REVISION OF THE PENNSYLVANIAN FLORA FROM VAL SANAGRA IN WESTERN PART OF THE SOUTHERN ALPS (ITALY)

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J. Pšenička, S. Opluštil, A. Ronchi & Z. Šimůnek, Revision of the Pennsylvanian flora from Val Sanagra

REVISION OF THE PENNSYLVANIAN FLORA FROM VALSANAGRA IN WESTERN PART OF THE SOUTHERN ALPS (ITALY)

Josef Pšenička1, Stanislav Opluštil2, Ausonio Ronchi3 & Zbyněk Šimůnek4

1 Palaeontology Department, West Bohemian Museum in Pilsen, Kopeckého sady 2, 30100 Plzeň, Czech Republic;[email protected]

2 Charles University in Prague, Faculty of Science, Albertov 6, 128 43 Praha 2, Czech Republic; [email protected] Dipartimento di Scienze della Terra e dell’Ambiente, Universita di Pavia, Via Ferrata 1, 27100 Pavia, Italy;

[email protected] Czech Geological Survey, Klárov 3/131, 118 21 Praha 1, Czech Republic; [email protected]

Abstract: The Pennsylvanian flora from the Alpe Logone/Val Sanagra locality in the Western Lombardy(Southern Alps, Italy) last underwent a taxonomic study in the mid 20th century. The main problem isgenerally poor preservation of the plant remains, which makes their identification problematic. Despitethis, the authors have identified 43 fossil species from this locality. Based on the sedimentary context ofthis paleoflora, we assume that the Val Sanagra sediments were deposited in a continental setting, ina fluvial environment with a well-developed and vegetated floodplain where where clastic substrates wouldlocally and occasionally change into peat swamp. The common presence of rhytidolepis and sub-rhytidol-epis sigillarias, as well of Eusphenopteris neuropteroides, Lobatopteris miltonii, Mariopteris latifolia,Paripteris linguaefolia, Pecopteris microphylla, Alethopteris sp. (cf. grandinii) indicates that the florafrom the Val Sanagra locality spans the interval between the Duckmantian and Bolsovian (middle Mos-covian), and thus represents one of the oldest Pennsylvanian floras of the Southern Alps.

Key words: Pennsylvanian, fossil flora, Val Sanagra, Italy, Southern Alps

INTRODUCTION

The Pennsylvanian Subsystem has among thebest-known fossil floras described from many lo-calities throughout the world, including Europe.Nevertheless, Pennsylvanian floras from Italy re-main at the periphery of interest of palaeobota-nists because of being scattered in only a few lo-calities which often represent tectonic units with-out clear stratigraphic and paleogeographic con-text, as well as providing remains that are not al-ways well-preserved. Despite of this, the Pennsyl-vanian floras of Italy could play an important rolewhen reconstructing the Late Carboniferous envi-ronments and paleogeography of this part of Eu-rope, which are masked by strong Alpine over-print. Outcrops of terrestrial sediments, whichhave yielded Pennsylvanian floras, occur in differ-ent parts of Italy, especially in the regions of West-ern Lombardy, Canton Ticino, the Carnic Alps,Tuscany, and south western, central and southeastern Sardinia (Ronchi et al., 2012).

This paper deals with a revision of the Pennsyl-vanian flora from the Alpe Logone/Val Sanagra lo-

cality (Figures 1, 2) in the Western Lombardy(Southern Alps, Italy) which was first discoveredin the mid 20th century. A rich collection of about2000 specimens is now stored in the Museo Civicodi Storia Naturale (Milan), and a minor but signif-icant additional collection is in the Museo Et-nografico e Naturalistico Val Sanagra (Grandolaed Uniti, Como).

The first report on Carboniferous macroflorasfrom this locality came from Magnani (1946), whodiscovered impressions and casts of CalamitesSuckow ex Sternberg, Sigillaria Brongniart andLepidodendron Sternberg. In the mid 20th centu-ry, Silvio Venzo and Luigi Maglia (Venzo and Ma-glia, 1947; Venzo, 1951) collected more than 2000leaf compressions and impressions from the an-thraciferous and silicified beds in Val Sanagra andVal Cavargna at Alpe Logone. These authors de-scribed, but never illustrated, about 22 genera and75 species belonging to the sphenopsids (Spheno-phyllum Brongniart, Calamites, AsterophyllitesBrongniart, Annularia Sternberg), lycopsids (e.g.,

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Lepidodendron, Knorria Sternberg, Lepidophyl-lum Brongniart, Sigillaria, Stigmaria Brongniartor Ulodendron Rhode), cycads, cordaitaleans,ferns (e.g. Pecopteris Brongniart) and pteri-dosperms (e.g. Mariopteris Zeiller, NeuropterisBrongniart, Linopteris Presl). Poorly preservedribbed fragments of Sigillaria were also reportedfrom the arenaceous layers.

Venzo and Maglia (1947) ascribed this flora tothe late Westphalian (Westphalian C/ Middle Mos-covian) and consider the flora of the Alpe Logone/Val Sanagra to be slightly older than the florafrom the Manno Conglomerate in Ticino, attribut-ed to the early Stephanian by Sordelli (1896).Some years later, Venzo (1951) attributed the ValManno flora (in agreement with Jongmans 1960)to the Westphalian B-C transition (early Moscov-

ian) and therefore considered it older than theAlpe Logone/Val Sanagra flora. Recently, Pittau etal. (2008) found and described very rich late West-phalian-early Stephanian palynomorph associa-tions in the Bedero locality (Luino, eastern side ofMaggiore Lake, W Lombardy). On the basis of areview of the old papers, these authors assignedboth the Logone and Manno’s macrofloristic as-semblages to the Moscovian (Bolsovian) or to thelower Moscovian (Duckmantian-Bolsovian) transi-tion.

This paper revises all the accessible materialfrom the Alpe Logone/Val Sanagra locality, identi-fies individual species and results based on a newidentification of the plant species.

GEOLOGICAL BACKGROUNDSparse and thin continental Pennsylvanian suc-cessions crop out in the western Southern Alps(Figure 2), to the west of Lake Como (Varesotto-Luganese area), across the border between Italyand Canton Ticino (Switzerland).

These deposits are the product of the erosion ofthe Variscan mountain chain and occur generallypinched as tectonic slices along major tectoniclines or in stratigraphic non-conformity above themetamorphic basement (Figure 3).

The age of all these Upper Palaeozoic sedimen-tary successions, scattered over a wide area be-tween Lake Como, Lake Maggiore and Ticino, has

Figure 1. Entrance into one of the two anthracite quarriesstill existing in Val Gariasca (a small tributary valley of themain Val Sanagra) and exploited around the first half ofthe 20th century.

Figure 2. Location of the Val Sanagra/Alpe Logone Penn-sylvanian plant fossil bearing deposits in the WesternSouthern Alps.

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long been debated as being between middle West-phalian and Stephanian (i.e. Moscovian/Kasimov-ian) (e.g. Lehner, 1952; Venzo and Maglia, 1947;Stadler et al., 1976; Casati, 1978; Gaetani et al.,1986).

Among the main localities, Val Sanagra (or moreprecisely Val Gariasca, near Alpe Logone) and sec-ondarily Val Cavargna close to western side ofLake Como, were the ones which yielded the rich-est fossil floras. A detailed description on histori-cal findings as well as the exploitation of the smallanthracite mines can be found in Selva (2009).

The thickness of the fine to coarse-grained clas-tic deposits (Figure 3) varies from missing to 15 m(Venzo and Maglia, 1947).

MATERIALAll fossil plant remains presented in this paperhave been studied from collections stored in theMuseo Civico di Storia Naturale (Milan) and alsoin the Museo Etnografico e Naturalistico Val San-agra (Grandola ed Uniti, Como).

Although the studied fossils are preserved ina wide spectrum of lithologies ranging from con-

Figure 3. Geological sketch section of the Pennsylvanian toLadian succession on the ravine of Val Sanagra-Logone,W of Lake Como (from Venzo & Maglia, 1947, mod).1. Metamorphic basement cloritic mica shists, 2. Transgres-sive conglomerates and sandstones, with coal seams, rich inplants 3. Layer of a porphyritic conglomerates (0.70-2 m);4. Coarse-grained sandstone, poorly fossiliferous (3 m);5. Triassic Dolostones, brecciated at the bottom; 6. Minedebris, with anthracitiferous and fossiliferous shists.

Figure 4. Syringodendron sp. preserved in the fine-grainedbiotite-rich sandstones. Scale bar is 100 mm.

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glomerates and feldspatic sandstones to mud-stones, they dominantly occur in fine-grained sed-iments. The specimens are mostly preserved indark grey to nearly black, massive to poorly bed-ded or laminated micaceous siltstone, locally witha clay admixture and/or roots, and mostly com-prise lycopsid appendices. Also identified wererare stigmarian axes. Fossils are either scatteredor often concentrated on particular bedding planesas drifted fragments, usually composed mostly ofisolated linopteroid pinnules, calamites axis or iso-lated lycospid (mostly sigillarian) leaves. Silt-stones often grade into grey fine-grained biotite-rich sandstones sometime with silty laminas. Theypreserve mostly pteridosperm or cordaites axes, orremains of decorticated (ribbed) sigillarian stems(Figure 4). The same fossils can be occasionallypreserved in even coarser facies represented bycoarse-grained feldspatic sandstones to finegrained conglomerates (about 3 mm grains ofquartz and feldspar). These lithologies preservemostly sphenopsid stems (Figure 5).

High coal rank, anthracite-coal deposit, preclud-ed usage of any maceration techniques. In addi-tion, some fossils are tectonically deformed. Theyare preserved as compressions or impressions (ad-pressions); some of them are rather “shadows”preserving the outline of former fossil remains.Also compressions are often poorly preserved, es-pecially foliage remains which makes their speciesidentification complicated, or even impossible.Dominating among plant fossils from Val Sanagraare unidentifiable axes of pteridosperms and/orcordaitalean plants, decorticated sigillarian stems,isolated linopteroid pinnules and calamites stems.Subordinate are remains of lycopsids and foliageof ferns and pteridosperms.

SYSTEMATIC PARTDivision LycopsidaLycopsid remains comprise a significant part ofthe Val Sanagra collection, although most of themare represented only by partly decorticated sig-illarian stems which exhibit pronounced verticalribs some of them with rows of double ellipticalscars of parichnos strands typical of Syringoden-dron Sternberg. Specimens preserving the outersurface with leaf scars assignable to the genusSigillaria are much less common, although sever-al relatively well-preserved specimens were found

Figure 5. Calamites sp. preserved in the coarsegrainedfeldspatic sandstone. Scale bar is 100 mm.

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in the collection. These can be attributed to sever-al species described below. Common, but occurringusually separately from stems are isolated long,single-veined grass-like lycopsid leaves, about 4 –8 mm wide, are attributed to Cyperites Lindleyand Hutton, a fossil genus which is generallythought to represent compressed sigillarianleaves. These leaves are mostly preserved in thefinest facies where they are associated with isolat-ed linopteroid pinnules or foliage fragments offerns and pteridosperms. Remains of other lycops-id genera are much less common compared to Sig-illaria, and are represented by few specimens ofLepidodendron, Lepidophloios Sternberg and per-haps Bothrodendron Lindley et Hutton. Also iden-tified were decorticated lepidodendrid stems rep-resented by a few specimens of the Knorria andAspidiariopsis type of preservation. Fertile organsof lycopsids are very rare and only Lepidocarponmajus sporophyll remains have been identified.Underground organs are represented by commonappendices and few specimens of Stigmaria.No subarborescent to herbaceous forms of lycops-ids have been identified in the studied material.

