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The Carboniferous floras of the Iberian Peninsula: A synthesis with geological connotations

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  • Botanical Garden of Cordoba

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A general review is presented of the Carboniferous floral records in the Iberian Peninsula in the context of the geological history and distribution of the different basins. Mississippian floras are found in Sierra Morena, where major strike-slip faults brought in terranes of diverse provenance. Lower Pennsylvanian floras are represented in the Peñarroya–Belmez–Espiel (Córdoba) and Villanueva del Río y Minas (Sevilla) coalfields of SW Spain (also strike-slip controlled), at La Camocha, near Gijón (Asturias), and in other parts of northern Spain. Middle Pennsylvanian is represented near Oporto, but more completely in the Central Asturian Coalfield, as well as other, more limited localities in NW Spain and the Pyrenees. Upper Pennsylvanian (Stephanian Stage–Cantabrian to Stephanian B substages) floras are splendidly represented in NW Spain. Uppermost Pennsylvanian (Stephanian C–Autunian) floras are present in the strike-slip controlled Douro and Buçaco basins of North Portugal, the Pyrenees, Central Spain (Ciudad Real, Guadalajara, Zaragoza), and SW Spain (Guadalcanal and Valdeviar in Sevilla province).A complete succession of megafloral zones is presented. This includes a new Annularia spicata Zone at the top of the Pennsylvanian (equivalent to middle to upper Autunian which has been often attributed to the Lower Permian). The information is summarised in charts compiled from a selection of the most significant species. A number of floral elements are illustrated including the zonal indices. Brief taxonomic comments are provided in the Appendix.
Map showing the different plant-bearing occurrences of Carboniferous age in the Iberian Peninsula (provincial name in brackets) (1a-enlarged portion of the same showing a more complete representation of localities in NW Spain; 1b-enlarged portion showing localities in Sierra Morena, SW Spain, in more detail): 1. Vale de Corvos (Faro); 2. Santa Susana (Setubal); 3. Guadalcanal (Sevilla); 4. Valdeviar (Sevilla); 5. Villanueva del Río y Minas (Sevilla); 6. Los Santos de Maimona (Badajoz); 7. Bienvenidas (Badajoz); 8. Casas de Reina (Badajoz); 9. Berlanga (Badajoz); 10. Valdeinfierno (Córdoba/Sevilla); 11. Cerrón El Hornillo/Puebla de los Infantes (Sevilla); 12. Virgen de Linares (Córdoba); 13. Benajarafe/ Matachel (Córdoba); 14. Guadiato (Córdoba); 15. Peñarroya-Belmez-Espiel (Córdoba); 16. Serra de Rates, Bougado (Porto); 17. Casais/Alvarelhos (Porto); 18. Ervedosa (Bragança); 19. Douro (Porto/Viseu); 20. Buçaco (Aveiro); 21. Sierra de San Pedro (Cáceres); 22. Guadalmez (Ciudad Real); 23. Puertollano (Ciudad Real); 24. Henarejos (Cuenca); 25. Valdesotos/Retiendas (Guadalajara); 26. Fombuena (Zaragoza); 27. Sierra de la Demanda (Burgos); 28. San Vitero (Zamora); 29. Arnao/San Juan de Nieva (Asturias); 30. Tineo (Asturias); 31. Cangas de Narcea (Asturias); 32. Rengos (Asturias); 33. Tormaleo (Asturias); 34. El Bierzo (León); 35. Carballo (Asturias); 36. Carrasconte/Villablino (León); 37. La Camocha (Asturias); 38. Monte Naranco (Asturias); 39. Puerto Ventana (León); 40. Teverga/San Emiliano (Asturias); 41. Quirós (Asturias); 42. Viadangos/Rodiezmo (León); 43. La Magdalena (León); 44. Ciñera-Matallana (León); 45. Central Asturian Coalfield (Asturias); 46. San Tirso (Asturias); 47. Canseco/Rucayo (León); 48. Tejerina/Ocejo de la Peña (León); 49. Prioro/Pando (León); 50. Sabero (León); 51. Guardo (Palencia/León); 52. Cueli (Asturias); 53. Sebarga (Asturias); 54. Gamonedo/Inguanzo (Asturias); 55. Arenas de Cabrales (Asturias); 56. Valdeón (León); 57. Sotres (Cantabria); 58. Cucayo/Dobres (Cantabria); 59. Curavacas Formation (Palencia); 60. La Pernía (Palencia); 61. Castillería (Palencia); 62. Peña Cildá (Palencia); 63. Barruelo/Redondo (Palencia); 64. Pico Cordel (Cantabria); 65. Puente Pumar (Cantabria); 66. Ibantelly (Navarra); 67. Collado de Izas (Huesca); 68. Béarnais (French Pyrenees); 69. Aguiró (Lleida); 70. Malpás (Lleida); 71. Baró/Arcalís (Lleida); 72. Coll de Jou (Girona); 73. Ogassa/Surroca (Girona); 74. El Priorat (Tarragona).
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The Carboniferous oras of the Iberian Peninsula: A synthesis with
geological connotations
Robert H. Wagner , Carmen Álvarez-Vázquez
Centro Paleobotánico, Jardín Botánico de Córdoba, Avenida de Linneo, s/n, 14004 Córdoba, Spain
abstractarticle info
Article history:
Received 18 January 2010
Received in revised form 16 May 2010
Accepted 6 June 2010
Available online 18 June 2010
Keywords:
Carboniferous
Permian
oristics
Iberia
Spain
Portugal
A general review is presented of the Carboniferous oral records in the Iberian Peninsula in the context of the
geological history and distribution of the different basins. Mississippian oras are found in Sierra Morena,
where major strike-slip faults brought in terranes of diverse provenance. Lower Pennsylvanian oras are
represented in the PeñarroyaBelmezEspiel (Córdoba) and Villanueva del Río y Minas (Sevilla) coalelds of
SW Spain (also strike-slip controlled), at La Camocha, near Gijón (Asturias), and in other parts of northern
Spain. Middle Pennsylvanian is represented near Oporto, but more completely in the Central Asturian
Coaleld, as well as other, more limited localities in NW Spain and the Pyrenees. Upper Pennsylvanian
(Stephanian StageCantabrian to Stephanian B substages) oras are splendidly represented in NW Spain.
Uppermost Pennsylvanian (Stephanian CAutunian) oras are present in the strike-slip controlled Douro
and Buçaco basins of North Portugal, the Pyrenees, Central Spain (Ciudad Real, Guadalajara, Zaragoza), and
SW Spain (Guadalcanal and Valdeviar in Sevilla province).
