382 C.BroglioLoriga,A.Fugagnoli, J.VanKonijnenburg-VanCittert,E.Kwsatscher,R.PosenatoGM.lXlachtler

(Fois & Gaetani 1984; De Zanche et al. 1.993; Senow- of the Voltago Conglomerate of the and the bari-Daryan et al. 1993). The Anisian outcrops of the "Vohzia beds" of Recoaro (De Zanche et aI. L993). Prags/Braies Dolomites have been famous in the The plant horizon is about one meter thick and palaeontological literature since the 19rh century, mainly situated iust above bed Kt 1 of Bechstàdt & Brandner on account of the Anisian brachiopods and ammonoids (1970, enclosure 2) (Fig. 1A). Plants are concentrated in of the "alpiner Mwschelbalk" (Loretz 1875; Mojsisovics some centimetres-thick layers of siltstone, marly silt- 1879, 1882; Bittner 1890) and calcareous algae (Pia stone and carbonate siltstone. The fossiliferous layers 1937), while plant remains have remained almost unno- are lens-shaped. Therefore, their number and thickness ticed (e.g. Pia 1937). The only illustrations of Anisian may change laterally. They alternate with silty and marly plants from this area were published by Bechstàdt & limestone layers, in which only sparse terrestrial plant Brandner (1,97A; pL.8, fig. 4; pl. 10, fig. 1). remains occur. Marine biota (bivalves, brachiopods, The preliminary taxonomic analysis of the numer- ammonoids, gastropods and fishes) are present, but not ous and well reserved specimens is based on both macro- abundant, throughout the horizon. Occurrence and morphology and some cuticles and in slla spores; a very preservation of terrestrial (plants and reptiles) and diverse terrestrial plant assemblage emerges which has marine fossils are presumably related to very rapid bur- no comparison with the scarce data hitherto available ial events caused by gravity flows within a marine basin, regarding the Anisian macroflora of the Dolomites, connected with heavy storms in the terrestrial domain which only mentions rare terrestrial plant remains (e.g. (Tintori et al. 2001; Broglio Loriga et aI.2aa2). Mo;'sisovics 1879;Pia 1,937; Ogrlvie Gordon 193a). The At the base of the deposit alayer of carbonate silt- Kùhwiesenkopf flora is the first rich Triassic flora of the stone, about 30 cm thick, represents a marker bed, easi- Dolomites. ly recognizable because its upper surface is rich in The aim of the report is to present the high diver- roughly N-S oriented trunks (trunk bed; Fig. 1B). Large sity of this Anisian plant assemblage which contains new but rare plant remains (trunks) are also present at the taxa both at generic and specific level. In addition, the top of a limestone bank situated at about tv/o metres results of the foraminiferal study of the stratigraphical below (brachiopod bed, Fig. 1B). This unit of nodular segment including the plant horizon are also presented. and massive wackestone and packstone contains a rich foraminiferal assemblage. At the top of the bank, byssate bivalves (Mysidioptera) and brachiopods [e.g. Puncto sp iriferella (Schlotheim) and Angwstothyris Stratigraphical setting and age fragilis angustaeformrs (Bóckh)] are frequent (Tintori et al. The plant horizon crops out for several hundred 2001). This brachiopod bed was already noticed by metres along the steep gullies furrowing the western Bechstàdt Er Brandner (197q, who named it Brachiopo- slope of Monte Prà della Yacca / Kúhwiesenkopf, inside denhorizont 2, and is characterized by the occurrence of the Braies/Prags Natural Park (Fig. 1). It occurs in the Mentzelia mentzeli (Dunker) and Coenothyris awlgaris lower part of the Anisian basinal succession (Fig. 1), (Schlotheim). now named the Dont Formation of the Braies Group The Dont Formation of the Dolomites is tradi- (Pisa et al. 1,978; Fois Er Gaetani 1984; De Zanche et aI. tionally considered Pelsonian-Illyrian in age (Delfrati et 1993; Senowbari-Daryan et al. 1993; Delfrati et al. 2000). al. 2OOO, with references). In the lower part of the Dont This unit is considered a hemipelagic carbonate-terrige- Formation, Upper Bithynian ammonoids have been nous sequence of a marginal basin environment (Senow- cited in the Braies area (Rio Schade - Belvedere sections; bari-Daryan et aI. 1993). DeZanche et aL.1.993; Gianolla et al. 1998). In this section, well-known through the detailed The Anisian formations of the Prags/Braies area study of Bechstàdt & Brandner (197A), the Dont For- were dated by Bechstàdt & Brandner (1,970) mostly on mation is more than 200 m thick. The plant-bearing beds the basis of the brachiopod assemblages; ammonoids, are situated about 75 m above a massive carbonate plat- crinoids and conodonts were also considered. However, form (sample K3, enclosrre 2 of Bechstàdt Ee Brandner the stratigraphic distribution of these latter groups was 1970; Fig. 1A), previously known as the Algen- not reported in detail, but referred only to their lithos- wellenkalk by Bechstàdt & Brandner (1.970), and now tratigraphic units. According to the above cited authors, attributed to the Gracilis Formation (De Zanche et al. the Algenroellenkalk interfingers with the siltitische 1992; Senowbari-Daryan et al. t9911 . The lower part of Mergel und Knollenkalke and its upper boundary ranges the Dont Formation is considered to be as an equivalent from the late Bithynian (eastward, where it may lacking)

