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Harris et al., eds., 2006, The Triassic-Jurassic Terrestrial Transition. New Mexico Museum of Natural History and Science Bulletin 37.
THE TETRAPOD FOSSIL RECORD FROM THE NORIAN-RHAETIAN OF FRIULI
(NORTHEASTERN ITALY)
FABIO M. DALLA VECCHIA
Museo Friulano di Storia Naturale, Via Marangoni 39-41, I-3310 Udine, Italy, E-mail: fabdalla@tin.it
Abstract—The latest Triassic tetrapods of northern Friuli (Carnian Pre-Alps, NE Italy) are represented by both
skeletal and ichnological remains. Pterosaurs, drepanosaurids, protorosaurians and a lepidosauromorph occur in
the Norian (middle Alaunian-Sevatian) Dolomia di Forni Formation. They include the holotypes of Preondactylus
buffarinii, Eudimorphodon rosenfeldi and Megalancosaurus preonensis, as well as the most complete skeleton of
the protorosaurian Langobardisaurus. Footprints of larger tetrapods, including dinosaurs, are known from the
inner carbonate platform facies of the Dolomia Principale Formation (late Carnian-Rhaetian), part of which is
laterally equivalent to the basinal Dolomia di Forni Formation. The tetrapods from the Dolomia di Forni are all
small terrestrial animals; no aquatic tetrapods are found. This is unlike the tetrapod fauna from the Norian basins
of Lombardy, where the overall tetrapod diversity also is higher, a difference possibly due to environmental
conditions. The larger animals of the platform (represented only by footprints) are not present in the basinal
sample. This could be due to selective transport, to a higher number of small tetrapods, to the small size of the
skeletal sample, or simply to a different age of the skeletal and ichnological records. While the Friulian and
Lombardian vertebrate samples do share some tetrapod and fish species, others are different. This is possibly due
to the slightly different ages of these deposits.
INTRODUCTION
The preserved remnants of relatively small, intraplatform marine
basins of the Norian crop out along the Pre-Alps of northern Italy. Dark
carbonate muds were deposited within these basins, and preserved within
the rocks derived from the sediments is a peculiar floral and faunal asso-
ciation. The basins were present in what are today the Lombardy and
Friuli regions. To date, tetrapod body fossils from these basinal units are
known from the Calcare di Zorzino (Zorzino Limestone) and Argillite di
Riva di Solto (Riva di Solto Shale) in Lombardy, and from the Dolomia di
Forni (Forni Dolostone) in Friuli.
The Lombardian faunas are better known, mostly because the
stratigraphically thin, vertebrate-rich intervals are easily accessible for
the purpose of field work. This has been undertaken by the University
of Milano and the Natural History Museum of Bergamo in some sites
(e.g., Cene and Zogno/Endenna in the Calcare di Zorzino, Berbenno/
Ponte Giurino in the Argillite di Riva di Solto).
The Dolomia di Forni of northern Friuli (Carnian Pre-Alps) is
several hundred meters thick, and lacks thin, vertebrate-rich intervals
easily accessible for the purpose of field work. Here tetrapods are rare
and only occasionally found by chance, often in the debris of small rock
falls, and sometimes reworked and transported for short distances by the
streams. Despite these factors, 17 tetrapod specimens have been col-
lected in the last 25 years, most of them of great scientific interest. They
include the oldest known pterosaurs (together with the roughly coeval
forms from Lombardy, Tyrol and eastern Greenland [Dalla Vecchia,
2003a]). The holotype of the bizarre Megalancosaurus, the only speci-
men preserving a complete skull, is part of the sample. Megalancosaurus
was once inferred to have avialan affinities (e.g., Feduccia and Wild,
1993) and has been the object of much debate (e.g., Padian and Chiappe,
1998; Ruben, 1998).The sample also includes the first distally complete
tail of Megalancosaurus, demonstrating the presence of a claw-like ter-
minal element like that seen in Drepanosaurus, as well as the best speci-
men of Langobardisaurus, the only one to preserve its highly special-
ized dentition.
As a consequence of the fact that the specimens were often de-
scribed in local journals of limited distribution, the Dolomia di Forni
tetrapods have frequently been confused with those from the Calcare di
Zorzino of Lombardy (e.g., Unwin, 2006), or ignored altogether.
The late Carnian-Rhaetian inner carbonate platform facies crop-
ping out just south-west of the basinal Dolomia di Forni preserves
ichnofossils of larger tetrapods, including dinosaurs (Dalla Vecchia and
Mietto, 1998; Dalla Vecchia, 2002a). The aim of this paper is to present
an overview of the latest Triassic tetrapod evidence from both the basi-
nal and the inner platform facies of the Carnian Pre-Alps.
Abbreviations: BSP, Bayerische Staatssammlung für Paläontologie
und historische Geologie, Munich, Germany; MFSN, Museo Friulano di
Storia Naturale, Udine, Italy; MPUM, Museo di Paleontologia,
Dipartimento di Scienze della Terra, University of Milano, Italy; S.N.,
Number of the Italian State inventory.
GEOLOGICAL AND STRATIGRAPHICAL FRAMEWORK
Most of the Late Triassic rocks of the Carnian Pre-Alps pertain
to the Dolomia Principale Formation (Main Dolostone; Hauptdolomit of
German authors) (Carulli et al., 2000) (Figs. 1-2). Within the western
Carnian Pre-Alps, this formation is informally subdivided into three
units: a lower peritidal unit, a middle subtidal unit and an upper peritidal
unit (Bosellini and Hardie, 1988). However, these units are not
chronostratigraphically equivalent in the eastern Carnian Pre-Alps, nor
can they be recognized everywhere (Carulli et al., 1998). The peritidal
units are composed of cycles comprised of a basal breccia with an ero-
sional base, middle subtidal muds sometimes including megalodontid
bivalves, and a stromatolite layer at the top. The subtidal unit is mainly
composed of thick beds of dolomitized muds with megalodontid bivalves
and algae, sometimes capped by a stromatolite layer, and with a thin
breccia layer at the base. The local thickness of the Dolomia Principale is
1500-2000 m (Carulli et al., 2000).
A lateral, basinward, equivalent of the upper part of the Dolomia
Principale is the Calcare del Dachstein (Dachstein Limestone Formation)
(Cozzi et al., 2000, p. 150, fig. 5), which appears to be the same as the
Dolomia Principale but not dolomitized. It is about 400 m thick at the
southern slope of Mt. Verzegnis (Carulli et al., 2000).
To the north (along the upper Tagliamento River Valley), the
basinal facies of the Carnian Basin (Dolomia di Forni and Calcare di
Chiampomano Formations) are laterally equivalent to the carbonate plat-
form facies (Figs. 1-2). Note that in this case “Carnian” does not have a
chronostratigraphic meaning, but refers, as does the Carnian Pre-Alps, to
that zone of northern Friuli, which is named Carnia.