A complete succession of megaoral zones is presented. This includes a new Annularia spicata Zone at the top
of the Pennsylvanian (equivalent to middle to upper Autunian which has been often attributed to the Lower
Permian). The information is summarised in charts compiled from a selection of the most signicant species.
A number of oral elements are illustrated including the zonal indices. Brief taxonomic comments are
provided in the Appendix.
© 2010 Elsevier B.V. All rights reserved.
Contents
1. Preamble ............................................................... 240
2. Brief historical account ......................................................... 240
3. Geological overview .......................................................... 241
4. Carboniferous oras in different parts of the Iberian Peninsula ...................................... 247
4.1. Mississippian oras ....................................................... 248
4.1.1. Tournaisian (Plate I, gs. 14; Plate II, gs.1,2)..................................... 250
4.1.2. Visean (Plate II, gs. 3, 4; Plate III, gs. 15a; Plate IV, g. 1)s............................... 250
4.1.3. Serpukhovian (Plate IV, gs. 23)............................................ 254
4.2. Pennsylvanian oras....................................................... 254
4.2.1. Middleupper Namurian (Plate V, gs. 2, 4, 5; Plate VI, g.1)............................... 254
4.2.2. Langsettian (Table 1; Plate V, gs. 1, 3; Plate VI, gs. 2, 2a; Plate VII, gs. 16; Plate VIII, g. 1; Plate IX, g. 2; Plate X, gs. 1, 2; Plate XII,
g.1).......................................................... 258
4.2.3. Duckmantian (Table 2; Plate VIII, gs. 24; Plate IX, g. 1; Plate X, gs. 35; Plate XI, gs. 13a; Plate XII, gs. 25; Plate XIII,
gs.1,3,6)...................................................... 262
4.2.4. Bolsovian (Table 3; Plate XIII, gs.2,4,5)........................................ 266
4.2.5. Asturian (Table 4a; Plate XIV, gs. 14; Plate XV, gs. 15; Plate XVI, gs. 1, 5 Linopteris obliqua Zone; and Table 4b; Plate XVI, gs. 2
4, 6 Lobatopteris vestita Zone) ............................................. 270
4.2.6. Cantabrian (Table 5; Plate XVII, gs. 16; Plate XIX, gs.1,2)............................... 274
4.2.7. Barruelian (incorporating Stephanian A as known from the Massif Central, France) (Table 6; Plate XVIII, g.1)........ 276
4.2.8. Saberian(corresponding, approximately, to Stephanian B sensu Carmaux) (Table 7; Plate XIX, g. 3; Plate XX, gs. 25; Plate
XXI, gs. 14)..................................................... 278
Review of Palaeobotany and Palynology 162 (2010) 239324
Corresponding author.
E-mail address: cr1wagro@uco.es (R.H. Wagner).
Plate X, gs. 1, 2; Plate XII, g. 1)
Plate XII, gs. 25; Plate XIII, gs. 1, 3, 6)
Plate XVI, gs. 24, 6 Lobatopteris vestita Zone)
Plate XX, gs. 25; Plate XXI, gs. 14)
0034-6667/$ see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.revpalbo.2010.06.005
Contents lists available at ScienceDirect
Review of Palaeobotany and Palynology
journal homepage: www.elsevier.com/locate/revpalbo
4.2.9. Stephanian B (i.e. the classical Stephanian B sensu Saint Étienne) (Table 8; Plate XIX, g. 4; Plate XX, g. 1; Plate XXII, gs. 1, 3; Plate
XXIII, gs. 15; Plate XXIV, gs. 1, 2; Plate XXV, gs. 14)................................. 282
4.2.10. Stephanian C (=lower Autunian) (Table 9; Plate XXII, g. 2; Plate XXVI, gs. 1, 2, 4; Plate XXIX, g. 2)s.......... 289
4.2.11. Mid to upper Autunian (Table 10; Plate XXVI, gs. 3, 57; Plate XXVII, gs. 15; Plate XXVIII, gs. 15; Plate XXIX, gs. 1, 3; Plate XXX,
gs. 18) ....................................................... 294
5. General considerations on the Carboniferous oras as found in the Iberian Peninsula . ........................... 302
6. Reections on extrabasinalelements .................................................. 304
7. Biostratigraphy: megaoral zones .................................................... 305
7.1. Mississipian and lower Pennsylvanian zones............................................ 305
7.2. Upper Pennsylvanian (Stephanian) zones ............................................. 309
7.2.1. Stratigraphic subdivisions of the Stephanian........................................ 309
7.2.2. Megaoral biostratigraphy of the Stephanian ....................................... 309
7.2.3. The higher Autunian: Annularia spicata megaoral zone (new) ............................... 312
8. Summary and conclusions........................................................ 314
Acknowledgements ............................................................. 314
Appendix. Brief taxonomic comments ..................................................... 315
References .................................................................. 315
1. Preamble
Modern work on the Carboniferous oras of Spain commenced with
the critical review published by Jongmans (1951), who also incorpo-
rated the preliminary results of his own investigations. Jongmans's
paper provided the starting point for more detailed investigations
carried out partly by himself (up to his death in 1957), partly by his
pupil, R.H. Wagner and his alumni, and also by different workers such as
F. Stockmans and Y. Willière from Brussels, J.-P. Laveine and his
associates in the University of Lille, J. Doubinger from Strasbourg,
J. Broutin from Paris, M.C. Álvarez-Ramis in Madrid, and several others,
as mentioned in the bibliography. Substantial collections were brought
together by the different workers. Perhaps the most important
collection available in Spain is beingcurated in the Centro Paleobotánico
(IMGEMA-Jardín Botánico de Córdoba), but other collections of note are
in the Museo Geominero (Instituto Geológico y Minero de España), the
Museo Nacional de Ciencias Naturales in Madrid, the Museu de Ciències
Naturals in Barcelona, and the Geology Departments of the universities
of Oviedo and Zaragoza. Outside Spain, there are additional collections
in the Nationaal Natuurhistorisch Museum (Naturalis) in Leiden, the
Musée Royale des Sciences Naturelles in Brussels, the Senckenberg
Museum in Frankfurt am Main, the Jardin des Plantes in Paris, and the
Musée d'Histoire Naturelle in Lille. Indeed, the collections are widely
scattered.
With regard to Portugal, the investigations of C. Teixeira, from
1937 to 1964, yielded important collections kept in the universities of
Oporto and Lisbon, as well as the Serviços Geologicos de Portugal
(presently INETI Geological Survey). These collections were revised by
Wagner and Lemos de Sousa (1983a).
