Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy

Cretaceous Research (2002) 23, 671–685doi:10.1006/cres.2002.1026

Late Cretaceous plants from the Bonarelli Levelof the Venetian Alps, northeastern Italy

*Bernard Gomez, †Frederic Thevenard, ‡Marco Fantin, and ‡Luca Giusberti

*School of Earth Sciences, University of Leeds, Leeds LS2 9JT, UK; e-mail: [email protected]†Paleobotanique, UMR 5125 du CNRS, Universite Claude Bernard Lyon 1, batiment Charles Darwin A,7, rue Dubois, 69622 Villeurbanne cedex, France; e-mail: [email protected]‡Dipartimento di Geologia, Paleontologia e Geologia, Via Giotto 1, I-35137 Padova, Italy;e-mail: [email protected]

Revised manuscript accepted 6 June 2002

Fossil plant macroremains were collected from the black shale of the outcrops of the Bonarelli Level near the village of Quero,Venetian Alps, NE Italy. This important radiolarian-ichthyolithic, organic-rich marker bed was deposited close to theCenomanian/Turonian boundary and represents the sedimentary expression of the global Oceanic Anoxic Event 2 (OAE2).Taphonomic evidence suggests that terrestrial areas were exposed near Quero during the deposition of the black shaleassemblage. The floristic assemblage consists of vegetative and reproductive remains belonging to ferns, conifers andangiosperms. Preservation of the material is poor and, therefore, only impressions and compressions have been preserved. Inmost cases, only gross morphological features were studied. However, cuticular analyses with light and scanning electronmicroscopy have aided in the description of the new species Frenelopsis veneta. This taxon shares with F. harrisii Doludenko& Reymanowna and F. kaneviensis Barale & Doludenko the narrow, inter-cell grooves on the outer internodal epidermalsurface, but it differs in cuticle thickness, stomatal diameter and stomatal density. An attempt is made to establishrelationships between extant families and genera, and the fossil specimens. � 2003 Published by Elsevier Science Ltd.

K W: fossil plants; taxonomy; Late Cretaceous; Bonarelli Level; OAE2.

1. Introduction

In order to analyze, describe and interpret the bioticand abiotic changes across and within the BonarelliLevel (BL) in Italy, an international multidisciplinaryproject was promoted and organized by R. Coccioniof Urbino University (Italy). Within the frameworkof the ‘Bonarelli Level of Italy Project’ (BLIP) wecarried out a study of the plant macroremains ofthis important marker bed. During the last tenyears studies of the OAE2 in Italy have gainedmomentum and essentially focused on the geochemi-cal, sedimentological and micropalaeontologicaldisciplines (e.g., Bellanca et al., 1996; Salvini &Marcucci Passerini, 1998; Luciani & Cobianchi,1999; Premoli Silva et al., 1999), while the macro-palaeontological aspects were neglected. In fact, onlythe rich fish fauna of the BL have been studied indetail (Sorbini, 1976, 1980; Astolfi & Colombara,1990; Bizzarini & Coccioni, 1990; Avanzini &Luciani, 2002).

0195–6671/03/$30.00/0

2. Geological setting

The Jurassic and Cretaceous sediments of the ItalianSouthern Alps are generally considered to have beendeposited on the northern passive continental marginof the Mesozoic Tethys, characterized by horst andgraben structures inherited from the rifting associatedwith the opening of the central North Atlantic(Winterer & Bosellini, 1981). The palaeogeographicreconstruction of this part of the continental marginshows an external carbonate platform, the FriuliPlatform, and two continental margin basins, theBelluno and the Lombardian basins, separated by anintervening chain of seamounts known as the TrentoPlateau. At the end of the Jurassic and during theEarly Cretaceous the platform-basin morphology be-came subdued and calcareous pelagic oozes, mostlyconsisting of nannofossils, blanketed much of theLombardian and Belluno basins as well as the TrentoPlateau (Channell et al., 1992). At the same time, reeflimestones characterized the margin of the Friuli

� 2003 Published by Elsevier Science Ltd.

672 B. Gomez et al.

Figure 1. Map showing the location of the exposures studied (A–C), and stratigraphic columns of sections B and C.The Bonarelli Level is folded and not well exposed at site A; for this reason only exposures B and C were sampled indetail.

Platform and limestones composed of re-sedimentedshallow-water debris with pelagic material weredeposited in the eastern part of the Belluno Basin(Winterer & Bosellini, 1981). At the beginning ofthe Aptian these limestones were succeeded every-where by repeated alternations of limestones, vari-coloured marlstones and organic, carbon-rich shaleswidely known in literature as the Scaglia VariegataFormation.

