treatments-xml/data/95/78/5F/95785F43378B5597B3153DB1E67E8A63.xml

<document ID-DOI="http://dx.doi.org/10.3897/phytokeys.10.2734" ID-PMC="PMC3310194" ID-Pensoft-Pub="1314-2003-10-25" ID-Pensoft-UUID="FFDEFFB2FFCC5E0D8269FFD7FFD28A31" ID-PubMed="22461730" ID-Zenodo-Dep="576110" ModsDocID="1314-2003-10-25" checkinTime="1451252035118" checkinUser="pensoft" docAuthor="Saarela, Jeffery M." docDate="2012" docId="95785F43378B5597B3153DB1E67E8A63" docLanguage="en" docName="PhytoKeys 10: 25-82" docOrigin="PhytoKeys 10" docPubDate="2012-03-21" docSource="http://dx.doi.org/10.3897/phytokeys.10.2734" docTitle="Spartina ×townsendii H. Groves &amp; J. Groves, Bot. Exch. Club Rep. 1880. 37. 1881." docType="treatment" docVersion="4" id="FFDEFFB2FFCC5E0D8269FFD7FFD28A31" lastPageNumber="51" masterDocId="FFDEFFB2FFCC5E0D8269FFD7FFD28A31" masterDocTitle="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" masterLastPageNumber="82" masterPageNumber="25" pageNumber="48" updateTime="1668138964632" updateUser="ExternalLinkService">
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<mods:title>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</mods:title>
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<mods:roleTerm>Author</mods:roleTerm>
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<mods:namePart>Saarela, Jeffery M.</mods:namePart>
<mods:affiliation>Research &amp; Collections, Canadian Museum of Nature, P. O. Box 3443 Stn. D, Ottawa, Ontario K 1 P 6 P 4, Canada</mods:affiliation>
<mods:nameIdentifier type="email">jsaarela@mus-nature.ca</mods:nameIdentifier>
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<mods:date>2012</mods:date>
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<mods:number>2012-03-21</mods:number>
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<subSection lastPageId="27" lastPageNumber="52" pageId="23" pageNumber="48" type="taxonomy">
<treatment ID-GBIF-Taxon="182229445" LSID="urn:lsid:plazi:treatment:95785F43378B5597B3153DB1E67E8A63" httpUri="http://treatment.plazi.org/id/95785F43378B5597B3153DB1E67E8A63" lastPageId="27" lastPageNumber="51" pageId="23" pageNumber="48">
<subSubSection pageId="23" pageNumber="48" type="nomenclature">
<paragraph pageId="23" pageNumber="48">
<taxonomicName LSID="95785F43-378B-5597-B315-3DB1E67E8A63" authority="H. Groves &amp; J. Groves, Bot. Exch. Club Rep. 1880. 37. 1881." authorityName="H. Groves &amp; J. Groves, Bot. Exch. Club Rep. 1880. 37. 1881." class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="Spartina ×townsendii" order="Poales" pageId="23" pageNumber="48" phylum="Tracheophyta" rank="species" species="×townsendii">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
H. Groves &amp; J. Groves, Bot. Exch. Club Rep. 1880. 37. 1881.
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="23" pageNumber="48" type="reference_group">
<paragraph pageId="23" pageNumber="48">
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="23" pageNumber="48" phylum="Tracheophyta" rank="species" species="xtownsendii">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</taxonomicName>
H. Groves &amp; J. Groves, Bot. Exch. Club Rep. 1880. 37. 1881. 
<emphasis bold="true" pageId="23" pageNumber="48">Type.</emphasis>
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.
</paragraph>
<paragraph pageId="23" pageNumber="48">
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="23" pageNumber="48" phylum="Tracheophyta" rank="species" species="xneyrautii">
Spartina 
<normalizedToken originalValue="×neyrautii">xneyrautii</normalizedToken>
</taxonomicName>
Fouc., Ann. Soc. Sci. Nat. Charente-Maritime 31: 8. 1894. 
<emphasis bold="true" pageId="23" pageNumber="48">Type.</emphasis>
France. 
<normalizedToken originalValue="près">pres</normalizedToken>
de Hendaye (
<normalizedToken originalValue="Basses-Pyrénnées">Basses-Pyrennees</normalizedToken>
), E-J.Neyraut s.n., 24 Jul 1892 (isotypes: P [P00753804!, P03457326!, P03457334!, P03457416!], US! (fragm. ex P)]).
</paragraph>
</subSubSection>
<subSubSection pageId="23" pageNumber="48" type="description">
<paragraph pageId="23" pageNumber="48">Description.</paragraph>
<paragraph pageId="23" pageNumber="48">
Culms 46-100 cm tall, thick, fleshy, rhizomatous. Sheaths glabrous; ligules 1-1.5 mm long; blades 6.5-37 cm long 
<normalizedToken originalValue="×">x</normalizedToken>
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 
<normalizedToken originalValue="×">x</normalizedToken>
7-25 mm wide at midpoint, erect, with (2)3-6(-10) branches; branches (6-)7.5-15(-18) cm long 
<normalizedToken originalValue="×">x</normalizedToken>
(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 
<normalizedToken originalValue="×">x</normalizedToken>
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 
<normalizedToken originalValue="×">x</normalizedToken>
0.5-0.7 mm wide, 1-veined, tips acuminate or obtuse; upper glumes 12.5-16.5 mm long 
<normalizedToken originalValue="×">x</normalizedToken>
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. 2 
<emphasis italics="true" pageId="23" pageNumber="48">n</emphasis>
= 62 (
<bibRefCitation author="Marchant, CJ" journalOrPublisher="Journal of Coastal Research" pageId="31" pageNumber="56" publicationUrl="10.1038/199929a0" refId="B90" refString="Marchant, CJ, 1963. Corrected chromosome numbers for Spartina xtownsendii and its parent species. Nature 199: 929. ., 10.1038/199929a0" title="Corrected chromosome numbers for Spartina xtownsendii and its parent species. Nature 199: 929." url="10.1038/199929a0" year="1963">Marchant 1963</bibRefCitation>
, 1968b).
</paragraph>
</subSubSection>
<subSubSection pageId="23" pageNumber="48" type="common name">
<paragraph pageId="23" pageNumber="48">Common name.</paragraph>
<paragraph pageId="23" pageNumber="48">
<normalizedToken originalValue="Townsend’s">Townsend's</normalizedToken>
cordgrass.
</paragraph>
</subSubSection>
<subSubSection pageId="23" pageNumber="48" type="etymology">
<paragraph pageId="23" pageNumber="48">Etymology.</paragraph>
<paragraph pageId="23" pageNumber="48">
The epithet 
<emphasis italics="true" pageId="23" pageNumber="48">townsendii</emphasis>
was given in honour of the English botanist Frederick Townsend (1822-1905).
