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. (Stonesfield) 2. S. Sweden England 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. tha t 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. gingkophyte of the Baiera-type for there is brauniana Zigno and C. heterophyUa Zigno. evidence of Some sort of articulation (remark­ The most interesting feature of the Veneto ed upon by De Zigno) which is suggestive flora is the abundance of bennettitalean of its having been a branching shoot system remains, especially fronds belonging to the rather than a much divided leaf. genera Otoza11'lites,Sphenoza11dtes and Ptilo­ One major elemtnt 01 the flora. cvmpnses phyllum and possibly also Zamites. The a number of thick-leaved fronds assigned to extreme limits of these genera are not parti­ the genera Cycadopter1:s and Dichopteris. cularly clear and intergradations occur Both types have thick cuticles, especially which often make their separation difficult. Cycadopteris, and for this reason it is sugge~t­Nevertheless, the species are usually well ed that they belonged to the pteridosperms defined. It is certain that Otozamites was and were not ferns as De Zigno considered the commonest representati ve with the them. Numerous isolated seeds, up to an largest number of specimens and species. inch in length occur, and there is a po~sibi­There are at least eight or nine good species lity that some of them may have come from and possibly more. Special interest per­ the plants which bere the fronds called tains to O. bunburyanus Zigno and O. Jeist­ Cycadopteris and Dichopteris. This is pure mantelii Zigno which have long been con­ assumption. however, since there is no sidered as occurring in other more northerly organic connection. I have evidence of the floras. Current work, as yet unpublished, presence of genuine cycads in the flora clearly shows that forms from elsewhere and these seeds could equally have been with closely similar gross morphology belong produced by such plan ts. to different species and that true O. bunbury­ Specimens of Dichopteris, often of large anus and O. Jeistmantelii were probably size and as much as a yard in length, are confined to Veneto. Another characteris­ typified by a single, dichotomous fork of the tic species, not recorded from elsewhere, main rachis. The lamina is pinnately is O. massalongianus Zigno. divided with the pinnae deeply pinnatifid The genus Sphenozann'tes is represented or even divided into discrete pinnules, so by two species of which one, S. rossii Zigno, that the frond may be described as truly is unique amongst the Bennettitales in bipinnate. In many ways Dichopteris ap­ having spinous dentate margins to the more proximates to both Pachypteris and Thinn­ basal pinnae. Jeldia, but it differs in the forking of the Ptilophyll1t111 is also represen ted by at main rachis. Many authors in the past least two species, one large and called have fused Dichopteris with these two P. grandiJoliu1?:/. Zigno, the other smaller, 128 THE PAL.-\EOBOTAl':!ST differing from previously described species, during the early part of the Lias or much and called P. triangulare Wesley in MS. later during the Bajocian s.s. and Bathonian Some Pterophyllum-like fronds have been and Upper Jurassic time. The Italian flora found, but their taxonomic position awaits is thus younger than the well known Thau­ investigation. Anomozamites is not repre­ 1'I1atoperis-f1ora of Greenland, S.V\!. Germany, sented at all. Typical ovulate organs of the Scania and certain parts of France, and Bennettitales have yet to be discovered, but older than the famous Inferior Ooli te flora there are several 'male inflorescences' which of Yorkshire, the Middle Jurassic flora of can be assigned to the genus Weltrichia. Sardinia, and the Upper Jurassic floras of I t is possible that some cycadophytes of Cirin (France) and Nussplingen and other uncertain affinity were present since some places in Germany. The only floras of large taeniopteroid fronds, as well as some approximately the same age are those of peculiar once-pinnate fronds called Ptero­ Bornholm. and Grojec in Poland. phyllum platyrachis Zigno (though not a Petrographic evidence from the limestones true Pterophyllum in any sense), have been in \vhich the plant remains are preserved found. There is also the peculiar entire, strap­ points to an environment like that of the shaped frond, Yuccites schirnperianus Zigno, seas around the Bahamas at the present of which the relationships are unknown. day, where finely crystalline aragonite is Conifers also occur with great frequency being precipitated from ocean waters in and are represented chiefly by Rrachyphyl­ areas of shallow water bordered by man­ lum (five species) and Pagiophyllum (seven groves. The size and mostly good preser­ species). There are also some shoots of vation of the specimens suggests that Ii ttle Elatocladus-type