Taxonomic revision of Trisetum sect. Trisetum (Poaceae: Pooideae: Aveninae) from Eurasia and North Africa

TAXONOMIC REVISION OF TRISETUM SECT. TRISETUM (POACEAE: POOIDEAE: AVENINAE) FROM EURASIA AND NORTH AFRICA 1 Patricia Barber´a, 2,4 Carlos Romero-Zarco, 3 and Carlos Aedo 2 ABSTRACT A taxonomic revision of Trisetum Pers. sect. Trisetum is presented. We include descriptions and synonyms of each taxon from a study of 894 vouchers from 45 herbaria. Detailed morphometric descriptions, illustrations, distribution maps, an identification key, and habitat data are given for each taxon. Morphometric variation of the main characters is shown by box plots. Thirty-one names are lectotypified. Two neotypes are designated. We recognize eight species of Trisetum in the section: T. alpestre (Host) P. Beauv., T. altaicum Stephan ex Roshev., T. bertolonii Jonsell, T. flavescens (L.) P. Beauv., T. fuscum Schult., T. glaciale (Bory) Boiss., T. gracile (Moris) Boiss., and T. laconicum Boiss. & Orph. Two infraspecific taxa of T. flavescens are recognized (T. flavescens subsp. flavescens and T. flavescens subsp. griseovirens (H. Lindb.) Dobignard). Six of the eight species of Trisetum sect. Trisetum are endemic to the different European mountain ranges, while T. altaicum grows in the Altai and Tian Shan Mountains, and in the mountains of northern Mongolia and southern Russia, and T. flavescens is widespread in temperate regions of Europe, western Asia, and northern Africa. Vegetative propagation by pseudoviviparism is observed for the first time in specimens of T. flavescens subsp. flavescens. Key words: Aveninae, Europe, lectotypification, neotype, northern Africa, taxonomy, Trisetum. Trisetum Pers. is one of the 18 genera of perennial grasses classified in the Aveninae J. Presl subtribe (Poeae tribe, subfamily Pooideae Benth.; cf. Tzvelev, 1989; GPWG, 2001; Soreng et al., 2003, 2007, 2015, 2017). This genus includes about 50 species that often form a significant part of the temperate and cold grasslands of the Northern Hemisphere, although they are also found in South America, Australia, and New Zealand. Their habitat varies according to their wide distributional range, and its species are found in open grasslands to shady areas of forests, high mountain meadows, and even the tundra (Hult´en, 1959; Chrtek, 1965; Clayton & Renvoize, 1986; Randall & Hilu, 1986; Watson & Dallwitz, 1992; Finot et al., 2004, 2005a, 2005b; Barber´a et al., 2017a, 2017b). Trisetum flavescens (L.) P. Beauv., one of the species included in the section studied here, is of great economic interest as a fodder plant for both wild and domestic livestock. To date, detailed taxonomic revisions of only the American and New Zealand species of this genus have been carried out (Edgar, 1998; Finot et al., 2004, 2005a, 2005b), in addition to those recently published by us on two Eurasian sections, Trisetum sect. Acrospelion (Besser) Pfeiff. and Trisetum sect. Sibirica (Chrtek) Barber´a (Barber´a et al., 2017a, 2017b). Six of the species of the section to which we dedicate the present work, Trisetum sect. Trisetum, are endemic to Europe, while both T. altaicum and the most ubiquitous T. flavescens reach eastern Asia. Traditionally, Trisetum has been divided into two sections: Trisetum sect. Trisetum and Trisetum sect. Trisetaera Asch. & Graebn. A review of the history of the genus can be found in Finot et al. (2005a) and Barber´a et al. (2017a). Chrtek (1965, 1967a, 1968) carefully studied the European species of Trisetum, proposing new infrageneric divisions of the genus. 1 This work was financed by Flora Iberica CGL2014–52787–C3–1–P, CGL2012–32914 and CGL2015–66161–P, projects of the Spanish government, supported by funds from the Fonds Europ´een de D´eveloppement ´ Economique et R´egional (FEDER). We are indebted to the curators and staff of the herbaria mentioned in the Materials and Methods section for kindly providing specimens. We thank the editor and anonymous reviewers for their critical reviews of the manuscript and for their helpful suggestions during the editing process. We thank Ram´on Morales for translations of manuscripts in German; Toni Buira and Marta Fernand´ez-Albert for their technical support; J. A. Alejandre, A. Quintanar, and R. Soreng for their helpful comments and suggestions during the development of this work; A. Managlia (BOLO), L. Guglielmone (TO), T. Khmil (LW), W. Paul (KRAM), and Carmen Ulloa (MO) for help in finding rare literature, type materials, and geographic localities; Roman Ufimov for Russian translations and help in finding rare literature; and other personnel from the Komarov Institute, N. N. Tzvelev, Dmitry Geltman, Elena Glazkova, Denis Melnikov, for their help and hospitality during our stay. We also are grateful to Ram´on Garc´ıa Mora for the illustrations. 2 Real Jard´ın Bot´anico, Consejo Superior de Investigaciones Cient´ıficas, Plaza de Murillo, 2, 28014 Madrid, Spain. 3 Departamento de Biolog´ıa Vegetal y Ecolog´ıa, Facultad de Biolog´ıa, Sevilla, Spain. 4 Author for correspondence: [email protected]. VERSION OF RECORD FIRST PUBLISHED ONLINE ON 14 SEPTEMBER 2018 AHEAD OF FALL 2018 ISSUE. doi: 10.3417/2018067 ANN.MISSOURI BOT.GARD. 103: 350–392. Chrtek (1965) divided the European species into four subgenera (Trisetum subg. Trisetum,Trisetum subg. Distichotrisetum Chrtek, Trisetum subg. Glaciotrisetum Chrtek, and Trisetum subg. Graciliotrisetum Chrtek), based mainly on the variation in vernation (conduplicate or convolute) and on anatomical characters of leaves and roots. At the same time, Chrtek (1965) divided the subgenus Trisetum into five sections (sections Trisetum, Trisetaera,Rigida Chrtek, Hispanica Chrtek, and Carpatica Chrtek). Apart from the anatomical characters, Trisetum sect. Trisetum was separated by its lax panicles and short ligules and included T. flavescens,T. alpestre (Host) P. Beauv., T. sibiricum Rupr., T. agrostideum (Laest.) Fr., and T. baregense Laffitte & Mi´egev. The monotypic Trisetum sect. Carpatica, established for T. fuscum Schult. of the Carpathian Mountains, was characterized by its longer ligules and by the hairiness of the lemmas, palea keels, and awns. Tzvelev (1976) simplified the classification of Trisetum by not addressing the anatomical characters used by Chrtek. Tzvelev (1976) recognized three sections (Rigida,Trisetaria, and Trisetum) in his treatment of Trisetum for Russia. He characterized the different sections by the type of growth of underground and vegetative shoots, leaf disposition, panicle density, and rachilla and callus hair length. He included seven species within Trisetum sect. Trisetum:T. ciliare (Kit. ex Schult.) Domin, T. turcicum Chrtek, T. sibiricum,T. flavescens,T. alpestre,T. altaicum Stephan ex Roshev., and T. agrostideum. Later, Probatova (1979), in her study of the genus from the Caucasus, arranged Trisetum sect. Trisetum into four subsections: the monotypic Trisetum subsect. Carpatica (Chrtek) Prob. (including T. ciliare); subsection Trisetum (including T. flavescens s.l., T. parvispiculatum (Tzvelev) Prob., and T. alpestre); subsection Sibirica (Chrtek) Prob. (including T. sibiricum and T. turcicum); and subsection Agrostidea Prob. (including T. altaicum and T. agrostideum). The subsections were characterized by the type of growth of underground and vegetative shoots, lemma apex, callus, awn and palea keel hair length, and anther length. Another relevant floristic study, which covers most of the species of Trisetum sect. Trisetum, is the Flora Europaea treatment by Jonsell (1980). However, he did not recognize any sections. After studying Trisetum sect. Acrospelion and Trisetum sect. Sibirica (Barber´a et al., 2017a, 2017b), we present here the taxonomic revision of Trisetum sect. Trisetum, which is the third of our ongoing monograph on the genus Trisetum in Eurasia and northern Africa. Trisetum sect. Trisetum is here characterized by having panicles lax to somewhat dense or compact, spikelets usually greenish or purplish, callus with short hairs, awns geniculate or bent, rarely straight, and ovaries usually hairy apically. The most serious taxonomic problems of the group have been the delimitation of the section and the treatment given to T. flavescens,a very variable species that historically has been split into many taxa. Our final classification is also based on our current studies on molecular data, which will be published in the near future. This study, based on a thorough review of herbarium material and living plants, is the first to provide a detailed taxonomic scheme of the whole section Trisetum as we currently understand it based on our morphological and molecular surveys of the whole subtribe Aveninae. MATERIALS AND METHODS This revision is based on the study of 894 herbarium specimens from the following herbaria: AAH, ARAN, B, BC, BCN, BM, BOLO, C, F, FI, G, GB, GDA, GH, GOET, H, HBG, JE, K, KRAM, L, LE, LW, M, MA, MW, MO, MPU, NY, O, P, PE, PH, PR, PRC, RO, S, SEV, UPS, US, W, and WU (acronyms according to Thiers, 2017). Photos of specimens from HNWP were also examined on the website of Chinese Virtual Herbarium (,www.cvh.ac.cn.), and specimens from CAI, DAO, FR, and KFTA on the website of Global Plants (JSTOR, ,https://plants.jstor.org/.). An index of numbered collections is presented in Appendix 1. One hundred-seventy specimens were used for the morphometric analyses, as operational taxonomic units, selected to represent, as far as possible, the geographic range and morphological variability of the taxa. Fiftynine quantitative characters were recorded using a Mitutoyo CD-15DCX digital vernier caliper (Mitutoyo, Kawasaki, Japan) on 20 specimens of each species when available. Spikelet measurements were taken on the distal spikelet from the longest branch of the second node of the panicle; floret measurements were taken from the basal floret of the selected spikelet. Leaf blade width was measured 1 cm above the ligule insertion. The rachilla internode between the first and second flowers is also referred to as “rachilla.”Each character was analyzed for its minimum, maximum, and lower and upper quartiles using the Statistica package. Quantitative and qualitative characters are also used in the identification key and descriptions. Minimum and maximum values are noted in brackets, and the lower and upper quartile values (between the 25th and 75th percentiles) are noted outside. Transverse sections of leaf blades were prepared by hand using the uppermost caulinar leaf; they were stained with FASGA (fucsina, alcian blue, safranina, glicerina, agua) (Tolivia & Tolivia,1987)andphotographedwithaNikon SMZ1000 optical microscope (Nikon, Tokyo, Japan). For leaf anatomy, the terminology of Ellis (1976, 1979) was used. Volume 103, Number 3 2018 Barber ´a et al. 351 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa Additional data on the habitat, distribution, and chromosome numbers were checked from literature and collection labels. Species distribution maps were made using ArcGis v.9.3 (ESRI). The material studied is listed under each species in the taxonomic treatment below. The geographic coordinates were derived, in almost all cases, using the GeoNames database (,www.geonames.org.). RESULTS MORPHOLOGY OF TRISETUM SECT.TRISETUM Habit and stem These are perennial grasses commonly with a loose habit, with short to long rhizomes and thus well-isolated shoots, sometimes densely tufted as in Trisetum glaciale and T. gracile. The stems are straight in most of the species, except in Trisetum gracile, which has stems often bent at its blackish nodes, and T. bertolonii, which sometimes has the upper part of the stems curved. The stems are usually shorter than 60 cm, except those in T. laconicum, which can reach more than 80 cm, and T. flavescens subsp. flavescens, which can rarely exceed these sizes. Furthermore, T. glaciale and T. gracile are the shortest species ([3–]8.8–14.4[–19.5] cm and [4.5–] 8.9–12.5[–18.2] cm, respectively). Most of the species of the section produce extravaginal tillers, being intravaginal in Trisetum bertolonii and T. laconicum, rarely in T. fuscum.Trisetum glaciale is the only species that always presents intravaginal branching. The culms are always glabrous in T. altaicum,T. fuscum, and T. laconicum, sometimes puberulous in some parts in T. alpestre,T. gracile, and T. glaciale, rarely so in T. flavescens.Trisetum bertolonii always has pubescent culms. The number of nodes varies between two and six (seven). The culm internodes are mostly concentrated in the lower part of the culm in most species, although T. flavescens,T. fuscum, and T. laconicum have relatively elongated lower internodes and nodes well separated along the culm. Leaves In Trisetum sect. Trisetum, as in the rest of the genus, the different characters of the leaf sheaths and leaf blades are variable in the same plant, depending on whether they are the basal or top culm leaf blades, or if they are young or mature (Barber´a et al., 2017a, 2017b). Leaf sheaths Basal leaf sheaths are sparsely to densely pubescent, sometimes glabrous in Trisetum bertolonii and T. gracile, rarely in T. alpestre and T. flavescens. The oldest basal leaf sheaths usually decay into fibers in most of the species, forming a dense layer of decomposed sheaths in T. glaciale and T. gracile. The latter two species have the leaf sheaths distinctly veined and are somewhat inflated in T. gracile. Top culm leaf sheaths are always glabrous in Trisetum bertolonii,T. gracile, and T. laconicum, sometimes with hairs on the upper part in T. fuscum and sometimes pubescent in T. alpestre and T. altaicum, rarely in T. flavescens.Trisetum glaciale always has pubescent top culm leaf sheaths. Sheath margins are glabrous in Trisetum bertolonii,T. glaciale, and T. gracile and always ciliate in T. altaicum and T. laconicum. Ciliate margins also sometimes develop in T. alpestre and T. fuscum and rarely in T. flavescens. Leaf blades The species of Trisetum sect. Trisetum have flat leaf blades in most of the species, although those of T. flavescens sometimes have convolute to revolute or conduplicate ones, rarely filiform. Most of the species of the section have non-rigid leaf blades, but T. glaciale and T. gracile have rigid ones with a thickened midrib, as well as the shortest blades (Fig. 1A), with the margins thickened and cartilaginous in T. glaciale. The indumentum of the leaf blades has traditionally been used as a distinctive character for taxa delimitation (Tzvelev, 1976; Jonsell, 1980). However, for Trisetum sect. Acrospelion and Trisetum sect. Sibirica, the leaf blade features have been considered less important than those of the spikelet morphology for the taxonomy of the widespread species, due to the high variability of leaf blade traits (Barber´a et al., 2017a, 2017b). The species of Trisetum sect. Trisetum have leaves varying from glabrous to pubescent adaxially and abaxially, usually with ciliate margins. The species of Trisetum sect. Trisetum, as the rest of the genus, are mesophytic or xerophytic grasses. While the abaxial surface of the leaf blade often has a flat surface, the adaxial surface has ribs of varying depth (Metcalfe, 1960; Watson & Dallwitz, 1992). The xerophytic species of the section have deeper intercostal zones and associated ribs than the mesophytic ones, and the sclerenchyma girders and strands are well developed (Barber´a et al., 2017a, 2017b). As indicated before, the midrib is not particularly marked in most of the species of the section but is strongly marked only in T. glaciale and T. gracile. Chrtek (1965) made the first detailed study of the histology of European Trisetum, including some of the species included in our work of section Trisetum (T. fuscum,T. glaciale, and T. gracile). The vascular bundles of the species of Trisetum sect. Trisetum can be accompanied or not by more or less developed sclerenchyma girders or strands that attach to the abaxial or adaxial sides of the leaf blade, 352 Annals of the Missouri Botanical Garden or to both sides. No transversal section of T. laconicum has been done because of the few specimens available. All of the species of the section have a cap of sclerenchyma at the margins. Trisetum altaicum and T. fuscum are mesophytic grasses, which explains their shallow intercostal zones or ribs, and the vascular bundles with no strands and with narrow girders adaxially and abaxially. Trisetum flavescens subsp. flavescens has vascular bundles with girders as deep as wide in both surfaces, alternating with bundles with only small girders Figure 1. Box plots of a selection of studied variables. —A. Top culm leaf blade length. —B. Ligule length. —C. Spikelet length. —D. Rachilla hair length. —E. Lower glume width. —F. Lower glume/upper glume length. —G. Lemma length. —H. Palea/lemma length. Numbers along x-axis correspond to the studied taxa: 1. Trisetum glaciale (Bory) Boiss. 