433
The Dolomia di Forni is composed mainly of dark, often cherty,
well-bedded dolostone that is frequently thinly laminated and some-
times slumped (for more details, see Dalla Vecchia, 1991). Thicker lami-
nae are graded and represent distal turbidites. Closer to the carbonate
platform margin the unit comprises pebbly dolostones and breccia beds
(slope facies). Its thickness exceeds 800 m in the Seazza Creek section.
The Dolomia di Forni is overlain by the Calcare di Chiampomano
(Chiampomano Limestone Formation), a blackish, well-bedded lime-
stone, that is often slumped, locally cherty and has thin marly levels and
rare breccia beds in the basal part. Its thickness is 400-500 m along the
NW slope of Mt. Verzegnis. It is considered laterally equivalent to the
Calcare del Dachstein (Carulli et al., 1998; Cozzi et al., 2000, p. 150, fig.
5).
The lower boundary of the basinal sequence along the upper
Tagliamento River Valley is disturbed by an important thrust zone (the
Mt. Dof-Mt. Auda thrust). This marks the northern part of the Norian
outcrops where it is overthrust by older Triassic formations. Some au-
thors (e.g., Carulli et al., 2000) recognize the Tuvalian (Roghi and Dalla
Vecchia, 1997) Formazione del Monticello (Monticello Formation) be-
low the Dolomia di Forni. I think that this is not the case, at least in the
Seazza Creek and Forchiar Creek sections where most of the tetrapod
fossils come from. Because the tetrapods occur only in the Dolomia di
Forni, and the dating of the fossil-bearing sections (see below) is based
on microfossils found in it, this different interpretation is of no impor-
tance for this paper.
The age of the Dolomia Principale is poorly resolved
biostratigraphically. According to Carulli et al. (2000), it spans the Norian-
Rhaetian interval in the southern part of the Carnian Pre-Alps (e.g., at
Mt. Raut), although no biomarkers are found in its lower part. This is
also the case in its upper part, and in the lower part of the overlying
carbonate platform facies of the Calcari Grigi del Friuli (Grey Lime-
stones of Friuli Formation). Where the Dolomia Principale is overlain by
the Calcare del Dachstein (e.g., northern flank of Mt. Verzegnis), Carulli
et al. (2000) consider it as only Norian because they date the latter as
Rhaetian. Actually, this dating is based on the presence of the Sevatian-
Rhaetian foraminifer Triasina hantkeni (Salaj et al., 1988; Reijmer and
Everaars, 1991), indicating that the lower part of the Calcare del Dachstein
could be late Norian in age as suggested by Carulli et al. (1998, fig. 13)
and Cozzi et al. (2000, p. 150, fig. 5). There, the top of the Dolomia
Principale could be latest Alaunian in age. A thin intercalation of dark
dolomite in the Dolomia Principale in the Mt. Pramaggiore area (Mt.
Valmenone Anoxic Event of Cozzi et al., 2000, p. 150, fig. 5) appears to
have originated in a shallow, intraplatform, anoxic depression. It is dated
to the uppermost Alaunian 3 because of its abundance of the benthic
foraminifer Gandinella falsofriedli and “the presence of the Gandinella
falsofriedli interval-zone (Alaunian-Sevatian….) in the bituminous unit,
just before the Sevatian-Rhaetian Triasina hatkeni [sic] range-zone” (Cozzi
FIGURE 1. Stratigraphy and geochronology of the latest Triassic in the
northern Carnian Pre-Alps, upper Tagliamento River Valley, Friuli Venezia
Giulia, northeastern Italy (based on Dalla Vecchia, 1991; Roghi et al., 1995;
Cozzi et al., 2000), with the positions of some tetrapod specimens. The
basinal sequence (Dolomia di Forni and Calcare di Chiampomano) is based
on the depocentral Seazza Creek section. The stratigraphic occurrences of
MFSN 19235, MFSN 1921, MFSN 1769 and 332466 are tentative and
based on information given by the discoverers. 1, Dolomie cariate (vuggy
dolostone, breccia beds). 2, Basinal facies of the Dolomia di Forni (cherty,
well-bedded dolostone, often thinly laminated, sometimes slumped). 3, Slope
facies of the Dolomia di Forni (well-bedded dolostone often slumped, pebbly
dolostones and breccia beds). 4, Slope and basinal Calcare di Chiampomano
(well-bedded limestone, often slumped, locally cherty and with rare breccia
beds in the basal part). F1 and F4 refer to the fossiliferous outcrops reported
in Dalla Vecchia (1991) and Roghi et al. (1995). Dating of the stage/age
limits is based on Gradstein and Ogg (2004).
FIGURE 2. Top: Outcrop area of the Dolomia di Forni Formation (dark
grey) bearing the tetrapod body fossils. Numbered points mark the zones
where the specimens were collected. 1, Forchiar Creek; MFSN 1797 (holotype
of Eudimorphodon rosenfeldi was found in a section coincident with the
point), the small Langobardisaurus (MFSN 24992) and the probable
Eudimorphodon (MFSN 26823) were found in the stones of the creek bed.
2, Seazza Creek; MFSN 21546 (Eudimorphodon sp.) was found in the
stones of the creek bed in the lower part of the valley. 3, Seazza Creek;
MFSN 1769 (holotype of Megalancosaurus preonensis), MFSN 1801
(Megalancosaurus preonensis), MFSN 1921 (Langobardisaurus tonelloi),
MFSN 1891 and MFSN 19864 (Pterosauria indet.) were found in the middle
part of the valley. 4, Seazza Creek; MFSN 19235 (“Langobardisaurus?
rossii”), MFSN 18443 (Megalancosaurus preonensis), MFSN 1770
(holotype of Preondactylus buffarinii), MFSN 25161 (possibly
Preondactylus buffarinii) and S.N. 332466 were found in the middle-upper
part of the valley. 5, Purone Creek; MFSN 1922 (Eudimorphodon sp.) was
found in a boulder in the creek bed. 6, Rovadia Creek; MFSN 24992
(Langobardisaurus sp.) and MFSN 19836 (Pterosauria indet.) were found
in boulders in the creek bed. Bottom: Interpretative reconstruction of the
carbonate platform/basin section along the A-B transect in the zone of Mt.
Pramaggiore (based on Cozzi et al., 2000).
434
and Jäger, 2000, p. 165). The lower part of the Dolomia Principale is
Tuvalian (late Carnian) in the eastern Dolomites (De Zanche et al., 1993),
as is its basal 300 m in the Julian Alps (Gianolla et al., 2003). In sum-
mary, the maximum stratigraphic range of this unit could actually be late
Tuvalian-latest Rhaetian.