The plant material was discovered near the villageof Quero (Belluno, NE Italy) by two of us (MF andLG in 1993) within the black shale of the BonarelliLevel (close to the Cenomanian/Turonian boundary),which represents the top of the Scaglia VariegataFormation and grades upward into the typicalred–pinkish cherty limestones of the Scaglia RossaFormation (Upper Cretaceous–Lower Eocene). Theexposures studied (Figure 1) are located along thevalley of the Tegorzo Creek, which crosses thesouthern flank of the Monte Grappa Anticline. Here a

Jurassic–Eocene pelagic succession is exposed andrepresents sedimentation in a deep-sea area tran-sitional between the Belluno Basin and the TrentoPlateau (Bizzarini et al., 1987).

The BL consists of yellowish-brown laminatedmudstones, alternating with olive-grey, silty shalesand olive-green, radiolarian-rich, silty–sandy layers.Limonite nodules, representing the alteration oforiginal Fe-sulphide nodules, are also present. Thefossil remains are mainly concentrated in the mud-stones of the lower–middle part of the level, and weremostly collected from exposure B. They consist ofplant macroremains and fish remains such as Pachy-rhizodus sp., Tselfatia formosa, Coelodus sp. and Pty-chodus decurrens teeth. Radiolaria, sponge spicules andsmall ichthyoliths are usually the main component ofthe microfossil assemblages of the BL, whereas plank-tonic foraminifera are generally absent. Calcareousnannoplankton occur exclusively in the lowermostpart of the BL and are moderately well preserved. The

Late Cretaceous plants from the Bonarelli Level 673

3. Material and methods

Fragments of Frenelopsis cuticle were separatedfrom the sediment surface with a scalpel. Theywere cleaned in 37% hydrochloric acid (HCl) andsubsequently macerated in Schulze’s reagent, theduration of the latter depending on the degreeof carbonization and preservation (Kerp, 1990).Adhering siliceous debris on the cuticle surfaces wasremoved by immersion in 40% hydrofluoric acid(HF).

No cuticle was found on the scale-like leaves;hence, only cuticles of the internodes were studied.The internodes were separated into two parts with twoneedles and mounted in glycerine jelly on glass slidesfor light microscope examination. A few fragmentswere stained with safranine before mounting; otherswere prepared for study under a scanning electronmicroscope (SEM).

Light microscope observations were made using aLeitz Aristoplan microscope. Photomicrographs weretaken with a Wild Photoautomat MPS 45–51S in theEquipe de Paleobotanique de l’Universite ClaudeBernard Lyon-1 (LPUL), Villeurbanne (Rhone,France). The cuticles were examined using a HitachiS800 SEM at the Centre of Microscopy Applied toBiology and Geology (CMABG) of the UniversityClaude Bernard Lyon-1. All the material studied ishoused in the Museo di Geologia e Paleontologia of

Padua University (Italy), collection numbers MGPD28740–28754, MGPD 28765–28780 and MGPD28781–28787.

4. Italian Cretaceous plant records

In Italy, Cretaceous beds bearing plant macroremainshave been known of since the 19th Century. However,few palaeobotanical studies have been carried out andthese have been scattered over a wide area (Table 1,Figure 2).

Figure 2. Map of Italy with locations of Cretaceous plantfossil outcrops indicated in Table 1.

5. Palaeobotany

Ferns. This group is represented by the part andcounterpart of a single, incomplete, sterile pinna(Figure 3a, b), 17 mm long and 9 mm wide. Pinnulesare oblong, c. 4.0 mm long and 1.5 mm wide, and areinserted in a typical pinnate arrangement on therachis. They show full basal attachment and are veryslightly decurrent. The margin is entire and the apexobtuse. The strong midrib seems branched in ananastomosed net of secondary veins.

According to its gross morphology the materialresembles a secondary pinna of the fossil matoni-aceous fern Weichselia reticulata (Stokes & Webb)

assemblage is composed of Corollolithion kennedyi,Helenea chiastia, Lithraphidites acutus and Rhago-discus asper, suggesting a late Cenomanian age(determinations by Eliana Fornaciari).

This radiolarian-ichthyolithic, organic-rich markerbed, ranging from 30 to 300 cm thick in the SouthernAlps (Kuhnt et al., 1990; Salvini & MarcucciPasserini, 1998), has been recognized in differentsettings throughout Italy, from the Dolomites toSicily, and in other parts of the world. It correspondsto the global OAE 2 (Oceanic Anoxic Event 2)(Schlanger & Jenkins, 1976), which is marked by amajor turnover in several fossil groups and a majorpositive shift in �13C of both bulk carbonate andorganic carbon. The biotic changes observed acrossthe BL point to an increase in primary productivityinduced by an exceptional increase in nutrients insurface and subsurface waters lasting for hundreds ofthousands of years (Arthur & Premoli Silva, 1982;Coccioni et al., 1995; Premoli Silva et al., 1999). TheOAE2 probably took place in a global transgressivecontext and was related to an exceptionally intenseupwelling (Jenkyns, 1980; Coccioni et al., 1991; Galeet al., 2000).

