Nova Hedwigia 56 1-2 173-178 Stuttgart, Februar 1993

Kolkwitziella Lindemann 1919 and Preperidinium Mangin 1913: correct genera names in the DiplopsaHs-group (Dinophyceae)

by

Malte Elbrachter* Biologische Anstalt Helgoland, Wattenmeerstation Sylt

Hafenstr. 43, D-2282 List/ Sylt , Germany

Elbrachter, M. (1993): Kolkwitziella Lindemann 1919 and Preperidinium Mangin 1913 : correct genera names in the Diplopsalis-group (Dinophyceae). -Nova Hedwigia 56: 173-178 .

Abstract: Kolkwitziella salebrosa Lindemann 1919 represents the cyst of the organism earl ier described as Glenodinium acutum and later asigned to the genera Diplopsalis and finally Entzia or Apsteinia. The genera names Entzia Lebour 1922 and Apsteinia Abe 1981 are later synonyms of Kolkwitziella Linde­mann . The combination Kolkwitziella acuta (Apstein) comb. nov. is introduced . The family name Di­plopsidaceae Matsuoka 1988 is a latter synonym of Kolkwitziellaceae Lindemann 1928.

Preperidinium Mangin 1913 is the correct name on genus level for the organism first described as Diplop­salis lenticula Bergh. fo. minor Paulsen 1907, if regarded distinct on genus level from Diplopsalis. Diplo­peltopsis Pavillard 1913 and Zygabikodinium Loeblich et Loeblich 1970 are synonyms based on the same type. The species now placed into the genus Zygabikodinium are transferred to Preperidinium. Kolk­witziella and Preperidinium both are legitime names in the plant and in the animal kingdom, bringing into accordance nomenclature in both kingdoms.

Kolkwitziella Lindemann 1919 Synonyms: Entzia Lebour 1922, non Daday 1883 (Foraminifera); Apsteinia Abe 1981 , non Schmeil (Crustaceae) nee Metcalf (Tunicata) .

Apstein (1896) described a freshwater dinoflagellate which he called Glenodinium acutum. This organism is quite different from the type species of Glenodinium Ehrenb. 1836, G. cinctum Ehrenb. 1836 (synonym: Sphaerodinium polonicum Wolosz. 1916) and consequently has been transferred to the genus Diplopsalis Bergh 1881 by Entz 1904. As this species has four apical plates in contrast to the type spe­cies of Diplopsalis, Diplopsalis lenticula Bergh 1881, Lebour (1922) removed it from Diplopsalis, creating a new genus Entzia Lebour 1922. This name is preoccupied in the zoological nomenclature by Entzia Daday 1883 (Foraminifera) . Therefore Abe (1981) replaced it by Apsteinia Abe 1981 which is also preoccupied by Apsteinia Schmeil (Crustacea) and Apsteinia Metcalf (Tunicata).

*Member of the "Taxonomische Arbeitsgruppe an der BAH".

173

0029-5035/ 93/ 0056-0173 $1 .50 © 1993 J. Cramer in der Gebrtider Bornt raeger

Veriagsbuchhandlung, D· IOOO Berlin · D-7000 Stuttgart

Lindemann (1919) described an unusual dinoflagellate, Kolkwitziella salebrosa Lin­demann 1919, and later a variety Kolkwitziella salebrosa var. gibbera (Lindemann 1924) which he later raised to species level (Lindemann 1928). Sebestyen (1935) showed that the cyst of Diplopsalis acuta (Apstein) Entz corresponds to Kolkwitziel­la salebrosa Lindemann and that the variety K. salebrosa var. gibbera is apparently not distinct from the type. If this species is regarded as distinct on genus level from Diplopsalis Bergh, as currently accepted, the correct genus name has to be Kolkwit­ziella Lindemann, and the species name Kolkwitziella acuta (Apstein) comb. nov. (see below). In any case Kolkwitziella Lindemann 1919 has priority over Entzia Le­hour 1922 and therefore also over Apsteinia Abe 1981. This brings botanical and zoological nomenclature into accordance.