2. Brief historical account
The early investigations in Spain mainly resulted in lists of
Carboniferous plant fossils being published. These included work by
Areitio y Larrinaga (1874), a civil engineer, and by Mallada (1892,
1898), a mining engineer. Another mining engineer, Patac (1920)
published photographs of Autunian plant fossils, e.g. Callipteris
conferta (Sternberg) Brongniart (now Autunia conferta (Sternberg)
Kerp). Most meritorious is the work by the two mining engineers, Ruíz
Falcó and Madariaga Rojo (193133), who embarked on a full
description of Carboniferous and Lower Permian plant remains in
the collection of the Instituto Geológico y Minero de España. Un-
fortunately, this work did not progress beyond the sphenopsids. A few
plant remains from Autunian strata were gured by the mining
engineers and geologists Gavala (1927) and de la Rosa, de Alvarado
and Hernández Pacheco (1929), when dealing with Geological Survey
sheets involving the basins of Valdeviar (Sevilla province) and
Puertollano (Ciudad Real), respectively. Early work published abroad
includes the annotated list of oral remains from the Asturias by Zeiller
(1882) and, above all, the records published by Grand'Eury (1877).Itis
noted that Zeiller commented on collections made by Barrois (1882),
whose stratigraphic work on the Palaeozoic strata in Asturias,
Cantabrian Mountains, was a landmark in the geological exploration
of this intricate area. Grand'Eury obtained his Spanish collections from
mining engineers working the coal deposits at PeñarroyaBelmez
(province of Córdoba), Puertollano (Ciudad Real), Surroca (Girona),
and several localities in the Asturias. None of this material was
illustrated, but the lists published by Zeiller and Grand'Eury are well
worth taking into consideration. A few species were listed from a
Culmora at Barcelona by de Saporta (1891), who also failed to
provide illustrations.
In Portugal, the early work includes a memoir by Gomes (1865),
and a well illustrated paper on Dicranophyllum by de Lima (1888),
who also published several notes on the fossil oras of the Douro,
Buçaco and Santa Susana basins. De Lima, an important wine grower
in the Douro Valley, who became Prime Minister of the last
monarchist government of Portugal, prepared an atlas of Carbonifer-
ous plants in Portugal, which was eventually published by Teixeira
(1942a). Several small papers by C.J.F. Bunbury (in Ribeiro, 1853),
Feistmantel (1875), Geinitz (1867), Heer (1881), and Sterzel (1903)
should also be mentioned (see Wagner and Lemos de Sousa, 1983a,
for an account of the literature).
During the rst half of the XXth century the lists of fossil ora
obtained from different parts of the Pyrenees (Dalloni, 1910, 1930;
Schmidt, 1931) were provided by R. Zeiller (Paris), P. Bertrand
(University of Lille) and W. Gothan (Berlin). Other lists, with
determinations by W. Gothan, were published in papers by Simon
(1941, 1943, 1950) dealing with the early Westphalian coal basin of
Villanueva del Río y Minas (province of Sevilla) and the Autunian
occurrences at Valdeviar and San Nicolás del Puerto (Sevilla). The
Autunian is presently (this paper) assigned to the highest Carbonif-
erous. The only palaeobotanical paper with descriptions and illustra-
tions of Visean and Lower Permian(recorded as Stephanian)
elements of SW Spain (province of Badajoz) was published by
Hartung (1941). A special mention deserves the general summary of
Carboniferous and Permian records in Spain by Ruíz Falcó and
Madariaga Rojo (1941). This general overview was produced from
manuscript notes found in the inheritance of these authors, and it may
well be that these notes were accumulated gradually and did not
represent the up-to-date knowledge at the time of publication.
Whilst the knowledge of Carboniferous and Lower Permianoras
languished in Spain, an important volume of papers dealing with the
Portuguese oras was created by Teixeira from 1937 to 1943, culminating
in his comprehensive work O Antracolítico continental português
Plate XXII, gs. 1, 3; Plate XXIII, gs. 15; Plate XXIV, gs. 1, 2; Plate XXV, gs. 14)
Plate XXIX, gs. 1, 3; Plate XXX, gs. 18)
317
240 R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
(Teixeira, 1945). Teixeira continued publishing palaeobotanical papers
from 1946 to 1964, albeit more sporadically in the later years.
Jongmans published between 1950 and 1957 a number of
accounts, generally but not always accompanied by illustrations, of
his collecting trips in the Carboniferous of Spain, partly in collabora-
tion with B. Meléndez (University of Madrid), who organised his
eldwork, and also with R.H. Wagner, as his assistant. He also pub-
lished in collaboration with J. Menéndez Amor (Menéndez Amor and
Jongmans, 1952, 1954). Jongmans ranged widely, from the Cantabrian
Mountains (Asturias, León and Palencia) to Andalusia, and including
central Spain (provinces of Cuenca and Ciudad Real) as well as the
Pyrenees (province of Girona). His work was continued by R.H.
Wagner and his alumni (J.A. Knight, E. Iwaniw, C.J. Cleal, C. Álvarez-
Vázquez and M.P. Castro) and, indirectly, by M.C. Álvarez-Ramis
(University of Madrid). Meanwhile, different schools of palaeobota-
nical research in Germany (Münster W. and R. Remy), Belgium
(Brussels F. Stockmans and Y. Willière), and France (J.-P. Laveine, C.
Brousmiche and S. Beckary in Lille; J. Doubinger in Strasbourg; and J.
Broutin and C. Vozenin-Serra in Paris), and the Netherlands (H.W.J.
van Amerom in Heerlen) became involved in a documentation of the
Spanish Carboniferous oras which are, by now, quite well known. On
the other hand, the work in Portugal, so ably carried out by Teixeira in
the earlier part of the twentieth century, was virtually allowed to
lapse until R.H. Wagner and M.J. Lemos de Sousa produced several
papers in the 1980s.
The major contributions to the different Carboniferous areas in
Spain and Portugal will be quoted in the following sections dealing
with these areas systematically.
3. Geological overview
Carboniferous up to and including Stephanian B, and later
Stephanian deposits (including the Autunian) correspond to two
quite different episodes in the geological history of the Iberian
Peninsula. During most of Carboniferous times (Tournaisian to
Stephanian B) the geological conguration was such that a Tethyan
connection was apparent for northern Spain, whereas the southern
part of the Peninsula amalgamated slowly by means of strike-slip
faulting. The southeastern part of the present-day Iberian Peninsula
(including the Balearic Islands) seems to be a much later addition
(Fig. 1). Indeed, the Iberian Peninsula, as we know it today, did not
exist as such before the Cenozoic (Fig. 1), and the Iberian Massif only
came into being in latest Carboniferous times. References to an Iberian
microplate are inappropriate for the Carboniferous, and any attempt
to place the different parts of Iberiain an Atlantic context must take
into account their quite disparate palaeogeographic origins (which
are not always easy to determine within reasonable limits of accuracy).
