Spartina alterniflora Loisel., Fl. Gall. ii. 719. 1807.

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 : 30-34

Spartina alterniflora Loisel., Fl. Gall. ii. 719. 1807.

Spartina glabra var. alterniflora (Loisel.) Merr., U.S.D.A. Bur. Pl. Industr. Bull. 9: 9. 1902. Spartina maritima var. alterniflora (Loisel.) St.-Yves. Candollea 5: 25, 53, pl. 2, f. 1-4, 1932. Spartina stricta var. alterniflora (Loisel.) A. Gray, Manual (ed. 2) 552. 1856. Trachynotia alterniflora (Loisel.) DC., Fl. Franc. (ed. 3) 5: 279. 1815. Type. France. Sur les bord de l’Odo[…?], à Bayonne, dans une prairie au bout des allées marines, 20 Jun 1803, n.c. (holotype: AV!, Appendix Fig. 1 ). Note: The location of the holotype in the herbarium (AV) at Museum Requien in Avignon, France, has not been reported previously in the literature.

Dactylis fasciculata Lam., Tabl. Encycl. 1: 180. 1791. Type. UNITED STATES OFAmerica. D.Richard s.n. (holotype: P [P00564318!] (hb. Lamarck); isotype: US [USA865637!] (fragm. ex P)).

Spartina glabra Muhl. ex Elliott, Sketch Bot. S. Carolina 1(1): 95, pl. 4, f. 2. 1816. Limnetis glabra (Muhl. ex Elliott) Eaton & Wright, Man. Bot. (ed. 8) 301. 1840. Spartina alterniflora var. glabra (Muhl. ex Elliott) Fernald, Rhodora 18: 178. 1916. Spartina maritima subsp. glabra (Muhl. ex Elliott) St.-Yves, Candollea 5: 48. 1932. Spartina maritima var. glabra (Muhl. ex Elliott) St.-Yves, Candollea 5: 24, 49, t. 1, f. b-2. 1932. Spartina stricta var. glabra (Muhl. ex Elliott) A. Gray, Manual ed. 2. 552. 1856. Type. United States of america.South Carolina or Georgia: S.Elliott s.n. (holotype: ChM [ChMBY4006!] (hb. Elliott), Appendix Fig. 2 ). Note: There is a specimen collected by S. Elliott, likely in 1810 or 1811, in the Elliott herbarium at the Charleston Museum (Charleston, South Carolina). Chauncey Beadle and Charles Boynton conducted a systematic inventory of the Elliott herbarium in the first decade of the 20th century; they determined this specimen to be Elliott’s " Spartina glabra of Sketch" and annotated it as such on the sheet (A.E. Sanders, personal communication 2011). This specimen is the holotype of Spartina glabra . Material seen by Muhlenberg is housed at PA (see Hitchcock 1932: 30).

Spartina brasiliensis Raddi, Agrostogr. Bras. 21-22. 1823. Spartina maritima var. brasiliensis (Raddi) St.-Yves, Candollea 5: 56, t. 2, f. 56. 1932. Type. Brazil. Rio de Janeiro, G.Raddi s.n. (holotype: PI; isotypes: FI (2 sheets), PI, US [US3376186!] (fragm. ex PI)).

Spartina glabra var. pilosa Merr., U.S.D.A. Bur. Pl. Industr. Bull. 9: 9. 1902. Spartina alterniflora var. pilosa (Merr.) Fernald, Rhodora 18: 179. 1916. Spartina maritima subvar. pilosa (Merr.) St.-Yves, Candollea 5: 51, t. 1. f. c-3. 1932. Type. UNITED STATES OF AMERICA. New Jersey: Aug 1895, F.L.Scribner s.n. (holotype: US [US81736!]). Note: Peterson (2001) identified the US specimen as an isotype, but I interpret it to be the holotype, as Merrill (1902) indicated his work was based entirely on specimens in US, and wrote type after citing this specimen in the protologue.