</paragraph>
</subSubSection>
<subSubSection pageId="23" pageNumber="48" type="illustration">
<paragraph pageId="23" pageNumber="48">Illustration.</paragraph>
<paragraph pageId="23" pageNumber="48">
<bibRefCitation author="Cope, T" journalOrPublisher="Botanical Society of the British Isles, London" pageId="28" pageNumber="53" refId="B38" refString="Cope, T, Gray, A, 2009. Grasses of the British Isles. Botanical Society of the British Isles, London, 612 pp." title="Grasses of the British Isles." year="2009">Cope and Gray 2009</bibRefCitation>
:547.
</paragraph>
</subSubSection>
<subSubSection pageId="23" pageNumber="48" type="distribution">
<paragraph pageId="23" pageNumber="48">Distribution.</paragraph>
<paragraph pageId="23" pageNumber="48">
This species is found in England, Wales, Scotland, Ireland (
<bibRefCitation author="Cope, T" journalOrPublisher="Botanical Society of the British Isles, London" pageId="28" pageNumber="53" refId="B38" refString="Cope, T, Gray, A, 2009. Grasses of the British Isles. Botanical Society of the British Isles, London, 612 pp." title="Grasses of the British Isles." year="2009">Cope and Gray 2009</bibRefCitation>
), Italy (
<bibRefCitation author="Scarton, F" editor="Ozhan, E" journalOrPublisher="Ravenna, Italy, Vol. 2" pageId="33" pageNumber="58" pagination="787 - 792" refId="B127" refString="Scarton, F, Ghirelli, L, Curiel, D, Rismondo, A, 2003. First data on Spartina xtownsendii 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" title="First data on Spartina xtownsendii in the lagoon of Venice (Italy)." volumeTitle="Proceedings of the Sixth International Conference on the Mediterranean Coastal Environment, MEDCOAST 03." year="2003">Scarton et al. 2003</bibRefCitation>
), United States (Washington), New Zealand (
<bibRefCitation author="Partridge, TR" journalOrPublisher="New Zealand Journal of Botany" pageId="32" pageNumber="57" pagination="567 - 575" refId="B107" refString="Partridge, TR, 1987. Spartina in New Zealand. New Zealand Journal of Botany 25: 567 - 575" title="Spartina in New Zealand." volume="25" year="1987">Partridge 1987</bibRefCitation>
), and Canada (British Columbia).
</paragraph>
</subSubSection>
<subSubSection lastPageId="26" lastPageNumber="51" pageId="23" pageNumber="48" type="comments">
<paragraph pageId="23" pageNumber="48">Comments.</paragraph>
<paragraph lastPageId="24" lastPageNumber="49" pageId="23" pageNumber="48">
In the mid to late 1800s an unknown cordgrass of restricted distribution appeared and spread rapidly along the shores of Southampton Water, England (
<bibRefCitation author="Stapf, O" journalOrPublisher="Proceedings of the Bournemouth Natural Science Society" pageId="34" pageNumber="59" pagination="76 - 82" refId="B133" refString="Stapf, O, 1914. Townsend's grass or ricegrass. Proceedings of the Bournemouth Natural Science Society 5: 76 - 82" title="Townsend's grass or ricegrass." volume="5" year="1914">Stapf 1914</bibRefCitation>
), which differed morphologically (particularly by its sterile pollen) from 
<taxonomicName authorityName="M.L.Fernald" authorityYear="1916" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="maritima">
<emphasis italics="true" pageId="24" pageNumber="49">
<pageBreakToken pageId="24" pageNumber="49" start="start">Spartina</pageBreakToken>
maritima
</emphasis>
</taxonomicName>
, the single cordgrass species native to the Atlantic coast of Europe and north Africa (
<bibRefCitation author="Marchant, CJ" journalOrPublisher="Journal of Ecology" pageId="31" pageNumber="56" pagination="287 - 291" publicationUrl="10.2307/2258222" refId="B96" refString="Marchant, CJ, Goodman, PJ, 1969c. Spartina maritima (Curtis) Fernald. Journal of Ecology 57: 287 - 291, 10.2307/2258222" title="Spartina maritima (Curtis) Fernald." url="10.2307/2258222" volume="57" year="1969 c">Marchant and Goodman 1969c</bibRefCitation>
), and the introduced 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="alterniflora">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina alterniflora</emphasis>
</taxonomicName>
, 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 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
from plants collected near Hythe. In the 1890s a second form of 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
, 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 
<bibRefCitation author="Goodman, PJ" journalOrPublisher="Journal of Ecology" pageId="29" pageNumber="54" pagination="651 - 677" publicationUrl="10.2307/2257297" refId="B58" refString="Goodman, PJ, Braybrooks, EM, Lambert, JM, 1959. Investigations into 'die-back' in Spartina Townsendii AGG.: I. The present status of Spartina Townsendii in Britain. Journal of Ecology 47: 651 - 677, 10.2307/2257297" title="Investigations into ' die-back' in Spartina Townsendii AGG.: I. The present status of Spartina Townsendii in Britain." url="10.2307/2257297" volume="47" year="1959">Goodman et al. 1959</bibRefCitation>
, 
<bibRefCitation author="Hubbard, CE" journalOrPublisher="University of Washington Press, Seattle and London" pageId="30" pageNumber="55" refId="B78" refString="Hubbard, CE, 1957. In report of British Ecological Society Symposium on Spartina. Journal of Ecology 45: 613." title="In report of British Ecological Society Symposium on Spartina. Journal of Ecology 45: 613." year="1957">Hubbard 1957</bibRefCitation>
, 
<bibRefCitation author="Hubbard, JCE" journalOrPublisher="Journal of Ecology" pageId="30" pageNumber="55" pagination="799 - 813" publicationUrl="10.2307/2257637" refId="B79" refString="Hubbard, JCE, 1965. Spartina marshes in Southern England: VI. Pattern of invasion in Poole Harbour. Journal of Ecology 53: 799 - 813, 10.2307/2257637" title="Spartina marshes in Southern England: VI. Pattern of invasion in Poole Harbour." url="10.2307/2257637" volume="53" year="1965">1965</bibRefCitation>
). For decades these two forms (one sterile, the other fertile) of 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
were referred to collectively as the 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
aggregate or 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
sensu lato. Because of its vigorous growth and ability to rapidly colonize and stabilize mud flats, 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
s.l. was considered to be a 
<normalizedToken originalValue="“useful”">&quot;useful&quot;</normalizedToken>
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., 
<bibRefCitation author="Oliver, FW" journalOrPublisher="Journal of Ecology" pageId="32" pageNumber="57" pagination="74 - 91" publicationUrl="10.2307/2255557" refId="B105" refString="Oliver, FW, 1925. Spartina townsendii: Its mode of establishment, economic uses and taxonomic status. Journal of Ecology 13: 74 - 91, 10.2307/2255557" title="Spartina townsendii: Its mode of establishment, economic uses and taxonomic status." url="10.2307/2255557" volume="13" year="1925">Oliver 1925</bibRefCitation>
, 
<bibRefCitation author="Harboard, W" journalOrPublisher="New Zealand Journal of Agriculture" pageId="30" pageNumber="55" pagination="507 - 508" refId="B67" refString="Harboard, W, 1949. Spartina townsendii: a valuable grass on tidal mudflats. New Zealand Journal of Agriculture 78: 507 - 508" title="Spartina townsendii: a valuable grass on tidal mudflats." volume="78" year="1949">Harboard 1949</bibRefCitation>
, 
<bibRefCitation author="Goodman, PJ" journalOrPublisher="Journal of Ecology" pageId="29" pageNumber="54" pagination="651 - 677" publicationUrl="10.2307/2257297" refId="B58" refString="Goodman, PJ, Braybrooks, EM, Lambert, JM, 1959. Investigations into 'die-back' in Spartina Townsendii AGG.: I. The present status of Spartina Townsendii in Britain. Journal of Ecology 47: 651 - 677, 10.2307/2257297" title="Investigations into ' die-back' in Spartina Townsendii AGG.: I. The present status of Spartina Townsendii in Britain." url="10.2307/2257297" volume="47" year="1959">Goodman et al. 1959</bibRefCitation>
, 
<bibRefCitation author="Ranwell, D" journalOrPublisher="Journal of Applied Ecology" pageId="32" pageNumber="57" pagination="239 - 256" publicationUrl="10.2307/2401421" refId="B114" refString="Ranwell, D, 1967. World resources of Spartina townsendii (sensu lato) and economic use of Spartina marshland. Journal of Applied Ecology 4: 239 - 256, 10.2307/2401421" title="World resources of Spartina townsendii (sensu lato) and economic use of Spartina marshland." url="10.2307/2401421" volume="4" year="1967">Ranwell 1967</bibRefCitation>
).