Carboniferous reconstructions using the present-day outline of the
Iberian Peninsula are essentially uninformed. It is noted that no real
evidence exists for considering the various depositional areas in the
Iberian Peninsula as forming part of a Hercynian (Variscan) Chain,
which would imply a direct link with Northern and Central Europe.
This is not immediately apparent.
The geological history of the Palaeozoic rocks in Galicia, and in
most of the Cantabrian Mountains (NW Spain), is apparently linked to
that of Armorica (Brittany) and, perhaps, parts of Normandy and the
Vendée in France, but the exact linkage is still open to question. The
southeastern part of the Palaeozoic outcrop area in the Cantabrian
Mountains, the Pyrenees and part of the Iberian Chain belong to an
area which probably includes also the Montagne Noire in southern
France. The most complete Carboniferous record is found in the
Cantabrian Mountains, forming part of the CantabricAsturian arcuate
fold belt, with a history from Cambrian to Stephanian B (sensu Saint
Étienne). Compressional tectonics are in evidence for this arcuate fold
belt, as are a number of successive palaeogeographic congurations
which, despite afrmations to the contrary (e.g. Pérez Estaún et al.,
1988), do not seem to be related directly to the tectonic deformation.
Marine deposits were laid down as well as terrestrial, coal-bearing
strata throughout most of the Pennsylvanian (Bashkirian to Gzhelian;
also mid-Namurian to Stephanian). The marine faunas show a Tethyan
connection, right up to the Gzhelian (Merino-Tomé et al., 2006). In this
respect, there is a marked contrast with the classical Stephanian of the
Massif Central in south-central France, where intramontane, strike-slip
controlled basins developed in a non-marine setting, with only tenuous
indications for a marine connection (Schultze, 2009). Nonetheless, the
Stephanian oras of the Massif Central are quite similar to those of the
Cantabrian Mountains. Lowland areas must have been involved in both
areas. This means that the Stephanian oras of the Massif Central are
unlikelyto have lived at a high altitude, as someauthors have postulated
(e.g. Becq-Giraudon et al., 1996).
The CantabricAsturian arcuate fold belt, shown in structural out-
line with an undifferentiated Palaeozoic succession up to and includ-
ing the lower Asturian, and overlying lower (post-Leonian) and upper
(post-Asturian) Stephanian strata (Fig. 2), has a complex geological
history. This area contains a number of successive Carboniferous basins
with different palaeogeographic congurations (Figs. 35). Although
each of these basins suffered compressional tectonic deformation,
leading to a progressive tightening of the arcuate fold belt, the differ-
ent successive palaeogeographic congurations seem to obey different
underlying patterns which are, to some extent, independent from the
supercial tectonic structure. The successive compressional tectonic
deformations, which took place at different times during the Pen-
nsylvanian, were sharply delimited and of relatively short duration.
They alternated with much longer periods of sedimentation with syn-
sedimentary tectonics marked by vertical movements. Successive
compressional tectonic events produced increasingly tighter arcuate
shapes, the superposition of which shows that (a) the structural defor-
mation was supercial, i.e. not involving basement, and (b) individual
episodes of compressional tectonic deformation were short-lived and
areally restricted.
The rst palaeogeographic reconstruction attempted (Fig. 3) cor-
responds to Duckmantian, Bolsovian and early Asturian. At this time, a
tectonic hinterland to the NW corresponded to an area of uplift which
advanced SE-wards. It was fringed by a coal-producing, mixed marine
and terrestrial area which shifted gradually in southeastern direction,
encroaching upon a wholly marine area. On the southeastern side of
this marine basin an active mountain front associated with important
tectonic deformation in latest Langsettian times, produced a large
volume of uviatile conglomerates (Curavacas Formation) grading
laterally and upwards into submarine fan deposits and siliciclastic
turbidites (Savage, 1967; Colmenero et al., 1988). Megaoral remains
from the Curavacas Conglomerate Formation (uppermost Langset-
tianlower Duckmantian) occur in the more uviatile area of northern
Palencia and southern Cantabria, and also in coal-bearing deposits
(alternating with marine sediments) in parts of the Central Asturian
Coaleld (Fig. 2). Bolsovian and lower Asturian coal-bearing strata in
the Asturias (compare Wagner et al., 2002) are contrasted with mainly
marine deposits in northern Palencia (with sparse plant remains),
whilst lower Asturian valley-ll deposits are found in the Sierra de la
Demanda, representing a mountainous relief on the southeastern
basin margin.
The subsequent palaeogeographic conguration (Fig. 4) involves
mainly marine strata but also some terrestrial deposits with oral
remains. This corresponds to a Palentian Basin of late Asturian, Canta-
brian and early Barruelian ages. This basin only occupied the east
southeastern part of the arcuate fold belt. It accumulated up to 6000 m
of sediment. A substantial palaeotopography existed on the NW basin
margin, which shifted NW-wards in time (Iwaniw, 1985). A major
expansion of the Palentian Basin in mid-Cantabrian times involved
tectonic movements on a faulted NE basin margin producing exo-
olistoliths and, subsequently, turbidites and debris ows. For more
detailed accounts see Wagner and Winkler Prins (1985), Iwaniw
241R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
(1985) and Wagner and Martínez-García (1998). Contemporaneous
carbonate deposits were laid down on a more stable foreland area in
the Picos de Europa (Merino-Tomé et al., 2009).
Finally, another, more extensive basin came into being after con-
siderablecompressional tectonic deformation (Asturian Phase), in mid-
Barruelian times. Upper Barruelian, Saberianand Stephanian B
sediments were laid down in this basin. A nal tightening of the arcuate
fold belt took place afterwards, in (early) Stephanian C times. This is
reected in the outcrop distribution of tectonic outliers (coalelds) as
shown in Fig. 2. Coal-bearing strata were laid down in a strongly
subsidentpost-Asturian basin which anked a mountainous hinterland,
whilst marine deposition took place on a more stable foreland (Fig. 5).
Floral remains correspond to an alluvial plain with access to the marine
area on the western edge of the Palaeotethys. Successive transgressions
on this foreland area apparently provoked non-marine ooding leading
to lacustrine intervals in the alluvial plain succession. These commence
with Leaia bands (Fig. 6). The record of Stephanian oras in the various
tectonic remnants of the alluvial plain area is unrivalled. (See Fig. 6 for
the correlation of coaleld successions representing westerly onlap of
the alluvial plain succession.)