2. Trisetum alpestre (Host) P. Beauv. 3. Trisetum fuscum Schult. 4. Trisetum bertolonii Jonsell. 5. Trisetum altaicum Stephan ex Roshev. 6. Trisetum gracile (Moris) Boiss. 7. Trisetum flavescens (L.) P. Beauv. subsp. flavescens.8.Trisetum flavescens subsp. griseovirens (H. Lindb.) Dobignard. 9. Trisetum laconicum Boiss. & Orph. Volume 103, Number 3 2018 Barber ´a et al. 353 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa adaxially. The rest of the species of the section are more or less xerophytic grasses. Trisetum glaciale has girders only on the midrib, which are abaxially well developed, while T. gracile has no girders and presents well-developed strands in every vascular bundle in both surfaces. Trisetum alpestre,T. bertolonii,andT. flavescens subsp. griseovirens present both, girders and strands, more or less developed. Ligules The ligule measurements used in this study were taken from the top culm leaf blades, which are usually longer than those of the basal leaves. Trisetum fuscum has the longest ligules ([3–]3.7–4.7[–4.8] mm), followed by T. laconicum (2.5–3.2 mm). The ligules are membranous, truncate or acute, dentate to laciniate, always glabrous in T. bertolonii and T. fuscum, and usually more or less hairy along the upper edge in the rest of the species (Fig. 1B). Inflorescences Inflorescences are paniculate, lax to somewhat dense or compact, narrowly elliptic to elliptic or lanceolate in outline, sometimes oblong, and always ovate in Trisetum glaciale and T. gracile.Trisetum laconicum has the longest panicles (8–19 cm), followed by T. flavescens and T. fuscum ([3.5–]6–10.5[–16.2] cm and [6.8–] 7.9–11.2[–13] cm, respectively), whereas T. glaciale and T. gracile have the shortest ones ([2.3–]3–4.2[–4.8] cm and [1.3–]1.6–2.5[–3.1] cm, respectively). Trisetum fuscum is characterized by having the longest basal branches ([1–]2.2–3.8[–5] cm), while T. glaciale and T. gracile have the shortest ones ([0.3–] 0.4–0.9[–1.1] cm and [0.2–]0.3–0.9[–1.1] cm, respectively). The inflorescence rachis is glabrous to sparsely pubescent or always pubescent in Trisetum bertolonii, with hairs up to 0.5 mm long. Spikelets Spikelet length in Trisetum sect. Trisetum, as in the rest of the genus, varies according to the lemma length and the number of florets. Trisetum laconicum and T. bertolonii have the longest spikelets, followed by T. flavescens subsp. griseovirens (8–9.2 mm, 7–9 mm, and [6.3–]6.5–7.3[–7.7] mm, respectively) (Fig. 1C). The awn is not included in the length of the spikelet. Furthermore, T. laconicum has the longest pedicels (5–6.3 mm). Rachilla segments are always pubescent; Trisetum laconicum,T. gracile, and T. bertolonii have the longest hairs (1.5–2.5 mm, [1.6–]2–2.4[–3] mm, and [0.8–] 1.3–2.2[–2.8] mm, respectively). Therefore, it is a useful diagnostic character for those three species (Fig. 1D). The number of florets is usually two to three per spikelet, except in Trisetum laconicum, which has three to four florets, rarely four in T. flavescens, and rarely one in T. glaciale. Vegetative progagation by pseudoviviparism was observed in a specimen of Trisetum flavescens for the first time. Glumes In section Trisetum, the glumes are unequal, except in T. glaciale and T. gracile, which have subequal glumes (lower glume length:upper glume length ratio of [0.75–] 0.85–0.92[–0.98] and [0.79–]0.82–0.89[–0.97], respectively) (Fig. 1F). Trisetum altaicum,T. gracile,and usually T. alpestre,T. fuscum,andT. glaciale have elliptic to oblong or lanceolate lower glumes, these wider than those of the rest of the species of the section, which have usually narrower ones, sometimes linear in T. bertolonii. Lower and upper glumes are acuminate to long-acuminate, sometimes acute, rarely with the upper one aristulate in T. alpestre, with some short hairs from the middle part to the top of the main nerve and along the margins. The lower glume in T. bertolonii and T. laconicum always has one nerve; other species rarely have one (T. gracile)ortwolateralnerves(T. alpestre,T. altaicum,T. flavescens,andT. fuscum)(Fig. 1E). Trisetum glaciale has lower glumes with three nerves, sometimes two. The upper glume has always three nerves, the central one reaching the tip, the lateral ones extending to the middle or the upper half. Lemma The lemma structure, as in the rest of the genus, is characterized by having a dorsal awn and a bifid apex with two apical teeth normally ending in two aristules separated by a more or less deep sinus. Aristules are the intermediate nerves protruding beyond each tooth apex (Nicora, 1978; Koch, 1979; Finot et al., 2006; Barber´a et al., 2017a, 2017b). Aristule length is a variable character at the species level, being longer in Trisetum bertolonii than in therestofthespecies(1.2–2.3[–4] mm), followed by T. flavescens subsp. griseovirens ([0.9–]1–1.4[–1.8] mm) (Fig. 2B). The length of the lemma, which includes the aristule length without taking the awn into account, is longer in T. bertolonii,T. laconicum,andT. flavescens subsp. griseovirens ([5.7–]6–8[–10] mm, 6.5–6.6 mm, and 6–6.2[–6.4] mm, respectively). However, the length of the lemma does not differ greatly among the rest of the species of the section (Fig. 1G). The lemma is laterally compressed, from narrowly or broadly lanceolate to elliptic or oblong, greenish to yellowish at maturity in the central part, surrounded by purplish tinges, except in T. bertolonii, which has hyaline (colorless) margins. Most of the species of Trisetum sect. Trisetum have glabrous lemmas with very short, adpressed hairs up to 0.1 mm long, except T. fuscum, which sometimes has longer hairs at the awn insertion, and T. gracile, which can have pubescent 354 Annals of the Missouri Botanical Garden lemmas with hairs (0.05–)0.6–1.4(–2) mm long between its base and the awn insertion. Awn The length and shape of the awn are important taxonomic characters for the classification of the species of this section. Trisetum bertolonii has the awn significantly longer than the rest of the species of the section ([11–]11.5–16 [–17] mm), but the length does not differ greatly in the rest of the species. At subspecific rank, the length of the awn is also a good character to differentiate T. flavescens subsp. griseovirens, with longer awns than T. flavescens subsp. flavescens ([8–]8.2–9[–10.6] mm and [4.2–]5.6–7[–8.3] mm, respectively) (Fig. 2A). The awn shape is slightly geniculate to geniculate, and with a twisted column up to the geniculation in T. flavescens,T. glaciale,andT. gracile. Trisetum altaicum,T. fuscum,andT. laconicum have more or less bent awns at the base, while T. alpestre and T. bertolonii have awns straight or bent at the middle. The awn is dorsal, inserted above or at the middle of the lemma, except in T. gracile, where the awn is inserted slightly below the middle (awn insertion from the base length: lemma length ratio of [0.34–]0.41–0.49[–0.51]). Callus The callus is elliptic to orbicular or sometimes oblong and always has short hairs. Trisetum bertolonii has the longest callus hairs (1.5–2.2[–2.6] mm), while the rest of species have hairs always shorter than 1.4 mm (Fig. 2C). Palea The palea disposition and shape are the same as in the other sections. The surfaces are hyaline. The two keels extend into two fine teeth (Barber´a et al., 2017a, 2017b). In this section, the palea of Trisetum laconicum is longer than that in the rest of species (5.8–6 mm). The upper part of the flange edges and keels of the palea normally have very short hairs, which can reach to 0.4 mm long in Figure 2. Box plots of a selection of studied variables. —A. Total awn length. —B. Aristule length. —C. Callus indument length. —D. Anther length. Numbers along x-axis correspond to the studied taxa: 1. Trisetum glaciale (Bory) Boiss. 2. Trisetum alpestre (Host) P. Beauv. 3. Trisetum fuscum Schult. 4. Trisetum bertolonii Jonsell. 5. Trisetum altaicum Stephan ex Roshev. 6. Trisetum gracile (Moris) Boiss. 7. Trisetum flavescens (L.) P. Beauv. subsp. flavescens.8.Trisetum flavescens subsp. griseovirens (H. Lindb.) Dobignard. 9. Trisetum laconicum Boiss. & Orph. Volume 103, Number 3 2018 Barber ´a et al. 355 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa T. fuscum. The length of both the palea and the lemma is always markedly unequal in T. bertolonii (palea length: lemma length ratio of [0.4–]0.58–0.67[–0.7]), followed by T. flavescens subsp. griseovirens (palea length:lemma length ratio of [0.53–]0.65–0.76[–0.77]) (Fig. 1H). In the rest of the species, the difference between the palea and the lemma is more or less subequal to equal. Lodicules As in the rest of the sections of Trisetum, the two lodicules flank the dorsal sides of the ovary or caryopsis, and have a more or less oblanceolate shape (Barber´a et al., 2017a, 2017b). The apices are irregularly dentate to laciniate or with (one) two to three teeth or lobules of equal or different depth, except in T. gracile, in which the apices are usually entire. Stamens The length of the anthers does not vary much among the species of this section, except in Trisetum altaicum,which has the shortest anthers ([0.8–]1–1.35 mm) (Fig. 2D). Ovary and caryopsis In this section, most of the species have a glabrous ovary. However, Trisetum glaciale has ovaries that range from sparsely to more densely puberulous on the upper part, rarely glabrous, and T. alpestre usually has an apical tuft of hairs or scattered hairs at the apex. Trisetum altaicum rarely has a single hair on the upper part of the ovary. The shape of the mature caryopsis is similar in all species, being narrowly elliptic to oblong, while Trisetum glaciale has a more or less fusiform one. The transversal section is narrowly elliptic to elliptic and the caryopsis is not sulcate, with a punctiform hilum. The endosperm is semi-liquid. TAXONOMIC TREATMENT Trisetum Pers., Syn. Pl. 1: 97. 1805. TYPE: Avena flavescens L. (current name T. flavescens (L.) P. Beauv.) (lectotype, designated by Hitchcock [1920: 108]). Trisetum Pers. sect. Trisetum. Trisetum sect. Gracilia Chrtek & V. Jir´asek, Webbia 17: 573. 1963. TYPE: Trisetum gracile (Moris) Boiss. Trisetum sect. Carpatica Chrtek, Bot. Not. 118(2): 222. 1965. Trisetum subsect. Carpatica (Chrtek) Prob., Novosti Sist. Vyssh.Rast.15:19.1979.TYPE:Trisetum fuscum Schult. Trisetum subg. Glaciotrisetum Chrtek, Bot. Not. 118(2): 223. 1965. TYPE: Trisetum glaciale (Bory) Boiss. Trisetum ser. Laconica Chrtek, Acta Univ. Carol., Biol. 1966: 94. 1967. TYPE: Trisetum laconicum Boiss. & Orph. Herbs with or without tufted habit, rhizomatous, sometimes shortly stooling; panicles lax to somewhat dense or compact; spikelets greenish or purplish, never goldish brown; callus with short hairs, rarely longer than 1.4 mm; awns distinctly geniculate or slightly bent at middle or base, rarely straight; ovaries glabrous, sometimes sparsely to densely pubescent apically. Distribution and habitat. Trisetum sect. Trisetum occurs from western Europe and the Maghrebian Mountains to western Asia, extending eastward into the mountains of Altai and Tian Shan and northern Mongolia and adjacent Russia, eastern Kyrgyz Republic, and Kazakhstan; it is introduced in North and South America and New Zealand. The richest areas are the different European mountain ranges. It grows on mountain meadows and floodplains, in rock fissures and forests, at elevations from sea level to more than 3000 m. KEY TO THE TAXA OF TRISETUM SECT.TRISETUM 1. Callus hairs 1.5–2.2(–2.6) mm long; awns (11–)11.5–16(–17) mm long . . . . .....................T. bertolonii Jonsell 19. Callus hairs (0.1–)0.3–0.5(–1.3) mm long; awns (3.8–)5.3–7(–10.7) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 2. Plants (3–)8.9–14.1(–19.5) cm tall; top culm leaf blades (0.2–)0.3–1.5(–2.6) cm long; panicles (1.3–)2.2– 3.8(–4.8) cm long, ovate in outline ...................................................................3 3. Nodes enfolded by sheaths; basal leaf sheaths not inflated, densely pubescent; basal leaf blades with margins thickened, cartilaginous; callus hairs (0.2–)0.3–0.6(–0.9)mmlong..............T. glaciale (Bory) Boiss. 39. Nodes not enfolded by sheaths, rarely enfolded; basal leaf sheaths somewhat inflated, glabrous to sparsely pubescent; basal leaf blades with margins not thickened; callus hairs (0.7–)1–1.3 mm long . . . T. gracile (Moris) Boiss. 29. Plants (14.4–)28–47(–141.5) cm tall; top culm leaf blades (1.3–)2.2–12.5(–14.5) cm long; panicles (3.4–)6– 9.4(–19) cm long, not or very rarely ovate .............................................................4 4. Spikelets 8–9.2 mm long; paleae 5.8–6 mm long . . . . . ......................T. laconicum Boiss. & Orph. 49. Spikelets (3.8–)5.3–6.3(–7.8) mm long; paleae (2.2–)3.6–4.3(–5.6) mm long . . .........................5 5. Ligules (3–)3.7–4.7(–4.8) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. fuscum Schult. 59. Ligules (0.5–)0.8–1.7(–4) mm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 6. Lower glumes (0.8–)1.2–1.6 mm wide; anthers (0.8–)1–1.3 mm long . . . T. altaicum Stephan ex Roshev. 69. Lower glumes (0.4–)0.6–1(–1.7) mm wide; anthers (1–)1.8–2.2(–3) mm long . . . . . . . . . . . . . . . . . 7 7. Awns straight or bent, very slightly twisted at base; ovaries usually with apical tuft of hairs 0.1–0.2(–0.3) mm long or with scattered hairs at apex . . .........T. alpestre (Host) P. Beauv. 