and a peculiar form called drifting of the organs had occurred after Dactylethrophyllum peristt'ctuln Wesley. falling from their parent plants, and that None of these leafy shoots bears reproctuctive sedimentation had been taking place in structures so that their real taxonomic localities away from tidal \v'aters, in calm position within the Coniferae remains un­ water free from wave action and surface known. Nevertheless, I can state that disturbances which would have damaged genuine Stachyotaxus and araucarian cone­ large, entire or pinnatifid leaves. The scales, not yet described, are present. absence of water-worn fragments suggests Summarizing the composition of the flora that the flora was almost autochthonous and as a \\'-hole then we see that it is dominated that it is preserved very near where it grew. by a variety of Bennettitales, with small­ The paucity of ferns and the relatively leaved conifers and plants with large, scrappy nature of their remains could be coriaceous leaves forming other major due to the failure of their delicate leaves to elements. Very much less strongly repre­ be preserved in marine sediments. Tough­ s'ented are the Caytoniales and the pteri­ leaved plants would tend to suffer much less dophytes, while the evidence for the presence from immersion in sea-water and conse­ of gingkophytes is very debatable. In quently their remains would tend to predo­ addition, there are two unusual genera, minate in the preserved record of the flora. Phyllotheca and Yuccites. However, since optimum conditions appear to have prevailed for the preservation of the AGE AND ENVIRONMENT OF THE FLORA plant remains and the flora was most likely autochthonous in origin, we may assume The remarkable composition of the flora that the composition of the flora was not may be nothing more than a reflection of so very different from the record preserved an incomplete record resulting from the in the rocks and that it represents the vege­ particular conditions unde I' which preser­ tation of a particular type of habitat in vation has occurred, but equally it may be which a number of the members were grow­ attributed to its being of an age not repre­ ing under the influence of saline waters. sented aJ;l1ong the plant bearing rocks of We may visualize a brackish swamp with neighbouring regions. thick-leaved plants growing near the shore­ The age of the flora is generally considered line. Quite near and probably on higher on geological grounds to be Upper Domerian ground were the conifers, while further (Upper Middle Lias), and in this respect is away from the influence of brackish water rather unusual since most of the other were the ferns, probably only in small European Jurassic floras flourished either numbers. WESLEY - F0SSIL FLORA OF THE GREY LIMESTO:-

Of course, it may be argued that the an increase in such characteristic genera of predominance of coriaceous types, some of the Oolite as Otozamites and Bracltyphytlum. them showing xeromorphic characters, An analysis of the Italian flora reveals a indicates a relatively arid climate, an idea combination of early Liassic elements such which gains significance in the light of the as Sagenopteris nilssoniana, Stachyotaxus almost certain absence of a gingkophytic and possibly Phlebopteris angustiloba, to­ elemen t. Yet undoubted fern remains do gether with a diversity and abundance of the occur which surely suggests that conditions oolitic genera Otozamites and BrachYPhyl­ of reasonable humidity existed. lum. Lingering on from much earlier time is Yuccites, while Phytlotheca is not recorded elsewhere in Europe. The complete absence RELATIONSHIPS TO OTHER EUROPEAN of Tltaumatopteris, but a combination of FLORAS early Liassic and oolitic elements, the latter A glance at the accompanying map will predominating, suggests that the Italian show that the nearest located early Jurassic flora is of slightly younger age still than the floras to that of Veneto are those of French and Polish floras. This is in accord S. W. Germany (Wurttemberg), S. Poland with its placement at the top of the Middle (Grojec), E. France (Lorraine) and S. France Lias on purely geological evidence. (Lozere), all of which are separated from A comparison of the generic and specific Northern Italy by the alpine arc. It is with composition of the flora with those of Ger­ these floras that immediate comparisons many, France and Poland reveals very few must be made for the equally near floras of similarities. The nearest approach is to the central E. France (Cote d'or and Ain) and French flora, where shared genera are Yuc­ those of Germany (Nussplingen and Solen­ cites, Otozamites, Cycadospadix, Cycadeosper­ hofen) are of much younger Upper Jurassic mum, Brachypltyllwn, Pagiophyllum and age. The only geographically near Middle Equisetites. These genera do not occur in Jurassic flora is that of Sardinia. the