South and southwest of the large area linked to western shore of
the Palaeotethys, and which is represented by the Cantabrian Moun-
tains, Pyrenees, Sierra de la Demanda, as well as part of the Central
Mountain System (including the Serranía de Cuenca), lies the vast
Palaeozoic outcrop area of the Iberian Massif. Recent data (Martínez-
García, 2002; Wagner, 2004a; Martínez-García, 2006) have shown that
the Iberian Massif is an assembly of several different terranes which
came together by continental strike-slip faulting during Pennsylvanian
times. The basic picture was rst proposed by Martínez-García (1986)
but the more solid evidence with regard to the timing of major strike-
slip movements came from a detailed study of the strike-slip con-
trolled basin of Peñarroya in Sierra Morena (Wagner, 1999a). This
basin mimics the classic Ridge Basin generated alongside the San
Andreas Fault in the Cenozoic of California (Crowell, 1974). Further
Fig. 1. Map showing the different plant-bearing occurrences of Carboniferous age in the Iberian Peninsula (provincial name in brackets) (1a enlarged portion of the same showing
a more complete representation of localities in NW Spain; 1b enlarged portion showing localities in Sierra Morena, SW Spain, in more detail): 1. Vale de Corvos (Faro); 2. Santa
Susana (Setubal); 3. Guadalcanal (Sevilla); 4. Valdeviar (Sevilla); 5. Villanueva del Río y Minas (Sevilla); 6. Los Santos de Maimona (Badajoz); 7. Bienvenidas (Badajoz); 8. Casas de
Reina (Badajoz); 9. Berlanga (Badajoz); 10. Valdeinerno (Córdoba/Sevilla); 11. Cerrón El Hornillo/Puebla de los Infantes (Sevilla); 12. Virgen de Linares (Córdoba); 13. Benajarafe/
Matachel (Córdoba); 14. Guadiato (Córdoba); 15. PeñarroyaBelmezEspiel (Córdoba); 16. Serra de Rates, Bougado (Porto); 17. Casais/Alvarelhos (Porto); 18. Ervedosa (Bragança);
19. Douro (Porto/Viseu); 20. Buçaco (Aveiro); 21. Sierra de San Pedro (Cáceres); 22. Guadalmez (Ciudad Real); 23. Puertollano (Ciudad Real); 24. Henarejos (Cuenca); 25.
Valdesotos/Retiendas (Guadalajara); 26. Fombuena (Zaragoza); 27. Sierra de la Demanda (Burgos); 28. San Vitero (Zamora); 29. Arnao/San Juan de Nieva (Asturias); 30. Tineo
(Asturias); 31. Cangas de Narcea (Asturias); 32. Rengos (Asturias); 33. Tormaleo (Asturias); 34. El Bierzo (León); 35. Carballo (Asturias); 36. Carrasconte/Villablino (León); 37. La
Camocha (Asturias); 38. Monte Naranco (Asturias); 39. Puerto Ventana (León); 40. Teverga/San Emiliano (Asturias); 41. Quirós (Asturias); 42. Viadangos/Rodiezmo (León); 43. La
Magdalena (León); 44. CiñeraMatallana (León); 45. Central Asturian Coaleld (Asturias); 46. San Tirso (Asturias); 47. Canseco/Rucayo (León); 48. Tejerina/Ocejo de la Peña (León);
49. Prioro/Pando (León); 50. Sabero (León); 51. Guardo (Palencia/León); 52. Cueli (Asturias); 53. Sebarga (Asturias); 54. Gamonedo/Inguanzo (Asturias); 55. Arenas de Cabrales
(Asturias); 56. Valdeón (León); 57. Sotres (Cantabria); 58. Cucayo/Dobres (Cantabria); 59. Curavacas Formation (Palencia); 60. La Pernía (Palencia); 61. Castillería (Palencia); 62.
Peña Cildá (Palencia); 63. Barruelo/Redondo (Palencia); 64. Pico Cordel (Cantabria); 65. Puente Pumar (Cantabria); 66. Ibantelly (Navarra); 67. Collado de Izas (Huesca); 68. Béarnais
(French Pyrenees); 69. Aguiró (Lleida); 70. Malpás (Lleida); 71. Baró/Arcalís (Lleida); 72. Coll de Jou (Girona); 73. Ogassa/Surroca (Girona); 74. El Priorat (Tarragona).
242 R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
evidence of strike-slip faulting has been forthcoming from Portugal
(Chaminé et al., 2003; Pinto de Jesús and Lemos de Sousa, 2003). In the
BadajozCórdoba fault zone, along which the Peñarroya Basin is
situated, it is noted that Visean and Serpukhovian strata overlie
strongly deformed, metamorphic Lower Palaeozoic and Precambrian
rocks. This is a Caledonian setting reminiscent of northern England.
Locally complex tectonic structures in the Carboniferous strata are due
to transpression associated with the strike-slip movements. It seems
quite likely that long-lasting strike-slip movements were instrumen-
tal in bringing part of Caledonian Europe to the Iberian Peninsula. This
Fig. 1 (continued).
243R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
Bierzo
N
Rengos
Carballo
Villablino
Tineo
Mieres
Central
Asturian
Coalfield
Puerto
Ventana
Canseco Rucayo
Sabero
La Robla
La Magdalena
C.M.
León
Ferroñes
Gijón
fault
Fo
Se
Ponga
area
Beleño
Lebeña
Valdeón
Picos fault
Potes
Palentian
area
Puente
Pumar
Pico
Cordel
Barruelo
Riaño
Cervera
de Pisuerga
fault
Carrionas
Guardo
Cabo Peñas
Ribadesella
Llanes
Arenas de
Direction of thrusting
Faults
C.M. = Ciñera-Matallana
Fo = Fontecha
Se = Sebarga
P.C. = Peña Cildá
Paleozoic undifferentiated
(incl. Precambrian)
Post-Leonian deposits
Post-Asturian deposits
Post-orogenic cover
Ven tani ella
Cabrales
Ca
Oviedo
Rubias
Punta
Arnao
Ca = Carrasconte
Avilés Mar Cantábrico
(Bay of Biscay)
0102030
40 50 km
Tormaleo
Cangas
Gamonedo
Fig. 2. Map showing the distribution of Stephanian outcrops in the Cantabrian Mountains. The Early Stephanianbasin established after the Leonian tectonic deformation event
involves upper Asturian, Cantabrian and lower Barruelian (substages). The Late Stephanianbasin (after the Asturian movements) involves upper Barruelian, Saberianand
Stephanian B (substages). The two successive Stephanian basins are mutually unconformable. After Wagner and Castro (in press).
Fig. 3. Palinspastic palaeogeographic reconstruction for NW Spain in Duckmantian to early Asturian times showing distribution of different facies areas. Oviedo and Sierra de la
Demanda are shown for reference.
