Spartina ×townsendii
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Ordo: Poales
Familia: Poaceae
Genus: Spartina
Name
Spartina ×townsendii H. Groves & J. Groves, Bot. Exch. Club Rep. 1880. 37. 1881. – Wikispecies link – Pensoft Profile
- Spartina ×townsendii H. Groves & J. Groves, Bot. Exch. Club Rep. 1880. 37. 1881. Type. England. Mud flats, near Hythe, South Hants, 1 Sep 1879, H.Groves s.n. (holotype: BM [BM001003965!]; isotypes: C, K [K000710272!], W [W19160030795!, W19160030798!] (ex hb. Groves), US [US1127161!] (fragm. ex W ex hb. Groves), US [US878793!]). Note: The location of the holotype has not been reported previously in the literature.
- Spartina ×neyrautii Fouc., Ann. Soc. Sci. Nat. Charente-Maritime 31: 8. 1894. Type. France. près de Hendaye (Basses-Pyrénnées), E-J.Neyraut s.n., 24 Jul 1892 (isotypes: P [P00753804!, P03457326!, P03457334!, P03457416!], US! (fragm. ex P)]).
Description
Culms 46–100 cm tall, thick, fleshy, rhizomatous. Sheaths glabrous; ligules 1–1.5 mm long; blades 6.5–37 cm long × 4–10 mm wide, flat proximally, often involute distally, divergent 30–40° from culms, adaxial surfaces glabrous, occasionally with very sparse hairs proximally, when present hairs to 0.2 mm long, abaxial surfaces glabrous, occasionally with sparse hairs proximally, when present hairs to 0.5 mm long, margins smooth. Inflorescences 10.5–24(–36) cm long × 7–25 mm wide at midpoint, erect, with (2)3–6(–10) branches; branches (6–)7.5–15(–18) cm long × (2.5–)3–4 mm wide, appressed or ascending, rachises 1–1.9 mm wide between spikelets, extending 2–10(–18) mm beyond the terminal spikelet, extension occasionally absent, glabrous, margins glabrous, occasionally with a few marginal hairs, when present hairs to 0.2 mm long. Spikelets 14–17.5 mm long × 1.5–2.5 mm wide, weakly appressed, weakly overlapping, calluses 0.6–1.5(–2) mm long. Glumes weakly to moderately pubescent, hairs 0.1–0.2 mm long, proximal hairs occasionally to 0.6 mm long, keels glabrous, ciliate or scabrous, when present hairs and teeth 0.2–0.5 mm long, usually longest proximally; lower glumes 7–13 mm long × 0.5–0.7 mm wide, 1-veined, tips acuminate or obtuse; upper glumes 12.5–16.5 mm long × 1–1.5 mm wide, 3-veined, tips acuminate or obtuse. Lemmas 9.5–13.5 mm long, 1-3–veined, pubescent distally, glabrous proximally, margins membranous, keels ciliate distally, hairs to 0.2 mm long, glabrous proximally. Paleas exceeding lemmas by ca. 1 mm, glabrous. Anthers 5–7(–8.5) mm long, not or incompletely exserted at maturity, indehiscent, medium to dark brown, pollen sterile; caryopses absent. 2n = 62 (Marchant 1963[1], 1968b).
Common name
Townsend’s cordgrass.
Etymology
The epithet townsendii was given in honour of the English botanist Frederick Townsend (1822–1905).
Illustration
Cope and Gray 2009[2]:547.
Distribution
This species is found in England, Wales, Scotland, Ireland (Cope and Gray 2009[2]), Italy (Scarton et al. 2003[3]), United States (Washington), New Zealand (Partridge 1987[4]), and Canada (British Columbia).
Comments
In the mid to late 1800s an unknown cordgrass of restricted distribution appeared and spread rapidly along the shores of Southampton Water, England (Stapf 1914[5]), which differed morphologically (particularly by its sterile pollen) from Spartina maritima, the single cordgrass species native to the Atlantic coast of Europe and north Africa (Marchant and Goodman 1969c[6]), and the introduced Spartina alterniflora, which had been present in the region since the early part of the 19th century. The brothers Henry and James Grove (1881) described this taxon as Spartina townsendii from plants collected near Hythe. In the 1890s a second form of Spartina townsendii, which was recognized and considered distinct by having fertile stamens, was collected at multiple localities in the region, and by the mid twentieth century it had expanded substantially on tidal flats across the British Isles (see Goodman et al. 1959[7], Hubbard 1957[8], 1965[9]). For decades these two forms (one sterile, the other fertile) of Spartina townsendii were referred to collectively as the Spartina townsendii aggregate or Spartina townsendii sensu lato. Because of its vigorous growth and ability to rapidly colonize and stabilize mud flats, Spartina townsendii s.l. was considered to be a “useful” species and was distributed and planted widely for land reclamation, coastal protection, and animal feed across the British Isles, Europe, and in New Zealand (e.g., Oliver 1925[10], Harboard 1949[11], Goodman et al. 1959[7], Ranwell 1967[12]).
