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these characters are insufficient to differentiate species of Ficus,
which are variable in the forms of stem, stipules and leaves, and in
the size of syconia in the same individual when young or mature.
Additionally, when we consider that Desvaux (l.c.: 311) compared
F. yoponensis with an uncertain specimen of F. insipida using only
vegetative characters, it is possible to understand his interpretation.
Currently, among Ficus specialists characters of the figs are emphasized in differentiating species (e.g., DeWolf, l.c. 1960; Carauta, l.c.;
Berg, l.c.; Pederneiras & al. in Rodriguesia 62: 86. 2011; Carauta &
al. in Albertoa 36: 277–284. 2012; Pederneiras & Romaniuc in Syst.
Bot. 37: 684–687. 2012).
Given this conceptual divergence regarding Ficus insipida and
F. yoponensis, we further analyzed these taxa, bringing together
current field observations and herbarium collections to clarify this
taxonomic problem. As a result, F. insipida is considered a wellcircumscribed species, with some minor adjustments to its description
proposed here based on our examination of a specimen (Venezuela, Caracas, Meier 2494, VEN) with the same locality data as the
type: stipules ca. 5 cm long, leaves with most of the secondary veins
straight, forming a right angle with the intramarginal vein, peduncle
1–1.3 cm long, figs with ostiole abruptly prominent, conical, 3–5 mm
tall and 2–3 mm in diameter. In our view, both names apply to the
same species (species A), and although commonly known as F. yoponensis its correct name would become F. insipida. Furthermore,
the name F. glabrata would need to be rehabilitated for the species
(species B) currently known as F. insipida. The two species differ in
that species A has leaves with most of the secondary veins straight,
forming a right angle with the intramarginal vein, and figs with ostiole
abruptly prominent, while species B has leaves with veins uniformly
curved and ostiole plane or shortly elevated when dry. This confirms
Dugand’s (l.c.) observation when he proposed that F. glabrata (species
B) was a well-defined species, differentiated from what he (correctly)
knew as F. insipida (species A).
As we indicated, since that time DeWolf’s (ll.c. 1960, 1965)
misapplication of the name F. insipida to the species (species B) correctly known as F. glabrata has been universally adopted by later
authors. Because of this, rehabilitating F. glabrata from synonymy
as the adopted name of this species and restoring the original concept
of F. insipida, including now F. yoponensis as its synonym, for the
other species (species A, formerly known as F. yoponensis) would
create nomenclatural confusion for both species. This can be avoided
through conservation of F. insipida with a different type selected
here that matches its current usage (species B). The choice of Cuatrecasas 16851, which contains fertile branches and was determined
as “F. glabrata var. typica Dugand” (species B) by Dugand, as type
of F. insipida makes clear the current concept.
Acknowledgements
We are grateful to Dra. Omaira Hokche (VEN) for permission
and facilities for the first author to consult the VEN collection, and
Dr. Jefferson Prado (SP) for his comments on a previous version of
the manuscript. This research was supported by Programa Nacional
de Apoio e Desenvolvimento da Botânica (PNADB), Coordenação de
Aperfeiçoamento de Pessoal de Nível Superior (CAPES).
(2127) Proposal to conserve the name Sphenozamites (Brongn.) Miq.
(Otozamites sect. Sphenozamites Brongn.) with a conserved type
(fossil Cycadophyta: Bennettitales)
Gea Zijlstra,1 Evelyn Kustatscher2 & Han van Konijnenburg-van Cittert1,3
1 Laboratory of Palaeobotany & Palynology, Budapestlaan 4, 3584 CD Utrecht, The Netherlands
2 Museum of Nature Southtyrol, Bindergasse 1, 39100 Bolzano, Italy; Department für Geo- und Umweltwissenschaften,
Paläontologie und Geobiologie, Ludwig-Maximilians-Universität, Richard-Wagner-Strasse 10, 80333 München, Germany
3 Naturalis Biodiversity Center, Postbus 9517, 2300 RA Leiden, The Netherlands
Author for correspondence: Gea Zijlstra, [email protected]
(2127)Sphenozamites (Brongn.) Miq. in Tijdschr. Wis- Natuurk.
Wetensch. Eerste Kl. Kon. Ned. Inst. Wetensch. 4: 210. 1851
(Otozamites sect. Sphenozamites Brongn. in Orbigny, Dict.
Univ. Hist. Nat. 13: 110. 1849), nom. cons. prop.
Typus: S. rossii Zigno (in Atti Reale Ist. Veneto Sci. Lett. Arti
ser. 3, 13: 14. 1868), typ. cons. prop.
The Committee for Fossil Plants (Herendeen in Taxon 60: 904.
2011) left undecided our previous proposal (Zijlstra & al. in Taxon
58: 1016. 2009) to conserve Sphenozamites with a conserved type.