244 R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
Fig. 4. Palinspastic palaeogeographic reconstruction for NW Spain in late Asturian to early Cantabrian times showing distribution of facies areas. A gradual expansion of the early
post-Leonian basin was followed in mid-Barruelian times by a sudden, tectonically induced, major expansion of the basin in both WNW- and E-ward directions. N.B. The Asturian
area is indicated for reference in the position it occupied before it swung into an EW strike.
Fig. 5. Palinspastic palaeogeographic reconstruction for NW Spain in Stephanian B times showing facies areas (after Wagner and Castro, in press). See Fig. 2 for reference.
245R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
would being as a corollary that the composition of Mississippian oras
in the area of the Iberian Massif (Sierra Morena) would be similar to
that in NW Europe. Terrane movement was indeed from the NNW.
The more recent information on the geological history of the
present-day Iberian Peninsula has shown that the classic subdivision
into South Portuguese, Ossa-Morena, LusitanianAlcudian, Galaico-
Castillian, West AsturianLeonese, and Cantabrian zones, as proposed
by Lotze (1945), needs to be modied in the light of plate tectonics.
This includes strike-slip faulting as an important component. By no
means can the Iberian Peninsula be taken as an entity in Carboniferous
times, and the various palaeogeographic reconstructions proposed in
the literature would do well to take this fact into account.
A marked change in tectonic regime occurred in latest Carbon-
iferous times. Stephanian C (equals early Autunian) and mid to late
Autunian (latest Pennsylvanian) times initiate what is, on the whole,
the Permian tectonic history of the Iberian Peninsula. This is charac-
terised by extensional tectonics and widespread volcanic activity.
The CantabricAsturian arcuate fold belt of NW Spain, together with
the Pyrenees and part of the present-day Central Mountain Sys-
tem, reached its nal stage of compressional tectonic deformation
in Stephanian C times. Likewise, the major strike-slip fault activity in
the South Portuguese, Ossa-Morena and East LusitanianAlcudian
areas (Lotze, 1945) ceased during the early Stephanian C, when total
assembly was achieved for the Iberian Massif, and the core of the
Iberian Peninsula was constituted for the rst time. The tensional
tectonic regime of latest Carboniferous and Early Permian times
matches that of western Europe in general, and so does the volcanic
activity. Generalised changes in the underlying plate tectonic mech-
anism must be assumed. Coincidentally, the Late Palaeozoic Ice Age
(mainly corresponding to Pennsylvanian times) was on the wane
throughout the Autunian and Early Permian, and this produced
environmental changes which affected oral composition and the
relative proportions of hygrophile and mesophilexerophile elements
in the areas of different pluvial regimes. Major palaeogeographic
changes linked to a change in tectonic regime were thus accompanied
by climatically induced environmental changes.
With the changeover to a tensional tectonic regime a number of
small sedimentary basins came into being, mostly of a terrestrial
Fig. 6. Correlation of the stratigraphic successions of the Sabero, CiñeraMatallana, La Magdalena and Carrasconte/Villablino coalelds in northern León, showing onlap on the
western basin margin. Horizontal scale different to vertical one. After Wagner and Castro (in press).
246 R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
nature. In latest Carboniferous and Permian times the area corres-
ponding to most of the present-day Iberian Peninsula (i.e. excluding
its southwestern part), formed part of a general West and Central
European region.
The geological record of Carboniferous strata in the Iberian Peninsula
and, more particularly, that of the CantabricAsturian arcuate fold belt,
where terrestrial oras occur in the same successions as varied marine
faunas, has allowed rening the international correlations (Fig. 7). In
particular, this refers to the positioning of the Yeadonian/Langsettian
(Namurian/Westphalian) boundary near the base of the Moscovian (as
explained most recently in Kullmann et al., 2007). It will be noted that
the chronostratigraphic correlation chart issued on behalf of the IUGS
Subcommission on Carboniferous Stratigraphy by Heckel (2004) and
Heckel and Clayton (2006) shows the base of the Moscovian in a higher
position, thus squeezing the various Namurian subdivisions into a
smaller time slot than seems to be warranted. Fig. 7 also shows the
inclusion of middle to upper Autunian in the highest Carboniferous
(Gzhelian). In Heckel's chart the Autunian was shown as a stage
following upon the Stephanian. The reasons for our inclusion of the
Autunian (sensu stricto) as the highest substage of the Stephanian
Regional Stages are explained in Section 7.
4. Carboniferous oras in different parts of the Iberian Peninsula
All the different ages of the Carboniferous Period are represented
in the megaoral record of the Iberian Peninsula. The various outcrop
zones
middle to upper
Autunian
(= lower Autunian)
"Saberian" Alethopteris zeilleri
Barruelian Lobatopteris lamuriana
Lobatopteris vestita
Linopteris obliqua
Bolsovian
Lonchopteris rugosa /
Alethopteris urophylla
Langsettian Lyginopteris hoeninghausii /
Neuralethopteris schlehanii
Marsdenian
Kinderscoutian
Alportian
Asbian
Holkerian
Hastarian Adiantites
M I S S I S S I P P I A N
Ivorian
Chadian
Arundian
Brigantian
Pendleian
VISEAN
UPPER
MIDDLE
Triphyllopteris
Chokierian
Neuralethopteris larischii /
Pecopteris aspera
Arnsbergian
Lyginopteris stangeri
SYSTEM
REGIONAL
STAGES
WESTPHALIAN
Asturian
Duckmantian
C A R B O N I F E R O U S
SUB-
SYSTEM
Cantabrian
Stephanian B
KASIMOVIAN
Odontopteris cantabrica
Yeadonian
GZHELIAN
SERIES
MOSCOVIAN
BASHKIRIAN
Sphenophyllum angustifolium
Annularia spicata
P E N N S Y L V A N I A N
UPPER
MIDDLE
LOWER
GLOBAL STAGES
SERPUKHOVIAN
VISEAN
TOURNAISIANLOWER
Paripteris linguaefolia
STEPHANIAN
(Spathulopteris Subzone )
Triphyllopteris
(Lepidodendropsis Subzone)
TOURNAISIAN
Neuropteris antecedens
NAMURIAN
Autunia conferta
Lyginopteris larischii
Megafloral
SUBSTAGES
Stephanian C
(W. Europe)
Fig. 7. Global chronostratigraphic subdivisions of the Carboniferous as correlated with the regional European stages and substages (after Heckel, 2004 and Heckel and Clayton, 2006,
with modied position of Westphalian base, after Kullmann et al., 2007). Megaoral zones after Wagner (1984), with adjustment of the Asturian (=Westphalian D)/Cantabrian
boundary as in Wagner and Winkler Prins (1985), and the addition of a new Annularia spicata Zone.