Fio | - Srretìorrnhìcrì selling o{ the plant deposir from Kúhwiesenkopf /Monte-Prà della Vacca. A - General column adapted and modified from Bechstàdt & Brandner (1970; enclosure 2). B - Detail of the segment bearing the plant deposit. Symbols: FS - Foraminifer sam- ples; BH - Brachiopod horizons (from Bechstàdt tr Brandner 1970); K3, K11 - bed numbers from Bechstàdt & Brandner (1970, enclo- sure 2) column. Rivista Italiana di Paleontologia e Stratigrafia volume 108 numero 3 tavole 1 pagine 381-390 Novembre 2002

THE ANISIAN MACROFLORA FROM THE NORTHERN DOLOMITES (MONTE PRÀ DELLA VACCA / KÚH\TIESENKOPF, BRAIES): A FIRST REPORT

CARMEN BROGLIO LORIGA" ANNA FUGAGNOLI" VAN KONIJNENBURG - JOHANNA VAN CITTERT,, EVELYN KUSTATSCHER', RENATO POSENATO, 8. MICHAEL WACHTLER,

Received. May 5,2A02; accepted August 9,2002

Key-zaords: macroflora, stratigraphy, Dont Formation, Anisian, to the Cycadophyta (Bj uaia, Taeniopteris, Dioonito catpid.iwm, Ptero - Dolomites, pbyllum / Nilssonia). Coniferophyta are represented by Vobzia, ?Voltzia and Albertia; the latter genus is recorded herein for the first Riassunto. Le conoscenze sulla macroflora triassica delle time in the Middle Triassic of the Dolomites. Besides, two Lycophyta Dolomiti riguardano principalmente il Ladinico, mentre sono scarsi in genera (?Isoetites, Lycophyta nes/ raxon), three Pteridospermae genera letteratura i dati relativi a quella anisica. Questa lacuna viene qui col- (?Sagenopteris, Scytophyllum and Pebaspermum), and one Sphenophy- mata con la scoperta di un giacimento a piante a Monte Prà della Vacca ta genus (Equisetítes) have also been recognized. / Kúhwiesenkopf (Dolomiti di Braies), località già nota in letteratura soprattutto per le faune a brachiopodi e le flore a dasicladali. Le piante si trovano alf interno di un orizzonter spesso circa 1 metro, posto nella Introduction parte inferiore della Formazione di Dont, un'unità bacinale formata prevalentemente da sedimenti emipelagici, terrigeno-carbonatici di età Triassic macrofloras of the Dolomites are rare, and pelsonico-illirica. L'orizzonte a pianre si inserisce in un segmento the majority of the literature refers to Ladinian plants. stratigrafico con foraminiferi e brachiopodi tradizionalmente riferiti al Pelsonico. I macroresti vegetali, numerosi e ben conservatì, compren- After the sparse. vague cirarions of the pioneering dono fronde, corpi fruttiferi, rizomi e legni. researchers (e.g. Wissmann & Mùnster 1841; Mojsiso- L'analisi preliminare, basata sulla macromorfologia, su cuticole vics 1879; Ogilvie Gordon 1927;Mutschlechner 1932), e su spore in situ, ha permesso di identificare 17 generi. Il maggior followed by the first monograph of Leonardi (1953) and numero di taxa appartiene alle Preridophyta (Anomopteris, Neu- brief updatings (e.g. Leonardi 1967; Calligaris 1986), the ropteridium, C ladophlebis, Crenatopteris, I Marattiop sis) ; seguono 