Spartina xmerrilli A.Chev. Bull. Soc. Bot. France 80: 787. 1933. Type. UNITED STATES OF AMERICA. New York: Long Island, Rockaway Point, 22 Oct 1908, E.P.Bicknell 11300 (holotype: NY [NY431515!]; isotype: PH [PH736377!] Note: Chevalier (1933: 788) lists nine specimens in the protologue, and indicates in a figure that Bicknell 11300 is the type; the NY specimen of this collection has ‘type’ handwritten on it.

Spartina maritima subvar. fallax St.-Yves, Candollea 5: 57. 1932. Type. BRAZIL. Bahia: Salzmann s.n. [sheet A] (lectotype: P [P02249890!], here designated). Note: St-Yves (1932) listed twelve specimens (syntypes) in the protologue, including the one designated here as lectotype.

Description.

Culms to 250 cm tall, rhizomatous. Sheaths glabrous; ligules 1-2 mm long; blades 5-63 cm long × 3-10 mm wide at base, usually flat proximally, involute distally, divergent from stems 15-18°, adaxial and abaxial surfaces glabrous, margins smooth, rarely with occasional scabrous teeth. Inflorescences (6-)11-33 cm long × (5-) 6-10(-15) mm wide at midpoint, erect, with 3-9(-12) branches; branches (4)5-15 cm long × 2-4 mm wide, appressed to main axis or ascending, rachises 0.4-1 mm wide between spikelets, extending 1-20 mm beyond terminal spikelet. Spikelets 8-14(-16.5) mm long × 1-2 mm wide, alternate, weakly appressed, weakly or moderately overlapping, calluses 0.5-1.5 mm long. Glumes glabrous or weakly pubescent, when present hairs to 0.2 mm long, proximal hairs sometimes denser and longer to 0.5 mm, keels glabrous or ciliate, when present hairs to 0.3 mm long, margins glabrous; lower glumes 4-9 mm long × 0.2-0.5 mm wide, 1-veined, tips acute; upper glumes 7-14 mm long × 1-1.2 mm wide, 5-7-veined, tips acuminate or obtuse. Lemmas 7-12 mm long, glabrous or scabrous; paleas exceeding lemmas by up to 1 mm; anthers 3-6 mm long, yellow, exserted at maturity, dehiscent, pollen fertile. 2 n = 62 ( Marchant 1963, 1968b).

Common name.

Smooth cordgrass, Atlantic cordgrass, Atlantic smooth cordgrass.

Etymology.

The epithet alterniflora means alternating flowers.

Illustrations.

Hitchcock 1935:492, Hitchcock 1951:512, Hitchcock et al. 1969:706, Holmgren et al. 1998:749, Barkworth 2003:245, Cope and Gray 2009:543, Scholz et al. 2009:172.

Distribution.

Native to the Atlantic and Gulf coasts of North America from Newfoundland and Labrador, Canada, to Texas, U.S.A. ( Mobberley 1956, Barkworth 2003), and Tamaulipas and Veracruz, Mexico ( Mora-Olivo and Valdés-Reyna 2011). Introduced in England, France, and Spain ( Campos et al. 2004), the west coast of the United States (Washington, Oregon, California), New Zealand ( Partridge 1987), and China ( Bixing and Phillips 2006, An et al. 2007, Scholz et al. 2009).

Comments.

Spartina alterniflora is often misspelled in the literature as "Spartina alternifolia". Spartina alterniflora was described from Bayonne in southwestern France ( Loiseleur-Deslongchamps 1807), where it has been introduced for over two centuries. The taxon was collected in France as early as 1803, and was known from southern England (Southampton Water) by 1829; it is presumed to have been introduced from North America to these European coastal regions in shipping ballast (see Bromfield 1836, Marchant 1968). Spartina alterniflora is one of the parents of the sterile F1 hybrid Spartina xtownsendii (see notes under this taxon), which was formed when this introduced species hybridized with the European Atlantic coast native species Spartina maritima . By 1969 Spartina alterniflora was extremely rare in Southampton Water, known from only a single locality ( Marchant and Goodman 1969a, 1969b), and it remains rare in Britain ( Cope and Gray 2009).