Once, the Dolomia di Forni was considered laterally equivalent to
the basal part of the Dolomia Principale and early Norian in age (e.g.,
Tintori et al., 1985). Subsequently, the dating of its lower-middle portion
in the sections at the Seazza Creek and Forchiar Creek was indicated as
Alaunian 2-3 (middle-upper middle Norian; Fig. 1) based on the presence
of the conodont Epigondolella slovakensis and its morphological trends
(Roghi et al., 1995). The same conodont species occurs in the western
sections of the Dolomia di Forni (Poschiadea Creek, Negro Creek, Caprizzi,
Libertan Creek, Mt. Rua), where some samples contain Epigondolella
postera associated with it (Carulli et al., 2000). A form transitional be-
tween Epigondolella postera and E. slovakensis is reported by Cozzi et
al. (2000, p. 162). E. slovakensis and E. postera are also reported in the
upper Rovadia Creek section (Carulli et al., 1998). E. postera occurs in
the middle Alaunian (Alaunian 2) E. postera Zone (Rigo et al., 2005).
Actually, the biostratigraphy of the Norian-Rhaetian is far from settled.
E. slovakensis has been shown to have a longer range, extending into the
Sevatian, and the in-progress revision of the conodont fauna from the
Dolomia di Forni has revealed an higher diversity, including the possible
presence of Epigondolella bidentata in the upper part of the formation
(Manuel Rigo and Guido Roghi, personal commun.). The E. bidentata
Zone represents the Sevatian (Rigo et al., 2005). Dalla Vecchia (1996,
2003a) hypothesized that the lithological transition from the Dolomia di
Forni to the Calcare di Chiampomano (as well as possibly that from the
Dolomia Principale to the Calcare del Dachstein on the platform) is
related to an event of dilution of marine waters and input of fine terrig-
enous sediment, possibly due to an humid climatic event (Stefani et al.,
1992; Jadoul et al., 1992, 1994; Cirilli, 1995; Carulli et al., 1998), “docu-
mented, at least in land areas by pollen and spores and the transition, all
over Europe, from gypsiferous and dolomitic horizons (Keuper) to lime-
stones and clays” (Jadoul et al., 1992, p. 39). Dalla Vecchia (1996, 2003a)
dated this event, under the assumption that it was synchronous, to the
Alaunian-Sevatian boundary, based on the dating of the uppermost Calcare
di Zorzino of Lombardy to the Alaunian-Sevatian boundary (Jadoul et
al., 1994). However, this event seems to occur inside the E. bidentata
Zone in the Lagonegro Basin of southern Italy (Rigo et al., 2005), and at
the Sevatian 1-2 boundary in the Northern Calcareous Alps (Donofrio et
al., 2003). Therefore the Dolomia di Forni most probably spans the
Alaunian 2 – Sevatian 1 interval (Fig. 1).
The basal part of the Calcare di Chiampomano is dated as Rhaetian
by Carulli et al. (2000) on the basis of a small palynomorph sample from
the northern slope of Mt. Verzegnis. This sample contains Corollina
torosus, Retritiletes semimuris, Lycopodiumsporites austroclavatoides,
Trachysporites fuscus, Araucariacites sp. and Lycopodiacidites sp. (Carulli
et al., 2000). However, it does not appear indicative of the Rhaetian:
Corollina torosus and Trachysporites fuscus occur also in the basal part
of the upper Norian, between the middle and the upper part of phase II
of Schuurman, according to Jadoul et al. (1994, table 1). Retritiletes
semimuris, Lycopodiumsporites austroclavatoides, Araucariacites sp.
and Lycopodiacidites sp. are not reported in Jadoul et al. (1994, table 1),
nor are they listed in the palynomorph association indicative of the
Rhaetian phase III in Carulli et al. (1998, fig. 13), in which Trachysporites
fuscus occurs only in the palynomorph association of the upper part of
the phase II (upper Norian). The ammonoid Choristoceras cf. C.
rhaeticum is present in the upper part of the Calcare di Chiampomano
(Paolo Mietto, personal commun., 2005). C. rhaeticum occurs in the
Choristoceras marshi Zone (upper Rhaetian), rendering the upper part
of the Calcare di Chiampomano probably late Rhaetian in age.
DEPOSITIONAL PALEOENVIRONMENT AND
PALEOGEOGRAPHIC REMARKS
The depositional environment of the zone during latest Triassic
times was, from present-day south to north, subdivided into: 1) inner,
tidal carbonate platform, 2) outer platform, 3) platform margin, 4) upper
slope, 5) lower slope, and 6) basin. This lateral succession has a NE-SW
orientation in the western part (Mt. Pramaggiore) (Cozzi et al., 2002)
(Fig. 2). The Dolomia di Forni includes the basinal and slope facies. The
depocenter of the preserved portion of the basin corresponds to the
eastern part of the outcrop area of the Dolomia di Forni (Seazza Creek,
Forchiar Creek). Along the Seazza Creek valley one goes up the strati-
graphic section, but also moves toward the platform margin – thus, both
the basinal and slope facies of the Dolomia di Forni cross (Fig. 1).
The Dolomia di Forni Formation was deposited under prevailing
anoxic conditions (Mattavelli and Rizzini, 1974; Dalla Vecchia, 1991;
Cozzi et al., 2000) in a basin with an estimated maximum depth of 300-
400 m (Cozzi et al., 2000). The basinal facies preserving the tetrapod
remains are composed mainly of distal turbidites and carbonate mud
deposited by fallout and sourced on the platform where it was put into
suspension by storms and tidal currents. Dolomitization of the original
calcite was syndiagenetic.
The presence of terrestrial tetrapods and plants (see below) in the
basinal facies, in addition to the discovery of vertebrate footprints in the
inner platform facies, demonstrates the local emergence of the platform
and the existence of a terrestrial ecosystem. Also, 96-99% of the organic
matter preserved in the shallow intraplatform depression (Mt. Valmenone
Anoxic Event) in the area of Mt. Pramaggiore is continental in origin
(Cozzi and Jäger, 2000).
According to the paleoenvironmental/paleogeographical maps of
Yilmaz et al. (1996) and Gaetani et al. (2000), during the Norian-Rhaetian
the zone corresponding to the present-day Carnian Pre-Alps was situ-
ated at the western end of the Tethys, and was part of a wide,
pericontinental carbonate platform bordering the Laurasian and
Gondwanan megacontinents (Fig. 3). The basin in which the Dolomia di
Forni was deposited (Carnian Basin) is considered the termination of the
larger Slovenian Basin by Cozzi et al. (2000), although no exposed evi-
dence exists today of a connection between the two basins. The origin
and evolution of the Carnian Basin was caused by extensional tectonics
involving the carbonate platform, and is linked to both Tethyan rifting
and the opening of the central Atlantic Ocean (Cozzi et al., 2000).
