Brachypodium distachyon, photographed by Neil Harris.
Belongs within: Poaceae.
Contains: Phalarideae,
Deyeuxia,
Calamagrostis,
Piptochaetium,
Polypogon,
Meliceae,
Stipa,
Poa,
Avena,
Agrostis,
Deschampsia,
Trisetum,
Triticum,
Hordeum,
Agropyrum,
Festuca,
Bromus.
The Pooideae is a clade of grasses characterised by the loss of stylar fusion (Grass Phylogeny Working Group 2001). Many pooids also possess multiple florets per spikelet and lack multicellular microhairs. Members of the clade include a number of significant grain species, such as wheat (
Triticum), rye (
Secale cereale), oats (
Avena) and barley (
Hordeum), and a number of major pasture grasses. In contrast, some species of
Nassella (needle grasses) are regarded as weeds in pasture because of their low palatability for livestock. The basalmost member of the subfamily is the genus
Brachyelytrum, a perennial grass with solitary spikelets borne in panicles, with persistent but very short glumes, found in Asia and North America.
Characters (from Grass Phylogeny Working Group 2001): Plants annual or perennial (rhizomatous, stoloniferous, or neither), herbaceous. Culms hollow (rarely solid). Leaves distichous; abaxial ligule absent; adaxial ligule scarious or membranous, the margin not or infrequently short ciliate fringed (rarely long ciliate); blades somewhat broad to usually narrow, rarely pseudopetiolate, venation parallel; sheaths sometimes auriculate. Inflorescences spicate, racemose, or paniculate, bracts outside of the spikelets absent or rarely present. Spikelets bisexual, infrequently unisexual or mixed, usually with two glumes (rarely without glumes or the first absent except on terminal spikelets), 1 to many female-fertile florets with apical or infrequently basal reductions, compressed laterally, infrequently not or dorsally compressed, disarticulating above the glumes (infrequently below the glumes or at the nodes of the inflorescence); lemma lacking uncinate macrohairs, if awned, the awn single; palea usually present and well developed, but variable and sometimes very reduced or absent; lodicules 2 (rarely 3, fused, or absent), usually lanceolate, broadly membranous apically (sometimes fleshy, truncate), often lobed, obscurely few-nerved, or infrequently more or less distinctly few-nerved, not or conspicuously ciliate on the margins; stamens usually 3 (infrequently 1 or 2); ovary glabrous or pubes- cent, rarely with an apical appendage or rostellum, haustorial synergids absent, styles usually 2, close, stigmas 2 (rarely 1 or 3). Caryopsis with the hilum linear and up to as long as the fruit, or subbasal and punctiform, linear, ellipsoidal, ovate, or circular and less than 1/3 the length of the fruit; endosperm hard or sometimes soft or liquid, with or without lipids, containing compound starch grains, or simple starch grains; embryo small, epiblast present (rarely absent), scutellar cleft absent (rarely present, but not deeply incised), mesocotyl internode absent (rarely short), embryonic leaf margins meeting (infrequently margins overlapping). Basic chromosome numbers:
x = 7, 2, 4, 5, 6, 8, 9, 10, 11, 12, 13.
Foliar anatomy: Mesophyll nonradiate, adaxial palisade layer absent, fusoid cells absent, arm cells absent; Kranz anatomy absent; midrib simple; adaxial bulliform cells present.
Foliar micromorphology: Stomata with parallel-sided subsidiary cells; bicellular microhairs absent (rarely present, where chloridoid or panicoid), unicellular microhairs absent (rarely present); papillae usually absent, when present rarely more than one per long cell. Photosynthetic pathway C
3.