Order LepidodendralesFamily SigillariaceaeGenus Sigillaria Brongniart, 1822Species Sigillaria rugosa Brongniart 1828 (PlateI, Figs 1-6)Three specimens which possess the scars on ribsseparated by prominent straight furrows havebeen identified in the collections. The ribs are 8 –10 mm wide on specimen B1722 (Plate I, Fig. 2),about 12 mm on specimen B1740 (Plate I, Fig. 1)and up to the 16 mm on specimen B3539 (Plate I,Fig. 4). Leaf scars are nearly oval to slightly sub-triangular with rounded lateral angles and onlygently convex upper end. Some leaf scars stillshow lateral parichnos as linear markings withpunctuate vein cicatricule in between, all situatedat about one third down the leaf scar. Length ofleaf scars varies mostly between 8 and 10 mm andwidth 4.5 and 5.5 mm (Plate I, Figs 3, 5), exceptspecimen B1740 which possess leaf scars 12.5 mmlong and 7.2 mm wide in the lower third of thescar length (Plate I, Fig. 6). The shape, however,is very similar to the other two specimens. Verti-cal distance between leaf scars also varies among

specimens from 22 mm (B1740) to 30 mm (B1722,3539). Surface between leaf scars is smooth(B1722, 1740) (Plate I, Fig. 3), however, specimenB3539 (Plate I, Fig. 5) displays fine vertically ori-ented closely spaced and slightly undulating linesalong the ribs. In addition, lateral angles of theleaf scars give rise to two fairly prominent mildlyundulating lines descending vertically to the leafscar next below (Plate I, Fig. 5). Specimen B1722is partly decorticated. These decorticated partspossess prominent ribs with vertically striated ar-eas between them and markings indicating posi-tion of leaf scars (Plate I, Fig. 3).Remarks: The specimens described above showthe same shape and size of leaf scars, and theirdensity and relatively poor ornamentation is typi-cal for Sigillaria rugosa documented by Němejc(1951) or Crookall (1966) or as can be seen fromoriginal Brongniart drawing of the type specimencoming from Pennsylvania. S. rugosa can be com-pared with several other eurhytidolepis speciessuch as S. voltzi Brongniart, S. cortei Brongniartor S. contracta Brongniart, some of them beingconsidered as synonymous to S. rugosa by variousauthors (e.g. Deltenre, 1924; Crookall, 1966).S. rugosa is common in Duckmantian to lowerBolsovian (lower/middle Moscovian) strata of mostcoalfields in Europe and its precise stratigraphicrange can vary slightly among basins. Conse-quently, this species is occasionally reported fromLangsettian (upper Bashkirian) strata and Crook-all (1966) mentioned it to be very rare even in theAsturian (upper Moscovian) of the Bristol andForest of Dean coalfields in UK.

Species Sigillaria cf. inferior Weiss 1881 (PlateI, Fig. 7)The only specimen (B1742) is a poorly preservedfragment of a ribbed bark with leaf scars. Furrowsseparating ribs are undulating and the distancebetween neighbouring ones varies from 3 to 5.5mm; the largest width is reached at the level ofthe leaf scar or slightly above it. Leaf scars arearound 3.5 – 4 mm wide and 2 mm long, about 4.5mm apart. Area between leaf scars appearssmooth, however, this could be also due to preser-vation. Leaf scars are generally lenticular inshape, although some of them (Plate I, Fig. 7)seem to be sub-triangular with an upper angle andconvex margin, thus being more bell-shaped and

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Plate I

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transversally asymmetrical. However, most leafscars have only a convex upper margin. Pointed orlinear impressions of parichnos tissues and/or veinin between are also seen on some leaf scars.Remarks: The specimen B1742 can be comparedwith several species among which the most simi-lar is S. inferior from the early middle Moscovianstrata (Duckmantian) of the Upper Silesian CoalBasin. The Weiss species has similar size and den-sity of lenticular to sub-hexagonal leaf scars andmarkedly undulating ribs. It possesses also trans-versally oriented wrinkles (striations) betweenleaf scars, which however are not observable onthe Val Sanagra specimen. Another species whichhas to be taken into account when comparing withthe above described specimen is S. canobiana Kid-ston 1903 from the Carboniferous of Scotlandwhich is of similar size and shows undulating ribsbut has generally more bell-shaped leaf scars.Comparison was also made with S. transversalisBrongniart (1828) originally described fromAachen coalfield (?Langsettian-Duckmantian) inNW Germany. This species has lenticular leafscars of similar size as the specimen describedhere, but the ribs are nearly straight. Finally, thedescribed specimen strongly resembles Sternberg’s“Rhytidolepis ocellata” from lower Bolsovian stra-ta of the Radnice Basin, Czech Republic. Bothhave undulating ribs, and smooth areas betweenlenticular to flat sub-triangular leaf scars. Němejc(1951) reinterpreted this Sternberg species, repre-sented only by a single specimen, as S. pachyder-ma sensu Corda non Brongniart. S. pachydermasensu Corda differs from the specimen describedhere mainly in having slightly undulating longitu-dinal wrinkles between the leaf scars. In any case,all the comparable sigillarian species are of theRhytidolepis-type and are stratigraphically locatedto the Bashkirian and Moscovian (late Namurianto about early-middle Bolsovian).

Species Sigillaria cf. ovata Sauver 1848 (Plate II)Two relatively well-preserved and partly decorti-cated specimens (B1769 and B1887) have been as-signed to this species. The specimen B1769 pre-serves fragments of two associated bands (ribs) ofbark situated at a low angle to each other. One ofthem is about 9.5 mm wide and smooth with leafscar(s) that shows rather straight only slightlyconvex upper edge and rounded lower edge andboth lateral angles (Plate II, Fig. 4). The scar is6.3 mm long and 5.8 mm wide slightly below themiddle of the scar length. The upper edge is 2.6 mmlong. Although only one leaf scar is well preserved,the outline of the position of the second scar isidentifiable which allows the distance betweenscars to be estimated at about 6.5 mm. A vascularbundle and parichnos cicatricules are situatedslightly above the mid of the scar length. The sec-ond bark band is 13.5 mm wide, vertically striatedwith preserved vascular bundle markings 14.5 mmapart. No leaf scar is present, which together withvertical striations across the whole rib indicatesthat it is a decorticated part of bark (Syringoden-dron stage).The second specimen (B 1887) is split into severalfragments which preserve several ribs of sigillari-an bark, partly with leaf scars, partly withoutthem. Ribs are all about 13.0 – 18.5 mm wide andare either smooth or vertically striated (Plate II,Fig. 1). Those which are smooth possess leaf scarswhich have a straight or nearly straight (or verygently convex) upper edge about 2.8 mm long, androunded lower edge and lateral angles. The leafscars of one fragment are 6.5 mm wide and5.0 mm long (Plate II, Fig. 1), however bark bandson Plate II, Fig. 5 have leaf scars which are about10 mm wide and 8.5 mm long. Vascular bundleand parichnos markings are in the upper half ofthe scars. Striated (?decorticated) bands (Plate II,Figs 1, 3, 6) display outline of leaf scars similar to

Plate I1) Sigillaria rugosa; Museo Civico di Storia Naturale (Milan); specimen B1740; scale bar 5 mm; 2) Sigillaria rugosa;Museo Civico di Storia Naturale (Milan); specimen B1722; scale bar 5 mm; 3) Sigillaria rugosa; Museo Civico di StoriaNaturale (Milan); specimen B1722; scale bar 5 mm; 4) Sigillaria rugosa; Museo Civico di Storia Naturale (Milan); speci-men B3539; scale bar 5 mm; 5) Sigillaria rugosa; Museo Civico di Storia Naturale (Milan); specimen B3539; scale bar5 mm; 6) Sigillaria rugosa; Museo Civico di Storia Naturale (Milan); specimen B1740; scale bar 5 mm; 7) Sigillaria cf.inferior; Museo Civico di Storia Naturale (Milan); specimen B1742; scale bar 5 mm.

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Plate II

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those of smooth bands also with apparent parich-nos and vascular bundle markings. The scar fig-ured on Plate II, Fig. 3 displays a concave upperedge.Remarks: The specimens described here can becompared with S. ovata Sauver and S. principisWeiss. Both have relatively wide pear-like leafscars with lateral and rounded lower edges. How-ever, S. principis has a prominent arched trans-verse mark above the leaf scar and subverticallines running down from the lateral angles whichhave not been observed in the specimens de-scribed here. Therefore we assigned them toS. ovata, although with some uncertainty result-ing from the upper edge, which is more straight inthe Val Sanagra specimens. S. ovata is a fairlycommon species in most basins of the belt ofparalic coalfields in western and central Europebetween the Langsettian to Duckmantian, but be-comes very rare in Bolsovian and Asturian floras.

Species Sigillaria tessellata Brongniart 1828(Plate III, Figs 1, 2)

Only a single specimen shows two straight ribsseparated by gently flexuous furrows. The ribs areabout 9 - 10 mm across. Leaf scars are hexagonal,occupying most of the rib width, about as wide aslong or slightly wider, with lateral angles sub-an-gular to sub-rounded (Plate III, Fig. 2). Margins ofscars are straight or slightly convex. Leaf scars aredensely spaced separated only by prominentstraight transversal furrow across the half of therib width. Most of leaf scars show the position ofthe parichnos and vascular cicatricules, which aresituated at about one third the way down the scar.Remarks: The specimen described here which isfrom the Museo Etnografico e Naturalistico ValSanagra can be compared also with S. boblayiBrongniart, S. micaudi Zeiller and S. barbataWeiss. All the mentioned species have leaf scars ofsimilar size and shape, especially in the case of S.

boblayi. S. barbata has a slightly longer transver-sal furrow separating the leaf scars than the spec-imen described here. In any case, all the men-tioned species occur in Langsettian to Duckman-tian strata, being less common in strata of lateMoscovian age.

Species Sigillaria cf. micaudi Zeiller 1886 (PlateIII, Fig. 3)One specimen in the Museo Etnografico e Natu-ralistico Val Sanagra consists of a fragment withtwo ribs showing several leaf scars separatedalong nearly the whole width of the ribs by astraight transversal furrow. Ribs are about 10 mmwide and separated by flexuous furrows. Thetransversal furrow is closer to the upper margin ofthe leaf scar than to the lower one. Leaf scars arehexagonal, slightly wider than long, and withrounded angles. Vascular and parichnos cicatri-cules are usually one third the way down the leafscar.Remarks: The specimen described here resemblesstrongly the specimen described above as S. tessel-lata, the only difference being that the leaf scarsare slightly less densely spaced. Nevertheless thismay be a result of a different position on the stemsince the shape of leaf scars of both specimens isvery similar. The stratigraphic range of S. micau-di is comparable with that of S. tessellata.

Species ?Asolanus camptotaenia Wood 1860(Plate IV, Fig. 1)The specimen is a fragment of a bark with denseand slightly undulating vertical wrinkles whichirregularly merge and/or split after some distance.Scars are not observable.Remarks: The specimen is probably a poorly pre-served bark of Asolanus camptotaenia, a sub-sig-illarian lycopsid. This species has a very widestratigraphic range and is known from at least thelate Bashkirian to late Gzhelian.

Plate II - Sigillaria cf. ovata; Museo Civico di Storia Naturale (Milan)1) Specimen B1887; scale bar 5 mm; 2) Specimen B1887; scale bar 5 mm; 3) Specimen B1887; scale bar 5 mm; 4) SpecimenB11769; scale bar 5 mm; 5) Specimen B1887; scale bar 5 mm; 6) Specimen B1887; scale bar 5 mm; 7) Specimen B1887;scale bar 5 mm.

