Review of Palaeobotany and Palynology 162 (2010) 231–238

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Review of Palaeobotany and Palynology

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Devonian floral assemblages and plant megafossils from the Iberian Peninsula:A review

Angel Montero a,⁎, Carmen Dieguez b

a I.M.G.E.M.A. Jardín Botánico de Córdoba, Avda. de Linneo s/n, 14004, Córdoba, Spainb Departamento de Paleobiología, Museo Nacional de Ciencias Naturales-CSIC, José Gutiérrez Abascal 2, 28006 Madrid, Spain

⁎ Corresponding author.E-mail address: paleo.amontero@jardinbotanicodeco

0034-6667/$ – see front matter © 2010 Elsevier B.V. Aldoi:10.1016/j.revpalbo.2010.02.008

a b s t r a c t

a r t i c l e i n f o

Article history:Received 29 July 2009Received in revised form 10 February 2010Accepted 17 February 2010Available online 26 February 2010

Keywords:Devonianfloral assemblagesIberian Peninsula

This paper presents a review and critical analysis of the literature on Devonian floras of the Iberian Peninsula.Although the known outcrops of Devonian strata in the Iberian Peninsula are marine, in some cases, a fewfragmentary remains of vascular plants are associated with faunal remains. Records include largelyspecimens from the Lower Devonian of Barrancos (Alentejo, Portugal) and the Upper Devonian of Sierra deHornachos in Badajoz province, southwest Spain; the remainder consists of drifted plant fragments fromscattered sites in the Iberian Peninsula ranging in age from Lochkovian to Upper Devonian–EarliestCarboniferous. The vegetation inferred for the Lower Devonian of the Iberian Peninsula is mainly based onpalynological data and corresponds to herbaceous types characterised by bryophytes, rhyniophytes(Horneophyton, Cooksonia, Rhynia), trimerophytes (Psilophyton, Pertica and Hostinella), primitive lycophytes(Drepanophycus) and incertae sedis such as Nothia and Chaleuria, all flora that developed near the coast inlow-lying and, at least periodically, wet areas. In the Middle Devonian, two vegetation strata can berecognised: herbaceous (Psilophyton) and semi-arboreal (Cladoxylales). Although three levels of vegetationin the Upper Devonian, have been described from outcrops worldwide, the scarce available data from theIberian Peninsula only indicate an arborescent lycopsid vegetation and species with uncertain botanicalaffinity such as Sphenopteridium keilhauii Nathorst.

rdoba.com (A. Montero).

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1. Introduction and historical overview

A variety of circumstances have led to the Devonian beingconsidered one of the most crucial periods on Earth for thedevelopment of terrestrial flora, since it saw not only the diversifi-cation and expansion of tracheophytes, which had already begun inthe late Silurian, but also other, frequently forgotten but equallyimportant events, such as the formation of soils (weathering andpedogenesis) and a change in the concentration of atmospheric CO2

with its important effects on ecosystems and global climates(Beerbower et al., 1992; Retallack, 1997; Berner, 1997, 2001; Algeoet al., 2001; Driese and Mora, 2001).

Devonian floras have been studied since the mid nineteenhundreds. To our knowledge the first remains were cited by Miller(1858) and Salter (1858) from Caithness (Scotland). However, thefirst mention of a complete Devonian flora corresponds to that of theLower Devonian in the Gaspé, Peninsula of Quebec (Canada) with thedescription and establishment of the genus Psilophyton (Dawson,1859). Subsequently, the floras of Maine and New York weredescribed (Dawson, 1862, 1863). Afterwards, the knowledge of