1e Cycadophyta (Bjuvia, Taeniopteris, Dioonitocarpidium, Pterophyllwm / biodiversity of the Ladinian rerresrrial plants has been Nilssonia). Alle Coniferophyta appartengono Voltzia, lVobzia ed presented in a recent systematic work by Wachtler & v. Albettìa: quest'ultimo genere è segnalato nel Triassico medio delle Konijnenburg - van Cittert (2000) and Kustatscher Dolomiti per la prima volta. Due generi appartengono alle Lycophyta (2001). (?Isoetites, Lycophyta nuovo taxon) e tre alle Pteridospermae The latter rwo papers are {rìostly based on the (?Sagenopterts, Scytophyllwm e Peltaspermum), un genere alle Spheno- discovery by one author (M. Wachtler) of new Ladinian t E^,,:-^t:'^-, ytt!^1,,.^ td \Lq4t)rLtLO ). plant deposits within the I a Valle/Vengen Formation flora Questa rappresenra la prima ricca documentazione della from the Braies/Prags a(ea and Badia Valley. This mate- vita vegetale terrestre dopo I'estinzione permo-triassica nel Sudalpino. rial, now housed in the Museo di Scienze Naturali del- Abstract. The knowledge of Triassic macroflora from the I'Alto Adige (), was idenrified using both Dolomites mosrly concerns the Ladinian, while literature data on macromorphology and curicular analysis. Further plant Anisian plants are scarce. This gap is filled by the discovery reported collections of the Cortina d'Ampezzo and Ortisei Muse- here, of a rich plant deposit from Monte Prà della Vacca Kúh- / ums have been also considered. wiesenkopf (Prags/Braies Dolomites). The fossils occur in a horizon, about 1 m thick, from the lower part of the Dont Formation, a basinal More recently, M. \fachtler discovered a new and unit mostly constituted by hemipelagic, terrigenous-carbonatic sedi- rich Anisian plant deposit located in the western slope ments of Pelsonian - Illyrian age._The strarigrafic interval with the of Monte Prà della Yacca / Kùhwiesenkopf (Braies/ plant horizon is Pelsonian in age. Prags Dolomites). The fossils occur in a basinal unit, A preliminary sysremaric analysis of the numerous and well which has been variously preserved specimens has allowed the identification of at least 1Z gen- named in the literaturei Prugser era. The taxa belong primarily to the Pteridophyta (A.nomopteris, Neu- Schichten by Pia (1937); siltitische Mergel wnd Knol- ropteridium, Cladopblebis, Cremdtopteris, I M arattiop sts),subordinately lenkalke (Bechstàdt Ec Brandner 1970); Dont Formation

1 Dipartimento delle Risorse Naturali e Culturali, Università di Ferrara, Porta Mare 2,441OOFerrara,Italy. e-mail [email protected] 2 Dipartimento di Scienze della Terra, Università di Ferrara, C.so Ercole I d'Este 32, 44100 Ferrara, Italy. e-mail [email protected] 3 Laboratory of Palaeobotany and Palynology, Budapestlaan 4, 35s4CD Utrecht, Netherlands. e-mai1 [email protected] 4 Rainerstrasse 11, 39038 , Italy. e-mail [email protected] Anisian macroflora fram Dolomites 383

