o 0.5-2 mm; lemma o he lowe lo e 8-9 mm, lanceola e, 7- ne ed, 2-den a e in he apex, glab ous; awn 10.5- 13.5 mm, a ising om he middle o he back, genicula e, wi h e e e, sho ly pubescen column and scab id b is le; lodicules c. 1.5 mm, na owly lanceola e, en i e; an he s 4-4.5 mm; o a y hai y; ca yopsis no seen. Flowe ing ime in July. Habi a : s eppe-subsaline g assland communi y o Lygeum spa um, c. 700 m abo e sea le el. The name is dedica ed o D . J.A. De esa Al- ca az, au ho o in e es ing axonomic s udies on Spanish g asses and composi es. The new species is known only om he ype locali y in Cen al Spain, whe e i has p obably no been collec ed since 1976. C. Rome o Za co IDENTIFICATION KEY FOR EUROPEAN SPECIES OF HELICTOTRI- CHON (SPECIES CIRCUMSCRIPTION ACCORDING TO RÖSER, 1996 PRO SUBGEN. HELICTOTRICHON;INFRASPECIFIC TAXA NOT ANALYZED) 1. Spikele s disa icula ing only abo e he glumes ............... 2 1. Spikele s disa icula ing abo e he glumes and below he lo e s ............................................................................ 4 2. Ligules 3-6 mm, oblong; blades pe sis en when dead ....... .................................................................... H. pa la o ei 2. Ligules 0.2-1.5 mm, unca e; blades soon deciduous when dead .............................................................................. 3 3. Basal lea es se aceous, 0.5-0.8 mm in diame e ................ .................................................................... H. se aceum 3. Basal lea es con olu e, 0.9-1.5 mm in diame e o la and 2-4 mm wide ........................................ H. sempe i ens 4. Basal lea es wi hou a con inuos subepide mal laye o scle- enchyma; non- lowe ing shoo s ex a aginal ................. 5 4. Basal lea es wi h a con inuous subepide mal laye o scle- enchyma; non- lowe ing shoo s mos ly in a aginal ...... 6 5. Lea es g eyish-g een abo e, da k g een benea h; pedicels sca cely cla a e; basal shea hs open o he base ................ ..................................................................... H. deco um 5. Lea es g eyish-g een on bo h su aces; pedicels clea ly cla a e; basal shea hs open only in he uppe hal ............. .................................................................. H. sedenense 6. Ligules o basal lea es 3-8 mm, lanceola e, acu e; basal lea es se aceous, up o 0.7 mm wide ....... H. dese o um 6. Ligules o basal lea es up o 1.5 mm, wide han longe , unca e; lea es se aceous o no , up o 4 mm wide when la ................................................................................. 7 7. Lowe glume 6-6.7 mm; spikele s c. 10 mm; basal lea es 0.5-0.6 mm in diame e ................................. H. de esae 7. Lowe glume 8.5-17 mm; spikele s 10-20 mm; basal lea es (0.5) 0.7-2 mm in diame e , up o 4 mm wide when la .... ....................................................................................... 8 8. Basal lea es densely pubescen benea h (hai s 0.5-1 mm), a leas when young; basal shea hs g eyish, pale b own o s aw-colou ed ..................................... H. sa aceno um 8. Basal lea es usually glab ous benea h, some imes he younge lea es pubescen in plan s wi h eddish-b own basal shea hs .................................................................. 9 9. Shea hs o basal lea es da k iole -b own o eddish-b own ..................................................................... H. ili olium 9. Shea hs o basal lea es g eyish o b ownish .................. 10 10. Hai s o hachilla up o 3.5 mm; lemma 9-11 mm .............. ................................................................ H. con olu um 10. Hai s o hachilla 4.5-5 mm; lemma 10-14 mm .................. ............................................................... H. can ab icum Discussion Lea blade ana omy in c oss-sec ion o he new species is e y simila o hose o o he g asses adap - ed o xe ic en i onmen s. Thus, he di e ences in lea ana omy be ween he meso-xe ophilic H. sedenense and he ex eme xe ophilic H. de esae a e pa allel o hose obse ed in o he g ass gene a, like he closely ela ed Pseuda hena he um. The Eu osibe ian Pseuda hena he um longi olium (Tho e) Rouy p e- sen s wide, so blades whe eas P. pallens (Link) J. 208 Anales del Ja dín Bo ánico de Mad id 64(2): 205-211, julio-diciemb e 2007. ISSN: 0211-1322 Helic o ichon de esae, a new endemic g ass species om Cas illa-La Mancha 209 Anales del Ja dín Bo ánico de Mad id 64(2): 205-211, julio-diciemb e 2007. ISSN: 0211-1322 Fig. 3. Helic o ichon de esae (España, Toledo, La Puebla de Almo adiel, SEV 217062): a, habi ; b, ligule o basal lea ; c, spikele ; d, dispe sal uni in la e al iew; e, lemma; , palea; g, lodicules and e ile pa s o lo e . Holub, an endemic species om cen al-wes e n Po - ugal limes one a eas, esembles ou new species in lea ana omy (Rome o Za co, 1985b). The e olu ion om meso-xe omo phic o s ic ly xe omo phic species in Medi e anean pe ennial oa s has also been documen ed by Röse (1989, 1996, 1998). Two o he non-Spanish Helic o ichon species ha e simila lea blade ana omy han ha o H. de esae: H. se aceum (Vill.) Hen a d, a Wes -Alpine species belonging o he H. pa la o ei g oup, and H. dese o- um (Lees.) Ne ski, om he s eppes and ocky hills o Eas e n Eu ope and Wes e n Asia (see Sain -Y es, 1931, sub A enas um). The o me di e s om ou species mainly in i s sinap ospe mic spikele s, whe e- as he la e di e s in i s elonga ed ligules. The spikele s o H. de esae show a simila mo - phology and s uc u e o hose o o he Spanish species (Table 1): he hachilla disa icula es below each lo e , so he dispe sal uni is a single ma u e lo- e . By con as , in he Alps-endemic H. pa la o ei g oup (Röse , 1989: 65), he spikele s disa icula e only abo e he glumes, and he dispe sal uni s a e sinap ospe mic. This cha ac e (p obably de i ed) occu s in o he A enea gene a, e.g. A ena s e ilis g oup, Pseuda hena he um and Holcus. Rega ding spikele size, H. de esae esembles H. sedenense and small-spikele indi iduals o H. sa a- ceno um (Table 1). I di e s om bo h, howe e , in i s smalle lea -blade diame e and scle enchyma pa - e n. The pe sis ing lea pubescence obse ed in young indi iduals o H. de esae lea es usually occu s in H. sa aceno um, bu in he o me he hai s a e sho e (0.1-0.2 mm) and sca e ed. Se aceous lea es a e p esen in bo h H. de esae and H. ili olium subsp. ili olium howe e , in H. ili olium a . ili o- lium om SE Spain (0,5-1 mm in diame e ), well-de- ined scle enchyma gi de s a e isible in c oss-sec- ion. Simila di e ences sepa a e H. de ease om H. can ab icum, hough he angula shape o he ibs is common in bo h species. Di e ences in lea ana o- C. Rome o Za co my ha e been shown o pe sis in cul i a ed plan s o Helic o ichon axa (Ge ais, 1973; Röse , 1989). The species mos closely ela ed o H. de esae is H. con olu um (C. P esl) Hen a d, a diploid, xe- ophilic, cen al o eas e n Medi e anean species p oposed by Röse (1996: 237) as a geog aphical ic- a ian o he H. ili olium-sa aceno um complex. The c oss-sec ion lea blade o H. de esae is e y simila o hose o H. can ab icum and H. con olu um (see Ge - ais, 1973, and Fig. 2), indica ing a close a ini y o he o me axon o he la e axa, p obably o igina ed in he Eu osibe ian pa o he Medi e anean basin, whe eas he H. ili olium-sa aceno um complex p obably o igina ed in he Be ic-No h A ican a ea. I is highly unlikely ha he new species could be a hyb id as he only possible ex an pa en al species in he egion is H. ili olium, and in e gene ic hyb ids do no occu be ween his species and o he congene s o non-congene s (Ge ais, 1983). Fu he mo e, he pollen g ains show no mal shape and hei size alls wi hin he known ange o o he Helic o ichon spp. (e.g. H. sa aceno um, 30-38 µm; Rome o Za co, 1984, sub H. ili olium subsp. elu inum). H. de esae g ows in alba dina plan communi ies (Senecio au iculae-Lygee um Ri as Goday & Ri as Ma ínez in Ri as Ma ínez & Cos a 1976). I is a g ass o ma ion on subsaline soils exposed o se e e hyd ic s ess du ing summe . No o he species in he genus is known o g ow in subsaline habi a s. The only axon p esen in s eppe-like, highly a id en i onmen s is H. ili olium a . ili olium which g ows in S ipa enacissima o ma ions on basic subs a e in SE Spain (S ipo-Helic o iche um ili oliae G. López 1973) and on di e se calca eous subs a es in he Se anía de Cuenca moun ain ange (López, 1976). Mo