Table 1. Summary of previous Italian Cretaceous palaeobotanical studies; see Figure 1 for geographical location; *age established from the presence of Miculaspp. and the absence of typical Campanian taxa (determined by Eliana Fornaciari).

References

Dalla Vecchia (2000a)

Dalla Vecchia (2000a)

ia, cf.s

Dalla Vecchia & Rigo (1998), Rigo(1999), Dalla Vecchia (2000a)

gracile,S.

yllitesua)

Bozzi (1888, 1891), Tommasi (1889,1891), Muscio & Venturini (1990),Dalla Vecchia (2000a)

Meleleo et al. (1984), Dalla Vecchia(2000a)

mensSorbini (1974), Astolfi & Colombara(1990)

nelopsissperms

Bizzarini & Coccioni (1990), Pigozzo(1996, 2002)

Bizzarini & Coccioni (1990), DallaVecchia (2000a)Pigozzo (2001)

Pigozzo (2001), hereinood Lehner et al. (1987)

Taramelli (1873), De Zigno (fideTaramelli, 1881)

zamites?sp.)

Bravi (1995), Bravi & Garassino(1998a), Dalla Vecchia (2000a)

and Bravi & Garassino (1998b), DallaVecchia (2000a)Venturini (1995), Dalla Vecchia(2000a)

and Bravi (1997), Dalla Vecchia (2000a)

scale-like Muscio & Venturini (1990), DallaVecchia (2000a)

674B

.G

omez

etal.

Age Formation/Locality Plant fossils

Late Maastrichtian (16) Trieste Karst, at Trebiciano (Trieste,Friuli Venezia-Giulia region)

Conifer remains

Late Santonian (15) Aurisina Limestones, at Villaggio delPescatore (Trieste, Friuli Venezia-Giuliaregion)

Undetermined fragments and rarecharophytes

Coniacian–Santonian (14) Aurisina Limestones, at Polazzo (CarsoIsontino, Friuli Venezia-Giulia region)

Conifers (Cunninghamites, cf. SequoAraucaria) and possible dicotyledon

Coniacian–Santonian* (13) Limestones, at Vernasso (Udine, FriuliVenezia-Giulia region)

Conifers (Araucaria macrophylla,Cunninghamites elegans, CyparissidiumFrenelopsis konigii, Sequoia ambigua,concinna) and angiosperms (Arundogroenlandica, Myrica vernassiensis, Phproteaceus, P. platanoides, Rhus antig

Cenomanian–Senonian (12) Limestones of Melissano, at Surbo(Lecce, Apulia region)

Cunninghamites? (Araucariaceae)

Cenomanian/Turonian (11) BL, at Cinto Euganeo (Euganei Hills,Padua, Veneto region)

Very abundant fragmentary plantmacroremains including some specidoubtfully ascribed to Equisetites

Cenomanian/Turonian (10) BL, at Carcoselle, near Possagno(Treviso, Veneto region)

Sequoia concinna (Taxodiaceae), Frealata (Cheirolepidiaceae) and angio(Sapindopsis alpina and Carcophyllumleogianense)

Cenomanian/Turonian (9) BL, at Furlo (Marche region) Undetermined fossil plants

Cenomanian/Turonian (8) BL, at Ca’ Trenta di Schio (Vicenza,Veneto region)

Frenelopsis

Cenomanian/Turonian (7) BL, at Quero (Belluno, Veneto region) Sequoia, hereinLate Cenomanian (6) Scaglia Bianca Fm, at Costa Valley

(Brescia, Lombardy region)Silicified fragment of gymnosperm w

Albian–Cenomanian? (5) Bituminous limestones exposed atFaierazzo (near Polcenigo, FriuliVenezia-Giulia region)

Ferns and monocotyledons

Late Albian (4) at Petina (Salerno, Campania region) Conifers (Pagiophyllum sp. and Podosp.) and other plants (e.g., Zamites

Early Albian (3) Limestones with ichthyoliths, at Pietraroja(Benevento, Campania region)

Cycadales?, Bennettitales (Zamites?)conifers (Brachyphyllum)

Middle Aptian (1) Marls, in the Cornappo Valley, nearTorlano (Udine, Friuli Venezia-Giuliaregion)

Charophytes

Early Aptian (2) Marls, at Profeti (Caserta, Campaniaregion)

Conifers such as Brachyphyllum sp.Podozamites sp.