Devonian palaeobotany increased substantially so much so thatArber (1921) was able to mention the respective floras of Scotland,England, Southern Ireland, Norway, Germany, Belgium, Ukraine, theCzech Republic, USA, Canada and Australia. However, apart from itsbeing one of the oldest floras known (Early Devonian), the diversityand extraordinary state of preservation make the fossil flora of theRhynie Chert in Aberdeenshire (Scotland) a reference in the study ofthe Devonian period. The flora from this locality has been extensivelystudied. Kidston and Lang (1917) provided the first description of apetrified plant from the Psilophyton flora: Rhynia gwynne-vaughaniKidston & Lang. In subsequent studies (Kidston and Lang, 1920a,b,1921a,b), the same authors widened the number of species describedbased on the material with preserved anatomical structures (e.g.Rhynia major Kidston & Lang, Hornea lignieri (Kidston & Lang)Barghoorn & Darrah, Asteroxylon mackiei Kidston & Lang) anddescribed in what today may be considered the taxonomic character-istics of the deposit. From these pioneer works to date, theinformation obtained from successive studies on the Rhynie Cherthas documented: ancient biodiversity (cyanobacteria, fungi, algae,lichen, macroplants and animals), morphology and life history offungi and land plants, and the interactions and associations amongthe organisms of this ancient ecosystem (Remy et al. 1994; Edwardset al., 1998; Roth-Nebelsick et al., 2000; Kerp et al., 2001; Taylor andKrings, 2005; Taylor et al., 2005, 2006). These studies increase the

Fig. 1. Devonian palaeogeography. Dotted areas correspond to land masses (based on Scotese and McKerrow, 1990).

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knowledge about the biodiversity of Devonian floras worldwide, andwidened the geographical distribution of the representative genusand species.

Similar studies in the Iberian Peninsula are scarce and limited toscattered finds of drifted plant remains, deposited in marine strataandmostly recovered from outcrops at coastal areas. Hermite (1879)records Archaeocalamites renaultii Hermite and Sphenophyllummaresii Hermite from Menorca in the Balearic Islands. The formerwas assigned to the Lower Devonian, and the latter to the MiddleDevonian. Although the Devonian is present in Menorca it is morethan likely that the fossils reported by Hermite belong to theMississippian, a Subsystem which is widely represented in thisIsland. The whereabouts of these unfigured specimens are unknown.Nery Delgado (1908) mentioned very fragmentary specimens in fewsites of the Barrancos Region (Alentejo, Portugal) and Teixeira(1951) figured and described few species from reputedly LowerDevonian strata at Barrancos. Also in this case the whereabouts ofspecimens cited and figured by Teixeira are currently unknown.

Gómez de Llarena (1950, 1956) figured and identified a fewmacroremains from sediments in the western Pyrenees of Navarra,and dated it as Mid-Devonian age.

Hartung (in Kullmann, 1960) identified Taeniocrada sp. from LowertoMiddle Devonian strata in northern Palencia, CantabrianMountains(Palentian Domain). A later revision (Kullmann, pers. comm., 2008)suggests that these may have been ichnofossils.

Very fragmentary remains, from NE Palencia of early Famennian age(Weyer & Blumenstengel, in Wagner et al., 1984: 17–19) were identifiedby Wagner (1966).

Teixeira (1970) mentioned indeterminable plant remains ofpresumed Silurian age from Guadramil, near Bragança, Portugal. Thewhereabouts of these specimens are unknown, and the age wassubsequently corrected (Teixeira and Pais, 1973) as Late Devonian orEarly Carboniferous.

In the 1970s several papers dealing with the Palaeozoic rocks ofthe province Zamora (Spain), near the Portuguese border recordedthe presence of fossil plant remains in strata overlying a Silurian/Devonian succession (Martínez García, 1972, 1973; Teixeira and Pais,1973). The whereabouts of these specimens are unknown.

Adeposit of driftedplant remainswas found in theprovinceof Badajozin SWSpain, not far from the Portuguese border. The long list of taxa fromthis deposit given by Álvarez-Ramis (1981) was accompanied byillustrations of axial fragments which recalled similar assemblages atBarrancos (Alentejo, Portugal). Originally assigned a Later Devonian ageby Álvarez-Ramis (1981), this opinion was subsequently corrected toeither Late Devonian or Early Carboniferous (Álvarez Ramis in HerranzAraujo, 1985: 793).

In a later paper, the same author (Álvarez-Ramis, 1988) describedmacroremains attributable to Sciadophyton steinmanni Kräusel &Weyland from another locality of the province Badajoz (Spain), towhich a Gedinnian age was attributed. The whereabouts of thismaterial are unknown.

Finally, Pardo Alonso (1997) mentioned the presence of Lepido-dendraceae among a mass of unidentifiable plant fragments from ahorizon in between upper Emsian and upper Givetian at Guadalmez(Ciudad Real province, Spain).

