Taxonomic reassessment of Polysiphonia foetidissima (Rhodomelaceae, Rhodophyta) and similar species, including P. schneideri, a newly introduced species in Europe

Figures & data

Fig. 1. Maximum likelihood tree for Polysiphonia and their relatives from plastid-encoded rbcL sequence data. The bootstrap values shown above the branches are from 1000 bootstrap resamplings.

Figs 2–6. Polysiphonia foetidissima: type material. 2. Lectotype (PC 0146017). 3. Cross section of an axis with 8 pericentral cells. 4. Rhizoids cut off from pericentral cells (arrow). 5. Apex showing a branch formed in the axil of a trichoblast (arrow) and trichoblasts or scar cells several segments apart (arrowheads). 6. Axes with a scar cell of a trichoblast (arrowhead). Scale bars = 1.5 cm (Fig. 2), 25 µm (Figs 3–5) and 90 µm (Fig. 6).

Figs 7–9. Polysiphonia foetidissima: vegetative morphology of Iberian specimens. 7. Habit of vegetative thallus. 8. Erect axis with trichoblasts three or four segments apart and branches seven or eight segments apart. 9. Creeping axis with rhizoids cut off from pericentral cells and two adventitious branches. Scale bars = 1 mm (Fig. 7) and 100 µm (Figs 8, 9).

Figs 10–17. Polysiphonia foetidissima: vegetative morphology of Iberian specimens. 10. Entangled decumbent axes on a turf of Rhodothamniella floridula. 11, 12. Erect axes with trichoblasts 4 segments apart and branches 7 or 8 segments apart. 13–17. Cross- section of axes with 6–9 pericentral cells. Scale bars = 1 cm (Fig. 10), 400 µm (Fig. 11), 60 µm (Fig. 12) and 50 µm (Figs 13–17).

Figs 18–21. Polysiphonia foetidissima: reproductive morphology of Iberian specimens. 18. Branch with tetrasporangia in spiral series. 19. Mature procarp showing the axial cell (ax), the supporting cell (su), one-celled second sterile group (st2) and four-celled carpogonial branch (1–4). 20. Erect axis with cystocarps. 21. Apical portion of an erect axis with cylindrical spermatangial branches. Scale bars = 100 µm (Figs 18, 20, 21), and 10 µm (Fig. 19).

Figs 22–25. Polysiphonia foetidissima: reproductive morphology of Iberian specimens. 22. Branches with tetrasporangia in spiral series. 23. Erect axes with cystocarps. 24. Apical portions of erect axes with cylindrical spermatangial branches. 25. Forked spermatangial branch. Scale bars = 400 µm (Figs 22–24) and 50 µm (Fig. 25).

Fig. 26. Distribution of Polysiphonia foetidissima (circles) and P. schneideri (squares) along the Atlantic coasts of the Iberian Peninsula.

Figs 27–36. Polysiphonia schneideri: vegetative and reproductive morphology of Iberian specimens. 27. Habit of vegetative thallus. 28. Rhizoids cut off from pericentral cells (arrow). 29, 30. Cross-sections of axes with 6 or 7 pericentral cells. 31. Branches growing in the axils of trichoblasts (arrow). 32. Trichoblasts borne several segments apart (arrowheads). 33. Tetrasporangia in straight series. 34. Procarp with four-celled carpogonial branch (1–4). 35. Mature cystocarp. 36. Spermatangial branch formed at one of the branches of the first dichotomy of a trichoblast. Scale bars = 0.5 cm (Fig. 27), 100 µm (Fig. 28), 50 µm (Figs 29–32, 36), 200 µm (Figs 33, 35) and 25 µm (Fig. 34).