<==Pooideae (see below for synonymy)
|--Brachyelytrum [Brachyelytreae] GPWG01
| `--B. erectum GPWG01
`--+--+--Nardus [Nardeae] GPWG01
| | `--N. stricta GPWG01
| `--Lygeum [Lygeeae] GPWG01
| `--L. spartum GPWG01
`--+--Meliceae GPWG01
`--+--+--+--Anisopogon avenaceus GPWG01
| | `--Phaenosperma [Phaenospermatideae] GPWG01
| | `--P. globosum GPWG01
| `--+--Ampelodesmos [Ampelodesmeae] GPWG01
| | `--A. mauritanica GPWG01
| `--+--+--Stipa GPWG01
| | `--Nassella GPWG01
| | |--N. leucotricha GPWG01
| | |--N. trichotoma HE80
| | `--N. viridula GPWG01
| `--+--Piptatherum GPWG01
| | |--P. coerulescens (Desfontaines) Beauvois 1812 PL04
| | |--P. miliaceum GPWG01
| | `--P. songaricum GPWG01
| `--Oryzopsis GPWG01
| |--O. hymenoides CL86
| |--O. lateralis O88
| |--O. miliacea AGF98
| |--O. munroi O88
| `--O. racemosa [=Piptatherum racemosum] GPWG01
`--+--Diarrhena [Diarrheneae] GPWG01
| |--D. americana GPWG01
| `--D. obovata GPWG01
`--+--+--Trachynia Link 1827 GR98
| | `--T. platystachya (Balansa) Scholz in Greuter & Raus 1998 (see below for synonymy) GR98
| `--Brachypodium [Brachypodieae] GPWG01
| |--B. distachyon (Linnaeus) Beauvois 1812 GPWG01, PL04 [=Trachynia distachya GR98]
| |--B. mexicanum GPWG01
| |--B. pinnatum (Linnaeus) Beauvois 1812 PL04
| |--B. ramosum C74
| |--B. retusum PT98
| `--B. sylvaticum S03
`--+--Poeae [Aveneae] GPWG01
| |--Poa GPWG01
| |--Avena GPWG01
| |--Agrostis GPWG01
| |--Deschampsia C06
| |--Trisetum C06
| `--Amphibromus C06
| |--A. fluitans Kirk 1884 C06
| |--A. neesii C06
| |--A. nervosus GK00
| `--A. vickeryae GK00
`--+--Triticeae [Hordeae] A05
| |--Triticum GPWG01
| |--Hordeum A05
| |--Australopyrum A05
| |--Agropyrum C06
| |--Asperella C06
| | |--A. gracilis Kirk 1895 [=Gymnostichum gracile] C06
| | `--A. laevis Petrie 1895 C06
| `--Lolium S06
| |--L. italicum C06
| |--L. multiflorum LNB03
| | |--L. m. var. multiflorum S06
| | `--L. m. var. muticum S06
| |--L. perenne LNB03
| |--L. rigidum Gaud. 1811 S06
| | |--L. r. var. rigidum S06
| | `--L. r. var. aristatum Hackel in Stuckert 1906 S06
| `--L. temulentum C55
`--Festuceae C06
|--Festuca GPWG01
|--Bromus GPWG01
|--Munroa S06
| |--M. benthamiana S06
| `--M. squarrosa [incl. Monroa argentina, Munroa mendocina] S06
|--Dactylis glomerata Linnaeus 1753 S06, PL04 [=Festuca glomerata S06]
| |--D. g. ssp. glomerata PL04
| `--D. g. ssp. hispanica (Roth) Nyman 1882 PL04
|--Koeleria C06
| |--K. albescens BBM02
| |--K. kurtzii Hackel in Kurtz 1899 (see below for synonymy) C06
| |--K. phleoides (Vill.) Pers. 1805 [=Festuca phleoides Vill. 1787] S06
| `--K. splendens Presl 1820 PL04
|--Diplachne S06
| |--D. carinata (Gris.) Hackel 1900 [=Atropis carinata Gris. 1879] S06
| |--D. chloridiformis Hackel in Stuckert 1906 S06
| |--D. dubia S06
| |--D. fusca [incl. Leptochloa fascicularis] S06
| |--D. latifolia S06
| |--D. spicata S06
| `--D. verticillata Nees & Meyen 1843 (see below for synonymy) S06
|--Atropis C06
| |--A. convoluta S06
| | |--A. c. var. convoluta S06