Late Barremian (1) In the Cornappo Valley, near Torlano(Udine, Friuli Venezia-Giulia region)

Unidentified conifer shoots bearingleaves


Fontaine emend. Harris (Reymanowna, 1965; Alvin,1971; Harris, 1981). Weichselia reticulata was a wide-spread component of tropical–subtropical florasduring the Early Cretaceous (Retallack & Dilcher,1986; Saward, 1992; Watson & Alvin, 1996).

Conifers

Vegetative remains. Based on external morphology,three types of vegetative leafy shoots of conifers havebeen identified:

1. The most abundant form consists of leafy shootsup to 290 mm long (MGPD 28741–28754; Figure 3c,d). The main axis of the largest specimen (9 mmwide) ends with a sheaf of more than 30 branches(each from 3 to 4 mm wide). Branches bear small(2.50 mm long, 0.75 mm wide), scale-like, spirallyarranged leaves (5+8 parastichies). Each leaf is in-flexed and has entire margins. The basal part isadpressed and decurrent upon the axis. The distalportion is free, short, and may be slightly archedinwards. The apex is mostly acute, and a keel occurson the abaxial side of the leaf. The basal cushion isrhombic in shape.

It is difficult to identify specimens that are devoid ofcuticle. However, the gross morphology of the leafyshoots is similar to that of several conifers usuallyencountered in the Cretaceous and Cenozoic such asSequoia fastigiata (Sternberg) Velenovsky (non Heer),Sequoia concinna Heer (cf. Bozzi, 1891), SequoitesBrongniart and Quasisequoia Srinivasan & Friis.Among these taxa, the specimens of Quero appearto be particularly close to Sequoia fastigiata (cf.Velenovsky, 1885, pl. 8, fig. 13; pl. 9, figs 3, 4, 9–11;pl. 11, figs 1,2; pl. 12, fig. 13). They greatly resemblethe specimen depicted by Bozzi (1891) in his plate 15,figure 2, and those drawn in figures 1 and 3 of thesame plate, but the specimen in plate 6, figure 3 in anearlier paper of his (Bozzi 1888), also attributed toSequoia concinna, appears to be morphologically dis-tinct. The extant genus Sequoia Endlicher is definedby both vegetative and reproductive organs. Given thefact that the leaves of the single modern species, S.sempervirens (D. Don) Endlicher, are never spirallyarranged along the shoots, the present fossil materialshould be distinguished from Sequoia. The inclusionof these Italian specimens in the fossil genus GeinitziaEndlicher (cf. Meyen, 1987) may be more appro-priate. Comparisons with modern conifers suggestmorphological resemblances with extant taxo-diaceous species such as Athrotaxis cupressoides D. Don(Gaussen, 1967, p. 49, fig. 452; Page, 1990a, pp. 356,357, fig. 193) and Taiwania flousiana Gaussen (Page,1990a, p. 357, fig. 195).

2. Unbranched conifer shoots (MGPD 28765–28776; Figure 3e) bear spirally arranged, inflexed,lanceolate leaves 5 mm long, 1 mm wide with obtuseapices. These fossils may be assigned to Geinitziareichenbachii (Geinitz) Hollick & Jeffrey. Although thistaxon is a typical component of mid–Late CretaceousEuropean and North American assemblages, no speci-mens in the present collections yielded cuticle. Thespecies is known from Bohemia, Saxony, the Atlanticside of the North America, southern Greenland andFrance.

Foliage similarities exist with extant Cupressaceaesuch as Fitzroya cupressoides (Mol.) I. M. Johnston(Phillips, 1981, p. 118; Page, 1990b, p. 313, fig. 162)and with modern Araucariaceae such as Araucariaexcelsa (Lamb.) R. Brown [=A. heterophylla (Salisb.)Franco], A. cunninghamii D. Don, and A. balansaeBrongniart & Gris (Gaussen, 1970). Moreover, mor-phological affinities are especially numerous withextant taxa such as Athrotaxis selaginoides D. Don(Phillips, 1981, p. 87), Cryptomeria japonica (L.f.) D.Don (Phillips, 1981, p. 109; Page, 1990a, p. 357, fig.196) and Taiwania cryptomerioides Hayata (Phillips,1981, p. 203; Page, 1990a, p. 357, fig. 195D),classically assigned to the Taxodiaceae.3. Cuticular study of the third coniferous form led tothe identification of a new species of Frenelopsis.

Order: ConiferalesFamily: Cheirolepidiaceae Takhtajan, 1963

Genus Frenelopsis (Schenk, 1869) emend. Watson,1977

Type species. Frenelopsis hoheneggeri (Ettingshausen,1852) Schenk, 1869, p. 13, pl. 4, figs 5–7; pl. 5, figs1–2; pl. 6, figs 1–6; pl. 7, fig. 1.