79. Awns geniculate to slightly geniculate, more or less twisted at base, rarely not twisted; ovaries always glabrous ............................................T. flavescens (L.) P. Beauv. 356 Annals of the Missouri Botanical Garden 1. Trisetum alpestre (Host) P. Beauv., Ess. Agrostogr.: 88. 1812. Avena alpestris Host, Icon. Descr. Gram. Austriac. 3: 27, tab. 39. 1805. Trisetaria alpestris (Host) Baumg., Enum. Stirp. Transsilv. 3: 264. 1816. Avena flavescens var. alpestris (Host) Duby, Bot. Gall. 1: 512. 1828. Trisetum flavescens var. alpestris (Host) Schrad., Linnaea 12: 443. 1838. Trisetum flavescens subsp. alpestre (Host) Figure 3. Trisetum alpestre (Host) P. Beauv. —A. Habit. —B. Transverse section of leaf blade. —C. Portion of sheath, ligule, and blade. —D. Portions of culm and node. —E. Spikelet. —F. Florets. —G. Lower glume, dorsal view. —H. Upper glume, dorsal view. —I. Floret. —J. Lemma, upper part, lateral view. —K. Palea, lateral view. —L. Stamens. —M. Lodicules. —N. Pistil. —O. Caryopsis. A, C–L based on Du Rietz s.n. (UPS-V-644660); B based on Bornm¨uller s.n. (B-10_0526398); N based on Du Rietz s.n. (UPS-V-644660) and Barber´a et al. 953PB (MA-872340); M based on Du Rietz s.n. (UPS-V-644660), Barber´a et al. 953PB (MA- 872340), Barber´a & Quintanar 1062PB (MA-876682), and Vestergren s.n. (S); O based on Neyraut s.n. (JE). Volume 103, Number 3 2018 Barber ´a et al. 357 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa Hack., Magyar Bot. Lapok 2: 106. 1903. TYPE: Austria. N. T. Host s.n. (lectotype, designated here, W-1885-0002400!; isolectotype, W-0024994!). Figure 3. Trisetum alpestre var. argentoideum Schur, Oesterr. Bot. Z. 10: 74. 1860. TYPE: Romania. Harghita, } Ocsem Teteje Mtn., 46°399N, 25°489E, July, P. J. F. Schur s.n. (lectotype, designated here, W!). Trisetum alpestre var. purpurascens Schur, Oesterr. Bot. Z. 10: 74. 1860. TYPE: Romania. Harghita, } Ocsem Teteje Mtn., 46°399N, 25°489E, 28 July 1853, P. J. F. Schur s.n. (lectotype, designated here, W!). Trisetum alpestre var. glabrescens Schur, Enum. Pl. Transsilv.: 759. 1866. Trisetum alpestre subsp. glabrescens (Schur) Tzvelev, Novosti Sist. Vyssh. Rast. 7: 64. 1971. TYPE: Romania. K¨onigstein Mtn. (Piatra Craiului), 45°389N, 25°369E, Aug., P. J. F. Schur s.n. (lectotype, designated here, LW-213277 image!). Trisetum alpestre var. macranthum Schur, Enum. Pl. Transsilv.: 759. 1866. TYPE: Romania. Harghita, } Ocsem Teteje Mtn., 46°399N, 25°489E, 28 July 1853, P. J. F. Schur s.n. (lectotype, designated here, LW-213278 image!). Trisetum baregense Laffitte & Mi´egev., Bull. Soc. Bot. France 21: 46. 1874. Avena subalpestris subsp. baregense [baregensis] (Laffitte & Mi´egev.) Nyman, Consp. Fl. Eur.: 813. 1882. Trisetum agrostideum subsp. baregense (Laffitte & Mi´egev.) Chouard, Bull. Soc. Bot. France 72: 340. 1925. Trisetum flavescens subsp. baregense (Laffitte & Mi´egev. ex Mi´egev.) O. Bol`os, Masalles & Vigo, Collect. Bot. (Barcelona) 17: 96. 1988. Trisetaria baregense (Laffitte & Mi´egev.) Banfi & Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat. Milano 135: 382. 1996. TYPE: France. Haut-Pyren´ees, H´eas valley, Gabi´edou peak, 42°429N, 0°59E, 13 July 1862, J. Mi´egeville s.n. (lectotype, designated here, P-2217672!). Trisetum alpestre f. calvescens Hack., Magyar Bot. Lapok 2: 108. 1903. TYPE: Romania. Harguita Distr., Nagy Hagym´as Mtn., 46°429N, 25°489E, 21 June 1901, A. de Degen, Gram. Hung. 121 (lectotype, designated here, JE!). Trisetum flavescens f. tirolense Hack., Allg. Bot. Z. Syst. 9: 189. 1904. TYPE: Italy. Trentino-Alto Adige, South Tyrol, Carbonin [Schluderbach], 46°379N, 12°139E, 21 Aug. 1903, A. Kneucker,Gram. Exsicc. 442 (lectotype, designated here, B-10_0366348!; isolectotypes, B-10_ 0366347!, C!, G-00443036!, G-00443093!, KFTA- 0000478 image!, L-0050287!, L-1346752!, MA-292728, MO-2879843!, MO-3057065!, NY!, O-V2126570!, O- V2126668!, PR-9070!, S!, US-557059!, WU!). Trisetum alpestre f. anomalum Zapał., Rozpr. Wydz. Mat.- Przyr. Akad. Umiejetn., Dzia B, Nauki Biol. 4: 111. 1904. TYPE: Tatra Mtns., A. Rehman s.n. (LW?, herbarium number 69/276, original material not located). Trisetum alpestre var. aureum Zapał., Rozpr. Wydz. Mat.-Przyr. Akad. Umiejetn., Dzia B, Nauki Biol. 4: 111. 1904. TYPE: Poland. Lesser Poland, Tatra Mtns., Jaworzynka valley, 49°169N, 19°599E, 1856, [F. I. Berdau]s.n. (lectotype, designated here, KRAM-74425, the specimen mounted lower left hand of the sheet, with yellowish spikelets, of the two marked with “1.”, image!). Trisetum alpestre f. majus Zapał., Rozpr. Wydz. Mat.-Przyr. Akad. Umiejetn., Dzia B, Nauki Biol. 4: 111. 1904. TYPE: Poland. Lesser Poland, Tatra Mtns., Dolina Ko´scieliska valley, 49°169N, 19°529E, B. Kotula s.n. (KRAM?, original material not located). Trisetum alpestre var. pulchrum Zapał., Rozpr. Wydz. Mat.- Przyr. Akad. Umiejetn., Dzia B, Nauki Biol. 4: 110. 1904. SYNTYPES: Tatra Mtns., B. Kotula s.n. (KRAM?, original material not located); Ukraine. Chyvchyn Mtn., 47°519N, 24°429E, E. Woloszczak s.n. (W?, original material not located); Romania. Suhard Mtns., Bucovina, 47°309N, 25°219E, A. Rehman s.n. (LW?, original material not located). Trisetum alpestre var. tatrense Zapał., Rozpr. Wydz. Mat.-Przyr. Akad. Umiejetn., Dzia B, Nauki Biol. 4: 111. 1904. TYPE: Poland. Lesser Poland, Tatra Mtns., Mt. Łysanki, 49°169N, 19°559E, 20 July 1887, [B. Kotula]s.n. (lectotype, designated here, KRAM-74418 image!). Herbs (14.4–)21.5–35.7(–53.5) cm tall, slightly caespitose to caespitose, shortly rhizomatous, culm 0.3–0.8 mm diam., straight, glabrous, sometimes puberulous at middle part or around nodes, with hairs up to 1.5 mm; nodes (2 to)3 to 4(to 5), mostly concentrated in lower part of culm, not included in sheaths, sometimes included, glabrous, yellowish to dark brownish. Basal leaf sheath surfaces and margins pubescent, rarely glabrous, with hairs (0.05–)0.3–0.6 mm, sometimes decaying into fibers, yellowish, sometimes greenish or brownish; basal leaf blades (1.9–)3–7.5(–12) cm 3 (1.1–)1.6–2.3(–4) mm, flat, slightly inrolled when dried, from sparsely to densely pubescent abaxially and adaxially, with hairs (0.05–)0.1–0.9(–1.2) mm, sometimes glabrous, margins also with hairs, yellowish or brownish to greenish; top culm leaf sheaths (5–)6.3–9.6(–16) cm, glabrous to pubescent, with or without cilia; top culm leaf blades (1.5–)2.2–5.2(–9.3) cm 3(1–)1.6–3(–4.1) mm, flat, sometimes with margins inrolled, parallel to culm, glabrous to pubescent adaxially and abaxially, with hairs (0.05–)0.1–0.8(–1.7) mm, with cilia, greenish to grayish, sometimes yellowish or brownish; inner collar region glabrous to pubescent on margins, with hairs (0.05–)0.4–0.7(–1.1) mm; ligules (0.5–)0.9–2(–4) mm, irregularly dentate, sometimes laciniate, glabrous, sometimes with scattered hairs 0.05–0.3 mm on apex. Basal node of panicle glabrous. Panicles (3.4–)5.4–7.5 (–11.5) 3(1.2–)1.8–2.5(–3.6) cm, lanceolate in outline, sometimes oblong to elliptic, rather lax; rachis internodes glabrous to sparsely pubescent, mostly on upper part, with hairs up to 0.2 mm; longest basal branches (0.4–)0.8–2(–3.3) cm. Spikelets (4.4–)5.5–6.3(–7.8) 3 (1.2–)2.2–3.2(–5) mm, 2- or 3-flowered; pedicels (1.5–) 2.4–4.1(–5.2) mm, glabrous to sparsely pubescent, mostly apically, with hairs up to 0.2 mm. Glumes unequal (lower glume length:upper glume length ratio of [0.5–]0.69–0.79[–0.9]); lower glume (2.5–)3.4–4.4 (–5.3) 3(0.4–)0.8–1.2(–1.7) mm, narrowly to broadly lanceolate, rarely elliptic to oblong (lower glume width: lower glume length ratio of [0.15–]0.19–0.31[–0.4]), acuminate to long-acuminate, 1(to 3)-nerved, glabrous, 358 Annals of the Missouri Botanical Garden Aug. 1985, Alanko et al. 53269 (H); Terektinskiy range, the Kostakhta River, 50°309N, 86°09E, 4 July 1931, Shishkin et al. s.n. (LE, NY); Kosh-Agachsky rayon, 17 km SW of Chibit, pass from Baksara to Eshtykol valley, 50°149N, 87°199E, 28 July 2008, Serogin & Serogin S-443 (MW); Seminsky Pass, 51°19N, 85°389E, 26 July 1996, Sukopp et al. 1667 (NY); Zmeinogorsk uyezd, valley of the Chernovaya River, 51°99N, 82°119E, 5 Aug. 1910, Tomin 313 (LE). Buryatia Republic: Tunkinsky Distr., near the selo of Mondy, bottom of the valley of the Khulugaysha River, 51°409N, 100°599E, 2 Aug. 1963, Alyanskaya et al. s.n. (LE); Lake Baykal, source of the Muzhinay River, 54°549N, 108°549E, Aug. 1967, Egorova & Siplivinskiy s.n. (AAH, GB, LE); Barguzin uyezd, basin of the Muya River, on top of the Kindikanskiy pass, 53°379N, 109°389E, 7 Aug. 1914, Korotkiy et al. 849 (LE); eastern Sayan, the Pogranichniy range, the Tengisin-Daban, 52°109N, 98°569E, 19 Aug. 1959, Malyshev & Pezhemskiy 56 (LE); Hangarulsky range, at the source of the Snezhnaya River, 51°59N, 102°439E, 20 Aug. 1912, Smirnow 615 (LE); Lake Baikal, Chivyrskuy, upper reach of the Khozhalvyy River, 53°529N, 109°179E, 1 Aug. 1928, Sukachev et al. 1652 (LE); Barguzin uyezd, mouth of the Kudalda River, 54°99N, 109°339E, 25 July 1939, Tyulina s.n. (LE). Irkutsk: Lake Baykal, Kulkut, on the way up to Khamar- Daban, 51°559N, 105°59E, 14 July 1915, Tsinzerling s.n. (LE). Khakassia Republic: Abakan River, Mt. Saman, 52°169N, 88°599E, 8 Aug. 1927, Plotnikov s.n. (PR). Krasnoyarsk Krai: Western Sayan, vic. of the Olenya Rechka station, 52°489N, 93°179E, 21 July 1964, Khabarov & Chayka s.n. (LE); Western Sayan, the Monysh range, 51°549N, 92°239E, 25 July 1968, Krasnoborov & Khanminchum 6812 (LE); Western Sayan, Sabinskiy range, at the upper reach of Sambyd River, 51°549N, 92°239E, 20 Aug. 1966, Krasnoborov & Kuklina s.n. (LE); Minusinsk Distr., in alpibus Sajanensibus ad fl. Kazyr super, 53°429N, 91°419E, 12 Aug. 1913, Kusnezow & Iljyn s.n. (K, LE, MO, MW, PR, S); basin of the Sydy, Sisma & Many rivers (tributaries of Enisey), 54°229N, 91°519E, 30 July 1908, Volkov 397 (LE). Tuva Republic: western slope of the Akademika Obrucheva range, basin of the Tapsa River, upper reach of its right tributary the Kara-Khem River, 53°29N, 95°49E, 10 July 1974, Krasnoborov & Khanminchun 749 (LE); Western Sayan, Uyukskiy range, upper reach of the Orto-Khem River, eastern part of Mt. Bedelig, 51°479N, 92°269E, 18 July 1974, Lomosonova 2590 (LE); Western Sayan, Kurtushibinskiy range, interfluve of the Khem & Ozhu rivers, 51°559N, 93°09E, 11 July 1979, Shaulo & Belskaya 1520 (LE). Zabaykalsky Krai: Khentey-Chikoyskoe upland, the Borshchovochniy range, 51°439N, 111°589E, 5 Aug. 1967, Maksimova s.n. (AAH, B, G, K, LE, M, S); Aksha uyezd, basin of the Onon River, near the source of the Balidzhi River, 50°169N, 113°179E, 1 Aug. 1913, Smirnow 1433 (LE). 3. Trisetum bertolonii Jonsell, Bot. J. Linn. Soc. 76 (4): 320. 1978. Avena villosa Bertol., Excerpta Re Bot.: 6. 1820 [Opusc. Sci. 4: 222. 1820, n.v.]. Trisetum villosum (Bertol.) Schult., Mant. 2: 368. 1824, nom. illeg., non Trisetum villosum Pers., Syn. Pl. 1: 97. 1805. Trisetaria villosa (Bertol.) Banfi & Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat. Milano 135: 386. 1996. TYPE: Italy. Teramo Prov., Monte Corno, Arapietra, 42°299N, 13°349E, 1818, A. Orsini s.n. (lectotype, designated by Jonsell [1978: 320], BOLO image!). Figure 7. Trisetum villosum var. glaberrimum Ces., Comp. Fl. Ital. 1(3): 64. 1869. Trisetum villosum subsp. glaberrimum (Ces.) Arcang., Comp. Fl. Ital.: 779. 1882. TYPE: Italy. Chiarino, G. Gussone s.n. (original material not located). Herbs (16.5–)17.7–27.8(–38) cm tall, caespitose, shortly rhizomatous, culm 0.4–07 mm diam., straight, sometimes curved at upper part, pubescent, with hairs 0.2–0.8 mm; nodes 2 to 3(to 4), mostly concentrated in lower part of culm, included or not in sheaths, glabrous, brownish. Basal leaf sheaths glabrous to pubescent, with hairs 0.1–0.3 mm, decaying into fibers, yellowish; basal leaf blades (2–)4.2–7.8(–12) cm 31.2–1.5(–2) mm, inrolled, glabrous abaxially and pubescent adaxially, with hairs up to 0.2 mm, also on margins, grayish to greenish; top culm leaf sheaths (3.6–)5.3–8.5(–11) cm, glabrous, without cilia on margins; top culm leaf blades (1–)2.5–3.8(–6.2) cm 31.4–2(–2.4) mm, conduplicate or flat, with margins inrolled, parallel to culm, pubescent abaxially, glabrous adaxially, with scattered hairs on margins, with hairs (0.05–)0.2–0.4 mm, greenish; inner collar region glabrous; ligules 0.5–1.3 mm, slightly irregularly dentate, glabrous. Basal node of panicle glabrous to pubescent, with hairs up to 0.5 mm. Panicles (3.5–)4.6–7.4(–8) 31.6–3 cm, narrowly elliptic, sometimes ovate to broadly lanceolate in outline, rather lax; rachis internodes pubescent, rarely sparsely puberulous, with hairs up to 0.5 mm; longest basal branches (0.7–)0.9–1.6(–1.7) cm. Spikelets 7–932.5–4(–5) mm, 2- to 3-flowered; pedicels 3.7–4.7 (–8.4) mm, pubescent, sometimes very sparsely pubescent, with hairs up to 0.2 mm. Glumes unequal (lower glume length:upper glume length ratio of [0.53–]0.63–0.7); lower glume (3.8–)4.6–5.3(–6) 3 (0.4–)0.6–1 mm, narrowly lanceolate, rarely linear (lower glume width:lower glume length ratio of [0.11–]0.14–0.17[–0.2]), long-acuminate to acuminate, 1-nerved, glabrous, usually with scattered hairs up to 0.1 mm on upper part of central nerve, greenish margins hyaline, rarely surrounded by purplish; upper glume 6.8–8(–9) 31.4–2(–2.4) mm, broadly lanceolate to oblong or elliptic, rarely narrowly lanceolate (upper glume width:upper glume length ratio of [0.19–] 0.22–0.28[–0.3]), acuminate to long-acuminate, 3- nerved, glabrous, sometimes with scattered hairs up to 0.1 mm on upper part of midrib, greenish, sometimes yellowish, with hyaline margins; rachilla segment between first and second florets 1–1.8 mm, with hairs (0.8–)1.3–2.2(–2.8) mm; rachilla segment to sterile floret 1.4–2(–2.3) mm, with hairs 1–1.5 mm. Lemmas (5.7–)6–8(–10) 3(0.4–)0.6–1 mm, oblong to narrowly lanceolate (lemma width:lemma length ratio of 0.13–0.23[–0.3]), glabrous, sometimes scabridulous, yellowish or greenish, with hyaline margins, sometimes surrounded by purplish flush; apical teeth 0.3–0.5 (–0.8) mm, with aristules 1.2–2.3(–4) mm; awn (11–) 11.5–16(–17) mm, inserted 2.8–4 mm from base (awn Volume 103, Number 3 2018 Barber ´a et al. 365 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa insertion from base length:lemma length ratio of [0.56–] 0.6–0.68[–0.75]), straight or slightly curved at middle, twisted, with short, adpressed hairs up to 0.05 mm; callus 0.2–0.3 mm, elliptic, rarely orbicular, with hairs 1.5–2.2(–2.6) mm. Paleae (3.2–)3.8–5.2 30.8–1.2 mm (palea length:lemma length ratio of [0.4–]0.58–0.67 [–0.7]), oblong to oblanceolate, margins glabrous or with scattered hairs; teeth 0.2–0.4(–0.6) mm, with short, Figure 7. Trisetum bertolonii Jonsell. —A. Habit. —B. Portion of culm internode and node. —C. Portion of sheath, ligule, and blade. —D. Transverse section of leaf blade. —E. Spikelet. —F. Florets. —G. Lower glume, dorsal view. —H. Upper glume, dorsal view. —I. Floret. —J. Lemma, upper part, lateral view. —K. Palea, lateral view. —L. Lodicules. —M. Stamens. —N. Pistil. —O. Caryopsis. A based on Bertoloni s.n. (P-03365075); B, C, E–K, M, N based on Merxm¨uller & Wiedmann 19570 (M-0223256); D based on Navarro et al. CN4234 (MA-699433); L based on Bertoloni s.n. (P-03365075) and Navarro et al. CN4234 (MA-699433); O based on Orsini s.n. (P-02320457). 