Recently a report on the Devonian vascular plants in the IberianPeninsula has been carried out (Montero, 2008).

This shortcomingmay be due to the lack of knowledge (or the non-existence) of continental outcrops of this age, given the palaeogeo-graphy of the Iberian Peninsula during the Devonian (Figs. 1 and 2).

2. Material and methods

In the present paper an exhaustive review of Devonian macroflorasfrom the Iberian Peninsula (Spain and Portugal) has been carried out viathe study of previous literature on Devonian vascular plants (Table 1).Wherever possible the specimens were studied (see Table 2 forrepositories).

After degagément, the examination of available specimenswas donewith a Stemi 2000-C Zeiss stereomicroscope. Hand samples werephotographed under low angle incident light with a Canon Eos 300 Ddigital camera.

A map was created in which the different outcrops and plant fossillocalities in the Iberian Peninsula have been pointed out. In addition tothis, a map with the distribution of continental masses duringDevonian timeswas redrawn based on Scotese andMcKerrow (1990).

Fig. 2. Distribution of Devonian outcrops in the Iberian Peninsula showing the position of plant fossil localities as reported in the literature, and discussed in the present paper.1. Navarrese Pyrenees (Gómez de Llarena, 1950, 1956). 2. Northern Palencia (Kullmann, 1960). 3. NE Palencia (Wagner, 1966). 4. Zamora (Teixeira and Pais, 1973). 5. Badajoz(Álvarez-Ramis, 1981, Álvarez-Ramis, 1995). 6. Alentejo (Teixeira, 1951). 7. Ciudad Real (Pardo Alonso, 1997).

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3. Palaeogeographical setting

The Devonian was a period of intense tectonic activity, duringwhich Caledonian and Variscan orogenies produced changes in thecontinental land masses and determined the evolution of Europethrough the Palaeozoic. In the Lower Palaeozoic, the continent ofLaurussia was formed as a result of the collision of the Laurentia,Baltica and Avalonia and Gondwana plates to the south, from whichsmall land masses separated and later amalgamated (Cocks, 2000).The palaeocontinent Laurentia was represented in Europe by Scotlandand Ireland, Spitsbergen and the highest layers of the NorwegianCaledonides. For its part, Baltica included more than half of present-day Europe. From the uppermost Arenig onwards, a series of terranesof Gondwana (Avalonia, Armorica, Perunica and Iberia, whichincluded part of the south of France, Sardinia and most of the IberianPeninsula, with the exception of the sub-Portuguese zone) broke offfrom the European side of Gondwana and changed position, movingtowards the north and gradually approaching Laurentia. Although theexact relationship between Iberia, France, Germany and Bohemia isunclear, palaeomagnetic data from the west of France (Perroud andvan der Voo, 1985) and Spain (Perroud 1983) indicate that these areaswere situated in high latitudes and adjacent to the northern part ofGondwana (Scotese and McKerrow, 1990). Faunal data suggests thatareas of the Iberian Peninsula formed part of the north of Gondwana(Robardet et al., 1990; Quesada, 1991; Robardet, 2002, Robardet andGutiérrez-Marco, 2004) while Gondwana and Laurussia were sepa-rated by the Rheic Ocean. Together, the palaeomagnetic and faunisticdata suggest that this ocean was not very large during the Devonian

(Scotese andMcKerrow, 1990) and that it became enclosed during theUpper Devonian.

Most continental masses were set around the tropical sea Proto-tethys, where a warm greenhouse-type climate prevailed (Becker andKirchgasser, 2007). Throughout this period the continents weregradually flooded and large epicontinental seas, which have nomodern-day equivalent, were formed.

4. Lower Devonian

Devonian macrofloras were recorded from the Iberian Peninsulasince Nery Delgado (1908, p. 175, 209), found atMonte das Eiras Altas,Volta dos Nogais and in Noudar (Barrancos, Alentejo, Portugal)fragmentary remains of axes which were considered to be from thehigher part of the Gotlandian (Silurian). A subsequent study of thisflora (Teixeira, 1951, 1982a, 1982b) resulted in attributions toDrepanophycus-type axes, resembling D. spinaeformis Göeppert, andfragments of narrower, smooth dichotomous axes dubiously assignedto Psilophyton and Hostimella (=Hostinella). The fossils were found inlevels of lutites and greywackes of the Terena Formation (Barrancos).Based on the flora found in this work, Teixeira (1951, 1982a,b)assigned these sediments to the Lower Devonian and postulated ashallow coastal environment.