Frenelopsis veneta sp. nov.Figures 4a–g, 5a–j, 6

Holotype. MGPD 28778, from the collection of theMGPD, Padua University (Italy).

Paratypes. MGPD 28779, slide number MGPD28783 and stub number MGPD 28784, from thecollection of the MGPD, Padua University (Italy).

Type locality. Locality B, the valley of Tegorzo Creek,near Quero, Belluno, NE Italy (Figure 1).

Stratigraphic horizon. Bonarelli Level, top of ScagliaVariegata Formation; age close to the Cenomanian/Turonian boundary.

676 B. Gomez et al.


Derivation of name. From venetus-a-um, referring tothe Veneto region (NE Italy) where the species wasfound.

Other material. MGPD 28780–28782.

Diagnosis. Frenelopsis shoots attaining at least threeorders of branching. Branches arising distichously atangles between 20 and 30� and the axis up to 3.5 mmwide. Three scale-like leaves at each node. Stomataldensity of 70–90 per mm2. Stomatal complex 60–90 �m in diameter, formed by 4–5 subsidiary cells,each with one outer and one inner papilla. Rosette ofouter papillae sunken below the cuticle surface andflanked by massive papillae of the encircling cells.Narrow grooves present between the epidermal cellson the outer cuticular surface.

Description. Branching occurs at irregular intervalsarising at 20–30�. The most complete shoot (Figure4a) measures about 170 mm long and 3.5 mm wide.Branches are produced alternately, and the specimentends to have one main axis. Axes are clearly seen tobe segmented, each node bearing a whorl of threescale-like leaves. The arrangement of leaves alternatesfrom one node to the next (Figure 4b). The cuticle ofthe internode is thick. The stomata are arranged inill-defined rows (Figures 4c, 5a). One to four ordinaryepidermal cells separate two adjacent stomatal rowsand 1–5 epidermal cells intercalate between two con-secutive stomata of the same row. Two stomata canoccasionally be in contact but they never share theirsubsidiary cells (Figure 5f). The stomatal densityvaries between 70 and 90 per mm2 while the stomatalindex is about 12.5. The stomatal complex is 60–90 �m in diameter and the pores 15–35 �m in maxi-mum diameter (SEM internal view). The haplocheilicstomata are monocyclic or incompletely dicyclic.They consist of 4–5 (Figures 4d–g, 5e, f, h, i), butusually four, subsidiary cells (20–40 �m long and10–20 �m wide). Each subsidiary cell bears one outerand one inner papilla (Figures 4e–g, 5e–j). Innerpapillae measure 10–15 �m long. Outer papillae arefused proximally to form a protruding rosette leavingan aperture of about 5–20 �m. The rosette of outerpapillae (4–12 �m thick) is sunken below the cuticlesurface and also flanked by massive papillae of the

encircling cells which delimit an area 25–45 �m indiameter (Figures 5g, j, 6). Narrow grooves occurbetween the epidermal cells on the outer cuticularsurface (Figures 5b, d, g, j, 6). Guard cells are notpreserved. Epidermal cells are square to polygonal(Figure 4c–d) and range between 15–45 �m long and15–35 �m wide (average size 25 �m). They have anti-clinal walls between 5 and 12 �m thick. A hypodermisis partly preserved but it appears to be highly degraded(Figure 5a, c).

Remarks. Among the species attributed to Frenelopsis(Gomez et al., 2002), the cuticles of the specimensfrom Quero are closest to F. kaneviensis Barale &Doludenko (1985) from the Albian of the Ukraineand F. harrisii Doludenko & Reymanowna (1978)from the Cenomanian of Tadzhikistan, with whichthey share the narrow grooves between the epidermalcells of the outer cuticular surface (Table 2). Thisfeature is, however, inconsistent in F. kaneviensis andhas not been considered as a specific character byBarale & Doludenko (1985). Moreover, the circular,non-papillate external outlines of stomata, and theunique papilla borne by each subsidiary cell inthe throat of the stomatal pit in F. kaneviensis,constitute additional differences in comparison tothe Italian specimens. F. kaneviensis also displays alower stomatal density (30 vs. 70–90 st./mm2) andsmaller stomatal diameter (35–50 vs. 60–90 �m). Thestomata can be composed of six subsidiary cellsin contrast to our material which possesses a maxi-mum of five subsidiary cells. In both F. harrisiiand F. veneta, each subsidiary cell bears a large,tongue-shaped papilla in the throat of thestomatal pit and usually one smaller papilla at the pitentrance. By way of contrast to F. harrisii, the surfaceof the periclinal walls in F. veneta never bulgeor extend into a conical hair (Doludenko &Reymanowna, 1978). Just as for F. harrisii, the pro-truding upper parts of subsidiary cells are similar tothose of ordinary epidermal cells but the dome overthe stomatal pit in F. veneta is formed by muchsmaller outer papillae and the lobes are fused togetherat the base in a coronal stomatal rim. Moreover,the cuticles of F. veneta are thinner than those ofF. harrisii (20–30 vs. up to 100 �m) and the stomatahave lower diameters (45�20 �m in F. harrisii).