ASSOCIATED FAUNA AND FLORA
The Dolomia di Forni has yielded an interesting and relatively rich
fauna and flora (Dalla Vecchia, 1991; Sirna et al., 1994). Echinoderms
FIGURE 3. Paleogeographic sketch of western Tethys based on the late
Norian (Sevatian) paleogeographic map by Gaetani et al. (2000). The position
of the Carnian Pre-Alps and the Norian Carnian Basin is marked with the
white asterisk. A, Apulia (southern Italy); AuA, Austroalpine; Ebr, Ebro
Massif (Spain); Hal, Hallstatt (Austria); Ibl, Iblean Plateau (Sicily); IM,
Iberian Meseta (Spain); L, Lombardy; MC, Massif Centrale (France); MM,
Moroccan Meseta; Sar, Sardinia. White, exposed land, continental clastics
and volcanic islands; gray, epicontinental sea; dark gray, oceanic basins.
435
(ophiuroids), cephalopods (sparse hooks), bivalves, and gastropods are
very rare, while crustaceans are the most common fossils. The decapod
peneidean Dusa longipes is the most common crustacean in the eastern,
depocentral part of the formation (Seazza Creek, Forchiar Creek) (Dalla
Vecchia, 1991; Garassino et al., 1996). Acanthinopus gibbosus, Antrimpos
noricus and small peneideans are also present (Garassino et al., 1996).
Benthic forms (Palinura and Astacidea) like Rosenfeldia triasica,
Pseudocoleia mazzoleni, Archaeopalinurus levis, Glyphaea sp. and
Glaessnericaris sp. (Garassino et al., 1996) are more common in the
western part (northern flank of Mt. Auda, Caprizzi, Rovadia Creek,
south of Forni di Sopra), probably because it was deposited closer to the
margin of the platform. Thylacocephalans (Clausiocaris pinnai,
Microcaris minuta) are found mainly in the Seazza Creek section (Dalla
Vecchia and Muscio, 1990).
Chondrichthyans are represented only by tooth batteries of the
diminutive shark Pseudodalatias barnstonensis, while osteichthyans are
much more diverse. However, the fish collection has been only partly
studied to date, so the diversity of the ichthyofauna will probably in-
crease in the future. Durophagous forms are represented by Sargodon
tomicus and rare, small pycnodonts. ?Colobodus, Dapedium and
Heterolepidotus are represented by single specimens, while Paralepidotus
is relatively common in the western part of the formation. Among preda-
tors, Saurichthys is common and ubiquitous, whereas the flying fish
Thoracopterus martinisi occurs mainly in the Seazza Creek section;
some specimens of a coelacanth and a few remains of Birgeria are also in
the sample. Possible members of the Furidae are reported from the west-
ern part of the formation. The most common osteichthyans are small
taxa mostly belonging to Pholidophoriformes (including
Pholidorhynchodon malzannii and Eopholidophorus forojuliensis;
Zambelli, 1990) and Peltopleurus. Relatively common is a still-
undescribed, small, scaleless osteichthyan resembling later, basal teleo-
sts (“leptolepids”).
Terrestrial plant remains are relatively abundant in the Seazza
Creek section, representing about 30% of the fossils collected (Dalla
Vecchia, 1991); they are also found in the western sections. They consist
of single leaves apparently similar to those of the Bennettitales (but
possibly belonging to a long-leafed, Araucaria-like plant), and short
branch fragments of Coniferales, mainly the form genera Brachyphyllum-
Cyparissidium-Pagiophyllum (Dalla Vecchia, 1991). Hygrophytic forms
like ferns and equisetaleans are absent.
SYSTEMATIC PALEONTOLOGY
Diapsida Osborn, 1903
Lepidosauromorpha Benton, 1983
“Langobardisaurus? rossii” Bizzarini and Muscio, 1995
“Langobardisaurus? rossii” (Bizarrini and Muscio, 1995) was
named for an articulated, complete skeleton (MFSN 19235; Fig. 4) that,
unfortunately, is heavily weathered. Furthermore, parts of the slab con-
taining most of the skull were lost. As in the case of Langobardisaurus?
tonelloi (see below), while the authors erected a new species, the generic
attribution of the specimen was tentative. Indeed, MFSN 19235 is not a
member of the Protorosauria (Renesto and Dalla Vecchia, 2000), but is
instead probably a lepidosauromorph (S. Renesto and F.M. Dalla Vecchia,
unpubl., 2005).
Archosauromorpha von Huene, 1946
Drepanosauridae Olsen and Sues, 1986
Megalancosaurus Calzavara, Muscio and Wild, 1981
Megalancosaurus preonensis Calzavara, Muscio and Wild, 1981
The Drepanosauridae is represented by three specimens of the
small and bizarre Megalancosaurus preonensis. The holotype (MFSN
1769; Fig. 5) was the first reptile found and described from the Dolomia
di Forni (Calzavara et al., 1981). It comes from an outcrop along the left
flank of the lower Seazza Creek valley and consists of a partial but
articulated skeleton that includes the skull and lower jaw, cervical verte-
brae, the left scapula and forelimb, and the neural spines of the anterior
dorsal vertebrae. The skull is strikingly bird-like; it convergently ac-
quired some features shared with basal pterosaurs, and lacks an antorbital
fenestra (Renesto and Dalla Vecchia, 2005).
An isolated caudal segment of the vertebral column, also from the
Seazza Creek section (MFSN 1801; Fig. 6 right) was first described by
Pinna (1988) as the tail of a juvenile of Drepanosaurus unguicaudatus
and later identified correctly by Renesto (1994b). A third specimen
(MFSN 18443; Fig. 6 left), described by Renesto (2000), is also an
isolated caudal segment of the vertebral column, but from a higher level in
the Seazza Creek section (Fig. 1). Whereas the terminal, claw-like ele-
ment of the vertebral column is only partly preserved in MFSN 1801,
and was not identified as such, it is complete in MFSN 18443, showing
that this unusual feature is shared with Drepanosaurus (Renesto, 2000).
Protorosauria Huxley, 1881
Langobardisaurus Renesto, 1994
Langobardisaurus tonelloi Muscio, 1997
The Protorosauria is represented by Langobardisaurus, a rela-
tively small animal (around 50 cm adult body length) similar in overall
morphology to the Middle Triassic Macrocnemus (Renesto, 1994a;
Renesto and Dalla Vecchia, 2000; Renesto et al., 2002). A nearly com-
plete, articulated and relatively large skeleton (MFSN 1921, Fig. 7; fe-
mur length is 49 mm) from the Seazza Creek section was first described
by Muscio (1997), who coined the new species Langobardisaurus?
tonelloi (sic). A question mark was used because the attribution to the
genus Langobardisaurus was regarded as tentative (Muscio, 1997). Sub-
FIGURE 4. “Langobardisaurus? rossii,” MFSN 19235, Seazza Creek.