</paragraph>
<paragraph pageId="24" pageNumber="49">
Soon after its formal description 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
was considered to be a species of hybrid origin. 
<bibRefCitation author="Foucaud, J" journalOrPublisher="Annales de la Societe des Sciences Naturelles de la Charente-Maritime" pageId="29" pageNumber="54" pagination="8 - 9" refId="B53" refString="Foucaud, J, 1894. Un nouveau Spartina. Annales de la Societe des Sciences Naturelles de la Charente-Maritime 31: 8 - 9" title="Un nouveau Spartina." volume="31" year="1894">Foucaud (1894)</bibRefCitation>
suggested that 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
was probably a hybrid of the native 
<taxonomicName authorityName="M.L.Fernald" authorityYear="1916" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="maritima">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina maritima</emphasis>
</taxonomicName>
and the introduced 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="alterniflora">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina alterniflora</emphasis>
</taxonomicName>
, a hypothesis later supported by 
<bibRefCitation author="Stapf, O" journalOrPublisher="Proceedings of the Bournemouth Natural Science Society" pageId="34" pageNumber="59" pagination="76 - 82" refId="B133" refString="Stapf, O, 1914. Townsend's grass or ricegrass. Proceedings of the Bournemouth Natural Science Society 5: 76 - 82" title="Townsend's grass or ricegrass." volume="5" year="1914">Stapf (1914)</bibRefCitation>
and Huskins (1930), who examined cytological evidence and hypothesized that fertile plants of 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
s.l. originated from chromosome doubling following hybridization between its parent species. 
<bibRefCitation author="Marchant, CJ" journalOrPublisher="Journal of Coastal Research" pageId="31" pageNumber="56" publicationUrl="10.1038/199929a0" refId="B90" refString="Marchant, CJ, 1963. Corrected chromosome numbers for Spartina xtownsendii and its parent species. Nature 199: 929. ., 10.1038/199929a0" title="Corrected chromosome numbers for Spartina xtownsendii and its parent species. Nature 199: 929." url="10.1038/199929a0" year="1963">Marchant (1963)</bibRefCitation>
confirmed this work, and reported chromosome numbers as 2 
<emphasis italics="true" pageId="24" pageNumber="49">n</emphasis>
= 62 for 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="alterniflora">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina alterniflora</emphasis>
</taxonomicName>
, 2 
<emphasis italics="true" pageId="24" pageNumber="49">n</emphasis>
= 60 for 
<taxonomicName authorityName="M.L.Fernald" authorityYear="1916" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="maritima">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina maritima</emphasis>
</taxonomicName>
, 2 
<emphasis italics="true" pageId="24" pageNumber="49">n</emphasis>
= 62 for sterile plants of 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
, and 2 
<emphasis italics="true" pageId="24" pageNumber="49">n</emphasis>
= 120, 122, 124 for fertile plants of 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
. The hybrid and chromosome doubling origins of the forms of 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
have been confirmed by multiple lines of molecular evidence (
<bibRefCitation author="Guenegou, M" journalOrPublisher="Canadian Journal of Botany" pageId="30" pageNumber="55" pagination="1830 - 1833" publicationUrl="10.1139/b88-249" refId="B65" refString="Guenegou, 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, 10.1139/b88-249" title="The hybrid status of Spartina anglica (Poaceae). Enzymatic analysis of the species and of the presumed parents." url="10.1139/b88-249" volume="66" year="1988">
<normalizedToken originalValue="Guénégou">Guenegou</normalizedToken>
et al. 1988
</bibRefCitation>
, 
<bibRefCitation author="Raybould, AF" journalOrPublisher="Biological Journal of the Linnean Society" pageId="33" pageNumber="58" pagination="111 - 126" publicationUrl="10.1111/j.1095-8312.1991.tb00588.x" refId="B116" refString="Raybould, AF, Gray, AJ, Lawrence, MJ, Marshall, DF, 1991. The evolution of Spartina anglica C. E. Hubbard (Gramineae): origin and genetic variability. Biological Journal of the Linnean Society 43: 111 - 126, 10.1111/j.1095-8312.1991.tb00588.x" title="The evolution of Spartina anglica C. E. Hubbard (Gramineae): origin and genetic variability." url="10.1111/j.1095-8312.1991.tb00588.x" volume="43" year="1991">Raybould et al. 1991</bibRefCitation>
, 
<bibRefCitation author="Ferris, C" journalOrPublisher="Molecular Ecology" pageId="29" pageNumber="54" pagination="185 - 187" publicationUrl="10.1046/j.1365-294X.1997.00165.x" refId="B52" refString="Ferris, C, King, RA, Gray, AJ, 1997. Molecular evidence for the maternal parentage in the hybrid origin of Spartina anglica C.E. Hubbard. Molecular Ecology 6: 185 - 187, 10.1046/j.1365-294X.1997.00165.x" title="Molecular evidence for the maternal parentage in the hybrid origin of Spartina anglica C. E. Hubbard." url="10.1046/j.1365-294X.1997.00165.x" volume="6" year="1997">Ferris et al. 1997</bibRefCitation>
, 
<bibRefCitation author="Ayres, DR" journalOrPublisher="American Journal of Botany" pageId="27" pageNumber="52" pagination="1863 - 1867" publicationUrl="10.1046/j.1365-294x.1999.00679.x" refId="B11" refString="Ayres, DR, Strong, DR, 2001. Origin and genetic diversity of Spartina anglica (Poaceae) using nuclear DNA markers. American Journal of Botany 88: 1863 - 1867, 10.1046/j.1365-294x.1999.00679.x" title="Origin and genetic diversity of Spartina anglica (Poaceae) using nuclear DNA markers." url="10.1046/j.1365-294x.1999.00679.x" volume="88" year="2001">Ayres and Strong 2001</bibRefCitation>
, 
<bibRefCitation author="Baumel, A" journalOrPublisher="Plant Systematics and Evolution" pageId="27" pageNumber="52" pagination="87 - 97" publicationUrl="10.1007/s00606-002-0251-8" refId="B21" refString="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 x neyrautii re-examined. Plant Systematics and Evolution 237: 87 - 97, 10.1007/s00606-002-0251-8" title="Genetic evidence for hybridization between the native Spartina maritima and the introduced Spartina alterniflora (Poaceae) in South-West France: Spartina x neyrautii re-examined." url="10.1007/s00606-002-0251-8" volume="237" year="2003">Baumel et al. 2003</bibRefCitation>
). 
