RESEARCH ARTICLE
Douropteris alvarezii gen. nov., sp. nov., a new medullosalean
pteridosperm from the Late Pennsylvanian of Portugal
Pedro Correia
1,5
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Zbynĕk Šimůnek
2
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Christopher J. Cleal
3
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Artur A. Sá
4,5
1
Institute of Earth Sciences of University of
Porto, Porto, Portugal
2
Czech Geological Survey, Prague, Czech
Republic
3
Department of Natural Sciences, National
Museum Wales, Cardiff, UK
4
Department of Geology, University of
Trás‐os‐Montes e Alto Douro, Vila Real,
Portugal
5
Geosciences Centre, University of Coimbra,
Coimbra, Portugal
Correspondence
P. Correia, Institute of Earth Sciences, Pole of
the Faculty of Sciences, University of Porto,
Rua do Campo Alegre, 687, 4169‐007 Porto,
Portugal.
Email: pedro.correia@fc.up.pt
Funding information
Fundação da Ciência e Tecnologia, Grant/
Award Number: SFRH/BD/78741/2011;
Fundação Calouste Gulbenkian; National
Museum Wales Research Program, Grant/
Award Number: NS49; Czech Geological Sur-
vey, Grant/Award Number: CGS 323100
Handling Editor: Ian D. Somerville
A new plant fossil‐genus, Douropteris alvarezii gen. nov., sp. nov., is described from the
Douro Basin (lower Stephanian C [lower Gzhelian], Upper Pennsylvanian) in north‐
western Portugal. Douropteris alvarezii has a remarkable morphology characterized
by an unusual combination of features seen in several different fossil‐genera belong-
ing to ferns and pteridosperms. Its unique and distinguishing features place it nearest
to the medullosalean pteridosperms Margaritopteris and Callipteridium, suggesting
close affinities with the fossil‐family Cyclopteridaceae. Morphologically,
Margaritopteris is the closest relative with basis on the strong resemblance to the
veining pattern of Douropteris gen. nov. Here, Douropteris represents a potential
“Lazarus taxon” or evolutionary descendant from the Margaritopteris lineage.
KEYWORDS
Douro Basin, Douropteris alvarezii gen. nov., Late Pennsylvanian, Medullosales, Portugal,
pteridosperm, sp. nov.
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INTRODUCTION
The medullosalean pteridosperms (Medullosales) are a group of
extinct seed‐bearing plants characterized by fern‐like foliage that
flourished widely in tropical habitats of Euramerica and China during
the late Carboniferous to early Permian (e.g., Cleal, 2008; Cleal &
Popa, 2015; Cleal & Shute, 2003, 2012; Mapes & Rothwell, 1980;
Seyfullah, Hilton, Wang, & Galtier, 2009; Taylor & Eggert, 1967).
They occupied mainly wetland habitats but during the Moscovian
occurred more abundantly in better drained clastic substrate habitats
(Cleal, 2008). These habitats generally were located in lowland and
upland intramontane basins (e.g., Cleal, 2008; Cleal & Popa, 2015;
Šimůnek & Cleal, 2004). The first medullosalean forms appeared dur-
ing the late Mississippian (Cleal & Shute, 2012; Dunn et al., 2003;
Laveine, Lemoigne, & Shanzhen, 1993; Mapes & Rothwell, 1980;
Taylor & Eggert, 1967), but they were especially abundant and
diverse in Westphalian and Stephanian times (Cleal & Shute, 2012).
The last representatives of the medullosalean pteridosperms became
extinct in Euramerica and China (Amerosinian Realm) in the Asselian
(early Permian; Cleal & Shute, 2012; Rößler et al., 2012; Seyfullah
et al., 2009). Most medullosalean taxa were monoaxial shrubs and
small trees (e.g., Cleal & Shute, 2012; Retallack & Dilcher, 1988;
Wnuk & Pfefferkorn, 1984), with fronds that ranged in length from
1 to 7 m (e.g., Cleal & Shute, 1991; Cleal & Shute, 2012; Laveine,
1986; Laveine & Belhis, 2007), although some Stephanian–early
Permian taxa (e.g., Lescuropteris Schimper and Odontopteris
[Brongniart] Sternberg) have developed a scrambling/climbing or
lianescent growth habit (described as vine‐ and liana‐like pterido-
sperms), with axillary branches arising from the stems and having
non‐arborescent forms with small fronds usually less than 0.5 m
(Cleal & Shute, 2012; Hamer & Rothwell, 1988; Krings et al., 2006;
Krings & Kerp, 1997; Krings, Kerp, Taylor, & Taylor, 2003). The
medullosaleans generally produced large ovules and had complex pol-
len‐organs, consisting of elongate pollen sacs (sporangial tubes)
arranged in a ring or aggregated into concentric rings (e.g., Anderson,
Anderson, & Cleal, 2007; Beck & Wight, 1988; Cleal, 2008).
Received: 8 December 2017 Revised: 4 May 2018 Accepted: 5 May 2018
DOI: 10.1002/gj.3251
Geological Journal. 2018;1–11. Copyright © 2018 John Wiley & Sons, Ltd. wileyonlinelibrary.com/journal/gj 1