Soon after its formal description Spartina townsendii was considered to be a species of hybrid origin. Foucaud (1894)[13] suggested that Spartina townsendii was probably a hybrid of the native Spartina maritima and the introduced Spartina alterniflora, a hypothesis later supported by Stapf (1914)[5] and Huskins (1930), who examined cytological evidence and hypothesized that fertile plants of Spartina townsendii s.l. originated from chromosome doubling following hybridization between its parent species. Marchant (1963)[1] confirmed this work, and reported chromosome numbers as 2n = 62 for Spartina alterniflora, 2n = 60 for Spartina maritima, 2n = 62 for sterile plants of Spartina townsendii, and 2n = 120, 122, 124 for fertile plants of Spartina townsendii. The hybrid and chromosome doubling origins of the forms of Spartina townsendii have been confirmed by multiple lines of molecular evidence (Guénégou et al. 1988[14], Raybould et al. 1991[15], Ferris et al. 1997[16], Ayres and Strong 2001[17], Baumel et al. 2003[18]). Hubbard (1957)[8] observed the type specimen of Spartina townsendii to be the sterile F1 hybrid, and the taxon was subsequently referred to as Spartina ×townsendii. The fertile plants remained without a name until Hubbard (1978)[19] later described them as Spartina anglica, and the two forms have since been recognized as distinct taxa. Molecular data have identified Spartina alterniflora as the female parent and Spartina maritima as the male parent in the origin of Spartina ×townsendii (e.g., Ferris et al. 1997[16]).
An independent origin of Spartina ×townsendii is documented in France. In 1894 Jules Foucaud described Spartina neyrautii Fouc. from southwestern France and northern Spain. Spartina neyrautii was initially considered to be a variant of Spartina maritima (e.g., Chevalier 1923, Saint-Yves 1932[20]), but was later recognized as a morphologically and cytologically distinct hybrid, Spartina ×neyrautii,with the same parentage as Spartina ×townsendii (e.g., Jovet 1941[21], Chevalier 1933[22], Marchant 1977[23]). Baumel et al. (2003)[18] confirmed this with molecular data, demonstrating that Spartina ×neyrautii and Spartina ×townsendii originated independently by hybridization between the same maternal (Spartina alterniflora) and paternal (Spartina maritima) taxa. Because both taxa apply to the hybrid Spartina alterniflora × S. maritima, the later name Spartina ×neyrautii is a synonym of Spartina ×townsendii, in accordance with article H.2 of the Vienna Code (McNeill et al. 2006[24]), and as noted earlier by Raybould et al. (1990)[25]. In recent decades it has been documented that these hybrid plants are highly restricted in distribution in France (Marchant 1977[23], Hubbard et al. 1978[26], Raybould et al. 1990[25], Baumel et al. 2003[18]). Minor morphological differences between Spartina ×neyrautii and Spartina ×townsendii were noted by Marchant (1977)[23]. Measurements of spikelet characters in Spartina ×neyrautii type material examined at US fall within the range of variation reported here for Spartina ×townsendii.
Spartina ×townsendii has been introduced into North America, where apparently only a single occurrence has been reported in the literature. Hitchcock et al. (1969)[27] noted a single known population of Spartina ×townsendii in Washington at Stanwood, Snohomish Co. At the time of that publication, the fertile (=Spartina anglica) and non-fertile forms of Spartina ×townsendii were not distinguished taxonomically, and it is not explicitly clear from the flora which form of the taxon was known from the site. A specimen collected in 1965 from this population [Austenson s.n. (WTU, Suppl. Fig. 35)] is here confirmed to be the F1 sterile hybrid Spartina ×townsendii. The determination of a more recent collection from the Stanwood area identified as Spartina ×townsendii requires confirmation (Snohomish Co., Davis Slough west of Stanwood, 25 Aug 1990, M.Arnot 254, WTU-317391, not seen). A 2005 collection from Washington originally identified as Spartina ×townsendii (Giblin & Legler 270 WTU) is here re-determined to be Spartina anglica. Barkworth (2003)[28] included Spartina ×townsendii in her treatment of Spartina for North America, but did not include a distribution map or otherwise indicate a range for the species, suggesting some confusion in the literature on its status in North America. Kozloff (2005)[29] also included Spartina ×townsendii (as well as Spartina anglica) in his Pacific Northwest flora, indicating only ‘coastal salt marshes’ for its distribution. There apparently are no published data on the current status of the Stanwood population. If the population at Stanwood persists, new collections should be made to document its continued existence at the site, and if other populations are known or discovered, herbarium collections should be made to document their existences.
