Proposals to conserve the names Cycadeoidea ...

Doweld • (2436–2437) Conserve Cycadeoidea and C. megalophylla

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(2436–2437) Proposals to conserve the names Cycadeoidea against Mantellia and C. megalophylla against M. nidiformis (fossil Spermatophyta: Cycadeoideopsida) Alexander B. Doweld National Institute of Carpology (Gaertnerian Institution), 21 Konenkowa Street, 127560, Moscow, Russian Federation; [email protected]; [email protected] DOI http://dx.doi.org/10.12705/652.29

(2436) Cycadeoidea Buckland ex Lindl. & Hutton, Foss. Fl. Gr. Brit. 1: xli. Apr 1832, nom. cons. prop. Typus: C. megalophylla Buckland, nom. cons. prop. (Mantellia nidiformis Brongn.). (≡) Mantellia Brongn., Prodr. Hist. Vég. Foss.: 96. Dec 1828; in Cuvier, Dict. Sci. Nat., ed. 2, 57: 101. Dec 1828, nom. rej. prop. (2437) Cycadeoidea megalophylla Buckland, O Sagovovid. Iskop. Rast. Portlanda: 11. 1834 (post 9 Feb), nom. cons. prop. Typus: Great Britain, Isle of Portland, Upper Jurassic (Tithonian) (Oxford Univ. Mus. Nat. Hist. No. J.23000).

(≡)

Mantellia nidiformis Brongn., Prodr. Hist. Vég. Foss.: 200. Dec 1828; in Cuvier, Dict. Sci. Nat., ed. 2, 57: 192. Dec 1828 (neotyp. des.: Watson & Lydon in Cretaceous Res. 25: 14. 2004), nom. rej. prop.

The fossil genus Cycadeoidea is one of the well-reconstructed and widely known fossil genera of modern palaeobotany, the name serving as type of a distinct group of pre-angiosperm extinct plants, bennettitaleans or cycadeoids, and formally classified in a separate class Cycadeoideopsida D.H. Scott (Stud. Foss. Pl., ed. 3: 320. 1923 (‘Cycadeoideae’)) or phylum Cycadeoideophyta T.N. Taylor

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(Paleobotany: 25. 1981 (‘Cycadeoidophyta’)), also known under alternative suprafamilial names such as Bennettitopsida Engler (Syllabus: 61. 1892 (‘Bennettitales’)) and Bennettitophyta Kravtsov & Poljarnaja (Paleobotanika: 43, 45. 1995). The nomenclature of the genus is very complex and confused (Doweld, l.c.) because of the numerous nomenclatural mistakes in the past that led to the current status of Cycadeoidea as an illegitimate, nomenclaturally superfluous generic name that should be rejected in favour of the nearly forgotten Mantellia, which disappeared from systematic palaeobotany at the end of 19th century. Cycadeoidea with two intended species names, C. megalophylla and C. microphylla, first appeared in a brief report by Buckland (in Proc. Geol. Soc. London 1: 80–81. Jun 1828) [Archive Records of the Geological Society of London for publication of No. 8 of the first volume of the for 1826–1833]. Buckland, being inspired and consulted by famo Proceedings us British botanists Robert Brown and George Loddiges, upon their influence provided description of an intended new fossil family Cycadeoideaceae (‘Cycadeoideae’) and detailed descriptions of two fossil species, but failed to provide a separate generic diagnosis. As a consequence, none of the names, those of the two species, of the family and of the genus itself, was validly published due to the lack of a description or diagnosis for Cycadeoidea. A similar brief summary of the Proceedings of the Geological Society was independently re-published (Buckland in Philos. Mag. Ann. Chem., ser. 2, 4: 225–226. 1 Sep 1828) with a distinct extensive description of a new family and two species, but again lacking the necessary generic description for validation of the names. The same taxonomic format was used by Buckland again in his main monographic paper (in Trans. Geol. Soc. London 2: 295–401. 28 Nov 1828) [exact date of publication was established from the Minutes of the Council, Archive of the Geological Society of London: LDGSL CM1/2]. In sum, in none of Buckland’s publications was Cycadeoidea validly published as a generic name. At the end of 1828, Adolphe Brongniart renamed these British fossils as Mantellia (in Cuvier, Dict. Sci. Nat., ed. 2, 57: 101. Dec 1828 & Prodr. Hist. Vég. Foss.: 96. Dec 1828). Brongniart was well-acquainted with Buckland’s 1828 publications (l.c.: 92, 199. 1828 [Prodr.]), but disagreed with the British geologist in the application of Cycadeoidea to these fossils: “nom qui nous paroît avoir l’inconvénient de pouvoir s’appliquer également à toutes les plantes fossiles de cette famille, qui doivent cependant constituer des genres bien distincts”. Although in the enumeration of taxa in the “Famille Cycadées” Mantellia (on p. 96) was associated with a generic description in French and thereby validly published, the two species listed there, M. nidiformis and M. cylindrica, were devoid of any original description or illustration that led subsequent researchers to have doubts as to whether Mantellia and its species were validly published. However, Brongniart (l.c.: 1828 [Dict.]: 192 & l.c. 1828 [Prodr.]: 200, both in footnotes) replaced (renamed) one of Buckland’s intended species names, “Cycadeoidea megalophylla”, by his new species name, Mantellia nidiformis, thus validating it through Buckland’s intended species description. The second species in the original protologue of Mantellia, “M. cylindrica Brongn.”, remained a nomen nudum with no description until that provided by Mantell (Medals Creat. 1: 159. 1844). In numerous palaeobotanical synonymies published for over 180 years one could not find the established nomenclatural fact that Mantellia and its type M. nidiformis are the earliest validly published names for Cycadeoidea and C. megalophylla. However, the generic name Cycadeoidea was soon validated by Lindley & Hutton (l.c.: xli), who provided a separate generic diagnosis in English, but unfortunately included the previously validly

