The Jurassic Plant Fossils of the Lovisato Collection: Preliminary Notes

Bollettino della Società Paleontologica Italiana, 51 (2), 2012, 71-84. Modena, 28 settembre 2012

The Jurassic plant fossils of the Lovisato Collection: preliminary notes

Giovanni Giuseppe Scanu, Evelyn Kustatscher & Paola Pittau

G.G. Scanu, Dipartimento di Scienze Chimiche e Geologiche, Università di Cagliari, via Trentino 51, I-09127 Cagliari, Italy; [email protected] E. Kustatscher, Museo di Scienze Naturali dell’Alto Adige, via dei Bottai 1, I-39100 Bolzano, Italy; [email protected] P. Pittau, Dipartimento di Scienze Chimiche e Geologiche, Università di Cagliari, via Trentino 51, I-09127 Cagliari, Italy; [email protected]

KEY WORDS - Palaeobotanical collection, Domenico Lovisato, Middle Jurassic, Sardinia, Museo Sardo di Geologia e Paleontologia “D. Lovisato”.

ABSTRACT - A preliminary account is presented herein of the revision of 473 slabs containing macrofossil plant remains from the Domenico Lovisato plant Collection. The latter is housed in the Museo Sardo di Geologia e Paleontologia “D. Lovisato” of the Università di Cagliari. The material examined encompasses palaeobotanical remains collected from the Genna Selole Formation and from the basal part of the Dorgali Formation, both of Jurassic age at a variety of localities in Barbagia and Sarcidano, eastern Sardinia. A reconstruction is given of the history of the collection, initiated in 1888, and the scientific studies of the flora by a variety of palaeobotanists. Based on a preliminary revision of the specimens the following genera are recognized: Phlebopteris Brongniart, 1836, Hausmannia Dunker, 1846, Coniopteris Brongniart, 1849, Cladophlebis Brongniart, 1849, Sagenopteris Presl in Sternberg, 1838, Cycadeospermum Saporta, 1875, Ptilophyllum Morris in Grant, 1840, Williamsonia Carruthers, 1870 emend. Harris, 1969, Weltrichia Braun, 1847 emend. Harris, 1969, Taeniopteris Brongniart, 1828, Czekanowskia Heer, 1876 emend. Harris et al., 1974, Brachyphyllum Brongniart, 1828, Elatocladus Halle, 1913 emend. Harris, 1979 and Carpolithes Brongniart, 1822. Some plant remains have been putatively assigned to the following genera: Ptilozamites Nathorst, 1878 emend. Antevs, 1914, Nilssonia Brongniart, 1825, Pterophyllum Brongniart, 1828 and Geinitzia Endlicher, 1847. Several of these genera are known also from the Jurassic flora of Yorkshire and from other Jurassic floras of Italy.

RIASSUNTO - [La flora giurassica della Collezione Lovisato: nota preliminare] - In questo lavoro vengono presentati i dati preliminari della revisione sistematica di 473 reperti contenenti resti macroscopici di piante fossili della Collezione Lovisato, conservata presso il Museo di Geologia e Paleontologia Domenico Lovisato, dell’Università degli Studi di Cagliari. La collezione comprende resti fossili raccolti in diverse località del Sarcidano e della Barbagia, nella Sardegna orientale, e provenienti dalla Formazione di Genna Selole e dalla porzione basale della Formazione di Dorgali, entrambe di età medio giurassica. Viene ricostruita la storia della collezione e degli studi effettuati su di essa dai vari studiosi che si sono succeduti nel tempo, a partire dal 1888, anno di inizio delle raccolte e dei campionamenti da parte del Prof. Domenico Lovisato. La revisione tassonomica preliminare consente di fornire una lista aggiornata a livello generico delle forme riconosciute nella collezione, che sono: Phlebopteris Brongniart, 1836, Hausmannia Dunker, 1846, Coniopteris Brongniart, 1849, Cladophlebis Brongniart, 1849, Sagenopteris Presl in Sternberg, 1838, Cycadeospermum Saporta, 1875, Ptilophyllum Morris in Grant, 1840, Williamsonia Carruthers, 1870 emend. Harris, 1969, Weltrichia Braun, 1847 emend. Harris, 1969, Taeniopteris Brongniart, 1828, Czekanowskia Heer, 1876 emend. Harris et al., 1974, Brachyphyllum Brongniart, 1828, Elatocladus Halle, 1913 emend. Harris, 1979 e Carpolithes Brongniart, 1822. Anche i generi Ptilozamites Nathorst, 1878 emend. Antevs, 1914, Nilssonia Brongniart, 1825, Pterophyllum Brongniart, 1828 e Geinitzia Endlicher, 1847 possono essere dubitativamente segnalati per questa flora. Un numero consistente di generi segnalati per il Giurassico della Sardegna sono noti sia nella flora giurassica dello Yorkshire (Inghilterra) sia nelle altre flore giurassiche del territorio italiano, come ad esempio presso la località di Rotzo, in Veneto.