Fig. 37. World distribution map of Polysiphonia foetidissima and related species. P. foetidissima (● type locality, ● reports with descriptions, ○ reports), P. isogona (▬ type locality, ▬ reports with descriptions), P. kappannae (♦ type locality), P. nizamudinii (▼ type locality), P. schneideri (* type locality), P. stuposa (■ type locality, □ reports) and P. tepida (▲ type locality, ▲ reports with descriptions, ∆ reports). References: 1, Kützing (1864); 2, Bornet (1892); 3, Collins & Hervey (1917); 4, Howe (1918); 5, Batten (1923); 6, Newton (1931); 7, Hollenberg (1958); 8, Lancelot (1966); 9, Sreenivasa Rao (1967); 10, Oliveira Filho (1969); 11, Hollenberg (1968); 12, Giaccone (1969); 13, Ardré (1970); 14, Cordeiro-Marino (1977); 15, Kapraun (1977); 16, Farooqui & Begum (1978); 17, Giaccone (1978); 18, Kapraun (1979); 19, Taylor (1960); 20, Schnetter & Schnetter (1981); 21, Weisscher (1983); 22, Audiffred & Prud´homme van Reine (1985); 23, Giaccone et al. (1985); 24, Conde & Soto (1986); 25, Silva et al. (1987); 26, Ballesteros (1990); 27, Adams (1991); 28, Schneider & Searles (1991); 29, Ballesteros (1993); 30, Lawson et al. (1995); 31, Silva et al. (1996); 32, Abbott (1999); 33, Coll & Oliveira (1999); 34, Rojas-González & Afonso-Carrillo (2000, 2008); 35, McDermid et al. (2002); 36, Womersley (2003); 37, Guimarães et al. (2004); 38, Suárez (2005); 39, Duncan & Lee Lum (2006); 40, Tsuda et al. (2006); 41, Dawes & Mathieson (2008); 42, Taskin et al. (2008); 43, Stuercke & Freshwater (2010); 44, Mamoozadeh & Freshwater (2011).

Figs 38–42. Polysiphonia stuposa: type material. 38. Holotype (MEL 2324435, collected by Sandri in Dalmatia, Croatia). 39. Prostrate axis with rhizoids cut off from pericentral cells. 40. Apex of an erect axis showing branch origin not associated with trichoblasts. 41. Cross section of an axis with 7 pericentral cells. 42. Erect axes. Scale bars = 1 cm (Fig. 38), 150 µm (Fig. 39), 30 µm (Fig. 40), 20 µm (Fig. 41) and 600 µm (Fig. 42).

Figs 43–47. Polysiphonia tepida: type material. 43. Holotype (MICH, Herbarium of W.R. Taylor, collected by H.L. Blomquist in 1940 in Beaufort, North Carolina, U.S.A.). 44. Cross-section of an axis with eight pericentral cells. 45. Creeping axis with a rhizoid cut off from a pericentral cell. 46. Apical portions of an erect axis with abundant trichoblasts borne 3 or 4 segments apart and branches arising from the axils of trichoblasts. 47. Scar cell of a trichoblast on an erect axis. Scale bars = 2 cm (Fig. 43) and 50 µm (Figs 44–47).

Figs 48–55. Polysiphonia exilis: type material. 48. Holotype (TCD 0012730, collected by Harvey at Key West, Florida). 49. Erect axes with secund branches. 50. Cross-section of thallus with nine pericentral cells. 51. Rhizoid cut off from a pericentral cell (arrow). 52. Detail of an erect axis with scar cells of trichoblasts on every segment (arrows) and pericentral cells in straight rows. 53. Detail of an erect axis with young cicatrigenous branches growing from scar cells (arrowheads); the arrow shows a scar cell of a trichoblast. 54. Detail of an axis bearing a branch with young tetrasporangia forming a spiral series; the arrowhead indicates a young cicatrigenous branch. 55. Detail of a prostrate axis bearing erect axes at irregular intervals. Scale bars = 2.5 cm (Fig. 48), 3 mm (Fig. 49), 50 µm (Figs 50–53), 250 µm (Fig. 54) and 650 µm (Fig. 55).

Figs 56–61. Polysiphonia brodiei: small and creeping forms in the Iberian Peninsula. 56. Habit of a tetrasporophyte growing on Codium adhaerens. 57. Spermatangial branches borne on every segment and without sterile apical cells. 58. Erect axes with scar cells of trichoblasts on every segment. 59–61. Upper portions of erect axes with branches arising in the axils of trichoblasts, 3 or 4 segments apart. Scale bars = 2 cm (Fig. 56), 80 µm (Fig. 57), 50 µm (Figs 58, 59), 500 µm (Fig. 60) and 200 µm (Fig. 61).

Table 1. Main taxonomic features of Polysiphonia foetidissima and similar species with 6–11 pericentral cells, rhizoids cut off from pericentral cells, and prostrate or creeping habit.