<bibRefCitation author="Hubbard, CE" journalOrPublisher="University of Washington Press, Seattle and London" pageId="30" pageNumber="55" refId="B78" refString="Hubbard, CE, 1957. In report of British Ecological Society Symposium on Spartina. Journal of Ecology 45: 613." title="In report of British Ecological Society Symposium on Spartina. Journal of Ecology 45: 613." year="1957">Hubbard (1957)</bibRefCitation>
observed the type specimen of 
<taxonomicName authorityName="H. Groves &amp; J. Groves" authorityYear="1881" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="townsendii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina townsendii</emphasis>
</taxonomicName>
to be the sterile F1 hybrid, and the taxon was subsequently referred to as 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
. The fertile plants remained without a name until 
<bibRefCitation author="Hubbard, CE" journalOrPublisher="Journal of Ecology" pageId="30" pageNumber="55" refId="B81" refString="Hubbard, CE, 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." title="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." year="1978">Hubbard (1978)</bibRefCitation>
later described them as 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina anglica</emphasis>
</taxonomicName>
, and the two forms have since been recognized as distinct taxa. Molecular data have identified 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="alterniflora">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina alterniflora</emphasis>
</taxonomicName>
as the female parent and 
<taxonomicName authorityName="M.L.Fernald" authorityYear="1916" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="maritima">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina maritima</emphasis>
</taxonomicName>
as the male parent in the origin of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
(e.g., 
<bibRefCitation author="Ferris, C" journalOrPublisher="Molecular Ecology" pageId="29" pageNumber="54" pagination="185 - 187" publicationUrl="10.1046/j.1365-294X.1997.00165.x" refId="B52" refString="Ferris, C, King, RA, Gray, AJ, 1997. Molecular evidence for the maternal parentage in the hybrid origin of Spartina anglica C.E. Hubbard. Molecular Ecology 6: 185 - 187, 10.1046/j.1365-294X.1997.00165.x" title="Molecular evidence for the maternal parentage in the hybrid origin of Spartina anglica C. E. Hubbard." url="10.1046/j.1365-294X.1997.00165.x" volume="6" year="1997">Ferris et al. 1997</bibRefCitation>
).
</paragraph>
<paragraph lastPageId="25" lastPageNumber="50" pageId="24" pageNumber="49">
An independent origin of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
is documented in France. In 1894 Jules Foucaud described 
<taxonomicName authorityName="Foucaud" authorityYear="1894" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="neyrautii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina neyrautii</emphasis>
</taxonomicName>
Fouc. from southwestern France and northern Spain. 
<taxonomicName authorityName="Foucaud" authorityYear="1894" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="neyrautii">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina neyrautii</emphasis>
</taxonomicName>
was initially considered to be a variant of 
<taxonomicName authorityName="M.L.Fernald" authorityYear="1916" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="maritima">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina maritima</emphasis>
</taxonomicName>
(e.g., Chevalier 1923, 
<bibRefCitation author="Saint-Yves, A" journalOrPublisher="Candollea" pageId="33" pageNumber="58" pagination="19 - 100" refId="B122" refString="Saint-Yves, A, 1932. Monographia Spartinarum. Candollea 5: 19 - 100" title="Monographia Spartinarum." volume="5" year="1932">Saint-Yves 1932</bibRefCitation>
), but was later recognized as a morphologically and cytologically distinct hybrid, 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xneyrautii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×neyrautii">xneyrautii</normalizedToken>
</emphasis>
</taxonomicName>
,with the same parentage as 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
(e.g., 
<bibRefCitation author="Jovet, P" journalOrPublisher="Bulletin de la Societe botanique de France" pageId="31" pageNumber="56" pagination="115 - 123" refId="B83" refString="Jovet, P, 1941. Notes systematiques et ecologiques sure les Spartines du Sud-Ouest. Bulletin de la Societe botanique de France 88: 115 - 123" title="Notes systematiques et ecologiques sure les Spartines du Sud-Ouest." volume="88" year="1941">Jovet 1941</bibRefCitation>
, 
<bibRefCitation author="Chevalier, A" journalOrPublisher="Bulletin de la Societe botanique de France" pageId="28" pageNumber="53" pagination="779 - 788" refId="B34" refString="Chevalier, A, 1933. Nouvelles observations sur les Spartina et specialement sur le Spartina Townsendi. Bulletin de la Societe botanique de France 80: 779 - 788" title="Nouvelles observations sur les Spartina et specialement sur le Spartina Townsendi." volume="80" year="1933">Chevalier 1933</bibRefCitation>
, 
<bibRefCitation author="Marchant, CJ" journalOrPublisher="Botanical Journal of the Linnean Society" pageId="31" pageNumber="56" pagination="289 - 296" publicationUrl="10.1111/j.1095-8339.1977.tb01182.x" refId="B93" refString="Marchant, CJ, 1977. Hybrid characteristics in Spartina X neyrautii Fouc., a taxon rediscovered in northern Spain. Botanical Journal of the Linnean Society 74: 289 - 296, 10.1111/j.1095-8339.1977.tb01182.x" title="Hybrid characteristics in Spartina X neyrautii Fouc., a taxon rediscovered in northern Spain." url="10.1111/j.1095-8339.1977.tb01182.x" volume="74" year="1977">Marchant 1977</bibRefCitation>
). 