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Plate III

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Family LepidodendraceaeGenus Lepidodendron Sternberg 1820Species Lepidodendron aculeatum Sternberg1820 (Plate III, Fig. 4)

This species is represented by only a single speci-men which is a fragment of a bark showing typicallongitudinally and transversally asymmetrical leafcushions with pointed upper and lower angles androunded lateral angles. Below the leaf scar is aprominent keel with notches across. Diamond-shaped leaf scars are as long as wide and some ofthem possess a vascular bundle and two parichnosimprints. Infrafoliar parichnos scars are presentlaterally next to the leaf scars.Remarks: The preservation of the specimen isgood enough to identify the main features typify-ing this species, and therefore its determination isreliable. This species is common from Early toearly Middle Pennsylvanian of Europe and NorthAmerica, and becomes very rare in the end of theMiddle Pennsylvanian.

Species Lepidodendron cf. acutum Presl inSternberg 1838 (Plate V, Fig. 4)Several specimens are preserved as bark adpres-sion. The specimen figured here shows narrow,slightly inflexed and longitudinally asymmetricalfusiform leaf cushions. They are about 18 mmlong and 3 – 3.5 mm wide reaching their maxi-mum width around half the way up the cushion.They display sharply pointed upper and lower anglesand rounded lateral angles. The keel is smoothand only slightly raised. A leaf trace rather thana leaf scar can be seen about three quarters up thecushion. This represents a false leaf scar indicat-ing that leaves were still attached to the stem;however leaves themselves are not preserved.No evidence of an infrafoliar parichnos is present.Remarks: The specimen represents the type oflepidodendrid species with fusiform leaf cushionsand false leaf scars without infrafoliar parichnos.

The most common and widespread representativeof this type is Lepidodendron acutum from earlyto middle Moscovian strata of Europe. This spe-cies corresponds in shape, size and smooth keel tohere described specimen. A similar species, L. sim-ile sensu Němejc (1947) non Kidston, from centraland western Bohemia, possesses similar but usu-ally a little smaller leaf cushions. However, itclearly differs from L. acutum in having muchsmaller and S-bent leaves. Because the leaves arenot preserved in the described specimen, we leaveits determination as uncertain.

Species Lepidophloios sp.(Plate V, Fig. 5)A single, poorly preserved specimen shows theoutline of slightly deformed rhombic and transver-sally symmetrical leaf cushions. These are about16 - 17 mm wide and about 7 – 8 mm long. Theirlower end is angular (about 1200) whereas the up-per end is rounded. Lateral angles are both about70-800. Some of the leaf cushions display imprinttraces above the lower end which might be a poor-ly preserved leaf scar.Remarks: Poor preservation of the leaf cushionswhich are wider than longer allows only for a ge-neric determination, although the size of leafcushions suggests it probably belongs to someforms with larger leaf cushions especially L. larici-nus Sternberg from Early to Middle Pennsylva-nian strata of Euramerica.

Species Halonia sp. (Plate IV, Fig. 1)The only specimen is about 20 cm long and about6 – 6.5 cm wide branch showing small helicallyarranged “ridges” representing poorly preservedleaf cushion outlines. Since no other structure canbe observed, it is assumed that the specimen wasdecorticated already prior the final burial in thesediment. Typical of this fossil genus is an oval10 x 12 mm large branch scar. Some shallow hollowselsewhere on the specimen may suggest the pres-ence of other very poorly preserved branch scars.

Plate III1) Sigillaria tessellata; the Museo Etnografico e Naturalistico Val Sanagra; scale bar 10 mm; 2) Sigillaria tessellata; theMuseo Etnografico e Naturalistico Val Sanagra; scale bar 10 mm; 3) Sigillaria cf. micaudi; the Museo Etnografico e Nat-uralistico Val Sanagra; scale bar 10 mm; 4) Lepidodendron aculeatum; Museo Civico di Storia Naturale (Milan); scale bar10 mm.

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Remarks: Although the specimen is quite poorlypreserved the clear presence of an oval branchscar is indicative of its attribution to the fossil ge-nus Halonia sp. which is most commonly connect-ed with Lepidophlois as the parent plant (e.g.Hirmer, 1927).

Species Lepidocarpon cf. majus (Brongniart)(Plate V, Figs 2, 3)Two specimens represent fragments of the laminapart of a large sporophyll. It displays a pointedapex passing downward into a part with entireparallel sides. A midrib is traceable nearly up tothe apical part of the lamina. The fragment, rep-resent about the apical half of a sporophyll blade,is about 52 mm long and 12 mm wide.Remarks: The size and shape of the lamina indi-cate that it belongs to the fossil genus Lepidocar-pon Allen 1961, a large sporophyll bearing seed-like megaspore and produced by the genus of ar-borescent lycopsids, Lepidophloios (e.g. Batemanet al. 1992). Its large size further suggests that itmost probably represents Lepidocarpon majuswhich is common throughout Early to MiddlePennsylvanian strata of Euramerica.

Genus Stigmaria Brongniart 1822Species Stigmaria ficoides (Sternberg 1820)Brongniart 1822 (Plate V, Fig. 6)The specimen figured is a fragment of the rhizo-phore of an arborescent lycopsid which displaysoval (due to a strong postdepositional deforma-tion) scars regularly arranged into a helix. Theoval scars are about 6 x 3 mm large with a smallcicatrice in the middle. The scars are spaced about6 mm apart.Remarks: Although the preservation is poor andthe specimen is only a small fragment, character-istic appendices scars allow for its generic deter-mination as Stigmaria, most probably represent-ing the most common species S. ficoides.

Species Knorria sp. (Plate IV, Figs 4, 5)Several remains of deeply decorticated lepidoden-drid stems of the Knorria (as well as Aspidiopsis)stage have been found in the museum collections.Two of them have been figured on Plate IV, Fig. 4and show diamond shaped ridges arranged in to ahelix. The space in between them is covered byvertically arranged straight but discontinuous lowand narrow ridges or wrinkles. Another specimen(Plate IV, Fig. 5) shows helically arranged anddensely spaced oval ridges, wider than long, prob-ably as a result of deformation.

Family BothrodendraceaeGenus Bothrodendron Lindley et HuttonSpecies ?Bothrodendron sp. (Plate IV, Figs 2, 3)The specimen preserves several thin and poorlypreserved leafy shoots (Plate IV, Fig. 2) with smallshort leaves thus resembling those of some repre-sentatives of the genus Bothrodendron. Theshoots are about 7 mm long, 2 mm wide andslightly bent in an S-curve (Plate IV, Fig. 3). Rep-resentatives of the genus Bothrodendron are typi-cal for Duckmantian to late Bolsovian strata ofmost European coalfields.

Division SphenopsidaOrder EquisetalesFamily CalamitaceaeGenus Calamites SuckowSpecies Calamites (Diplocalamites) carinatusSternberg 1823 (Plate VI)Calamites (Diplocalamites) carinatus is a commonfossil at the Val Sanagra locality. The species isrepresented by casts of various widths which varyfrom 15 mm to more than 100 mm. The distancebetween two nodes is possible to observe only inone specimen (B1887; Plate VI, Fig. 4); the otherspecimens only have one node preserved. Longitu-

Plate IV1) Asolanus camptotaenia; Museo Civico di Storia Naturale (Milan); specimen B1999; scale bar 5 mm; 2) Bothrodendron sp.;Museo Civico di Storia Naturale (Milan); specimen B1686; scale bar 5 mm; 3) Bothrodendron sp.; Museo Civico di StoriaNaturale (Milan); specimen B1686; scale bar 5 mm; 4) Knorria sp.; Museo Civico di Storia Naturale (Milan); specimenB1784; scale bar 5 mm; 5) Knorria sp.; Museo Civico di Storia Naturale (Milan); specimen B1772, scale bar 5 mm.

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Plate V1) Halonia sp.; Museo Civico di Storia Naturale (Milan); specimen B1747; scale bar 10 mm; 2) Lepidocarpon cf. majus;Museo Civico di Storia Naturale (Milan); specimen B1766; scale bar 5 mm; 3) Lepidocarpon cf. majus; Museo Civico diStoria Naturale (Milan); specimen B1587; scale bar 5 mm; 4) Lepidodendron cf. acutum; Museo Civico di Storia Naturale(Milan); specimen B1787; scale bar 5 mm; 5) Lepidofloios sp.; Museo Civico di Storia Naturale (Milan); specimen B1924;scale bar 5 mm; 6) Stigmaria ficoides; Museo Civico di Storia Naturale (Milan); scale bar 10 mm.

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Plate VI - Calamites (Diplocalamites) carinatus; Museo Civico di Storia Naturale (Milan).1) specimen B1974; scale bar 5 mm; 2) specimen B1693; scale bar 10 mm; 3) specimen B1974; scale bar 5 mm; 4) specimenB1887; scale bar 5 mm; 5) specimen B1619; scale bar 10 mm.

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dinal canals are straight, 1 - 2 mm broad. The el-liptic leaf scares are rarely seen in the nodal part.The leaf scars are 0.8 - 1.5 mm in diameter (Plate VI,Fig. 3 arrow). Specimen B1693 (Plate VI, Fig. 2 ar-row) shows a round branch scar 22 mm in diameter.Remarks: This species has a large stratigraphicrange from Bashkirian (Chokerian) to Gzhelian(Stephanian) (Němejc, 1953). However, the speciesis mainly known from the middle Bashkirian(Langsettian) to lower Moscovian (Bolsovian)(Boureau, 1964). This long stratigraphic rangecould be explained by the fact that this kind of stemin itself probably includes several natural species.The species is in need of detail taxonomic revision.

Species Calamites (Crucicalamites) cruciatusSternberg 1825 (Plate VII, Fig. 1)Only one specimen is observed from the collectionfrom this locality in the Museo Civico di StoriaNaturale. The species is easily distinguished fromother calamitalean species because the carinal ca-nals converge slightly on the branch scars. Branchscars are varied from 5 mm to 10 mm. The intern-odes are wider than high. The internodes are 20 mmhigh and 60 mm wide. The carinal canals arestraight except near the large branch scares (nodes).Both kinds of branch scars are placed in nodes, from3 to 4 in number. Leaf scars are not present.Remarks: This species has a stratigraphic rangefrom middle Bashkirian (Langsettian) to Gzhelian(Stephanian), however it mainly occurs in lowerMoscovian (Bolsovian) to Kasimovian (Barruelan)floras (Boureau, 1964).

Species Calamites (Stylocalamites) cisti Brongn-iart 1828 (Plate VII, Fig. 2)Like Calamites cruciatus, one specimen was recog-nized from the collection from this locality.The stem is 40 mm broad. Internodes are 35 mmlong. Carinal canals are narrow and straight, havingelliptical scars in the upper part (in the nodal re-gion). At the bottom of the carinal canals are onlya small scar represented by „dots“ (again nodal area).Carinal canals have a character of flattened straightcanals that are separated by a narrow groove.Remarks: This species has a large stratigraphicrange from Pennsylvanian to Permian (Němejc,1953), but is mainly known from the lower Mos-covian (Bolsovian) (Němejc, 1953; Boureau, 1964).The long stratigraphic range could be explained by

the fact that this kind of stem in itself probablyincludes several natural species. The species needsdetail taxonomic revision.