366 Annals of the Missouri Botanical Garden antrorse hairs. Lodicules 0.4–0.7(–0.8) mm, with 2 lobules of the same length, rarely with 3 shallow ones. Anthers (1.4–)2.2–2.3(–2.6) mm. Ovaries 0.5–0.9(–1.1) mm, glabrous. Caryopses ca. 3.2 30.9 mm, oblong. Chromosome number: 2n514 (Favarger, 1973). Additional illustrations. Chrtek & Jir´asek (1963: 577, fig. 3, 3, sub Trisetum villosum); Chrtek (1967c: 176, tab. I, 2; 178, tab. II, 3–4, sub Trisetum villosum); Fiori & Paoletii (1895: 27, 231, sub Trisetum villosum). Phenology. Trisetum bertolonii has been collected in flower and fruit from May to August. Distribution and habitat. This species is endemic to the Apennines. It is found in limestone rocks on mountain cliffs at 170–1630 m. Figure 4. Discussion. Trisetum bertolonii is easily differentiated from the rest of the species of the section by its habit, with long and convolute leaves or with margins inrolled, and a combination of different characters of the spikelets. This species has the longest callus hairs (1.5–2.2[–2.6] mm), upper glumes (6.8–8[–9] mm), and awns ([11–]11.5–16[–17] mm), which are straight or slightly curved at the middle, and lemmas with hyaline margins. Chrtek (1967c) considered Trisetum bertolonii to be closely related to T. griseovirens, from the High Atlas Mountains in Morocco, because of their similar habits, shape and anatomy of the leaves, and awn length. Later, Jonsell (1978) agreed that T. bertolonii is close to, but not conspecific with, T. griseovirens. We consider T. griseovirens to be a subspecies of T. flavescens. The material studied of both species shows some similarities in habit and other characters of the spikelets, which are more pronounced in T. bertolonii.Trisetum bertolonii and T. flavescens subsp. griseovirens have narrow leaves (convolute or filiform and inrolled, respectively), longer aristules (1.2–2.3[–4] mm and [0.9–]1–1.4[–1.8] mm, respectively), longer awns ([11–]11.5–16[–17] mm and [8–]8.2–9[–10.6] mm, respectively), and a more marked difference between the palea and the lemma length (palea length:lemma length ratio of [0.4–]0.58–0.67 [–0.7] and [0.53–]0.65–0.76[–0.77], respectively). However, both taxa are easy to differentiate, because T. bertolonii has longer callus hairs (1.5–2.2[–2.6] mm and 0.4–0.5[–0.7] mm, respectively) and straight or slightly curved awns, whereas T. flavescens subsp. griseovirens has geniculate ones. Additionally, our molecular data (Barber´a et al., unpublished) show that both taxa are not closely related. Thus, we do not agree with Jonsell (1978) who states that T. bertolonii and T. griseovirens may constitute an isolated pair of species with an interesting phytogeographical disjunction. Due to the long aristules of Trisetum bertolonii, the aristules were not taken into account for calculating the ratio of the awn insertion from the base length/lemma length. Additional specimens examined. ITALY. Ascoli Piceno: Monte Vettore, 42°499N, 13°169E, 21 July 1886, Orsini 863 (L); Monte Volubrio, 42°559N, 13°169E, 1847, Bertoloni s.n. (P); Furca del Cristo sub Monte Vettore, 42°499N, 13°169E, 6 July 1856, Caruel s.n. (K); Monte della Sibilla, 42°549N, 13°159E, 1843, Marzialetti s.n. (FI); Monte Vettore, 42°499N, 13°169E, July 1856, Rostan s.n. (BM). L’Aquila: Gran Sasso, Campo Imperatore, Observatory, 42°269N, 13°339E, 3 Aug. 1960, Segelberg s.n. (GB, S). Macerata: in montibus di Camerino, 43°89N, 13°49E, 1831, Bertoloni s.n. (B). Pesaro and Urbino: Gola die Furlo an der Via Flaminia, zw. Fossombrone und Acqualagna, 43°389N, 12°429E, 16 May 1964, Merxm¨uller & Wiedmann 18/64 (M). Pescara: Vado di Sole, 42°239N, 13°479E, 2 July 2002, Navarro et al. 4234 (MA). Teramo: Monte dei Fiori, 42°469N, 13°359E, s. coll. s.n. (H); Monte Corno, 42°289N, 13°339E, Orsini s.n. (FI, P); 1833, Orsini s.n. (P); Monte Corno ad Arapietra, 42°299N, 13°349E, Orsini s.n. (FI). 4. Trisetum flavescens (L.) P. Beauv., Ess. Agrostogr. 88. 1812. Avena flavescens L., Sp. Pl.: 80. 1753. Trisetum pratense Pers., Syn. Pl. 1: 97. 1805, nom. illeg. Trisetaria flavescens (L.) Baumg., Enum. Stirp. Transsilv. 3: 263. 1816. Avena flavescens var. pratensis Neilr., Fl. Nieder.-Oesterreich: 56. 1859, nom. illeg. Avenastrum flavescens (L.) Jess., Deutschl. Gr¨aser: 215. 1863. Trisetum flavescens subsp. pratense Asch. & Graebn., Syn. Mitteleur. Fl. 2: 265. 1899. TYPE: Herb. A. van Royen N° 913. 7-458 (lectotype, designated by Cope in Cafferty et al. [2000: 247], L-221378 image!). Herbs (20.2–)30.9–58.1(–141.5) cm tall, not to slightly caespitose or caespitose, rhizomatous or with stoloniferous rhizome, culm (0.3–)0.6–0.8(–1.2) mm diam., straight, glabrous, rarely sparsely pubescent, with hairs 0.1–0.5 mm; nodes (3 to)4 to 5(to 7), separated along culm, not included in sheaths, glabrous, yellowish to brownish. Basal leaf sheaths sparsely pubescent to pubescent, rarely glabrous, with hairs (0.1–)0.4–1(–1.7) mm, decaying into fibers, yellowish to brownish, sometimes greenish; basal leaf blades (1.4–)3.9–7(–18) cm 3(0.6–)1.5–2.7(–5.6) mm, flat, sometimes convolute, rarely filiform, inrolled when dried, from glabrous to pubescent abaxially, pubescent, rarely glabrous adaxially, with hairs (0.05–)0.1–0.8(–1) mm, usually with cilia, brownish to yellowish, sometimes greenish or grayish; top culm leaf sheaths (4.5–) 7.5–11(–20) cm, glabrous, rarely pubescent, rarely with cilia, usually with hyaline margins; top culm leaf blades (1.3–)4–9(–14.5) cm 3(1–)1.8–3.4(–6) mm, from flat to convolute or filiform, rarely parallel to culm, glabrous to pubescent abaxially, pubescent to densely pubescent adaxially, rarely glabrous, with hairs (0.05–)0.2–1(–1.5) mm, sometimes with cilia, greenish to grayish; inner collar region glabrous, rarely with scattered hairs or Volume 103, Number 3 2018 Barber ´a et al. 367 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa sparsely pubescent, with hairs 0.05–1 mm; ligules (0.5–)0.7–1.2(–2) mm, irregularly dentate to laciniate, rarely entire, glabrous, sometimes with adpressed hairs on surface, glabrous or with hairs (0.05–)0.2–0.4 (–0.6) mm on upper part. Basal node of panicle glabrous, rarely sparsely puberulous, with hairs 0.05–0.2 mm. Panicles (3.5–)6–10.5(–16.2) 3(0.9–) 1.5–2.8(–5) cm, narrowly elliptic or elliptic to narrowly lanceolate or lanceolate in outline, rarely oblong or ovate, lax to somewhat dense; rachis internodes glabrous to sparsely pubescent, mostly on upper part and branches, rarely pubescent, with hairs up to 0.2(–0.3) mm; longest basal branches (0.5–)1.1–2.1(–4) cm. Spikelets (3.8–)5.2–6.3(–7.7) 3(0.8–)1.8–3(–3.8) mm, 2- to 3(4)-flowered; pedicels (1.5–)2.6–3.7(–5) mm, sparsely pubescent, sometimes glabrous or pubescent, with hairs 0.05–0.1(–0.3) mm. Glumes unequal (lower glume length:upper glume length ratio of [0.54–] 0.6–0.7[–0.78]); lower glume (2–)3–3.7(–4.6) 3(0.4–) 0.6–0.7(–1.1) mm, narrowly lanceolate, sometimes broadly lanceolate (lower glume width:lower glume length ratio of [0.1–]0.17–0.22[–0.37]), acuminate, sometimes long-acuminate, rarely acute, 1(3)-nerved, glabrous, usually with short hairs from center to upper part of main nerve, up to 0.1 mm, greenish, rarely purplish or brownish, with hyaline margins; upper glume (3.2–)4.6–5.3(–7.4) 3(1.1–)1.5–1.8(–2.2) mm, elliptic, usually oblong or broadly lanceolate, sometimes slightly oblanceolate (upper glume width: upper glume length ratio of [0.25–]0.3–0.36[–0.47]), acuminate to long-acuminate, 3-nerved, glabrous, usually with short hairs from center to upper part of central nerve, with hairs up to 0.1 mm, greenish, rarely purplish, usually with hyaline margins; rachilla segment between first and second florets (0.6–)0.8–1.1(–1.3) mm, with hairs (0.2–)0.6–1.1(–2.1) mm; rachilla segment to sterile floret (0.7–)1–1.2(–1.5) mm, with hairs (0.2–)0.4–0.7(–1.2) mm. Lemmas (3.2–)4.4–5.4 (–7.4) 3(0.4–)0.6–0.7(–1.1) mm, narrowly to broadly lanceolate, sometimes oblong or elliptic, rarely oblanceolate (lemma width:lemma length ratio of [0.19–] 0.23–0.3[–0.43]), glabrous to scabridulous, with short hairs up to 0.1 mm on upper part and along midrib, greenish to yellowish on central part, sometimes with purple; apical teeth (0–)0.2–0.3(–0.6) mm, with aristules (0.1–)0.4–0.7(–1.8) mm; awn (4.2–)5.7–7.2 (–10.6) mm, inserted (1.5–)2.4–3.3(–4) mm from base (awn insertion from base length:lemma length ratio of [0.45–]0.54–0.59[–0.67]), geniculate to slightly geniculate, 6twisted at base, rarely not twisted, with hairs up to 0.05 mm; callus (0.1–)0.2 mm, orbicular, sometimes rounded or elliptic, with hairs (0.1–)0.2–0.4(–0.7) mm. Paleae (2.2–)3.4–4.2(–5.5) 3(0.6–)0.8–1(–1.3) mm (palea length:lemma length ratio of [0.53–]0.72–0.82 [–0.95]), elliptic to narrowly elliptic, sometimes slightly oblanceolate, margins with hairs on upper part, sometimes glabrous; teeth (0.05–)0.2–0.3(–0.6) mm, with short, antrorse hairs mostly on upper part, rarely glabrous. Lodicules (0.3–)0.4–0.6(–0.9) mm, with (1 to) 2 to 3 teeth or lobules of equal or different depths, sometimes entire. Anthers (1–)1.7–2.2(–3) mm. Ovaries (0.3–)0.4–0.7(–1.1) mm, glabrous. Caryopses (2.3–) 2.4–2.9(–3.7) 30.3–0.5 mm, narrowly elliptic, sometimes linear. KEY TO THE SUBSPECIES OF TRISETUM FLAVESCENS 1. Lemmas (3.2–)4.3–5.1(–7.4) mm long; aristules (0.1–)0.3–0.6(–1.4) mm long; awns (4.2–)5.6–7(–8.3) mm long . . . ................................................................ T. flavescens (L.) P. Beauv. subsp. flavescens 19. Lemmas 6–6.2(–6.4) mm long; aristules (0.9–)1–1.4(–1.8) mm long; awns (8–)8.2–9(–10.6) mm long . . . . . . . . . . . ....................................................... T. flavescens subsp. griseovirens (H. Lindb.) Dobignard 4a. Trisetum flavescens subsp. flavescens. Avena purpurascens DC., Cat. Pl. Horti Monsp.: 82. 1813. Trisetum flavescens subsp. purpurascens (DC.) Arcang., Comp. Fl. Ital.: 779. 1882. Trisetum flavescens subvar. purpurascens (DC.) J.Zimm.inHegi,Ill.Fl.Mitt.-Eur.1,ed.2:337.1936. Trisetum flavescens f. purpurascens (DC.) Borza, Consp. Fl. Roman. 1: 24. 1947. Trisetaria flavescens subsp. purpurascens (DC.) Banfi & Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat. Milano 135: 383. 1996. TYPE: France. Savoie, Cenis Mtn., 45°159N, 6°549E, 1809, J. L. Bonjean s.n. (lectotype, designated here, G-DC-00418509 image!). Trisetum splendens C. Presl, Cyper. Gramin. Sicul.: 30. 1820. Avena splendens (C. Presl) Guss., Fl. Sic. Prodr. 1: 126. 1827. Trisetum flavescens var. splendens (C. Presl) Parl., Fl. Ital. 1: 261. 1848. Trisetum flavescens subsp. splendens (C. Presl) Arcang., Comp. Fl. Ital.: 779. 1882. Trisetaria flavescens subsp. splendens (C. Presl) Banfi & Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat. Milano 135: 383. 1996. TYPE: Italy. Sicily, San Martino, 37°599N, 15°99E, June 1817, C. Presl s.n. (lectotype, designated here, PRC-450870 image!; possible isolectotypes, PRC-450868 image!, W-1889-0242136!). Trisetum flavescens var. biflorum Gray, Nat. Arr. Brit. Pl. 2: 129. 1821. TYPE: Great Britain. S. F. Gray s.n. (no original material found). Avena flavescens var. depauperata Mert. & W. D. J. Koch, Deutschl. Fl., ed. 3, 1: 563. 1823. Trisetum flavescens var. depauperatum (Mert. & W. D. J. Koch) R. Uechtr., Fl. Schlesien: 512. 1881. TYPE: Germany. Alps, F. K. Mertens & W. D. J. Koch s.n. (LE?) (no original material found). Avena flavescens var. variegata Gaudin ex Mert. & W. D. J. Koch in R¨ohl., Deutschl. Fl. 1, ed. 3, 1: 562. 1823. Trisetum flavescens var. alpinum Parl., Fl. Ital. 1: 261. 1850, nom. illeg. Trisetum flavescens var. variegatum (Gaudin ex Mert. & W. D. J. Koch) Schur, Enum. Pl. Transsilv.: 758. 1866. Trisetum flavescens [b]variegatum (Gaudin ex Mert. & W. D. J. Koch) Asch. & Graebn., Syn. 368 Annals of the Missouri Botanical Garden Mitteleur. Fl. 2: 266. 1899. Trisetum flavescens f. variegatum (Gaudin ex Mert. & W. D. J. Koch) Beetle, Phytologia 38(3): 176. 1978. TYPE: Switzerland. J. F. Gaudin s.n. (lectotype, designated here, L-1346419!). Avena flavescens var. major Spenn., Fl. Friburg. 1: 108. 1825. TYPE: Germany. Friburg, F. C. L. Spenner s.n. (FB?, original material not located). Avena flavescens var. capillacea Gaudin, Fl. Helv. 1: 337. 1828. Trisetum flavescens var. capillaceum (Gaudin) Ducommun, Taschenb. Schweiz. Bot.: 857. 1869. TYPE: Switzerland. Vaud, Chˆateaux d’Oex, June & July, J. F. Gaudin s.n. (original material not located). Avena flavescens var. maior Gaudin, Fl. Helv. 1: 337. 1828, nom. illeg., non Spenn., Fl. Friburg. 1: 108. 1825. Trisetum flavescens var. latifolium Schur, Enum. Pl. Transsilv.: 758. 1866, nom. illeg. TYPE: Switzerland. J. F. Gaudin s.n. (original material not located). Avena flavescens var. lutescens Rchb., Fl. Germ. Excurs. 1: 51. 1830. Trisetum flavescens var. lutescens (Rchb.) Druce, Fl. Oxfordshire, ed. 2: 478. 1927. Trisetum flavescens subvar. lutescens (Rchb.) Maire, Cat. Maroc.: 48. 1931. Trisetum flavescens f. lutescens (Rchb.) Buia & Morariu, Fl. Republ. Socialiste Romˆania 12: 292. 1972. TYPE: (lectotype, designated here, Rchb., Icon. Fl. Germ. Helv. 1, tab. XCIX, fig. 1695. 1834). Trisetum burnoufii Req. ex Parl., Fl. Ital. 1: 263. 1850. Avena burnoufii (Req. ex Parl.) Nyman, Syll. Fl. Eur.: 413. 1855. Trisetum flavescens var. burnoufii (Req. ex Parl.) Hack in Briq., Prodr. Fl. Corse 1: 104. 1910. Trisetum flavescens subsp. burnoufii (Req. ex Parl.) P. Fourn., Quatre Fl. France, ed. 2: 60. 1946. Trisetaria burnoufii (Req. ex Parl.) Banfi & Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat. Milano 135: 382. 1996. TYPE: France. Corse, Haute-Corse, Corte, 42°189N, 9°89E, 7 Oct. 1847, E. Requien s.n. (lectotype, designated by Signorini & Ricceri [1996: 215], FI not seen; isolectotypes, G-00100184!, P-03365150!). Avena candollei M. Serres, Bull. Soc. Bot. France 4: 440. 1857. Trisetum candollei (M. Serres) B. Verl., Cat. Pl. Vasc. Dauphin´e: 365. 1872. TYPE: France. Hautes-Alpes, Gap, 44°349N, 6°59E, B. Blanc s.n. (original material not located). Trisetum flavescens var. glabratum Asch., Fl. Brandenburg 1: 830. 1864. TYPE: Germany. Brandenburg, s. coll. (original material not located). Trisetum flavescens var. biflorum Schur, Enum. Pl. Transsilv.: 757. 1866, nom. illeg., non Trisetum flavescens var. biflorum Gray, Nat. Arr. Brit. Pl. 2: 189. 1821. TYPE: Romania. Kronstadt [Brasxov], P. J. F. Schur s.n. (original material not located). Trisetum flavescens var. triflorum Schur, Enum. Pl. Transsilv.: 758. 1866. TYPE: Romania. Sibiu [Hermannstadt], 45°489N, 24°99E, June 1852, P. J. F. Schur s.n. (lectotype, designated here, LW-213271 image!). Trisetum bungei Boiss., Fl. Orient. 5: 535. 1884. TYPE: Iran. Golestan, Ziarat, 36°419N, 54°319E, 13 May 1858, A. A. v. Bunge 68 (lectotype, designated here, P-03644568!; isolectotypes, B not seen, K image!). Trisetum pratense var. villosum ˇ Celak., Sitzungsber. K¨onigl. B¨ohm. Ges. Wiss. Prag., Math.-Naturwiss. Cl. 1887: 180. 1887. Trisetum flavescens var. villosum ( ˇ Celak.) Fiek, Jahresber. Schles. Ges. Vaterl. Cult. 67: 167. 