Several introductions of Spartina alterniflora have occurred along the west coast of North America where it is not native. The species was reported in 1945 from a single estuarial location in Willapa Bay, Washington, where occasional plants had been observed since around 1911, and thought to have been introduced in the early twentieth century with oyster culture ( Scheffer 1945). The earliest known record documenting Spartina alterniflora at Willapa Bay is a photograph taken in the early 1940s by T. Scheffer, housed at the California Academy of Sciences Herbarium ( Civille et al. 2005). Based on a review of historical records of the oyster trade, Civille et al. (2005) concluded that Spartina alterniflora was probably introduced to Willapa Bay between 1893 and the late 1910s via seeds attached to oysters during their railway transport from the Atlantic coast to the Pacific coast. Since the mid-twentieth century Spartina alterniflora has expanded dramatically in Willapa Bay and northwards to Grays Harbour and the Copalis River estuary in Washington ( Stiller and Denton 1995), radically transforming these tidal ecosystems from mudflats to dense cordgrass swards (e.g., Feist and Simenstad 2000, Davis et al. 2004a, 2004b). Civille et al. (2005) used historical records and remote sensing to reconstruct the dramatic expansion of Spartina alterniflora that occurred in Willapa Bay between 1945 and 2000, and concluded that its rapid colonization likely originated from multiple introductions over several decades, as opposed to the long-standing hypothesis of post-establishment colonization following a single introduction to the region (e.g., Scheffer 1945, Stiller and Denton 1995, Feist and Simenstad 2000). Considerable research has been conducted on understanding the invasion dynamics of Spartina alterniflora in Willapa Bay (e.g., Davis et al. 2004a, 2004b, Taylor and Hastings 2004, Grosholz et al. 2009), and substantial efforts and resources have been directed at its management, control and eradication (e.g., Grevstad et al. 2003, Taylor and Hastings 2004). Spartina alterniflora is also known from the southern end of Padilla Bay in northwestern Washington, where it was apparently introduced independently between 1941 and 1945 to aid in beach stabilization (see Wiggins and Binney 1987, Riggs 1992). Stiller and Denton (1995) confirmed the genetic distinctiveness of this stand compared with the Willapa Bay population, supporting its putative independent origin.

In Oregon, Spartina alterniflora has been reported from the Siuslaw River estuary and Coos Bay ( Howard 2005; Wilson et al. 2012). In California Spartina alterniflora occurs in southeastern San Francisco Bay (Alameda Creek Flood Control Channel and along the shoreline some three km south) where it was planted for restoration in the late 1970s ( Spicher and Josselyn 1985, Ayres et al. 2003). By some 20 years later it had spread considerably, and was known from seven different areas in San Francisco Bay ( Callaway and Josselyn 1992). Spartina alterniflora has also been reported from Bolina’s Lagoon and Point Reye’s National Seashore north of San Francisco Bay; these occurrences are thought to have originated from floating seed originating in San Francisco Bay, but this hypothesis has not been confirmed ( Ayres et al. 2003).

The only native Spartina taxon in California is Spartina foliosa , and by the 1990s it was known that Spartina alterniflora was in the process of competitively excluding Spartina foliosa ( Callaway and Josselyn 1992). Substantial evidence has accumulated indicating that the two taxa have successfully interbred producing a highly invasive fertile hybrid capable of backcrossing with its parental taxa, resulting in a genetically heterogeneous hybrid cordgrass swarm (i.e., Spartina alterniflor a × Spartina foliosa ) that is much more invasive than either parental taxon ( Daehler and Strong 1997, Ayres et al. 1999, 2003, 2004, Anttila et al. 2000). Morphological variation of these hybrids is not well characterized in the literature.