INTRODUCTION from Cagliari. In 1957 all the palaeontological and mineralogical collections returned to the University and The collection of the Sardinian Jurassic plant remains, were housed in the Dipartimento di Scienze Chimiche e housed at the Geological and Palaeontological Museum Geologiche. During the fifties and sixties of the twentieth of Cagliari University (Sardinia, Italy), consists of century, small additions were made to the original several hundred specimens coming from fossiliferous Lovisato Collection. levels distributed over a wide area (Fig. 1) representing From this collection, 473 slabs were chosen from lacustrine-fluvial to coastal transitional environments. those collected by Lovisato and Tornquist and specifically From 1888 to 1916 Domenico Lovisato (Fig. 2), identified by Krasser (1912, 1913, 1920) and Edwards Professor of Mineralogy and Geology and Director of the (1929). Regio Museo Mineralogico e Geologico dell’Università di The aim of this study is to show the historical Cagliari, created a collection based on plant remains from background of one of the most important Italian floras of several localities in the Sarcidano and Barbagia areas of the Jurassic and to give an overview on previous studies central-eastern Sardinia; his original collection numbered carried out on the historical collection of Lovisato and about 1200 specimens. The collection was allocated in the on other specimens collected from the same area. A brief, Natural History section of the Regio Museo housed in the preliminary overview on the flora is given. Finally, this Belgrano Palace in Cagliari, property of the University. paper will show the high potential of this flora in order to Bombings in the 2nd World War severely damaged the gain a better picture of the Jurassic floras of Italy and of the collections, that were transferred to a building in Ghilarza palaeoenvironmental and palaeoclimatic reconstruction (province of Oristano) in 1946, some tens of kilometres during this time period.

ISSN 0375-7633 doi:10.4435/BSPI.2012.9 72 Bollettino della Società Paleontologica Italiana, 51 (2), 2012

Fig. 1 - Schematic map of the Tacchi area with the locations of sampling: a) Dispensa Genna Selole; b) Fermata Cignoni; c) Nuraghe Sutta Corongiu (from Dieni & Massari, 1985 and Costamagna et al., 2007, modified).

LOVISATO’S PLANT COLLECTION 1902, 1904a, b; Krasser, 1912, 1913, 1920; Edwards, 1929) comprise a set of about 1200 rock slabs and only Domenico Lovisato was born at Isola d’Istria (Izola, in 73 index cards of taxa (Fig. 3). The astonishing difference Slovenian) in 1842 and came to Sardinia after he returned between the number of fossils and the small number of from the Patagonia and Tierra del Fuego expedition of the cards is due to their loss during the 2nd World War. The Società Geologica Italiana (1881-83), since the Austrian plant remains were kept in cardboard boxes and had administration declared him politically inconvenient for been numbered. From 1960 onwards, Comaschi Caria his fervid Italian patriotic feeling. He gained the position created a new inventory list when she rearranged the as Professor of Geology and Mineralogy at the Regia whole museum collection after the war. Very recently, Università di Cagliari where he died on February 1916 293 slabs, stored at the the Lovisato Collection of the after a fruitful life spent for the geological research in Museo Sardo di Geologia e Paleontologia “D. Lovisato” the field. He was very interested in Sardinian geology, (MGPDL), have been numbered in accordance to their palaeontology and mineralogy and, together with Gennari position in the ICCD General Catalogue (Central Institute (Botanist), added numerous collections of fossils and of the Catalogue and the Documentation of the Italian minerals to the Regio Museo dell’Università di Cagliari. Heritage Board). The name given in the original card often Part of the collection will be discussed here. disagrees with the taxon name reported in the inventory The materials collected by Lovisato and Tornquist and list; some cards are annotated with titles of literature, described by Tornquist, Krasser and Edwards (Tornquist, and Lovisato’s or Krasser’s or Edwards’s remarks on the G.G. Scanu et alii - Jurassic flora of Sardinia 73

Yuccites, Caulopteris and Pterophyllum (Lovisato, 1903). Sterzel suggested a Triassic age for this flora. Tornquist (1902), in his paper on the geology of Sardinia, identified Jurassic genera such as Ptilophyllum. In 1904 Tornquist studied specimens collected by Lovisato and, partly, by himself (Tornquist, 1904a, b). In these papers he identified Jurassic species such as Ptilophyllum (Williamsonia) pecten, Otozamites beani and Coniopteris cf. arguta and confirmed the Jurassic age. Few years later, Krasser (1912, 1913, 1920) studied the Lovisato Collection coming from Laconi’s sandstones and organic shales. He attributed many remains of male flowers and seeds to Cycadeospermum impressum based on the affinity to Williamsonia leckenbyi. New species, such as Cycadeospermum sardinicum, Cycadeospermum persica and Cycadeospermum lovisatoi were erected as well. The holotypes and paratypes of these species are indicated as stored at the University of Prague (Comaschi Caria, 1959) but are missing today (pers. comm., Jiří and Zlatko Kvaček, 2012). Edwards (1929) revised Krasser’s taxa and moved Krasser’s species Laconiella sardinica to Gristhorpia nathorsti, Cycadeospermum sardinicum to Araucarites sardinicum, Otozamites lovisatoi to O. veronensis and Cheirolepis setosa to Cryptomerites divaricatus. Sagenopteris goeppertiana was considered close to S. phillipsi and Podozamites lanceolatus to Eretmophyllum spp., also known from the Yorkshire flora (Edwards, 1929; Comaschi Caria, 1959). The revision of the whole Fig. 2 - Portrait of Professor Domenico Lovisato (1842-1916). taxa with descriptions of the published and unpublished taxa (Fig. 3).