German flora, though Otozamites does The German, French and Polish floras make its appearance in the Upper Lias of clearly belong to the first half of the Lias, vVurttemberg together with Pagiophyllum. but whether they are exactly coaeval is not Apart from Otozamites and Bracltyphytlum, quite certain. The differences which exist they do not occur in the Polish flora either. suggest that the French and Polish floras The Italian flora shares Ph.lebopteris and might be slightly younger than that of Sagenopteris with the German and Polish Germany. The German flora is of Lower floras and Stachyotaxus (absen t from Ger­ Liassic age and can be clearly identified many) with the Polish, the three genera not with the well-known Thaumatopteris-flora of being represen ted in the French flora. It Greenland and elsewhere in N. W. Europe, differs from all three trans-alpine floras in even though it contains species which occur lacking Clathopteris, Ctenopteris, Tltinnfeldia, outside the zone in other parts 01 Europe Thaumatopteris and Cladophlebis which are and Greenland (HARRIS, 1937). The Grojec common to all of them. It further agrees flora lacks the large Thinnfeldia element with the French flora in lacking some of the which is so prominent in both Germany and German and Polish common genera such as France. By reason of the fact that it con­ Neocalamites. Marattiopsis, Nilssonia and tains good Thau11'latopteris zone-fossils like Ctem's as well as certain strong coniferous Thaumatopteris itself, Phlebopteris angusti­ and gingkophytic elements loba (Presl) Hirmer & Hbrhammer, Phi. In fact the distinctiveness of the Italian muensteri (Schenk) Hirmer & Hbrhammer, flora is quite remarkable, for not only are Pterophyllum subaequale Hartz and Otozami­ there these dissimilarities, with such genera tes obtusus L. & H., side by side w'ith oolitic as Ctenis, Ano11!ozamites, Nilssonia, Clado­ genera such as Pachypteris, Brachyphyllum phiebt's, Todites and the gingkophytes com­ and Marskea, the Grojec flora must be pletely omitted, but there are also present considered as of slightly younger age, prob­ such forms as Gleichenites, Dt'cltopteris, ably Middle Lias (REYMANOWNA, 1963). Cycadopteris, Cycadospadix, Sphenozamites, The French flora is also probably slightly Dactylethrophytlum, Yuccites and Ph'ytlotheca, younger than the German flora for, though all of which impart an aspect to the flora they are very much alike, it lacks some of which is quite unlike that of any other in the more delicate leaved types and there is Europe. 130 THE PAL.A.EOBOTANJST

The flora of Sardinia, quite clearly of present location of the fossiliferous strata Middle Jurassic age, is also different in aspect is in the Dinaric fold system of the alpine from the Veneto flora, though the presence are, and that these strata are generally con­ of Yuccites and Sagenopteris points to a sidered to have been derived from the remote relationship. folding of deposits that were originally What the antecedents of the Veneto flora accumulated in areas bordering the southern were, or to what new vegetation it gave rise shores of the Tethyan Mediterranean. The are questions which remain unanswered. A intervening body of water would have been small Triassic florule from the Continental a very effective barrier between the deve­ Ladinic stage has been found in the nearby loping floras of the Lias. The idea of the Val Gardena (LEONARDI & PAN, 1953), as flora inhabiting an area on the southern well as an even earlier Permian one (LEO­ shores of the ancient Mediterranean becomes NARDI, 1948). The Ladinic florule com­ more plausible when the presence of Phyl­ prises Yuccites, two species of Pagiophyllum, lotheca is considered. This genus is not Cycadospadix and a V oltzia-like leafy shoot. recorded in floras elsewhere in Europe, and Yuccites, Cycadospadix and Pagiophyllum in general is confined to the former Gond­ are cornman to the Veneto and French wana region whose northern borders formed Lorraine early Jurassic and the Sardinian the southern coastlines of Tethys. Highly Middle Jurassic floras, and it may that a provoking is the recent hypothesis, based connection through some early distribution on palaeomagnetic data, that the Veneto centre in the Mediterranean region can be region occupied a point somewhere in the inferred. The Veneto centre soon became eastern part of the modern Ylediterranean effectively isolated by some barrier from the during Jurassic time (RUTTEN, 1964). Such French region, thus leading to the develop­ ideas could indeed account for the many ment of a flora peculiar in its composi bon peculiarities of the Veneto flora, but very and differing in many ways from other floras much stronger evidence is awaited before of much the same age. It is a fact that the they can be accepted. REFERENCES (This list is purposely restricted in length and is in .no way complete.)