Spartina alterniflora is not known from British Columbia, Canada. Daehler and Strong (1996) identified estuaries along the Pacific coast extending to the Canada / United States border, which they predicted to be potentially vulnerable to invasion by Spartina species, including Spartina alterniflora . Workers should be aware of the potential for its invasion and keep watch for the species in British Columbia. In a recent study of saltmarsh diatoms in central mainland British Columbia some 40 km northeast of Vancouver Island, Roe et al. (2009) reported Spartina alterniflora as a dominant species in low and mid marsh components of Waump (51°11'15N, 126°55'15W) and Wawwat’l saltmarshes (51°11'36N, 126°40'5W) in Seymour Inlet, based on fieldwork conducted in 2002. If the species is correctly identified as Spartina alterniflora in this study, this would be the first known site of the taxon in British Columbia. Alternatively, the taxon may be Spartina anglica , but it is not possible to further confirm its identification based on the study (voucher specimens are not mentioned). Spartina has not otherwise been reported from Seymour Inlet. This region should be re-visited for further field study and collection of voucher specimens. Whatever the identity of the species is, this report suggests that by 2002 invasive Spartina was present in coastal British Columbia at sites further north than those first documented in 2003 near the international border south of Vancouver.

Morphology.

Spartina alterniflora and the European species Spartina maritima are the parents of the sterile F1 hybrid Spartina xtownsendii ; unsurprisingly, Spartina alterniflora is morphologically similar to Spartina xtownsendii and the amphidiploid Spartina anglica . It can be distinguished from these taxa by its shorter spikelets [8-14(-16.5) mm vs. 14-25 mm], narrower branch rachises [0.4-1 mm wide between spikelets vs. 1-2.2 mm wide], glumes glabrous or weakly pubescent [vs. glumes weakly to densely pubescent], and leaf blades erect, forming an angle of 15-18° with the culm [vs. leaf blades spreading, forming an angle of 30-60° with the culm]. Spikelets of Spartina alterniflora are shown in Fig. 1 , and an exemplar specimen is shown in Fig. 2 . Glumes in Spartina alterniflora vary from glabrous to pubescent (details on this variation are given in Mobberley 1956). In individuals with glabrous glumes, this is a good character for distinguishing Spartina alterniflora from Spartina xtownsendii and Spartina anglica , which consistently have pubescent glumes, but the character is more difficult to interpret and less reliable in Spartina alterniflora individuals with pubescent glumes. Spartina alterniflora also tends to have narrower inflorescences than Spartina xtownsendii and Spartina anglica [(5-)6-10(-15) mm wide vs. 7-25 mm wide], but there is considerable overlap in this character. Recent taxonomic keys ( Barkworth 2003, Kozloff 2005) distinguishing these three taxa emphasize variation in number of panicle branches (3-25 in Spartina alterniflora vs. 1-12 in the other taxa). I found this character unreliable in material examined here, and do not include it in the key. Spartina alterniflora can further be distinguished from Spartina anglica by its shorter anthers [3-6 mm long vs. 7-10 mm long], and from Spartina xtownsendii by its fully exserted, dehiscent anthers at anthesis and fertile pollen [vs. anthers not or incompletely exserted, indehiscent and sterile pollen]. Spartina alterniflora is readily distinguished from Spartina densiflora , Spartina gracilis , Spartina patens and Spartina pectinata by it glabrous leaf blade margins [vs. scabrous leaf blade margins].

There is considerable morphological variation in Spartina alterniflora throughout its native range with northern plants from Canada and Maine tending to have looser inflorescences, weakly overlapping spikelets, and less glume pubescence, and southern plants tighter inflorescences, more strongly overlapping spikelets, and more pubescent glumes. This variation has been recognized taxonomically in the past at the species and infraspecific levels; however, Mobberley (1956) demonstrated the variation to be clinal and therefore taxonomically inconsistent, and only a single taxon is now generally accepted (e.g., Barkworth 2003). Specimens examined from the Pacific coast in Washington have moderately overlapping spikelets.

Specimens examined.