Species Annularia radiata (Brongniart 1822)Sternberg 1825 (Plate VII, Figs 6, 7)A few specimens of Annularia radiata were ob-served in the collection from the studied localitywhich is housed in the Museo Civico di Storia Nat-urale (Milan) and also in the Museo Etnografico eNaturalistico Val Sanagra. The ultimate shoot is1 mm broad. Three whorls are presented on thespecimen. Two adjacent whorls on the ultimateshoot is 12 mm broad. One whorl has up to 15leaves. Leaves are laminar, lanceolate, 5 mm longand 0.8 - 1 mm wide (in the middle part of leaf)with more or less sharp tip. The specimen figuredon the Plate VII, figure 7 probably representsa juvenile stage of this species. Leaves are still un-developed (Figure 6).

Figure 6. Probably a juvenile stage of Annularia radiatawith undeveloped leaves; camera lucida from specimenB1559.

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Remarks: This species occurs from the Ser-pukhovian to Gzhelian, nevertheless the mostabundace of the species is from the middleBaskhirian (Langsettian) to lower Moscovian(Bolsovian) (Boureau, 1964; Němejc, 1953).

Species Asterophyllites longifolius (Sternberg1825) Brongniart 1828 (Plate VII, Fig. 3)

One specimen shows interesting calamitaleanleaves having diagnostic features of Asterophyl-lites longifolius. It represents part of leafy branch,more than 60 mm long. The shoot axis is 2 - 3 mmbroad, and is slightly thickened at the nodes. Thedistance between the nodes is 15 mm. Character-istic, very long linear leaves are 35 - 40 mm longand 0.8 - 1 mm wide with a sharp tip. The leavesare fused together in a small area (ca. 0.2 mm)near their attachment point to the nodes. Morethan 11 leaves are attached to each node.Remarks: This species has a wide stratigraphicrange from Moscovian to the Permian (Boureau,1964; Němejc, 1953).

Species Calamostachys cf. tuberculata (Stern-berg 1825) Jongmans 1915 (Plate VII, Figs 4, 5)

The studied material included several specimensof calamitalean cones assigned to the same spe-cies. The cones are more than 50 mm long and 5 -8 mm wide (Plate VII Fig. 4). Internodes are 3 -4 mm long with the axis 2 mm broad. Sterilebracts are lanceolate, 5 - 7 mm long. Cones arereally poorly preserved which makes it impossibleto distinguish sterile and fertile parts, or to deter-mine the character of the sporangia.

Remarks: This species is mainly known fromthe Moscovian (Bolsovian - Asturian), although italso occurs in the Kasimovian and Gzhelian stag-es, as well as in Permian deposits (Boureau, 1964;Němejc, 1953).

Species Asterophyllites equisetiformis (Schlo-theim 1820) Brongniart 1828 (Plate VII, Fig. 8)

The collection contains only a single, poorly pre-served specimen of this species. The leaves aretypically short, acicular and arranged in whorls.The sample could not be photographically docu-mented.Remakrs: The species is not stratigraphically sig-nificant.

Division Pteridopsida

Order MarattialesFamily AsterothecaceaeSubfamily MiltonoideaeSpecies Lobatopteris cf. simonii Zeiller 1888(Plate VIII)Several specimens of Pecopteris cf. simonii wereidentified within the material from the study lo-cality. Nevertheless, it is not an abundant species.The specimens represent fragments of sterile pen-ultimate pinnae, most of them with incompleteultimate pinnae (Plate VIII, Figs 1, 5). The penul-timate rachises are not visible. Ultimate rachisesare 2.5 mm broad and form an angle of 50° withthe penultimate rachis. The ultimate pinnae aremore than 40 mm long and 7 mm wide (Plate VIII,Figs 3, 4). Due to poor preservation of specimens,it is impossible to describe any ornamentation onthe rachises. Pinnules are attached to the ulti-mate rachis at between 90° and 45°. Pinnules are3 mm long and 2.5 mm wide. Two adjacent leavestouch along about half the pinnule length. Theirmargins converge quickly to their end, which isarched. The midrib is strong, up to about 1/3 - 1/2the length of the pinnule, slightly curved at thebase. Lateral veins dichotomize once or twice, orare rarely simple (Figure 7). The apical pinnule ofthe ultimate pinna has a triangular shape withrounded apex (Plate VIII, Fig. 2).

Figure 7. Venation diagram from Lobatopteris icf. simoniiwith margins converge quickly to their end, which isarched; camera lucida from specimen B1519.

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Remarks: These specimens can be compared withthe specimen published by Dalinval (1960, plate56, fig. 1), which also has a strong ultimate rachis,small pinnules and similar venation pattern. Dal-inval (1960) mentioned that pinnules of his speci-men (Pecopteris cf. simonii) are attached perpen-dicular to the ultimate rachis, while the pinnulesof studied species often form an angle of 45° withthe ultimate rachis. However, fossil remains fromVal Sanagra locality are really very poorly pre-served and partly tectonically deformed, and so itis impossible to use the angle of attachment of thepinnules as a reliable taxonomical character.Pecopteris (Lobatopteris) simonii occurs in upper-most Bashkirian (Duckmantian) to middle Mos-covian (Bolsovian, Asturian?) strata (Dalinval,1960).

Species Lobatopteris miltonii (Artis 1825) Wag-ner 1958 (Plate IX, Figs 1, 3, 4)Several specimens from Val Sanagra locality havebeen attributed to this species.Specimens show sterile fragments of separated ul-timate pinnae (Plate IX, Figs 1, 3, 4). Ultimaterachises are 0.8 mm broad and probably coveredby trichomes. Ultimate pinnae are 15 mm wideand more than 30 mm long, parallel-sided for mostof their length and have blunt (70–90°) terminals.Pinnules alternate on the ultimate rachis, are pin-natifid, 7 - 8.4 mm long and 2 - 2.4 mm wide, usu-ally with no more than 5 pairs of lobes along theirlength (Plate IX, Fig. 4). Venation is not visiblebecause of poor preservation.Remarks: The studied specimens compare wellwith the Lobatopteris miltonii holotype describedby Pšenička et al. (2009).Lobatopteris miltonii occurs in middle Bashkirian(Langsetian) to early Moscovian (Bolsovian) stra-ta (Němejc, 1953; Dalinval, 1960).

Species Pecopteris microphylla Corsin 1951(Plate IX, Fig. 2)Several species from collections housed in theMuseo Civico di Storia Naturale and MuseoEtnografico e Naturalistico Val Sanagra belong tothis species. The lateral veins once divided, closeto the midvein, into two arms which are decurringagainst each other, straight the pinnule marginwith a rounded/triangular tip.Remarks: Lobatopteris microphylla occurs fromMoscovian (Bolsovian) to the lowermost Kasimov-ian (Corsin, 1951).

Order ZygopteridalesFamily ZygopteridaceaeSpecies: Corynepteris (Alloiopteris) sp. (Plate IX,Figs 5, 6)Only two sterile specimens of Corynepteris werediscovered in the collections from Val Sanagra lo-cality.Both specimens show small fragments of penulti-mate pinnae with three incomplete ultimate pin-nae attached (Plate IX, Fig. 5). The penultimaterachis is 2 mm broad and hirsute. The distancebetween two adjacent ultimate rachises is 10 mm.Ultimate pinnae are linear, 4 - 4.5 mm broad andmore than 15 mm long. The ultimate rachises are0.6 mm broad. Although the pinnules are reallypoorly preserved, it appears that they are free orpartly merged, asymmetrical, 2.2 mm long and1.1 mm wide (Plate IX, Fig. 5).Remarks: The studied specimens show a similari-ty with Corynepteris sternbergii Ettingshausensensu Zeiller described from Újezd u SvatéhoKříže, Radnice Basin, Czech Republic (Pšeničkaand Opluštil, 2011). However, a detail comparisoncannot be done because of the absence of diagnos-tic features.

Plate VII1) Calamites (Crucicalamites) cruciatus; Museo Civico di Storia Naturale (Milan); specimen B1960; scale bar 10 mm;2) Calamites (Stylocalamites) cisti; Museo Civico di Storia Naturale (Milan); specimen B1886; scale bar 5 mm; 3) Astero-phyllites longifolius; Museo Civico di Storia Naturale (Milan); specimen B1761; scale bar 5 mm; 4) Calamostachys tubercu-lata; Museo Civico di Storia Naturale (Milan); specimen B1902; scale bar 5 mm; 5) Calamostachys tuberculata; Museo Civ-ico di Storia Naturale (Milan); specimen B1887; scale bar 5 mm; 6) Annularia radiata; Museo Civico di Storia Naturale(Milan); specimen B1888; scale bar 5 mm; 7) the juvenile stage of Annularia radiata; specimen B1559; scale bar 5 mm;8) Asterophyllites equisetiformis; apical part; the Museo Etnografico e Naturalistico Val Sanagra; scale bar 10 mm.

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Plate VIII - Lobatopteris cf. simoni; Museo Civico di Storia Naturale (Milan).1) specimen B1519; scale bar 5 mm; 2) specimen B1519; scale bar 5 mm; 3) specimen B1519; scale bar 5 mm; 4) specimenB1546; scale bar 5 mm; 5) specimen B1546; scale bar 5 mm.

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Corynepteris sternbergii ranges stratigraphicallyfrom the middle Bashkirian (Yeadonian) to middleMoscovian (Asturian), but is most common in stra-ta of the early Moscovian (Duckmantian, Bolso-vian) age (Němejc, 1953; Galtier and Scott, 1979).

Order: Filicales (leptosporangiate ferns)Family: TedeleaceaeSpecies: Senftenbergia plumosa (Artis 1825)Stur 1877 (Plate X)This common Euramerican species occurs abun-dantly at the studied locality. Nevertheless, allspecimens are really poor preserved.The largest specimen B1634 (Plate X, Fig. 1)shows part of antepenultimate rachis more than80 mm long with 2 mm thick straight rachis. Thepenultimate pinnae alternate on the antepenulti-mate rachis, and are lanceolate, more than 40 mmlong 15 - 18 mm wide, with a straight rachis0.7 mm wide. The distance between adjacentpenultimate pinnae is 17, 6 mm. The ultimate pinnaeare linear lanceolate, 10 mm long and 2 mm wide,slowly tapering toward the acute apex (Plate X,Figs 3, 4). The pinnules alternate, are more or lessoblique, triangular, elongate, and are attached tothe ultimate rachis by the whole base. The pin-nules are 1 - 5 mm long and 1 - 3 mm wide (PlateX, Fig. 2). The midvein extends to the apex. Lat-eral veins are not visible.Remarks: The penultimate pinna is too narrowand some pinnules are relatively small (specimenB163) in comparison with measurements of theSenftenbergia plumosa specimens published byPšenička and Bek (2003). It probably, therefore,represents a part of an immature pinna.Senftenbergia plumosa stratigraphically rangesfrom uppermost middle Bashkirian (Yeadonian)(Carboniferous) to Cisuralian (Permian) strata(Bek and Pšenička, 2001).

Division: Gymnospermopsida

Order: LyginopteridalesFamily: Insertae sedisSpecies: Sphenopteris sp. (Plate XI, Figs 1, 2)Specimen B1671 is a fragment of sphenopterid fo-liage, 20 mm long, with a rachis with several in-complete and one complete sphenopterid pinnules

(Plate XI, Figs 1, 2). The rachis is 1 mm broad andslightly curved. The complete pinnule is lobate,6 mm long and 4 mm wide with a serrate/dentatemargin (Figure 8). The pinnule base is slightlyconstricted. The distance between adjacent pin-nules is 5 mm. The midvein is strong and slightlyS-curved. Lateral veins bifurcate several times.

Figure 8. Venation of pinnules of Sphenopteris sp. Pinnuleshas dentate margin; camera lucida from specimen B1671.