1889. TYPE: Czech Republic. Pt´aˇckovna, P´ısek, 49°199N, 14°109E, H. Ciboch s.n. (original material not located). Trisetum flavescens var. nodosum Chabert, Bull. Soc. Bot. France 36: 31. 1889. TYPE: Algeria. Nador supra Medeah, 36°159N, 2°459E, 1871–1875, A. C. Chabert s.n. (FI?, original material not located). Trisetum flavescens var. tenue Hack. ex Form´anek, Verh. Naturf. Vereins Br¨ unn 35: 155. 1897. Trisetum tenue (Hack. ex Form´anek) Chrtek, Acta Univ. Carol., Biol. 1971: 228. 1973, nom. illeg., non Trisetum tenue Roem. & Schult., Syst. Veg. (ed. 15 bis) 2: 657. 1817. Trisetum tenuiforme Jonsell, Bot. J. Linn. Soc. 76: 320. 1978. Trisetum flavescens subsp. tenue (Hack. ex Form´anek) Strid, Mount. Fl. Greece 2: 808. 1991. Trisetaria tenuiformis (Jonsell) Banfi & Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat. Milano 135: 385. 1996. TYPE: Greece. Thessalia, Agrapha, Monoster Korona, 39°89N, 21°389E, 1 Aug. 1896, E. Form´anek s.n. (lectotype, designated by Jonsell [1978: 320], W-1916-0031810!). Trisetum flavescens var. ponticum Velen., Fl. Bulg., Suppl. 1: 295. 1898. TYPE: Bulgaria. Burgas, 42°309N, 27°289E, 1893, S. Jablonowsky s.n. (lectotype, designated here, PRC-451827 image!). Trisetum flavescens var. serbicum Velen., Fl. Bulg., Suppl. 1: 295. 1898. Trisetum flavescens subsp. serbicum (Velen.) Hayek, Repert. Spec. Nov. Regni Veg. Beih. 30(2): 312. 1933. TYPE: Serbia. Vranja, 42°339N, 21°549E, 1898, L. L. Adamovi´cs.n. (lectotype, designated here, PRC-451825 image!). Avena flavescens f. major Hausskn., Mitth. Th¨uring. Bot. Vereins 13: 42. 1899, nom. illeg., non Avena flavescens var. major Spenn., Fl. Friburg. 1: 108. 1825. TYPE: Greece. Thessaly, Pharsalus, 39°179N, 22°239E, June 1885, C. Haussknecht s.n. (lectotype, designated here, JE-18858!). Avena flavescens var. pindica Hausskn, Mitth. Th¨uring. Bot. Vereins 13: 43. 1899. TYPE: Greece. Monte Karava, 39°189N, 21°339E, July 1885, C. Haussknecht s.n. (lectotype, designated here, JE-00018850!; isolectotype, JE-00018851!). Trisetum corsicum Rouy, Rev. Bot. Syst. G´eogr. Bot. 1: 139. 1904. Trisetum flavescens var. corsicum (Rouy) Briq., Prodr. Fl. Corse 1: 103. 1910. Trisetum flavescens subsp. corsicum (Rouy) Cif. & Giacom., Nomencl. Fl. Ital., 1: 30. 1950. TYPE: France. Corse, Corte, 42°189N, 9°99E, July 1898, J. Foucaud 924 (lectotype, designated by Signorini & Ricceri [1996: 216], LY not seen; isolectotypes, G- 00100185!, P-319106!, P-2217837!, P-2221569!). Trisetum flavescens var. paczoskii Zapał., Bull. Int. Acad. Sci. Cracovie, Cl. Sci. Math. 3: 167. 1904. TYPE: Tatra Mtns., J. K. Paczoski s.n. (KRAM?; original material not located). Trisetum flavescens f. scabriusculum Zapał., Rozpr. Wydz. Mat.- Przyr. Akad. Umiejetn., Dzia B, Nauki Biol. 4: 107. 1904. TYPE: Ukraine. Dublyani, 49°559N, 24°99E, J. K. Paczoski s.n. (KRAM?, original material not located). Trisetum flavescens f. subpilosum Zapał., Rozpr. Wydz. Mat.- Przyr. Akad. Umiejetn., Dzia B, Nauki Biol. 4: 107. 1904. TYPE: Poland. Lesser Poland, Wola Batorska, 50°039N, 20°159E, J. Krupa s.n. (KRAM?, original material not located). Trisetum flavescens var. barcinonense Sennen, Bull. G´eogr. Bot. 23: 50. 1913. TYPE: Spain. Barcelona Prov., Tibidabo Mtns., Santa Cruz de Olorde, 41°259N, 2°069E, June 1911, F. Sennen 1238 (lectotype, designated here, MA-7940!; isolectotypes, B-10_0470051!, BCN-82824!, DAO-000465368 image!, FR-0036078 image!, JE- 00021246!, JE-00021247!, JE-00021248!, JE-00021249!, MPU!, NY!, SEV-6701!, W-19120005494 image!, W-19230017014 image!, W-20080017999 image!, WU!). Volume 103, Number 3 2018 Barber ´a et al. 369 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa Trisetum handelii Vierh., Oesterr. Bot. Z. 73: 128. 1924. TYPE: Italy. Veneto, Ampezzo, 46°259N, 12°479E, 1905–1907, F. K. M. Vierhapper s.n. (lectotype, selected by W. Till in sched., designated here, WU-0073386 image!). Trisetum flavescens var. africanum H. Lindb., Acta Soc. Sci. Fenn., Ser. B, Opera Biol. 1(2): 25. 1932. Trisetum flavescens subsp. africanum (H. Lindb.) Dobignard, J. Bot. Soc. Bot. France 28: 44. 2004. TYPE: Morocco. Marrakesh-Safi, Amizmiz, 31°139N, 8°139W, 24 May 1926, H. Lindberg 2719 (lectotype, designated by V¨are [2012: 89], H-1168497!; isolectotypes, CAI-000243 image!, H-1168500!). Trisetum flavescens var. clausonii Maire, Bull. Soc. Hist. Nat. Afrique N. 23: 219. 1932. TYPE: Algeria. Djelfa Prov., Kef Roumia, 34°239N, 3°109E, 30 May 1855, Th. Clauson s.n. (lectotype, designated here, MPU-3030!; isolectotype, P-83440!). Trisetum flavescens f. hirticulme H. Lindb., Acta Soc. Sci. Fenn., Ser. B, Opera Biol. 1(2): 25. 1932. TYPE: Morocco. Marrakesh-Safi, Asni, Rera¨ıa River valley, 31°109N, 7°549W, 2 June 1926, H. Lindberg 3483 (lectotype, designated by V¨are [2012: 89], H-1168498!). Trisetum macratherum Maire & Trab., Bull. Soc. Hist. Nat. Afrique N. 23: 218. 1932. Trisetum flavescens var. macratherum (Maire & Trab.) Maire & Weiller, Fl. Afrique N. 2: 247. 1953. Trisetum flavescens subsp. macratherum (Maire & Trab.) Dobignard, J. Bot. Soc. Bot. France 28: 44. 2004. TYPE: Algeria. Near Oran, 35°419N, 0°379W, L. C. Trabut s.n. (lectotype, designated here, MPU-3029!). Trisetum flavescens subsp. tatricum Chrtek, ˇ Cas. N´ar. Mus., Odd. Pˇr´ır. 35(2): 81. 1966. TYPE: Slovakia. Belansk´e Tatry, Bujaci, 49°139N, 20°169E, 15 Aug. 1933, K. Domin s.n. (holotype, PRC-454454 image!). Trisetum flavescens subsp. parvispiculatum Tzvelev, Spisok Rast. Gerb. Fl. S.S.S.R. Bot. Inst. Vsesojuzn. Akad. Nauk 18: 6. 1970. Trisetum parvispiculatum (Tzvelev) Prob., Bot. Zhurn. (Moscow & Leningrad) 60: 674. 1975. TYPE: Russia. Dagestan Republic, Makhachkala, Tarkitau Mtn., 42°569N, 47°279E, 17 May 1953, J. I. Prokhanov 407 (holotype, LE-01042366!). Herbs (20.2–)36.4–63.1(–141.50) cm tall. Basal leaf blades (1.5–)4.2–7(–18) cm 3(0.6–)1.5–2.8(–5.6) mm, flat, sometimes convolute, inrolled when dried. Top culm leaf blades (1.6–)4.9–9.2(–14.5) cm 3(1–) 2–3.5(–6) mm, from flat to convolute, rarely filiform or parallel to culm. Panicles (3.6–)6.9–10.6(–16.2) 3 (0.9–)1.6–2.9(–5) cm. Spikelets (3.8–)5–6.1(–7.6) mm. Lemmas (3.2–)4.3–5.1(–7.4) mm; aristules (0.1–)0.3–0.6(–1.4) mm; awn (4.2–)5.6–7(–8.3) mm. Palea length:lemma length ratio of (0.66–)0.73–0.83 (–0.95). Anthers (1–)1.6–2.1(–2.5) mm. Chromosome numbers: 2n512 (Sokolovskaya & Probatova, 1975), 24 (Frey, 1992), 28 (Sorokin, 1990), 36 (Strid & Andersson, 1985). Figures 8A, 9A, 10A, and 11A. Additional illustrations. Fiori & Paoletii (1895: 28, 234 sub Trisetum burnoufii, 235, sub T. flavescens); Chrtek & Jir´asek (1963: 577, figs. 3, 4); Sãvulescu (1972: 289, Plansxa 56, 2a–c); Frey (1992: 453, fig. 6); Signorini & Ricceri (1996: 219, fig. 1, sub Trisetum corsicum); Gabrielian (2010: 176, tab. 44, 1, 1a). Phenology. Trisetum flavescens subsp. flavescens has been collected in flower and fruit from May to October. Distribution and habitat. Trisetum flavescens subsp. flavescens is widespread throughout Europe, extending to western Asia and northern Africa, and it has been introduced in many other temperate zones in Asia, North and South America, and New Zealand; on meadows, forest clearings, among shrubs, or on stony places, growing from sea level to 2900 m. Figure 12. Discussion. Trisetum flavescens is one of the most polymorphic and widespread taxa of the genus. The morphology and hairiness of leaves, as well as the shape and color of the panicles, are highly variable. This species is characterized as having short ligules ([0.5–]0.7–1.2[–2] mm) and short callus hairs ([0.1–] 0.2–0.4[–0.7] mm), narrow lower glumes ([0.4–]0.6–0.7 [–1.1] mm), and always geniculate awns and glabrous ovaries. Barber´a et al. (2017b) pointed out that it frequently has been confused with T. sibiricum, which belongs to Trisetum sect. Sibirica. Due to the polymorphism of Trisetum flavescens, many subspecies and varieties have been described by different authors to classify its morphological variation. Presl (1820) described T. splendens, with flat leaves and contracted golden panicles, which was later reduced to the rank of subspecies by Arcangeli (1882); this was also accepted as such by many authors (Jonsell, 1980; Pignatti, 1982; Signorini & Ricceri, 1996). In the same way, T. purpurascens, with wider leaves, slightly longer ligules, and dense, purplish spikelets, was described by de Candolle (1813) from the Alps and then reduced to subspecies by Arcangeli (1882) and, at that rank, accepted by Jonsell (1980), Frey (1992; subvariety purpurascens), and Tison and de Foucault (2014). Strid and Tan (1991) indicated that the differences between T. flavescens subsp. splendens and T. flavescens subsp. flavescens are rather obscure. Chrtek (1966) described T. flavescens subsp. tatricum specimens from the Carpathian Mountains, but these are hardly distinguishable from T. flavescens subsp. purpurascens. Three additional species have been described because of their more or less pubescent and convolute leaves and culm hairiness, namely T. tenuiforme,T. burnouffi, and T. corsicum.Trisetum burnouffi and T. corsicum are endemic to Corsica, the latter with intermediate characters between T. burnouffi and T. flavescens. Signorini and Ricceri (1996) studied these species, including T. splendens, and accepted all these taxa at the species rank. Later, Jeanmonod and Gamisans (2007) 370 Annals of the Missouri Botanical Garden recognized T. corsicum as a subspecies of T. flavescens, and T. burnouffi as a variety of this subspecies. Apart from these taxa from the Mediterranean region and Europe, other taxa were described from Africa and Asia. Tzvelev (1970) published T. flavescens subsp. parvispiculatum, with small spikelets and lemmas, from the Caucasus and mountain ranges of Iran and Turkmenistan. The detailed study of the morphology of the specimens belonging to this taxon indicates that these supposed diagnostic characters are not exclusive to them, and that they also occur indiscriminately in other specimens from Europe and the Mediterranean region. Maire and Trabut (Maire, 1932) and Lindberg (1932) described other species and varieties from northern Africa, namely Trisetum macratherum,T. flavescens var. africanum,andT. griseovirens. Chrtek (1967c) considered that T. griseovirens was the only one well characterized, mainly by the characters of the spikelet (length of spikelet, lemma, and awn). However, Dobignard (2004) treated these three taxa as subspecies, making clear that he did so only to reflect intraspecific variability of T. flavescens and awaiting a more rigorous taxonomic revision in the future. As has already been mentioned, the leaf blade traits of Trisetum flavescens and its associated taxa are highly variable, much more so than the characters of the spikelets and, therefore, we recommend giving them a secondary diagnostic value. Consequently, we Figure 8. Transverse section of leaf blade. —A. Trisetum flavescens (L.) P. Beauv. subsp. flavescens.—B. Trisetum flavescens subsp. griseovirens (H. Lindb.) Dobignard. —C. Trisetum fuscum Schult. A based on Aedo et al. 8571 (MA-700192); B based on Jury et al. 8847 (MA-391300); C based on Rechinger s.n. (S). Figure 9. Portion of sheath, ligule, and blade. —A. Trisetum flavescens (L.) P. Beauv. subsp. flavescens.—B. Trisetum flavescens subsp. griseovirens (H. Lindb.) Dobignard. —C. Trisetum fuscum Roem. & Schult. —D. Trisetum laconicum Boiss. & Orph. A based on D¨oring & Holtz s.n. (GOET); B based on Cabezas et al. FJC419 (MA-746349); C based on Ny´ar´ady s.n. (L-1346389); D based on Orphanides 2890 (B-10_0508993). Volume 103, Number 3 2018 Barber ´a et al. 371 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa decided to synonymize all these taxa with more labile foliar morphology, and to maintain only two subspecies in T. flavescens,T. flavescens subsp. griseovirens and the typical subspecies. Vegetative propagation by pseudoviviparism has been noted in a specimen of Trisetum flavescens subsp. flavescens collected by de Wilde (de Wilde s.n.,L- 1346507) (Fig. 13). Until now, this type of propagation has been observed only in T. rigidum (M. Bieb.) Roem. & Schult., T. tamonanteae Marrero Rodr. & S. Scholz, and T. velutinum Boiss., all belonging to Trisetum sect. Acrospelion (Marrero Rodr´ıguez & Scholz, 2013; Barber´a et al., 2017a). Additional specimens examined. ALBANIA. Gjirokast¨er: Mali Lunxheris, N-H¨ange, 40°49N, 20°89E, 13 July 1980, Krendl s.n. (C); Nemerçka, ¨uber Permet, s¨udwestlich vom Qafa e Dhembelit, 40°129N, 20°209E, 12 July 1959, Meyer s.n. (JE). Korç¨e: near Ersek¨e, 40°209N, 20°409E, 29 June 1933, Alston & Sandwith 1994 (K). Vlor¨e: Mali i Gjer (Mali Gjinezh), Delvina, zwischen Qafa Gradishtit und Qafa Piloit, 40°09N, 20°59E, 26 June 1959, Meyer s.n. (JE). ALGERIA. Algiers: `a Maison Carr´ee, pr`es d’Alger, 36°439N, 3°89E, 31 May 1930, Faurel s.n. (P). Batna: Pic des C´edres, 35°339N, 6°29E, 6 July 1935, Faurel s.n. (P). Blida: `a Chrea, 36°259N, 2°529E, 5 July 1930, Faurel s.n. (P). Bou¨ıra: environs de Tala Rana, versant meridional du Lalla Khelidja, 36°259N, 4°139E, 30 June 1875, Duhamel s.n. (G). Constantine: Djebel Cougour pr`es de Batna, 35°339N, 6°109E, 22 May 1899, Cosson s.n. (L). Oran: pente sud du Djebel-Santo, 35°449N, 0°439W, 11 June 1852, Balansa s.n. (L). S´etif: Massif de Bou-Talef, pr`es de Char`ene, 35°459N, 5°199E, 21 May 1937, Faurel s.n. (P). Tiaret: Massif de B´elezma, de Bocdem, 35°139N, 0°459E, 16 June 1937, Faurel s.n. (P). Tizi Ouzou: Grande Kabylie, National Park Djurdjura, bei Tala Guilef, 36°279N, 4°139E, 30 June 1983, Scholz & Baillargeon s.n. (B). Tlemcen: au-dessus de Tlemcen, 34°529N, 1°189W, 19 June 1856, Bourgeau s.n. (UPS). ANDORRA. Pr. Soldeu, 42°349N, 1°409E, 29 Aug. 2002, Aedo et al. 8571 (MA). ARMENIA. Aragatsotn: monte Aragat, Ghazaravan, carretera de subida al lago Kari, 40°239N, 44°159E, 30 June 2005, Medina et al. 2606 (MA); Ashtarak Distr., in declivibus montis Arailer, in vicinitate pagi Egvart, 40°249N, 44°289E, 15 July 1975, Vasak s.n. (B, G). Gegharkunik: pr. Sevan, 40°359N, 44°579E, 19 June 2005, Aedo et al. 11722 (MA); Vardenis Distr., montis Vardenisskij Khrebet, in declivibus montis Kungukh-tar, 40°109N, 45°439E, 29 July 1975, Vasak s.n. (B, G, MA, NY). Kotayk: pr. Tsahkadzor village, Tehenis Mtn., 40°329N, 44°419E, 22 June 2005, Castroviejo et al. 17687 (MA). Lori: Akhtala, above village Ghochkan (Tchotchkan), 41°119N, 44°499E, 24 June 2009, Vitek et al. 09-1870 (B). Shirak: Akhuryan Distr., Shirak range, N of the Dzhadzhur pass, 40°519N, 43°599E, 23 June 1960, Tsvelev & Cherepanov 215 (LE). Syunik: Meghri Distr., on a slope to a right tributary of the Mergiget River, near Vardanadzor, 38°589N, 46°129E, 29 June 1956, Egorova et al. 1512 (LE); Pirdaudan, 39°159N, 46°89E, 23 June 1893, Lipskiy s.n. (LE). Tavush: Haghartsin monastery, 40°499N, 44°539E, 21 June 2005, Castroviejo et al. 17555 (MA). Vayots Dzor: pr. Ger-Ger, 39°479N, 45°329E, 24 June 2005, Herrero et al. 2841 (MA). AUSTRIA. Carinthia: Mauthen im Gailthal, 46°399N, 12°599E, 22 June 1934, Ronninger s.n. (NY). Lower Austria: St. Poelten, 48°129N, 15°389E, June 1879, Hackel s.n. (B, PR); Gloggnitz, 47°409N, 15°569E, 28 June 1861, Hal´acsy s.n. (WU); Semmering ad stationem in prato, 47°389N, 15°499E, 29 June 1922, Vestergren s.n. (NY, S). Salzburg: Gaisberg, 47°489N, 13°69E, 22 May 1922, Pettersson s.n. (H). Styria: in ruderatis prope urbem Graz, 47°49N, 15°279E, June 1808, Fritsch s.n. (H, L, MA, O, WU). Tyrol: Obergurgl, 46°529N, 11°19E, 3 Aug. 1992, Emanuelsson 1319 (S); im Figure 10. Upper and lower glume, dorsal view. —A. Trisetum flavescens (L.) P. Beauv. subsp. flavescens.