13.032   ALOPECURUS L.

DRAFT TREATMENT. Please send comments to Mary Barkworth.
William J. Crins

Plants annual or perennial; sometimes shortly rhizomatous, cespitose or occasionally the culms solitary. Culms 5-110 cm, erect or decumbent, occasionally cormlike at the base; nodes glabrous. Leaves inserted mostly on the lower 1/2 of the culms; sheaths open; auricles absent; ligules 0.6-6.5 mm, truncate to acute, membranous, dorsally puberulent or glabrous, entire to lacerate; blades 0.7-12 mm wide, flat or involute, glabrous or scabrous, those of the uppermost leaves sometimes short or absent. Inflorescences terminal panicles, spikelike, capitate to cylindrical; branches usually less than 5 mm, lower branches sometimes to 2 cm; disarticulation below the glumes. Spikelets 1.8-7 mm, pedicellate, strongly laterally compressed, with 1 floret; rachilla prolongations absent. Glumes equaling or exceeding the florets, membranous or coriaceous, free or connate basally to more than 1/2 their length, 3-veined, keeled, keels ciliate, at least basally, apices obtuse to acute or shortly awned; calluses blunt, glabrous; lemmas membranous, margins often connate basally to 1/2 their length, keeled, indistinctly 3-5-veined, apices truncate to acute, awned dorsally, awns arising from just above the base to about midpoint, geniculate or straight; paleas absent or greatly reduced; lodicules absent; anthers 3, 0.3-4.1 mm; ovaries glabrous; styles fused, with 2 branches. Caryopses glabrous; hila short. x = 7. Name from the Greek alopex, fox, and oura, tail, referring to the cylindrical panicles.

Alopecurus is a genus of 36 species that grow primarily in open, mesic habitats, native to the northern temperate zone and South America. Four species are native to the Flora region, four were introduced and have become established, and two were introduced and are not known to persist. Some species, including some native to the Flora region, were introduced as pasture grasses outside of their native ranges. Of these, only A. pratensis has become widely naturalized.

Some species of Alopecurus can appear similar to Phleum, which has truncate glumes that are abruptly awned or mucronate, lemmas without awns and keels, and well-developed paleas; Alopecurus has glumes that are obtuse to acute and gradually awned or unawned, lemmas with both awns and keels, and paleas that are absent or greatly reduced.


SELECTED REFERENCES Dogan, M. 1999. A concise taxonomic revision of the genus Alopecurus L. (Gramineae). Turk. J. Bot. 23:245-262; Moyer, J.R. and A.L. Boswall. 2002. Tall fescue or creeping foxtail suppresses foxtail barley. Canad. J. Pl. Sci. 82:89-92; Tzvelev, N.N. 1995. Alopecurus. Pp. 106-114 in J.G. Packer (ed., English edition). Flora of the Russian Arctic, vol. 1, trans. G.C.D. Griffiths. University of Alberta Press, Edmonton, Alberta, Canada [English translation of A.I. Tolmachev, ed. (1964). Arkticheskaya Flora SSSR, vol. 2. Nauka, Leningrad, Russia].