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Plate IX

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Remarks: The species could be compared withSphenopteris andraeana Roehl which was de-scribed from the middle Moscovian (Bolsovian) ofthe Saar Basin. Pinnules of both species havesphenopterid pinnules with a serrate/dentate mar-gin. However, S. andraeana tends to have moreovate pinnules with a strongly constricted base(Alvaréz-Ramis et al., pl. 1, fig. 1) while the stud-ied Sphenopteris sp. has elongate pinnules with aslightly constricted base (Plate XI, Fig. 1).The studied specimen could represent a new spe-cies but the material is too fragmentary and poor-ly preserved to be suitable for typifying of a newtaxon.

Species: Eusphenopteris cf. neuropteroides(Boulay 1876) Novik 1947 (Plate XI, Figs 3, 4)This species is represented by only one specimen(B 1893) which shows part of a penultimate pinnawith several incomplete ultimate pinnae (Plate XI,Fig. 4). The penultimate rachis is 0.8 mm broad,bearing alternating ultimate pinnae. The distancebetween adjacent ultimate pinnae is 6.8 mm. Theultimate rachis is 0.5 mm broad and makes anangle of 55° with the penultimate rachis. The ulti-mate pinnae are more than 25 mm long and 13 mmwide (Plate XI, Figs 3, 4). Eusphenopterid pin-nules (Figure 9) alternate on the ultimate rachis(Plate XI, Fig. 3), spaced as a distance of 3.7 mm.Pinnules are lobate (4-5 lobes), 5.7 - 8.2 mm longand 3.5 - 4 mm wide, slightly elongated, and makean angle of 55° with the ultimate rachis. The pin-nule base is strongly constricted. The tip of thepinnules is more or less triangular or rounded(Plate XI, Figs 3, 4). Venation is not well pre-served but a marked midvein is present.

Remarks: The studied specimen corresponds toEusphenopteris neuropteroides as described byBoulay (1876) and van Ameron (1975). This is notthe same as the species that most authors (e.g.Kidston) have tended to name as E. neuropteroi-des. Small differences can be observed in the tip ofpinnules. Our studied species has more triangular

apical part of pinnules while pinnules in Eusphe-nopteris neuropteroides have more rounded apicalpart. Venation pattern cannot be compared becauseof poor preservation of the studied specimen.

Eusphenopteris neuropteroides stratigraphicallyranges from the middle Bashkirian (Langsettian)to middle Moscovian (Bolsovian) (Ameron, 1975).

Plate IX1) Lobatopteris miltoni; Museo Civico di Storia Naturale (Milan); specimen B1894; scale bar 10 mm; 2) Pecopteris microphylla;the Museo Etnografico e Naturalistico Val Sanagra; scale bar 10 mm; 3) Lobatopteris miltoni; Museo Civico di Storia Naturale(Milan); specimen B1894; scale bar 5 mm; 4) Lobatopteris miltoni; Museo Civico di Storia Naturale (Milan); specimenB1894; scale bar 5 mm; 5) Corynepteris sp.; Museo Civico di Storia Naturale (Milan); specimen B1788; scale bar 5 mm;6) Corynepteris sp.; Museo Civico di Storia Naturale (Milan); specimen B1788; scale bar 5 mm.

Figure 9. The pinnule shape of Eusphenopteris cf.neuropteroides; camera lucida from specimen B1893.

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Plate X - Senftenbergia plumosa; Museo Civico di Storia Naturale (Milan).1) specimen B1634; scale bar 5 mm; 2) specimen B1662; scale bar 5 mm; 3) specimen B1634; scale bar 5 mm; 4) specimenB1662; scale bar 10 mm.

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Order: Callistophytales

Family: Insertae sedisSpecies: Mariopteris cf. lobatifolia Kidston1925 (Plate XII, Fig. 1)

Only one specimen (B1904) could be referred tothis species. It shows the distal part of a pinnawith several ultimate pinnae. The pinna rachis is2.3 mm broad. The distance between adjacentultimate pinnae is 9.3 mm. Ultimate rachises are0.6 mm broad and more than 40 mm long. Pin-nules are robust, obtuse, 6 - 7 mm long 3 - 3.5 mmwide (Plate XII, Fig. 1). Venation is not well pre-served but the midvein is well visible.Remarks: Mariopteris lobatifolia Kidston resem-bles Mariopteris nervosa (Brongniart) Zeiller, butthe pinnules of the latter species are almost in-variably united to each other at the base, even onthe ultimate pinnae in basal positions in thefrond. In Mariopteris nervosa only the basal pos-terior pinnule has a lobe at the base of their distalmargin, while in Mariopteris lobatifolia the suc-ceeding pinnules on both sides of the rachis havea lateral basal lobe. The presence of these laterallobes also separates M. lobatifolia from Mariopter-is sauveuri (Brongniart) Frech. Lateral lobes onMariopteris sauveuri, when present, are usuallyfeebly developed.Mariopteris lobatifolia Kidston was described byKidston (1925) from Great Britain, YorkshireCoalfield, Barnsley, Peacock Coal of the late Bash-kirian (Langsettian) age. It has been also men-tioned by Danzé-Corsin (1953) from France, fromPas de Calais, Assise de Bruay, also of Moscovian(Bolsovian) age.

Species: Mariopteris cf. sauveurii (Brongniart,1833) Frech, 1899 (Plate XII, Figs 2, 3)

This species is represented by two specimens.Specimen B1932 (Plate XII, Fig. 2) shows the up-permost part of a pinna and specimen B1923(Plate XII, Fig. 3) shows the middle part of an ex-terior ultimate pinna (2P3ext) of interior pinna(P2 int) (based on terminology of Boersma 1972).The pinna rachis is straight, 0.7 mm broad. Differ-ent pinnules occur in the acroscopic and basiscop-ic parts (Plate XII, Fig. 3). Acroscopic pinnules arelinguaeform, with an obtuse apex, 8 mm long and4 mm wide, and lie at 70° to the rachis (Plate XII,Fig. 3). Basiscopic pinnules are elongated, 12 -

15mm long and 3 - 3.7 mm wide, and lie at 40° tothe rachis (Plate XI, Fig. 3). The midvein is prom-inent, slightly S-curved in basiscopic pinnules, butthe lateral veins are not preserved due to poorpreservation.Remarks: Fragments of Mariopteris sauveuriifrond are difficult to distinguish from M. nervosaand M. lobatifolia. The pinnules of comparablesize with Mariopteris nervosa are more elongateand have an obtuse to acute apex, while the apicesin Mariopteris sauveurii are rounded. Mariopterislobatifolia hardly shows fusion of the pinnules,even in small fronds, whereas pinnule fusion isa characteristic for M. sauveurii (Boersma, 1972).Mariopteris sauveuri has a very wide geographicdistribution e.g. Great Britain, Belgium, France,Germany, Algeria and Turkey (Jongmans andDijkstra, 1960). It starts in the Duckmantian, oc-curs most commonly in the Bolsovian, and disap-pears in the Asturian.

Species: Karinopteris dernoncourti (Zeiller1886) Boersma 1972 (Plate XII, Figs 4-6)

This species is represented by several small frag-ments of ultimate pinna at the studied locality.Their rachises are 1.2 mm broad. Sphenopteridlobate pinnules are 5 - 11 mm long and 4.3 - 5.5mm wide, with a slightly constricted base (PlateXI, Figs 4-6). Pinnules make an angle of 65° withthe rachis. The distance between adjacent pin-nules is 6 mm. The midvein is prominent andslightly S-curved (Plate XI, Fig. 4). Lateral veinsare not preserved.Remarks: Kariopteris dernoncourtii looks like K.acuta with rounded pinnule apices and thereforedetermination of “intermediate” pinnules can beproblematical. The pinnules of K. dernoncourtiiare more “sphenopteroid” and more divided thanin K. acuta. The prominent proximal basal lobesare an important diagnostic feature of Karinopter-is dernoncourtii. The pinnule margin can beslightly undulated (Josten, 1991).Boersma (1972) erected the new genus Karinopt-eris to accommodate “quadripinnate” forms ofMariopteris. Not all researchers accepted thistransfer; e.g. Josten (1991) used the traditionalname Mariopteris for this species. The species hasgot a wide geographical and stratigraphical distri-bution. It occurs usually in the Langsettian andDuckmantian (Great Britain, Netherlands, Bel-

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gium, France, Germany, Russia and Turkey)(Jongmans and Dijkstra, 1960). It has been foundalso in the Namurian – Yeadonian in Poland andYeadonian and Kindercontian in the Czech Repub-lic (Upper Silesian Basin) Purkyňová (1976).

Order: MedullosalesFamily: Insertae sedisSpecies: Alethopteris sp. [? cf. grandinii(Brongniart, 1828) Goeppert, 1836] (Plate XIII,Figs 1, 5, 7)

Specimen B1894 represents an upper part of apinna with apical long pinnules with more or lessstraight (Plate XIII, Fig. 5) or lobate margins(Plate XIII, Fig. 5). Pinnules are more than 10 mmlong and 4 mm wide, and the rachis is 1 mmbroad. Specimen B1902 (Plate XIII, Fig. 1) showsthe apical part of an ultimate pinna with apicalpinnules 3 - 4 mm long and 1.8 - 2 mm wide andlarger pinnules 8 mm long and 3 mm wide (Figure10). Pinnules are linguaeform. The rachis is 1 mmbroad. Pinnules lie at 45° to the rachis. The mid-vein is marked and extends for 2/3 of the pinnulelength. Lateral veins are not visible.Remarks: Identifying the Alethopteris speciesfrom Val Sanagra is difficult because the venation,which is a critical feature in determining Alethop-teris species, is not preserved. We considered thatspecimen B1902 (Plate XIII, Fig. 1) could belongeither to Alethopteris grandinioides Kessler orAlethopteris grandinii Brongniart (Goeppert)based on their relatively small pinnules. Alethopt-eris grandinioides Kessler, 1916 is poorly docu-mented, the original illustration baing a re-touched photograph and its holotype has beenlost. Therefore Wagner (1968) selected a neotypebut, according to Zodrow and Cleal (1988), this didnot reflect the original characteristics of Kessler‘s(1916) specimen. Zodrow and Cleal (1998) solvedthis problem by erecting a new species, Alethopt-eris pseudograndinioides Zodrow and Cleal, to re-place A. grandinioides Wagner (non Kessler).

Plate XI1) Sphenopteris sp.; Museo Civico di Storia Naturale (Milan); specimen B1671; scale bar 5 mm; 2) Sphenopteris sp.; MuseoCivico di Storia Naturale (Milan); specimen B1671; scale bar 5 mm; 3) Eusphenopteris cf. neuropteroides; Museo Civico diStoria Naturale (Milan); specimen B1893; scale bar; 4) Eusphenopteris cf. neuropteroides; Museo Civico di Storia Naturale(Milan); specimen B1893; scale bar 5 mm.

Figure 10. Ultimate pinna of Alethopteris sp.(cf. grandinii);camera lucida from specimen B1902.

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Alethopteris grandinii, A. grandinioides andA. pseudograndinioides have holotypes of West-phalian D (Asturian) age, but they can also occurstratigraphically lower, in the Bolsovian.The reason, why we decided for Alethopteris gran-dinii is in its pinnule shape. Alethopteris grandi-nii has gibbous pinnules that are also on sampleB1902. The pinnules near the pinna apex becomeshorter and more slender, and the terminal is alsoshort and small in both A. grandinii and sampleB1902 (not preserved in full length (Pl. XIII, Fig.1). A. grandinioides Kessler has usually “flat” pin-nules, slightly prominent midvein, and relativelywider and longer terminal pinnules. On the otherhand, sample B1902 (Fig. 10) has rounded pinnuleapices that are rather more characteristic for Ale-thopteris pseudograndinioides Zodrow and Cleal(A. grandinioides Wagner) than for Alethopterisgrandinii, the latter having rather bluntly acumi-nate to rounded apices. According to these notes,it is very difficult to decide to which species thespecimens on Pl. XIII belong. We identified themas Alethopteris sp. with a little probability, thatthe specimen on Pl. XIII, Fig. 1 could belong toAlethopteris grandinii.