| | `--A. c. var. mendozina Hackel in Stuckert 1906 S06
| |--A. distans [=Glyceria distans] C06
| |--A. novae-zealandiae (Petrie) Hackel in Cheeseman 1906 (see below for synonymy) C06
| `--A. stricta (Hook.f.) Hackel in Cheeseman 1906 [=Glyceria stricta] C06
| |--A. s. var. stricta C06
| `--A. s. var. suborbicularis Hackel in Cheeseman 1906 C06
`--Briza S06
|--B. elegans [=Calotheca elegans; incl. Briza brizoides, Bromus brizoides] S06
|--B. humilis Bieberstein 1808 PL04
|--B. maxima OS04
|--B. media C55
|--B. minor Linnaeus 1753 S06
|--B. stricta S06
`--B. triloba S06
|--B. t. f. triloba S06
`--B. t. f. pumila S06
Pooideae incertae sedis:
Lamarckia aurea GPWG01, K10
Echinaria capitata GPWG01, Y98
Cynosurus GPWG01
|--C. cristatus C06
`--C. echinatus Linnaeus 1753 PL04
Anthoxanthum GPWG01
|--A. horsfieldii H03
`--A. odoratum S03
Phalarideae GPWG01
Secale cereale GPWG01, E-SM02
Sesleria GPWG01
|--S. coerulea R91
|--S. rigida H09
`--S. sphaerocephala PS98
Ammochloa GPWG01
Rostraria GPWG01
|--R. cristata PT98
|--R. pumila AGF98
`--R. salzmannii GR98
|--R. s. ssp. salzmannii [=Koeleria pubescens var. salzmanni] GR98
`--R. s. ssp. maroccana (Domin) Scholz in Greuter & Raus 1998 (see below for synonymy) GR98
Brylkinia [Brylkinieae] GPWG01
Echinopogon C06
|--E. caespitosus B00
`--E. ovatus [incl. Agrostis ovata, Cinna ovata; incl. Hystericina alopecuroides] C06
Alopecurus C06
|--A. aequalis BTA75
|--A. agrestis C06
|--A. alpinus D03
|--A. geniculatus CA27
|--A. myosuroides TG88
`--A. pratensis C55
Deyeuxia C06
Dichelachne C06
|--D. crinita (see below for synonymy) C06
| |--D. c. var. crinita C06
| `--D. c. var. intermedia Hack. in Cheeseman 1906 C06
`--D. sciurea (see below for synonymy) C06
|--D. s. var. sciurea C06
`--D. s. var. inaequiglumis Hack. in Cheeseman 1906 C06
Simplicia laxa Kirk 1897 C06
Calamagrostis C06
Piptochaetium S06
Muehlenbergia S06
|--M. diffusa Willd. 1797 S06
|--M. nana Benth. 1846 S06
`--M. peruviana [=Clomena peruviana Beauv. 1812; incl. M. clomena] S06
Lycurus S06
|--L. alopecuroides S06
`--L. phleoides S06
Phleum S06
|--P. alpinum Lin. 1753 S06
|--P. arenarium G88
|--P. graecum PT98
| |--P. g. ssp. graecum PT98
| `--P. g. ssp. aegaeum PT98
|--P. paniculatum H91
`--P. pratense LNB03
Epicampes S06
|--E. arundinacea [=Sporobolus arundinaceus] S06
`--E. coerulea S06
|--E. c. var. coerulea S06
`--E. c. var. submutica Hackel in Stuckert 1906 S06
Polypogon S06
Atropis novae-zealandiae (Petrie) Hackel in Cheeseman 1906 [=Glyceria novae-zealandiae Petrie 1901; incl. Poa walkeri Kirk 1885] C06
Dichelachne crinita [=Agrostis crinita, Anthoxanthum crinitum; incl. Dichelachne forsteriana Trin. & Rupr. 1842, D. hookeriana Trin. & Rupr. 1842] C06
Dichelachne sciurea [=Agrostis sciurea; incl. Stipa micrantha, D. montana, D. sieberiana Trin. & Rupr. 1842] C06
Diplachne verticillata Nees & Meyen 1843 [incl. Leptochloa imbricata Turb. 1880, D. imbricata (Thurb.) Scribn. 1891, Rhabdochloa imbricata (Thurb.) OK. 1891] S06