436
sequently, Renesto and Dalla Vecchia (2000) redescribed the specimen,
focusing on the peculiar dentition (slender incisiform teeth, followed by
bulky, tricuspid cheek teeth and a large, molariform, grinding tooth that
has a distal occlusion). The latter indicates a very specialized feeding
habit and a method of food processing rather unusual for a reptile. Renesto
and Dalla Vecchia (2000) showed both that MFSN 1921 does belong to
Langobardisaurus and that it is probably conspecific with L. pandolfii
from the Norian of Lombardy. Pending a detailed comparison to the
Lombardian specimens, however, they retained the original specific de-
nomination. Muscio (1997) failed to identify any truly diagnostic fea-
tures for L. tonelloi and I retain the species here only because a formal
revision of Langobardisaurus is yet to be published. Langobardisaurus
probably fed on hard or tough food (crustaceans, large insects, small
“ganoid” fishes) and could have adopted a facultative bipedal gait (Renesto
and Dalla Vecchia, 2000; Renesto et al., 2002; Renesto, this volume).
Langobardisaurus sp.
A second partial and, as yet, incompletely prepared specimen
(MFSN 26829), was discovered on a stone at the river bed of the Rovadia
Creek where it had fallen from the rocky walls of the valley, later having
been weathered by the stream. It consists mainly of hind limbs (Fig. 8)
that were described by Renesto et al. (2002), but other parts of the
skeleton are still covered by rock. It represents a relatively large indi-
vidual just slightly smaller than the holotype of L. pandolfii (femur
length is 54 mm versus 56 mm in the L. pandolfii holotype).
A third Langobardisaurus specimen (MFSN 24992), discovered
on a stone found on the river bed of Forchiar Creek, is still unprepared
and undescribed. It consists of an articulated and possibly complete
skeleton (the skull is apparently absent, but could be obscured). MFSN
24992 is substantially smaller than other Langobardisaurus specimens
and is very probably a juvenile. Its femur is only 18 mm long whereas
that of the juvenile specimen of L. pandolfii is 27 mm long.
Pterosauria Kaup, 1834
Preondactylus Wild, 1984
Preondactylus buffarinii Wild, 1984
The most common and systematically diverse tetrapods in the
Dolomia di Forni are pterosaurs: their fossils represent 10 of the 17
tetrapod specimens thus far discovered. The holotype of Preondactylus
buffarinii (Wild, 1984; MFSN 1770, Fig. 9) is a nearly complete and
articulated skeleton from the Seazza Creek section. Unfortunately, most
of the bones split away when the specimen was discovered, so its skel-
etal elements were identified on a silicon rubber mold (Wild, 1984; Dalla
Vecchia, 1998). Preondactylus was considered the most basal pterosaur
by Unwin (2003a), but was regarded as more derived than anurognathids
and Sordes by Kellner (2003). MFSN 25161 is a partial skull also from
the Seazza Creek section. It was preliminarily attributed to Preondactylus
by Dalla Vecchia (2003a).
Eudimorphodon Zambelli, 1973
Eudimorphodon rosenfeldi Dalla Vecchia, 1995a
Eudimorphodon appears to be the most common and widespread
Triassic pterosaur, occurring in Lombardy, Friuli, Tyrol, eastern Greenland
and possibly SW USA, Switzerland, Luxembourg and France (Dalla
Vecchia, 2003b). The only unambiguous character shared by the speci-
mens attributed to this genus is the possession of tricuspid and
quinticuspid teeth (Dalla Vecchia, 2003a). However, it is likely that, in
the future, this character will prove diagnostic of a more inclusive clade
(Dalla Vecchia, unpubl., 2005).
The holotype of Eudimorphodon rosenfeldi (MFSN 1797a-b;
Fig. 10), from Forchiar Creek, is a nearly complete, articulated skeleton
with traces of the patagium. It was briefly described by Dalla Vecchia
(1995a) and a detailed description is in progress. It was referred to
Eudimorphodon because it shares quinticuspid teeth and other features
with specimens of this genus from Lombardy (Dalla Vecchia, 1995a,
2003a). Kellner (2003) proposed that E. rosenfeldi is not closely related
to other Eudimorphodon species, but is instead closer to Peteinosaurus.
The character evidence for this is weak, however, with the only pro-
posed synapomorphy for the E. rosenfeldi-Peteinosaurus clade being
the presence of multicusped teeth. Curiously, the same synapomorphy
was indicated by Unwin (2003b) as being the only one uniting
Eudimorphodon and Austriadactylus.
Eudimorphodon sp.
A partial, disarticulated skeleton from Purone Creek in the west-
ern part of the Dolomia di Forni (MFSN 1922, Fig. 11) was described in
Dalla Vecchia (2004). It consists of part of the vertebral column and ribs,
a humerus, part of the second wing phalanx, part of a mandibular ramus
with teeth, and traces of the patagium. It was referred to Eudimorphodon
on the basis of its multicusped dentition and the squared deltopectoral
crest of its humerus.
Specimen MFSN 21545 (Fig. 12, left), found in the stones at the
river bed of Seazza Creek (and consequently of unknown stratigraphical
position in the Dolomia di Forni), was preliminarily attributed to the
genus Eudimorphodon on the basis of its dentition (Dalla Vecchia,
2003a,b). It consists of most of a highly disarticulated skeleton, notably
with a skull and lower jaw bearing tricuspid and quinticuspid teeth.
Specimen MFSN 26823 consists of three articulated wing phalan-
ges, though other parts of the skeleton may be present but covered by
rock. It also comes from the stones at the river bed of Forchiar Creek, and
its stratigraphic position in the Dolomia di Forni is likewise unknown. It
is still unprepared, but, on the basis of the length ratios of the wing
FIGURE 5. Megalancosaurus preonensis, holotype, MFSN 1769, Seazza
Creek. Detail of the skull and hand.
437
phalanges, seems to be referable to Eudimorphodon (Dalla Vecchia,
2003b).
Pterosauria indet.
MFSN 1891 is a pellet of bones (Fig. 13) from the Seazza Creek
section interpreted as a gastric eject (Dalla Vecchia et al., 1989). It is
composed mainly of several long limb bones, some of them broken at
their extremities, and vertebrae (at least one elongate caudal element and
a couple of procoelous dorsals can be clearly identified). The specimen
was first reported as cf. Preondactylus buffarinii (Dalla Vecchia et al.,
1989) at a time when no other pterosaur was known from the Dolomia di
Forni, and Megalancosaurus was the only other reptile discovered in it.