—B. Trisetum flavescens subsp. griseovirens (H. Lindb.) Dobignard. —C. Trisetum fuscum Schult. —D. Trisetum laconicum Boiss. & Orph. A based on Barber´a & Quintanar 897PB (MA-871565); B based on Cabezas et al. FJC419 (MA-746349); C based on Ny´ar´ady s.n. (L-1346389); D based on Orphanides 2890 (B-10_0508993). 372 Annals of the Missouri Botanical Garden Paznauntal bei Galt¨ur am Eingang des Jamtales, 46°589N, 10°119E, 27 July 1978, Holtz 03.892 (GOET). Upper Austria: ca. Aistersheim, 48°119N, 13°449E, June 1875, Keck s.n. (B, F, G, GOET, H, MO, MW, NY, WU); Lichtenberg, N v. Ort bei Kaiserberg 5, 48°229N, 14°159E, 18 June 1995, Kleesadl 178 (W, WU). Vienna: Neuwaldegg, 48°149N, 16°179E, 27 July 1921, Fiedler s.n. (B). Vorarlberg: Illufer unter Feldkirch, 47°139N, 9°369E, 3 Aug. 1869, Boetz s.n. (W). AZERBAIJAN. Daglig-Shirvan: rd. Shemakha–Aksu, 40°349N, 48°249E, 1 June 1949, Beydeman s.n. (LE). Ganja-Gazakh: Dashkasan Distr., Khachbulag, 40°289N, 46°09E, 16 July 1970, Lyatifova 24 (LE). Lankaran: a pass betw. Lerik & the village of Bozuchar, 38°469N, 48°249E, 3 July 1931, Shipchinskiy 565 (LE). BELGIUM. East Flanders: Gent, “Snepkaai,”51°39N, Figure 11. Florets. —A. Trisetum flavescens (L.) P. Beauv. subsp. flavescens.—B. Trisetum flavescens subsp. griseovirens (H. Lindb.) Dobignard. A based on Barber´a & Quintanar 897PB (MA-871565); B based on Cabezas et al. FJC419 (MA-746349). Volume 103, Number 3 2018 Barber ´a et al. 373 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa 3°439E, 14 June 1957, Michiels s.n. (C). Li`ege: Ardennen, tussen Eupen en Baraque Michel, 50°319N, 6°39E, June 1950, Borssum Waalkes 5114 (L). Limburg: prairies `a Maeseyck, 51°59N, 5°479E, 24 June 1871, Cogniaux s.n. (NY). Namur: Rochefort, Croise St. Jean, 50°99N, 5°149E, 15 June 1934, Mosseray-Staner s.n. (C). Walloon Brabant: Waterloo, 50°429N, 4°239E, 1 July 1923, Lambert s.n. (US). BOSNIA AND HERZEGOVINA: Homami do pr. cast. Jablan, 43°329N, 18°89E, Sep. 1910, Vandas s.n. (PR). BULGARIA. Plovdiv: Central Rhodopes Mtns., pr. Kosovo, 41°549N, 24°429E, 29 June 2004, Aedo et al. 10371 (MA). Smolyan: Central Rhodope Mtns., Smolyan Distr., Gela village, 41°399N, 24°339E, 28 June 2004, Aedo et al. 10141 (MA). CROATIA. Zagreb Co.: Zagreb, 24 km E of Zagreb, by the rd. Zagreb–Beograd, 45°489N, 15°589E, 13 June 1971, Suominen 472 (H). CZECH REPUBLIC. Central Bohemian: Pˇr´ıbram Distr., na luk´ach pred Novou Hospodou [Nov´a Hospoda], 49°419N, 14°29E, June 1904, Tocl s.n. (G, H). South Moravian: Str´aˇznice, Velk´a, locis “na Kuk´ach,”supra V´apenky, 48°529N, 17°379E, 20 June 1930, Podpera et al. s.n. (C, G, H, K, MO, S, UPS, US, WU). ´ Ust´ınad Labem: Dˇeˇc´ın Distr., ˇ Sluknov, 51°09N, 14°279E, 16 June 1964, Chrtek & Pistulkov´a s.n. (G, H, K, L, MA, MO, S, UPS). DENMARK. Capital Region: Bornholm, Rønne ˚ Avangskolen, 55°69N, 14°429E, 17 June 1978, L¨utken s.n. (MO). Central Jutland: Aabyhoj, Ruderat ved Frichs fabrikker, 56°99N, 10°99E, 21 June 1964, Øllgaard s.n. (MO). Region Zealand: Bastrup, 55°499N, 12°169E, 25 June 1944, Lange s.n. (US). FINLAND. ˚ Aland Islands: Maarianhamina, Badhusberget, 60°69N, 19°559E, 21 Sep. 1984, Hæggstr¨om 5224 (H). Lapland: M¨annikk¨o prope opp. Willmanstrand, 66°39N, 26°559E, 12 July 1887, Sælan s.n. (H). Northern Savonia: Kuopio, Vanhan tulitikkentehtaan puisto, 62°539N, 27°409E, 25 July 1931, Toivanen s.n. (H). Ostrobothnia: Vaasa, Vasklot, 63°59N, 21°349E, 28 June 1954, Malmberg s.n. (H). Pirkanmaa: Satakunta, Karkku, 61°269N, 23°29E, 2 Sep. 1870, Hjelt s.n. (H). South Karelia: Lappeenranta, Linnoitus, 61°39N, 28°109E, 20 June 1948, Eurola s.n. (H). Southern Savonia: Savonlinna, Saaminki, Aholahti, 61°539N, 28°459E, 27 June 1920, Linkola s.n. (H). Southwest Finland: Turku, Paimio, Juntola, E bank of Paimio River at the bridge above the power station, 60°279N, 22°419E, 3 July 1982, Kukkonen 11193 (H). Uusimaa: Helsinki, Nordsj¨o, Rastila, 60°129N, 25°79E, 3 July 1964, Ruotsalo s.n. (H). FRANCE. Allier: Arouer, Les Damariats, 46°419N, 3°179E, June 1894, Lassimonne s.n. (C). Alpes-de-Haute-Provence: Lautaret, 44°269N, 6°219E, July 1905, Jeanpert s.n. (F). Alpes- Maritimes: Cipi`eres village, 43°469N, 6°579E, 1 July 1992, Hepper 9464 (B). Ard`eche: Saint-Julien-du-Serre, 44°399N, 4°249E, 5 June 1902, Girod s.n. (G). Ari`ege: Haies `a Pamiers, 43°79N, 1°369E, 12 June 1877, Huet s.n. (UPS). Bouches-du- Rhˆone: Martigues, 43°249N, 5°39E, May 1871, Autheman s.n. (UPS). Charente-Maritime: La Rochelle, 46°109N, 1°99W, 6 June 1955, Galiano s.n. (SEV). Corse: Upper Corsica, Massif du San Petrone, a l’WNW d’A¨ıti, 42°239N, 9°149E, 30 June 1990, Gamisans 14681 (G); Corte, au dessus de Casanova, ravin de Taola, 42°159N, 9°109E, 8 July 1979, Gamisans s.n. (G); Upper Corsica, Calacuccia, 42°209N, 9°09E, 8 July 1911, Petry s.n. (BM, H, O, PR). Cˆotes d’Armor: Cˆote du Nord, Pl´eh´erel, 48°399N, 2°229W, 11 June 1967, Bakker & Beglel s.n. (L). Dordogne: Sainte-Orse, 45°129N, 1°49E, 16 June 1891, Mouillefarine s.n. (US). Haute-Garonne: Bagn`eres-de- Luchon, 42°479N, 0°359E, 6 July 1885, Peyron s.n. (G). Haute-Saˆone: Champagney, 47°429N, 6°409E, 6 June 1868, Vendrely s.n. (JE). Haute-Savoie: Mt. Veyrier, Col Figure 12. Distribution of Trisetum flavescens (L.) P. Beauv. subsp. flavescens (dots) in Eurasia and North Africa. 374 Annals of the Missouri Botanical Garden length ratio of 0.5–0.68[–0.83]), strongly curved basally, not or slightly twisted at base, with adpressed hairs 0.05–0.2 mm, the longest ones at base; callus (0.1–)0.2(–0.3) mm, elliptic to orbicular, with hairs (0.4–)0.5–1.1(–1.2) mm. Paleae (3.6–)3.9–4.5(–5.5) 3 1–1.5(–1.7) mm (palea length:lemma length ratio of [0.77–]0.87–0.95[–1]), lanceolate or elliptic to slightly oblanceolate, margins with hairs from middle to upper part; teeth 0.2(–0.3) mm, with short, antrorse hairs sometimes up to 0.4 mm. Lodicules (0.4–)0.5–0.8 (–0.9) mm, with 2 or 3 lobules or teeth of the same length. Anthers 1.7–2.1(–2.5) mm. Ovaries (0.4–) 0.5–0.8(–1.1) mm, glabrous. Caryopses 1.6–2.8 3 0.5–0.6 mm, oblong to elliptic. Chromosome number: 2n528 (Frey, 1991). Additional illustrations. Chrtek & Jir´asek (1963: 577, fig. 3, 3, sub Trisetum ciliare); Chrtek (1965: 213, Abb. 1, 2; 214, Abb. 3, 1; 217, Abb. 4, 1); Sãvulescu (1972: 289, Plansxa 56, 3a–c, sub T. ciliare); Frey (1992: 464, fig. 12). Phenology. Trisetum fuscum has been collected in flower and fruit from June to October. Distribution and habitat. This species is endemic to the Carpathian Mountains; it grows on scree of mountain meadows and floodplains in wet rock fissures, at 1500–2000 m. Figure 16. Discussion. Trisetum fuscum is differentiated from the rest of species of the section mainly by its longer ligules ([3–]3.7–4.7[–4.8] mm). Trisetum fuscum is similar to T. alpestre, with which it frequently has been confused. Apart from the length of the ligules, T. fuscum is separated from T. alpestre by its larger size ([26.8–] 44.9–54[–72] cm vs. [14.4–]21.5–35.7[–53.5] cm) and not or more loosely tufted habit, longer and wider top culm leaf blades ([5.7–]6.5–12.5[–13.7] cm 3[2.1–] 3.4–5[–6.1] mm vs. [1.5–]2.2–5.2[–9.3] cm 3[1–] 1.6–3[–4.1] mm), and longer panicles ([6.8–]7.9–11.2 [–13] mm vs. [3.4–]5.4–7.5[–11.5] mm). Concerning spikelet characters, T. fuscum has wider upper glumes than T. alpestre ([1.8–]2–2.4[–2.6] mm vs. [1.5–]1.8–2 [–2.3] mm), awns strongly curved basally and sometimes with long hairs at the base (up to 0.2 mm), paleae sometimes also with longer hairs (up to 0.4 mm apically), and glabrous ovaries. Trisetum fuscum is easy to differentiate from T. flavescens, with which it sometimes has been confused. The ligules are significantly shorter in T. flavescens ([0.5–]0.7–1.2[–2] mm). Some authors, such as Chrtek (1968), Jonsell (1980), and Frey (1992), give much taxonomic importance to the hairiness of the lemma and the palea to differentiate T. fuscum from T. alpestre and T. flavescens. In particular, Chrtek (1968) published a detailed morphological study of T. fuscum, comparing each of those characters among the three species. In our study of T. fuscum, the hairiness of both the palea and the lemma exhibited great variability in length and density, suggesting that this feature is of minor taxonomic utility. Jonsell (1980) described the ovary of Trisetum fuscum as having a few apical hairs. Later, Frey (1992) also observed two to three long papillae (atypical hairs) on the top of the ovary of T. fuscum. However, in all the specimens of T. fuscum studied by us, the ovaries were glabrous. Arduino (1789) proposed the name Avena fusca in an illustration (tab. 4) for his A. hispanica, which was described in the same publication (pp. 111–113). Thus, A. fusca should be considered as a valid published name (alternative name), according to Article 36.2 of the ICN (McNeill et al., 2012). Consequently, A. fusca Kit. ex Schult. is a later homonym (Art. 53.1), spelled exactly like Arduino’s name for a taxon of the same rank, but based on a different type. We have decided to conserve the epithet “fuscum”as a replacement name with the same type as the illegitimate name in order to avoid disadvantageous nomenclatural changes. Additional specimens examined. POLAND. Lesser Poland: Morskie Oko, 49°119N, 20°49E, 19 July 1929, Nilsson s.n. (S); Kociol pod Rysami, Rysy, 49°119N, 20°49E, 20 Aug. 1931, Pawlowski s.n. (GB, GH, K, LE, S, UPS). SLOVAKIA. Bansk´a Bystrica: Djumbier [ ˇ Dumbier], 48°569N, 19°389E, Gustav s.n. (MO). Preˇsov: Mlynica, 49°69N, 20°189E, 29 July 1966, Dahl & Hadac s.n. (O); Csorbam, J´egt´oi v¨olgy valley & Furkota v¨olgy, 49°109N, 20°19E, 9 Aug. 1904, Degen s.n. (BM, GB, JE, LE, WU); J´egt´oi, 49°109N, 20°19E, July 1904, Degen s.n. (BM, GB, JE, WU); Dolina Litworowa valley, 49°109N, 20°79E, 29 Aug. 1988, Frey s.n. (L); Nowy, 49°159N, 20°119E, Aug. 1972, Fritze s.n. (K); July 1971, Fritze s.n. (JE, M, PR); pr. Jaworina, 49°359N, 19°249E, 11 Sep. 1865, Fritze s.n. (K); Zelene Pleso, 49°109N, 20°79E, 1 Aug. 1964, Hadac & Dahl s.n. (O); Omlad´ek-v¨olgy valley (Tr¨ummertal), 49°109N, 20°49E, 2 Aug. 1932, Lengyel s.n. (GB, HBG, M, S); Mengusovsk´y valley, 49°109N, 20°49E, 18 Sep. 1910, Ny´ar´ady s.n. (L, UPS). ˇ Zilina: Jarzabczy Wierch, 49°119N, 19°479E, 10 Aug. 1922, Pawlowski s.n. (S); Arva, lacum inferiorem Roh´aˇc see, 49°129N, 19°459E, 25 July 1928, Rechinger s.n. (S); Raczkowa sj¨o, 49°119N, 19°489E, 3 Aug. 1813, Wahlenberg s.n. (UPS). 6. Trisetum glaciale (Bory) Boiss., Elench. Pl. Nov.: 87. 1838. Avena glacialis Bory, Ann. G´en. Sci. Phys. 3: 6. 1820, as “Glacialis.”Trisetaria glacialis (Bory) Paunero, Anales Jard. Bot. Madrid 9: 514. 1950. TYPE: Spain. Granada, Sierra Nevada, Aug. 1837, P. E. Boissier 186 (neotype, designated here, G-00176302 image!; isoneotypes, B- 10_0250162!, B-10_0347071!, BM-001134049 image!, G-00176303!, G-00176305 image!, GH!, GOET!, JE!, K!, L-1346396!, L-1346397!, L- 1346398!, L-1346399!, L-1346401!, LE!, M-0210838!, MA-8022!, MO-1604193!, MPU!, MW!, NY-431723!, O- V2126600!, O-V2126602!, P-00731516!, P-02219569!, Volume 103, Number 3 2018 Barber ´a et al. 381 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa P-02219574!, P-02219579!, P-03364420!, RO image!, US!, W-0025052 image!, W-1889-0242183!). Figure 17. Trisetum antonii-josephii Font Quer & Muñoz Med., Anales Jard. Bot. Madrid 6: 495. 1946, as “Antonii-Josephii.” Trisetaria antonii-josephii (Font Quer & Muñoz Med.) Paunero, Anales Jard. Bot. Madrid 9: 516. 1950. TYPE: Spain. Granada, Sierra Nevada, Jerez del Marquesado, Barranco del Nacimiento del r´ıo Alhor´ı, 37°079N, 3°149W, 19 July 1923, P. Font Quer & J. M. Muñoz Medina s.n. (holotype, BC-81358!). Herbs (3–)8.8–14.4(–19.5) cm tall, caespitose to densely caespitose, shortly rhizomatous, culm 0.3–0.7 mm diam., straight, glabrous, with some dispersed hairs mostly on upper part, sometimes puberulous, with hairs 0.1–0.6(–1.2) mm; nodes (2 to)3(to 4), concentrated in lower part of culm, included in sheaths, pubescent, yellowish. Basal leaf sheaths distinctly veined, not inflated, densely pubescent, with hairs up to 1.2 mm, with dense layer of oldest ones decomposed into fibers, yellowish to greenish; basal leaf blades (0.9–)1.4–4.2(–8.4) cm 3(0.7–)1.6–2.5(–3) mm, rigid, flat, sometimes conduplicate, with central nerve and margins thickened, cartilaginous, densely pubescent, with patent hairs 0.1–0.5(–0.9) mm abaxially and adaxially, grayish to greenish; top culm leaf sheaths (1.8–) 2.5–3.7(–4.6) cm, involving most of culm, distinctly veined, pubescent, with long and short hairs, without cilia on margins; top culm leaf blades (0.2–)0.5–1.5 (–2.6) cm 3(0.6–)1.2–2(–2.5) mm, flat to conduplicate, densely pubescent abaxially and adaxially, with short and some scattered long hairs (0.05–)0.1–0.5(–0.6) mm, greenish to grayish, sometimes slightly yellowish; inner collar region densely pubescent, with hairs 0.1–0.3 (–0.7) mm; ligules (0.3–)0.5–0.7(–1) mm, acute, laciniate, slightly strigose, puberulous on upper part, with hairs 0.2–0.4(–0.9) mm. Basal node of panicle glabrous, sometimes with dispersed hairs 0.1–0.2(–0.3) mm. Panicles (2.3–)3–4.2(–4.8) 3(0.7–)1–1.4(–1.8) cm, ovate in outline, from compact to somewhat lax; rachis glabrous to sparsely pubescent, with hairs 0.1–0.3 mm; longest basal branches (0.3–)0.4–0.9(–1.1) cm. Spikelets (4.2–)4.8–6(–7) 3(1–)1.6–2.4(–3.8) mm, (1)2(3)-flowered; pedicels (1–)1.5–2.2(–3.6) mm, from sparsely pubescent to pubescent, with hairs 0.1–0.3 mm. Glumes unequal to subequal (lower glume length:upper glume length ratio of [0.75–]0.85–0.92[–0.98]); lower glume (3.2–)4.2–5(–5.7) 3(0.6–)0.8–1.2 mm, broadly lanceolate, sometimes narrowly lanceolate (lower glume width:lower glume length ratio of [0.1–]0.2–0.25[–0.3]), acuminate to long-acuminate, (2 to)3-nerved, glabrous, sometimes with very short hairs up to 0.06 mm on upper part of margins and central nerve, greenish on central basal part, sometimes purplish, laterals and upper part Figure 16. Distribution of Trisetum fuscum Schult. (triangles) and T. laconicum Boiss. & Orph. (dots). 