Study of the palynofloras of successive horizons of the TerenaFormation (Lochkovian–Emsian), and of the Russiana (Pragian) and“Xistos Raiados” (Pragian–Emsian) Formations (Pereira, et al., 1999;Oliveira et al., 2007) confirms Teixeira's dating and provides check-lists of sporae dispersae from each horizon. Taking into account the

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correlation between species of dispersed and “in situ” Devonianspores, carried out by several authors in an attempt to determine thebotanical affinity of dispersed spores (Allen, 1980, Gensel, 1980,Edwards and Richardson, 1996, Wellman, 2004), the palynologicalassociations found in these Portuguese formations indicate, mostlikely, such megafossil taxa as the following: several species ofrhyniophytes (Horneophyton, Cooksonia) accompanied by trimero-phytes (Psilophyton, Pertica), and incertae sedis elements like Nothiaand Chaleuria.

In Spain, the plant megafossils collected correspond to bryophyte-like plants and specimens attributed to the species Sciadophytonsteinmanni Kräusel and Weyland. In the first case, the only specimencollected comes from Bruguers (Gavà, Barcelona) and shows amorphology similar to Thallites sp. (Plate I, 1); it needs a detailedstudy before it may be accurately identified. The specimen wascollected from marine sediments and was dated as Lockhovian,according to the associated graptolite faunas (Gutiérrez-Marco et al.,1999).

The remains assigned to Sciadophyton steinmanni Kräusel andWeyland, synonymouswith Sciadophyton laxum (Dawson) Steinmann(see Schweitzer, 1980), were collected from the Gedinnian sedimentsof Monterrubio de la Serena (SE Badajoz) and are preserved ascompressions (Álvarez-Ramis, 1988, 1995), and correspond to axes ina rosette-like arrangement with simple or bifurcate branches. Theaxes present a central strand spreading at their end structures a roundto slightly lobed outline. Edwards andWellman (2001)mentioned thefirst reference of Álvarez-Ramis (1988) in a study of terrestrial plantsfrom the Ordovician to the Lower Devonian.

Morphogenus Sciadophyton has been considered the gametophytestage of primitive terrestrial plants. Sciadophyton has been placedwith the Zosterophyllopsida (Bell and Hemsley, 2000) or therhyniophytes (Gerrienne et al., 2006), but its true relationship isstill open (see Taylor et al., 2008).

Overall, the Lower Devonian floras from Iberia correspond to aherbaceous vegetation developed across coastal marshes as indicatedby sedimentological data. As was expressed by Schweitzer (1983),these littoral areas probably suffered frequent flooding, giving rise to amultitude of intermediate biotopes between the continental andmarine types, whereby individuals of the same species formed strandsin which groups of various species intermingled.

5. Middle Devonian

Middle Devonian vegetation is represented by assemblages fromQuinto Real and Olaberri (Navarra, northern Spain) and one isolatedremain from Guadalmez (SE Ciudad Real, central Spain) (Table 1).

The associations from Quinto Real and Olaberri were collectedfrom black slates and dated as Mid-Devonian in the presence ofCalceola sandalina (Cnidaria). The fragmentary remains, preserved asimpressions, were cautiously identified by Gómez de Llarena (1950,1956) as Psilophyton sp., Haliserites sp., and Calamophyton sp. Thesespecimens were never described. Due to this, the author attributionsare problematic.

Niklas and Banks (1990) suggested that zosterophylls might beseparated into two groups depending on the position of the sporangia.At present, examples of Psilophyton with lateral sporangia are

Plate I.

1. Thallites sp. Gavà (Barcelona), Lower Devonian. Specimen lodged in Seminari2. Stem fragment of Lycophyta, Cangas de Onís (Asturias), Upper Famenian–Tou3. Strobile of probable Sphenophyta, Valle de Santullán (Palencia), Lower Famen4. Sphenopteridium keilhaui Nathorst, 1902, Valle de Santullán (Palencia), Lower5. Lepidodendraceae indet, Guadalmez (Ciudad Real), Upper Emsian–Upper Givetian

5 cm.6. Stem fragment of cf. Precyclostigma sp. Upper Famenian–Tournaisian. Specim

included in the genus Sawdonia, while all other zosterophylls withterminal sporangia (e.g. Psylophyton princeps Dawson) have beenmaintained in the trimerophytes (Taylor et al., 2008, p. 259).