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published Mantellia in its synonymy rendering their Cycadeoidea Buckland ex Lindl. & Hutton nomenclaturally superfluous and illegitimate when published (Art. 52 of the ICN, McNeill & al. in Regnum Veg. 154. 2012). Therefore, Cycadeoidea, being a superfluous nomenclatural synonym of Mantellia, is automatically typified by M. nidiformis under Art. 7.5. Perhaps the British authors also overlooked the lack of a separate generic diagnosis in Buckland’s 1828 publications and simply subsumed Brongniart’s name into synonymy on the grounds of its later publication. Lindley & Hutton (l.c.) did not mention any species within Cycadeoidea, and C. megalophylla and C. microphylla were not validly published until 1834 when they were validated in Buckland’s paper from the Transactions that was translated and published with annotations in Russian by Obodovsky (Bokland, V., O sagovovidnykh iskopaemykh rastenijakh […] Portlanda. St.-Petersburg, N. Grech 1834. 23 pp.; Censor’s permission for publication of 28 Jan 1834 [Old Style] = 9 Feb 1834 [New Style], presumably shortly published after censorship, but precise date of publication was not established due to the gaps in archival records of Russian printing). Mantellia has incorrectly been viewed as a superfluous later replacement or synonymic name for Buckland’s original name, even though it is not so, because Buckland’s generic name was not validly published until 1832. No literature exists between 1828 and 1832, in which Cycadeoidea might be validated, although the name was in some use in geological treatises (Doweld, l.c.). However, soon Cycadeoidea was withdrawn by Buckland (Geol. Mineral. 1: 496. 24 Sep 1836) from palaeobotanical systematics, by stating that: “in my paper, just quoted [in Trans. Geol. Soc. London, l.c. 1828], I applied to them [the fossils] the provisional name of Cycadeoidea megalophylla and Cycadeoidea microphylla; but Mr. Brown is of opinion, that until sufficient reasons are assigned for separating them from the genus Cycas or Zamia, the provisional name of Cycadites is more appropriate, as expressing the present state of our knowledge upon this subject.” As a consequence, both species of Cycadeoidea were recombined in 1836 as Cycadites megalophyllus (Buckland) Buckland, nom. illeg. & superfl. [Mantellia nidiformis Brongn. incl. as synonym] and C. microphyllus (Buckland) Buckland. Mantellia then came into taxonomic literature and became widely used (Bronn, Lethaea Geogn.: 153. 1835 & 227. 1836; Endlicher, Gen. Pl.: 72. 1836; Brongniart in Orbigny, Dict. Univ. Hist. Nat. 4: 485. 1844; Jussieu in Orbigny, Dict. Univ. Hist. Nat. 7: 758. 1846; Pomel in Amtl. Ber. Versamml. Deutsch. Naturf. Ärzte 25: 341, 340. 1849; Carruthers in Rep. Brit. Assoc. Advancem. Sci. 37(2, Trans. Sect.): 80. 5 Aug 1868 [‘1867’] & in Trans. Linn. Soc. London 26: 678, 701. 1870; Mansel-Pleydell in Proc. Dorset Nat. Hist. Antiq. Field Club 2: 8. 1878; Zigno, Fl. Foss. Oolith. 2: 192. 1885; Engler & Gilg, Syllabus, ed. 8: 107. 1920 [‘1919’]). However, Mantellia was noted (Buckland, l.c. 1836) as being a homonym of the fossil Cretaceous sponge, Mantellia Parkinson (Intr. Foss. Organ Remains: 53. 1822). The independence of botanical nomenclature from zoological establishes that there are no grounds for treating the fossil plant name Mantellia as an illegitimate later homonym of the earlier published fossil animal name. But at that time Brongniart himself (in Orbigny, Dict. Univ. Hist. Nat. 13: 108, 156. 1849) withdrew the name Mantellia and superseded it by Cycadeoidea (misspelled as “Cycadoidea”), although just a few years earlier he (in Orbigny, l.c. 4: 485. 1844) still preferred to use his Mantellia. Cycadeoidea was finally restored in systematic palaeobotany by Schimper in his influential treatise (Traité Paléontol. Vég. 3: 556. 1874, misspelled again as “Cycadoidea”) and since then the generic name has come into wide and unquestioned use in botany, including associated geological works and even popular literature (Capellini