Dalla Vecchia (2003b) showed that it is better to consider MFSN 1891 as
an indeterminate pterosaur rather than a Preondactylus specimen. The
bones cannot belong to Megalancosaurus because this taxon has elon-
gate cervical vertebrae markedly different from the elongate vertebra
preserved in MFSN 1891 (e.g., Renesto, 2000, fig. 2). Also,
Megalancosaurus does not have long bones matching in relative size and
number those in the pellet. The more recent discovery of
Langobardisaurus, which also possesses elongate cervical vertebrae
(Renesto, 1994a, figs. 2-3), procoelous dorsal vertebrae and long hind
limb elements (Renesto, 1994a), might now cast doubt on the proposed
pterosaurian nature of the specimen. However, the number and relative
length of the long bones in MFSN 1891, when compared to the size of
the vertebrae, are unlike those of Langobardisaurus. Also, in the
protorosaurian taxon, the radius and ulna are relatively short, the hu-
merus is substantially expanded at the extremities, the straight femur
tapers proximodistally, and the fibula is significantly more gracile than
the closely appressed tibia (Figs. 7-8). A pterosaurian identification of
MFSN 1891 therefore remains the most plausible.
Other pterosaur fossils recovered from the Dolomia di Forni
are known, but lack features allowing reliable generic identification. MFSN
19864 (Fig. 14), from the Seazza Creek section, consists of most of the
articulated caudal segment of a vertebral column, two fourth wing pha-
langes, and a small portion of the third wing phalanx. It is important
because it, together with other specimens, shows that the tails of some
basal pterosaurs lacked the sheath of elongate pre- and postzygapophyses
characteristic of Jurassic long-tailed pterosaurs. This absence of elongate
zygapophyses is also a feature observed in Eudimorphodon (at least in
FIGURE 6. Megalancosaurus preonensis, isolated segments of the caudal vertebral column. Right: MFSN 1801, left: MFSN 18443. Both from the Seazza
Creek. The claw-like terminal element of the vertebral column is indicated.
FIGURE 7. Langobardisaurus tonelloi, MFSN 1921, Seazza Creek.
FIGURE 8. Langobardisaurus pandolfii, MFSN 26826, partial hind limbs,
Rovadia Creek. as, astragalus; c, centrale; ca, calcaneum; d1-5, pedal digit 1
to 5; dt, distal tarsal; f, femur; fi, fibula; mt I, metatarsal I; mt V, metatarsal
V.
438
specimen MPUM 6009 from Cene, Lombardy, and BSP 1994 I 51 from
the Norian of Seefeld Beds, Austria), Preondactylus (MFSN 1770), and
Austriadactylus (from the Seefeld Beds as well) (Dalla Vecchia, 2002b).
MFSN 19836 (Fig. 15) is an isolated fourth wing phalanx from
Rovadia Creek in the western part of the Dolomia di Forni. With a length
of 137 mm, it shows that large “rhamphorhynchoids” were already present
in the Late Triassic (Dalla Vecchia, 2000).
Specimen S.N. 332466 (Fig. 12, right), from the Seazza Creek
FIGURE 9. Preondactylus buffarinii, holotype, MFSN 1770, Seazza Creek. The interpretative drawing obtained from the cast (reversed) is modified from
Dalla Vecchia (1998).
FIGURE 10. Eudimorphodon rosenfeldi, holotype, MFSN 1797, Forchiar Creek.
FIGURE 11. Eudimorphodon sp., MFSN 1922, Purone Creek. Modified from Dalla Vecchia (2003). cv, cervical vertebra; dr1, first dorsal rib; h, humerus;
lj, lower jaw; pdr, proximal dorsal rib; ppvc, posterior portion of the dorsal vertebral column; wp, remains o fthe wing patagium; wph2, wing phalanx 2.
439
section, consists of part of an articulated skeleton including the skull and
the lower jaw, all of which is in a poor state of preservation. It was
tentatively attributed to Preondactylus by Dalla Vecchia (1994, 2003a),
but actually represents a genus different from either Eudimorphodon and
Preondactylus (Dalla Vecchia, unpubl., 2005).
TETRAPOD ICHNOFOSSILS
During the last 20 years, abundant paleoichnological evidence of
terrestrial tetrapods has been discovered in the Dolomia Principale of the
Dolomites (Mietto, 1988, 1991; Dalla Vecchia, 1995b). Footprints have
also been found in the inner carbonate platform facies of the western
Carnian Pre-Alps, a few kilometers south and southwest of the basinal
unit with tetrapod skeletal remains (compare Figs. 2 and 16). These
footprints are found only on the surfaces of boulders that have fallen
from the mountain walls because erosion rarely exposes the bedding
surfaces, and when this occurs, the surface in often weathered and
karstified. Footprints were preliminarily described by Dalla Vecchia &
Mietto (1998) and Dalla Vecchia (2002a). The field work involved in
these studies was difficult because the boulders are mostly located in
remote, dangerous sites. To date, 10 or 11 footprint-bearing boulders
have been identified (one is doubtful); they occur in Andreis, Claut and
Cimolais municipalities of Pordenone province (Fig. 16).
Medium-sized, tridactyl, mesaxonic footprints (lengths between
15 and 25 cm) with narrow digital prints and a median digit substantially
more prominent than the other two digits, are the most common (upper
Susaibes Creek, boulder A, Fig. 17B; upper Susaibes Creek, boulder B;
Ciol del Tramontin, boulder A, Fig. 17C; Ciol de la Fratta, boulder A, Fig.
17D; Ciol del Tramontin, boulder B, Fig. 17E; southern flank of Mt.
Caserine Basse). Some occur in short trackways (Ciol de la Fratta, boul-
der A, upper Susaibes Creek, boulder A) left by bipedal track makers.
The footprint shape, if accurately reflecting the actual foot morphology,
suggests that the track makers could be “theropods” sensu lato (i.e.,
including more basal saurischians). Larger, tridactyl and mesaxonic foot-
prints, up to 35 cm in length and also probably “theropodan,” are rarer
(Casera Cjasevent, Fig. 17A; Settimana Valley).
A single, 18 cm-long footprint is tetradactyl (Ciol de la Fratta,
boulder A, Fig.17F). Dalla Vecchia and Mietto (1998) noted that it re-
sembles Sphingopus of the French Middle Triassic and tentatively iden-
tified it as the pedal print of a small prosauropod; alternatively, it could
belong to an herrerasaurid basal saurischian.
A single, large, but poorly preserved footprint in boulder A of
upper Susaibes Creek appears to be pentadactyl (Fig. 17G). If actually
FIGURE 12. Left: Eudimorphodon sp., MFSN 21545, Seazza Creek. Right: Pterosauria (Dalla Vecchia, unpubl., 2005), S.N. 332466, Seazza Creek; scale
bar equals 50 mm.
FIGURE 13. Pterosauria indet., MFSN 1891, Seazza Creek. Gastric eject composed of pterosaur bones. Drawing after Dalla Vecchia et al. (1989), modified.
cv, caudal vertebra; dv, dorsal vertebra; lb, long limb bone.