<bibRefCitation author="Baumel, A" journalOrPublisher="Plant Systematics and Evolution" pageId="27" pageNumber="52" pagination="87 - 97" publicationUrl="10.1007/s00606-002-0251-8" refId="B21" refString="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 x neyrautii re-examined. Plant Systematics and Evolution 237: 87 - 97, 10.1007/s00606-002-0251-8" title="Genetic evidence for hybridization between the native Spartina maritima and the introduced Spartina alterniflora (Poaceae) in South-West France: Spartina x neyrautii re-examined." url="10.1007/s00606-002-0251-8" volume="237" year="2003">Baumel et al. (2003)</bibRefCitation>
confirmed this with molecular data, demonstrating that 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xneyrautii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×neyrautii">xneyrautii</normalizedToken>
</emphasis>
</taxonomicName>
and 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
originated independently by hybridization between the same maternal (
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="alterniflora">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina alterniflora</emphasis>
</taxonomicName>
) and paternal (
<taxonomicName authorityName="M.L.Fernald" authorityYear="1916" class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="maritima">
<emphasis italics="true" pageId="24" pageNumber="49">Spartina maritima</emphasis>
</taxonomicName>
) taxa. Because both taxa apply to the hybrid 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="variety" species="alterniflora" subSpecies="x" variety="maritima">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina alterniflora 
<normalizedToken originalValue="×">x</normalizedToken>
S. maritima
</emphasis>
</taxonomicName>
, the later name 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xneyrautii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×neyrautii">xneyrautii</normalizedToken>
</emphasis>
</taxonomicName>
is a synonym of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="24" pageNumber="49" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="24" pageNumber="49">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
, in accordance with article H.2 of the Vienna Code (
<bibRefCitation author="McNeill, J" journalOrPublisher="New Phytologist" pageId="31" pageNumber="56" refId="B98" refString="McNeill, J, Barrie, FR, Burdet, HM, Demoulin, V, Hawksworth, DL, Marhold, K, Nicoloson, DH, Prado, J, Silva, PC, Skog, JE, Wiersema, JH, Turland, NJ, 2006. International Code of Botanical Nomenclature (Vienna Code). Regnum Vegetabile 146. A.R.G. Gantner Verlag KG." title="International Code of Botanical Nomenclature (Vienna Code). Regnum Vegetabile 146. A. R. G. Gantner Verlag KG." year="2006">McNeill et al. 2006</bibRefCitation>
), and as noted earlier by 
<bibRefCitation author="Raybould, AF" journalOrPublisher="Watsonia" pageId="32" pageNumber="57" pagination="207 - 209" refId="B115" refString="Raybould, AF, Gray, AJ, Lawrence, MJ, Marshall, DF, 1990. The origin and taxonomy of Spartina xneyrautii Foucaud. Watsonia 18: 207 - 209" title="The origin and taxonomy of Spartina xneyrautii Foucaud." volume="18" year="1990">Raybould et al. (1990)</bibRefCitation>
. In recent decades it has been documented that these hybrid plants are highly restricted in distribution in 
<pageBreakToken pageId="25" pageNumber="50" start="start">France</pageBreakToken>
(
<bibRefCitation author="Marchant, CJ" journalOrPublisher="Botanical Journal of the Linnean Society" pageId="31" pageNumber="56" pagination="289 - 296" publicationUrl="10.1111/j.1095-8339.1977.tb01182.x" refId="B93" refString="Marchant, CJ, 1977. Hybrid characteristics in Spartina X neyrautii Fouc., a taxon rediscovered in northern Spain. Botanical Journal of the Linnean Society 74: 289 - 296, 10.1111/j.1095-8339.1977.tb01182.x" title="Hybrid characteristics in Spartina X neyrautii Fouc., a taxon rediscovered in northern Spain." url="10.1111/j.1095-8339.1977.tb01182.x" volume="74" year="1977">Marchant 1977</bibRefCitation>
, 
<bibRefCitation author="Hubbard, J" journalOrPublisher="Documents Phytosociologiques" pageId="30" pageNumber="55" pagination="273 - 282" refId="B82" refString="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" title="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." volume="2" year="1978">Hubbard et al. 1978</bibRefCitation>
, 
<bibRefCitation author="Raybould, AF" journalOrPublisher="Watsonia" pageId="32" pageNumber="57" pagination="207 - 209" refId="B115" refString="Raybould, AF, Gray, AJ, Lawrence, MJ, Marshall, DF, 1990. The origin and taxonomy of Spartina xneyrautii Foucaud. Watsonia 18: 207 - 209" title="The origin and taxonomy of Spartina xneyrautii Foucaud." volume="18" year="1990">Raybould et al. 1990</bibRefCitation>
, 
<bibRefCitation author="Baumel, A" journalOrPublisher="Plant Systematics and Evolution" pageId="27" pageNumber="52" pagination="87 - 97" publicationUrl="10.1007/s00606-002-0251-8" refId="B21" refString="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 x neyrautii re-examined. Plant Systematics and Evolution 237: 87 - 97, 10.1007/s00606-002-0251-8" title="Genetic evidence for hybridization between the native Spartina maritima and the introduced Spartina alterniflora (Poaceae) in South-West France: Spartina x neyrautii re-examined." url="10.1007/s00606-002-0251-8" volume="237" year="2003">Baumel et al. 2003</bibRefCitation>
). Minor morphological differences between 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xneyrautii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×neyrautii">xneyrautii</normalizedToken>
</emphasis>
</taxonomicName>
and 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
were noted by 
<bibRefCitation author="Marchant, CJ" journalOrPublisher="Botanical Journal of the Linnean Society" pageId="31" pageNumber="56" pagination="289 - 296" publicationUrl="10.1111/j.1095-8339.1977.tb01182.x" refId="B93" refString="Marchant, CJ, 1977. Hybrid characteristics in Spartina X neyrautii Fouc., a taxon rediscovered in northern Spain. Botanical Journal of the Linnean Society 74: 289 - 296, 10.1111/j.1095-8339.1977.tb01182.x" title="Hybrid characteristics in Spartina X neyrautii Fouc., a taxon rediscovered in northern Spain." url="10.1111/j.1095-8339.1977.tb01182.x" volume="74" year="1977">Marchant (1977)</bibRefCitation>
. Measurements of spikelet characters in 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xneyrautii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×neyrautii">xneyrautii</normalizedToken>
</emphasis>
</taxonomicName>
type material examined at US fall within the range of variation reported here for 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph pageId="25" pageNumber="50">
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
has been introduced into North America, where apparently only a single occurrence has been reported in the literature. 
