Chloridoideae

Spartina anglica, photographed by Jürgen Howaldt.


Belongs within: Poaceae.
Contains: Eragrostis.

The Chloridoideae is a clade of mostly C4 grasses supported by molecular analysis, but as yet with no identified morphological synapomorphies. However, the clade of chloridoids excluding 'Merxmuellera' rangei and Centropodia is supported by the presence of 'chloridoid-type' microhairs with short and wide apical cells (Grass Phylogeny Working Group 2001). Major genera within the Chloridoideae include Spartina, the cordgrasses, which are found mostly in coastal salt marshes on either side of the Atlantic, and Sporobolus, the dropseed grasses, which are found in prairies and savannahs.

Characters (from Grass Phylogeny Working Group 2001): Plants annual or perennial (rhizomatous, stoloniferous, caespitose or decumbent), herbaceous (rarely woody), of dry climates, especially in the tropics and subtropics, also found in the temperate zone. Culms solid or hollow. Leaves distichous; abaxial ligule usually absent, rarely present as a line of hairs; adaxial ligule a fringed or less commonly unfringed membrane; blades relatively narrow, without pseudopetioles, venation parallel; sheaths usually non-auriculate. Inflorescences paniculate, paniculate with spicate branches, racemose, or spicate, bracts outside of the spikelets absent. Spikelets bisexual or sometimes unisexual (if so the plants dioecious or monoecious), with glumes 2, rarely a sterile lemma, and female-fertile florets 1 to many, apical reduction usually present, usually laterally compressed, sometimes dorsally compressed, usually disarticulating above the glumes (below in a few Eragrostis species); lemma lacking uncinate macrohairs, if awned, the awns single or if multiple, lacking a basal column; palea well developed; lodicules 2 or absent, fleshy, glabrous; stamens 1 to 3; ovary glabrous, apical appendage absent, haustorial synergids absent, styles 2, free, close, stigmas 2. Caryopsis with the pericarp often free or loose; hilum short; endosperm hard, without lipid, containing simple or compound starch grains; embryo large or rarely small, epiblast present or rarely absent, scutellar cleft present, mesocotyl internode elongated, embryonic leaf margins meeting or rarely overlapping. Foliar mesophyll usually radiate, without an adaxial palisade layer, fusoid cells absent, arm cells absent; Kranz anatomy present; midrib simple; adaxial bulliform cells present. Foliar stomata with dome-shaped or triangular subsidiary cells; bicellular microhairs present, usually chloridoid-type; papillae absent or present.

<==Chloridoideae [Eragrostoideae, Pappophoroideae]
    |--+--‘Merxmuellera’ rangei GPWG01
    |  `--Centropodia glauca GPWG01
    `--+--+--Pappophorum [Pappophoreae] GPWG01
       |  |    `--P. bicolor GPWG01
       |  `--+--Eragrostis GPWG01
       |     `--Uniola paniculata GPWG01
       `--+--Distichlis GPWG01
          |    |--D. scoparia CLB06
          |    `--D. spicata (Linnaeus) Greene 1887 BF89
          |         |--D. s. ssp. spicata GPWG01
          |         `--D. s. ssp. stricta GPWG01
          `--+--Zoysia matrella GPWG01
             `--+--Sporobolus GPWG01
                |    |--S. africanus P82
                |    |--S. airoides GPWG01
                |    |--S. capensis G60
                |    |--S. giganteus GPWG01
                |    |--S. helvolus PP07
                |    |--S. heterolepis B75
                |    |--S. indicus GPWG01
                |    |    |--S. i. var. indicus S03
                |    |    `--S. i. var. purpureosuffusus S03
                |    |--S. littoralis C55
                |    |--S. minutiflorus C55
                |    |--S. nebulosus CV06
                |    |--S. spicata BB01
                |    |--S. tenuissimus PP07
                |    |--S. virginicus SM06 [=Agrostis virginica C55]
                |    `--S. wallichi Munro ex von Trinius 1889 SY02
                `--Spartina GPWG01
                     |--S. alterniflora Loisel. 1807 BF89
                     |--S. anglica C-S98
                     |--S. foliosa CH97
                     |--S. gracilis GPWG01
                     |--S. juncea C55
                     |--S. maritima C-S98
                     |--S. michaxiana B75
                     |--S. patens (Ait) Muhl 1817 BF89
                     |--S. pectinata GPWG01
                     |--S. spartinae M83
                     |--S. stricta D37
                     `--S. townsendii D37