PREVIOUS WORKS

Meneghini (1857) was the first to note fossil plants from the Jurassic basin of central Sardinia that had been collected by Lamarmora, published and figured in the famous scientific diary “Voyage en Sardaigne” (Lamarmora, 1857). Meneghini kept in Pisa the studied material that was collected at Laconi and at San Antonio near Nuraghe Adòni (Sarcidano area) and near Tonara (Barbagia). He reported the presence of Brachyphyllum majus and Calamites lehmannianus. The flora was considered of Jurassic age. In a short note De Stefani (1891), Director of the Regio Museo di Fisica e Storia Naturale di Firenze, reported conifers (Voltzia sp.) and Equisetales (Equisetum cf. arenaceum) at the base of the dolomitic massifs at Tonara and Laconi, central Sardinia. This taxonomic interpretation led him to attribute the basal fluvial conglomerate with flora remains to the Triassic, analogous to the “Muschelsandstein” of Vosges. Lovisato never published the material collected by him, but many preliminary descriptions, with new species as well, were written by him in small cards added to the samples (Fig. 3). In 1902 Lovisato sent samples, collected in the Fig. 3 - Selection of some original taxa index cards (from the Sarcidano area, to Sterzel in Chemnitz, who indicated Historical Archive of the Museo Sardo di Geologia e Paleontologia the following genera for the flora: Albertia, Calamites, “D. Lovisato”). 74 Bollettino della Società Paleontologica Italiana, 51 (2), 2012

genus Laccopteris (junior synonym of Phlebopteris) led 1. massive quartz conglomerate and stratified the author to confirm the presence of L. polypoidioides sandstone lenses representing deposits of distal alluvial and L. woodwardii. Finally, Comaschi Caria (1959) gave fans and braided rivers are spread in the Laconi-Gadoni an overview on the Lovisato’s Collection and published territories; a list of all fossil plants coming from Jurassic sediments 2. well-stratified fine sandstone, clayey siltites with of Sardinia. She indicated also several taxa from a new organic shales and coal are very productive in plant outcrop near Laconi (Nuraghe Enna): Chladophlebis remains. This facies crops out all over the surroundings denticulata, Equisetum arenaceum, Pachyphyllum of Nurri-Escalaplano territories and is the expression of araucarinum, Ptilophyllum (Williamsonia) pecten and alluvial, palustrine and upper delta plain sedimentation; Taeniopteris vittata. In the 60s, Prof. Charrier revised a 3. well stratified sandstones alternating with marls very small part of the Lovisato Collection, studying only and marly limestones with thin wood coal beds yielding the specimens shown at the museum. He did not publish a lagoonal brackish and marine fauna including bivalves, the systematic revision, but annotated the revised names brachiopods, gastropods, ostracods, foraminifers, on the small index taxa cards. The latest findings are charophytes and echinoderms. These facies develop from reported in Dieni et al. (1983) in a paper on the Genna Ussassai - Perdasfogu - Baunei up to Tavolara representing Selole Formation at Baunei (Ogliastra Province), where deposits of littoral and deltaic setting, transitional to the only Ptilophyllum hirsutum and Nilssonia orientalis are marine Dorgali Formation. Its major development is to the mentioned. For a detailed list of the species indicated by south, in the Sarcidano area (Laconi and Nurallao plant the various authors, see Tab. 1. fossiliferous sites) where it reaches a maximum thickness Del Rio (1976, 1984) studied the palynomorphs of 50 m. The conglomerate and sandstone of the Genna revealing a higher number of taxa than recognized in the Selole Formation are of industrial interest; thus, quarries macro-remains. The general picture of the microflora offer good sections to collect fossil plant remains. corresponds to a Middle Jurassic flora developed in The fossil plant remains of the Lovisato Collection alluvial plains and coastal environments. come from several quarries and natural sections at the base