Lindemann (1928) created the family Kolkwitziellaceae for his genus Kolkwitziella. If the Diplopsalis-group is regarded to represent an own family, the correct family name has to be Kolkwitziellaceae Lindemann. Diplopsidaceae Matsuoka 1988 is a later synonym of Kolkwitziellaceae, as Matsuoka (1988) included in his family the genus Entzia Lebour, a later synonym of Kolkwitziella, the type of the Kolwitziella­ceae. Silva (1980) included into the family Ptychodiscaceae (Schiitt) Lemmermann 1899 the family Kolkwitziellaceae apparently regarding Kolkwitziella as a vegetative stage and not as the cyst of Entzia acuta, which he placed in the Index Nominorum Genericorum (ING; Farr et al. 1979) into the Dinosphaeraceae, a family accepted by Silva (1980) as well. If Dinosphaera Kofoid et Michener 1912 and Kolkwitziella should be placed into the same family, this would be, in my opinion, the Peridinia­ceae Ehrenberg 1831 .

The genera of the Diplopsalis-group easily fit into the order Peridiniales Haeckel 1894, therefore the order Kolkwitziellales Lindemann 1928 is regarded as a later synonym of Peridiniales Haeckel.

Ko1kwitziella acuta (Apstein 1896) comb. nov.

Basionym : Glenodinium acutum Apstein 1896; Apstein , C.: Das Siil3wasserp1ankton . Methode und Re­su1tate der quantitativen Untersuchungen . Lipsius & Tisch1er , Kie1 , Leipzig, p. 152, fig. 54.

Synonyms: Diplopsalis acuta (Apstein) Entz 1904; Peridinium latum Pau1sen 1908; Kolkwitziella salebro­sa Lindemann 1919; Entzia acuta (Apstein) Lebour 1922; Kolkwitziella salebrosa var. gibbera Linde­mann 1924; Kolk witziella gibbera (Lindemann) Lindemann 1928; Apsteinia acuta Abe 1981 (nom . nudum) .

Preperidinium Mangin 1913

Synonyms : Diplopeltopsis Pavillard 1913, non Hennings ex von Hoehn1 1911 ; Zygabikodinium Loeb1ich et Loeb1ich 1970.

When describing briefly the genus, Mangin (1913) included two species: Preperidini­um assymmetricum (Mangin) Mangin and Preperidinium paulseni (Mangin) Mangin without designating a type species. Nevertheless, the genus name is validly published.

174

fa salebrosa Lin­•era (Lindemann )ebestyen (1935) s to Kolkwitziel­?ra is apparently genus level from ts to be Kolkwit­ein) comb. nov. over Entzia Le­

~s botanical and

1s Kolkwitziella. te correct family tsuoka 1988 is a n his family the the Kolwitziella­t) Lemmermann 'a as a vegetative :lex Nominorum family accepted nd Kolkwitziella n, the Peridinia-

jiniales Haeckel arded as a later

n. Methode und Re-2, fig. 54.

·atkwitziella salebro­var. gibbera Linde­ta Abe 1981 (nom.

>ikodinium Loeblich

:ies: Preperidini­\'langin) Mangin 1ame is validly

The basionym of Preperidinium assymmetricum is Peridiniopsis asymetrica Mangin 1911, the correct spelling should be therefore Preperidinium asymetricum. It has been later taken into synonymy with Diplopelta bomba Stein ex Joergensen.

Preperidinium paulseni has as basionym Peridinium paulseni Mangin 1911 (C.R. Acad. Sci. 153: 648), which is a nomen novum for Peridinium lenticulatum Mangin 1911 (C.R. Acad. Sci. 153: 27; non Faure-Fremiet 1908). In all 3 publications Man­gin (1911a, 1911b, 1913) explicitely states that the organism he is dealing with is identical to the species Paulsen (1907) has described as Diplopsalis lenticula fo. mi­nor. This is corroborated by the fact that Mangin investigated also material from the type locality of that taxon, which Paulsen had sent to him. Also the figures (Mangin 1911a, figs 3, 5; Mangin 1911b, figs I-III, V; Mangin 1913 fig. 12, I-III, V) clearly show, that he was dealing with the species Paulsen had described. Thus Preperidinium paulseni Mangin 1913 is a validly published name for the species ori­ginally described as Diplopsalis lenticula fo. minor Paulsen 1907.