Therefore, we supply now more information, along with a major revision of the earlier proposal. We even considered the possibility of
conserving Sphenozamites Zigno 1881, an artificial homonym under
Art. 48.1 of the ICN (McNeill & al. in Regnum Veg. 154. 2012).
The Melbourne addition to Art. 14.1 allowing conservation of basionyms at the rank of subdivision of a genus (or at infraspecific rank),
however, now enables us to propose conservation with retention of
Brongniart’s basionym. [Editor’s note: Article 14.1 actually provides
for conservation of Otozamites sect. Sphenozamites Brongn., but until
the lay-out of these entries in the Appendices to the Melbourne Code
are finalised, the familiar layout presenting the generic name for
conservation is retained.]
There is no other generic name available for Sphenozamites, even
though Weber (in Revista Mex. Ci. Geol. 13: 201–220. 1996) restored
Macropterygium Schimp. (in Traité Paléont. Vég. 2: 132. 1870) as an
“almost forgotten genus” for a few species that in our opinion can be
placed in Sphenozamites (see below).
When Brongniart (l.c.) created Otozamites sect. Sphenozamites
(see Zijlstra & al., l.c. for more details) he already stated that this
taxon might eventually deserve generic rank. Therefore, it is not
surprising that there is a strong tradition to ascribe the generic name
to Brongniart alone. The fact that Miquel’s raising of this taxon to
Version of Record (identical to print version).
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Zijlstra & al. • (2127) Conserve Sphenozamites
TAXON 62 (1) • February 2013: 185–186
generic rank was published in a Dutch journal (in the Dutch language)
and seems to have escaped notice of all classical authors, apparently
contributed to this tradition. Besides this, Brongniart did something
that was rather unique in that period, particularly for fossil plants: he
designated a type, Cyclopteris beanii Lindl. & Hutton. Of two more
species, Brongniart stated that they also should be placed in this section. When Miquel (l.c. 1851) raised this section to generic rank, he
published new combinations for the three species. In his monograph
(Prodr. Syst. Cycad. 1861), Miquel added six more species, four of
which were based on names in Zamites that Bornemann had given to
fragmentary specimens.
Later on, several authors (Saporta in Paléontol. Franç. Pl. Jurass. 2. Cycadées: 182. 1873; Zigno in Fl. Foss. Oolith. 2: 105. 1881;
Seward in Foss. Pl. 3: 587. 1917; Linnell in Svensk. Bot. Tidskr. 26:
254. 1932) concluded that the concept of Sphenozamites was too wide:
Otozamites includes species with auriculate leaflets, and a species of
which the leaflets are slightly auriculate, should no more be retained
in Sphenozamites, so S. beanii was assigned to Otozamites. Of the two
other original Brongniart species, S. oblongifolius (Kurr) Miq. was
soon qualified as doubtful, e.g., Saporta already said it might better
be placed in Glossozamites; S. undulatus (Sternb.) Miq. got various
questioning comments, yet it was retained in Sphenozamites by many
later authors, e.g., Zigno (l.c.: 108) and Linnell (l.c.: 259).
A few of the oldest authors (Saporta, l.c.: 182; Zigno, l.c.: 106) realized that under this taxonomic treatment the type of Sphenozamites
was excluded from the genus, but they did not draw the conclusion that
this implies that Sphenozamites thus could not any more be used as
a genus alongside Otozamites. Later authors who were already more
familiar with the type concept (Seward, l.c.: 587; Linnell, l.c.: 242,
259—both authors who stated that Zigno had raised Sphenozamites
to generic rank!) apparently did not realize that the type of Sphenozamites was excluded; maybe they have not even seen Brongniart—
they wrongly wrote that Brongniart treated it as a subgenus (not as a
section). Not even Wesley (in Mem. Ist. Geol. Mineral. Univ. Padova
21: 1–56, t. 1–3. 1958), who studied Zigno’s rich collections from the
Jurassic of N Italy, seemed to be aware of the exclusion of the original
type. Albeit that since Wesley’s publication, some authors started to
give the authorship as “Brongn. emend. Wesley”. More recent authors
(e.g., Harris, Yorkshire Jurassic Fl. 3: 12–16. 1969; Wang & al. in
Palaeoworld 17: 222–234. 2008; Yamada & Uemura in Paleontol. Res.
12: 7. 2008: Crane & Herendeen in Amer. J. Bot. 96: 284–295. 2009)
continued to treat S. beanii as a species of Otozamites.
Doludenko (in Bot. Zhurn. (Moscow & Leningrad) 59: 560–564.
1974) stated (p. 560) that according to Andrews (in Bull. U.S. Geol.