	P. foetidissima	P. schneideri	P. brodiei	P. confusa	P. exilis	P. isogona	P. kappannae	P. nizamuddinii	P. stuposa	P. tepida
Type locality	Plymouth, Cornwall, UK	Wrightsville Beach, North Carolina, USA	Bantry Bay, Cork, UK	California, USA	Key West, Florida	Blind Bay, New Zealand	Okga Port, India	Paradisepoint, Pakistan	Dalmatia, Croatia	Beaufort, North Carolina USA
Distribution	Atlantic Europe and North America	West Atlantic, southern Spain	North Atlantic, Mediterranean, PacificNorth America, Australia, New Zealand	Pacific America	Bermuda Caribbean Brazil, Australia Pacific islands, Maldives	New Zealand, Southern Australia	India	Pakistan	Mediterranean Sea	Atlantic North America
Pericentral cells	(6 or) 7or 8 (or 9)	(5 or) 6 (or 7)	(6 or) 7 or 8	8–10	8–11	(7–) 9 or 10 (–12)	7–9	(8 or) 9 (or 10)	7 or8	(6 or) 7 or 8
Cortication	ecorticate	ecorticate	slight to heavy	ecorticate	ecorticate	ecorticate	ecorticate	ecorticate	ecorticate	ecorticate
Branch origin	axil of trichoblast	axil of trichoblast	axil of trichoblast	axil of trichoblast	not associated withtrichoblast	axil of trichoblast	axil of trichoblast	axil of trichoblast	not associated with trichoblast	axil of trichoblast
Trichoblasts (abundance; segments between successive; divergence)	abundant; 3–5; 1/4	few; not one per segment; irregular	abundant; one per segment; 1/7–1/8	abundant; oneper segment; 1/4	abundant; one persegment; 1/4	variable	abundant; several; 1/4	abundant; 3 or 4; –	absent	abundant to scarce; 3–5; 1/4
Carpogonial branch cells; Cystocarp	4; globose	4; globose	4; globose	–	–	–; subspherical	–; spherical	–; globose		–; ovate to urceolate
Spermatangial branches	basal branch of trichoblasts, sometimes forked	basal branch of trichoblasts	basal branch of trichoblasts	–	basal branch of trichoblastsor replacing them	basal branch of trichoblasts	–	basal branch of trichoblasts	–	basal branch of trichoblasts, sometimes forked
Tetrasporangia	spiral series	linear series	spiral series	spiral series	spiral series	spiral series	linear series	linear series	–	spiral series
References	This study	Stuercke & Freshwater (2010); this study	Maggs & Hommersand (1993); this study	Hollenberg (1944 as P. inconspicua Hollenberg nom. illeg.; 1961)	Harvey (1853); Hollenberg (1968); Kapraun & Norris (1982); Schneider & Searles (1997); Abbott (1999); this study	Adams (1991); Womersley (2003).	Sreenivasa Rao (1967).	Farooqui & Begum (1978).	Kützing (1864); this study	Hollenberg (1958), Kapraun (1977), this work.

Table S1. Materials collected during this study, including herbarium codes and Genbank accession numbers.