1
Glume keels winged, ciliate near the base, the glumes glabrous or the veins pubescent (2)
Glume keels without wings, ciliate, the glumes sparsely to densely pubescent elsewhere (3)
2
Glumes 4.5-7.5 mm long, connate to 1/2 of their length, the apices acute, convergent to parallel; lemma apices acute ..... 9. A. myosuroides
Glumes 3-4.5 mm long, connate for more than 1/2 of their length, the apices obtuse, mucronate, divergent; lemma apices truncate ..... 10. A. creticus
3
Plants annual, without rhizomes, not rooting at the lower nodes; blades 1-16 cm long, 0.9-4 mm wide; culms 5-50 cm tall (4)
Plants perennial, often rhizomatous, sometimes rooting at the lower nodes; blades 2-40 cm long, 1-12 mm wide; culms 5-110 cm tall (6)
4
Glumes 5-6.4 mm long, coriaceous and dilated in the lower 1/2; glume and lemma apices acute to acuminate; anthers about 3 mm long ..... 6. A. rendlei
Glumes 2.1-5 mm long, membranous, not dilated below; glume and lemma apices obtuse; anthers 0.4-1 mm long (5)
5
Upper sheaths conspicuously inflated; glumes 3-5 mm long; lemmas 3-5 mm long, awns exceeding the lemmas by 3-6 mm; panicles 5.5-10 mm wide (excluding the awns) ..... 7. A. saccatus
Upper sheaths not inflated; glumes 2.1-3.1 mm long; lemmas 1.9-2.7 mm long, awns exceeding the lemmas by 1.6-4 mm; panicles 3-6 mm wide (excluding the awns) ..... 8. A. carolinianus
6
Glumes 1.8-3.1(3.7) mm long, the apices obtuse; anthers 0.5-2.2 mm long (7)
Glumes 3-6 mm long, the apices acute; anthers 2-4 mm long (8)
7
Awns geniculate, exceeding the lemmas by 1.2-4 mm; anthers (0.9)1.4-2.2 mm long ..... 4. A. geniculatus
Awns straight, not exceeding the lemmas, or exceeding them by less than 2.5 mm; anthers 0.5-1.2 mm long ..... 5. A. aequalis
8
Lemma margins connate only at the base; glumes densely covered with long wooly hairs ..... 3. A. borealis
Lemma margins connate for at least 1/3 of their length; glumes with long hairs mainly restricted to the veins (9)
9
Lemma apices acute; glume apices parallel or convergent ..... 1. A. pratensis
Lemma apices obtuse to truncate; glume apices divergent ..... 2. A. arundinaceus


1.   Alopecurus pratensis L.
Meadow Foxtail, Vulpin des Prés

Plants perennial; shortly rhizomatous. Culms 30-110 cm, erect. Ligules 1.5-3 mm, obtuse to truncate; blades 6-40 cm long, 1.9-8 mm wide; upper sheaths not or scarcely inflated. Panicles 3.5-9 cm long, 6-10 mm wide. Glumes 4-6 mm, connate in the lower 1/4, membranous, sides pubescent, keels not winged, finely ciliate, apices acute, parallel or convergent; lemmas 4-6 mm, connate in the lower 1/3, glabrous or the keels sometimes ciliate apically, apices acute, awns 5-10.5 mm, geniculate, exceeding the lemmas by (1)2.2-5.5 mm; anthers 2-4 mm, yellowish, orange, reddish, or purplish, varying within a population. Caryopses about 1-1.2 mm. 2n = 28, 42.

Alopecurus pratensis is native from temperate northern Eurasia south to North Africa. It is now widely naturalized in temperate regions throughout the world. It grows in poorly to somewhat drained soils in meadows, riverbanks, lakesides, ditches, roadsides and fence rows. It has been widely introduced as a pasture grass; it may also have become established from ballast or imported hay. The earliest collections are from coastal New England; it is now known from most provinces and states.


2.   Alopecurus arundinaceus Poir.
Creeping Meadow Foxtail, Vulpin Roseau

Plants short-lived perennials; rhizomatous. Culms 30-110 cm, erect. Ligules 1.3-5 mm, truncate; blades 6-40 cm long, 3-12 mm wide; upper sheaths somewhat inflated. Panicles 3-10 cm long, 7-13 mm wide. Glumes 3.6-5 mm, connate in the lower 1/3, membranous, sparsely pubescent, keels not winged, ciliate, apices acute, diverging, pale green to lead-gray; lemmas 3.1-4.5 mm, connate for at least 1/3 of their length, glabrous or sometimes with scattered hairs near the apices, apices truncate to obtuse, awns 1.5-7.5 mm, geniculate, exceeding the lemmas by 0-3 mm; anthers 2.2-3.5 mm. 2n = 26, 28, 30.

Alopecurus arundinaceus is native to Eurasia, extending north of the Arctic Circle and south to the Mediterranean. It grows on wet, moderately acid to moderately alkaline soils, on flood plains, vernal ponds, and along rivers, streams, bogs, potholes, and sloughs. It was introduced for pasture in North Dakota and now occurs more widely, having been promoted as a forage species, and is sometimes used in seed mixtures for revegetation projects. It was evaluated for revegetation in Alberta, but there is no evidence that it was ever actually used in that province. Alopecurus arundinaceus was found to suppress Hordeum jubatum, a troublesome, unpalatable, weedy species, in irrigated pastures (Moyer and Boswall 2002).