GEOLOGY AND STRATIGRAPHY: BRIEF NOTES

The Genna Selole Formation in the Tacchi area (Fig. 1) ranges from 5 to 50 m and contains the most abundant plant fossils found in Sardinia, where the entire Jurassic System can reach about 1000 m of thickness. During the Middle Jurassic (Bathonian) extensive areas in eastern Sardinia were subjected to a marine transgression that covered a thin succession of continental and transitional deposits (Genna Selole Formation), unconformably lying over the folded and flattened Palaeozoic Basement (Fig. 4). The Genna Selole Formation is continental at the base and evolves towards transitional and coastal sediments upwards. It starts with alluvial fan deposits consisting of conglomerates made of Permian volcanites and other Palaeozoic rocks; then, fluvial conglomerates mainly built by well-rounded quartz pebbles deriving from the erosion of the Palaeozoic phyllitic basement follow. These likely represent deposits of the middle and distal portion of alluvial fans (Dieni et al., 1983). The quartz sandstones and mudstones with grey and black pelitic layers are rich in macro-remains such as leaves, seeds, fructifications, small amber drops, and micro-remains like pollen grains, spores and foraminiferal linings. These sediments are interpreted as abandoned channels or temporary ponds (Dieni et al., 1983). The following transitional sediments are lagoonal tidally- influenced and include layers rich in brackish-water molluscs. The sandy-marly oolitic limestones of the basal Dorgali Formation with interlayered thick layers of black organic shales also hold abundant plant remains. The continental succession is overlain by a marine oolitic Fig. 4 - Schematic representation of the Genna Selole Fm. and the limestone of the Dorgali Formation that ranges 200-300 basal part of the Dorgali Fm. (not in scale). Explanation of the legend: 1: Hercynian basement; 2: conglomerates; 3: pebbly sandstones; 4: m in thickness (Fig. 4). sandstones; 5: pelites; 6: marly limestones; 7: limestones; 8: sandy The succession is laterally discontinuous and overall, oolitic dolostones; 9: oolitic dolostones; 10: dolostones; 11: seeds; in the area, three lithofacies-types are recognized 12: ferns and pteridosperms; 13: cycadophytes; 14: ginkgoes and (Costamagna et al., 2007): conifers; 15: molluscs; 16: bioturbations. G.G. Scanu et alii - Jurassic flora of Sardinia 75

TAXA Tornquist (1902, 1904b) Krasser (1912, 1913, 1920) Edwards (1929) Comaschi Caria (1959) Dieni et al. (1983)

Albertia brauni Schimper, 1837 x Albertia latifolia Schimper, 1837 x Araucarites sardinicus (Krasser) Krasser, 1920 x x x Baiera phillipsi Nathorst, 1880 x x x Brachyphyllum expansum (Presl in Sternberg) Seward x x emend. Kendall, 1949 Brachyphyllum mammillare Brongniart, 1828 x x x Calamites cistii Brongniart, 1828 x Calamites sp. x Carpolithes sp. x Caulopteris sp. x x Cladophlebis denticulata (Brongniart) Fontaine, 1889 x x x Coniopteris arguta (Lindley et Hutton) Seward, 1900 cf. x x cf. Coniopteris hymenophylloides (Brongniart) Seward x x x emend. Harris, 1961 Cryptomerites divaricatus Bunbury, 1851 x x Cycadeospermum impressum Nathorst, 1879 x Cycadeospermum lovisatoi Krasser, 1912 x x x Cycadeospermum persica Krasser, 1912 x x Cycadeospermum sardinicum Krasser, 1912 x Czekanowskia murrayana (Lindley et Hutton) Seward, 1900 x x x Dadoxilon sp. x x Dictyophyllum rugosum Lindley et Hutton, 1831 x x x Equisetites columnaris Brongniart, 1928 x x x Equisetum arenaceum (Jaeger) Schenk, 1864 x x Equisetum mougeoti Brongniart, 1928 x x Eretmophyllum lovisatoi Edwards, 1929 x x Gristhorpia nathorsti Thomas, 1925 x x Klukia exilis (Phillips) Raciborski, 1890 x x x Laccopteris woodwardi (Leckenby) Seward, 1899 x x x Nageiopsis anglica Seward, 1900 x x x Neocalamites lehmanianus Goeppert, 1836 x Nilssonia compta Brongniart, 1828 x x x Nilssonia orientalis Heer, 1878 x Otozamites beani (Lindley et Hutton) Brongniart, 1849 x x x Otozamites lovisatoi Krasser, 1913 x Otozamites veronensis Zigno, 1881 x x Pachyohyllum araucarinum (Pomel) Saporta, 1870 x Pagiophyllum williamsoni Brongniart, 1828 x x x Phlebopteris polypoidioides Brongniart, 1836 x x Pterophyllum cf. braunianum Goeppert, 1844 x Ptilophyllum hirsutum Thomas et Bancroft, 1913 x Ptilophyllum pecten (Phillips) Morris, 1841 x x Sagenopteris goeppertiana Zigno, 1885 x x x Sagenopteris phillipsii (Brongniart) Presl in Sternberg, 1838 x Sagenopteris rhoifolia Presl, 1838 var. elongata Braun x x Sardoa robitschekii Krasser, 1920 x x x Taeniopteris sp. x Taeniopteris vittata Brongniart, 1831 x x x Thuites expansus Sternberg, 1823 x x Todites williamsoni (Brogniart) Seward emend. Harris, 1961 x x x Voltzia sp. x x Williamsonia acuminata (Zigno) Krasser, 1920 x x Williamsonia leckenbyi Nathorst, 1911 x x x Williamsonia pecten Phillips, 1829 x Williamsonia sewardi Krasser, 1913 x x x Williamsonia whitbiensis Nathorst, 1911 x x Yuccites dubius Schimper et Mougeot, 1844 x Yuccites vogesiacus Schimper et Mougeot, 1844 x x Zamites sp. x x x