382 Annals of the Missouri Botanical Garden purplish, sometimes yellowish; upper glume (4.2–)4.7–5.6 (–6.5) 3(1–)1.2–1.6(–2.1) mm, broadly lanceolate, sometimes lanceolate (upper glume width:upper glume length ratio of [0.17–]0.24–0.33[–0.48]), acuminate to longacuminate, 3-nerved, glabrous, sometimes with scattered andveryshorthairsupto0.07mmonmidribandupper part of margins, greenish on central basal part, sometimes purplish, margins and upper part purplish, sometimes yellowish; rachilla segments between first and second florets 1–1.5(–2.2) mm, with hairs (0.7–)1–1.4(–2) mm; rachilla segments to sterile floret (1–)1.3–1.7(–1.9) mm, with hairs (0.5–)0.8–1.1(–1.7) mm. Lemmas (3.5–)4–4.5 (–5.6) 3(0.6–)0.8–1.2 mm, broadly lanceolate (lemma width:lemma length ratio of [0.24–]0.28–0.37[–0.45]), glabrous to strigulose, greenish, sometimes yellowish, purplish on laterals and upper part; apical teeth (0.2–) 0.4–1(–2) mm, with aristules 0.2–0.5(–0.9) mm, rarely without; awn (4.5–)4.8–7.4(–10.7) mm, inserted (1.9–) 2.3–2.6(–3) mm from base (awn insertion from base length:lemma length ratio of [0.45–]0.54–0.63[–0.71]), Figure 17. Trisetum glaciale (Bory) Boiss. Image of an isoneotype (B-10_0250162). Volume 103, Number 3 2018 Barber ´a et al. 383 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa slightly geniculate, twisted at base when mature, with very short, adpressed hairs up to 0.05 mm; callus (0.15–)0.2 (–0.3) mm, elliptic, sometimes elliptic-oblong, with hairs (0.2–)0.3–0.6(–0.9) mm. Paleae (3–)3.6–4.1(–4.6) 3 (0.6–)0.8–1(–1.4) mm (palea length:lemma length ratio of 0.8–1.05), narrowly elliptic, margins with short, antrorse hairs; teeth (0.2–)0.3–0.6(–0.8) mm, with short, antrorse hairs less than 0.1 mm. Lodicules 0.4–0.7(–0.9) mm, irregularly dentate. Anthers (0.9–)1.7–2.6(–2.9) mm. Ovaries (0.3–)0.5–0.8(–2) mm, from sparsely puberulous to puberulous on upper part, with hairs 0.1–0.3 mm, rarely glabrous. Caryopses 2.5–3.1 30.5–0.8 mm, 6fusiform. Chromosome number: 2n514 (K¨upfer, 1968). Illustrations. Paunero (1950: 548, l´am. XII, sub Trisetaria glacialis; 549, l´am. XIII, sub T. glacialis; 550, l´am.XIV,subT. antoni-josephii; 551, l´am. XV, sub T. antoni-josephii; 552, l´am.XVI,subT. antoni-josephii and T. glacialis; 553, l´am. XVII, sub T. antoni-josephii and T. glacialis;554,l´ am. XVIII, sub T. antoni-josephii and T. glacialis); Chrtek & Jir´asek (1963: 575, figs. 2, 4–6); Chrtek (1965: 214, Abb. 2, 1; 215, Abb. 3, 8; 217, Abb. 4, 4–5; 218, Abb. 5, 4–5). Phenology. Trisetum glaciale has been collected in flower and fruit from June to September. Distribution and habitat. Trisetum glaciale is endemic to the Sierra Nevada in southern Spain, at 2400–3400 m elevation, in mountain pastures and in fissures of schist soils. Figure 14. Discussion. Burdet et al. (1981: 580) proposed Boissier 186 as the lectotype, which seems to be a mistake as that was not original material. After searching in Bory’s collection, no original material has been found. Nevertheless, the specimens collected by Boissier come from the type locality and match the current concept of Trisetum glaciale. Therefore, it is appropriate to consider the Burdet proposal as a neotypification (Art. 9.9 of ICN). The IPNI website has two entries for Avena glacialis, one attributed to Bory (1820) and the other to Boissier (1838). However, Boissier (1838) made a combination of Bory’s name in the genus Trisetum,notAvena,and explicitly referred to the basionym as “Avena glacialis Bor. Fl. n° 20.”Consequently, the name “Avena glacialis Boiss.”has no standing and should be disregarded. Trisetum glaciale is characterized by its densely tufted habit and short height ([3–]8.8–14.4[–19.5] cm), and by having short and rigid leaf blades with a thickened midrib and margins, the margins also cartilaginous, pubescent top culm leaf sheaths, panicles ovate in outline, subequal glumes (lower glume length:upper glume length ratio of [0.75–]0.85–0.92 [–0.98]), palea almost or as long as the lemma (palea length:lemma length ratio of 0.8–1.05), and ovaries from sparsely to densely puberulous apically, rarely glabrous. Font Quer and Muñoz Medina (Font Quer, 1946) described another species from Sierra Nevada, Trisetum antonii-josephii. It was separated from T. glaciale by its narrower and 1-veined lower glumes and lanceolate lemmas with longer awns. Paunero (1950) made a more detailed study indicating the difference in the lemma apex (acute in T. antonii-josephii and obtuse in T. glaciale), noting that the leaf structure, which varies widely among the species of the genus, as well as the ovary indumentum, is identical in both species. Romero (2009) recognized both species and indicated that specimens with intermediate characters have been found, probably of hybrid origin. In the material studied, we determined that the number of nerves in the lower glumes depends on the stage of development of the specimen, and is, therefore, not a good taxonomical character state for discriminating these taxa. Awn length is also variable in specimens of the same population. Therefore, we consider T. antonii-josephii to be a synonym of T. glaciale. Additional specimens examined. SPAIN. Granada: Sierra Nevada, ascensi´on al Veleta desde la parada del microb´us, bordes del Corral del Veleta, 37°39N, 3°229W, 24 July 2013, Barber´a et al. 937PB (MA); ascensi´on al Veleta, trincheras junto a la parada del microb´us, 37°39N, 3°229W, 24 July 2013, Barber´a et al. 939PB (MA); Caldera du Mulhacen, 37°39N, 3°189W, 25 July 1851, Bourgeau 1517 (G, GOET, K, P); Valle de Lanjar´on, Peñ´on Elorrieta, 37°29N, 3°239W, 6 July 1930, Ceballos & Vicioso s.n. (MA); Monachil, inter Laguna de Las Yeguas et Pico Veleta, 37°39N, 3°229W, 26 July 1978, Charpin et al. 15297 (G); autour des lagunas de R´ıo Seco, 37°39N, 3°209W, 15 July 1983, Defferrard 4317 (G); Monte Pelado, 37°69N, 3°159W, 24 Aug. 1923, Font Quer s.n. (BC, BM, GDA, JE, K, MA, S, UPS); Laguna de la Caldera, 37°39N, 3°199W, 8 Aug. 1924, Ginzberger s.n. (W); Picacho de Veleta, 37°39N, 3°229W, 1 Aug. 1879, Huter et al. 1172 (B, BM, G, GB, HBG, K, LE, M, MPU, P, PR, W, WU); Raspones de R´ıo Seco, 37°39N, 3°209W, 4 Aug. 1997, Mart´ınez Lirola & Vargas s.n. (GDA); entre 1 a y2 a laguna del r´ıo Lanjar´on, 37°29N, 3°249W, 28 July 1997, Mart´ınez Lirola & Vargas s.n. (GDA); Picacho de Veleta, 37°39N, 3°229W, 22 Aug. 1974, Merxm¨uller & Lippert 29582 (M); Capileira, alrededores de la Laguna de Majano, 37°29N, 3°199W, 13 Aug. 1978, Molero Mesa s.n. (GDA); Pico del Veleta, 37°39N, 3°229W, 25 July 1947, Muñoz Medina & Serrano s.n. (ARAN, BC, F, GDA, MA); Picacho de Veleta, 37°39N, 3°229W, Aug. 1891, Porta & Rigo 630 (B, G, GB, JE, K, M, MA, NY, P, S, W, WU); subida al Pico Mulhac´en, 37°39N, 3°189W, 18 June 2014, Quintanar et al. 5226 (MA); subida al Moj´on Alto, entre la loma de Maitena y el Calvario, 37°79N, 3°179W, 21 June 2014, Quintanar et al. 5273 (MA); al pie del Collado de las Buitreras, 37°69N, 3°169W, 21 June 2014, Quintanar et al. 5275 (MA); bajada del Collado de las Buitreras en el camino hacia Cerro Pelao, 37°69N, 3°159W, 21 June 2014, Quintanar et al. 5277 (MA); Cerro Pelao, 37°69N, 3°159W, 21 June 2014, Quintanar et al. 5278 (MA); Barranco del R´ıoAlhor´ıy Ventisquero de los Caballones, 37°69N, 3°159W, 21 June 2014, Quintanar et al. 5279 (MA);LagunadelaCalderahastaladeMajano,37°29N, 384 Annals of the Missouri Botanical Garden 3°199W, 4 Aug. 1981, Romero s.n. (GDA);Barranco de San Juan, 37°49N, 3°229W, 18 July 1981, Romero s.n. (GDA); alrededores de la Laguna de Aguas Verdes, 37°29N, 3°229W, 22 July 1980, Romero & Morales s.n. (GDA); Refugio de Elorrieta, 37°29N, 3°239W, 24 July 1981, Romero & S´anchez s.n. (GDA); Mulhac´en, Laguna La Caldera, 37°39N, 3°229W, 28 July 1967, Segura Zubizarreta s.n. (MA); Sierra Nevada, 4 Aug. 1845, Willkomm 1281 (MPU, P); Picacho de Veleta, 37°39N, 3°229W, 31 July 1876, Winkler s.n. (B, M, MPU, P, WU). 7. Trisetum gracile (Moris) Boiss., Voy. Bot. Espagne 2: 654. 1844. Avena gracilis Moris, Stirp. Sard. Elench. 1: 50. 1827. Trisetum morisii Trin., Bull. Sc. Acad. Petersb. 1: 66. 1836, nom. illeg. superfl. TYPE: Italy. Sardinia, Nuoro Prov., Gennargentu Mtns., 40°019N, 9°209E, G. G. Moris s.n. (lectotype, designated here, LE-01042367!; possible isolectotypes, JE-00018855!, P-02222289!, P- 02222295!). Figure 18. Trisetum conradiae Gamisans, Candollea 26: 322. 1971. Trisetaria conradiae (Gamisans) Banfi & Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat. Milano 135: 382. 1996. TYPE: France. Corsica, Incudine Massif, 41°519N, 9°129E, 14 July 1968, J. Gamisans 68 (holotype, Gamisans’private herbarium; isotype, G- 00100183!). Trisetum gracile var. morisianum Gamisans, Candollea 28: 54. 1973. TYPE: Italy. Sardinia, Ogliastra Prov., Punta La Marmora Mtn., 39°599N, 9°199E, 24 June 1970, J. Gamisans & M. Reille S-2399 (holotype, G- 00386420!). Herbs (4.5–)8.9–12.5(–18.2) cm tall, caespitose to densely caespitose, shortly rhizomatous, culm (0.2–) 0.4–0.6 mm diam., often bent at nodes, glabrous, sometimes with hairs 0.2–0.4 mm below basal node; nodes 2 to 3(to 4), mostly concentrated in lower part of culm, not included in sheaths, rarely included, glabrous, blackish. Basal leaf sheaths distinctly veined, somewhat inflated, glabrous to sparsely pubescent, with hairs up to 0.1 mm, with dense layer of oldest ones decomposed into fibers, yellowish to brownish, rarely greenish; basal leaf blades 1–2(–4) cm 3(0.6–)1.3–1.7 (–2.6) mm, rigid, flat to convolute or revolute, with central nerve thickened, glabrous to sparsely puberulous adaxially, pubescent abaxially, also in margins, with hairs (0.05–)0.1–1.2(–1.8) mm, greenish to grayish; top culm leaf sheaths (1.4–)2–3.2(–4) cm, distinctly veined, somewhat inflated, glabrous, without cilia on margins; top culm leaf blades (0.2–)0.3–1(–1.8) cm 3 (0.6–)0.8–1.2 mm, conduplicate, rarely flat, parallel to culm, glabrous to sparsely pubescent abaxially and adaxially, with hairs up to 0.2 mm on margins, brownish to yellowish, sometimes grayish; inner collar region glabrous, sometimes with few scattered hairs up to 0.3 mm on margins; ligules 0.5–0.9 mm, laciniate to shortly irregularly toothed, with hairs 0.1–0.3 mm on upper part. Basal node of panicle glabrous. Panicles (1.3–)1.6–2.5(–3.1) 3(0.7–)1–1.5(–1.8) cm, ovate in outline, somewhat dense; rachis internodes glabrous to sparsely pubescent, with hairs 0.05–0.2 mm; longest basal branches (0.2–)0.3–0.9(–1.1) cm. Spikelets (4–) 5.2–6.4(–6.7) 3(1.7–)2.4–4(–5.2) mm, 2- to 3-flow- ered; pedicels (1.4–)1.8–3.2(–4.4) mm, sparsely puberulous, with hairs up to 0.2 mm. Glumes slightly subequal to almost equal (lower glume length:upper glume length ratio of [0.79–]0.82–0.89[–0.97]); lower glume (3.6–)4–5.6(–6.5) 31.2–1.6(–2.4) mm, elliptic to broadly lanceolate (lower glume width:lower glume length ratio of [0.28–]0.3–0.35[–0.4]), acuminate to long-acuminate, 1(or 2)-nerved, glabrous, greenish surrounded by purple, margins hyaline; upper glume (4–) 4.7–6.6(–6.8) 3(1.8–)2–2.4 mm, elliptic to oblong, sometimes narrowly elliptic or narrowly lanceolate (upper glume width:upper glume length ratio of 0.33–0.43 [–0.49]), acuminate, sometimes long-acuminate, 3- nerved, glabrous, greenish surrounded by purple, margins hyaline; rachilla segment between first and second florets 0.6–1.2 mm, with hairs (1.6–)2–2.4(–3) mm; rachilla segment to sterile floret (0.8–)1–1.3 mm, with hairs (0.7–)1–1.5(–2) mm. Lemmas (3–)4–5(–5.6) 3 1.2–1.6(–2.4) mm, narrowly elliptic to narrowly lanceolate, sometimes broadly lanceolate (lemma width: lemma length ratio of [0.26–]0.3–0.36[–0.47]), glabrous to pubescent, with hairs (0.05–)0.6–1.4(–2) mm between awn and base, greenish, sometimes yellowish, surrounded by purplish flush; apical teeth (0.1–) 0.2–0.4 mm, with aristules 0.3–0.6(–1) mm; awn (4.5–) 5–6.6(–9.1) mm, inserted (1.3–)1.8–2.2(–2.8) mm from base (awn insertion from base length:lemma length ratio of [0.34–]0.41–0.49[–0.51]), geniculate and slightly twisted at base, with very short, adpressed hairs up to 0.05 mm; callus 0.1–0.4 mm, elliptic to orbicular, with hairs (0.7–) 1–1.3 mm. Paleae (2.7–)3.7–4.2(–4.6) 30.7–1.3 mm (palea length:lemma length ratio of [0.66–]0.76–0.84 [–1.1]), elliptic or narrowly elliptic to slightly oblanceolate, margins glabrous; teeth (0.1–)0.2–0.4 mm, glabrous or with short, antrorse hairs. Lodicules (0.6–)0.9–1.2(–1.8) mm, entire, sometimes with 2 lobules or irregularly dentate. Anthers (1.4–)1.8–2.3(–3) mm. Ovaries 0.5–1(–2) mm, glabrous. Caryopses 1.5–2.6 30.4–0.7 mm, oblong or narrowly elliptic. Chromosome number: 2n514 (Gamisans, 1973). Additional illustrations. Fiori & Paoletii (1895: 27, 230); Chrtek & Jir´asek (1963: 575, fig. 2, 1–3); Chrtek (1965: 214, Abb. 2, 4–5; 215, Abb. 3, 8; 217, Abb. 4, 6–7; 218, Abb. 5, 1–3); Gamisans (1971: 323, fig. 1, 1–5; 325, fig. 2, 1–3); Jeanmonod & Gamisans (2007: XXX, fig. 32, b). Phenology. Trisetum gracile has been collected in flower and fruit from May to August. Volume 103, Number 3 2018 Barber ´a et al. 385 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa Distribution and habitat. This species is endemic to the mountains of southern Corsica and Sardinia. It grows in poor soils of granitic sands at 1300–2100 m. Figure 14. Discussion. Moris’s herbarium is kept at TO, but Dr. Laura Guglielmone kindly informed us that the Moris collection was closed for security reasons. Figure 18. Trisetum gracile (Moris) Boiss. —A. Habit. —B. Portion of culm and node. —C. Portion of sheath, ligule, and blade. —D. Transverse section of leaf blade. —E. Spikelet. —F. Florets. —G. Lower glume, dorsal view. —H. Upper glume, dorsal view. —I, J. Floret. —K. Lemma, upper part, lateral view. —L. Palea, lateral view. —M. Lodicules. —N. Stamens. —O. Pistil. —P. Caryopsis. A based on Gamisans 14148 (G-00220037); B–H, J–O based on Reverchon 312 (S); I based on Barber´a & Quintanar 1441PB (MA); P based on M¨uller s.n. (M-0223271). 