<bibRefCitation author="Hitchcock, CL" journalOrPublisher="University of Washington Press, Seattle and London" pageId="30" pageNumber="55" refId="B75" refString="Hitchcock, CL, Cronquist, A, Ownbey, M, Thompson, JW, 1969. Vascular plants of the Pacific Northwest. Part 1: Vascular cryptogams, gymnosperms, and monocotyledons. University of Washington Press, Seattle and London, 914 pp." title="Vascular plants of the Pacific Northwest. Part 1: Vascular cryptogams, gymnosperms, and monocotyledons." year="1969">Hitchcock et al. (1969)</bibRefCitation>
noted a single known population of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
in Washington at Stanwood, Snohomish Co. At the time of that publication, the fertile (= 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina anglica</emphasis>
</taxonomicName>
) and non-fertile forms of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
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 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
. The determination of a more recent collection from the Stanwood area identified as 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
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 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
(Giblin &amp; Legler 270 WTU) is here re-determined to be 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina anglica</emphasis>
</taxonomicName>
. 
<bibRefCitation author="Barkworth, ME" editor="Barkworth, ME" journalOrPublisher="Oxford University Press" pageId="27" pageNumber="52" pagination="240 - 250" refId="B17" refString="Barkworth, ME, 2003. Spartina Schreb. In: Barkworth, ME, Capels, KM, Long, S, Piep, MB, Eds., Flora of North America North of Mexico, Volume 25: Magnoliophyta: Commelinidae (in part): Poaceae, part 2. Oxford University Press: 240 - 250" title="Spartina Schreb." volumeTitle="Flora of North America North of Mexico, Volume 25: Magnoliophyta: Commelinidae (in part): Poaceae, part 2." year="2003">Barkworth (2003)</bibRefCitation>
included 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
in her treatment of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina</emphasis>
</taxonomicName>
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. 
<bibRefCitation author="Kozloff, EN" journalOrPublisher="Timber Press, Portland, Oregon" pageId="31" pageNumber="56" refId="B85" refString="Kozloff, EN, 2005. Plants of western Oregon, Washington &amp; British Columbia. Timber Press, Portland, Oregon, 512 pp." title="Plants of western Oregon, Washington &amp; British Columbia." year="2005">Kozloff (2005)</bibRefCitation>
also included 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
(as well as 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina anglica</emphasis>
</taxonomicName>
) in his Pacific Northwest flora, indicating only 'coastal salt 
<normalizedToken originalValue="marshes’">marshes'</normalizedToken>
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.
</paragraph>
<paragraph pageId="25" pageNumber="50">
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
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, 
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Photograph of a specimen of Spartina xtownsendii collected in Boundary Bay, British Columbia (Taylor 80, UBC). This is the first record of the taxon for British Columbia. Image published with the permission of the University of British Columbia Herbarium, Beaty Biodiversity Museum." httpUri="https://binary.pensoft.net/fig/10026" pageId="25" pageNumber="50">Fig. 11</figureCitation>
) and Saarela &amp; Percy 791 (CAN, 
<figureCitation captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Photograph of a specimen of Spartina xtownsendii collected in Boundary Bay, British Columbia (Saarela and Percy 791, CAN). This is the second record of the taxon for British Columbia." httpUri="https://binary.pensoft.net/fig/10027" pageId="25" pageNumber="50">Fig. 12</figureCitation>
, 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 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina anglica</emphasis>
</taxonomicName>
and 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="alterniflora">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina alterniflora</emphasis>
</taxonomicName>
, since 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
was not expected in British Columbia. Subsequent study of this material, in combination with the 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina</emphasis>
</taxonomicName>
taxonomic literature and comparisons with specimens of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina anglica</emphasis>
</taxonomicName>
and Old World specimens of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
at CAN and UBC, confirmed the specimens to be 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
, prompting the current taxonomic study. Pollen in these specimens is sterile, as determined by pollen staining (see discussion under 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="25" pageNumber="50">Spartina anglica</emphasis>
</taxonomicName>
, 
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Pollen stained with lactophenol cotton blue. a, b Spartina anglica, fertile pollen [Canada, British Columbia, Williams 2004 - 3 (CAN)] c Spartina xtownsendii, sterile pollen [England, Hampshire, 1877, Groves s. n. (CAN)] d Spartina xtownsendii, sterile pollen [Canada, British Columbia, Saarela &amp; Percy 791 (CAN)]" httpUri="https://binary.pensoft.net/fig/10020" pageId="25" pageNumber="50">Fig. 5</figureCitation>
), further confirming their identities as 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
. 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.
</paragraph>
<caption httpUri="https://binary.pensoft.net/fig/10026" pageId="25" pageNumber="50" start="Figure 11" startId="F11">
<paragraph pageId="25" pageNumber="50">
<emphasis bold="true" pageId="25" pageNumber="50">Figure 11.</emphasis>
Photograph of a specimen of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
collected in Boundary Bay, British Columbia (Taylor 80, UBC). This is the first record of the taxon for British Columbia. Image published with the permission of the University of British Columbia Herbarium, Beaty Biodiversity Museum.
</paragraph>
</caption>
<caption httpUri="https://binary.pensoft.net/fig/10027" pageId="25" pageNumber="50" start="Figure 12" startId="F12">
<paragraph pageId="25" pageNumber="50">
<emphasis bold="true" pageId="25" pageNumber="50">Figure 12.</emphasis>
Photograph of a specimen of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
collected in Boundary Bay, British Columbia (Saarela and Percy 791, CAN). This is the second record of the taxon for British Columbia.
</paragraph>
</caption>
<paragraph lastPageId="26" lastPageNumber="51" pageId="25" pageNumber="50">
The origin of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
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 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="25" pageNumber="50" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="25" pageNumber="50">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
has persisted in 
<pageBreakToken pageId="26" pageNumber="51" start="start">British</pageBreakToken>
Columbia since collected some five years ago. Since major efforts are ongoing to remove 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="26" pageNumber="51">Spartina</emphasis>
</taxonomicName>
plants from Boundary Bay where 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
was 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 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
somewhereto the south of Boundary Bay. Workers searching Puget Sound for invasive 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="genus">
<emphasis italics="true" pageId="26" pageNumber="51">Spartina</emphasis>
</taxonomicName>
(e.g., 
<bibRefCitation author="Benbrook, R" journalOrPublisher="Phytologia" pageId="27" pageNumber="52" refId="B23" refString="Benbrook, R, 2011. Spartina grass: friend and foe? Sea Kayaker August 2011: 13-17." title="Spartina grass: friend and foe? Sea Kayaker August 2011: 13 - 17." year="2011">Benbrook 2011</bibRefCitation>
) should be aware that 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
may also be present.