Spartina ×townsendii has not previously been reported from British Columbia. It is here reported as new for the province on the basis of two collections made in 2006 in in Boundary Bay at sites separated by some 4.4 kilometers (by air)[Taylor 80 (UBC, Fig. 11) and Saarela & Percy 791 (CAN, Fig. 12, UBC, Suppl. Fig. 34)]. These appear to be the most recent confirmed reports of the taxon in North America since it was collected at Stanwood, Washington. Herbarium specimens of these collections were initially determined (incorrectly) as Spartina anglica and Spartina alterniflora, since Spartina ×townsendii was not expected in British Columbia. Subsequent study of this material, in combination with the Spartina taxonomic literature and comparisons with specimens of Spartina anglica and Old World specimens of Spartina ×townsendii at CAN and UBC, confirmed the specimens to be Spartina ×townsendii, prompting the current taxonomic study. Pollen in these specimens is sterile, as determined by pollen staining (see discussion under Spartina anglica, Fig. 5), further confirming their identities as Spartina ×townsendii. Specimens from which pollen was extracted and stained with lactophenol cotton blue to assess fertility are identified with the symbol † in the Specimens Examined below.
The origin of Spartina ×townsendii in British Columbia is not known, and there are no data on the extent of the Boundary Bay sub-populations in 2006 aside from notes on the Saarela and Percy collection label indicating a single clump of the grass approximately one meter in diameter. It is not known if Spartina ×townsendiihas persisted in British Columbia since collected some five years ago. Since major efforts are ongoing to remove Spartina plants from Boundary Bay where Spartina ×townsendiiwas found, it is possible the original stands from which the specimens were collected have been removed. The region should be studied to determine if the taxon is present. Since the taxon is sterile and does not set seed, it must have been introduced into Boundary Bay by vegetative reproduction, probably from rhizome fragments transported in tidal currents. It is possible that the British Columbia plants originated from the stand at Stanwood, Washington, if it persists, or there may be other extant occurrences of Spartina ×townsendii somewhereto the south of Boundary Bay. Workers searching Puget Sound for invasive Spartina (e.g., Benbrook 2011[30]) should be aware that Spartina ×townsendii may also be present.
Morphology
The description here is based on the first known collection from Washington, the two collections from British Columbia, and Old World material housed at CAN and UBC (see Specimens Examined), including collections made by H. & J. Groves who first described the taxon over a century ago. The North American specimens of Spartina ×townsendii are morphologically similar to the Old World specimens examined. Spartina ×townsendii is distinguished from Spartina anglica by its shorter spikelets [(14–17.5 mm long vs. (15–)16.5–25 mm long]; shorter anthers [5–7(–8.5) mm long vs. 7–10 mm long]; indehiscent anthers that are not or incompletely exserted with sterile pollen [vs. dehiscent anthers that are usually fully exserted with fertile pollen; see Figs 4, 5]; shorter ligules [1–1.5 mm long vs. 1–3 mm long]; upper glumes 3-veined [vs. upper glumes 3–6-veined]; and shorter upper glumes [12.5–16.5 mm long vs. 13–22 mm long]. The angle of the leaf blade with the stem is 30–40° in Spartina ×townsendii, compared to 30–60° in Spartina anglica (Marchant 1968a[31]). This character can be difficult to evaluate on herbarium specimens depending on how they were pressed, but should be more reliable in the field for distinguishing the taxa, particularly if they occur together. If stands of Spartina ×townsendii are relocated in British Columbia or elsewhere in the region, the taxonomic utility of this character should be carefully evaluated. Marchant (1968a)[31] noted swards of Spartina ×townsendii to be distinct in appearance from swards of Spartina anglica in England, having high culm density and high tiller density (ca. 96/100 cm2 vs. ca. 52/100 cm2). It is not clear how Spartina ×townsendii differs morphologically from Spartina alterniflora × S. foliosa hybrids that have been documented in California (Daehler and Strong 1997[32], Ayres et al. 1999[33], 2003[34], 2004, Anttila et al. 2000[35]).