Given that the samples collected by Gómez de Llarena have notbeen revised and that this author did not observe (or did not describe)the fertile parts of the specimens described as belonging toPsilophyton sp., they cannot be definitively assigned to zosterophyllsor trimerophytes.

As for the genusHaliserites, it was first described from Cenomanianof Saxony (Germany) with the species H. reichii Sternberg. In 1847and 1852, Göeppert applied this generic term to H. dechenianusGöeppert from the Lower Devonian, which was subsequently used inconnection with various fragmentary and/or badly preserved remainsfrom the Devonian until 1903, when White (1903) introduced thenew name Taeniocrada (see Höeg, 1937, p. 565, Höeg in Boureau,1967, p. 222).

Due to the lack of description and figures of specimens identifiedas Haliserites, and Calamophyton it is not possible to know if theattribution of this material is appropriate.

Hartung (in Kullmann, 1960, p. 467) identified Taeniocrada sp.from Siegenian–Eifelian sediments in the Arruz valley (northernPalencia, Cantabrian Mountains; Spain). Hartung recognised that thematerial was deficient and did not allow a specific identification. Laterrevision suggested that ichnofossils were involved (Jürgen Kullmann,2008, written communication).

The deposits at Guadalmez (SE Ciudad Real) have been dated asranging between the Late Emsian and Late Givetian (Pardo Alonso,1997). The only specimen was collected in the red quartzosesandstones of marine origin which laterally present fluvial sedimentsinterspersed. This plant macroremain is a fragment of the externalmould of a small stem determined as Lepidodendraceae indet. (PardoAlonso, 1997, p. 132). Although its morphological characters suggest alycopsid, a more detailed determination is required (Plate I, 5). Due tothe scarcity of Mid-Devonian floras in Spain this occurrence becomesimportant.

The paucity of remains available for study makes it impossible toreconstruct the vegetation existing in the Iberian Peninsula during theMid Devonian, and it is only possible to postulate that at least twolevels of vegetation existed— a ground cover formed by zosterophyllsor trimerophytes (Psilophyton) and a semi-arborescent vegetationformed by Cladoxylales (Calamophyton) and lycophytes of small-treehabit.

6. Upper Devonian

The study of the vegetation of the Upper Devonian in the IberianPeninsula is challenging for several reasons. Firstly, because littlematerial is available and, secondly, because the identification of anyfragments and, in some cases, the stratigraphic dating is doubtful.

The best dated, and oldest, remains are from the villa of Valle deSantullán (Palencia, northern Spain). Associated faunal remains place thislocality in the Lower Famennian (Wagner et al., 1984, p.18). Floristically,the outcrop would be included in the Biozone VI of Archaeopteris (Banks,1981). The scantyavailablematerial consists of three small fragments, twoof which are part and counterpart (Wagner, 1966) and assigned toAneurophyton sp. (Progymnospermopsida, Aneurophytales) and

o de Barcelona Geological Museum. Scale bar 1 cm.rnaisian. Specimen lodged in IMGEMA Jardín Botánico de Córdoba. Scale bar 5 mm.ian. Specimen lodged in IMGEMA Jardín Botánico de Córdoba. Scale bar 5 mm.Famenian. Specimen lodged in IMGEMA Jardín Botánico de Córdoba. Scale bar 5 mm.. Specimen lodged in the Museum of the Geology Department, València University. Scale bar

en lodged in IMGEMA Jardín Botánico de Córdoba. Scale bar 2 cm.

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Table 1Species cited or figured of Devonian vascular plants reported from the Iberian Peninsula.