Figure 3. Ferns and conifers from the Cenomanian–Turonian succession of Quero. a, b, part and counterpart of Weichseliareticulata; pinnules arranged opposite each other on the rachis and with an apparent midrib, MGPD 28740a–b; �3.c, Geinitzia sp.; main axis bearing a sheaf of more than 30 branches, MGPD 28741; �0.7. d, Geinitzia sp.; spiralarrangement of the appressed leaves on the twigs, MGPD 28742; �1.2. e, Geinitzia reichenbachii; spiral arrangement ofthe falcate leaves on a twig, MGPD 28765; �1.6. f, poorly preserved sporangiate cone, MGPD 28777; �3.3.

678 B. Gomez et al.


Reproductive remains of conifers

One small (8.5 mm long�6.0 mm wide), spherical,poorly preserved sporangiate cone (Figure 3f) displaysan uncertain number of spirally arranged sporophylls.The 9.5-mm-long cone is positioned terminally on ashort leafy axis.

Although the poor preservation makes it difficult toestablish the morphology of the sporophylls, somefeatures are comparable to fossil taxodiaceous generafrom the Jurassic and Cretaceous of Europe. Forexample, there are some similarities with cones ofQuasisequoia Srinivasan & Friis and unassigned femalecones from the Santonian/Campanian of Sweden.

6. Discussion

It has been difficult to establish the affinities of theplant organs recovered from the Bonarelli Level atQuero, but some may be related to Matoniaceae,Taxodiaceae and Cheirolepidiaceae. Owing to thepoor preservation of the material, only Frenelopsisveneta sp. nov. can be considered satisfactorily de-scribed. The most numerous specimens are conifers,whereas ferns and angiosperms are sparse. In otherCenomanian–Turonian palaeofloras, angiosperms aremore numerous and diverse, commonly with largerleaf sizes (Hickey & Doyle, 1977; Retallack & Dilcher,1981). Both Weichselia and Frenelopsis, which are verycommon elsewhere in the Lower Cretaceous (Saward,1992), are sparse in the beds examined for this paper.

Taphonomic analyses of the beds bearing the fossilplants allow us to speculate on the necrobiosis(i.e., production), biostratinomic (i.e., transport) andfossil-diagenetic processes and, consequently, on thepalaeoenvironment of Quero deposits. From thenecrobiosis point of view, most of the conifer shootsfound in the BL are probably not the result ofcatastrophic events such as storms, which can removelarge plant fragments but are probably the result ofphysiological processes. Shoots such as those collectedare naturally shed, for example, in Cryptomeria

Flowering plants

Angiosperms are represented in the assemblage, asreflected by the presence of a few isolated leavesand winged seeds. Most leaves are fragmentary andvenation patterns are difficult to distinguish. Thepoor preservation of these features prevents theiridentification or comparison to fossil extant taxa.

The best-preserved specimen is represented by partand counter-part of an oval leaf or leaflet (Figure 7a),40 mm long, 11 mm wide, with an entire margin and7-mm-long petiole. The venation is ill-defined exceptfor a relatively prominent midrib, which narrows inthe distal part. The coriaceous aspect explains whythe lateral veins are indistinct.

These features can be found in many angiospermleaves. Among the Early Cretaceous angiosperms,Acaciophyllites Berry (Fabaceae), Sapindopsis Fontaine(Sapindaceae) and Dicotilophyllum spatulatium Pons,for which Pons (1988) evoked affinities with severalfamilies such as Apocynaceae, Lauraceae and Myrsi-naceae, broadly resemble the specimen studied. Thereare also strong resemblances with Phyllites proteaceusBozzi (1891, pl. 16, figs 6, 7), especially in the generalleaf shape, the short petiole, the strong midrib and thecoriaceous lamina, which hides the secondary netvenation.

Another dicotyledon is represented by the proximalportion of a leaf (Figure 7b). It is 25 mm long and16 mm wide, with a petiole 5 mm long and 1 mm

wide, and a prominent midrib. The lamina has acuneate, asymmetrical base. The margin is entire andslightly waved. The secondary venation is not pre-served. The centre of the leaf bears an irregular holethat may represent an example of insect herbivory.The specimen is too poorly preserved to assign it to anexisting taxon.