Surv. 1300: 201. 1970), Miquel in 1851 had raised Brongniart’s
subgenus to generic rank, with Cyclopteris beanii as its type (yes,
Andrews also wrongly wrote “subgenus”). After discussion of the
conclusions of the later authors (Saporta, l.c.; Zigno, l.c.; Seward, l.c.;
Linnell, l.c.; Wesley, l.c.), Doludenko (l.c.: 561) designated another
type: S. rossii Zigno.
Cleal & Rees (in Palaeontology 46: 758. 2003) were the first
authors to notice that conservation is necessary, and anticipating
that this should be formally done, they continued to use the current
concept of Sphenozamites: “Wesley’s revision is nevertheless the preferred option and we have followed it here.” We hope that with the current revised proposal, this conservation can actually be undertaken.
186
As for Macropterygium Schimp. that was recently restored (see
above), according to Weber (l.c.) this genus was restricted to western
and far-eastern Laurasia. Weber included four species: M. bronnii
(Schenk) Schimp. (this is one of Schimper’s original two species,
selected as lectotype of Macropterygium by Andrews in Bull. U.S.
Geol. Surv. 1013: 183. 1955; Weber agreed with that choice). Weber
added M. rogersianum (Fontaine) R. Weber, M. truncatum (Zeiller)
R. Weber and M. mexicanum R. Weber. Weber (l.c.: 216–217) excluded
several species that had been placed in Macropterygium by later authors (and that we also do not consider species of Sphenozamites).
Weber also mentioned a few species of Sphenozamites, a genus that
he accepted besides the revived Macropterygium; of S. changii Sze
(1956) he stated that “the original assignment to Sphenozamites seems
to be reasonable”. For the two species that have not yet been included
in Sphenozamites, we now publish the new combinations:
Sphenozamites truncatus (Zeiller) Zijlstra, Kustatscher & van Konijnenb., comb. nov. ≡ Zamites truncatus Zeiller, Fl. Foss. Charbon
Tonkin, Text: 166. 1903) ≡ Macropterygium truncatum (Zeiller)
R. Weber, l.c.: 210 – Holotype: specimen figured by Zeiller,
Fl. Foss. Charbon Tonkin, Atlas: pl. 43, fig. 4, 4a. 1902 – Type
locality: Hongaÿ Mine, eastern valley of l’Oeuf, Shaft Léonice,
Vietnam.
Sphenozamites mexicanus (R. Weber) Zijlstra, Kustatscher & van
Konijnenb., comb. nov. ≡ Macropterygium mexicanum R. Weber
in Revista Mex. Ci. Geol. 13: 210. 1996 – Holotype: Specimen
IGM-PB-521B-889 (pl. 5, fig. 1) from loc. 521B, CLP-IG-UNAM
– Type locality: Hermosillo-Tonichi highway, approx. km 152,
La Barranca Section 2 (measured by Potter & al., 1980), Unit 42,
“LB 3 stratum” and “Semillas” stratum (loc. 521B and C, CLPIG-UNAM; see Weber [pp. 107–124] in Weber (ed.), III Congr.
Latinoamericano Paleont. México: fig. 4, tab. 1. 1985), road-cut;
28°34′40″ N/109°40′06″ W, Sonora, Mexico.
Sphenozamites includes 24 species; it is found in Triassic and
Lower Jurassic strata of the Northern Hemisphere, with the main
distribution centers in Europe and Asia (especially China). European
species are: S. angustipinnatus Passoni & van Konijnenb., S. bellii
Seward, S. bronnii (Schenk) Passoni & van Konijnenb. (also in Mexico), S. geylerianus Zigno, S. latifolius (Brongn.) Schimp., S. rochei
Renault 1882, S. rossii (Zigno) Zigno (with two synonyms: S. adiantifolius Zigno, S. lanceolatus Zigno), S. sphenozamioides (Turutanova-Ketova) Doludenko, S. vachrameevii Doludenko, S. wengensis
Wachtler & van Konijnenb. Asian species are: S. changii H.C. Sze,
S. donggongensis F.S. Meng, S. drepanoides B.X. Li, S. evidens
F.S. Meng, S. fenshuilingensis F.S. Meng, S. marionii Counillon,
S. rhombifolius F.S. Meng, S. surakaicus Pryn. ex Brik, S. truncatus (Zeiller) Zijlstra & al. (also in Mexico), S. yungjenensis J. Hsü
& S.Y. Tuan. Only in Central America: S. mexicanus (R. Weber)
Zijlstra & al. (Mexico) and S. robustus Newb. (Honduras). Only in
North America: S. oblanceolatus Penh. (Canada) and S. rogersianus
Fontaine (U.S.A.).
Acknowledgement
We wish to thank John McNeill for suggestions to improve the text.
Version of Record (identical to print version).