Alexandrium Halim 1960

compiled by Mona Hoppenrath & Marina Montresor

last update: 19 December 2023

Species names in bold = validly published names
Species names in normal font = invalid names

 

Alexandrium Halim 1960, Vie Milieu 11: 102.
Type: Alexandrium minutum Halim 1960, Vie Milieu 11: 102-104, figs 1a-j.

 

Alexandrium acatenella (Whedon et Kofoid) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 19.
Basionym: Gonyaulax acatenella Whedon et Kofoid 1936, University of California Publications in

                  Zoology 41(4): 31, figs 8-13, 15.
Synonyms: Protogonyaulax acatenella (Whedon et Kofoid) Taylor 1979, in DL Taylor & HH Seliger
                  (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 51.

Gessnerium acatenellum (Whedon et Kofoid) Loeblich III et Loeblich 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 44.

 

Alexandrium affine (Inoue et Fukuyo) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 38.
Basionym: Protogonyaulax affinis Inoue et Fukuyo in Fukuyo et al. 1985, in Anderson et al. (eds) Toxic
                 dinoflagellates, Elsevier: 30, figs 1E, 3A-C, 24-29.

Synonyms: Alexandrium fukuyoi Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier:
                   fig. 6 (Balech 1995: nomen nudum).

Alexandrium andersonii Balech 1990, Helgoländer Meeresunters. 44: 394-395, figs 34-50.

Alexandrium australiense Sh.Murray 2014, in John et al. 2014, Protist 165: 797, fig. 8.

genetically-defined Group V of the species complex*

Synonym: Alexandrium tamarense (Lebour) Balech sensu Murray et al. (2012), 98, figs 1, 2.

Alexandrium balechii (Steidinger) Balech 1995, The genus Alexandrium Halim (Dinoflagellata), Sherkin Island Marine Station: 70, pl. 14 figs 1-23.

Basionym: Gonyaulax balechii Steidinger 1971, Phycologia 10(2/3): 183, 185, fig. 1.

Synonyms: Gessnerium balechii (Steidinger) Loeblich III et Loeblich 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 44.

Pyrodinium balechii (Steidinger) Taylor 1976, Bibliotheca Botanica 132: 110.

Alexandrium camurascutulum MacKenzie et Todd 2002, Harmful Algae 1(3): 296, figs 1-12.

Alexandrium catenella (Whedon et Kofoid) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37.

genetically-defined Group I of the species complex*

Basionym:  Gonyaulax catenella Whedon et Kofoid 1936, University of California Publications in Zoology 41(4): 25, figs 1-7, 14.

Synonyms: Protogonyaulax catenella (Whedon et Kofoid) Taylor 1979, in DL Taylor & HH

Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 51.

Gessnerium catenellum (Whedon et Kofoid) Loeblich III et Loeblich 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 44.

Alexandrium fundyense Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 18.

Alexandrium cohorticula (Balech) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 5.

Basionym: Gonyaulax cohorticula Balech 1967, Rev. Mus. Argent. Cienc. Nat. “Bernardino

Rivadavia”, Hidrobiologia 2(3): 111, pl. 6 figs 117-119, pl. 7 figs 120-122.

Synonyms: Protogonyaulax cohorticula (Balech) Taylor 1979, in DL Taylor & HH

Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 51.

Gessnerium cohorticulum (Balech) Loeblich III et Loeblich 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 44.

Alexandrium compressum (Fukuyo, Yoshida et Inoue) Balech 1995, The genus Alexandrium Halim (Dinoflagellata), Sherkin Island Marine Station: 51, pl. 12 figs 1-9.

Basionym: Protogonyaulax compressa Fukuyo, Yoshida et Inoue 1985, in Anderson et al.

(eds) Toxic dinoflagellates, Elsevier: 30, 32, figs 3D-F, 30-33.

 

Alexandrium concavum (Gaarder) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37-38.

emend. Nguyen-Ngoc et Larsen 2004, in Larsen & Nguyen-Ngoc 2004, Opera Botanica 140: 86-87.

Basionym: Goniodoma concavum Gaarder 1954, Report on the Scientific Results of the “Michael Sars” north Atlantic deep-sea expedition 1910, vol. II no 3: 27-28, fig. 32.

non:  Gonyaulax concava (Gaarder) Balech 1967, figs 108-116, auct. non Balech 1967

nec:   Alexandrium concavum (Gaarder) Balech 1985 , fig. 17, auct. non Balech 199.