Tab. 1 - Summary of the taxa list of the Jurassic Sardinian flora, as enlisted in the historical papers (Tornquist, 1902, 1904b; Krasser, 1912, 1913, 1920; Edwards, 1929; Comaschi Caria, 1959; Dieni et al., 1983). 76 Bollettino della Società Paleontologica Italiana, 51 (2), 2012

of the dolomitic massifs of central Sardinia, Barbagia and well-preserved cycadophytes. Conifers, ginkgophytes and Sarcidano. Quarries for the extraction of quartz are located ferns are abundant, while horsetails and seed ferns are rare. in the surroundings of the villages of Laconi, Nurri, Belvì The sphenophytes (Pl. 1, fig. 1) are documented by and Nurallao (Fig. 1) and the natural sections crop out some fragmentary stem impressions with distinct vascular in the same area. They belong to the same basin and are bundles and nodes. The specimens have been attributed accounted in the following localities: Dispensa Genna to the order Equisetales but due to the bad preservation Selole, Fermata Cignoni, Nuraghe Sutta Corongiu (Fig. 1). of these remains, no attribution on generic level has been possible. Fern fragments have been attributed to the genera PRESERVATION OF THE FOSSILS Phlebopteris Brongniart, 1836 (Pl. 1, fig. 2),Cladophlebis Brongniart, 1849 (Pl. 1, figs 3, 6), Hausmannia Dunker, The plant fossils are preserved in different ways 1846 (Pl. 1, fig. 4) andConiopteris Brongniart, 1849 (Pl. according to the environments in which they have been 1, fig. 5). Some frond fragments with a basal dichotomy buried. Compressions and impressions are the most belong to the genus Phlebopteris. The pinnules are narrow common preservation types. The compression fossils are and totally or broadly attached to the rachis. This genus in most cases represented by leaves with cuticles and are has already been reported from Jurassic sediments with usually associated with fine-grained sediments such as two species, Laccopteris (=Phlebopteris) polypoidioides silt, clay and marl, and/or with organic deposits like peat and L. (=Phlebopteris) woodwardii (Edwards, 1929). The or coal. Mostly the seed ferns, cycadophytes and conifers Cladophlebis fronds are bipinnate with smooth-edged yield well preserved cuticles for most species. pinnules. The veins extend for 2/3 of the lamina. One Impressions are found frequently in the fine-grained additional frond fragment has bigger pinnae and shows rock matrix where remarkable details of the original a very distinct midvein; unfortunately the secondary form are preserved. Occasionally, impressions are also veins are too badly preserved to say if this specimen the counterparts of compression fossils containing belongs to the genus Cladophlebis or perhaps to the the organic material when rock slabs are split apart. genus Dictyophyllum. Krasser (1920), Edwards (1929) Sometimes the impressions are preserved in dirty coarse- and Comaschi Caria (1959) enlisted already the genus grain sandstones mixed with silt and clay. In this case, Cladophlebis from the Jurassic of Sardinia. A specimen the rock slab is generally not well consolidated and has been attributed to the genus Hausmannia. It shows is formed in river and floodplain environments. The a wedge-shaped structure with marked midribs. From organic material is not preserved and has been replaced the midrib secondary veins arise and form rectangular by a clay film that retains a detailed impression of the or square meshes measuring about 2 x 2 mm. At least plant part. Dirty sand impressions are often record of two types of Coniopteris have been identified in the leaves, shoots, fructifications and seeds. Compression flora, with sterile and fertile pinnules. One frond type and impression fossil plants, preserved in fine-grained shows elongated, narrow pinnules, in the other one the and organic sediments, are found in the middle and upper pinnules are broad and with a rounded apex. The fertile part of the Genna Selole Formation and in the basal fronds show terminal sporangia. Krasser (1920) reported Dorgali Formation, whereas impression fossils in coarse C. hymenophylloides (Brongniart) Seward, 1900 emend. grained-sediments are recorded in the basal continental Harris, 1961 and C. cf. arguta (Lindley et Hutton) Seward, lithofacies (Fig. 4). 1900 from the Jurassic flora of Sardinia, whereas Edwards (1929) indicated only C. hymenophylloides. One of the two frond types observed might indeed belong to C. THE FLORA OF THE LOVISATO COLLECTION hymenophylloides (Pl. 1, fig. 5) but only a detailed study will enable us to understand to which species the second A systematic revision of the Jurassic plants of the sterile and type of fertile pinnae fragments belongs. Lovisato Collection is actually in progress. An updated list Two different types of seed ferns have been identified of the flora of the Lovisato Collection consists of 18 genera as well, Sagenopteris Presl in Sternberg, 1838 (Pl. 1, belonging to 7 divisions. The collection is dominated by fig. 7) and perhaps Ptilozamites Nathorst, 1878 emend.