</paragraph>
</subSubSection>
<subSubSection pageId="26" pageNumber="51" type="morphology">
<paragraph pageId="26" pageNumber="51">Morphology.</paragraph>
<paragraph pageId="26" pageNumber="51">
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. &amp; J. Groves who first described the taxon over a century ago. The North American specimens of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
are morphologically similar to the Old World specimens examined. 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
is distinguished from 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="26" pageNumber="51">Spartina anglica</emphasis>
</taxonomicName>
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 
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Anthers of a Spartina anglica [U. S. A., Washington, Pacific Co., Zika 17595, WTU], bar = 3 mm. b Spartina xtownsendii [England, Hythe, Southampton, Marchant s. n., UBCV 221074], 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 xtownsendii are not or incompletely exserted at anthesis, indehiscent, and the pollen is sterile. Photos: J. M. Saarela." httpUri="https://binary.pensoft.net/fig/10019" pageId="26" pageNumber="51">Figs 4</figureCitation>
, 
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Pollen stained with lactophenol cotton blue. a, b Spartina anglica, fertile pollen [Canada, British Columbia, Williams 2004 - 3 (CAN)] c Spartina xtownsendii, sterile pollen [England, Hampshire, 1877, Groves s. n. (CAN)] d Spartina xtownsendii, sterile pollen [Canada, British Columbia, Saarela &amp; Percy 791 (CAN)]" httpUri="https://binary.pensoft.net/fig/10020" pageId="26" pageNumber="51">5</figureCitation>
]; 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 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
, compared to 30-60° in 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="26" pageNumber="51">Spartina anglica</emphasis>
</taxonomicName>
(
<bibRefCitation author="Marchant, AD" journalOrPublisher="Journal of the Linnean Society (Botany)" pageId="31" pageNumber="56" pagination="1 - 24" publicationUrl="10.1111/j.1095-8339.1967.tb00076.x" refId="B91" refString="Marchant, AD, 1968a. Evolution in Spartina (Gramineae): I. The history and morphology of the genus in Britain. Journal of the Linnean Society (Botany) 60: 1 - 24, 10.1111/j.1095-8339.1967.tb00076.x" title="Evolution in Spartina (Gramineae): I. The history and morphology of the genus in Britain." url="10.1111/j.1095-8339.1967.tb00076.x" volume="60" year="1968 a">Marchant 1968a</bibRefCitation>
). 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 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
are relocated in British Columbia or elsewhere in the region, the taxonomic utility of this character should be carefully evaluated. 
<bibRefCitation author="Marchant, AD" journalOrPublisher="Journal of the Linnean Society (Botany)" pageId="31" pageNumber="56" pagination="1 - 24" publicationUrl="10.1111/j.1095-8339.1967.tb00076.x" refId="B91" refString="Marchant, AD, 1968a. Evolution in Spartina (Gramineae): I. The history and morphology of the genus in Britain. Journal of the Linnean Society (Botany) 60: 1 - 24, 10.1111/j.1095-8339.1967.tb00076.x" title="Evolution in Spartina (Gramineae): I. The history and morphology of the genus in Britain." url="10.1111/j.1095-8339.1967.tb00076.x" volume="60" year="1968 a">Marchant (1968a)</bibRefCitation>
noted swards of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
to be distinct in appearance from swards of 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="anglica">
<emphasis italics="true" pageId="26" pageNumber="51">Spartina anglica</emphasis>
</taxonomicName>
in England, having high culm density and high tiller density (ca. 96/100 cm2 vs. ca. 52/100 cm2). It is not clear how 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="species" species="xtownsendii">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina 
<normalizedToken originalValue="×townsendii">xtownsendii</normalizedToken>
</emphasis>
</taxonomicName>
differs morphologically from 
<taxonomicName class="Liliopsida" family="Poaceae" genus="Spartina" higherTaxonomySource="GBIF" kingdom="Plantae" lsidName="" order="Poales" pageId="26" pageNumber="51" phylum="Tracheophyta" rank="variety" species="alterniflora" subSpecies="x" variety="foliosa">
<emphasis italics="true" pageId="26" pageNumber="51">
Spartina alterniflora 
<normalizedToken originalValue="×">x</normalizedToken>
S. foliosa
</emphasis>
</taxonomicName>
hybrids that have been documented in California (
<bibRefCitation author="Daehler, CC" journalOrPublisher="American Journal of Botany" pageId="28" pageNumber="53" pagination="607 - 611" publicationUrl="10.2307/2445896" refId="B40" refString="Daehler, CC, Strong, DR, 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, 10.2307/2445896" title="Hybridization between introduced smooth cordgrass (Spartina alterniflora; Poaceae) and native California cordgrass (Spartina foliosa) in San Francisco Bay, California, USA." url="10.2307/2445896" volume="84" year="1997">Daehler and Strong 1997</bibRefCitation>
, 
<bibRefCitation author="Ayres, D" journalOrPublisher="Molecular Ecology" pageId="27" pageNumber="52" pagination="1179 - 1186" publicationUrl="10.1046/j.1365-294x.1999.00679.x" refId="B10" refString="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, 10.1046/j.1365-294x.1999.00679.x" title="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)." url="10.1046/j.1365-294x.1999.00679.x" volume="8" year="1999">Ayres et al. 1999</bibRefCitation>
, 
<bibRefCitation author="Ayres, D" journalOrPublisher="American Journal of Botany" pageId="27" pageNumber="52" refId="B12" refString="Ayres, D, Strong, DR, Baye, P, 2003. Spartina foliosa - a common species on the road to rarity? Madrono 50: 209-213." title="Spartina foliosa - a common species on the road to rarity? Madrono 50: 209 - 213." year="2003">2003</bibRefCitation>
, 2004, 
<bibRefCitation author="Anttila, C" journalOrPublisher="Molecular Ecology" pageId="27" pageNumber="52" pagination="765 - 770" publicationUrl="10.1046/j.1365-294x.2000.00935.x" refId="B8" refString="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, 10.1046/j.1365-294x.2000.00935.x" title="Reciprocal hybrid formation of Spartina in San Francisco Bay." url="10.1046/j.1365-294x.2000.00935.x" volume="9" year="2000">Anttila et al. 2000</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection lastPageId="27" lastPageNumber="52" pageId="26" pageNumber="51" type="specimens examined">
<paragraph pageId="26" pageNumber="51">Specimens examined.</paragraph>
<paragraph lastPageId="27" lastPageNumber="52" pageId="26" pageNumber="51">
<emphasis bold="true" pageId="26" pageNumber="51">Canada. British Columbia: Greater Vancouver Regional District:</emphasis>
Boundary Bay Regional Park, Boundary Bay, S of Richmond along trail off 12 Avenue in Tsawwassen, near 1st viewing platform, 
<geoCoordinate degrees="49" direction="north" minutes="01" orientation="latitude" precision="15" seconds="28" value="49.024445">49°01'28&quot;N</geoCoordinate>
, 
<geoCoordinate degrees="123" direction="west" minutes="03" orientation="longitude" precision="15" seconds="14" value="-123.053894">123°03'14&quot;W</geoCoordinate>
, ca. 0 m, 28 Nov 2006, J.M.Saarela &amp; D.M.Percy 791 (CAN [CAN590439†, 
<figureCitation captionStart="Figure 12" captionStartId="F12" captionText="Figure 12. Photograph of a specimen of Spartina xtownsendii collected in Boundary Bay, British Columbia (Saarela and Percy 791, CAN). This is the second record of the taxon for British Columbia." httpUri="https://binary.pensoft.net/fig/10027" pageId="26" pageNumber="51">Fig. 12</figureCitation>
, UBC [UBCV228476†, Suppl. Fig. 34]); Greater Vancouver in marsh close to Undersea marshes and Pacific Flyway displays, Boundary Bay, 
<geoCoordinate degrees="49" direction="north" minutes="03" orientation="latitude" precision="15" seconds="34" value="49.059444">49°03'34&quot;N</geoCoordinate>
, 
<geoCoordinate degrees="123" direction="west" minutes="01" orientation="longitude" precision="15" seconds="27" value="-123.02417">123°01'27&quot;W</geoCoordinate>
[secondary], 8 Nov 2006, T.Taylor 80 (UBC [UBCV222939†, 
<figureCitation captionStart="Figure 11" captionStartId="F11" captionText="Figure 11. Photograph of a specimen of Spartina xtownsendii collected in Boundary Bay, British Columbia (Taylor 80, UBC). This is the first record of the taxon for British Columbia. Image published with the permission of the University of British Columbia Herbarium, Beaty Biodiversity Museum." httpUri="https://binary.pensoft.net/fig/10026" pageId="26" pageNumber="51">Fig. 11</figureCitation>
). 