CARlA,!. COMASCHI (1959) Le piante fossili della Idem (1903). Bidrag till Bornholms fossila flora· Sarctegna. Riv. I tal. Paleant . Strat. M em., 7 :7 -177. Gymnospermer. 1<. svenska VetenskAkad. EDWARDS, W. N (1929). The Jurassic Flora of Sar­ Handl., 36: 1-56. dinia. Ann. Mag. nat. His!, N. S 10 (4): 385-394. REYMANOWNA, M. (1963). Review of investiga­ G.OTHAN, W. (1914). Die Unter-Jiassische (Rha­ tions oi Polish Jurassic Floras. Acta Palaeobot., tische) Flora der Umgegend von Numberg. 4: 1-48 Abh. naturh. Ges. Niirnberg, 19: 1-98. RUTTEN, TIL G. (1964). Paleomagnetism and Idem (1935). Die Unterscheidung der Lias- und Tethys. Geo!. Rdsch., 53: 9-16. RhiHflora. Z. dtsch. geo!. Ges., 87: 691. SALFELD, H. (1907). Fossile Land-pflanzen der GRANDORI, L. (1913). La Flora dei Calcari Gri~i Rat- und J uraformation Siidwestdeutschlands. del Veneto, Pt. 1. Mem. Ist. geol. Univ. Pado/'a, Palaeontogl'aphica, 54: 163-204. 2: 45-112. SAPORTA, G. DE (1873-1891). Plantes Jurassiques, HARRIS, T. 1If. (1937). The fossil flora of Scoresby Pal. Franc. 2e. SCr., Vegetaux, 1-4. Sound, East Greenland. Part 5, Stratigraphic SCHENK. A. (1867) Die fossile Flora der Grenz- relations of the plant beds. Medd. Gronland, schichten des Keupers und Lias Frankens. 112: 1-114. Wiesbaden. Idem (1961). The Yorkshire jurassic Flora, 1 ­ SEWARD, A. C. (1900). The Jurassic Flora. 1. Thallophyta and Pteridophyta. British Musewn The Yorkshire Coast. British lViuseum (Natural (Natural History), London. History), London. IDEM (1964). The Yorkshire jurassic Flora, 2­ VAKHRAMEEV, V. A. (1964). Jurassic and early Caytoniales, Cycad ales, and Pteridosperms. Ibid. Cretaceolls floras of Eurasia and the paleoflo­ LEONARDI, P. (1948). Contribllti alia conoscenza ristic provinces oi this period (in russian). della flora delle arenarie di Val Gardena (Per­ Trans. Ceo!. Inst. Acad. Sci. USSR, 102: 1-264. miano n;ledio-inL) dell' Alto Adige. Mftm. I st. WESLEY, A. (1956). Contributions to the Know­ geo!. Min. Un-iv. Padova, 16: 1-16. ledge of the Flora of the Grey Limestones of LEONARDI, P. & PAN, C. (1953) Flora continen­ Veneto, Part 1 Mem. 1st. geol. Min. Univ. tale Ladinica delle Dolomiti. Ibid., 18: 1-23. Padova, 19: 1-69. LUNDBLAD, A. B. (1950). Studies in the Rhaeto­ IDEM (1958). Contributions to the Knowledge Liassic floras of Sweden, 1. K. svenska of the Flora of the Grev Limestones of Veneto, VetenskAkad. Handl., Ser. 4, 1: 1-82. Part 2 Ibid, 21: 1-57. MOLLER, H. (1902). Bidrag till Bornholms fossila ZIGNO. A. DE (1856-1895) Flora fossilis forma- flora. Pteridofyter.Lunds Univ. A'rsskr., 38:1-63. tionis oolithicae, vol. 1-2. Padova.