Species: Alepthopteris sp.A (Plate XIII, Figs 2, 6)Specimen B1788 shows several very poorly pre-served fragments of pinnae. Pinnules are small,elongated, linguaeform, subtriangular, 5 mm longand 1 mm wide and united in lower part. The ra-chis is 0.5 mm broad. The midvein is prominentbut the lateral veins are not visible.Remarks: It is difficult to make a comparison,when venation is not preserved. Only one specieshas such narrow pinnules (1 mm wide) – Alethop-teris decurrens (Artis) Frech. However, it tends tohave much longer pinnules.

Species: Alepthopteris sp.B (Plate XIII, Figs 3, 8)Specimens B1771 (Plate XIII, Fig. 3) and part ofspecimen B1902 (Plate XIII, Fig. 8) show a smallfragment of what is probably an alethopterid pinna.

Pinnules are elongated, linguaeform, 8 mm longand 2 mm wide; adjacent pinnules are united intheir basal parts. The rachis is 0.7 mm broad. Themidvein is prominent for 3/5 of the pinnule length.Remarks: As with the previous species, the vena-tion is very poorly preserved and pinnule apicesare not preserved, and so any comparison is diffi-cult. The pinnules are a little longer and widerthan in the previouslyand becauseof its speciesand with its dimension it could belong to variousalethopteroid species.

Species: Neuralethopteris cf larischii (Šusta,1930) Laveine, 1967. (Plate XIII, Fig. 4)Several specimens could be assigned to the genusNeuralethopteris, most notably specimen B1893(Plate XIII, Fig. 4). Pinnules are linguaeform,elongated, 12 mm long and 2.3 mm wide, and at-tached to the rachis by a width of 0.8 mm. Thepinnules have a weakly cordate base, are parallel-sided in their proximal part, and taper in their dis-tal part to an bluntly acute apex (Figure 11). Themidvein lies in a pronounced longitudinal furrowthat extends for most of the length of the pinnule.Lateral veins are not visible.

Figure 11. Ultimate rachis with pinnules of Neuralethopt-eris cf. larischii; camera lucida of the specimen B1893.

Plate XII1) Mariopteris cf. lobatifolia; Museo Civico di Storia Naturale (Milan); specimen B1904; scale bar 10 mm; 2) Mariopteris cf.sauveurii; Museo Civico di Storia Naturale (Milan); specimen B1932; scale bar 5 mm; 3) Mariopteris cf. sauveurii; MuseoCivico di Storia Naturale (Milan); specimen B1923; scale bar 5 mm; 4) Karinopteris dernoncourti; Museo Civico di StoriaNaturale (Milan); specimen B1887; scale bar 5 mm; 5) Karinopteris dernoncourti; Museo Civico di Storia Naturale(Milan); specimen B1897; scale bar 5 mm; 6) Karinopteris dernoncourti; Museo Civico di Storia Naturale (Milan); specimenB1897; scale bar 5 mm.

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Remarks: Neuralethopteris larishii is very similarto Alethopteris decurrens except that the pinnuleshave a “neuropteroid” base. It differs from otherspecies of Neuralethopteris Cremer by the form ofthe pinnules (Josetn, 1983).The species Neuralethopteris larischii occursmainly in the Bashkirian Stage, possibly rangingvery rarely into the basal Moscovian (Duckman-tian) Stage (Tenchov and Cleal, 2010).

Species: Cf. Odontopteris sp. (Plate XIV, Fig. 1)Only one species shows any similarity with thegenus Odontopteris based on the shape of pin-nules. Pinnules are more or less triangular, at-tached to the rachis by their whole base, and makean angle of 50° to the rachis. The pinnules are rel-atively small (4.7 mm long and 2.8 mm wide)which is not typical for odontopterid species. Thevenation is not visible which makes the identifica-tion of the specimen rather speculative.Remarks: The pinnule shape resembles those ofthe Stephanian species Odontopteris nemejcii(Němejc) Šimůnek and Cleal, O. brardii Brongn-iart or O. minor Brongniart. However, withoutknowledge of the venation a detailed comparisoncannot be provided.

Species: Paripteris linguaefolia (P. Bertrand1927) Gothan 1941. (Plate XIV, Figs 2 - 4, 6 - 7)Remains of this species are fairly common at thestudied locality, represented by isolated tongue-shaped pinnules, 40 - 50 mm long and 10 - 14 mmwide. Variation in the size and shape of the pin-nules can be explained by their different positionsin the frond. The midvein is not preserved.Remarks: Paripteris linguaefolia differs fromParipteris gigantea, P. pseudogigantea and P.schuetzei in the absence of a strong midrib andhaving a fan-shaped venation. Paripteris schuetzeiand P. pseudogigantea also usually have smaller

pinnules, a more prominent, coarser venation, andoften a vaulted pinnule blade (Šimůnek, 2010).According to Wagner (1984), Paripteris linguaefo-lia is an index species for the Bolsovian linguaefo-lia Zone. Zhang et al. (1993) argue that Paripetrislinguaefolia and other paripterid species are onlyforms of Paripteris gigantea (Sternberg) Gothan.Their opinion is based on the co-occurrence ofboth these “forms” and others at the same locali-ties in China. On this evidence, they argue thatParipteris gigantea forma linguaefolia occurs alsoin older strata than Bolsovian is association with“true” Paripteris gigantea.

Species: Linopteris obliqua (Bunbury 1847)Zeiller 1899 (Plate XIV, Fig. 5; Plate XV; PlateXVI, Plate XVII, Fig. 1)

Linopteris obliqua represents one of the mostcommon plant remains at the studied locality. It isrepresented by isolated pinnules that vary inshape from round, linguaeform, straight to subtri-angular (Plate XV, Figs 1, 2). Pinnule length rang-es from 6 mm (Plate XVI, Figs 1,2) to 13 mm(Plate XVI, Figs 4, 7, 9) and width from 4.5 to 6.2mm. The pinnules have asymmetric cordate (PlateXVI, Figs 4, 5, 8) or straight bases (Plate XVI,Figs 1, 2, 6, 7, 9), and margins that are longitudi-nally arched with curled margins (Plate XVI, Fig5). The abaxial midvein occupies 80 per cent of thepinnule length (Plate XVI, Figs 4, 5), except in theshorter pinnules where it is relatively shorter(Plate XVI, Figs 1, 2). The venation is anastomos-ing, with lines-of-areolae. Tailed, elongate areolaelie lengthwise with the midvein (Figure 12). Adja-cent to these are elongate areolae of variableshape, including hexagonal and septangular mesh-es and toward the pinnule-margin areolae gradu-ally decrease in size. Areolae at the pinnule mar-gin are consistently elongate, open-ended.

Plate XIII1) Alethopteris sp. [? cf. grandinii (Brongniart, 1828) Goeppert, 1836]; specimen B1902; Museo Civico di Storia Naturale(Milan); scale bar 5 mm; 2) Alethopteris sp. A.; Museo Civico di Storia Naturale (Milan); specimen B1788; scale bar 5 mm;3) Alethopteris sp. B; Museo Civico di Storia Naturale (Milan); specimen B1771; scale bar 5 mm; 4) Neuralethopteris cf.larischii; Museo Civico di Storia Naturale (Milan); specimen B1893; scale bar 5 mm; 5) Alethopteris sp. [? cf. grandinii(Brongniart, 1828) Goeppert, 1836]; Museo Civico di Storia Naturale (Milan); specimen B1894; scale bar 5 mm; 6) Aletho-pteris sp. A.; Museo Civico di Storia Naturale (Milan); specimen B1913; scale bar 5 mm; 7) Alethopteris sp. [? cf. grandinii(Brongniart, 1828) Goeppert, 1836]; Museo Civico di Storia Naturale (Milan); specimen B1913; scale bar 5 mm; 8) Aleptho-pteris sp. B; specimen B1902; the Museo Etnografico e Naturalistico Val Sanagra; scale bar 10 mm.

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The association of pinnules of different shapes in-dicates that these pinnules come from the samespecies, but from different parts of the plant. Thisphenomenon is also shown in Zodrow et al. (2007).Figure 1 on the Plate XVII shows two poorlypreserved pollen organs in organic connection toa rachis with some Linopteris obliqua pinnulesstill attached.Pollen organs consist of several elongated,banana-shaped microsporangia, 7 mm long and0.7 mm wide. The number of microsporangia perpollen organ cannot be exactly given but it ap-pears number more than 10. Laveine et al. (1993)reported a great abundance of Potoniea Zeillerassociated with Linopteris pinnules. Nevertheless,our pollen organ compared with the genus Poto-niea is too small.

Remarks: Linopteris obliqua stratigraphicallyranged from lower Moscovian (Uppermost Duck-mantian) (Fissunenko and Laveine, 1984) to Upper-most Moscovian (Asturian) (Zodrow et al., 2007).Specimen B1714 shows a very interesting pinnuleform with anastomosed venation, with the lines-of-areolate. The pinnule is very long (20 mm) andnarrow (3.2 mm) (Figure 13). The pinnule couldrepresent an aberrant for of Linopteris obliqua, anopinion that would be supported by the venationpattern which corresponds with that species.Remarks: The specimen B1714 (Plate XVI, Fig. 3)is comparable with pinnule dimensions to Linopterislinearis Wagner from near the Westphalian/Stephanian boundary strata of NW Spain (Wagner,1964). The venation of both species is very similar,however on Spanish species is better preserved.

Plate XIV1) Cf. Odontopteris sp.; specimen B1894; Museo Civico di Storia Naturale (Milan); scale bar 5 mm; 2) Paripteris linguaefo-lia; Museo Civico di Storia Naturale (Milan); specimen B1714; scale bar 5 mm; 3) Paripteris linguaefolia; Museo Civico diStoria Naturale (Milan); specimen B1887; scale bar 5 mm; 4) Paripteris linguaefolia; Museo Civico di Storia Naturale (Mi-lan); specimen B1887; scale bar 5 mm; 5) Linopteris obliqua; Museo Civico di Storia Naturale (Milan); specimen B1895;scale bar 5 mm; 6) Paripteris linguaefolia; Museo Civico di Storia Naturale (Milan); specimen B1895; scale bar 5 mm;7) Paripteris linguaefolia; Museo Civico di Storia Naturale (Milan); specimen B1887; scale bar 5 mm.

Figure 12. The anastomosed venation of Linopteris obliqua, with the lines-of-areolate. Tailed, elongate areolae lie length-wise with the midvein. The elongate areolae are variable in shape, including hexagonal and septangular meshes andtoward the pinnule-margin areolae gradually decrease in size. Areolae at the pinnule margin are consistently elongate,open-ended; camera lucida.

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Plate XV1) Linopteris obliqua; Museo Civico di Storia Naturale (Milan); specimen B1660; scale bar 10 mm; 2) Linopteris obliqua;Museo Civico di Storia Naturale (Milan); specimen B1995; scale bar 10 mm.

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Species: Hexagonocarpus sp. (Plate XVII, Fig. 2)One specimen compares with the Hexagonocarpusfigured by Zodrow et al. (2007, fig. 23, A). Zodrowlinked this seed with Linopteris obliqua foliage,which could agree with the fact that this species isthe commonest type of foliage at the present local-ity. The seed has a diameter of 15 mm and threenarrow longitudinal ribs (sclerotesta).