247R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
areas for the palaeobotanical record are indicated in Figs. 1,1a, 1b, and
the different geological ages for these areas are stated in Fig. 8.
The enormously varied geological history of the component parts
of what presently constitutes the Iberian Peninsula, means that the
Carboniferous oral record of the different areas must be discussed
accordingly.
The variousrecords will be dealt within stratigraphicorder. It is clear
that only lists of taxa can be presented in a general account like the
present one, and that taxonomic problems may only be referred to
occasionally (see Appendix). The emphasis is on stratigraphic occur-
rence, oral composition, and general comparisons with regard to
the classic oral assemblages of northern Europe. The few records of
Mississippian and earliest Pennsylvanian oras are listed in the text,
whereas the more comprehensive records of later Pennsylvanian ages
(Langsettian to Autunian) are summarised in Tables 110.Numbers
have been assigned to the different outcrop areas (Fig. 1) and the same
numbers are referred to in brackets where the relevant areas
are discussed in the text and mentioned in the tables. Equivalents
to global stages are given in Fig. 7, which are correlated with the
West European regional stages and substages as well as the megaoral
zones. A full selection of taxa is represented, but a complete
representation must be left to more detailed papers.
4.1. Mississippian oras
The outcrop areas for Mississippian oral remains are widely
scattered, and the locations are generally quite different to their
original palaeogeographic position. Best preserved are the upper
Tournaisian oras which appear identical to those elsewhere in
Plate I.
Fig. 1. Psygmophyllum sp. (×3). Outcrop no 11, Puebla de los Infantes (loc. 9909), upper Tournaisian.
Fig. 2. Triphyllopteris collombiana Schimper (×3). Outcrop no 10, Valdeinerno (loc. 2813), upper Tournaisian.
Fig. 3. Sphenopteridium pachyrrhachis (Göppert) Schimper (×3). Outcrop no 10, Valdeinerno (loc. 4343), upper Tournaisian.
Fig. 4. Eusphenopteris sp. (×3). Outcrop no 10, Valdeinerno (loc. 4343), upper Tournaisian.
SUBSTAGES
(W. Europe)
Autunian
(middle to upper)
Stephanian C
(= lower Autunian)
"Saberian" El Bierzo (34), Puerto Ventana (39), La Magdalena (43), Ciñera-Matallana (44), Canseco/Rucayo (47),
Sabero (50)
Barruelian Sabero (50), Barruelo/Redondo (63), Valdeón (56)
Santa Susana (2), Guardo (51), La Pernía (60), Aguiró (69)
Ervedosa (18), Riosa/Olloniego (45a) and Aller/Nalón (45b) areas of the Central Asturian Coalfield, Castillería
(61), Sierra de la Demanda (27)
Bolsovian Serra de Rates (16), Casais-Alvarelhos (17), Riosa/Olloniego (45a) and Mina Inés (45c) areas of the Central
Asturian Coalfield, Prioro/Pando (49)
Langsettian
Vale de Corvos (1), Villanueva del Río y Minas (5), western part of Peñarroya-Belmez-Espiel Coalfield (15),
La Camocha (37), Monte Naranco (38), Teverga/San Emiliano (40), Quirós (41), Cucayo/Dobres (58),
Cardaño de Arriba/Triollo (59)
STAGESW E S T P H A L I A N
Asturian
Duckmantian
Cantabrian
S T E P H A N I A N
Stephanian B
Tejerina/Ocejo de la Peña (48), Guardo (51), Gamonedo/Inguanzo (54), Barruelo/Redondo (63)
Plant-bearing areas (numbers refer to map - Fig. 1)
Douro (19), Puertollano (23), Puente Bergueres, San Tirso (46), Sotres (57), Ibantelly (66), Malpás (70),
Ogassa/Surroca (73)
Guadalcanal (3), Valdeviar (4), Buçaco (20), Valdesotos/Retiendas (25), Rillo de Gallo, Fombuena (26),
Cueli (52), Baró/Arcalís (71), Coll de Jou (72), Sauquillo (Soria)
Punta de la Rubia, Ferroñes, Arnao (29), Tineo (30), Cangas del Narcea (31), Rengos (32), Tormaleo (33),
El Bierzo (34), Carballo (35), Villablino (36), Sebarga (53), Cabrales (55), Peña Cildá (62), Pico Cordel (64),
Puente Pumar (65), Henarejos (24)
Central and eastern part of Peñarroya-Belmez-Espiel Coalfield (15), Sierra de San Pedro (21), Curavacas
Formation in Los Cintos (59)
Fig. 8. Plant-bearing occurrences identied by name and number as shown in Fig. 1, with reference to the West European stages and substages.
248 R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
249R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324
Europe. Given the rather limited number of species involved (for
adpressions), this may well mean a fairly high ecological tolerance.
4.1.1. Tournaisian (Plate I,gs. 14; Plate II,gs. 1, 2)
Lower Tournaisian plant remains may be represented in a
turbiditic basin which has been fragmented by later tectonic move-
ments associated with strike-slip faulting (Martínez-García, 2006).
Although indeterminate axial remains are the norm, in one locality
(San Vitero (28), in Zamora province) a presumed isoetalean fertile
apex (Eoacanthocarpus) and a small fragment of Stigmaria coides
(Sternberg) Brongniart have been identied. These were illustrated
by Montero (2008), who left open the possibility that uppermost
Famennian might be involved.
Much more signicant late Tournaisian to earliest Visean plant
assemblages have been recorded from three different localities in SW
Spain, viz. a small, intramontane, pull-apart basin at Valdeinerno
(10), near Fuenteobejuna in Córdoba province (Jongmans and
Meléndez, 1950; Wagner, 1978a; Wagner et al., 1983a; Wagner,
2001); a terrestrial interval in a marine succession at Cerrón El
Hornillo, near La Puebla de los Infantes (11) in the province of Sevilla
(Laveine in Robardet et al., 1986, and an additional collection in the
Centro Paleobotánico, Córdoba); and also a 60 km long strip with
upper Tournaisian coal-bearing strata at BenajarafeMatachel (13)
(Garrote and Broutin, 1979; Wagner et al., 1983a). The latter are in
stratigraphic continuity with marine and volcanic deposits of Visean
age (see Coquel in Wagner et al., 1983a). All three localities are in the
Ossa-Morena Zone of Sierra Morena, Andalusia/Extremadura. The
geological context suggests that Valdeinerno is an individual basin,
whereas the other two occurrences are remnants of one or more
larger basins torn apart by strike-slip faulting of some magnitude,
during Pennsylvanian times. It seems likely that their original position
was quite remote from the present one, probably at a thousand or
more kilometres to the NNW, i.e. close to the British Isles. Indeed, the
geological relationships suggest that part of Caledonian Europe is
involved. The composition of these oras is very similar to that of
oras of the same age in France and Germany. Most characteristic
elements are as follows: Triphyllopteris collombiana Schimper (Plate I,
g. 2), Fryopsis frondosa (Göppert) Wolfe, Neurocardiopteris broilii
Lutz (Plate II,g. 1), Sphenopteris cf. foliolata Stur, Sphenopteridium cf.