Specimens examined
Canada. British Columbia: Greater Vancouver Regional District: Boundary Bay Regional Park, Boundary Bay, S of Richmond along trail off 12 Avenue in Tsawwassen, near 1st viewing platform, 49°01'28"N, 123°03'14"W, ca. 0 m, 28 Nov 2006, J.M.Saarela & D.M.Percy 791 (CAN [CAN590439†, Fig. 12, UBC [UBCV228476†, Suppl. Fig. 34]); Greater Vancouver in marsh close to Undersea marshes and Pacific Flyway displays, Boundary Bay, 49°03'34"N, 123°01'27"W [secondary], 8 Nov 2006, T.Taylor 80 (UBC [UBCV222939†, Fig. 11). United States of America. Washington: Snohomish Co.:near Stanwood, ca. 48°14'N, 122°21'W, 26 Aug 1965, H.M.Austenson s.n. (WTU [WTU229915†, Suppl. Fig. 35]). England. Hampshire Co.: Hythe, South Hants, 9 Oct 1883, H.Groves s.n. (US [US555778]); Hayling Island, 13 Sep 1900, E.S.Marshall s.n. (CAN [CAN585633, Suppl. Fig. 36]); Southampton, 50°53'49"N, 01°24'15"W, Sep 1904, H.Groves & J.Groves 4596 (CAN [CAN251679†, Suppl. Fig. 37], US [US1535531]); Lymington, Keyhaven, 50°47'N, 00°58'W, 28 Aug 1977, G.Halliday 457/77 (CAN [CAN522593†, Suppl. Fig. 38]); Keyhaven, 50°43'22"N, 01°34'10"W, 30 Jul 1966, G.Halliday 100/66 (CAN [CAN301583, Suppl. Fig. 39]); Hants, Aug 1877, J.Groves s.n. (CAN [CAN421009†, Suppl. Fig. 40]); Hythe, Southampton, central marshes, male sterile, 50.8667°N, 01.3999°W uncertainty 7193 m, 10 Sep 1959, C.Marchant s.n. (UBC [UBCV221074, Suppl. Fig. 41]); N Hayling Island, Duckard Point, male sterile, 50.8051°N, 0.9778°W uncertainty 7194 m, 4 Aug 1960, C.Marchant s.n. (UBC [UBCV221101, Suppl. Fig. 42]); N Hayling Island, male sterile, 50.8051°N, 0.9778°W uncertainty 7194 m, 17 Aug 1961, C.Marchant s.n. (UBC [UBCV221098, Suppl. Fig. 43]); Eling, male sterile, 50.8999°N, 1.4833°W uncertainty 7193 m, 29 Aug 1961, C.Marchant s.n. (UBC [UBCV221100, Suppl. Fig. 44, UBC221095, Suppl. Fig. 45]); Hythe, south marshes, male sterile, 50.8666°N, 1.3999°W uncertainty 7193 m, 28 Oct 1959, C.Marchant s.n. (UBC [UBCV221093, Suppl. Fig. 46, UBCV221099, Suppl. Fig. 47]); Hythe, central marsh, near Sylvan Villa, giant male sterile, 50.8666°N, 1.3999°W uncertainty 7193 m, 16 Aug 1961, C.Marchant s.n. (UBC [UBCV221097, Suppl. Fig. 48, UBCV221096, Suppl. Fig. 49]); Hythe, Hants, male sterile, 50.8666°N, 01.3999°W uncertainty 7193 m, 10 Sep 1959, C.Marchant s.n. (UBC [UBCV221094, Suppl. Fig. 50]).