CANADA. New Brunswick: Charlotte Co.:E of Biological Station, St. Andrews, 45°04'N, 67°03'W, Aug 1929, M.O.Malte 798/29 (CAN [CAN33914]); Grand Manan, Ross Island, 44°42'N, 66°48'W, 10 Aug 1927, C.A.Weatherby & U.F.Weatherby 5784 (CAN [CAN33957]); Grand Manan, Thoroughfare, 44°42'N, 66°48'W, 14 Aug 1944, C.A.Weatherby & U.F.Weatherby 7314 (CAN [CAN33913]). Gloucester Co.: Bathurst and vicinity, 47°37'N, 65°39'W, 2 Aug 1926, M.O.Malte 732 (CAN [CAN124678]). Restigouche Co.: Dalhousie, 48°04'N, 66°22'W, 4 Aug 1955, H.J.Scoggan 12682 (CAN [CAN240149]). Saint John Co.: St. John, 45°16'N, 66°04'W, 17 Aug 1877, J.Macoun 28971 (CAN [CAN33915]); Saints’ Rest Beach, on W side of St. John, St. John Harbour, NE of Lorneville, ca. 45°15'N, 66°02'W, 24 Aug 1975, P.M.Catling & S.M.McKay s.n. (CAN [CAN396499]). Westmoreland Co.: Moncton, 46°08'N, 64°46'W, 18 Sep 1912, M.O.Malte 108312 (CAN [CAN206832]); 1 mi E of Cape Bimet, 5 mi E of Shediac, 46°14'N, 64°27'W, 7 Aug 1981, M.Shchepanek & A.Dugal 3615 (CAN [CAN474764]). Newfoundland and Labrador: Chapel Island, Rocky Point, 3 mi SE of Summerford, 49°28'N, 54°45'W, 13 Aug 1977, M.J.Shchepanek & D.White 3051 (CAN [CAN436102]). St. Georges District, Stephenville Crossing, 48°31'N; 58°25'W, 4 Aug 1986, L.Brouillet & I.Saucier 86178 (CAN [CAN546706]); St. Georges, 48°31'11"N, 58°54'51"W, M.L.Fernald, K.M.Wiegand & J.Kittredge 2598 (CAN [CAN33900]); St. Georges, 48°31'11"N, 58°54'51"W, 4 Aug 1986, L.Brouillet & I.Saucier 86173 (CAN [CAN546843]). Port-au-Port District,West Bay Center, 4 Aug 1986, I.Saucier & A.Leduc 86185 (CAN [CAN546845]). Nova Scotia: Annapolis Co.:Granville, 44°47'02"N, 65°26'59"W, 18 Jul 1921, M.L.Fernald & N.C.Fassett 23294 (CAN [CAN33905]). Cape Breton Co.: Louisburg, Cape Breton, 18 Aug 1898, J.Macoun 21129 (CAN [CAN33902]). Digby Co.: Meteghan River, Clare Municipality, 44°13'N, 66°08'W, 27 Jul 1975, A.W.Dugal 75-66 (CAN [CAN475739]); 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 19972 (CAN [CAN33903]); Digby, 44°37'20"N, 65°45'38"W, 27 Aug 1910, J.Macoun 82104 (CAN [CAN33901]). Guysborough Co.:Canso, 45°20'12"N, 60°59'40"W, 15 Aug 1901, J.Fowler s.n. (CAN [CAN390991]); Canso, 45°20'12"N, 60°59'40"W, 1416 Aug 1930, J.Rousseau 35509 (CAN [CAN33907]); Guysborough, 45°23'N, 61°29'57"W, 6-7 Aug 1930, J.Rousseau 35357 (CAN [CAN33908]). Kings Co.: Avonport, 45°06'01"N, 64°15'27"W, 23 Jul 1957, H.J.Scoggan 13849 (CAN [CAN255571]). Lunenburg Co.: LaHave River, 44°17'37"N, 64°21'27"W, 6 Aug 1910, J.Macoun 82103 (CAN [CAN33904]). Pictou Co.: Pictou, 45°40'33"N, 62°42'33"W, 12 Aug 1880, McKay 28969 (CAN [CAN33906]); same locality, 31 Jul 1880, McKay 28972 (CAN [CAN33910]). Richmond Co.: Cape Breton Island, Richmond Municipality, Fullers River Salt Marsh, 3 km W