Several tiny seeds (Figure 7c, d) are formed fromtwo laminae arising from a common coriaceouspointed area. Each lamina is oval to lanceolate, andtheir size is 3.5 mm long and 2.5 mm wide. One sideof the lamina margin seems thicker and the laminaeshow several short, coriaceous strings converging tothe proximal area. These specimens may be inter-preted as winged seeds but their identification has notbeen possible.

Figure 4. Frenelopsis veneta sp. nov.; conifer from the Cenomanian–Turonian succession of Quero. a, paratype, shootbranched several times with branches arising at 20–30�, MGPD 28779; �0.5. b, holotype, detail of the branching andinternodes (arrows indicate some of the whorls of three leaves), MGPD 28778; �1.7. c, stomata arranged in ill-definedrows, MGPD 28783; �120. d, isodiametric, square to polygonal epidermal cells and surface ornamentation discernible,MGPD 28783; �250. e, two stomata side by side not sharing their subsidiary cells, and with four and five subsidiarycells respectively, each bearing one inner and outer papilla, MGPD 28783; �450. f, stomata with four subsidiary cellseach bearing one inner and outer papilla, MGPD 28783; �450. g, stomata with five subsidiary cells each bearing oneinner and outer papilla, MGPD 28783; �500.

680 B. Gomez et al.


Figure 6. Reconstruction of Frenelopsis veneta sp. nov.showing cross-section of the cuticle at the level of astoma.

japonica (L.f.) D. Don during periods of drought. Inthe same way, angiosperm leaves and winged seedswere probably deciduous, with an abscission arealocated at the most proximal part of the petiole for theformer. All these plant remains might have resided forsome time in the dried litter without disarticulation. Ithas been pointed out in the literature that damage ofplant macroremains is greatly related to the distance,duration and energy of the transport (e.g. Spicer,1981; Spicer & Greer, 1986; Ferguson, 1996).Frenelopsis probably could not have endured long orhigh energy transport without the shoots becomingfragmented (Gomez et al., 2001, 2002). Owing to theminor damage observed in the material, most of theplant macroremains were probably derived from anearby terrestrial source area. Exposed portions of theFriuli Platform are the likely source areas of the plantdebris. Additional evidence for exposure of portions ofthe Friuli and contiguous Adriatic Platform duringLate Cretaceous occur in the form of palaeokarsticdeposits, palaeosol horizons and dinosaur track sites(Sartorio, 1989; Swinburne & Noacco, 1993; Sartorioet al., 1997; Avanzini et al., 2000; Dalla Vecchia,2000b; Melis et al., 2000; Tunis & Venturini, 2000;Dalla Vecchia et al., 2001). Dalla Vecchia (2000a) hasalso pointed out that the relative abundance of terres-trial plants in basinal sediments around the Periadri-atic carbonate platforms of Italy, could be consideredas further evidence that they were periodically exposedand colonized by vegetation. The distance betweenthe Quero area and the Friuli Platform margin, as

deduced from palaeogeographic maps (e.g., Cati et al.,1989), was about 20–25 km whereas the taphonomicevidence of BL plants seems to favour a shorterdistance (not more than 1 or 2 km) between the plantsites and the emergent platform margin. Unfortu-nately, few details are available on the proximity of thesouthwestern part of the Friuli Platform, partly buriedunder the Cenozoic sediments of the Venetian Alps(Cati et al., 1989). We propose a reconstruction of thearea (Figure 8), but further investigation is needed toverify the palaeogeography.

Palaeomagnetic studies indicate that the ‘Apulianblock’ (or ‘Adriatic promontory’) was located between20 and 30�N during the Cenomanian–Turonian inter-val. Therefore, the Southern Alps were probably in asemi-arid, warm, subtropical zone during the depo-sition of most of the Cretaceous pelagic sequences(Arthur & Premoli Silva, 1982). Data from the floris-tic assemblage at Quero are consistent with thesepalaeoclimatic conditions. Both of the modern famil-ies Matoniaceae and Taxodiaceae are mainly confinedtoday to warm areas of the planet. Additionally, thefrenelopsids are considered to have been predomi-nantly xeromorphic plants probably adapted to aridity(e.g., Gomez et al., 2001, 2002), based on suchfeatures as reduced leaves, heavily protected stomataand thick cuticles, as evidenced in Frenelopsis venetasp. nov. Finally, as suggested by Watson & Fisher(1984) for specimens of the genus Glenrosa from theLower Cretaceous Glen Rose Formation, Texas,the narrow grooves between the epidermal cells ofF. veneta sp. nov. might have contained a spread ofsurface moisture droplets, thus reducing the meansurface area available for evaporation/transpiration.