e p ecise da a abou he ype-locali y o H. de- esae we e p o ided by D . S. Ci ujano (in li .). The new Helic o ichon species was collec ed in he ansi- ion be ween Senecio au icula-Lygee um and Schoeno- Plan agine um ma i imae Ri as-Ma ínez, 1984 phy- 210 Anales del Ja dín Bo ánico de Mad id 64(2): 205-211, julio-diciemb e 2007. ISSN: 0211-1322 Species Spikele Lowe glume Uppe glume Rhachilla Lemma Awn (1 h segmen ) (1 h lo e ) H. sedenense 7.5-14 6-10 (7.5) 8-12 2-2.5 (7.5) 8-10 (11) 12-20 H. de esae 9.6-10.3 6-6.7 9-9.8 1.8-2.5 8-9 10.5-13 H. con olu um 11-17 9-12 10-14 c. 3 9-11 ? H. can ab icum (10) 11-17 8.5-13 10-15 3-4 10-14 15-20 H. sa aceno um 10-15 8.5-11 11-13 2-2.5 9-12 15-22 H. ili olium 14-20 9-17 11-20 2.5-4 9-17 18-30 Table 1. Size ange o spikele s and lo al elemen s (in mm) in 5 Helic o ichon species p esen in Spain and in H. con olu um. Acco ding o Holub (1980), Rome o Za co (1984) and Röse (1989). ocenoses ( he plan communi ies included, among o he s, Schoenus nig icans, Plan ago ma i ima, Do ic- nium g acile, Linum ma i imum, Ag os is s oloni e a and Cynodon dac ylon). These da a sugges ha H. de esae could be a new and no ewo hy example o g ass specia ion ela ed o ecogeog aphic adap a ions. The o iginal popula ion (leco ype) o H. de esae, was des oyed by ag icul u al p ac ices some yea s ago (D . Ci ujano, in li .). Though he plan was seen (bu no collec ed) a o he Schoenus nig icans si es in he La Mancha a ea. Fu he sea ch on popula ions o H. de esae om Cen al Spain (Ri e Guadiana Valley and he su ounding moun ains) a e needed o es ab- lish he ecology, dis ibu ion and ploidy le el o his species. Acknowledgemen s This pape is de o ed o he memo y o he la e D . Camille Ge - ais (Mon eal, 1933, Québec, 2002), who guided my i s s eps h ough he g oup's axonomy wi h his ine s udy on pe ennial oa s. Thanks a e gi en o D . Ma in Röse (Halle, Ge many) who sen me many use ul publica ions. The He ba ium o he Royal Bo anic Ga den o Mad id (Spain) p o ided he ma e ial used o desc ibe he new species. D . S. Ci ujano (Mad id) kindly o e ed in o ma- ion abou he new species collec ion si es. Pollen g ain obse a- ions we e made by D . Regina Be jano om he Uni e si y o Se ille (Spain). La in diagnosis was kindly e ised by M. Laínz. Re e ences Ge ais, C. 1973. Con ibu ion à l’é ude cy ologique e ax- onomique des a oines i aces. Densch i en de Schweiz- e ischen Na u o schenden Gesellscha 88: 1-166; 1-56 (Figu es e Ca es). Ge ais, C. 1983. Wide hyb idiza ion a emp s in he ibe A e- neae Nees. Bo anica Hel e ica 93: 195-212. Helic o ichon de esae, a new endemic g ass species om Cas illa-La Mancha Holub, J. 1980. Helic o ichon Besse . In: Tu in, T.G. & al. (eds.), Flo a Eu opaea 5: 208-210. Camb idge. López, G. 1976. Con ibución al conocimien o i osociológico de la se anía de Cuenca, I. Anales del Ja dín Bo ánico de Mad id 33: 5-87. Ma ín-Blanco, C.J. & Ca asco, M.A. 2005. Ca álogo de la lo a ascula de la p o incia de Ciudad Real. Monog a ías de la Aso- ciación de He ba ios Ibe o-Maca onésicos ol. 1: 581 pp. Mad id. Rome o Za co, C. 1984. Re isión axonómica del géne o Helic- o ichon Bess. ex Schul es & Schul es il. (G amineae) en la Península Ibé ica. I. Es udio axonómico. Anales del Ja dín Bo ánico de Mad id 41: 97-124. Rome o Za co, C. 1985a. Re isión axonómica del géne o Helic- o ichon Bess. ex Schul es & Schul es il. (G amineae) en la Península Ibé ica. II. Es udios expe imen ales. Anales del Ja dín Bo ánico de Mad id 42: 133-154. Rome o Za co, C. 1985b. Es udio axonómico del géne o Pseuda- hena he um Rouy en la Península Ibé ica. Lagascalia 13(2): 255-273. Röse , M. 1989. Ka yologische, sys ema ische und cho ologische Un e suchungen an de Ga ung Helic o ichon Besse ex Schul es & Schul es (Poaceae) im wes lichen Mi elmee gebie . Disse a iones Bo anicae 145, 250 pp. Röse , M. 1996. Ecogeog aphy o he g ass genus Helic o ichon (Poaceae: A eneae) in he Medi e anean and adjacen egions. Plan Sys ema ics and E olu ion 203: 181-281. Röse , M. 1997. Pa e ns o di e si ica ion in Medi e anean oa g asses (Poaceae: A eneae). Lagascalia 19(1-2): 101-120. Röse , M. 2006. 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