Species	Locality	Date	Collectors	Herbarium	GenBank
Polysiphonia brodiei creeping froms	Spain: Guipúzcoa, Igeldo (43º19’06’’N; 2º02’14’’W)	13.iii.2006	A. Secilla	BIO-4478	-
	Spain: Vizcaya, Elantxobe (43º23’59’’N; 2º37’54’’W)	7.x.2006	A. Secilla	BIO-4477	-
	Spain: Vizcaya, S. Juan Gaztelugatxe (43º26’41’’N; 2º46’41’’W)	15.iv.2007	A. Secilla	SANT-Algae 21651	JQ653296
	Spain: Asturias, La Franca (43º23’39’’N; 4º34'18''W)	23.iii.2011	I. Bárbara & P. Díaz	SANT-Algae 25600	-
	Spain: Lugo, San Ciprián (43º42’02’’N; 7º27’25’’W)	30.ix.2004	P. Díaz & I. Bárbara	SANT-Algae 26259	-
	Spain: A Coruña, Cambre (43º18’00’’N; 8º46’04’’W)	12.i.2009	P. Díaz & I. Bárbara	SANT-Algae 21623	JQ653297
	Spain: Cádiz, Cala Encendida (36º18’40’’N; 6º09’12’’W)	18.ii.2011	I. Bárbara & P. Díaz	SANT-Algae 26644	-
Polysiphonia brodiei erect forms	Spain: A Coruña, Sada (43º23’18’’N; 8º16’49’’W)	17.iv.2009	J. Cremades	SANT-Algae 22808	JQ653295
Polysiphonia elongata	Spain: Pontevedra, Tragove (42º31’35’’N; 8º49’39’’W)	30.iv.2010	J. Cremades	SANT-Algae 24097	JQ653299
Polysiphonia foetidissima	France: Biarritz (43º29’03’’ N; 1º33’46’’ W)	19.iii.2011	P. Díaz & I. Bárbara	SANT-Algae 25433	JQ653294
	Spain: Vizcaya, Laida (43º24’28’’N; 2º40’54’’W)	31.iii.2006	P. Díaz	SANT-Algae 20286	-
	Spain: Vizcaya, La Arena (43º21’16’’N; 3º06’53’’W)	07.ix.2006	P. Díaz	SANT-Algae 20971	-
	Spain, Vizcaya, La Arena (43º21’16’’N; 3º06’53’’W)	22.iii.2011	P. Díaz, I. Bárbara & A. Secilla	SANT-Algae 25638, 25657	-
	Spain, Vizcaya, La Arena (43º21’16’’N; 3º06’53’’W)	22.iii.2011	P. Díaz, I. Bárbara & A. Secilla	SANT-Algae 25644	JQ653290
	Spain: Santander, Langre (43º28’37’’N; 3º41’37’’W)	06.xi.2010	P. Díaz	SANT-Algae 24620, 24628	-
	Spain: Santander, Virgen del Mar (43º28’40’’N; 3º52’31’’W)	07.xi.2010	P. Díaz	SANT-Algae 24648	JQ653285
	Spain: Asturias, La Franca (43º23’39’’N; 4º34'18''W)	23.iii.2011	I. Bárbara & P. Díaz	SANT-Algae 25611	JQ653291
	Spain: Asturias, Niembro (43º26’33’’N, 4º50’20’’W)	08.x.2006	P. Díaz & C. Peteiro	SANT-Algae 21644	-
	Spain: Asturias, Niembro (43º26’33’’N, 4º50’20’’W)	28.v.2010	P. Díaz & I. Bárbara	SANT-Algae 24162	-
	Spain: Asturias, Sarello (43º33’27’’N; 6º58’39’’W)	28.vii.2010	P. Díaz	SANT-Algae 25166	-
	Spain: Lugo, Catedrais (43º33’16’’N; 7º09’16’’W)	04.xi.2002	P. Díaz, I. Bárbara & T. Novo	SANT-Algae 16525	-
	Spain: Lugo, Catedrais (43º33’16’’N; 7º09’16’’W)	20.ix.2005	P. Díaz	SANT-Algae 16577	-
	Spain: Lugo, Catedrais (43º33’16’’N; 7º09’16’’W)	01.iii.2006	P. Díaz & V. Lagos	SANT-Algae 21622	JQ653292
	Spain: Lugo, Catedrais (43º33’16’’N; 7º09’16’’W)	01.iii.2006	P. Díaz & V. Lagos	SANT-Algae 21650	-
	Spain: Lugo, Catedrais (43º33’16’’N; 7º09’16’’W)	15.vii.2010	P. Díaz & I. Bárbara	SANT-Algae 24421	JQ653286
	Spain: Lugo, Castro (43º33’47’’N; 7º10’35’’W)	24.iv.2009	P. Díaz & I. Bárbara	SANT-Algae 24656	-