EXPLANATION OF PLATE 1

Horsetails, ferns and seed ferns from the Jurassic of Sardinia.

Fig. 1 - Sphenophyta indet. (NI - MGPDL 9350). Fig. 5 - Coniopteris sp. (NI - MGPDL 9032). Fig. 2 - Phlebopteris sp. (NI - MGPDL 9415). Fig. 6 - Cladophlebis sp. (NI - MGPDL 9418). Fig. 3 - Cladophlebis sp. (NI - MGPDL 9211). Fig. 7 - Sagenopteris sp. (NI - MGPDL 9425). Fig. 4 - Hausmannia sp. (NI - MGPDL 9415).

Scale bar = 10 mm G.G. Scanu et alii - Jurassic flora of Sardinia Pl.77 1 78 Bollettino della Società Paleontologica Italiana, 51 (2), 2012

Antevs, 1914 (Pl. 2, fig. 1).Sagenopteris is characterized fossils with a round-oval shape and a scale-like structure in leaflets with a marked midrib, although the preservation have been attributed to Cycadeospermum Saporta, 1875 of the fragments does often not enable us to see the midrib (Pl. 2, fig. 7); sometimes also small fragments of organic clearly. Comaschi Caria (1959) enlisted two species for material have been preserved. Krasser (1912) described the Jurassic of Sardinia, S. goeppertiana De Zigno, 1885 three different species from the Jurassic of Sardinia: and S. rhoifolia var. elongata Presl in Sternberg, 1838. Cycadeospermum sardinicum, Cycadeospermum Two leaf fragments show rhombic to slightly falcate persica and Cycadeospermum lovisatoi. So far we could segments inserted laterally to the rachis. The veins arise not identify different forms in this group. The genus from the axis to the apex without forking. These specimens Williamsonia Carruthers, 1870 emend. Harris, 1969 (Pl. 2, could be attributed to the seed fern Ptilozamites because fig. 8) includes bennettitalean female fructifications with a of the thick cuticle with isodiametric cells or even to the gynoecium with a minutely reticulated surface surrounded bennettitalean leaf genus Anomozamites Schimper, 1870. by interseminal scales (Pl. 2, figs 5-8). Krasser (1912, Both genera have never been reported in Sardinia before. 1913, 1920), Edwards (1929) and Comaschi Caria (1959) At least seven genera of cycadophytes have been described Williamsonia leckenbyi Nathorst, 1880 from distinguished in the collection. Several segment fragments the Jurassic of Sardinia. Some cup-like structures with with parallel, undivided secondary veins might belong to extensive lobes can be attributed to the genus Weltrichia the genus Nilssonia Brongniart, 1825 (Pl. 2, fig. 6). Only Braun, 1847 emend. Harris, 1969 (Pl. 3, fig. 1). cuticle analyses will permit a more detailed attribution. As already indicated by Krasser (1913), some leaf The leaves attributed putatively to the genus Pterophyllum fragments attributed to Czekanowskia Heer, 1876 emend. Brongniart, 1828 show elongated segments with parallel Harris et al., 1974 (Pl. 3, fig. 2) have also been identified in veins and slightly expanded bases. The segments arise on the flora. These elongated and linear leaves are sometimes the side of the rachis at almost right angle (Pl. 2, fig. 2). dichotomously divided, but not more than one dichotomy In the genus Ptilophyllum Morris in Grant, 1840 (Pl. 2, has been observed in each segment, so far. fig. 3) the segments are attached to the upper surface of Conifers are well represented in the flora, with at least the rachis at an angle of 55-80°; the venation frequency three different genera. In Brachyphyllum Brongniart, is about 9-12 per segment. Tornquist (1904b), Krasser 1828 (Pl. 3, fig. 3) the leaves are generally wider than (1920) and Comaschi Caria (1959) enlisted Ptilophyllum long and helically arranged. The shoots with larger leaves pecten (Phillips) Seward, 1904 from the Jurassic of (1.5-2 x 1.2 x 1.5 mm) could belong to Brachyphyllum Sardinia; a detailed cuticular analysis will be necessary mamillare Brongniart, 1828 enlisted already by Krasser to confirm this attribution. One leaf fragment can be (1913, 1920), Edwards (1929) and Comaschi Caria (1959). assigned to Taeniopteris Brongniart, 1828 (Pl. 2, fig. 4). Only cuticular analyses can confirm this attribution as The entire-margined leaf shows a typical taeniopterid well as the presence of a second species, Brachyphyllum venation arising from the thick midrib. Unfortunately the expansum (Sternberg) Seward, 1919 in the Jurassic of margin is not preserved. Fructifications belonging to the Sardinia. Shoot fragments with elongated, falcate leaves cycadophytes have been identified as well. Impression with an acute apex (up to 5 mm long and 1 mm wide)

EXPLANATION OF PLATE 2

Seed ferns and cycadophytes from the Jurassic of Sardinia.