Taxon Proposed age in the first reference Age in later references Locality (province) References

Psilophyton sp.a Middle Devonian Quinto Real (Navarra, Spain). Gómez de Llarena (1950)Haliserites sp.a

Drepanophycus sp.(aff. D. spinaeformis)

Devonian Lower Devonian Barrancos (Alentejo, Portugal) Teixeira (1951)

Psilophyton ? sp.Hostimella ? sp.Psilophyton sp.a Middle Devonian Quinto Real (Navarra, Spain) Gómez de Llarena (1956)Haliserites sp.a Olaberri (Navarra, Spain)Calamophyton sp.a

Taeniocrada sp.a Siegenian–Eifelian Valle del Arruz (Palencia, Spain) Kullmann (1960)Protolepidodendron wahnbachenseKräusel & Weylanda

Middle Devonian Lower Famennian Valle de Santullán (Palencia, Spain) Wagner (1966)

Aneurophyton sp.a

Hyenia sp.a Givetian? Lower Famennian Valle de Santullán (Palencia, Spain) Wagner (1971)Aneurophyton sp.a

cf. Calamites sp.a Upper Devonian–Lower Carboniferous Alcañices (Zamora, Spain) Teixeira and Pais (1973)cf. Sublepidodendron sp.Hyenia sp.a Lower Famennian Valle de Santullán (Palencia, Spain) Wagner et al. (1984)Sphenopteridium keilhauii NathorstSporogonites sp. (A)b Upper Devonian Upper Devonian–Lower

CarboniferousSierra de Hornachos (Badajoz, Spain) Álvarez-Ramis (1981)

Sporogonites sp. (B)b

Zosterophyllum aff. australianumLang & Cooksonb

Psilophyton sp.b

Thursophyton sp.b

Psilophytites sp.b

Hostimella sp.b

Dawsonites sp. (A)b

Dawsonites sp. (B)b

Ginkgophytopsis sp.b

Drepanophycus sp.b

Archaeosigillaria cf. vanuxemi(Göeppert, 1847, 1852) Kidstonb

Fragmento de Lycophytab

Cyclostigma sp.b

Anisopteris cf. circularis Walton?b

Archaeocalamites sp.b

Rhacophyton mirabilis (Nathorst)Leclerqb

Rhacophyton sp.?b

Archaeopteris roemeriana Goeppertb

Archaeopteris sp.?b

cf. Neurocardiopteris broilii Lutz?b

Sciadophyton steinmanniKräusel & Weyland

Gedinnian–Siegenian Monterrubio de la Serena (Badajoz, Spain) Álvarez-Ramis (1995)

Lepidodendraceae indet. Upper Emsian–Upper Givetian Guadalmez (Ciudad Real, Spain) Pardo Alonso (1997)

a Doubtful or incorrect identifications.b Not revised material.

Table 2Specimens of Devonian vascular plants from Spain lodged in Spanish museums.

Museum Taxon Suggested age Locality Number ofspecimens

Specimens

Botanical Garden of Córdoba(Spain)

Hyenia sp. Lower Famennian Valle de Santullán(Palencia)

2 Strobilus?

Sphenopteridium keilhauii Nathorst,1902

Lower Famennian Valle de Santullán(Palencia)

1 Axes

Lycopsida Upper Famennian–Tournaisian Cangas de Onís (Asturias) 1 Lycopsidaxe

cf. Precyclostigma sp. Upper Famennian–Tournaisian Cangas de Onís (Asturias) 1 Lycopsidaxe

Eoacanthocarpus sp. Upper Devonian–LowerCarboniferous

San Vitero (Zamora) 2 Fertile apex

Stigmaria ficoides Sternberg, 1825 Upper Devonian–LowerCarboniferous

San Vitero (Zamora) 2 Rootletscars

Geological Museum of Seminar ofBarcelona (Spain)

Thallites sp. Lower Devonian Bruguers, Gavà, Barcelona. 1 Branches

Geological Museum of University ofValència (Spain)

Lepidodendraceae indet. Upper Emsian–Upper Givetian. Guadalmez, Ciudad Real 1 Stem

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Protolepidodendron wahnbachense Kräusel and Weyland (Lycopsida,Protolepidodendrales. Subsequently (Wagner, 1971), the specimendetermined as Protolepidodendron wahnbachense Kräusel and Weylandwas later determined as Hyenia sp. (Pteropsida, Cladoxylales).