440
pentadactyl and not the result of the overlap of two footprints, it could
have been produced by a large dicynodont therapsid (Lockley and Meyer,
2000). If tetradactyl, it would resemble ichnotaxa like Evazoum (Nicosia
and Loi, 2003) and Otozoum (Lockley et al., 1996). Rainforth (2003) did
not consider it referable to Otozoum (an Early Jurassic ichnotaxon), but
rather to Pseudotetrasauropus (= Evazoum), an ichnotaxon regarded by
her as having been produced by a crurotarsan archosaur contra Nicosia
and Loi (2003) who considered it prosauropodan in origin. Boulder B
from upper Susaibes Creek contains small (13 cm long) tetradactyl prints
(Fig. 17H). Misshapen, shallower, and larger depressions associated
with the tetradactyl prints were interpreted as possible pedal prints, and
the tetradactyl tracks interpreted as the manual ones, of a single track-
way produced by a quadrupedal track maker (Dalla Vecchia and Mietto,
1998).
A trackway with footprints 18 cm long preserved in the Scandoler
Valley boulder (trampled by many track makers) was left by a bipedal
track maker (Fig. 18, upper). The pace angulation is relatively low (140-
160°, based on photographs), the stride is around 80 cm, and the foot-
prints do not seem to be tridactyl (direct observation was not possible
because the footprint-bearing surface was vertical, and the huge boulder
collapsed before I could visit the site). Dalla Vecchia and Mietto (1998)
hypothesized a small, bipedal prosauropod as the possible track maker.
A wide-gauge trackway of a quadrupedal tetrapod is preserved on
boulder B of Ciol de la Fratta (Fig. 18, lower). Its pace angulation is low
(102-111°) and pedal stride is 70-75 cm. The pes print is elliptical, 27-28
cm long, with slight or no rotation; the manual print is smaller, circular,
and impressed in front of, or slightly medial to, the pedal one. According
to Dalla Vecchia and Mietto (1998) it may have been impressed by a
prosauropod moving on all four limbs, or by a large, relatively wide-
bodied crurotarsan archosaur like an aetosaur.
This paleoichnological sample testifies to the passage of large to
medium-sized tetrapods across the tidal flats of the inner carbonate
FIGURE 14. Pterosauria indet., MFSN 19864, Seazza Creek long segment
of the caudal vertebral column and wing phalanges. cv, caudal vertebra; wph
3,4, wing phalanges 3, 4. Numbers do not refer to the actual position of the
vertebrae as the proximal elements are missing.
FIGURE 15. Pterosauria indet., MFSN 19836, Rovadia Creek. Large wing
phalanx 4.
FIGURE 16. Location of the boulders with footprints of terrestrial tetrapods
discovered in the western Carnian Pre-Alps. 1, Near Casera Cjasevent; 2,
Scandoler Valley; 3, Ciol de la Fratta, boulder A; 4, Ciol de la Fratta, boulder
B; 5, Forcella delle Pregoiane; 6, Ciol del Tramontin, boulder A; 7, Ciol del
Tramontin, boulder B; 8, Upper Susaibes Creek (boulders A and B); 9,
Settimana Valley; 10, Mt. Caserine Basse. Modified from Dalla Vecchia
(2002a).
FIGURE 17. Tetrapod footprint sample from the Dolomia Principale of
the western Carnian Pre-Alps. “Theropod” footprints: A, From Casera
Cjasevent. B1-2, from upper Susaibes Creek, boulder A. C, From Ciol del
Tramontin, boulder A. D, From Ciol de la Fratta, boulder A. E1-2, From Ciol
del Tramontin, boulder B. F, Tetradactyl footprint from Ciol de la Fratta,
boulder A. G, Pentadactyl footprint from upper Susaibes Creek, boulder A.
H1-2, Tetradactyl manual print from upper Susaibes Creek, boulder B.
Modified from Dalla Vecchia and Mietto (1998).
441
platform. This fauna was probably relatively diverse and included large
“theropods.” It is, together the report of Eubrontes in the Dolomia
Principale of the close eastern Dolomites (Mietto, 1991), further evi-
dence that large bipedal dinosaurs (theropods or basal saurischians) were
already present in the Late Triassic (Tanner et al., 2004; contra Olsen et
al., 2002). Because the footprints occur on fallen boulders, pinpointing a
precise location of the footprint-bearing levels in the stratigraphic col-
umn is not possible. As noted by Dalla Vecchia (2002a), it is unlikely
that ten boulders found randomly in different localities contain parts of
just a single footprint-bearing surface. Therefore, the footprints come
from several different stratigraphic levels. Dalla Vecchia (2002a) sug-
gested that the boulders from the upper Susaibes Creek, Ciol del Tramontin
and probably the Ciol de la Fratta localities are from the middle to upper
part of the Dolomia Principale (Norian-Rhaetian). This is because the
boulders had fallen from levels relatively high in the local stratigraphic
section (where the nearly vertical mountain walls are hundred of meters
high). Footprints have, in some cases (Casera Cjasevent and upper
Susaibes Creek), been impressed on “algal” mats, i.e., they are preserved
in the stromatolite intervals of the intertidal facies.
DISCUSSION
Keeping in mind the relatively small size of the sample, some
general features of the latest Triassic tetrapod faunas of northern Friuli
can be discussed. The tetrapods from the basinal Dolomia di Forni were
all small animals that inhabited the emergent areas of the carbonate plat-
form, and this is probably where the plant remains also came from.
Megalancosaurus displays scansorial adaptations and arboreal habits
(Calzavara et al., 1981; Renesto, 1994b, 2000), Langobardisaurus was a
possible facultative biped with no evident adaptation to aquatic life
(Renesto et al., 2002), and “Langobardisaurus? rossii” also lacks adap-
tations for living in the water. The dominant pterosaurs were obviously
linked to the presence of an emergent area.
Although a detailed taphonomic study has yet to be undertaken,
the articulated state of many of the skeletons suggests limited transpor-
tation after death. It also appears that disarticulation of some specimens
FIGURE 18. Tetrapod trackways found in the Dolomia Principale of the western Carnian Pre-Alps. Above: Bipedal trackway and other footprints in the
boulder of Scandoler valley (now destroyed). Below: Quadrupedal trackway from Ciol de la Fratta, boulder B. m, manual print; p, pedal print. Modified from
Dalla Vecchia and Mietto (1998).
(MFSN 1922; MFSN 21545) probably occurred after deposition at the
basin bottom by slow decay and very weak bottom currents.
No marine or semi-aquatic tetrapods, such as those found in the
Calcare di Zorzino and Argillite di Riva di Solto of Lombardy (e.g., the
thalattosaur Endennasaurus, the phytosaur Mystriosuchus planirostris
and the placodont Psephoderma alpinum) have been found in the Dolomia
di Forni. Sauropterygians (placodonts and eosauropterygians), common
in the shallow marine Carnian of northern Friuli (Sirna et al., 1994;
Rieppel and Dalla Vecchia, 2001), are absent. This could mean that the
basin and its margins were not suitable for shallow-water marine tetra-
pods. Furthermore, the absence of pelagic reptiles (ichthyosaurs) and
invertebrates (ammonoids), indicates a separation of the Carnian Basin
from the open sea and further suggests that the northern margin of the
basin was bordered by the carbonate platform.