Loeblich & Loeblich (1966) designated as type species P. paulsenii (Pavillard) Man­gin and gave as basionym Preidinium (sic!) paulsenii Pavillard 1909. Peridinium paulsenii Pavillard is transferred by Balech (1974) to the genus Protoperidinium Bergh and is quite different from the species, Mangin was dealing with. Thus this typification has to be rejected. Silva in Farr et al. (1979) cite as lectotype "Preperidi­nium paulsenii" Mangin ("paulseni"). According to article 64 of ICBN (Greuter et al. 1988) the epithet "paulseni" has to be rejected and to be replaced by the oldest legitimate epithet available, see below.

There are two other genera with the type species based on Diplopsalils lenticula Bergh fo. minor Paulsen, that is Diplopeltopsis Pavillard 1913 and Zygabikodinium Loeblich et Loeblich 1970.

Pavillard (20. July 1913) created the genus Diplopeltopsis Pavillard with the type species Diplopeltopsis minor (Paulsen) Pavillard. As synonyms he gives: Diplopsalis lenticula forma minor and Peridinium pau/seni Mangin. It thus has the same type as Preperidinium Mangin.

Pavillard handled the "peridiniens" always as plants, in particular in his publica­tions of that time (e.g. Pavillard 1910, 1916). Explicitly, he treated Diplopeltopsis minor with its synonym Peridinium pau/seni Mangin, as plant (Pavillard 1916). The­re is no doubt, that he regarded Diplopeltopsis also in the original publication as plant. The name Diplopeltopsis Pavillard 1913 is a later homonym of Diplopeltopsis Hennings ex von Hoehnel 1911 (Lichenes) and therefore illegitime. Consequently, it has been replaced by Zygabikodinium (Loeblich & Loeblich 1970). As type species of the new genus they designate Zygabikodinium lenticulatum Loeblich et Loeblich and cite as synonyms Peridinium lenticulatum Mangin 1911a, p. 30, 32; Diplopsalis lenticula Bergh f. mjlnor Paulsen 1907 and Peridinium paulsenii Mangin 1911.

Preperidinium Mangin 1913 (exact date not stated, no further information avail­able) has been published as plant and is available in the botanical and zoological nomenclature. Thus Preperidinium Mangin 191J is the correct genus name for the organism Paulsen (1907) described as Diplopsalis lenticula Bergh fo. minor, regard­less it is handled as plant or animal.

175

Finally, the question of the correct species name has to be discussed . Introducing the combination Preperidinium paulseni Mangin (1913) used as basionym the illegit­imate name paulseni as it is a later homonym of Peridinium paulseni Pavillard 1909 (ICBN art. 64.1). Therefore this combination has to be rejected. Four species names unequivocally have been published for the species under duscussion. These are:

lenticulatum (as Peridinium lenticulatum Mangin 1911 , non Faure Fremiet 1908; and Zygabikodinium lenticulatum Loeblich et Loeblich 1970), paulseni (as Peridinium paulseni Mangin 1911, non Pavillard 1909 and Preperidini­um paulseni Mangin 1913), meunierii (as Peridinium meunierii Pavillard 1912, nom. nov. proP. paulseni Man­gin 1911), minor (as Diplopeltopsis minor Pavillard 1913).

In addition, Pavillard (1912) stated, that Diplopsalis sphaerica Meunier (Meunier 1910) is identical to Peridinium paulseni Mangin 1911, a view followed by Lebour 1922. Meunier himself (1919) regarded his Diplopsalis sphaerica as Diplopsalis lenti­cula fo. sphaerica. But he lumped together some other species of the Diplopsalis­group like Peridiniopsis asymetricum and others. His fig. 8 on PI. 29, which repre­sents D. lenticula fo. sphaerica clearly shows only 6 precingular plates, in accord­ance with his text on p. 64 (Meunier 1919). Also the original figure of D. sphaerica (Meunier 1910, PI. 4, fig. 10) clearly shows only 6 precingular plates . This is in con­tradiction to the descriptions and figures of Mangin (1911a, 1911b, 1913), of Pavil­lard (1913) and of Lebour (1922). Thus it is most unlikely that Diplopsalis sphaerica is conspecific with Preperidinium paulseni Mangin and Diplopeltopsis minor Pavil­lard. At the present state of knowledge it is advisable not to include Diplopsalis sphaerica Meunier 1910 into the genus Preperidinium Mangin and to exclude it from the discussion of the name of the type species of this genus.