3.   Alopecurus borealis Trin.
Alpine Foxtail, Boreal Foxtail, Vulpin Alpin, Vulpin Boréale

Plants perennial; shortly rhizomatous. Culms (6)10-80 cm, erect or decumbent. Ligules 1-2 mm, truncate; blades 4-22 cm long, 2.5-7 mm wide; upper sheaths inflated. Panicles 1-5 cm long, 8-14 mm wide. Glumes 3-5 mm, connate at the base, membranous, densely pilose throughout, keels not winged, ciliate, apices acute and parallel; lemmas 2.5-4.5 mm, connate at the base, glabrous below, finely pubescent apically, apices usually obtuse, occasionally truncate, awns 2-6(8) mm, geniculate, exceeding the lemmas by 0-5 mm; anthers 2.3-3 mm, yellow. Caryopses 0.7-2 mm. 2n = 98, 100, 105, 112, 117, 119, ca. 120.

Alopecurus borealis has an arctic-alpine to subalpine circumpolardistribution, but it has not been found in Scandinavia or Iceland. It grows primarily in wet soils in tundra, meadows, along streams, shorelines, gravelbars, and floodplains, and occasionally in somewhat drier forest openings, in fine or silty to stony soils or moss. It is sometimes co-dominant with Dupontia fisheri in the arctic and subarctic portion of its range. The anthocyanic tint of the plant as a whole greatly increases to the north.

The morphological variability in Alopecurus borealis has prompted recognition of several segregate taxa. Alopecurus stejnegeri Vasey and A. occidentalis Scribn. & Tweedy are two of the more conspicuous extremes. The former are small plants occurring on enriched sites in the Arctic, usually around sea bird or seal colonies where high nutrient levels produce lush vegetative growth; the latter refers to tall-stemmed plants found in the Rocky Mountains. Because such plants are simply extremes in a continuum of variation, they do not merit taxonomic recognition.

Alopecurus borealis is listed in many floras as A. alpinus Sm., a name that had previously been applied to another species. The correct name is A. borealis.


4.   Alopecurus geniculatus L.
Water Foxtail, Vulpin Géniculé

Plants perennial; cespitose. Culms (5)10-60 cm, erect or decumbent, rooting at the nodes. Ligules 2-5 mm, obtuse; blades 2-12 cm long, 1-4(7) mm wide; upper sheaths somewhat inflated. Panicles 1.5-7 cm long, 4-8 mm wide. Glumes 1.9-3.5 mm, connate atthe base, membranous, pubescent, keels not winged, ciliate, apices obtuse, parallel, often purplish; lemmas 2.5-3 mm, connate in the lower 1/2, glabrous or with a few scattered hairs at the apices, apices truncate to obtuse, awns 3.5-5(6) mm, geniculate, exceeding the lemmas by (1.2)2-4 mm; anthers (0.9)1.4-2.2 mm, yellow. Caryopses 1-1.5 mm. 2n = 28.

Alopecurus geniculatus is native to Eurasia and parts of North America, growing in shallow water, ditches, open wet meadows, shores, and stream banks from the lowland to montane zones. It has been naturalized in eastern North America. The status of populations in the west, including the Queen Charlotte Islands, British Columbia, is less certain. Many occur in moist sites within native rangeland, but these areas have also been affected by European settlement, although less intensively and for a shorter period than those in eastern North America.

Alopecurus haussknechtianus Asch. & Graebn. is a hybrid between A. geniculatus and A. aequalis, which occurs fairly frequently in areas of sympatry, particularly in drier midcontinental areas in Alberta to Saskatchewan, south to Arizona and New Mexico. The hybrids appear to have 2n = 14; they are sterile.


5.   Alopecurus aequalis Sobol.

Plants perennial; cespitose. Culms 9-75 cm, erect or decumbent. Ligules 2-6.5 mm, obtuse, puberulent dorsally; blades 2-10 cm long, 1-5(8) mm wide; upper sheaths not inflated. Panicles 1-9 cm long, 3-9 mm wide. Glumes 1.8-3.7 mm, connate near the base, membranous, pubescent on the sides, keels not winged, ciliate, apices obtuse, sometimes erose, pale green, occasionally purplish; lemmas 1.5-2.5(3.5) mm, connate in the lower 1/3-1/2, glabrous, apices obtuse, awns 0.7-3 mm, straight, exceeding the lemmas by 0-2.5 mm; anthers 0.5-1.2 mm, usually pale to deep yellow or orange, rarely purple. Caryopses 1-1.8 mm. 2n = 14, 28.