 

Alexandrium diversaporum S.A.Murray, Hoppenrath et al. 2014, Harmful Algae 31: 56, figs 1-7.

 

Alexandrium foedum Balech 1990, Helgoländer Meeresunters. 44: 392-394, figs 19-33.

 

Alexandrium fragae S.Branco et M.Menezes 2020, in Branco et al. 2020, Harmful Algae 95: 3, figs 2-7. [13 May 2020]

Alexandrium fraterculus (Balech) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 4.

Basionym: Gonyaulax fratercula Balech 1964, Bol. Inst. Biol. Mar. Mar del Plata 4: 31-34, pl. 4 figs 47-58.

Synonyms: Gessnerium fraterculum (Balech) Loeblich III et Loeblich 1979, in DL Taylor & HH Seliger

(eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 44.

Protogonyaulax fratercula (Balech) Taylor 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 51.

 

Alexandrium gaarderae Nguyen-Ngoc et Larsen nom. nov. in Larsen & Nguyen-Ngoc 2004, Opera Botanica 140: 90, 92, pl. 6 figs 1-8.

Synonyms: Gonyaulax concava (Gaarder) Balech 1967, Rev. Mus. Argent. Cienc. Nat. “Bernardino Rivadavia”, Hidrobiologia 2(3): 108-111, pl. 6 figs 108-116.

Alexandrium concavum (Gaarder) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37-38, fig. 17.

 

Alexandrium globosum Nguyen-Ngoc et Larsen in Larsen & Nguyen-Ngoc 2004, Opera Botanica 140: 93, pl. 7 figs 1-8.

 

Alexandrium hiranoi Kita et Fukuyo 1988, Bulletin of Plankton Society of Japan 35(1): 2-4, Figs 1a-f.

Synonyms: Goniodoma pseudogonyaulax Biecheler sensu Silva 1965, Notas e Estudos do Instituto de Biologia Marítima 30: 11-12, figs 1-4.

Goniodoma pseudogoniaulax Biecheler sensu Kita et al. 1985, Bulletin of Marine Science 37(2): 646,

fig. 3.

non:  Goniodoma pseudogonyaulax Biecheler 1952, Bull. Biol. France Belgique Suppl. 36:   55-59, figs 30-32.

          Alexandrium pseudogonyaulax (Biecheler) Horiguchi 1983, PhD thesis: 64, fig. 10.

 

Alexandrium insuetum Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 1.

 

Alexandrium kutnerae (Balech) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 8.

Basionym: Gonyaulax kutnerae Balech 1979, Bol. Inst. Oceanogr., S Paulo 28(2): 61-62, pl. 3 figs 1-11.

 

Alexandrium leei Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 9.

 

Alexandrium limii S.T.Teng, Tillmann, N.Abdullah, S.Nagai et Leaw 2023, in Abdullah et al. 2023, Harmful Algae 127, 102475: 6, figs 1-5. [online 1 July 2023]

 

Alexandrium margalefii Balech 1994, Trans. Am. Microsc. Soc. 113(2): 220, figs 12-15.

 

Alexandrium mediterraneum U.John 2014, in John et al. 2014, Protist 165: 795-796, fig. 7.

genetically-defined Group II of the species complex*

 

Alexandrium minutum Halim 1960, Vie Milieu 11: 102-104, figs 1a-j.

Synonyms: Pyrodinium minutum (Halim) Taylor 1976, Bibliotheca Botanica 132: 110.

Alexandrium ibericum Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 15.

Alexandrium angustitabulatum Taylor in Balech 1995, The genus Alexandrium Halim (Dinoflagellata), Sherkin Island Marine Station: 70, pl. 14 figs 1-23. (after Hansen et al. 2003, Harmful Algae 2: 317-335; Lilly et al. 2005, Harmful Algae 4: 1004-1020)

A proper re-definition of the species diagnosis is still missing.

Alexandrium lusitanicum Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 16. (after Lilly et al. 2005, Harmful Algae 4: 1004-1020; Anderson et al. 2012, Harmful Algae 14: 10-35, Table 1)

   A proper re-definition of the species diagnosis is still missing.

 

Alexandrium monilatum (Howell) Balech 1995, The genus Alexandrium Halim (Dinoflagellata), Sherkin Island Marine Station: 67, pl. 15 figs 1-16.

Basionym: Gonyaulax monilata Howell 1953, Trans. Am. Microsc. Soc. 72(2): 153-154, figs 1-5.