386 Annals of the Missouri Botanical Garden Fortunately, we have found original material at some herbaria. There is a specimen at LE with a label handwritten by Moris and collected in the Gennargentu Mountains, one of the localities mentioned in the protologue, which has been selected as lectotype. P and JE specimens are potential isolectotypes, although their localities are less specific (the labels read only Sardinia). Trisetum gracile and T. glaciale are two species of the section Trisetum with a similar habit, being densely tufted, having the shortest height ([4.5–]8.9–12.5 [–18.2] cm and [3–]8.8–14.4[–19.5] cm, respectively), rigid leaves with a thickened midrib, short upper leaf blades ([0.2–]0.3–1[–1.8] cm and [0.2–]0.5–1.5[–2.6] cm, respectively), panicles ovate in outline, and subequal glumes. Nevertheless, T. gracile is clearly differentiated from T. glaciale by its blackish nodes usually exerted from the sheaths, with culms often bent at the nodes and more or less glabrous, sheaths somewhat inflated, long rachilla hairs ([1.6–]2–2.4[–3] mm and [0.7–]1–1.4[–2] mm, respectively), lemmas usually hairy between the base and the awn insertion, awn inserted slightly below the middle of the lemma (awn insertion from base length:lemma length ratio of [0.34–] 0.41–0.49[–0.51]), long callus hairs ([0.7–]1–1.3 mm and [0.2–]0.3–0.6[–0.9] mm, respectively), and glabrous ovaries. Gamisans (1971) described a new species, Trisetum conradiae, from Corsica that was very close taxonomically to T. gracile, based on its glabrous lemmas, among other characters. After collecting more material, Gamisans (1973) realized that this character was the only one that separated both species. Later, Jeanmonod and Gamisans (2007) recognized T. conradiae at subspecific rank. In our own observations, specimens with and without hairy lemmas have been collected in the same populations, varying in the density and the length of the hairs ([0.05–]0.6–1.4[–2] mm). In other species of the section, such as T. flavescens subsp. griseovirens, specimens with hairy lemmas have been found, hairs sometimes appearing only in the proximal lemma. Consequently, we agree with Tison and de Foucault (2014) that this character does not have any taxonomical importance here, and T. conradiae is considered a synonym of T. gracile. Additional specimens examined. FRANCE. South Corsica: Monte Incudine, Plateau de Cuscione, 41°509N, 9°109E, 5 July 2014, Barber´a & Quintanar 1439PB (MA); Barber´a& Quintanar 1440PB (MA); Barber´a & Quintanar 1441PB (MA); crˆete Monte Incudine, Tintennaga, 41°519N, 9°139E, 24 July 1985, Gamisans 11703 (G); Pianu di Cuscione, Fonda di Serra Longa, 41°509N, 9°89E, 1 July 1989, Gamisans 14148 (G); Monte Incudine, Foce Aperta, 41°529N, 9°129E, 6 July 1989, Gamisans 14203 (G); Massif de l’Incudine, Coscione, crˆete au S de la Punta Tozzarella, 41°519N, 9°119E, 1 Aug. 1972, Gamisans 2242 (C); Massif de l’Incudine, crˆete `a 100 m au SE de Bocca Chiralba, versant N, 41°509N, 9°119E, 1 Aug. 1972, Gamisans 2247-I (MPU); crˆete au S de la Punta Tozzarella, 41°519N, 9°109E, 25 July 1974, Gamisans 3591 (G); crˆete au S de Bocca Chiralba, 41°509N, 9°119E, 25 July 1974, Gamisans 3598 (G); Macizo del monte Incudine, meseta de Coscione, 41°509N, 9°89E, 6 July 2014, Quintanar & Barber´a 5390AQ (MA). ITALY. Sardinia: Nuoro Prov., Fonni, Gennargentu, Bronco Spina, 40°19N, 9°189E, 3 June 2003, Castroviejo et al. 17028 (MA); Nuoro Prov., Gennargentu, Monte di Scudo, versant SW, 40°19N, 9°199E, 23 June 1970, Gamisans S-2394bis (G); Nuoro Prov., Monte Gennargentu, 40°19N, 9°199E, July 1827, M¨uller s.n. (BM, H, HBG, JE, LE, M, PR, WU); Sassari Prov., Monte Limbara, lors d’un chemin pr`es de l’extremit´e de la route, 40°519N, 9°109E, 30 May 1983, Charpin et al. AC17871 (G); Sassari Prov., Tempio, Monte Limbardo [Limbara], 40°519N, 9°109E, 4 July 1882, Reverchon 312 (B, BM, G, JE, K, LE, M, MA, MO, MPU, NY, P, PR, UPS, WU); Reverchon s.n. (B, G, GB, GOET, H, HBG, JE, L, M, MPU, NY, P, PR, RO, S, UPS, US, WU). 8. Trisetum laconicum Boiss. & Orph. in Boiss., Diagn. Pl. Orient., ser. 2, 4: 129. 1859. Avena laconica (Boiss. & Orph.) Nyman, Syll. Fl. Eur. Suppl.: 71. 1865. Trisetaria laconica (Boiss. & Orph.) Banfi & Soldano, Atti Soc. Ital. Sci. Nat. Mus. Civico Storia Nat. Milano 135: 383. 1996. TYPE: Greece. Peloponnese: Malev´os Mtn., near Vrancopigadou, 37°379N, 22°319E, 19 July 1850, T. G. Orphanides s.n. (lectotype, designated by Tan & Iatrou [2001: 450], G-BOIS image!; isolectotypes, B-10_0508993!, JE-18853!, P-3365069!, W-1916-0031855!, WU-0079593 image!, WU- 0079594 image!). Figures 9D, 10D, 15B. Trisetum rechingeri Chrtek, Acta Univ. Carol., Biol. 1966: 92. 1967. TYPE: Greece. Central Greece: Euboea, Dirfi Mtn., 38°379N, 23°509E, 7 Aug. 1858, T. H. H. von Heldreich 3522 (holotype, P not seen; isotype, W-1916-0031796!). Herbs 38.5–83.5 cm tall, not caespitose, rhizomatous, culm 0.5–1.1 mm diam., straight, glabrous; nodes 4 to 6, separated along culm, included or not in sheaths, glabrous, yellowish or brownish. Basal leaf sheaths pubescent, with hairs 0.5–1 mm, decaying into fibers, brownish; basal leaf blades 5–6cm33.6–5mm, flat, pubescent, with hairs 0.3–1 mm, also on margins, brownish to grayish; top culm leaf sheaths 9.7–25 cm, glabrous, ciliate; top culm leaf blades 3.1–11.2 cm 31.3–4 mm, flat, sometimes slightly inrolled and parallel to culm, pubescent abaxially, glabrous adaxially, with hairs 0.2–0.6 mm, also on margins, greenish to grayish; inner collar region glabrous, sometimes with scattered hairs up to 0.1 mm on margins; ligules 2.5–3.2 mm, irregularly dentate to laciniate, glabrous or with scattered hairs up to 0.2 mm on surface and upper part. Basal node of panicle glabrous to sparsely pubescent with adpressed hairs up to 0.05 mm. Panicles 8–19 31.5–6 cm, narrowly elliptic to oblong in outline, somewhat dense; rachis internodes glabrous to sparsely pubescent, with hairs up Volume 103, Number 3 2018 Barber ´a et al. 387 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa to 0.05 mm; longest basal branches 1.3–2.4 cm. Spikelets 8–9.2 32.3–4.8 mm, 3- to 4-flowered; pedicels 5–6.3 mm, sparsely pubescent, with hairs up to 0.1 mm. Glumes unequal (lower glume length:upper glume length ratio of 0.69–0.74); lower glume 4.3–5.4 3 0.8–1.1 mm, narrowly lanceolate (lower glume width: lower glume length ratio of 0.18–0.2), long-acuminate, 1-nerved, glabrous, with hairs up to 0.05 mm on upper part and margins, yellowish to greenish; upper glume 6.2–7.3 31.4–2.1 mm, narrowly elliptic to lanceolate (upper glume width:upper glume length ratio of 0.22–0.29), acuminate, 3-nerved, glabrous, with scattered hairs up to 0.05 mm on upper part of central nerve and margins, yellowish to greenish; rachilla segment between first and second florets 1.2–1.7 mm, with hairs 1.5–2.5 mm; rachilla segment to sterile floret 1.3–1.6 mm, with hairs 0.7–1.1 mm. Lemmas 6.5–6.6 30.8–1.1 mm, narrowly elliptic to narrowly lanceolate (lemma width:lemma length ratio of 0.21–0.23), scabridulous, with hairs up to 0.05 mm, greenish to yellowish, sometimes with purplish flush; apical teeth 0.2 mm, with aristules 0.4–0.5 mm; awn 6.3–7.2 mm, inserted 4.1–4.5 mm from base (awn insertion from base length:lemma length ratio of 0.62–0.65), strongly bent near base, 6twisted at base, with very short, adpressed hairs up to 0.05 mm; callus 0.3 mm, elliptic to oblong, with hairs 0.6–0.9 mm. Paleae 5.8–631.1–1.5 mm (palea length:lemma length ratio of 0.89–0.9), narrowly elliptic, ciliate on upper part; teeth 0.3–0.4 mm, with short, antrorse hairs. Lodicules 0.6 mm, with regular teeth or laciniate. Anthers 2.4–3.1 mm. Ovaries 0.8–0.9 mm, glabrous. Caryopses not seen. Chromosome number not known. Phenology. Trisetum laconicum has been collected in flower from June to August. Distribution and habitat. Trisetum laconicum is endemic to Greece and is known from only two collections: Malevos Mountains in the south of the Peloponnese and Dirfi Mountains in Euboea Island; on limestone rocks and cliffs. It has been found at elevations from 1220 to 1500 m. Figure 16. Discussion. Boissier and Orphanides (Boissier, 1859) separated the Malevos Mountains species, Trisetum laconicum, from T. flavescens mainly by its densely hairy leaves, larger spikelets, and longer rachilla hairs. Apart from those characters, we have also discerned that T. laconicum has longer ligules (2.5–3.2 mm), pedicels (5–6.3 mm), lemmas (6.5–6.6 mm), callus indumentum (0.6–0.9 mm), and paleae (5.8–6 mm) than T. flavescens. Another distinguishing characteristic is the awn shape, which is strongly bent near the base in T. laconicum and geniculate to slightly geniculate in T. flavescens.We found no or only minor overlap with T. flavescens in any of the previously mentioned characters. For this reason, and considering that only the type material is available, T. laconicum is treated as a separate species. Chrtek (1967a) described Trisetum rechingeri from Euboae Island based on a single sheet (Heldreich 3522, P). He indicated that T. laconicum was the taxonomically closest species to T. rechingeri, separating them mainly by its smaller size and shorter rachilla hairs and spikelets. Chrtek (1967a) included both species in a series within Trisetum sect. Rigida, named Trisetum ser. Laconica, and characterized by its general habit and shorter rachilla hairs. Jonsell (1980) decided to consider these species as conspecific until more material is available. Furthermore, Strid (2016) indicated that no more material has been collected on Mt. Dirfis and considered that T. rechingeri represents individual variation within T. laconicum. We agree with these latter authors and therefore consider T. rechingeri to be a synonym of T. laconicum. MISCELLANEOUS DOUBTFUL,INVALID,AND EXCLUDED (FROM TRISETUM)NAMES Avena alpestris DC. The IPNI website attributed the name Avena alpestris to de Candolle. De Candolle (Lamarck, 1815), however, when recording A. alpestris, indicated “A. Alpestris.Host. gram.3,p.27,”suggesting that he was not publishing a nomen novum. Consequently, the isonym “Avena alpestris DC.”should be disregarded (Art. 6, Note 2). Avena flavescens var. vulgaris Alef., Landw. Fl.: 350. 1866, nom. inval. (Art. 24.3). Alefeld (1866) divided Avena flavescens into three varieties, and he used the epithet “vulgaris”for the first, purporting to indicate the taxon containing the type of A. flavescens. Avena sesquitertia Host The IPNI website attributed the name Avena sesquitertia to Host (1797).Host (1797), however, when he recorded A. sesquitertia, repeated the polynomial of Linn´e (1767), “A. paniculata, calycibus subtrifloris, flosculis omnibus aristatis, receptaculis barbatis. Syst. veget. p. 122,”suggesting that he was not publishing a new taxon name. Consequently, the isonym “Avena sesquitertia Host”should be disregarded (Art. 6, Note 2). Avena tenuis Moench, Methodus: 195. 1794, nom. illeg. superfl. for Avena dubia Leers [5Ventenata dubia (Leers) Coss.]. 388 Annals of the Missouri Botanical Garden Rebentischia flavescens Opiz, Lotos 4: 104. 1854, nom. inval., syn. of Trisetum flavescens. Trisetum alpestre var. viviparum-macranthum Schur, Oesterr. Bot. Z. 10: 74. 1860, nom. inval. (Art. 23.6). Trisetum ciliare (Kit. ex Schult.) Domin, Preslia 13–15: 41. 1935, nom. inval. The combination proposed by Domin (1935) is based on Avena ciliaris Kit. ex Schult., Oestr. Fl., ed. 2, 1: 268. 1814, nom. nud. Trisetum debile Chrtek, Folia Geobot. Phytotax. 25: 334. 1990. TYPE: China. Yunnan Prov., Dschungdien, 27°499N, 99°429E, 22 Aug. 1915, H. R. E. Handel- Mazzetti 7739 (holotype, W-1940-0014741!; isotype, WU-0033964 image!). The type is the only original material that could be located. It is a fragment that cannot be identified to species, but it may belong to one of the taxa allied to the genus Trisetum. Its slender habit may be due to the fact that the specimen was immature. Trisetum flavescens subsp. agrostideum (Laest.) Douin in Bonnier & Douin, Fl. Ill. France 12: 19. 1934, syn. of Trisetum subalpestre (Hartm.) Neuman. The epithet “agrostideum”was attributed to Fries (1842), but it was previously published by Laestadius (1839). Trisetum flavescens var. bulbosum Holla ex Asch., Fl. Brandenburg 1: 830. 1864. TYPE: Germany. Brandenburg, Holla s.n. (original material not located). Ascherson’s description seems to refer to Arrhenatherum bulbosum (Willd.) C. Presl. Trisetum flavescens subsp. copiosum Printz, Veg. Siber.- Mongol. Front.: 124, 125, fig. 74. 1921. TYPE: Russia. Krasnoyarsk Krai, Minusinsk Distr., July 1914, K. Printz s.n. (lectotype, designated here, C!; syn. of Trisetum sibiricum Rupr. subsp. sibiricum). This name was not included in Barber´a et al. (2017b). Trisetum flavescens var. dimorphantha Maire & Weiller, Fl. Afrique N. 2: 248. 1953, nom. inval. (Art. 39.1). Trisetum flavescens subsp. fuscum Domin, Preslia, 13–15: 41. 1935, nom. inval., syn. of Trisetum ciliare (Kit. ex Schult.) Domin. Trisetum flavescens var. latifolium-majus Schur, Oesterr. Bot. Z. 10: 77. 1860, nom. inval. (Art. 23.6). Trisetum flavescens var. sibiricum (Rupr.) Ostenf. ex B. Fedtsch., Izv. Imp. Bot. Sada Petra Velikago 14(2): 64. 1913, syn. of Trisetum sibiricum Rupr. subsp. sibiricum. This name was not included in Barber´a et al. (2017b). Trisetum flavescens var. subtriflorum-transsilvanicum Schur, Oesterr. Bot. Z. 10: 76. 1860, nom. inval. (Art. 23.6). Trisetum gracile subsp. conradiae (Gamisans) Gamisans, Fl. Corsica: 255. 2007, nom. inval. (Art. 41.5). Trisetum miegevillii Duval-Jouve ex Mi´egev., Bull. Soc. Bot. France 21: 47. 1874, nom. nud., syn. of Trisetum baregense Laffitte & Mi´egev. Trisetum tarnowskii Zapał., Rozpr. Wydz. Mat.-Przyr. Akad. Umiejetn., Dzia B, Nauki Biol., III, 4: 167. 1904. TYPE: Romania. Suceava, Petrile Doamnei, 47°279N, 25°349E, A. Rehman s.n. (lectotype, designated here, KRAM-074724 image!; syn. of Trisetum macrotrichum Hack.). This name was not typified in Barber´a et al. (2017a). Trisetum tarnowskii f. albovirens Zapał., Kosmos (Lviv) 34: 1176. 1909. TYPE: Romania. Suceava, Petrile Doamnei, 47°279N, 25°349E, 10 Aug. 1909, H. Zapałowicz s.n. (lectotype, here designated, KRAM-074594 image!; isolectotypes, KRAM- 074450 image!, KRAM-074595 image!; syn. of Trisetum macrotrichum Hack.). This name was not included in Barber´a et al. (2017a). Trisetum tenue Roem. & Schult., Syst. Veg., ed. 15 bis [Roemer & Schultes] 2: 657. 1817, nom. illeg. According to Jonsell (1978: 320), Trisetum tenue Roem. & Schult. is a nomen superfluum for Avena dubia Leers [Ventenata dubia (Leers) Cosson]. Trisetum transsilvanicum Schur, Oesterr. Bot. Z. 10: 77. 1860, nom. inval., syn. of Trisetum flavescens var. subtriflorum-transsilvanicum Schur, Oesterr. Bot. Z. 10: 76. 1860. Trisetum varium Schur, Oesterr. Bot. Z. 10: 75. 1860, nom. illeg. superfl. This forgotten name is based on Avena varia Schur, Sert. Fl. Transsilv.: 85. 1853, which is a superfluous name for A. carpatica Host, Icon. Descr. Gram. Austriac. 4: 18, tab. 31. 1809, Trisetum carpaticum (Host) Roem. & Schult., Syst. Veg. 2: 663. 1817, T. flavescens var. carpaticum (Host) Zapał., Rozpr. Wydz. Mat.-Przyr. Akad. Umiejetn., DziałB Nauki Biol. 4: 107. 1904. Host’s illustration clearly represents Avenula pubescens (Huds.) Dumort. Trisetum yunnanense Chrtek, Folia Geobot. Phytotax. 25: 335. 1990. TYPE: China. Yunnan Prov., near Volume 103, Number 3 2018 Barber ´a et al. 389 Trisetum sect. Trisetum (Poaceae) from Eurasia and North Africa Lidjiang, 26°529N, 100°149E, 18 July 1914, H. R. E. Handel-Mazzetti 4212 “(H-M Diar. Nr. 664)”(holotype, W-1926-0004138!; isotype, WU-0033963!). Chrtek (1990) described this species as remarkable for its robust habit, relatively short, rigid leaves, and large spikelets. Wu and Phillips (2006) included this species in the Flora of China, indicating that the hairy ovary is unusual in Trisetum and that they did not see the type. The type material has a thicker, scarious palea, a densely hairy ovary with long hairs, and an immature caryopsis with a long, linear hilum. Those characters fit well with the genus Helictotrichon, suggesting that this species belongs to that genus. Literature Cited Alefeld, F. 1866. Landwirthschaftliche Flora. Wiegandt & Hempel, Berlin Alejandre, J. A., V. J. Ar´an, P. Barbadillo, J. J. Barredo, J. Benito, M. J. Escalante, J. M. 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Berling, Lund. Gabrielian, E. 2010. Trisetum Pers. Pp. 172–177 in E. Gabrielian & M. E. Oganesian (editors), Flora of Armenia, Vol. 11. A. R. G. Gantner Verlag KG, Ruggell, Liechtenstein. Galland, N. 1985. Chromosome number reports LXXXVII. Taxon 34: 346–351. Galland, N. 1988. Recherche sur l’origine de la flore orophile du Maroc. Etude caryologique et cytogeographique. Trav. Inst. Sci. Univ. Mohammed V, S´er. Bot. 35: 1–168. Gamisans, J. 1971. Contribution `al’´etude de la flore de la Corse. II. Candollea 26: 309–358. 390 Annals of the Missouri Botanical Garden