Chloridoideae incertae sedis:
  Chloris SY02
    |--C. cucullata GPWG01
    |--C. dolichostachya SR07
    |--C. flabellata CV06
    |--C. gayana BB99
    |--C. neglecta M83
    |--C. pycnothrix von Trinius 1824 SY02
    |--C. roxburghiana S03
    `--C. virgata VB02
  Tridens flavus GPWG01
  Muhlenbergia capillaris GPWG01, M83
  Eustachys GPWG01
  Cynodon [Cynodonteae] GPWG01
    |--C. dactylon (Linnaeus) Persoon 1805 PL04
    `--C. transvaali BC01
  Lepturus [Leptureae] GPWG01
    `--L. repens C78
  Orcuttia [Orcuttieae] GPWG01

*Type species of generic name indicated

REFERENCES

[BB01] Bauer, R., D. Begerow, A. Nagler & F. Oberwinkler. 2001. The Georgefischeriales: A phylogenetic hypothesis. Mycological Research 105 (4): 416-424.

[BB99] Blumenthal, M. J., A. M. Bowman, A. Cole, R. M. Jones, W. M. Kelman, T. E. Launders & H. I. Nicol. 1999. Establishment, growth and persistence of greater lotus (Lotus uliginosus) at six sites in eastern Australia. Australian Journal of Experimental Agriculture 39: 819-827.

[B75] Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa. Special Publications, The Museum, Texas Tech University 9: 1-184.

[BC01] Briese, D. T., & J. M. Cullen. 2001. The use and usefulness of mites in biological control of weeds. In Acarology: Proceedings of the 10th International Congress (R. B. Halliday, D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff, eds) pp. 453-463. CSIRO Publishing: Melbourne.

[BF89] Burnham, B. H. & P. E. Fell. 1989. Distribution of Melampus bidentatus (Say) and Succinea wilsoni (Lea) within a tidal marsh in eastern Connecticut. Nautilus 103 (3): 109-112.

[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.

[CH97] Castro, P., & M. E. Huber. 1997. Marine Biology, 2nd ed. WCB McGraw-Hill: Boston.

[C-S98] Cavalier-Smith, T. 1998. A revised six-kingdom system of life. Biological Reviews 73: 203-266.

[C78] Clunie, N. M. U. 1978. The vegetation. In Handbooks of the Flora of Papua New Guinea vol. 1 (J. S. Womersley, ed.) pp. 1-11. Melbourne University Press: Carlton South (Australia).

[CV06] Craven, P., & P. Vorster. 2006. Patterns of plant diversity and endemism in Namibia. Bothalia 36 (2): 175-189.

[CLB06] Cutrera, A. P., E. A. Lacey & C. Busch. 2006. Intraspecific variation in effective population size in talar tuco-tucos (Ctenomys talarum): the role of demography. Journal of Mammalogy 87 (1): 108-116.

[D37] Dobzhansky, T. 1937. Genetics and the Origin of Species. Columbia University Press: New York.

[G60] Gillham, M. E. 1960. Vegetation of Little Brother Island, Cook Strait, in relation to spray-bearing winds, soil salinity, and pH. Transactions of the Royal Society of New Zealand 88 (3): 405-424.

[GPWG01] Grass Phylogeny Working Group. 2001. Phylogeny and subfamilial classification of the grasses (Poaceae). Annals of the Missouri Botanical Garden 88 (3): 373-457.

[M83] Myers, R. L. 1983. Site susceptibility to invasion by the exotic tree Melaleuca quinquenervia in southern Florida. Journal of Applied Ecology 20: 645-658.

[PP07] Pandey, R. P. & P. M. Padhye. 2007. Studies on phytodiversity of Arid Machia Safari Park-Kailana in Jodhpur (Rajasthan). Bulletin of the Botanical Survey of India 49 (1-4): 15-78.

[P82] Pickard, J. 1982. Catastrophic disturbance and vegetation on Little Slope, Lord Howe Island. Australian Journal of Ecology 7: 161-170.

[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.

[SY02] Salunkhe, C. B., & S. R. Yadav. 2002. Additions to the grasses of Maharashtra. Journal of the Bombay Natural History Society 99 (3): 569-573.

[SR07] Sankar, R. V., K. Ravikumar, N. M. Ganesh Babu & D. K. Ved. 2007. Botany of Anapady MPCA, Palghat district, Kerala with special emphasis on species of conservation concern. Bulletin of the Botanical Survey of India 49 (1-4): 165-172.

[SM06] Semeniuk, C. A., L. A. Milne, P. Ladd & V. Semeniuk. 2006. Pollen in the surface sediments of wetlands in the Becher Point area, southwestern Australia: a baseline for use in interpreting Holocene sequences. Journal of the Royal Society of Western Australia 89 (1): 27-43.

[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.

[VB02] Vijay, S. K., & T. N. Bhardwaja. 2002. Vegetation and phenodynamics of wetlands of central Rajasthan. Journal of the Bombay Natural History Society 99 (3): 573-581.

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