Taxon Treatment
- Saarela, J; 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 ×townsendii for British Columbia, Canada PhytoKeys, 10: 25-82. doi
Images
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![Figure 4. Anthers of a Spartina anglica [U.S.A., Washington, Pacific Co., Zika 17595, WTU], bar = 3 mm. b Spartina ×townsendii [England, Hythe, Southampton, Marchant s.n., UBCV221074], bar = 3 mm. Anthers of Spartina anglica are fully exserted at anthesis, dehiscent, and the pollen is fertile. The longitudinal splitting of the anthers is a good indicator of dehiscence. Anthers in Spartina ×townsendii are not or incompletely exserted at anthesis, indehiscent, and the pollen is sterile. Photos: J.M. Saarela.](/wiki/File:PhytoKeys-010-025-g004.jpg)
![Figure 5. Pollen stained with lactophenol cotton blue. a, b Spartina anglica, fertile pollen [Canada, British Columbia, Williams 2004[36]-3 (CAN)] c Spartina ×townsendii, sterile pollen [England, Hampshire, 1877, Groves s.n. (CAN)] d Spartina ×townsendii, sterile pollen [Canada, British Columbia, Saarela & Percy 791 (CAN)]](/wiki/File:PhytoKeys-010-025-g005.jpg)
Other References
- ↑ 1.0 1.1 Marchant C (1963) Corrected chromosome numbers for Spartina ×townsendii and its parent species. Nature 199: 929. doi: 10.1038/199929a0
- ↑ 2.0 2.1 Cope T, Gray A (2009) Grasses of the British Isles. Botanical Society of the British Isles, London, 612 pp.
- ↑ Scarton F, Ghirelli L, Curiel D, Rismondo A (2003) First data on Spartina ×townsendii in the lagoon of Venice (Italy). In: Ozhan E (Ed). Proceedings of the Sixth International Conference on the Mediterranean Coastal Environment, MEDCOAST 03. Ravenna, Italy, Vol. 2: 787-792.
- ↑ Partridge T (1987) Spartina in New Zealand. New Zealand Journal of Botany 25: 567-575.
- ↑ 5.0 5.1 Stapf O (1914) Townsend’s grass or ricegrass. Proceedings of the Bournemouth Natural Science Society 5: 76-82.
- ↑ Marchant C, Goodman P (1969c) Spartina maritima (Curtis) Fernald. Journal of Ecology 57: 287-291. doi: 10.2307/2258222
- ↑ 7.0 7.1 Goodman P, Braybrooks E, Lambert J (1959) Investigations into ‘die-back’ in Spartina Townsendii AGG.: I. The present status of Spartina Townsendii in Britain. Journal of Ecology 47: 651-677. doi: 10.2307/2257297
- ↑ 8.0 8.1 Hubbard C (1957) In report of British Ecological Society Symposium on Spartina. Journal of Ecology 45: 613.
- ↑ Hubbard J (1965) Spartina marshes in Southern England: VI. Pattern of invasion in Poole Harbour. Journal of Ecology 53: 799-813. doi: 10.2307/2257637
- ↑ Oliver F (1925) Spartina townsendii: Its mode of establishment, economic uses and taxonomic status. Journal of Ecology 13: 74-91. doi: 10.2307/2255557
- ↑ Harboard W (1949) Spartina townsendii: a valuable grass on tidal mudflats. New Zealand Journal of Agriculture 78: 507-508.
- ↑ Ranwell D (1967) World resources of Spartina townsendii (sensu lato) and economic use of Spartina marshland. Journal of Applied Ecology 4: 239-256. doi: 10.2307/2401421
- ↑ Foucaud J (1894) Un nouveau Spartina. Annales de la Société des Sciences Naturelles de la Charente-Maritime 31: 8-9.
- ↑ Guénégou M, Citharel J, Levasseur J (1988) The hybrid status of Spartina anglica (Poaceae). Enzymatic analysis of the species and of the presumed parents. Canadian Journal of Botany 66: 1830-1833. doi: 10.1139/b88-249
- ↑ Raybould A, Gray A, Lawrence M, Marshall D (1991) The evolution of Spartina anglica C. E. Hubbard (Gramineae): origin and genetic variability. Biological Journal of the Linnean Society 43: 111-126. doi: 10.1111/j.1095-8312.1991.tb00588.x
- ↑ 16.0 16.1 Ferris C, King R, Gray A (1997) Molecular evidence for the maternal parentage in the hybrid origin of Spartina anglica C.E. Hubbard. Molecular Ecology 6: 185-187. doi: 10.1046/j.1365-294X.1997.00165.x
- ↑ Ayres D, Strong D (2001) Origin and genetic diversity of Spartina anglica (Poaceae) using nuclear DNA markers. American Journal of Botany 88: 1863-1867. doi: 10.1046/j.1365-294x.1999.00679.x
- ↑ 18.0 18.1 18.2 Baumel A, Ainouche M, Misset M, Gourret J, Bayer R (2003) Genetic evidence for hybridization between the native Spartina maritima and the introduced Spartina alterniflora (Poaceae) in South-West France: Spartina × neyrautii re-examined. Plant Systematics and Evolution 237: 87-97. doi: 10.1007/s00606-002-0251-8
- ↑ Hubbard C (1978) Gramineae. (290) Spartina Schreb. In: Heywood AVH (Ed) Flora Europaea Notulae Systematicae ad Floram Europaeam spectantes no. 20. Botanical Journal of the Linnean Society 76: 384.