Alopecurus aequalis is native to temperate zones of the Northern Hemisphere. It generally grows in wet meadows, forest openings, shores, springs, and along streams, as well as in ditches, along roadsides, and in other disturbed sites, from sea level to subalpine elevations.

Alopecurus aequalis is the most widespread and variable species of Alopecurus in the Flora region. Despite its variability, only the phenotype found in the low marshes of Marin and Sonoma counties, California merits formal recognition. Some high elevation plants of the Sierra Nevada have unusually long awns (exserted by up to 1.5 mm) and anthocyanic spikelets but, like the semi-aquatic ecotype A. aequalis var. natans (Wahlenb.) Fernald, they do not warrant taxonomic recognition.

Alopecurus haussknechtianus Asch. & Graebn. is a hybrid between A. aequalis and A. geniculatus, which occurs fairly frequently in areas of sympatry, particularly in drier midcontinental areas in Alberta to Saskatchewan, south to Arizona and New Mexico. The hybrids apparently have 2n = 14; they are sterile.

1
Panicles 3-6 mm wide; glumes 1.8-3 mm long; awns not exceeding the lemmas or doing so by less than 1 mm; anthers 0.5-0.9 mm long ..... var. aequalis
Panicles 4-9 mm wide; glumes to 3.7 mm long; awns exceeding the lemmas by 1-2.5 mm; anthers 1-1.2 mm long ..... var. sonomensis


Alopecurus aequalis var. aequalis
Shortawn Foxtail, Vulpin à Courtes Arêtes

Culms 9-75 cm. Blades 1-5(8) mm wide. Panicles 1-9 mm long, 3-6 mm wide. Spikelets usually not purplish-tinged; glumes 1.8-3 mm; awns not exceeding the lemmas or exceeding them by less than 1 mm; anthers 0.5-0.9 mm.

Alopecurus aequalis var. aequalis is the widespread variety in the Flora region. It is listed as threatened in the state of Connecticut.


Alopecurus aequalis var. sonomensis P. Rubtzoff
Sonoma Shortawn Foxtail

Culms 30-75 cm. Blades to 7.5 mm wide. Panicles 2.5-9 cm long, 4-9 mm wide. Spikelets violet-gray to purplish-tinged, especially towards the apices; glumes to 3.7 mm; awns exceeding the lemmas by 1-2.5 mm; anthers 1-1.2 mm.

Alopecurus aequalis var. sonomensis is endemic to Marin and Sonoma counties, California, where it grows in shallow water and marshy or moist ground, usually in the open. It is listed as an endangered species by the U.S. Fish and Wildlife Service.


6.   Alopecurus rendlei Eig
Rendle's Meadow Foxtail

Plants annual; tufted. Culms 8-30(40) cm, erect or decumbent. Ligules 0.6-3 mm, obtuse; blades 1-16 cm long, 1-3 mm wide; upper sheaths inflated. Panicles 1-3.2 cm long, 5-11 mm wide. Glumes 5-6.4 mm, connate in the lower 1/4-1/3, dilated and coriaceous in the lower 1/2, pilose in the lower 1/2, indurate and constricted above the middle, keels not winged, ciliate on all veins, apices acute, slightly divergent; lemmas 5.6-6.4 mm, connate basally, glabrous or puberulent apically, apices coriaceous, acuminate, awns 6.5-18 mm, geniculate, exceeding the lemmas by 0.8-7.2 mm; anthers about 3 mm. Caryposes 3.5-4.5 mm. 2n = 14.

Alopecurus rendlei is native to wet meadows, and adventive in roadsides and waste places, in southern and western Europe. It was found growing on ballast in Philadelphia, Pennsylvania in 1880; it has not been collected in North America since that time.