of Fourchu, off Hwy. 327, 45°43'N, 60°18'W, M.J.Shchepanek & A.W.Dugal 6419 (CAN [CAN521694]). Queens Co.: N, end of Summerville Beach, Summerville Center, 43°57'N, 64°49'W, 28 Sep 1979, D.F.Brunton & H.L.Dickson 2092 (CAN [CAN452659]). Yarmouth Co.: Port Maitland, 43°59'03"N, 66°09'03"W, 24 Aug 1913, M.O.Malte s.n. (CAN [CAN206823]); Wedgeport, 43°42'58"N, 65°58'45"W, 18 Jul 1953, W.L.Klawe 1204 (CAN [CAN298544]); Lower Argyle, 43°43'44"N, 65°50'01"W, 11 Aug 1920, M.L.Fernald, C.H.Bissell, C.B.Graves, B.Long, D.H.Linder 19971 (CAN [CAN33909]). Prince Edward Island: Brackley Point, 46°23'N, 63°11'W, 4 Aug 1888, J.Macoun 28968 (CAN); Prince Co.: Tignish, 46°57'N, 64°02'W, 6 Aug 1912, M.L.Fernald, B.Long & H.St.John 6877 (CAN). Queens Co.: ¼ mi E of Pond Point, Long Creek salt marsh, 56°03'N, 62°57'W, M.Shchepanek & A.Dugal 4128 (CAN). Quebec: Bas-Saint-Laurent Region, Rocher blanc, 48°25'19"N, 68°36'24"W, 19 Jul 1949, Fr.Claude s.n. (CAN [CAN388720]); Cap a la Carre, St. Fabien, Gaspé, 48°17'N, 68°52'W, 9 Aug 1970, J.K.Morton NA3907 (CAN [CAN359898]); Cacouna Harbour development ca. 8 km NE of Rivière-du-Loup, 47°55'N, 69°30'W, 12 Aug 1980, D.F.Brunton 2519 (CAN [CAN455714]); Notre-Dame-du-Portage, 47°46'N, 69°37'W, 30 Aug 1970, G.Lemieux 13633 (CAN [CAN444192]); Rankin Point near Kamouraska, 24 Aug 1947, J.H.Soper & D.A.Fraser 3650 (CAN [CAN257673]). Capitale-Nationale Region, Baie-St.-Paul, 47°25'N, 71°20'W, 29 Jul 1984, S.G.Aiken 2962 (CAN [CAN484859]); Murray Bay, 47°38'60"N, 70°07'60"W, 14 Aug 1905, J.Macoun 68994 (CAN [CAN33921]). Côte-Nord Region, Anticosti Island, Ellis Bay, 49°30'N, 63°00'W, 7 Sep 1883, J.Macoun 28970 (CAN [CAN33920]); Betchwan [sic] [Betchouane], 50°14'27"N, 63°11'01"W, 25 Aug 1928, H.F.Lewis 130615 (CAN [CAN33919]); Havre des Canadiens, Natashquan, 50°10'59"N, 61°49'W, 7 Sep 1915, H.St.John 90141 (CAN [CAN33922]). Gaspésie-Îles-de-la-Madeleine Region, Parc de Forillon, Penouille, 48°51'N, 64°26'W, 17 Jul 1971, M.M.Grandtner G158 (CAN); Magdalen Islands, Coffin Island, East Cape, 47°33'N, 61°31'W, 17 Aug 1912, M.L.Fernald, B.Long & H.St.John 6878 (CAN [CAN33918]); Coin du Banc, 48°33'28"N, 64°17'34"W, 31 Jul 1939, H.J.Scoggan 646 (CAN [CAN33916]); House Harbour, Alright Island, 47°25'59"N, 61°46'W, 17 Aug 1917, F.Johansen s.n. (CAN [CAN33917]); Carleton, 48°05'N, 66°08'W, 11 Aug 1930, F.Marie-Victorin, F.Rolland-Germain & E.Jacques 33291 (CAN [CAN513103]); Gaspé Peninsula, Bonaventure River, 48°03'N, 65°29'W, 12 Aug 1940, H.J.Scoggan 1214 (CAN [CAN220299]); près du pont de la Grosse Ile, Iles-de-la-Madeleine, 47°37'N, 61°33'W, 0.3 m, 7 Aug 1966, M.M.Grandtner 10801-V (CAN [CAN519301]). United States of America. Connecticut: New London Co.: Norwich, 41°31'N, 72°04'W, G.R.Lumsden s.n., 11 