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_ '5\ "? LJ w Plsnt - 55 ,zî\ -*G Y' Arîmonoid @ Algee -5ìK3* a --1* __t"* J -cu -l- -_L 384 C. Broglio Loriga,A. Fugagnoli, J. Van Konijnenburg- Van Cittert, E. Kustatscher, R. Posenato €" M. Wachtler to the Pelsonian (westwards). ature, Pawlbronnimannia judicariensis, a species first dis- Inthe sihitische Mergel wnd Knollenkalke of Monte covered in the "orizzonte a binodosus s. str." (Premoli Prà della VaccalKùhwiesenkopf section, five brachiopod Srlva 1.971. p. 316), is generally recorded after the disap- horizons were reported by Bechstàdt & Brandner (1970) pearànce of M. dinarica even if their coexistence has not (Fig. 1A). The horizons 1 to 3 were considered Pelson- to be excluded (e.g. Premoli Silva 1971,; Gadzicki et al. ian in age, the horizon 4 Pelsonian - early Illyrian, while 1975;Trtfonova 1.979; Oravecz-Sch effer 1,987; Rettori et the horizon 5, occurring about 15 m above a thin bed of aL.1994; Angiolini etaL.1,994;Zanrnerti et al. 1994). This conglomerate (at the top of section in Fig. 1A), was record consequently enables the stratigraphic distribu- dated to the early lllyrian. More recently, Senowbari- tion of P. jwdicariensls to be lowered. All these raxa a.re Daryan et aI. (1.993) suggested a Pelsonian age also for still occurring in the upper part of the section (which the brachiopod assemblage of the horizon 4, because the includes the brachiopod horizons 3 and 4) in which same species are present throughout the Recoaro For- Pawlbronnimannia jwdicariensis becomes the dominating mation in the Braies Dolomites, where iate Pelsonian species (samples 97 to 110). Paulbronnimannella whit- ammonoids have been cited in the upper part (De takeri, which is considered a late Pelsonian marker (De Zanche et al. llsz). Zanche et al. 1,992;Zanrnetl 1994), occws only in the Therefore, according to Bechstàdt & Brandner samDleL.^ I tu. (1970), the plant horizon can be referred to the Pelson- ian as it occurs between the brachiopod assemblages 2 and 3. The macroflora The benthic foraminifera of the examined section A preliminary systematic analysis of the terrestri- are mainly represented by agglutinated, microgranular al plants contained in more than 1OO0 samples, has made and porcelaneous species belonging to Ammodiscidae it possible to determine at least 17 genera belonging to and Cornuspiridae. the following divisions: Lycophyta, Sphenophyta, Pteri- The most significant species are Meandrospira dophyta, Pteridospermae, Cycadophyta and Conifero- dinarica Kochansky-Devidé & Pantic, Pilammina densa phyta. The fossil plant association is dominated by ferns, Pantic, Pilamrninella semiplana (Kochansky-Devidé & conifers and cycadales, while horse-tails and seed-ferns Pantic), Pawlbronnimannia jwdicariensis (Premoli Silva), are rarer. and Pawlbronnimannella wbittaheri Rettori. The majori- The Lycophyú are represented by two genera. ty of these raxa. are well known from several areas of the Tethys domain and their acme has been traditionaily One genus (?Isoetites, Tab,1; Pl. 1, Fig. 2) is character- Isoetites Múnster. referred, sometimes by means of ammonites and con- izedby elongated leaves, similar to It is also rappresented nearly complete individuals with odonts, to the Pelsonian (e.g. Premoli Silva 1'971'; Gadz- by articulated roots, stem and elongated leaves covered, icki et al. tgZS; Salay et al. 1988; DeZanche et al. 1992; still Angiolini et al. 1992; Rettori et aI. 1994; Rettori 1995; at least on one side, by thin hairs (trichomes). The genus Zaninetrí et al. 1994). Isoetites is also known from the Lower tiassic of Ger- In particular, Pilammina densa shows the widest many (Kelber Er Hansch tlls; and China (Wang vertical distribution in the sections. Up to about 50 m z.1,ee6). from the base of the Dont Formation, the above quoted The other form is represented mostly by speci- least species is associated with Meandrospira dinarica and Pil- mens with stems and elongated leaves, covered, at amminella semiplana; this foraminiferal association is on one side, by very thin hairs (trichomes). At first they typícally recognized in the Tethys realm as indicative of were thought to belong to Pleuromeia sternbergii (Mùn- Pelsonian. The plant horizon is included in the strati- ster) Corda (Kustatscher, van Konijnenburg - van Cit- graphical interval bearing this association. tert & Vachtler 20A2), a typical plant of the Anisian of It is worth noting that Pawlbronnimannia jwdi- the Vosges and the Germanic Buntsandstein (Grauvo- cariensis occurs already in the lowest part of the section gel-Stamm,1978; Sander Ec Gee 1,99a); this species is (samples 65, 66). together with Pilammina densa and known from corm, leaves and undivided stem remains before the appearance of M. dinarica. According to liter- with elongated scars up to1 cm in length, giving the stem

Lvcopuvre SpHeNopuvrl PrrruoopttYtR PTERIoospgRMAE CvceoopHYrn CoNrrERopHvrR ? Isoetites Equisetites Anomopteris ? Sagenopteris Bjuvia Voltzia Lycophyta Neuropteridium Scytophyllum Taeniopteris ? Voltzia new taxon Cladophlebis Peltaspermum Dioonitocarpídium Albertia Crematopteris Pterophyllum ? Marattiopsis /Nilssonia