Synonyms: Gessnerium mochimaensis Halim 1967, Int. Revue ges. Hydrobiol. 52: 729, pl. 4 fig. 49, pl. 8 figs 101-108.

Gessnerium mochimaensis Halim 1969, Int. Revue ges. Hydrobiol. 54: 619.

Gessnerium monilata (Howell) Loeblich III 1970, Proc. N. Amer. Paleont. Conv. Part G: 903.

    Pyrodinium monilatum (Howell) Taylor 1976, Bibliotheca Botanica 132: 110.

 

Alexandrium ostenfeldii (Paulsen) Balech et Tangen 1985, Sarsia 70: 338.

Basionym: Goniodoma ostenfeldii Paulsen 1904, Medd. Komm. F. Havundersogelser, Ser. Plankton, I. Kobenhavn: 20, fig. 2.

Synonyms: Gonyaulax ostenfeldii (Paulsen) Paulsen 1949, Det Kongelige Danske Videnskabernes Selskab, Biologiske Skrifter 6(4): 62-63.

Heteraulacus ostenfeldii (Paulsen) Loeblich III 1970, Proc. N. Amer. Paleont. Conv. Part G: 904.

Gessnerium ostenfeldii (Paulsen) Loeblich III et Loeblich 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 44.

Triadinium ostenfeldii (Paulsen) Dodge 1981, Br. Phycol. J. 16: 279.

? Pyrodinium phoneus Woloszynska et Conrad 1939, Bull. Mus. Royal Hist. nat. Belgique 15: 1-5, figs 1-12.

Gonyaulax tamarensis var. globosa Braarud 1945, Avh. Utgitt. Nor. Vidensk. Akad. Oslo: 12-14, pl. 3 fig. 5B.

Gonyaulax globosa (Braarud) Balech 1971, Republica Argentina, Armada Argentina, Servicio de Hidrografia Naval 654: 29.

Protogonyaulax globosa (Braarud) Taylor 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 52.

Gonyaulax trygvei Parke 1976 in Parke & Dodge 1976, in Parke & Dixon, J. Mar. Biol. Ass. UK: 546.

Alexandrium peruvianum (Balech et B.R.Mendiola) Balech et Tangen 1985, Sarsia 70: 342. (after Kremp et al. 2014, J Phycol 50: 81-100)

A proper re-definition of the species diagnosis is still missing.

Basionym: Gonyaulax peruviana Balech et Mendiola 1977, Neotropica 23(69): 49-52, figs 1-12.

Gonyaulax dimorpha Biecheler 1952, Bull. Biol. France Belgique Suppl. 36: 50-54, figs 26-29. (after Kremp et al. 2014, J Phycol 50: 81-100)

A proper re-definition of the species diagnosis is still missing.

 

Alexandrium pacificum Litaker 2014, in John et al. 2014, Protist 165: 793-795, fig. 6.

genetically-defined Group IV of the species complex*

 

Alexandrium pohangense A.S.Lim et H.J.Jeong 2015, in Lim et al. 2015, Harmful Algae 46: 60, figs 3-7.

 

Alexandrium pseudogonyaulax (Biecheler) Horiguchi ex Yuki & Fukuyo 1992, Journal of Phycology 28: 395, 398.

Basionym: Goniodoma pseudogonyaulax Biecheler 1952, Bull. Biol. France Belgique Suppl. 36: 55-59, figs 30-32.

 

Alexandrium satoanum Yuki et Fukuyo 1992, Journal of Phycology 28: 396, fig. 1.

 

Alexandrium tamarense (M.Lebour) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 38.

emend. U.John in John et al. 2014, Protist 165: 792-793.

genetically-defined Group III of the species complex*

Basionym: Gonyaulax tamarensis Lebour 1925, Marine Biological Association UK, Plymouth: 95, pl. 14 figs 1a-d.

Synonyms:    Gessnerium tamarensis (Lebour) Loeblich III et Loeblich 1979, in DL Taylor & HH Seliger

eds),Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 44.

Protogonyaulax tamarensis (Lebour) F.J.R.Taylor 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: 51.

Gonyaulax tamarensis var. excavata Braarud 1945, Avh. Nor. Vidensk. Akad. Oslo: 10-12, 14, pl. 2 fig. 5A.

Gonyaulax excavata (Braarud) Balech 1971, Republica Argentina, Armada Argentina, Servicio de Hidrografia Naval 654: 28 (not figs 119-124).

