Spartina patens (Aiton) Muhl., Descr. Gram. 55. 1817.

Saarela, Jeffery M., 2012, Taxonomic synopsis of invasive and native Spartina (Poaceae, Chloridoideae) in the Pacific Northwest (British Columbia, Washington and Oregon), including the first report of Spartina xtownsendii for British Columbia, Canada, PhytoKeys 10, pp. 25-82 : 43-44

Spartina patens (Aiton) Muhl., Descr. Gram. 55. 1817.

Spartina juncea var. patens (Aiton) St.-Yves, Candollea 5: 86. 1932. Dactylis patens Aiton, Hort. Kew. 1: 104. 1789. Type. United Kingdom. Cultivated, native of North America, introduced 1781, by Mr. William Curtis (holotype: BM-001010621!).

Trachynotia juncea Michx., Fl. Bor.-Amer. 1: 64. 1803. Limnetis juncea (Michx.) Rich., Syn. Pl. 1: 72. 1805. Spartina juncea (Michx.) Willd., Enum. Pl. 81. 1809. Spartina patens var. juncea (Michx.) Hitchc., Rhodora 8: 210. 1906. Spartina juncea subvar. americana St.-Yves, Candollea 5: 84. 1932. Type. United States of America. bord des Creeks salés Basse Caroline, Michaux s.n. (holotype: P, not seen). Note: See Hitchcock (1908:153) for notes on the type specimen. Treatment of Trachynotia juncea as a synonym of Spartina patens follows Mobberley (1956).

Limnetis juncea var. monogyna M.A.Curtis, Boston J. Nat. Hist. 1: 136. 1837. Spartina patens var. monogyna (M.A.Curtis) Fernald, Rhodora 49: 114. 1947. Type. United States of America. North Carolina: M.A.Curtis s.n. (holotype: GH [GH00295532!], Appendix Fig. 3 ). Note: Fernald (1947:115) referred to this specimen as "The isotype of Curtis’s variety in the Gray Herbarium …” and "The Curtis type or isotype ….”; it likely is the holotype.

Spartina pumila Roth, Catal. Bot. 3: 10. 1806. Type. United States of America. New York, 1794, Schreber s.n. (holotype: B-Willd.!).

Description.

See Mobberley (1956), Barkworth (2003), Douglas et al. (2001), and Lonard et al. (2009). 2 n = 40 ( Marchant 1968b).

Common names.

Saltmeadow cordgrass, saltmeadow grass, marsh hay, wiregrass, foxgrass, couchgrass, rush saltgrass, spartine étalée ( Lonard et al. 2010).

Etymology.

The Latin epithet patens means "spreading, outspread" ( Stearn 1992), given in reference to the general habit of the species, which forms dense, monotypic stands.

Illustrations.

Hitchcock 1935: 494; Hitchcock 1951:514; Holmgren et al. 1998:750, Douglas et al. 2001:271, Barkworth 2003:251.

Distribution.

Native to the east coast of North America and Central America, distributed along the Atlantic coast of Newfoundland and Labrador to Texas, the Atlantic coast of Mexico, and throughout the Caribbean Islands (e.g., Barkworth 2003, Beetle 1977, Howard et al. 1989, Liogier and Martorell 2000, Peterson 2001). It has been introduced to coastal regions in Spain ( Campos et al. 2004, SanLeón et al. 1999, Prieto et al. 2011), China ( An et al. 2007), North Africa, and the west coast of the United States (Oregon, Washington), and Canada (British Columbia).

Comments.

Spartina patens grows in coastal salt marshes and brackish waters, where it usually forms dense stands above the intertidal zone and into higher and drier areas of the salt marsh ( Mobberley 1956). In Europe this taxon is sometimes treated as a separate species, Spartina versicolor Fabre, a traditional circumscription that some contemporary authors maintain (e.g., Sánchez Gullón 2001; Clayton et al. 2002 onwards; see SanLeón et al. 1999 and Prieto et al. 2011 for summaries of this taxonomic problem). In North America Spartina patens hybridizes with Spartina pectinata in areas where they grow together, forming the hybrid taxon Spartina xcespitosa . This hybrid is known from scattered locations along the Atlantic coast from Prince Edward Island to Maryland ( Mobberley 1956, Dore and Marchant 1968, Barkworth 2003), but has not been recorded along the Pacific coast. The biology of Spartina patens is reviewed by Lonard et al. (2010).