Tab. 1 - Preliminary list of the fossil plant genera discovered in the plant horizon from Kùhwiesenkopf /Monte Prà della Vacca Anisian macroJlora from Dolomites 385

a more or less reticulate structure. F{owever, recent find- the past already been found associated with New- ings at Kùhwiesenkopf of thick rhizomes giving off the ropterid,iwrn leaves (Schimper & Mougeot 1.844, Blanck- stems and stem fragments which show an apparenr divi- enhorn 1885). \fle are novv certain that Crematopteris sion now make this attribution impossible as Pleuromeia represents the fertile parts of Neuropteridium. Although is characterizedby a four-lobed corm and an undivided there is almost no macromorphological difference, we stem. The presenr material will probably have to be may be dealing with 2 species ol Crematopferis as we assigned to a new genus (Lycophyta new taxon, Tab. 1). found two different types of sporangia. The genus Cre- The way the rhizome divides is characteristic for e.g. matopteris has also been cited in the Upper Buntsand- primitive Lycophyres but can also still be found in living stein of Germany (Blanckenhorn 1885; Sander & Gee Lycopodiwm. 1994) and from the Early Triassic of China (Vang The division Sphenophyta is documented by some 2.1996). The first species, with small and roundish spo- stems, generally with only the internodes, isolated stro- rangia, ís Crematopteris typica Schimper Er Mougeot, bili and rare microsporophylls belonging to Eqwisetites also known from the Vosges and Germany (Schimper & Schimper, a genus well known all over the Triassic of Mougeot 1844, Blanckenhorn 1885) and belongs to Europe (i.e. Germany, Kelber & Hansch 1995). Neuropteridium elegans as we have found both attached The Pteridophyta contain five genera: to the same rhizome. The second species of Cre- An om op teris Bron gniart, N e wropteri di um S chimp er, C re - matopteris with the larger, oval sporangia probably mdtopteris Schimper & Mougeot, Cladophlebis Brong- belongs to N. bergense but it has not been found niart and ?Marattiopsis Schimper (Tab. 1). The fern attached to a rhizome. Spores from both Crematopteris genus Anornopteris is probabiy represented by two species are identical trilete globular spores of the type species; one is ,4. mowgeotìi Brongniart, formerly cited found in the Osmundaceae. in the Bunrsandstein of Vosges (Grauvogel-Stamm\978, Cladopblebis specimens are very similar ro rhose Grauvogel-Stamm & Grauvogel 1980) and of the Ger- described in the Ladinian of the Dolomites (\Tachtler & man Basin (Sander & Gee 1994), in the Muschelkalk of van Konijnenburg - van Cittert 2000), from which they France (Depape Er Doubinger 1963), in the Early Trias- differ by their larger pinnules (up to 3 cm in length). sic of China (Wang Z. 1996) and in the Ladinian of the Some incomplete specimens with microsporangia may Dolomites (\Tachtler & van Konijnenburg - van Cittert beiong ro Marattiopsls (see e.g. Wang Y. i,999). 2000). Its leaves may reach 70 cm in length and bear At least two leaf genera have been determined both sterile and fertile pinnules. The other, probably a within Pteridospermae. One (?Sagenopteris, Tab. t) new species of Anomopteris (P1.1, Fig. 5), is rather simi- comprises a few small leaf fragments with dichotomised lar on first sight to A. mowgeotii but differs- in the veins which tend to anastomise, as in Sagenopteris Presl absence of aphlebia; moreover, it is probabiy tripinnate or Neoglossopteris (WangZ. 1996). The other genus has whereas A. mowgeotii is bipinnate. pinnate leaves, ranging from elongated to lanceolate, Newropteridium (Pl. 1, Fig. 4) is represented by a simiiar to Sqttophyllwm Bornemann; it has been found large number of specimens. According to the literature associated with macrosporophyll heads of peltaspermwm they could be assigned to at leasr four species lN. elegans Flarris indicating that these fossils belong to the pteri- (Brongniart) Schimper & Mougeot, N. intermediwrn dosperm order of the Peltaspermales. Some leaves show Schimper & Mougeot, l,'1. voltzii (Brongniart) Schimper traces of insect chewi ng. Scytophyllum and Pelaspermwm & Mougeot and ìy'. bergense Blanckenhornf . N. elegans is are also recorded from the Keuper of Germany (Kelber characterized by diverging and forking veins and pin- & Hansch 1995). nules as wide as they are long, while N. intermedium is At least three genera of Cycadophyte leaves have characterízed by a fusiform disposition of the pinnules, been recognized. One, with the largest leaves (50 cm with the widest located in the middle part. InN. z;oltzii long and 20 cm wide), probably belongs to Bjuaia Florin the two lobes, which cover rhe rachis, are asymmetrical, (Tab. 1; Pl. 1, Fig. 6); its morphology is close to .8. while N. bergense has very large pinnules attached to a dolomitica Wachtler & van Koni;'nenburg - van Cittert rachis broader rhan in the other Neuropteridiwrn materí- (2000) from the Ladinian of the Dolomìtes. Theniopteris al. However, the occurrence of specimens with transi- Brongniart (Tab. 1), a form-genus for undivided cycado- tional characters at least between the first three species phyte leaves, is represented by three species, whose dis- suggests they should be considered as a singie sp..i... tinction is based on leaf size and outline and on vena- Further analysis will be necessary to resolve these taxo- tion; one is Theniopteris angustifulia Schenk also known