Fig. 1 - ?Ptilozamites sp. (NI - MGPDL 9328). Fig. 5 - Williamsonia sp. (NI - MGPDL 9223). Fig. 2 - ?Pterophyllum sp. (NI - MGPDL 9430). Fig. 6 - ?Nilssonia sp. (NI - MGPDL 9349). Fig. 3 - Ptilophyllum sp. (NI - MGPDL 246). Fig. 7 - Cycadeospermum sp. (NI - MGPDL 9050). Fig. 4 - Taeniopteris sp. (NI - MGPDL 9334). Fig. 8 - Williamsonia sp. (NI - MGPDL 9419).

Scale bar = 10 mm

EXPLANATION OF PLATE 3

Cycadophytes, ginkgophytes and conifers from the Jurassic of Sardinia.

Fig. 1 - Weltrichia sp. (NI - MGPDL 9345). Fig. 4 - ?Geinitzia sp. (NI - MGPDL 9416). Fig. 2 - Czekanowskia sp. (NI - MGPDL 9274). Fig. 5 - Elatocladus sp. (NI - MGPDL 9373). Fig. 3 - Brachyphyllum sp. (NI - MGPDL 9375). Fig. 6 - Carpolithes sp. (NI - MGPDL 9442).

Scale bar = 10 mm G.G. Scanu et alii - Jurassic flora of Sardinia Pl.79 2 Pl.80 3 Bollettino della Società Paleontologica Italiana, 51 (2), 2012 G.G. Scanu et alii - Jurassic flora of Sardinia 81

Yorkshire (Van Konijnenburg-van Cittert Roverè di Velo Lovisato’s Collection TAXA Rotzo (Wesley, 1966) & Morgans, 1999) (Bartiromo & Barone Lumaga, 2009) (this paper)

Androstrobus Schimper, 1870 x Angiopteris Hoffmann, 1796 x Anomozamites Schimper, 1870 x Araucarites Presl in Sternberg, 1838 x Aspidistes Harris ,1961 x Baiera Braun, 1843 x Bilsdalea Harris, 1952 x Brachyphyllum Brongniart, 1828 x x x x Carpolithes Brongniart, 1822 x Caytonia Thomas,1925 x Cladophlebis Brongniart, 1849 x x Coniopteris Brongniart, 1849 x x x Ctenis Lindley et Hutton, 1834 x Ctenozamites Nathorst, 1886 x Cycadeospermum Saporta, 1874 x Cycadolepis Saporta, 1873 x Cycadopteris Schimper, 1869 x x x Cyparissidium Heer, 1874 x Czekanowskia Heer, 1876 x x Dactylethrophyllum Wesley, 1956 x Deltolepis Harris, 1942 x x Desmiophyllum Lesquereux, 1878 x Dichopteris de Zigno, 1864 x Dicksonia L’Héritier, 1789 x x Dictyophyllum Lindley et Hutton, 1834 x x Eboracia Thomas, 1911 x Elatocladus Halle, 1913 x x x Equisetites Sternberg, 1833 x Equisetum Linnaeus, 1753 x Eretmophyllum Thomas, 1913 x Geinitzia Endlicher, 1847 x ? Ginkgo Linnaeus, 1771 x Gleichenites Bolchovitina, 1959 x Hausmannia Dunker, 1846 x Hepaticites Walton, 1925 x Hirmeriella Hörhammer, 1933 x Hymenophyllites Göppert, 1836 x Klukia Raciborski, 1890 x Leptostrobus Heer, 1876 x Lindleycladus Harris, 1979 x Lomatopteris Schimper, 1879 x Marattia Presl, 1845 x Marskea Florin, 1958 x Marzaria de Zigno, 1867 x Matonia Stephenson et Churchill, 1831 x Nilssonia Brongniart, 1825 x ? Nilssoniopteris Nathorst, 1909 x Osmundopsis Harris, 1931 x Otozamites Braun in Münster, 1843 x x x Pachypteris Karelin et Kirilow, 1842 x Pagiophyllum Heer, 1881 x x x Paracycas Harris, 1964 x Phlebopteris Brongniart, 1837 x x x Phyllotheca Brongniart, 1828 x Pityanthus (Nathorst) Seward, 1919 x Protorhipis Andrae, 1855 x Pseudoctenis Seward, 1911 x Pseudosagenopteris Potonié, 1900 x Pteroma Harris, 1964 x

Tab. 2 - Comparison at generic level of the Jurassic floras from Veneto (after Wesley, 1966; Bartiromo & Barone Lumaga, 2009), Sardinia (this paper) and Yorkshire (after Van Konijnenburg-van Cittert & Morgans, 1999). 82 Bollettino della Società Paleontologica Italiana, 51 (2), 2012

Pterophyllum Brongniart, 1825 x x ? Ptilophyllum Morris in Grant, 1840 x x x x Ptilozamites Nathorst, 1878 ? Sagenopteris Presl in Sternberg, 1838 x x x x Scarburgia Harris, 1979 x Solenites Lindley et Hutton, 1834 x Sphenobaiera Florin, 1936 x Sphenozamites (Brongniart) Miquel, 1851 x Stenopteris Saporta, 1872 x Taeniopteris Brongniart, 1828 x Todites Seward, 1900 x Weltrichia Braun, 1847 emend. Harris, 1969 x x Williamsonia Carruthers, 1870 x x Yuccites Schimper et Mougeot, 1844 x Zamites Brongniart, 1828 x