The sample identified asAneurophyton sp.was re-determined byH. J.Schweitzer (Wagner et al., 1984) as Sphenopteridium keilhauii Nathorst(Plate I, 4). Although there is little doubt concerning the correctness ofthis assignation, the specimen assigned to Hyenia sp. has a morphologythat does not fit either with the morphological characteristics of thisgenus nor with those of Calamophyton. (Plate I, 3) (Table 2).

Plant fossils, belonging to either the Upper Famennian or EarlyTournaisian fromCangasdeOnís (Asturias, northernSpain), and comingfrom the Ermita Formation, is regarded as having been deposited on ashallow platform (García-Alcalde, 1995; García-Alcalde et al., 2002).These are impressions of stem fragments attributable to Lycophyta(Plate I, 2). After our revision of thematerial one of the fragments seemsto be related to Precyclostigma sp. (Plate I, 6).

The floristic associations of six horizons from Sierra de Hornachos,Badajoz province (Álvarez-Ramis, 1981), include fragments of axesand reproductive structures of bryophytes, zosterophylls, trimero-phytes, lycophytes, sphenophytes and progymnosperms. The remainswere attributed to twenty one different taxa mostly belonging to theflora of Psilophyton (e.g. Psilophyton, Hostinella, Drepanophycus,Thursophyton) although some elements correspond to Archeopteris(Archaeopteris, Racophyton,Neurocardiopteris) (See Table 1). Althoughseveral of these taxa are widely distributed stratigraphically andcoexisted during the Devonian, this floristic association does not seemto be representative of Upper Devonian. It seems that more work isneeded, as a result of the poor illustrations and the fact that thespecimens are not available for a revision.

Other associations from the Iberian Peninsula with an age betweenthe Late Devonian and the earliest Carboniferous are those collected inSanVitero (Zamora, northernSpain) andGuadramil (Bragança, Portugal).

The Spanish material (Martínez García, 1972, 1973) (Table 2)corresponds to an incomplete and badly preserved apex of a lycophyte,determined by R. H. Wagner as Eoacanthocarpus sp. (Lycopsida,Isoetales), a taxon only known from the Tournaisian, but whosebeginning in the fossil record is unknown, and two hand samplesshowing examples of rootlet scars attributed to Stigmaria ficoidesSternberg. Themorphogenus Stigmaria (Lycophyta), typical taxon of theCarboniferous flora, also existed in the Devonian (see Lejal-Nicol andMassa, 1980). Other specimens collected in the same locality have beenattributed to cf. Calamites sp. and cf. Sublepidodendron sp. (Teixeira andPais, 1973), and the locality dated as probably Upper Devonian based onthe stratigraphy and the flora (Teixeira and Pais, 1973). The descriptionof the specimen attributed to cf. Calamites corresponds to a ribbed axialfragmentwithout nodes that could also be assigned to Archaeocalamitesgenus.

The flora from Guadramil (Teixeira, 1970; Teixeira and Pais, 1973)comprises indeterminable axial fragments.

The information obtained from single specimens related tolycopsids and Sphenopteridium keilhauii is clearly insufficient toreconstruct the Upper Devonian vegetation in the Iberian Peninsula.Nevertheless, some indications about the environment are provided.Lycophytes, whether herbaceous or arborescent, coincide as regardsof their habitat preferences. Analysis of plant-bearing sediments fromthe Iberian Peninsula suggests that both groups occupied floodplainsor channel margins adjacent to flowing water and lakes.

7. Conclusions

Our knowledge of the Devonian plants from the Iberian Peninsuladates from themiddle of the twentieth century. However, the remainsof the vascular Devonian plants found have frequently beendetermined erroneously and lack good stratigraphic control, whilethe majority has been given an open nomenclature due to the poor

state of conservation or the fragmentary nature of the specimens. Themacrofloras found correspond to the Lower Devonian of Barrancos(Alentejo, Portugal) and Upper Devonian Sierra of Hornachos(Badajoz, Spain). The rest consists of single finds from Lochkovianto Late Devonian–Early Carboniferous, as drifted fragments in shallowmarine strata.

Although the data are scarce or disseminated, they are important forrepresenting the only data available for the Iberian terrane. As a whole,they suggest preponderance or a greater conservation potential oflycophytes, and agree with the sequential appearance of different typesof vegetation: herbaceous – a mixture of herbaceous and arborescentvegetation – arborescent.

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