nomical questions. Neuropteridium has also been from the Keuper of Germany (Kelber & Hansch l9g5). described from the Upper Buntsandstein of the Vosges Moreover, leaves morphologically similar to either the (Gall & Grauvogel-Stamm 1.999), Germany (Blancken- bennettitalean genus Pteropltyllwm Brongniart or rhe horn 1885; Sander & Gee 1994) and the Lower Triassic cycadalean genus ÀIilssoniaBrongniarr have been found; of China (Wang 2.1996). Creruatopterls (Pl. 1, Fig. 3) is cuticular analysis indicates ar least for part of the mate- a genus for fertile, once pinnate fern-leaves that had in rial a possible attribution to Nilssonia which would be 386 C. Brr,,glio Loriga,A. Fugagnoli, J. Van Koniinenburg- Van Cittert, E. Kustarsclter, R. Posenato €t 14. \X/achtler Pl. 1 Anisian macroflora from Dolomites 387 the earliest record of this genus so far (Pterophyllum / This analysis will be applied to all those taxa in which the Nilssonia, Tab.1). cuticle is preserved. Almost complete female fructifications are also The fourth type of conifer leaves present in the present. These consist of a series of macrosporophylls association can be attributed ro rhe genus Albertia with an apical sterile parr, up to 4 cm long, and a basal Schimper (Tab. 1) that is also known from the Anisian fertile part that is up to 3 cm long, on which as many as flora of the Vosges (Grauvogel-Stamm 1928) and the .12 seeds are inserted in two rows. The single scale has a German Keuper (Kelber & Hansch 1995). morphology close to Dioonitocatpidiwm Lilienstern Some male cones have been also found but thev (Tab. 1; Pl. 1, Fig. Z) quoted from the Keuper of the are isolated; it is therefore impossible to derermine to Germanic Basin (Rùhle von Lilienstern 1928; Kelber & which conifer they belong. At least two differenr rypes Hansch 1995), Carnian of Lunz (Kràusel te53; and of isolated female ovuliferous cones have been found as from the Ladinian of the Dolomites (Leonardi 1953; well. Wachtler Er van Konijnenburg - van Cittert 2OOO) even if Research for pollen and spores in sitw also will be generic names sometimes differ. Other macrosporo- performed too. phylls are similar to rhose of the recent cycad Zamia or the fossii genus Beania Carruthers. Conclusions Four different forms have been determined among the Conipherophyta. The first (?Voltzia, Tab. t; Pl. i, The plant assemblage of Kùhwiesenkopf/Monte Fig. 1) shows a wide variability of leaf outlines, ranging Prà della Vacca is the first important record of the Meso- from short needles (1 cm long) with a rounded apex to zoìc flora from the Dolomites after the Permian-Triassic long needles (3-4 cm long) with a pointed apex. Falci- extinction. It occurs in the lower part of the Dont For- form leaves, strongly diverging from the axis, occur mation, a basinal unit of Pelsonian-Illyrian age. The Pel- between those two extremities. On the basis of leaf sonian age of the stratigraphical interval including the morphology, this material might be attributed rc Voltzia plant horizon, already dated by means of brachiopods heterophylla Brongniart known also from the Anisian of (Bechstàdt & Brandner D7A), is confirmed by Spain (Diez et al. 1996), from the Buntsandsrein and the foraminiferal analysis. The material is numerous and Muschelkalk of Germany (Blanckenhorn 1885; Sander well preserved, often yielding cuticles and in sitw spores. & Gee 1994), and France (Schimper Er Mougeot 1844). This assemblage represenrs an important flora Flowever, the female cones found in the deposit, with with several new taxa (e.g. Lycophyta ney/ raxon, ovuliferous scales divided in several lobes and only Anomopteris new species) and some other taxa that are slightly fused at the base, show similarities with those of recorded for the first time in the Dolomite region (e.g. Psewdor:ohzia Florin, a Late Permian conifer generally Albertia).In particular, the Lycophyres are found for the reported from Germany (Schweitzer 1962) and the first time in the Triassic of the region, whereas in China Dolomites (Clement-Vesterhof 1987). For this reason, and in Germany they are already known since the Lower the attribution of these plant remains to Vohzia Brong- Triassic. Some of the new raxa may even give a better niart is questionable. A second form (Vohzia, Tab. 1) is insight in the evolution of certain groups (e.g., Lyco- characterized by elongated triangular needles which are phyta new taxon, cycad female fructifications). inserted perpendicularly to the axes, as in Vohzia The flora has several taxa in common with the recwbariensis Schenk, well-known from the Anisian flora Anisian floras of e.g. the Vosges (e.g. Brongníart 1,828; of Recoaro (Schenk 1868). A third species seems to be Schimper & Mougeot 1844; Grauvogel-Stamm 1978), identical to Voltzia,u,alchiaeformis Fliche (Grauvogel- Recoaro and the German Buntsandstein (e.g. Blancken- Stamm 1978). k is not clear if this form is to be consid- horn 1885; Màgdefrau 1931; Schlùter & Schmidr 1,927). ered as belonging to the first conifer species mentioned In particular, the macroflora reported here is much rich- above, which is characterised by a strong heterophylly, er than that of the historical Vohzia beds of Recoaro, or if it represents a different raxon. Probably only cutic- which are the most famous Anisian plant deposit of the ular analysis will provide an answer to this question. Southern Alps (e.g. Catullo 1827, 1847; v. Schauroth