According to Art. 6.4, 64.1 and 72.1 of ICBN only legitimate names can be used for a taxon. A name rejected as it was illegitimate by its publication has to be re­placed by the name that has priority in the rank concerned (ICBN, art. 72.1) . If we exclude the epithet sphaerica - see above - the earliest legitimate combination available at species rank is Peridinium meunierii. The older epithets lenticulatum (Mangin 1911) and paulseni Mangin 1911 both are illegimate as they are junior ho­monyms of validly published species in the genus Peridinium (ICBN art. 64.1). Thus the type species of the genus has to get the name:

Preperidinium meunierii (Pavillard) Elbrachter, comb. nov. (lectotype)

Basionym: Peridinium meunieriiPavillard 1912, C.R. Sean . Acad. Se. 155: 177; non Peridinium meunie-

rii Peters 1930. f f"-"iJ!<J(Z/IJ Synonyms: Diplopeltopsis mino1P'avillard 1913; Diplopsalis lenticula Bergh. f. minor Paulsen 1907, p. 9, fig. 9; Diplopsalis minor (Paulsen) Lindemann 1927; Diplopsalis minor (Paulsen) Paulsen 1931; Di­plopsalis minor (Paulsen) Margalef 1973; Diplopsalopsis minor (Paulsen) A be 1981 ; Dubridinium cape­ratum Reid 1977; Glenodinium lenticula (Bergh) Schiller f. minor Paulsen; Peridiniopsis minor (Paulsen) Starmach 1974; Peridinium lenticulatum Mangin 1911 , non Faure-Fremiet 1908; Peridinium paulseni Mangin 1911, non Pavillard 1909; Zygabikodinium lenticulatum Loeblich et Loeblich 1970.

176

la salebrosa Lin­•era (Lindemann )ebestyen (1935) .s to Kolkwitziel­era is apparently genus level from ts to be Kolkwit­ein) comb. nov. over Entzia Le­;s botanical and

s Kolkwitziella. ~ correct family suoka 1988 is a 1 his family the he Kolwitziella-1 Lemmermann · as a vegetative ex Nominorum amily accepted j Kolkwitziella . the Peridinia-

niales Haeckel ded as a later

Vlethode und Re­'ig. 54.

witziella salebro­. gibbera Linde-1\be 1981 (nom.

tinium Loeblich

Preperidini­tgin) Mangin e is validly

As Zygabikodinium Loeblich et Loeblich is a later synonym of Preperidinium Man­gin, the species so far included into Zygabikodinium have to be transferred to Preperidinium.

Preperidinium granulosum (Balech) Elbrachter, comb. nov.

Basionym: Diplopeltopsis granulosa Balech 1958, Physis 21(60), p. 84, PI. 2, figs. 32-44; synonym Zyga­bikodinium granulosa (Balech) Loeblich Ill, 1970.

Preperidinium per la turn (Balech) Elbrachter, comb. nov.

Basionym: Diplopeltopsis perlata Balech 1971, Revta M us . argent. Cienc. nat. "B. Rivadavia", Hydro­bioi. 3(1), p. 81, PI. 13, figs. 198-207; synonym Zygabikodinium perlatum (Balech) Elbrachter 1975.

Preperidinium pseudooblea (Elbrachter) comb. nov . Basionym: Zygabikodinium pseudooblea Elbrachter 1975, Kieler Meeresforsch . 31(1), p. 62, fig. 5.

Acknowledgements

Partly supported by the Deutsche Forschungsgemeinschaft (DFG El 59/4). Essentiel help I got from A. Sournia, Paris - merci beaucoup!

References

ABE, T. (1981): Studies on the family Peridiniidae. An unfin ished monograph of the armoured dino­flagellata. - Acad. Scient . Book Inc. Tokyo, 412 pp.

APSTEIN, C. (1896): Das Siil3wasserplankton. Methode und Resultate der quantitativen Untersuchung. - Lipsius & Tischler, Kiel, Leipzig.

BALECH, E. (1958): Plancton de la campana antarctica argentina 1954-1955. - Physis 21: 75-108.

BALECH, E. (1971): Microplancton de la campana oceanografica, Productividad Ill.- Revta Mus. Ar­gent. Cienc. Hydrobiol. Ill, 1: 1-202.

BALECH, E. (1974): El genero Protoperidinium Bergh 1881 (Peridinium Ehr., 1831 partim). - Revta Mus. Argent. Cienc . Hydrobiol. IV(!): 1-79.

