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The Fossil Flora of the Grey Limestones of Veneto, Northern Italy, and Its Relationships to the Other

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The Fossil Flora of the Grey Limestones of Veneto, Northern Italy, and Its Relationships to the Other THE FOSSIL FLORA OF THE GREY LIMESTONES OF VENETO, NORTHERN ITALY, AND ITS RELATIONSHIPS TO THE OTHER . EUROPEAN FLORAS OF SIMILAR AGE ALAN WESLEY Department of Botany, The University of Leeds ABSTRACT De Zigno's almost exclusive devotion to an investigation of the flora from 1852 until The magnificent fossil flora of the Venetian Alps, in rocks of Jurassic age, includes a rich 1891, many new fossiliferous localities were assemblage of bennettitalean, pteridosperm and discovered in the surrounding region and a conifer remains. Pteridophytes and the Cayto­ fine collection of specimens \vas amassed. niales are also represented, but it is extremely During this period De Zigno described and doubtful that the Ginkgoales formed part of the flora. In this latter respect, the Italian flora is illustrated many of the plants in a series of easily distinguished from the majority of other publications which culminated in the two­ European floras of a similar age in which the Ging­ volume Flora .jossilis formationis oolithicae koales usually forms a conspicuous element. (1856-85). This work remained uncompleted It is also remarkable for the presence of Phyllo­ theca, a genus usually considered as confined to the at his death and the projected third volume former Gondwana continent. This unusual occur­ was never published. rence of a typically southern element on the modern There was negligible attempt by De European continent, north of the Mediterranean Zigno to examine the fossils microscopically, sea, considered together with certain geological data, is suggestive that the present location of the and hardly more by Grandori who intro­ fossil flora may be different from that where it duced a revision of the Italian species actually grew in Jurassic times. described in the first volume of the Flora fossil£s (GRANDORI, 1913). This was un­ INTRODUCTION fortunate, since many of the specimens still retain organic matter from which it is HE presence of a fossil flora in the often possible to prepare good cu ticle an d to Jurassic rocks of Northern Italy at learn something about the microscopic T a latitude of approximately 46°N structure of the plants. In the course of has been known since 1764 when the first re-examination of De Zigno's specimens by fragments of plants were discovered at modern methods I have been able to fill Rotzo. Nevertheless, almost a century was certain lacunae in our knowledge of the Grey to elapse before any serious study of the Limestone plants. To date I have been flora was to be undertaken. As a result of able to clarify the systematic position of --+ Geographical distribution of Jurassic floras in Europe and Greenland (the Spanish and Portuguese localities are omitted) Lower Jurassic: Middle Jurassic: 1. Greenland (Scoresby' Sound) 16. England (Yorkshire) 17. England (Stonesfield) 2. S. Sweden 3. Denmark (Bornholm) 18. France (Mamers) 4. Roumania (Anina) 19. Sardinia 20. W. Ukraine (Kamenka) 5. England (Lyme Regis) 6. France (Normandy) 21. Crimea 7. France (St. Honorine, Orne) 8. France (Lorraine) Upper Jurassic: 9. France (Vendee, Deux Serves) 22. Scotland (Sutherland) 10. France (Lozere) 23. England (Portland) 11. Italy (Veneto) 24. France (Chateauroux. Indre) 12 Hungary (Pees) 25. France (C6te d'Or) 13. S. W Germany (\Vurttemherg) 26. France (Dordogne) 14. N. W Germany (Brunswick) 27. France (Cirin, Orbagnoux, Ain) 15. S. Poland (Grojec) JR. Germany (Nussplingen, Solenhofen) 12! " .... ..:: N X :. ~:...,... :",' ...•... ...... ()o 126 THE PALAEOBOTANIST Sphenozamiles (WESLEY, 1958), and also to ceae is thought to be represen ted by the give the first descriptions and illustrations Coniopteris-like Hymenophyllites leckenbyi of the coniferous shoots which occur in the ligno [known elsewhere as Sphenopteris Italian flora (WESLEY, 1956), Further leckenbyi (ligno) Halle]. Its somewhat work, at present in progress, is producing wedge-shaped pinnules, as well as the some interesting results which it is hoped strongly anadromic first order of branching will be published before long. of the frond, set it apart from the most The present survey of the flora and its closely similar species, Coniopteris simplex relationships to other floras of similar age (L. &. H.) Harris (formerly known as Tym­ in Europe must inevitably be somewhat panophora racemosa L. & H.). Fragments tentative, because much still remains to be of Phlebopteris polypodioides Brongn. and done before all the species have been revised. specimens of Laccopteris rotzoana ligno, the Further investigations of the plant fossils latter more correctly assigned to M aton-i­ will quite likely lead to some new inter­ dium, point to the occurrence of the Mato­ pretations and comparisons and some modi­ niaceae. The specimens called M arzaria fication of the views presented here could paroliniana ligno are clearly parts of fern follow. It should also be remembered fronds and belong to the Phlebopteris-Mato­ that many of the determinations of species nidium complex. There is absolutely no and their occurrences in a number of the evidence whatsoever, for regarding M. other European floras date from the latter paroliniana as a detached whorl of equiseta­ part of the last century. A badly needed lean leaves, even though Harris (1961) has reworking of these would doubtless lead to found that the generic name was misused further modification and reassessmen t of for some Yorkshire specimens now referred views. -to Annulariopsis simpsoni (Phillips) Harris. Two reasonably well defined species of COMPOSITION OF THE FLORA Equisetites are also present representing another group of pteridophytic plants. E. The whole aspect of the flora is certainly bunb2tryanu,s ligno, in general, is a slender different from that as vi.sualized by De form, while the other, larger form with many ligno, due, of course, to the later recogni­ leaf teeth (E. veronensis ligno), recalls the tion of such extinct taxa as the Caytoniales, well known Equisetum columnare Brongn., Bennettitales and pteridosperms. Thus but appears to be quite distinct from it. many of the presumed ferns have been A quile unexpected articulate plant, Phyl­ transferred to other groups with the result lotheca brongniartiana ligno, also occurs. that the true ferns appear to have consti­ Another less common group in the flora tuted a very minor element in the flora. is the Caytoniales. De ligno assigned his The Filicales are represented by relatively specimens of Sagenopteris to four species, few specimens vl'hich are for the most part but these vl'ere reduced to two by Grandori fragmentary and not securely identified. - S. nilssoniana (Brongn,) Ward and The generic determinations which have been Pseudosagenopteris angustifolia (ligno) made suggest that only five families occurred. Grandori. The larger of these two, S, Gleichenites elegans ligno unfortunately nilssonicma (syn: S. rhoifolia Presl), origi­ is non-fertile so that its attribution to the nally called S. goeppertiana by De ligno Gleicheniacefle is by no means certain. and vl'hich name has been retained by However, its appearance and mode of Vakhrameev (1964), typically has four branching is very typical of the family and leaflets and is much like the Yorkshire S. I have little reason to doubt the correctness colpodes Harris (HARRIS, 1964) but differs of the iden tification. One small piece of in having broader leaflets with very obtuse Dictyophyllum (originally called C. a111ptop­ apices. It is not known yet whether there teris jurassica Goepp.) indicates the presence are any cuticle characters by which it of the, Dipteridaceae, as does .possibly may be further distinguished. The smaller Protorhipis asarifolia ligno. Nothing is form characteristically has five leaflets, known of the soral or sporangial characters but sometimes four, which caused Grandori of Protorhipis, but its entire, orbicular to adopt the combination Pseudosagenopteris lamina is quite distinct from the divided angustifolia. While the presence of a fifth forms which are usually called H a'ltsmannia leaflet may be nothing more than an abnor­ lind referred to this family. The Dicksonia- mality of a typical Sagenopteris, where four WESLEY - FOSSIL FLORA OF THE GREY LIMESTONES OF VENETO 127 leaflets are a generic character, it seems Mesozoic genera, but 1 believe there is useful for the moment to continue to use adequate reason for maintaining it as a this distinction. The species is very much separate taxon. De Zigno described five like the small forms of Sagenopteris phillipsi species, but Grandori reduced these to a (Brongn.) Pres!, but the cuticle characters single group, D. visianica Zigno. I believe of the Italian form still remain unknown. that neither was quite right and that two An in teresting poin t is the almost certain species may be defined. Finally it may be absence of any representative of the Gink­ mentioned that D1:chopteris shares with goales. Grandori considered that two of Dicroidium the common feature of a forked the specimens may be referable to this rachis. group, but a preliminary examination leads Cycadopten's is also a problematical genus, me to believe that insufficien t evidence is it having been retained as distinct from to hand. Cyclopteris minor Zigno, which L017latopteris, Odontopteris, J(irclmeria and De Zigno regarded as a fern, is a small ThinnJeldia by some authors: but merged specimen that is not securely identified with with one or more of these genera by others. GingllOites, even though the fan-shaped, In my opinion it should retain its rank as entire and petiolate, lamina shows dichoto­ a distinct genus, even though it has much mous venation. There is no trace of the in common with Lomatopteris. Again I lamina having bew bilob€.d. Nor does the cannot quite agree with De Zigno, who dichotomously forked Tre.visania Jurcellata drfined four species, or Grandori, who reduc­ Zigno (renamed Trichopitys lindleyana ed these to one under a different name Schimp. by De Zigno in an unpublished [Lcmatopteris jurensis (Kurr) Schimp.]. I MS of later date) appear to be a slender think that two species can be defined, C.
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