<emphasis bold="true" pageId="26" pageNumber="51">United States of America. Washington:</emphasis>
Snohomish Co.:near Stanwood, ca. 
<geoCoordinate degrees="48" direction="north" minutes="14" orientation="latitude" precision="925" value="48.233334">48°14'N</geoCoordinate>
, 
<geoCoordinate degrees="122" direction="west" minutes="21" orientation="longitude" precision="925" value="-122.35">122°21'W</geoCoordinate>
, 26 Aug 1965, H.M.Austenson s.n. (WTU [WTU229915†, Suppl. Fig. 35]). 
<emphasis bold="true" pageId="26" pageNumber="51">England.</emphasis>
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]
<pageBreakToken pageId="27" pageNumber="52" start="start">)</pageBreakToken>
; Southampton, 
<geoCoordinate degrees="50" direction="north" minutes="53" orientation="latitude" precision="15" seconds="49" value="50.896946">50°53'49&quot;N</geoCoordinate>
, 
<geoCoordinate degrees="01" direction="west" minutes="24" orientation="longitude" precision="15" seconds="15" value="-1.4041667">01°24'15&quot;W</geoCoordinate>
, Sep 1904, H.Groves &amp; J.Groves 4596 (CAN [CAN251679†, Suppl. Fig. 37], US [US1535531]); Lymington, Keyhaven, 
<geoCoordinate degrees="50" direction="north" minutes="47" orientation="latitude" precision="925" value="50.783333">50°47'N</geoCoordinate>
, 
<geoCoordinate degrees="00" direction="west" minutes="58" orientation="longitude" precision="925" value="-0.96666664">00°58'W</geoCoordinate>
, 28 Aug 1977, G.Halliday 457/77 (CAN [CAN522593†, Suppl. Fig. 38]); Keyhaven, 
<geoCoordinate degrees="50" direction="north" minutes="43" orientation="latitude" precision="15" seconds="22" value="50.72278">50°43'22&quot;N</geoCoordinate>
, 
<geoCoordinate degrees="01" direction="west" minutes="34" orientation="longitude" precision="15" seconds="10" value="-1.5694444">01°34'10&quot;W</geoCoordinate>
, 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, 
<geoCoordinate degrees="50.8667" direction="north" orientation="latitude" precision="5" value="50.8667">50.8667°N</geoCoordinate>
, 
<geoCoordinate degrees="01.3999" direction="west" orientation="longitude" precision="5" value="-1.3999">01.3999°W</geoCoordinate>
uncertainty 7193 m, 10 Sep 1959, C.Marchant s.n. (UBC [UBCV221074, Suppl. Fig. 41]); N Hayling Island, Duckard Point, male sterile, 
<geoCoordinate degrees="50.8051" direction="north" orientation="latitude" precision="5" value="50.8051">50.8051°N</geoCoordinate>
, 
<geoCoordinate degrees="0.9778" direction="west" orientation="longitude" precision="5" value="-0.9778">0.9778°W</geoCoordinate>
uncertainty 7194 m, 4 Aug 1960, C.Marchant s.n. (UBC [UBCV221101, Suppl. Fig. 42]); N Hayling Island, male sterile, 
<geoCoordinate degrees="50.8051" direction="north" orientation="latitude" precision="5" value="50.8051">50.8051°N</geoCoordinate>
, 
<geoCoordinate degrees="0.9778" direction="west" orientation="longitude" precision="5" value="-0.9778">0.9778°W</geoCoordinate>
uncertainty 7194 m, 17 Aug 1961, C.Marchant s.n. (UBC [UBCV221098, Suppl. Fig. 43]); Eling, male sterile, 
<geoCoordinate degrees="50.8999" direction="north" orientation="latitude" precision="5" value="50.8999">50.8999°N</geoCoordinate>
, 
<geoCoordinate degrees="1.4833" direction="west" orientation="longitude" precision="5" value="-1.4833">1.4833°W</geoCoordinate>
uncertainty 7193 m, 29 Aug 1961, C.Marchant s.n. (UBC [UBCV221100, Suppl. Fig. 44, UBC221095, Suppl. Fig. 45]); Hythe, south marshes, male sterile, 
<geoCoordinate degrees="50.8666" direction="north" orientation="latitude" precision="5" value="50.8666">50.8666°N</geoCoordinate>
, 
<geoCoordinate degrees="1.3999" direction="west" orientation="longitude" precision="5" value="-1.3999">1.3999°W</geoCoordinate>
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, 
<geoCoordinate degrees="50.8666" direction="north" orientation="latitude" precision="5" value="50.8666">50.8666°N</geoCoordinate>
, 
<geoCoordinate degrees="1.3999" direction="west" orientation="longitude" precision="5" value="-1.3999">1.3999°W</geoCoordinate>
uncertainty 7193 m, 16 Aug 1961, C.Marchant s.n. (UBC [UBCV221097, Suppl. Fig. 48, UBCV221096, Suppl. Fig. 49]); Hythe, Hants, male sterile, 
<geoCoordinate degrees="50.8666" direction="north" orientation="latitude" precision="5" value="50.8666">50.8666°N</geoCoordinate>
, 
<geoCoordinate degrees="01.3999" direction="west" orientation="longitude" precision="5" value="-1.3999">01.3999°W</geoCoordinate>
uncertainty 7193 m, 10 Sep 1959, C.Marchant s.n. (UBC [UBCV221094, Suppl. Fig. 50]).
</paragraph>
</subSubSection>
</treatment>
</subSection>
</document>