	Spain: Lugo, Llas (43º34’47’’N; 7º15’27’’W)	14.vii.2010	P. Díaz	SANT-Algae 24477	-
	Spain: Lugo, Llas (43º34’47’’N; 7º15’27’’W)	19.iv.2011	P. Díaz, A. Secilla & V. Peña	SANT-Algae 27332	JQ653284
	Spain: Lugo, Peinzás (43º35’09’’N; 7º16’13’’W)	15.vii.2010	P. Díaz & I. Bárbara	SANT-Algae 24412	JQ653289
	Spain: Lugo, Peinzás (43º35’09’’N; 7º16’13’’W)	19.iv.2011	P. Díaz, A. Secilla & V. Peña	SANT-Algae 26245	-
	Spain: Lugo, San Román (43º43’17’’N; 7º37’39’’W)	14.xi.2008	P. Díaz & I. Bárbara	SANT-Algae 21647	-
	Spain: A Coruña: Xilloe (43º44’41’’N; 7º39’02’’W)	10.ix.2002	P. Díaz & I. Bárbara	SANT-Algae 24910	-
	Spain: A Coruña, Barrañán (43º18’44’’N; 8º33’22’’W)	18.x.2002	P. Díaz & I. Bárbara	SANT-Algae 21645	-
	Spain: A Coruña, Barrañán (43º18’44’’N; 8º33’22’’W)	16.vi.2003	P. Díaz & I. Bárbara	SANT-Algae 21646	-
	Spain: A Coruña, Cambre (43º18’00’’N; 8º46'04''W)	14.iv.2008	P. Díaz & I. Bárbara	SANT-Algae 21648	-
	Spain: A Coruña, Lago (43º05’53’’N; 9º11’50’’W)	29.ix.2003	P. Díaz & I. Bárbara	SANT-Algae 17963	-
	Spain: A Coruña, Arnela (42º42’35’’N; 9º00’47’’W)	01.viii.2011	P. Díaz & V. García	SANT-Algae 26444	-
	Spain: A Coruña, Basoñas (42º37’22’’N; 9º03’10’’W)	18.iv.2011	P. Díaz & I. Bárbara	SANT-Algae 26267	JQ653288
	Spain: A Coruña, Basoñas (42º37’22’’N; 9º03’10’’W)	02.viii.2011	P. Díaz & V. García	SANT-Algae 26441	-
	Spain: A Coruña, El Puntal (42º36’37’’N; 8º53’40’’W)	04.ii.2005	V. Peña & I. Bárbara	SANT-Algae 21649	-
	Portugal: Estremadura, Baleal (39º22’25’’N, 9º19’56’’W)	14.vi.2010	P. Díaz & I. Bárbara	SANT-Algae 24255	JQ653287
	Portugal: Baixo Alentejo, Queimado (37º49’34’’N; 8º47’34’’W)	24.iv.2005	P. Díaz	SANT-Algae 24658	-
	Portugal: Baixo Alentejo, Almograve (37º39’54’’N; 8º48’04’’W)	25.v.2005	P. Díaz	SANT-Algae 24657	-
	Portugal: Algarve, Ingrina (37º02’46’’N; 8º52’43’’W)	09.v.2005	P. Díaz	SANT-Algae 24659	-
	Portugal: Algarve, Ingrina (37º02’46’’N; 8º52’43’’W)	20.ii.2011	P. Díaz & I. Bárbara	SANT-Algae 25470	JQ653293
	Portugal: Algarve, Armaçao de Pera (37º06’04’’N; 8º22’17’’W)	17.x.2005	P. Díaz	SANT-Algae 24840	-
	Portugal: Algarve, Olhos d'Agua (37º05’20’’N; 8º11’27’’W)	20.ii.2011	P. Díaz & I. Bárbara	SANT-Algae 26464	JQ653283
	Spain: Cádiz, Puerto de Santa María (36º34’48’’N; 6º15’51’’W)	16.xi.2005	P. Díaz	SANT-Algae 26069	-
	Spain: Cádiz, Punta Plata (36º06’28’’N; 5º49’41’’W)	19.i.2011	P. Díaz & I. Bárbara	SANT-Algae 26517	-
Polysiphonia schneideri	Spain: Cádiz, Santa Leocadia (36º25’15’’ N, 6º13’02’’ W)	9.vi.2010	M. Fernández-Lora	SANT-Algae 26813	-
	Spain: Cádiz, Santa Leocadia (36º25’15’’ N, 6º13’02’’ W)	23.ii.2011	M. Fernández-Lora	SANT-Algae 26065	JQ653298
	Spain: Cádiz, Barbate (36º11’03’’ N, 5º55’39’’ W)	18.ii.2011	P. Díaz & I. Bárbara	SANT-Algae 26572	-

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