Aug 1885 (CAN [CAN162186]). Massachusetts: Barnstable Co.: Provincetown, Provincetown Harbor, 42°02'N, 70°10'W, 9 Oct 1988, S.G.Aiken & S.R.Johnstone 88-488 (CAN). Essex Co.: salt marsh between Briar Neck and Bass Rocks, Gloucester, 42°36'N, 70°39'W, 26 Aug 1945, L.B.Smith 1318 (CAN [CAN162185]); same locality, 42°36'N, 70°39'W, 19 Aug 1945, L.B.Smith 1317 (CAN [CAN162182]); Salem, 42°31'N, 70°53'W, 188-, J.Sears 47174 (CAN [CAN162187]). Maryland: seashore, Sep 1863, Wm.M.Canby s.n. (CAN [CAN162184, CAN162183]). Washington: Skagit Co.: Dike Island in Padilla Bay, 7 mi W of Mt. Vernon, 19 Nov 1964, R.G.Jeffrey 64-1 (US [US2580778]). Pacific Co.: Willapa Refuge, tide water marsh, 28 Sep 1945, M.L.Hinshaw s.n. (US [US1867453]); Willapa Bay, W and N Long Island, some five small patches, scattered, apparently spreading, 10 Aug 1942, W.G.McFarland s.n. (US [US2436000]); T1N R10W, Section 22, mouth of Naselle River at SE end of Hwy. 101 bridge, 46.4291°N, 123.9078°W, 16 Sep 2000, C.L.Maxwell 1575 (WTU [WTU342828], Suppl. Fig. 1); The Nature Conservancy’s Ellsworth Creek Preserve, mouth of Ellsworth Creek on S bank of Naselle River mouth, entrance to Yellow Gate Road, T11N R10W S22, 46°25.6'N, 123°53.675'W, 0 m, 4 Sep 2003, P.F.Zika, D.Giblin, S.Rodman et al. 18936 (WTU [WTU357790], Fig. 2 ); Willapa Bay, sea level, 15 Sep 1994, W.Lebovitz s.n. (WTU [WTU344373], Suppl. Fig. 2); below Bruceport County Park campground, T14N R10W S22, 46°41.4'N, 123°53.1'W, 3 Sep 2003, P.F.Zika 18935 (WTU [WTU371783], Suppl. Fig. 3). England. Hampshire Co.: Southampton, 50°53'49"N, 01°24'15"W, s.d., Dalington s.n. (CAN [CAN134028]). Isle of Wight Co.:Isle of Wight, ca. 50°40'51"N, 01°16'51"W, 9 Oct 1871, F.Stratton s.n. (CAN). France. Bayonne, 43°29'N, 01°28'W, s.d., collector illegible (CAN [CAN134029]); same locality, 43°29'N, 01°28'W, Sep 1899, E.Mouillefarine s.n. (CAN [CAN560721]); Biarritz, 43°29'N, 01°33'W, Oliver s.n. (CAN [CAN421008]); Landes, near Capbreton, 43°38'N, 01°25'E, 24 Jun 1954, A.E.Porsild 18851 (CAN [CAN244668]).

Figures

Figure 1. Spikelets of Spartina alterniflora (U. S. A.: Washington, Pacific Co., Zika 18935, WTU). Bar = 3 mm. Photo: J. M. Saarela.

Figure 2. Photograph of a specimen of Spartina alterniflora collected at Ellsworth Creek Preserve, Pacific County, Washington, where the species is introduced (Zika et al. 18936, WTU). Image published with the permission of University of Washington Herbarium, Burke Museum.

Appendix figure 1. Holotype specimen of Spartina alterniflora (without collector, AV). Photo: Museum Requien. Specimen image reproduced with the permission of Museum Requien.

Appendix figure 2. Holotype specimen of Spartina glabra (Elliott s. n., ChMBY 4006). Photo: Sean Money, Charleston Museum.