Tab. 2 - continue

could belong to the genus Geinitzia Endlicher, 1847 (Pl. In 1929, Edwards revised Krasser’s work and 3, fig. 4). The shoot fragments ofElatocladus Halle, 1913 discussed the strong resemblance with the plant remains emend. Harris, 1979 (Pl. 3, fig. 5) are characterized by from England (21 out of 31 taxa). Edwards indicated also elongated leaves (4 x 1.3 mm) arranged in a large spiral. that on a generic level, Sardoa is the only taxon lacking Additionally various seeds, putatively attributed to the in both floras. genus Carpolithes Brongniart, 1822 (Pl. 3, fig. 6), occur This preliminary study makes a new comparison in the sediments. at generic level between the floras of Yorkshire, Rotzo and Roverè di Velo, the latter two from northern Italy, necessary (Tab. 2). THE FLORA OF SARDINIA AND The Yorkshire Jurassic flora is one of the classic OTHER EUROPEAN AND ITALIAN FLORAS and most abundant Jurassic floras in the world. Fossil plants buried in Yorkshire sediments have been studied The Early and Middle Jurassic floras, owing to for example by Harris (1961, 1964, 1969, 1979), Harris their close inter-relations, share the same set of plant et al. (1974), Van Konijnenburg-van Cittert & Morgans communities over a wide geographical area (Vakhrameev, (1999) and Cleal et al. (2001) since the early XX century. 1964, 1991). From England in the west, France and Comparing this flora with the Sardinian one, ten genera Sardinia in the south, throughout the Germanic Basin, have been found in common in both floras after the first Poland, western Ukraine and the Volgi area, Donbas, preliminary revision: Brachyphyllum, Cladophlebis, the southern Urals (Vakhrameev, 1991) and Romania Coniopteris, Czekanowskia, Elatocladus, Phlebopteris, (Dragastan & Barbulescu, 1977-78; Popa & Van Ptilophyllum, Sagenopteris, Weltrichia and Williamsonia. Konijnenburg-van Cittert, 2006) the fossil floras share Only a detailed study will enable us to understand if the a high number of genera in common with the flora of doubtful genera (indicated with the “?” in Tab. 2) are Yorkshire (Harris, 1961, 1964, 1969, 1979; Harris et al., present in the Lovisato Collection or not. These taxa are: 1974; Van Konijnenburg-van Cittert & Morgans, 1999) Geinitzia, Nilssonia, Pterophyllum and Ptilozamites. and differences are mainly at a specific level (Vakhrameev, The historical flora from the Oolite limestone of 1991). Rotzo is the most famous flora from Italy. It has been A first comparison between the Jurassic flora of studied mostly in the 19th century and became famous Sardinia and Yorkshire was done by Krasser (1920). after the studies by De Zigno (1856-67, 1873-85). Later, The author showed a close relationship between the two Grandori (1913), Wesley (1956, 1958, 1966, 1974), Barale floras concluding that of the 37 taxa from the Jurassic (1982) and Thévenard et al. (2005), revised partially of Sardinia, 23 were in common with the Yorkshire the flora, with special interest in the bennettitaleans and Jurassic flora and only seven “endemic” from Sardinia, pteridosperms. Additionally, the historical collection from such as Otozamites lovisatoi Krasser, 1913, Zamites sp., Roverè di Velo, housed at the Museo di Paleontologia of Laconiella sardinica Krasser, 1920, Cycadeospermum the Università di Napoli “Federico II” , has recently been lovisatoi Krasser, 1912, Cycadeospermum persica revised (Bartiromo & Barone Lumaga, 2009). Krasser, 1912, Araucarites sardinicus (Krasser) Krasser, The comparison of the Sardinian flora with the 1920 and Sardoa robitschekii Krasser, 1920 (Comaschi combined list of the revised flora of Wesley (1966) and Caria, 1959). Krasser (1920) considered Sagenopteris Bartiromo & Barone Lumaga (2009) shows differences goeppertiana Zigno, 1885 and Williamsonia acuminata in the floral compositions that could be due to different (Zigno) Krasser, 1920, ancestral forms of the flora of growing conditions and environmental features. Conifers the Oolites of the Veneto Alps and the genus Otozamites with scale-like leaves such as Brachyphyllum or falcate characteristic for the Jurassic flora of Yorkshire, France leaves (Pagiophyllum) dominated the Venetian Alps and of northern Italy. flora but are rare elements in Sardinia. Bennettitales of G.G. Scanu et alii - Jurassic flora of Sardinia 83 the Southern Alps account Otozamites and Ptilophyllum REFERENCES and the seed fern genera Cycadopteris, Dichopteris and Sagenopteris. Only Ptilophyllum and Sagenopteris have Antevs E. (1914). The Swedish species of Ptilozamites Nathorst. been found also in the Sardinian flora. 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