In Oregon, Spartina patens grows in the Siuslaw estuary on Cox Island (Lane Co.), where it has been introduced since at least 1939 and has expanded considerably since that time ( Frenkel and Boss 1988). It is not reported from elsewhere in Oregon ( Wilson et al. 2012). In Washington it has been reported from Dosewallips State Park on the west shore of Hood Canal ( Frenkel 1987). In California it is present in Southampton Marsh in San Francisco Bay ( Spicher and Josselyn 1985, Baird and Thieret 2012).

Spartina patens was the first of the invasive cordgrasses to be collected in British Columbia. It was discovered in 1979 in the Comox Estuary on Vancouver Island (Brayshaw 79-1143, V); nearly a decade later, in 1988, it was collected on the adjacent mainland coast in Burrard Inlet, North Vancouver (Lomer 88-140, UBC, Fig. 9 ). Spartina patens was recognized as part of the provincial flora by Douglas et al. (1994) based on collections from these two localities, and is reported for the region in recent floras ( Barkworth 2003, Douglas et al. 2001). Large stands of the species were reported in 2000 at the Comox locality in the Courtenay River estuary at the delta of Brooklyn Creek to the embayed area of Goose Spit ( Asp and Adams 2000). Spartina patens is also reported from Port Moody Arm of Burrard Inlet in British Columbia (Dresen et al. 2010). Voucher specimens from these latter two sites should be collected and deposited in herbaria.

Morphology.

Spartina patens exhibits considerable morphological variation and several authors have recognized two infraspecific taxa (see Mobberley 1956). Mobberley (1956) studied this variation across the native range of the taxon and found plants at the northern and southern extremes of the taxon’s range to be distinct, but considerable overlap and clinal variation from north to south in areas between the extremes. Based on these data he argued against the recognition of infraspecific taxa in Spartina patens , a circumscription followed by subsequent authors.

Based on specimens of Spartina patens examined here collected in Oregon, Washington, and British Columbia, introduced plants in these areas are similar morphologically to those in the eastern Canada and the northeastern United States, which tend to be smaller than plants distributed further south ( Mobberley 1956).

Spartina patens can be distinguished from Spartina densiflora by the following combination of characters: branches distinctly one-sided, appressed, ascending or spreading from main axis, distant or weakly overlapping, and approximately the same length within an inflorescence [vs. branches not distinctly one-sided, appressed, strongly overlapping, and conspicuously decreasing in length towards the inflorescence apex], rhizomes wiry, plants forming dense mats [vs. rhizomes absent, rarely short, plants cespitose, forming distinct clumps], upper glumes distinctly 3-veined [vs. upper glumes 1-veined], and ligules 0.5-1 mm long [vs. ligules 1-2 mm long].

Spartina patens can be distinguished from Spartina alterniflora , Spartina anglica and Spartina xtownsendii by: blade margins and adaxial surfaces scabrous [vs. blade margins and adaxial surfaces glabrous], blades 0.5-4 mm wide at base, involute for most or all of their length [vs. blades 3-10 mm wide at base, often involute distally], branches distinctly one-sided, distant or weakly overlapping [vs. branches not distinctly one sided, strongly overlapping], rhizomes thin and wiry [vs. rhizomes thick and fleshy], upper glumes conspicuously 3-veined [vs. upper glumes 1-3-veined, veins inconspicuous], and spikelets usually purple-tinged [vs. spikelets rarely or never purple-tinged].

Specimens examined.