ELBRACHTER, M. (1975): Taxonomical notes on North Sea dinoflagellates I. - Kieler Meeresf. 31: 58-64 .

ENTZ, G. (1904): Beitrage zur Kenntnis der Peridinee. - Math. naturw. Ber. Ung. 20: 96-144.

FARR, E.R., J.A. LEUSSINK & F.A. STAFLEU (1979): Index Nominorum Genericorum (Plantarum). - Regnum Vegetab. 100-102.

GREUTER, W. (ed.) (1988): International Code of Botanical Nomenclature. - Regn. Vegetabile 118: 328 pp.

LEBOUR, M. V. (1922): Plymouth Peridinians I. Diplopsalis lenticula and its relatives. - J. mar. bioi. Ass. U.K. 12: 795-812. •

LINDEMANN, E. (1919): Untersuchungen iiber Stil3wasserperidineen und ihre Variationsformen . -Arch. Protistenk. 319: 209-262.

LINDEMANN, E. (1924): Mitteilungen tiber nicht geniigend bekannte Peridineen. -Arch. Protistenk. 47: 431-439.

177

LINDEMANN, E. (1928): Abteilung Peridineae (Dinoflagellatae).- pp. 3-104 in: Engler, A. & Prantl, k. (eds.): Die natiirlichen Pflanzenfamilien .. . 2. Aufl., 2. Band, Leipzig.

LOEBLICH, A.R. Jr. & A.R. LOEBLICH IJl (1966): Index to the Genera, Subgenera, und Sections . of the Pyrrhophyta. - Stud. Trop. Oceanogr. 3: 94 pp.

LOEBLICH, A.R. Jr. & A.R . LOEBLICH Ill (1970): Index to the Genera, Subgenera and Sections of the Pyrrophyta, IV.- J. Paleontol. 44: 536-543.

MANGIN, L.A. (19lla): Sur !'existence d'individus dextres et senestres chez certains peridiniens.- C.R. Acad. Sci. Paris 153: 27-32.

MANGIN, L.A. (191Ib): Sur le Peridiniopsis asymetrica et le Peridinium paulseni.- C.R. Acad. Sci. Paris 153: 644-649.

MANGIN, L.A. (1913): Sur la flore planctonique de la rade de Saint-Vaast-LaHougue 1908-1912. -Nouv. Arch. Mus. Hist. nat. Paris, ser. 5, 5: 147-241.

MATSUOKA, K. (1988): Cyst-theca relationships in the Diplopsalid group (Peridiniales, Dinophyceae). - Rev . Palaeobot. Palynol. 56: 95-122.

MEUNIER, A. (1910): Microplancton des Mers de Barents et de Kara. -In: Due d'Orlean, Campagne Arctique de 1907, 355 pp.

MEUNIER, A. (1919): Microplancton de la Mer Flamande. Pat. Ill . Les Peridiniens.- Mem. Mus. Hist. nat. Belg. 8(1): 3-116.

PAULSEN, 0. (1907): The Peridiniales of the Danish waters.- Medd . Komm. Havundersog., Kbh., ser. Plankton 1(5): 1-26.

PAVILLARD, J. (1909): Sur les Peridiniens du Golf de Lion.- Soc. bot. France 56: 277-284.

PAVILLARD, J. (1910): Etat actuel de la Protistologie vegetale.- Progr. Rei Botan. 3: 474-544.

PAVILLARD, J. (1912) : Apropos du Diplopsalis lenticula Bergh.- C.R. Acad . Sci. Paris 155: 173-175 .

PAVILLARD, J. (1913): Le genre Diplopsalis Bergh et les genres voisins.- Montpellier, The Author, p. 1-12.

PA VILLARD, J. (1916): Recherches sur les Peridiniens du Golf de Lion. - Trav . Inst. Bot. Univ. Mont­pellier, Ser. Mixte, Mem. 4: 9-68 .

REID, P.C . (1977): Peridiniacean and Glenodiniacean dinoflagellate cysts from the British Isles.- Nova Hedwigia 29: 429-463.

SEBESTYEN, 0. (1935) : Studies on Diplopsalis acuta (Apstein) Entz with remarks on the question of Kolkwitziel/a salebrosa. - Arch. Protistenk. 85: 20-32 .

SILVA, P. (1980): Names of classes and families of living algae.- Regnum Vegetabile 108: 156 pp .

178