Protogonyaulax excavata (Braarud) F.J.R.Taylor 1979, in DL Taylor & HH Seliger (eds), Toxic dinoflagellate blooms, Elsevier, Developments in Marine Biology 1: ??.

Alexandrium excavatum (Braarud) Balech et Tangen 1985, Sarsia 70: 334.

 

Alexandrium tamiyavanichii Balech 1994, Trans. Am. Microsc. Soc. 113(2): 217, 219, figs 1-6.

Synonym: Protogonyaulax cohorticula (Balech) Taylor sensu Kodama et al. 1988, Toxicon 26(8): 707-712.

non:  Alexandrium cohorticula (Balech) Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37.

Basionym: Gonyaulax cohorticula Balech 1967, Rev. Mus. Arg. Cienc. Nat. “Bernardino Rivadavia”, Hidrobiologia 2(3): 111, pl. 6 figs 117-119, pl. 7 figs 120-122.

 

Alexandrium tamutum Montresor, Beran et John in Montresor et al. 2004, Journal of Phycology 40: 402, figs 1-4.

 

Alexandrium taylori Balech 1994, Trans. Am. Microsc. Soc. 113(2): 219, figs 7-11.

 

Alexandrium tropicale nom. nov. Balech 1985, in Anderson et al. (eds) Toxic dinoflagellates, Elsevier: 37, fig. 7.
Synonym: Gonyaulax excavata (Braarud) sensu Balech 1971, Republica Argentina, Armada Argentina,
                 Servicio de Hidrografia Naval 654: 28, pl. 7 figs 119-124.
non:  Gonyaulax tamarensis var. excavata Braarud 1945, Avh. Nor. Vidensk. Akad. Oslo: 10-12, 14, pl. 2 fig. 5A.

 

*Note on Alexandrium tamarense species complex
The Alexandrium tamarense species complex has been reviewed by John et al. (2014a). Based on morphological data, phylogenies of multiple regions of the rDNA operon, and toxicity data, the authors described three new species, A. mediterraneum (genetically-defined Group II), A. australiense (Group V), A. pacificum (Group IV), emended the diagnosis of A. tamarense (Group III), and used the name of A. fundyense for the isolates belonging to Group I.

 

The authors also proposed to the Nomenclature Committee for Algae of the International Code of Nomenclature for algae, fungi, and plants to reject the name Alexandrium catenella based on Art. 56 of the Code (John et al. 2014b). The species was originally described from material collected off the California and Oregon coasts but cells ‘morphologically’ identified as A. catenella from this region fall within the genetically-defined Group I that also includes strains isolated  from the eastern coast of the US that were traditionally identified as A. fundyense. The proposal by John et al. (2014b) was followed by a counterproposal by Fraga et al. (2015) who posed arguments against the rejection of the name A. catenella. The Nomenclature Committee for Algae took the decision to not reject the name Alexandrium catenella (Gonyaulax catenella). This implies that the name A. fundyense (Type I) has to be considered a synonym of A. catenella (Prud'homme van Reine 2017). For a practical guide to this new nomenclature see Litaker et al. (2018).

 

John, U., Litaker, W., Montresor, M., Murray, S., Brosnahan, M. & Anderson, D.M. (2014a). Formal revision of the Alexandrium tamarense species complex (Dinophyceae) taxonomy: the introduction of five species with emphasis on molecular-based (rDNA) classification. Protist 165:779–804.

John, U., Litaker, W., Montresor, M., Murray, S., Brosnahan, M.L. & Anderson, D.M. (2014b). Proposal to reject the name Gonyaulax catenella (Alexandrium catenella) (Dinophyceae). Taxon 63, 932-933.

 

Fraga, S., Sampedro, N., Larsen, J., Moestrup, Ø. & Calado, A.J. (2015). Arguments against the proposal 2302 by John & al. to reject the name Gonyaulax catenella (Alexandrium catenella). Taxon 64, 634-635.

 

Prud'homme van Reine, W.F. (2017). Report of the Nomenclature Committee for Algae: 15. Taxon 66, 191-192.

 

Litaker, W., Fraga, S., Montresor, M., Brosnahan, M., Hoppenrath, M., Murray, S., Wolny J., John, U., Sampedro N., Larsen, J. & Calado, A.J. (2018) A practical guide to new nomenclature for species within the “Alexandrium tamarense species complex”. Harmful Algae News 61: 13-15.