Canada. British Columbia: Vancouver Island, Goose Spit, Comox, 49°40'N, 124°54'W, 14 Sep 1979, T.C.Brayshaw 79-1143 (V [V95308]); Vancouver Island, based of Comox Spit, 49°40'N, 124°54'W, 7 Aug 1984, T.C.Brayshaw 84-139 (V [V127117]); N shore of Burrard Inlet, E of Second Narrows Bridge, North Vancouver, 49.3099°N, 123.1032°W uncertainty 4052 m, sea level, Jul 1987, F.Lomer 87-001 (UBC [UBCV194265, Suppl. Fig. 29]); North Vancouver, Maplewood Mudflats, S of Dollarton Highway and E of Second Narrows bridge, 49.3042°N, 123.0009°W uncertainty 500 m, 27 Aug 1988, F. Lomer 88-140 (UBC [UBCV195726, Fig. 9 ); North Vancouver, Maplewood mud flats, 49.3042°N, 123.0009°W uncertainty 500 m, 16 Sep 1993, F.Lomer s.n. (V [V169504, V169505]). New Brunswick: St. Andrew’s, 16 Aug 1900, J.Fowler s.n. (CAN [CAN390994]). Charlotte Co.: Grand Manan, 44°41'53"N, 66°49'20"W, 31 Jul 1944, C.A.Weatherby & U.F.Weatherby 7300 (CAN [CAN33948]). Westmorland Co.: 1 mi E of Cape Bimet, 5 mi E of Shediac, 46°14'N, 64°27'W, 7 Aug 1981, M.Shchepanek & A.Dugal 3657 (CAN [CAN474807]); W of upper cape, 7 Aug 1964, P.R.Roberts & N.Bateman 64-2564 (CAN [CAN305984]); Memramcook, 45°58'15"N, 64°35'36"W, 21 Aug 1919, F.Rolland-Germaine 8022 (CAN [CAN332081]); Shediac, 46°13'17"N, 64°32'23"W, 5 Aug 1904, J.Fowler s.n. (CAN [CAN391711]); Moncton, 46°05'58"N, 64°47'59"W, 18 Sep 1912, M.O.Malte 108313 (CAN [CAN206830]). Restigouche Co.: Dalhousie, 48°2'55"N, 66°23'25"W, 4 Aug 1955, H.J.Scoggan 12683 (CAN [CAN240148]). Newfoundland and Labrador: Bonavista South District, Newman Sound Marsh, 48°32'15"N, 53°58'06"W, R.Charest, L.Brouillet, A.Bouchard & S.Hay 96-2065 (CAN [CAN58446]); St. George’s, 48°24'55"N, 58°29'40"W, 13 Aug 1910, M.L.Fernald & K.M.Wiegand 2597 (CAN [CAN33945]); St. George’s District, St. George’s, 48°24'55"N, 58°29'40"W, 4 Aug 1986, L.Brouillet & I.Saucier 86170 (CAN [CAN546790]; St. George’s District, Stephenville Crossing, saltmarsh NE of Main Gut bridge, 48°31'44"N, 58°27'19"W, 4 Aug 1986, L.Brouillet & I.Saucier 86183 (CAN [CAN546704]. Nova Scotia: N end of Summerville Beach, Summerville Center, 43°57'N, 64°49'W, 28 Sep 1979, D.F.Brunton & H.L.Dickson 2089 (CAN [CAN452656]; LeHave River, 6 Aug 1910, J.Macoun 82102 (CAN [CAN33949]. Cape Breton Co.: Grand Narrows, 45°57'24"N, 60°47'32"W, 27 Jul 1893, J.Macoun 21127, (CAN [CAN33952]); near mouth of George River, 27 Aug 1920, C.H.Bissell & D.H.Linder 19976 (CAN [CAN33951]. Digby Co.: Clare Municipality, Meteghan River, 44°13'N, 66°08'W, 30 Jul 1975, A.W.Dugal 75-131 (CAN [CAN475807]; Sissiboo River, Weymouth, 