dissectum (Göppert) Schimper (Plate II,g. 2), Sphenopteridium
pachyrrhachis (Göppert) Schimper (Plate I,g. 3), Rhodeopteridium
stachei (Stur) Purkyňová, Paracalathiops cf. plauensis (Gothan)
Wagner, Rhacopteris cf. robusta Kidston, cf. Rhacopteris paniculifera
Stur, Eusphenopteris sp. (Plate I,g. 4), Psygmophyllum sp. (Plate I,
g. 1), Lepidodendropsiscf. hirmeri Lutz, Lepidodendropsiscf.
vandergrachtii Jongmans, Gothan & Darrah, Eoacanthocarpus fei-
litzschianus (Gothan & Schlosser) Daber, cf. Archaeosigillaria vanuxemi
Kidston, Archaeocalamites scrobiculatus (Schlotheim ex Sternberg)
Zeiller, and Sphenophyllum saxifragaefolioides Leyh.
4.1.2. Visean (Plate II,gs. 3, 4; Plate III,gs. 15a; Plate IV,g. 1)s
Floral assemblages of Visean and early Namurian (Serpukhovian)
ages occur in several small, isolate outcrop areas in a NWSE striking
terrane, the BadajozCórdoba fault zone, in the province of Córdoba
(Andalusia) and part of neighbouring Badajoz (Extremadura). The
Visean and preceding uppermost Tournaisian strata overlie previously
deformed, partly metamorphic, Precambrian and Lower Palaeozoic
rocks unconformably. These areas of Mississippian outcrop are
probably remnants of one or more basins disrupted by strike-slip
faulting of Pennsylvanian age, but their geological reconstruction
may be difcult to achieve. Mention has been made already of the
BenajarafeMatachel strip in BadajozCórdoba where Visean strata
dated on miospores follow upon upper Tournaisian coal-bearing
deposits (Wagner et al., 1983a).
About 5 km NE of Córdoba, near the chapel (Ermita) of Virgen de
Linares (12), a small Visean ora has been found in generally marine
siliceous shales, containing Neurocardiopteris broilii (specimen gured
in Wagner, 2001,g. 51), Sphenopteridium bidum (Lindley & Hutton)
Benson (Plate III,g. 2), Paracalathiops sp., Lepidodendronlossenii
Weiss, Lepidostrobus sp., and Archaeocalamites sp.
The best studied outcrop area, containing mainly shallow marine
deposits of Visean age, but also a plant-bearing terrestrial interval, is at
Los Santos de Maimona (6), near Zafra, Badajoz province (Rodríguez
et al., 1992). From this area, Jongmans (1956) gured Lepidodendron
lossenii (see also Plate III,gs. 5, 5a of the present paper), Archaeo-
calamites scrobiculatus, and Cyclostigmazafrensis Jongmans (see
taxonomic comments in the Appendix). A single specimen of
Adiantites rhombifolius (Daber) Wagner, Coquel & Broutin (Plate IV,
g. 1) may be added, as well as Fryopsis frondosa (Plate III,g. 1),
Rhacopteris inaequilaterale Göppert, cf. Rhacopteris dichotoma Kidston
(Plate III,g. 3), Sphenopteridium crassum (Lindley & Hutton) Kidston,
Sphenopteridium dissectum (Plate III,g. 4), Sphenopteridium cf.
schimperi (Göppert) Schimper (Plate II,g. 3), Cardiopteridium
nanum (Eichwald) Nathorst, and Sphenophyllum sp.
Poorly diversied Visean oras have been recovered from mixed
marine and terrestrial strata in unconformable outliers in the Badajoz
Córdoba fault zone, occurring at Bienvenidas (7), Casas de Reina (8),
and Berlanga (9) (all Badajoz province) (Hartung, 1941; Jongmans,
1956; Wagner et al., 1983a). Lepidodendronlossenii and Archaeo-
calamites scrobiculatus are the most common elements, but Genselia
rarinervis (Read) Knaus (Plate II,g. 4) (previously recorded as
Adiantites cf. rhombifolius by Wagner et al., 1983a), occurs as well.
Plate II.
Fig. 1. Neurocardiopteris broilii Lutz (×3). Outcrop no 11, Puebla de los Infantes (loc. 9909), upper Tournaisian.
Fig. 2. Sphenopteridium cf. dissectum (Göppert) Schimper (×3). Outcrop no 10, Valdeinerno (loc. 4343), upper Tournaisian.
Fig. 3. Sphenopteridium cf. schimperi (Göppert) Schimper (×3). Outcrop no 6, Los Santos de Maimona (loc. 2896), Visean.
Fig. 4. Genselia rarinervis (Read) Knaus (×3). Outcrop no 9, Berlanga (loc. 4232), Visean.
Plate III. (see on page 252)
Fig. 1. Fryopsis frondosa (Göppert) Wolfe (×3). Outcrop no 6, Los Santos de Maimona (loc. 6887), Visean.
Fig. 2. Sphenopteridium bidum (Lindley & Hutton) Benson (×3). Outcrop no 12, Virgen de Linares (loc. 9819), Visean.
Fig. 3. cf. Rhacopteris dichotoma Kidston (×3). Outcrop no 6, Los Santos de Maimona (loc. 6887), Visean.
Fig. 4. Sphenopteridium dissectum (Göppert) Schimper (×3). Outcrop no 6, Los Santos de Maimona (loc. 6887), Visean.
Fig. 5. Lepidodendronlossenii Weiss (×6). Outcrop no 6, Los Santos de Maimona (loc. 5300), Visean.
Fig. 5a. Part of the same specimen ×9.
Plate IV. (see on page 253)
Fig. 1. Adiantites rhombifolius (Daber) Wagner, Coquel & Broutin (×3). Outcrop no 6, Los Santos de Maimona, Visean. Coll. Instituto Geólogico y Minero, Madrid.
Fig. 2. Lepidodendronvolkmannianum Sternberg (×1). Outcrop no 14, El Couce, Guadiato (loc. 6016), Serpukhovian.
Fig. 2a. Another part of the same specimen ×3.
250 R.H. Wagner, C. Álvarez-Vázquez / Review of Palaeobotany and Palynology 162 (2010) 239324