7.   Alopecurus saccatus Vasey
Pacific Meadow Foxtail

Plants annual; tufted. Culms 12-45 cm, erect or decumbent. Ligules 1.5-5.5 mm, obtuse; blades 4-12 cm long, 1.2-4 mm wide; upper sheaths conspicuously inflated. Panicles 1.5-6.5 cm long, 5.5-10 mm wide, often dense. Glumes 3-5 mm, connate at the base, not dilated, membranous, pubescent, keels not winged, ciliate, apices obtuse; lemmas 3-5 mm, connate in the lower 1/3-1/2, glabrous, apices obtuse, awns 6-10 mm, geniculate, exceeding the lemmas by 3-6 mm; anthers 0.7-1.8 mm, yellow to rusty-brown. Caryopses 1.5-2 mm. 2n = unknown.

Alopecurus saccatus is a native annual that inhabits moist, open meadows, valley plains, and vernal pools at elevations below 700 m from Washington to California. Segregates have been treated as species in the past, but the variation between them appears to be continuous, and no habitat differentiation is evident.


8.   Alopecurus carolinianus Walter
Tufted Foxtail

Plants annual; tufted. Culms 5-50 cm, erect or decumbent. Ligules 2.8-4.5 mm, obtuse; blades 3-15 cm long, 0.9-3 mm wide; upper sheaths not inflated. Panicles 1-7 cm long, 3-6 mm wide, always dense. Glumes 2.1-3.1 mm, connate at the base, membranous throughout, sparsely pubescent, not inflated below, keels not winged, ciliate, apices obtuse, pale green to pale yellow; lemmas 1.9-2.7 mm, connate in the lower 1/2, glabrous, apices obtuse, awns 3-6.5 mm, geniculate, exceeding the lemmas by 1.6-4 mm; anthers 0.3-1 mm, yellow or orange. Caryopses 1-1.5 mm. 2n = 14.

Alopecurus carolinianus is native to the central plains, Mississippi valley, and southeastern United States, where it is common in wet meadows, ditches, wetland edges, and other moist, open habitats; it is occasionally a weed of rice fields. At the northern limit of its range it is clearly adventive, growing in gardens and nurseries. It also occurs in arid areas of the prairies and southwest, growing sporadically along sloughs and in ditches and vernal pools. Whether such populations are native or naturalized is not clear.


9.   Alopecurus myosuroides Huds.
Blackgrass, Slender Meadow Foxtail

Plants annual; tufted. Culms (10)40-85 cm, erect. Ligules 2-6 mm, obtuse; blades (2)3.5-6 mm wide; upper sheaths somewhat inflated. Panicles 4-12 cm long, 3-7 mm wide. Glumes 4.5-7.5 mm, connate in the lower 1/2, coriaceous, sides glabrous, keels winged, ciliate, scabrous apically, lateral veins ciliate basally or glabrous, apices acute, convergent to parallel; lemmas 4-7 mm, connate in the lower 1/3-1/2, glabrous, apices acute, awns to 12 mm, geniculate, exceeding the lemmas by 3-6 mm; anthers 2.4-4.1 mm, yellow. 2n = 14, 28.

Alopecurus myosuroides is native to Eurasia and grows in moist meadows, deciduous forests, and cultivated or disturbed ground. A significant weed species in temperate cereal crops, it is one of the most damaging weeds of winter cereals in England. It has been introduced repeatedly as a weed of cultivation into many parts of the Flora region, but apparently has not spread to a large degree out of cultivation. Alopecurus myosuroides has been listed as a noxious weed in the state of Washington, one of the states where winter wheat is a major crop.


10.   Alopecurus creticus Trin.
Cretan Meadow Foxtail

Plants annual; tufted. Culms 7-21(40) cm, erect. Ligules 1.4-2.3 mm, obtuse; blades 1-8 cm long, 1-3.5 mm wide; upper sheaths usually inflated. Panicles 2-3.5 cm long, 4-5.5 mm wide, dense. Glumes 3-4.5 mm, connate for 1/2-4/5 of their length, coriaceous, keels winged, ciliate, obtuse, mucronate, divergent; lemmas 2.5-3.5 mm, connate at the base, glabrous, apices truncate, awns 3-6 mm, geniculate, exceeding the lemmas by 0-4 mm; anthers 1.2-2.5 mm. 2n = unknown.

Alopecurus creticus is native to marshes and wet places in the southern part of the Balkan Peninsula. It was discovered on ballast dumps in Philadelphia, Pennsylvania in the nineteenth century; it has not persisted in the Flora region.