Figure 5. SEM micrographs of Frenelopsis veneta sp. nov. from the Cenomanian–Turonian succession of Quero. a, internalview of cuticle showing rows of stomata and anticlinal walls of epidermal cells, MGPD 28784; �90. b, external view ofcuticle showing narrow grooves between the epidermal cells, MGPD 28784; �90. c, internal view of cuticle showingstomata and anticlinal walls of epidermal cells, MGPD 28784; �170. d, external view of cuticle showing narrow groovesbetween the epidermal cells, MGPD 28784; �180. e, internal view of cuticle with four stomata, MGPD 28784; �206.f, internal view of cuticle showing three stomata side by side, but not sharing their subsidiary cells, MGPD 28784;�400. g, external view of cuticle showing stomata with a rosette of outer papillae and narrow grooves on outer cuticularsurface, MGPD 28784; �270. h, internal view of cuticle showing one stoma with four subsidiary cells and papillae,MGPD 28784; �530. i, internal view of cuticle showing a stoma with five subsidiary cells and papillae, MGPD 28784;�530. j, external view of cuticle showing a stoma with four outer papillae and the narrow grooves of epidermal cells,MGPD 28784; �530.

Acknowledgements

BG thanks his wife for her patience, assistance andadvice during the writing of this paper. BG and FT aregrateful to UMR 5125 and the programme ECLIPSEof the French CNRS, and for a European CommunityMarie Curie Fellowship (HPMF-CT-2002-01584)

Table s of genus Frenelopsis (modified from Gomez et al., 2002).

Charac

. harrisiiludenko &ymanowna

F. kaneviensisBarale & Doludenko

F. venetaGomez et al. sp. nov.

Branch te Profuse ModerateInterno 8–10 ?Interno 1.7–3.5 up to 4Leaf nu ing whorls of 3 Alternating whorls of 3 Alternating whorls of 3Maxim .5 1–0.7 ?Depth Up to 1 ?Leaf up Short hairs up to 20

�m long?

Interno 00 �m 15–20 20–30Stomat fined rows Ill-defined rows Ill-defined rowsDensity 12–16 8–11Density 30 70–90Diamet (elliptical) 35–50 60–90Numbe 4–6, usually 4 4–5, usually 4Orienta ? VariableSurface Non-papillate PapillatePapillae Present PresentRim of Circular StellateNarrow t Variable ConstantDistribu istan Ukraine ItalyStratigr nian Albian Cenomanian–Turonian

682B

.G

omez

etal.

2. Comparison with the nearest species described to date and with the type specie

ters/species

F. hoheneggeriemend. Reymanowna

& Watson(type-species)

FDoRe

ing Moderate Moderade length (mm) 8 8–12de width (mm) 3 1–4mber per node Alternating whorls of 3 Alternat

um length of free leaf (mm) 1.5 Up to 1of sheathing base (mm) 1 ?per margin Scarious ?

de and abaxial leaf cuticle thickness (�m) 40 Up to 1al arrangement Well-defined rows Well-deof stomatal rows per mm 10–12 9–10of stomata per mm2 90–100

er of stomatal apparatus (�m) 60–70 45�20r of subsidiary cells 4–6 usually 4 4–5tion of stomatal aperture Transverse ?around pit Thickened ring Papillatein throat of stomatal pit Large hollow papillae Present

stomatal pit Stellate Stellategrooves on epidermal cells None Constantion Poland, Czech Republic Tadzhik

aphic range Hauterivian Cenoma


Figure 7. Unidentified angiosperm leaves and winged seeds from the Cenomanian–Turonian succession of Quero. a, MGPD28785a–b; �2.5. b, MGPD 28786; �3. c, d, MGPD 28787; �11.

Figure 8. Palaeogeographic reconstruction showing the two hypothesized sources of the plant remains that make up the Queroassemblage. The dotted area is hypothesized from the taphonomic examination (by BG and FT) of the plant macroremains,while the Friuli Platform source area is supported by much palaeontological and sedimentological evidence.

for the financial support of our study. MF and LG areindebted to P. Grandesso, R. Coccioni and F. M.Dalla Vecchia for their enthusiastic support and help-ful discussions. They are deeply grateful to R. Astorifor her essential help during field work. Thanks arealso due to E. Fornaciari for providing unpublisheddata on the calcareous nannoplankton of Vernassoand Quero. Stefano Castelli is acknowledged forphotographic work. MF and LG acknowledge the

financial support of the MURST (ex 60% grants toI. Dieni and R. Gatto). We all acknowledge Dr C.Blanc-Louvel for having emphasized in a preliminaryreport the interest of the Cenomanian/Turonianpalaeoflora of Quero. We are most obliged to AndreasSandersen who helped by providing numerous com-ments to improve the manuscript and corrected theEnglish version. Thanks are also due to the refereesand editor, D. J. Batten, for similar help.

684 B. Gomez et al.

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Late Cretaceous plants from the Bonarelli Level of the Venetian Alps, northeastern Italy