Pteridospem seedsSpecies: Trigonocarpus spp. (Plate XVII, Figs 3, 4)Seeds were probably radially symmetrical (pres-ently preserved as compressions), elongate, andlonger than broad. The testa is differentiated intoa soft outer sarcotesta and hard inner sclerotesta.Only the sclerotesta is preserved with prominentlongitudinal, equidistant grooves (Crookall, 1976).Both illustrated specimens represent differentspecies. The seed on Plate XVII, Fig. 3 is 21 mmlong and 5.5 mm wide; the seed on Plate XVII,Fig. 4 is 15 mm long and 5 mm wide.

Species Carpolithus sp. (Plate XVII, Fig. 5)This seed is 20 mm long and 10 mm wide. It is of aquite different type in comparison with the Trigo-nocarpus species, as the inner sclerotesta does notshow prominent, longitudinal equidistant grooves.

Order: CordaitanthalesFamily: CordaitantaceaeSpecies: Cordaites sp.(Plate Plate XVIII)Determination of species of the genus Cordaites is al-ways difficult without knowledge of cuticles; as wasdocumented by Šimůnek (2007), the venation alone isnot sufficient for species determination. The pre-served venation is a result of taphonomy and modeof preservation of Cordaites leaves before burial.Such long leaves underwent some maceration beforeburial that could influence the state of preservation.It seems that two types occur in the Val Sanagralocality. The first (Plate XVIII, Figs 1, 2) has ve-nation of the “palmaeformis” type. with relativelydense, fine veins (about 40 vein per 10 mm). Thesecond (Plate XVIII, Figs 3, 4) has a venation con-sisting of alternating thick and thin veins (Josten,1991) of the “principalis” type. The density ofthick veins is about 15 veins per 10 mm.Remarks: These venation types have no strati-graphical significance, and range from the upperNamurian to the Permian.

Figure 13. Leave of Linopteris sp. with anastomosed vena-tion and lines-of-areolate. Pinnule is very long and narrow.

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Plate XVI1) Linopteris obliqua; Museo Civico di Storia Naturale (Milan); specimen B1605; scale bar 3 mm; 2) Linopteris obliqua;Museo Civico di Storia Naturale (Milan); specimen B1605; scale bar 3 mm; 3) Linopteris obliqua; Museo Civico di StoriaNaturale (Milan); specimen B1714; scale bar 1 mm; 4) Linopteris obliqua; Museo Civico di Storia Naturale (Milan); speci-men B1962; scale bar 5 mm; 5) Linopteris obliqua; Museo Civico di Storia Naturale (Milan); specimen B1659; scale bar2 mm; 6) Linopteris obliqua; Museo Civico di Storia Naturale (Milan); specimen B1659; scale bar 3 mm; 7) Linopteris obli-qua; Museo Civico di Storia Naturale (Milan); specimen B1962; scale bar 5 mm; 8) Linopteris obliqua; Museo Civico diStoria Naturale (Milan); specimen B1995; scale bar 5 mm; 9) Linopteris obliqua; Museo Civico di Storia Naturale (Milan);specimen B1782; scale bar 5 mm.

DISCUSSIONSince the first report of Pennsylvanian macroflo-ras (Calamites, Sigillarie and Lepidodendron)from the studied locality (Magnani, 1946), manyspecies have been described. Venzo & Maglia(1947) greatly upgraded the list of species fromhere, describing around 70 fossil species (Table 1),again with a clear dominance of lycopsids. LaterVenzo (1951) revised material from the localityand added several new species to the list (Table 1).In this present revision we have distinguishedabout 43 fossil species, which is more than Sordel-li (1896) described from Manno (Canton Ticino,Switzerland) but less than those identified by Ven-zo (1951) in Val Sanagra. The high number of spe-cies identified by Venzo could not be confirmed byour revision, mainly because the generally poorpreservation makes identification of the fossilsproblematic. Venzo probably took small differenc-es caused either by deformation of the plant re-mains due to a strong tectonic overprint of the fos-sil-bearing strata, or simply related to naturalvariability within species, as a justification foridentification of different species. Due to this fact,the number of species (especially among the lycop-sids and sphenopsids) described by Venzo & Ma-glia (1947) and Venzo (1951) has been reduced af-ter a careful revision. Nevertheless, there aresome species that are mentioned in all the papersincluding the present one. These common speciesare as follows: Sigillaria rugosa, Sigillaria tessel-lata, Lepidodendron aculeatum, Lepidocarpon cf.majus, Stigmaria ficoides, Calamites (Crucicalam-ites) cruciatus, Calamites (Stylocalamites) cisti,Annularia radiata, Asterophyllites equisetiformis,Lobatopteris miltonii, Senftenbergia plumosa andLinopteris obliqua. In addition, following speciesare described for the first time from the locality:Sigillaria cf. inferior, Sigillaria cf. ovata, Asola-nus camptotaenia, Lepidodendron cf. acutum,Bothrodendron sp., Calamites (Diplocalamites)

carinatus, Asterophyllites longifolius, Calamo-stachys cf. tuberculata, Lobatopteris cf. simonii,Pecopteris microphylla, Corynepteris (Alloiopteris)sp., Eusphenopteris cf. neuropteroides, Mariopter-is cf. lobatifolia, Mariopteris cf. sauveurii,Karinopteris dernoncourti, Alethopteris sp. [? cf.grandinii (Brongniart, 1828) Goeppert, 1836],Neuralethopteris cf. larischii, Paripteris linguaefoliaor Hexagonocarpus sp.

Zygopterids, which as a group of ferns, havenever been mentioned previously from the ValSanagra locality. We discovered only two poorlypreserved specimens of the genus Corynepteris butit is clear evidence that this type of fern waspresent in the plant assemblage.

Previous authors described two species of Li-nopteris - L. obliqua and L. neuropteroides. Never-theless, our research shows that all pinnule re-mains of the linopterid species probably belong toone species L. obliqua. Its pinnules are commonlyassociated with the other species from differentplant group. The pinnules identified as L. neu-ropteroides are mixed with pinnules, which arereferred to L. obliqua on the same specimen. Ve-nation diagrams of the pinnules made by cameralucida show some differences between pinnulespreserved showing the adaxial and abaxial sides,as already described by Zodrow et al. (2007). Dueto this fact we assume that only one linopteridspecies inhabited the original environment. Thepresence of pollen organ attached directly to ulti-mate rachis and accompanied by pinnules is inter-esting because it represents quite a different mod-el from that described by Laveine (1993). He re-constructed fertile pollen part as isolated fertile„fronds“ that lacked pinnules.

Collected rock samples containing fossils and asimple graphic documentation and description ofthe Val Sanagra section allow only a very generalconclusion concerning the sedimentary paleoenvi-ronment and habitats colonized by the plants.Available data indicate the existence of a wide

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Plate XVII1) The pollen organs of Linopteris obliqua; Museo Civico di Storia Naturale (Milan); specimen B1995; scale bar 5 mm;2) Hexagonocarpus sp.; Museo Civico di Storia Naturale (Milan); specimen B1962; scale bar 5 mm; 3) Trigonocarpus sp.;Museo Civico di Storia Naturale (Milan); specimen B1985; scale bar 5 mm; 4) Trigonocarpus sp.; the Museo Etnograficoe Naturalistico Val Sanagra; scale bar 5 mm; 5) Carpolithus sp.; Museo Civico di Storia Naturale (Milan); specimen B1985;scale bar 5 mm.

range of lithologies from mudstones to conglomer-ates deposited in environments with variable dy-namics. Fine-grained sediments are representedby massive and bedded mudstones, often withmica. Bedded mudstones are dark grey and oftencontain drifted plant remains, either concentratedon a single bedding plane, or scattered. Roots arerare or absent. Massive mudstones and siltstonesare either nearly without plant remains or usuallycontain appendices and rhizomes of arborescentlycopsids thus representing stigmarian root hori-zons probably underlying coal seams. Both thesetexturally different lithologies probably representfloodplain (or delta plain) sediments. Those with-out roots with drifted flora represent sediments ofshallow floodplain lakes, whereas rooted mud-stones represent aerially exposed parts of flood-plains. Sandy admixture in some mudstones indi-cates the influence of a clastic source, probably anactive channel. Locally common biotite may indi-cate a volcanic admixture, or may be simply ofdetrital origin.

Siltstones laminae often alternate with fine-grained sandstone or grade to fine-grained lami-nated sandstone. Rootlets are common as well assome aerial plant remains. Coarse-grained sand-stones and conglomerates rich in feldspar withclasts commonly around 3 – 4 mm and scatteredpebbles around 10 mm in diameter contain re-mains of massive plant axes. The affinities ofthese massive axes are not always determinableeven within the collected specimens, but thosewhich can be determined are mostly decorticatedribbed sigillarian and much less commonly lepido-dendrid stems. Those without “ornamentation”may represent also cordaites or pteridosperms.These coarse lithologies with coalified trunks sug-gest the presence of vegetation in proximity to thebed-load dominating fluvial channels. Dominationof certain plant taxa remains (e.g. isolated sigillar-ian leaves or Linopteris pinnules) indicate the ex-istence of partitioned habitats colonized by vari-ous plant assemblages dominated by pteri-dosperms and calamites, or by arborescent lycops-

ids, mostly sigillarians or less commonly lepido-dendrids (mostly Paralycopodites). Such ecologicaldifferentiation of vegetation is a common featuredescribed from many coalfields and we assume itexisted also in the Val Sanagra locality (e.g.DiMichele and Philips, 1994). The Carboniferousdeposits at Val Sanagra are only a thin tectonicslice of a once thicker sedimentary complex. It istherefore difficult to make any serious conclusionsabout the basin type and its development. Howev-er, the absence of marine fauna in the locality sug-gests this was formerly a part of a continental ba-sin. This is also in agreement with relatively highcontent of feldspatic sandstones indicating a closeproximity of feldspar source. In other Europeancoalfields, feldspatic sandstones are typical forcontinental basins rather than for the paralicones. We therefore assume that Val Sanagra sedi-ments represent part of a succession deposited ina continental setting in a fluvial environment witha well-developed and vegetated floodplain withclastic substrates that locally/occasionally changedinto peat swamps.

STRATIGRAPHICAL RELEVANCE

Venzo and Maglia (1947) placed the flora fromVal Sanagra locality in the late Westphalian,(Westphalian C, i.e. Middle Moscovian). Our revi-sion of the Val Sanagra flora shows that it con-tains elements which have their last appearance inthe uppermost Duckmantian. The best examplesare representatives of the genus Neuralethopterislarischii, Karinopteris dernoncourti and Sigillariaovata. The Duckmantian Substage is also indicat-ed by the rich occurrence of rhytidolepis and sub-rhytidolepis sigillarias although most of the iden-tified species range up into the Bolsovian. Theytend not to be so common there as in stratigraph-ically lower levels. There are some other species,which are typical for the Duckmantian Substage,however, their last appearance occurs within theBolsovian Substage (Table 2). These are Sigillariainferior, Lepidodendron acutum, Eusphenopteris

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Plate XVIII1) Cordaites sp.; Museo Civico di Storia Naturale (Milan); specimen B1858; scale bar 10 mm; 2) Cordaites sp.; MuseoCivico di Storia Naturale (Milan); specimen B1858; scale bar 5 mm; 3) Cordaites sp.; Museo Civico di Storia Naturale(Milan); specimen B1919; scale bar 10 mm; 4) Cordaites sp.; Museo Civico di Storia Naturale (Milan); specimen B19195;scale bar 5 mm.

neuropteroides and Lobatopteris miltonii. Typicalof the Bolsovian Substage are Mariopteris latifoliaand Paripteris linguaefolia, whereas Pecopterismicrophylla or Alethopteris sp. [? cf. grandinii(Brongniart, 1828) Goeppert, 1836] start their oc-currence in the Bolsovian Substage. Calamostach-ys tuberculata is usually associated with late Mos-covian floras, but is normally accompanied by An-nularia stellata, which is absent in Val Sanagra;the determination of C. tuberculata is, therefore,not very reliable here. As a whole, the predomi-nance of rhytidolepis sigillarias and species typicalfor Duckmantian or Bolsovian substages, suggestthat the Val Sanagra flora spans the interval be-tween the Duckmantian and Bolsovian Substages(middle Moscovian).