44°24'44"N, 65°59'43"W, 21 Aug 1920, M.L.Fernald, C.H.Bissell, C.B.Graves, B.Long & D.H.Linder 19974 (CAN [CAN33950]. Guysborough Co.: Canso, 45°20'12"N, 60°59'40"W, 15 Aug 1901, J.Fowler s.n. (CAN [CAN391709]. Hants Co.: mouth of Rennie Brook, East Walton, 17 Sep 1958, E.C.Smith, W.J.Curry, R.E.Clattenburg 18581 (CAN [CAN296579]). Kings Co.: Avonport, 45°06'01"N, 64°15'27"W, 23 July 1957, H.J.Scoggan 13850 (CAN [CAN255570]. Queens Co.: Port Mouton, 43°55'38"N, 64°50'55"W, C.H.Bissell & C.B.Graves 19978 (CAN [CAN33953]. Richmond Co.: Cape Breton Island, Fullers River Salt Marsh, 3 km W of Fourchu, off Hwy. 327, 45°43'N, 60°18'W, 23 Aug 1984, M.J.Shchepanek & A.W.Dugal 6426 (CAN [CAN521701]). Shelbourne Co.: Gunning Cove, 43°41'30"N, 65°20'45"W, 4 Oct 1982, S.J.Darbyshire 1790 (CAN [CAN487055]. Yarmouth Co.: Wedgeport, 43°44'23"N, 65°58'48"W 31 July 1953, W.L.Klawe 1278 (CAN [CAN298545]; Sand Beach, 43°48'43"N, 66°07'15"W, 7 Sep 1920, M.L.Fernald, B.Long, D.H.Linder 19977 (CAN [CAN33954]. Ontario: Essex Co.: Windsor, Windsor Salt Factory, 42°17'N, 83°06'W, 21 Sep 1975, P.M.Catling & S.M.McKay s.n. (CAN [CAN396523]); Windsor salt works, 24 Aug 1977, W.Botham 2011 (CAN [CAN459521], CAN [CAN459519]); Windsor, near salt factory, 29 Jul 1979, W.Botham 2182 (CAN [CAN459520]); Windsor, E side of Euclid Avenue bordering Detroit River, just S of Prospect Street, Ojibway Park, near salt plant, 42°17'N, 83°05'W, 3 Nov 1975, P.D.Pratt 18 (CAN [CAN440539]); Windsor, 50 m W of Prospect Avenue, along Euclid Road, E shore Detroit River, 42°17'N, 83°05'W, 4 Sep 1979, D.F.Brunton & P.D.Pratt 1915 (CAN [CAN452513]). Prince Edward Island: Prince Co.: Tignish, 46°57'02"N, 64°02'01"W, 6 Aug 1912, M.L.Fernald, B.Long & H.St-John 113172 (CAN [CAN33946]). Queens Co.: Long Creek salt marsh, ¼ mi E of Pond Point, 46°03'N, 62°57'W, 15 Aug 1981, M.Shchepanek & A.Dugal 4119 (CAN [CAN475274]); Brackley Point, 46°23'04"N, 63°11'06"W, 3 Aug 1888, J.Macoun 28967 (CAN [CAN33947]). Quebec: Maria Co.: Bonaventure, 48°03'N, 65°29'W, 11 Aug 1930, F.Marie-Victorin, F.Rolland-Germaine & E.Jacques 33799 (CAN [CAN33955]); Magdalen Islands, sandy sea strand at the Narrows, Alright Island, 21 Aug 1912, M.L.Fernald, B.Long & H.St-John 6880 (CAN [CAN33956]); Rivière-du-Loup Co.: Pointe-à-la-Loupe, L’Isle-Verte, 48°4'38"N, 69°16'28"W, 2 Sep 1957, E.Lepage 13956 (CAN [CAN252814]); Rivière-du-Loup, Hwy. 20, N shore of bay, 47°49'N, 69°32'W, 29 Sep 1979, H.L.Dickson & D.F.Brunton 3271 (CAN [CAN457594]). Kamouraska Co.: Rankin Point near Kamouraska, 24 Aug 1947, J.H.Soper & D.A.Fraser 3647 (CAN [CAN257674]); Rimouski Co.: Rimouski, 48°27'N, 68°32'W 30 Sep 1950, E.Lepage 13215 (CAN [CAN206869]); Le Bic, Cap aux Corbeaux, 48°23'N, 68°43'W, 30 Aug 1970, G.Lemieux 13630 (CAN [CAN444255]); L’Isle-Verte, 48 °01'N, 69°20'W, 24 Aug 1951, L.McI.Terrill s.n. (CAN [CAN337522]); Tobin, 23 Aug 1951, L.McI.Terrill 6636 (CAN [CAN337521]); Iles-de-la-Madeleine, Cap de l’Est, 47°36'47"N, 61°27'35"W, s.d., M.M.Grandtner s.n. (CAN [CAN519307]); Dune du Nord, près de la Grande Lagune, 47°29'N, 61°45'W, 66.08.06, M.M.Grandtner 10697-V (CAN [CAN519268]); Ile-aux-Coudres, La Baleine, pointe E de l’île, 25 Aug 1977, J.Cayouette J77-133 (CAN [CAN466657]). United State of America. Florida: near Jacksonville, 17 Jul 1894, A.H.Curtiss 4948 (CAN [CAN373389]). Louisiana: Cameron Parish, along the Gulf of Mexico, S of an unnamed shell road which runs E from Cameron Parish Road 3106, on the E edge of Cameron, T15S, R9W, 30 Jun 1984, B.E.Button & D.W.Pritchett 2536 (CAN [CAN495018]); Jefferson Parish, roadside at Elmer’s Island, 2 Oct 1976, J.Guider 5023 (CAN [CAN432238]). Maryland: sea coast, Sep 1863, Wm.M.Canby s.n. (CAN [CAN162200]). Massachusetts: Barnstable Co.: Sandy Meck, Cape Cod, 41°44'00"N, 70°19'58"W, 28 Oct 1939, J.H.Soper 1109 (CAN [CAN257821, CAN316382]). Dukes Co.: Katama Bay, Edgartown, Marthas Vineyard, 41°21'15"N, 70°28'58"W, 11 Sep 1901, M.L.Fernald s.n. (CAN [CAN162201]). New Jersey: Atlantic City, 1880, C.D.Fretz s.n. (CAN [CAN556282]). New York: Nassau Co.:Jones Beach, 40°35'40"N, 73°30'10"W, 18 Aug 1932, H.A.Gleason & A.C.Smith 149 (CAN [CAN162198]). Washington: Jefferson Co.: mouth of Dosewallips River, E of Route 101, SW of Sylopash Point, 1 m, T25N R2W S2, 47°41.4'N, 122°53.5'W, 9 Sep 2004, P.F.Zika & F.Weinmann 20160 (WTU [WTU359724, Suppl. Fig. 30]). Oregon: Lane Co.: Cox Island, Siuslaw Estuary, 2.5 km E of Florence, 43.9716°N, 124.0672°W, 9 Aug 1983, R.E.Frenkel 3060 (UBC [UBCV196070, Suppl. Fig. 31]); Lane Co.: center of Cox Island in Siuslaw River estuary, 3.5 km E of Florence, SW corner Sec 30, T. 18S, R. 11 W., W.M., 43.9716°N, 124.0672°W, 22 Oct 1977, R.E.Frenkel s.n. (WTU [WTU286900, Suppl. Fig. 32]).

Figures

Figure 9. Photograph of a specimen of Spartina patens collected at Maplewood mudflats, North Vancouver, British Columbia (Lomer 88 - 140, UBC). Image published with the permission of the University of British Columbia Herbarium, Beaty Biodiversity Museum.

Appendix figure 3. Holotype specimen of Limnetis juncea var. monogyna (Curtis s. n., GH [GH 00295532]). Photo: Gray Herbarium of Harvard University Herbarium.