Koeleria kurtzii Hackel in Kurtz 1899 [incl. K. cristata Gris. 1874 (preoc.), K. bergii var. fallacina Domin 1906, K. micrathera Griseb. 1879 non Trisetum micratherum] C06
Pooideae [Agrostideae, Agrostidoideae, Anthoxanthoideae, Avenoideae, Cynosuroideae, Echinarioideae, Festucoideae, Glycerioideae, Hordeoideae, Phalaroideae, Secaloideae]
Rostraria salzmannii ssp. maroccana (Domin) Scholz in Greuter & Raus 1998 [=Koeleria salzmannii var. maroccana Domin 1907, K. pubescens var. maroccana] GR98
Trachynia platystachya (Balansa) Scholz in Greuter & Raus 1998 [=Brachypodium distachyon var. platystachyon Balansa ex Cosson in Cosson & Durieu 1855, B. distachyon subvar. platystachyon (Balansa) St.-Yves 1934] GR98
*Type species of generic name indicated
REFERENCES
[A05] Aagesen, L. 2005. Direct optimization, affine gap costs, and node stability.
Molecular Phylogenetics and Evolution 36: 641–653.
[AGF98] Abd El-Ghani, M. M., & A. G. Fahmy. 1998. Composition of and changes in the spontaneous flora of Feiran Oasis, S Sinai, Egypt, in the last 60 years.
Willdenowia 28: 123–134.
[BTA75] Baker, E. W., D. M. Tuttle & M. J. Abbatiello. 1975. The false spider mites of northwestern and north central Mexico (Acarina: Tenuipalpidae).
Smithsonian Contributions to Zoology 194: 1–23.
[BBM02] Bonte, D., L. Baert & J.-P. Maelfait. 2002. Spider assemblage structure and stability in a heterogeneous coastal dune system (Belgium).
Journal of Arachnology 30 (2): 331–343.
[B00] Braby, M. F. 2000.
Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).
[C55] Candolle, A. de. 1855.
Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 2. Librairie de Victor Masson: Paris.
[C06] Cheeseman, T. F. 1906.
Manual of the New Zealand Flora. John Mackay, Government Printer: Wellington.
[CA27] Cockayne, L., & H. H. Allan. 1927. Notes on New Zealand floristic botany, including descriptions of new species, &c. (No. 5).
Transactions and Proceedings of the New Zealand Institute 57: 48–72.
[C74] Coineau, Y. 1974. Éléments pour une monographie morphologique, écologique et biologique des Caeculidae (Acariens).
Mémoires du Muséum National d'Histoire Naturelle, nouvelle série, Série A, Zoologie 81: 1–299, 24 pls.
[CL86] Collins, E. I., & R. W. Lichvar. 1986. Vegetation inventory of current and historic black-footed ferret habitat in Wyoming.
Great Basin Naturalist Memoirs 8: 85–93.
[D03] Dusén, P. 1903. The vegetation of western Patagonia.
In: Scott, W. B. (ed.)
Reports of the Princeton University Expeditions to Patagonia, 1896–1899 vol. 8.
Botany pp. 1–34. The University: Princeton (New Jersey).
[E-SM02] Ellneskog-Staam, P., & A. Merker. 2002. Screening for resistance to powdery mildew and brown rust in wheat-
Leymus racemosus and wheat-
Thinopyrum junceiforme alloploids.
Acta Agricultura Scandinavica Section B—Soil and Plant Science 52: 158–161.
[GK00] Gibson, N., & G. J. Keighery. 2000. Flora and vegetation of the Byenup-Muir reserve system, south-west Western Australia.
CALMScience 3 (3): 323–402.