Val Sanagra flora thus belongs among the oldestPennsylvanian floras of Southern Alps. Interest-ing is the biostratigraphical comparison of the ValSanagra flora with that found in the Manno local-ity (Canton Ticino, Switzerland), since the age-at-tribution of both has been a matter of debate for along time. The first discoveries from the above-cit-ed Swiss fossiliferous locality were made by Heer(1876), who listed Calamites cisti, Sigillaria elon-gata and S. elegans. Some years later Sordelli(1896), recognized 13 species (see Table 1), ten ofwhich are shared in common with the flora fromthe Alpe Logone/Val Sanagra, including five spe-cies of Sigillaria, and another five species of Ca-lamites, Lepidodendron and Cordaites. He attrib-uted the Manno flora to the early Stephanian, butwith many Westphalian elements. Further studieswere made later by Escher (1911) who confirmedthe presence of Calamites cisti, and Sigillariaelongata, but attributed the last species to S. tes-sellata.

Venzo & Maglia (1947) compared their taxonom-ic list with that of Sordelli (1896) and concludedthat the Val Sanagra flora was older than thatfrom Manno. Venzo (1951) later ascribed the Man-no flora to the transition between the WestphalianB and C substages, thus suggesting it to be slight-ly older than the flora from Alpe Logone. Jong-

mans (1950), related the floras of various Swisslocalities (including Manno), first to the Westpha-lian B-C due to the presence of some elementssuch as Linopteris neuropteroides, Pecopteridium,Sigillariophyllum, Cordaites cf. borassifolius, butlater considered it to be slightly younger, mostlikely Westphalian C (“Mittleres Westphalien”,Jongmans, 1960, p. 95). The taxonomic revisionmade in this paper, together with lithostratigraph-ical affinities of the two localities, permits a clearchronostratigraphic correlation, hence denyingany age-difference between them.

ACKNOWLEDGEMENTThe authors thank Paolo Arduini, Alessandro

Garassino, Giorgio Teruzzi (Museo Civico di Sto-ria Naturale di Milano) and Attilio Selva (MuseoEtnografico e Naturalistico Val Sanagra, Grandolaed Uniti) for their help with the collections andobtaining information. G. Cassinis (Pavia Univer-sity) is also acknowledged by AR for a fruitful dis-cussion on the geology of the area and to tirelesslystimulating us in revising this flora and re- inves-tigating the Pennsylvanian outcrops of W South-ern Alps. G. Santi (Pavia University) gave a fun-damental contribution in the picturing of samplesin the Museum of Milano and Val Sanagra. R. Gi-anotti (Pavia University) is thanked for histhoughtful suggestions and for the time spent onexcursion on some Pennsylvanian outcrop to theW of the area. Thanks also go to E. L. Zodrow(Nova Scotia, Canada) for the consultation of li-nopterid species. The visit of the collection in theMuseo Civico di Storia Naturale di Milano by twoof the authors (Stanislav Oplištil and Josef Pšenič-ka) was enabled with financial support of thegrant project of the Grant Agency of the CzechRepublic (GACR) P210/11/1431. This paper is acontribution to the IGCP 575 project, Pennsylva-nian terrestrial habitats and biotas of southeast-ern Euramerica. We also thank the reviewersChris Cleal and Erwin L. Zodrow for their usefulcomments and grammar revision.

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Table 1List of species published by several authors and the actual list of species. The gray fields represent thecommon species.

VenzoSordelli and Venzo present

1896 Maglia 1951 work1947

Lycopsida Sigillaria rugosa x x xSigillaria cf. inferior xSigillaria cf. ovata xSigillaria tessellata x x x xSigillaria pachyderma xSigillaria cf. micaudi x xSigillaria ocellata xSigillaria elongata x xSigillaria elongata f. minor xSigillaria deutschiana x xSigillaria cf. transversalis x xSigillaria cf. saulli xSigillaria cf. diploderma x xSigillaria reniformis xSigillaria principis xSigillaria principis f. reticolata xSigillaria scutellata xSigillaria scutellata f. elliptica xSigillaria laevigata xSigillaria sauveri xSigillaria intermedia xSigillaria nudicaulis xAsolanus camptotaenia xLepidodendron aculeatum x x xLepidodendron cf. acutum xLepidodendron cf. dichotomum xLepidodendron distatus xLepidodendron cf., obovatum xLepidodendron cf. nathorsti xLepidodendron cf. ophiurusSublepidodendron cf. elegans xLepidophloias cf. laricinus x xLepidophloios macrolepidotus xLepidophloios cf. acerosus xLepidophloios sp. xUlodendron cf. minus xPinacodendron cf. ohmanni xHalonia cf. tortuosa xHalonia sp. x

73




Arctodendron kidstoni xLepidocarpon cf. majus x x xLepidocarpon sternbergi xLepidocarpon lanceolatum xLepidostrobus sp. xStigmaria ficoides x x xStigmaria ficoides f. rimosa xStigmaria ficoides f. laevis xStigmariopsis xKnorria imbricata xKnorria sp. x xBergeria feistmanteli xAspidiaria sp. xBothrodendron sp. x

Sphenopsida Calamites (Diplocalamites) carinatus xCalamites (Crucicalamites) cruciatus x x xCalamites (Stylocalamites) cisti x x x xCalamites (Stylocalamites) suckowi x xCalamites (Stylocalamites) jubatus xC. ramosus forma monobrachiatus xCalamites (Calamitina) schützeiformis xCalamites (Calamitina) schützeiformisf. intermedius xCalamites (Calamitina) schützeiformisf. waldenburgensis xCalamites (Calamitina) undulatus x xAnnularia sphenophylloides xAnnularia radiata x x xAsterophyllites longifolius xCalamostachys cf. tuberculata xAsterophyllites equisetiformis x x xArthropitys cannaeformis x xArthropitys approximates x xArthropitys gigas xSphenophyllum sp. x xPalaeostachya elongata xPalaeostachya aff. schültzi xCalamostachys germanica xCalamostachys ramosa x xCalamostachys sp. cfr. tubercolata x

ferns Lobatopteris cf. simonii xLobatopteris miltonii x x xPecopteris microphylla x

74

Folia vol. 46, No. 1-2, 2012



Pecopteris cf. daubreei xPecopteris cf. arborescens x xCorynepteris (Alloiopteris) sp. xSenftenbergia plumosa x x x

Noeggeratiopsida Noeggheratia cf. tenuistriata xPteridospermopsida Sphenopteris sp. x

Eusphenopteris cf. neuropteroides xDicksoniites cf. pluckeneti xMariopteris latifolia xMariopteris cf. lobatifolia xMariopteris cf. sauveurii xKarinopteris dernoncourti xNeuropteris attenuata xNeuropteris flexuosa xLaveinopteris tenuifolia xAlethopteris sp. (cf. grandinii) xAlepthopteris sp.A xAlepthopteris sp.B xNeuralethopteris cf. larischii xCf. Odontopteris sp. xParipteris gigantea xParipteris linguaefolia xLinopteris obliqua x xLinopteris neuropteroides x xCyclopteris sp. xPalaeoweichselia defrancei xAulacopteris vulgaris xHexagonocarpus sp. xTrigonocarpus bernardi x xTrigonocarpus sp. xCarpolithus sp. xCarpolithes coniformis xCodonospermum sp. xRhabdocarpus costatus x

Cordaitopsida Cordaites sp. xCordaites principalis x xCordaites palmaeformis x xCordaites borassifolius xDorycordaites palmaeformis xCordaicarpus sp. xAspidiopsis aff. coniferoides xCordaicarpus cordai x

75


Tab

le 2

. St

rati

grap

hica

l ex

tens

ion

of i

ndiv

idua

l sp

ecie

s w

hich

are

pre

sent

on

the

stud

ied

loca

lity.

The

dar

k fi

elds

rep

rese

nt t

he t

ypic

al s

trat

igra

phic

exte

nsio

n, t

he l

ight

er f

ield

s re

pres

ent

the

occa

sion

ally

occ

urre

nce.

Bas

hkir

ian

Mos

covi

anK

asim

ovia

n

Yead

onia

nL

angs

etti

anD

uckm

antia

nB

olso

vian

Ast

uria

nC

anta

bria

nB

arru

elan

Step

hani

an B

Lyc

opsi

daS

igil

lari

a ru

gosa

Sig

illa

ria

cf.

infe

rior

Sig

illa

ria

cf.

ovat

aS

igil

lari

a te

ssel

lata

Sig

illa

ria

cf.

mic

audi

Aso

lanu

s ca

mpt

otae

nia

Lep

idod

endr

on a

cule

atum

Lep

idod

endr

on c

f. ac

utum

Lep

idop

hloi

os s

p.H

alon

ia s

p.L

epid

ocar

pon

cf.

maj

us

Sti

gmar

ia f

icoi

des

Kno

rria

sp.

Bot

hrod

endr

on s

p.

Sph

enop

sida

Cal

amit

es(D

iplo

cala

mit

es)

cari

natu

s

Cal

amite

s (C

ruci

cala

mite

s)cr

ucia

tus

Cal

amit

es (

Sty

loca

lam

ites)

cist

iA

nnul

aria

rad

iata

Ast

erop

hyll

ites

lon

gifo

lius

Cal

amos

tach

yscf

. tub

ercu

lata

Ast

erop

hyll

ites

equi

seti

form

isfe

rns

Lob

atop

teri

s cf

. si

mon

iiL

obat

opte

ris

mil

toni

iPe

copt

eris

mic

roph

ylla

Cor

ynep

teri

s sp

.S

enft

enbe

rgia

plu

mos

aP

teri

dosp

erm

opsi

daS

phen

opte

ris

sp.

76

Folia vol. 46, No. 1-2, 2012

Bas

hkir

ian

Mos

covi

anK

asim

ovia

n

Yead

onia

nL

angs

etti

anD

uckm

antia

nB

olso

vian

Ast

uria

nC

anta

bria

nB

arru

elan

Step

hani

an B

Eus

phen

opte

ris

cf.

neur

opte

roid

esM

ario

pter

is c

f. lo

bati

foli

aM

ario

pter

is c

f. sa

uveu

rii

Kar

inop

teri

s de

rnon

cour

tiA

leth

opte

ris

sp.

(cf.

gran

dini

i)A

lept

hopt

eris

sp.

AA

lept

hopt

eris

sp.

BN

eura

leth

opte

ris

cf.

lari

schi

iC

f. O

dont

opte

ris

sp.

Par

ipte

ris

ling

uaef

olia

Lin

opte

ris

obli

qua

Hex

agon

ocar

pus

sp.

Tri

gono

carp

us s

p.C

arpo

lith

us s

p.C

orda

itop

sida

Cor

dait

es s

p.

77


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REVISION OF THE PENNSYLVANIAN FLORA FROM VAL SANAGRA IN WESTERN PART OF THE SOUTHERN ALPS (ITALY)

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