PLATE 1

Some fossil plant specimens from the Anisian of Kúhwiesenkopf /Monte Prà della Vacca plant horizon. Scale bar is 2 cm.

Fig. 1) - Primary shoot of conifer bearing secondary shoots, both with spirally arranged leafes; the specimen is macromorfologically similar to Vobzia heteropbylla Brongniart (KÚHOO5 8). Fig. l) - ? Isoetites Múnster, nearly complete individual with roots (below), ,horr st"- and long nar- rorvleaves(KÚHOO35A). Fig.3)-Cremat-opterisSchimperEe Mougeot,fragmentof asimplypinnatefertilefern-leaf (KúH0642).Fig.+)-ln.r- roPteridium Schimper, leaf-fragment (KÙH0665). Fig. 5) - Anomopterís Brongniart, bipinnate leaf-fragment wìth small oblate pinnules (KUH0089). Fig. 6) - Specimen of a large Cycad leaf similar to Bjutia dolomitica'illachtler & van Konijnenburg - van Cirtert 1KùHO55I). Fig. /) - An almost complete female Cvcad fructification with macrosporophvlls (KÙH0658) of the Dioonitocarpidium Lllienstern - type. 388 C.BroglioLoriga,A.Fugagnoli, J.VanKoníjnenburg-VanCittèrt,E.Kustatscher,R.Posenato&M.Wachtler

1855; De Ztgno 1,862; Schenk 1868; GùmbeI 1879). sion of the manuscript. Thanks go to P Gianolla and R. Rettori for Only when the systematic study will be completed, the their suggestions on the stratigraphy and foraminiferal analysis, and to Nebelsick for the English revision. comparison with the other classical European Anisian James floras and Ladinian floras of the Dolomites is possible, This work represents a preliminary contribution of the project "Anisico- Kúhwiesenkopf (Prà della Vacca): paleoecologia e paleocli- also with paleoclimatic aims. matologia" supported by the Museo di Scienze Naturali of Alto Adige Acknouledgemenrs. The authors are grateful to H. Kerp. M. (Bolzano) under the leadership of Pro{. C. Broglio Loriga from the Gaetani and A. Tintori for their constructive reviews of an earlier ver- Ferrara lJniversity.

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