Bones of larger terrestrial tetrapods, whose existence in the inner
carbonate platform is verified by their footprints, were not found in the
basin. If the track makers and small tetrapods from the Dolomia di Forni
were contemporary, then size-biased transport of the carcasses into the
basin must have occurred, possibly because a barrier existed between the
inner carbonate platform and the basin which prevented the transporta-
tion of carcasses into the basin. Small reptiles could be transported by
the wind during hurricanes, and the pterosaurs could fly above the basin.
However, the total absence of trunks and large branches in the plant
sample, and the prevalence of single leaves and small (2-15 cm) conifer
branches, suggests a transportation by relatively mild winds rather than
storms or hurricanes. Possibly the transport of small animals into the
basin by tidal currents was easier than that of heavier-bodied archosaurs.
Also, small tetrapods could have been more numerous than larger ones,
and it may simply be that the tetrapod sample in the basinal facies is just
too small to be representative.
Alternatively, perhaps larger tetrapods lived on the emergent part
of the platform either before or after the time of deposition of the tetra-
pod-bearing part of the Dolomia di Forni. As reported above, no defini-
tive stratigraphic and biostratigraphic data are available to date the foot-
print-bearing boulders, and their ages could range from late Carnian to
442
Rhaetian. During the Alaunian 2-Sevatian 1, the size of the emergent part
of the carbonate platform could have been reduced as a result of local
variations in subsidence and/or eustatic sea level changes (such changes
would have had major effects on the flat carbonate platform, even if
minimal). The reduced landmass (probably an island or an archipelago)
may have been capable of supporting only small tetrapods like
drepanosaurids, protorosaurians and pterosaurs. Large and medium to
small-sized theropods, other dinosaurs, and/or other relatively large tet-
rapods, could have reached the marginal tidal flats when the emergent
part of the platform was wider (Dalla Vecchia, 2002a). According to the
global eustatic curve based on the relative change of coastal onlap, the
Alaunian 2-3 coincided with a marine highstand, whereas the Carnian-
Norian and Alaunian-Sevatian boundaries were marked by major drops
in sea level (Haq et al., 1988). However, according to Cozzi et al. (2000),
tectonics influenced the local sequence stratigraphy along the margin of
the platform/basin complex, which shows a highstand system tract that
extends to the upper part of the Alaunian 3, a transgressive system tract
in the ?uppermost Alaunian 3–Sevatian and a drowning of the marginal
part of the platform during the Rhaetian. However, nothing is known
how global eustasy and local tectonics influenced the emergence of either
the southern or northern portions of the platform.
The tetrapod diversity represented in the latest Triassic basins of
Lombardy is higher than that found in the Friulian (Carnian) basin (see
Renesto, this volume). This is obviously due to the fact that the
Lombardian sites were the object of organized field work at fossil-rich
sites. However, the number of Lombardian tetrapod specimens is only
double that of specimens from Friuli. About 30 specimens come from the
Calcare di Zorzino and three from the Argillite di Riva di Solto (Renesto,
this volume). Therefore, the differences (mainly, the absence of aquatic
taxa) could be indicative of different paleoecological conditions.
Some tetrapod taxa are shared by the Friulian and Lombardian
samples. Megalancosaurus preonensis is found in both, although com-
parison between the Lombardian and Friulian specimens is limited by
the fact that no skulls are preserved in the Lombardian sample, whereas
no hind limbs or pelvic girdles occur in the Friulian specimens.
Langobardisaurus probably represents a single species (L. pandolfii).
The pterosaur faunas are not so similar. Peteinosaurus is not found in
Friuli and Preondactylus has not been unambiguously reported in Lom-
bardy (Dalla Vecchia, 2003a, b; contra Wild, 1984). Eudimorphodon
rosenfeldi differs from the holotype of E. ranzii, but resemble another
Lombardian specimen attributed to Eudimorphodon (Dalla Vecchia,
unpubl. 2005).
It should also be noted that, while some fishes are shared by both
regions (e.g., Pseudodalatias barnstonensis, Sargodon tomicus,
Paralepidotus ornatus), others belong to different species. The flying
fish Thoracopterus is represented by T. magnificus in Lombardy, T.
martinisi in Friuli (Tintori and Sassi, 1992), while Eopholidophorus
forojuliensis occurs only in Friuli (Zambelli, 1990). T. martinisi is con-
sidered more primitive than T. magnificus (Tintori and Sassi, 1992).
Differences between the Lombardian and Friulian tetrapods could
be due to different environmental conditions, as well as to slightly differ-
ent ages. The Cene and Endenna/Zogno sites, in the uppermost part of
the Calcare di Zorzino (just below the boundary with the Argillite di Riva
di Solto; [Renesto, this volume]), and the Ponte Giurino/Berbenno site in
the lower part of the Argillite di Riva di Solto, contain palynomorphs of
the middle-late Norian Phase II of Schuurman (1979) (Jadoul et al.,
1994). Jadoul et al. (1994, p. 8) specify that the “palynological assem-
blage documents the Middle-Late Norian boundary” in the fossiliferous
horizon of the uppermost part of the Calcare di Zorzino, but do not
discuss in detail this dating. No conodonts are reported from the Calcare
di Zorzino and Argillite di Riva di Solto, therefore the dating of the
Friulian and Lombardian basins cannot be compared in absence of a
crossed control of the conodont and palynomorph biostratigraphy.
A thorough comparative study of both fish and tetrapods should
be undertaken to reveal systematic similarities and differences. Further
stratigraphic (paleomagnetic, isotopic, etc.) and micropaleontological
investigations could improve the correlations among the latest Triassic
basins.
The work on the vertebrate fauna from the latest Triassic of Friuli
is just in its earliest stages.
ACKNOWLEDGMENTS
I thank Dr. Carlo Morandini and Dr. Giuseppe Muscio, re-
spectively Director and Curator of the Geo-paleontological section of
the Museo Friulano di Storia Naturale, Udine, for the access to the
collections under their care. Thanks to Prof. Paolo Mietto and Dr. Manuel
Rigo (University of Padua) and Dr. Guido Roghi (CNR Padua) for the
personal communications. I am grateful to Prof. Silvio Renesto (Univer-
sity of Insubria, Varese, Italy) for the information, to Dr. Jerry D. Harris
(Dixie State College, St. George, Utah), Dr. Spencer Lucas (New Mexico
Museum of Natural History & Science, Albuquerque) and Darren Naish
(University of Portsmouth, United Kingdom) for their reviews of the
manuscript.
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