[GPWG01] Grass Phylogeny Working Group. 2001. Phylogeny and subfamilial classification of the grasses (Poaceae).
Annals of the Missouri Botanical Garden 88 (3): 373–457.
[GR98] Greuter, W., & T. Raus (eds.) 1998. Med-Checklist Notulae, 17.
Willdenowia 28: 163–174.
[G88] Grubb, P. J. 1988. The uncoupling of disturbance and recruitment, two kinds of seed bank, and persistence of plant populations at the regional and local scales.
Annales Zoologici Fennici 25: 23–26.
[H03] Heads, M. 2003. Ericaceae in Malesia: vicariance biogeography, terrane tectonics and ecology.
Telopea 10 (1): 311–449.
[HE80] Healy, A. J., & E. Edgar. 1980.
Flora of New Zealand vol. 3.
Adventive cyperaceous, petalous and spathaceous monocotyledons. P. D. Hasselberg, Government Printer: Wellington (New Zealand).
[H09] Heltmann, H. 2009. Der Königstein (Piatra Craiului), die Perle der Burzenländer Gebirge.
Mauritiana 20 (3): 515–527.
[H91] Hubálek, Z. 1991. Biogeographic indication of natural foci of tick-borne infections.
In: Dusbábek, F., & V. Bukva (eds)
Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 255–260. SPB Academic Publishing: The Hague.
[K10] Keighery, G. 2010. The naturalised vascular plants of the Pilbara region, Western Australia.
Records of the Western Australian Museum Supplement 78 (1): 299–311.
[LNB03] Ledeganck, P., I. Nijs & L. Beyens. 2003. Plant functional group diversity promotes soil protist diversity.
Protist 154 (2): 239–249.
[OS04] Obbens, F. J., & L. W. Sage. 2004. Vegetation and flora of a diverse upland remnant of the Western Australian wheatbelt (Nature Reserve A21064).
Journal of the Royal Society of Western Australia 87 (1): 19–28.
[O88] Ohba, H. 1988. The alpine flora of the Nepal Himalayas: an introductory note.
In: Ohba, H., & S. B. Malla (eds)
The Himalayan Plants vol. 1.
The University Museum, University of Tokyo, Bulletin 31: 19–46.
[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi.
Willdenowia 28: 95–116.
[PL04] Pohl, G., & I. Lenski. 2004. Zur Verbreitung und Vergesellschaftung von
Pennisetum orientale Rich. in Nordeuböa (Griechenland) (Poaceae, Paniceae).
Senckenbergiana Biologica 83 (2): 209–223.
[PS98] Prosser, F., & S. Scortegagna. 1998.
Primula recubariensis, a new species of
Primula sect.
Auricula Duby endemic to the SE Prealps, Italy.
Willdenowia 28: 27–46.
[R91] Reutimann, P. 1991. Alpine oribatid species and plant decomposition: feeding and faeces production.
In: Dusbábek, F., & V. Bukva (eds)
Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 417–422. SPB Academic Publishing: The Hague.
[S03] Singh, J. N. 2003. Grasses and their hydro-edaphic characteristics in the grassland habitat of Nilgiris Biosphere Reserve, Tamil Nadu.
Bulletin of the Botanical Survey of India 45 (1–4): 143–164.
[S06] Stuckert, T. 1906. Segunda contribución al conocimiento de las gramináceas Argentinas.
Anales del Museo Nacional de Buenos Aires, serie 3, 6: 408–555.
[TG88] Tepfer, D., A. Goldmann, N. Pamboukdjian, M. Maille, A. Lepingle, D. Chevalier, J. Dénarié & C. Rosenberg. 1988. A plasmid of
Rhizobium meliloti 41 encodes catabolism of two compounds from root exudate of
Calystegium sepium. Journal of Bacteriology 170 (3): 1153–1161.
[Y98] Yannitsaros, A. 1998. Additions to the flora of Kithira (Greece) I.
Willdenowia 28: 77–94.
Last updated: 8 December 2017.