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& Solms-Laubach in Mem. Reale Accad. Sci. Ist. Bologna, ser. 5, 2: 187. 1892; Ward in Proc. Biol. Soc. Wash. 9: 79. 1894; Wieland, Amer. Foss. Cycads 2: 17. 1916; Seward, Foss. Pl. 3: 367. 1917; Darrah, Princ. Paleobot.: 133. 1939; Kryshtofovich, Paleobotanika, ed. 4: 508. 1957; Delevoryas in Amer. J. Bot. 50: 45. 1963; Němejc, Paleobotanika 3: 265. 1968; Crepet in Palaeontographica, Abt. B, Paläophytol. 148: 145. 1974; Watson & Lydon in Cretaceous Res. 25: 11. 2004). Up to now, more than 120 species names have been published under Cycadeoidea (Jongmans & Dijkstra in Jongmans, Foss. Cat. Pl. 38: 867. 1959 & 67: 3836. 1967; Dijkstra & Van Amerom in Dijkstra, Foss. Cat. Pl. 88: 132. 1981; The International Fossil Plant Names Index, 2014–); more than 100 of them would need to be recombined into the disused Mantellia on account of priority. In order to stabilize palaeobotanical nomenclature in current use, and to legitimize the illegitimate Cycadeoidea, it is formally proposed to conserve this generic name against its earlier nomenclatural synonym, Mantellia. The species epithet of the generic type of Cycadeoidea, C. megalophylla (Andrews in Bull. U.S. Geol. Surv. 1013: 140. 1955), has entered so solidly into palaeobotanical systematics that nobody has recognised its illegitimate, superfluous status up to now. I propose also to conserve the species name C. megalophylla against its disused senior taxonomic synonym, M. nidiformis Brongn., providing the necessary typification of the latter name with the same specimen by which C. megalophylla was recently neotypified due to loss of original fossil specimens of 1828 (Watson & Lydon, l.c.: 14). As noted above, C. megalophylla was validated in the Russian translation of Buckland’s 1828 paper from the Transactions of the Geological Society of London (Bokland, l.c. 1834) The conservation of Cycadeoidea becomes more important and needed to legitimize (under the revised provisions in Art. 6.4, 18.3, & 19.6 of the Melbourne Code) the current use of a distinct family name for this group of fossils, Cycadeoideaceae Buckland

ex Fitton (Geol. Sketch Hastings: 77. 26 Jan 1833 (‘Cycadeoideae’)). Although the other available suprageneric names in current use such as: ordinal (Cycadeoideales E.W. Berry in Pl. World 19: 36. 1916 (‘Cycadeiodales’)), subordinal (Cycadeoideineae Němejc, l.c. 1968: 265), subclass (Cycadeoideidae Němejc, l.c. 1968: 275), class (Cycadeoideopsida D.H. Scott (l.c.) (‘Cycadeoideae’)) and phylum names (Cycadeoideophyta T.N. Taylor (l.c.) (‘Cycadeoidophyta’)) are not made illegitimate, by the illegitimacy of the generic name from which they are formed, it would be undesirable to have all the suprafamilial nomenclature of this fossil plant group based on a different genus from that of the family and changed into that derived from the alternative fossil cycadeoid generic name, Bennettites Carruth., which is also available and sometimes alternatively used in modern palaeobotany such as the family Bennettitaceae Lignier (in Nature 50: 595. 1894 (‘Bennettiteae’) (even although Cycadeoideaceae has priority), suborder Bennettitineae D.H. Campbell (Univ. Text-book Bot.: 347. 1902 (‘Bennettiteae’)), order Bennettitales Bessey (in Trans. Amer. Microscop. Soc. 29: 94. 1910), subclass Bennettitidae Davitashvili (Kurs Paleontol.: 701. 1949 (‘Bennettitales’)), class Bennettitopsida Engler (l.c. (‘Bennettitales’)) and phylum Bennettitophyta Kravtsov & Poljarnaja (l.c.). The legitimization of Cycadeoidea-derived suprageneric nomenclature will provide stabilization in the whole classification of extinct gymnosperms. Acknowledgements It is a pleasure to thank Valentina Bublik (Fundamental Botanical Library of the National Institute of Carpology, Moscow) for bibliographic assistance. Thanks also go to the librarians of the Geological Society of London for providing archive records for the exact date of publications of the Society (Sheila Meredith and Wendy Cawthorne). The research is a contribution to the Palaeoflora Europaea Project and Palaeoflora of Russia (Palaeoflora Rossica) Project (NOM-10-19).

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