- ↑ Saint-Yves A (1932) Monographia Spartinarum. Candollea 5: 19-100.
- ↑ Jovet P (1941) Notes systematiques et ecologiques sure les Spartines du Sud-Ouest. Bulletin de la Société botanique de France 88: 115-123.
- ↑ Chevalier A (1933) Nouvelles observations sur les Spartina et spécialement sur le Spartina Townsendi. Bulletin de la Société botanique de France 80: 779-788.
- ↑ 23.0 23.1 23.2 Marchant C (1977) Hybrid characteristics in Spartina X neyrautii Fouc., a taxon rediscovered in northern Spain. Botanical Journal of the Linnean Society 74: 289-296. doi: 10.1111/j.1095-8339.1977.tb01182.x
- ↑ McNeill J, Barrie F, Burdet H, Demoulin V, Hawksworth D, Marhold K, Nicoloson D, Prado J, Silva P, Skog J, Wiersema J, Turland N (Eds) (2006) International Code of Botanical Nomenclature (Vienna Code). Regnum Vegetabile 146. A.R.G. Gantner Verlag KG.
- ↑ 25.0 25.1 Raybould A, Gray A, Lawrence M, Marshall D (1990) The origin and taxonomy of Spartina ×neyrautii Foucaud. Watsonia 18: 207-209.
- ↑ Hubbard J, Grimes B, Marchant C (1978) Some observations on the ecology and taxonomy of Spartina neyrautii and Spartina alterniflora growing in France and Spain and comparison with Spartina townsendii and Spartina anglica. Documents Phytosociologiques 2: 273-282.
- ↑ Hitchcock C, Cronquist A, Ownbey M, Thompson J (1969) Vascular plants of the Pacific Northwest. Part 1: Vascular cryptogams, gymnosperms, and monocotyledons. University of Washington Press, Seattle and London, 914 pp.
- ↑ Barkworth M (2003) Spartina Schreb. In: Barkworth M Capels K Long S Piep M (Eds). Flora of North America North of Mexico, Volume 25: Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press: 240-250.
- ↑ Kozloff E (2005) Plants of western Oregon, Washington & British Columbia. Timber Press, Portland, Oregon, 512 pp.
- ↑ Benbrook R (2011) Spartina grass: friend and foe? Sea Kayaker August 2011: 13-17.
- ↑ 31.0 31.1 Marchant A (1968a) Evolution in Spartina (Gramineae): I. The history and morphology of the genus in Britain. Journal of the Linnean Society (Botany) 60: 1-24. doi: 10.1111/j.1095-8339.1967.tb00076.x
- ↑ Daehler C, Strong D (1997) Hybridization between introduced smooth cordgrass (Spartina alterniflora; Poaceae) and native California cordgrass (Spartina foliosa) in San Francisco Bay, California, USA. American Journal of Botany 84: 607-611. doi: 10.2307/2445896
- ↑ Ayres D, Garcia-Rossi D, Davis H, Strong D (1999) Extent and degree of hybridization between exotic (Spartina alterniflora) and native (Spartina foliosa) cordgrass (Poaceae) in California, USA determined by random amplified polymorphic DNA (RAPDs). Molecular Ecology 8: 1179-1186. doi: 10.1046/j.1365-294x.1999.00679.x
- ↑ Ayres D, Strong D, Baye P (2003) Spartina foliosa – a common species on the road to rarity? Madroño 50: 209–213.
- ↑ Anttila C, King R, Ferris C, Ayres D, Strong D (2000) Reciprocal hybrid formation of Spartina in San Francisco Bay. Molecular Ecology 9: 765-770. doi: 10.1046/j.1365-294x.2000.00935.x
- ↑ Williams G (2004) Discovery of a new salt marsh invasive to British Columbia, English cordgrass (Spartina anglica C.E. Hubb.) and management initiatives in 2003. Botanical Electronic News 324. http://www.ou.edu/cas/botany-micro/ben/ben324.html [accessed 17 Dec 2010] (Archived by WebCite® at http://www.webcitation.org/5v3Kz4Nzl)
