Societn Paleontologic a - Società Paleontologica Italiana

rssN0375 7633
Bollettino
della
a
SocietnPaleontologic
Italiana
Pubblicatosottogli auspicie col contributo
del ConsiglioNazionaledelle Ricerche
Volume 4lrr. 1r2002
MTJCCHI- MODENA
Bollettino delh Societi Paleontologica ltaliana
4r (r),2oo2 rssN0375-7633 4r-50
I pl.
Modena,Aprile 2002
Isocltirotherium inferni ichnosp. n.
in the Illyrian (Late Anisian, Middle Thiassic)of Adi
Giuseppe LEoNARDI
Marco AvnNzlNt
Naturali
MuseoThidentinodi Scienze
Thento,Italy
Morbiac Dark Lt
KEY WORDS - Tbtrapod
footprints, Isochirotherium, Icbnosystematics,
ABSTRACT - In the lower bedsof the Morbiac Dark Limestone(Ilfirian, Late Anisian,
which are still under study.
Bolzano) seueralarchosaurfootprints hiue beenfound, someof
"shortened
morphology thay sugg
Some large chirotheilld' Tootprints shoi a peculiar
Isochirotheritim inferni. Cokporiton sugl{eststiot the Termeno ichnoipecieTcould oPh
Ig4STut ort that is shorterand moreiobust (L/W ratio =
IsochirotheriummarshalliPeaSody,
RIASSUNTO - llsochirotheriuminferni n.sp. nell'lllirico (Anisicosuperiore,ThiasMed
'-::"':::i!;::f;;::,f;S;f:;,';t;:i:#,:;!'t;{!!!r:::^l:'i'::;tH#',f
,Vl'f,{il:{:,f;,;'^
pe
comecontroimpronte(epirilieui), mostrunocaratteristiche
Alcunegrandi or-ri conseruAte
Isochirotheriun
Isochi?otheriuminflerni n. ichnosp.Tra i ,ipprrttntanti dell'icnogenere
sDecie
"Landmark
per le orme di TbrmeVo,
l'attribuzionead ur
analysis"pArrebbisuggerire,'
Iioo detk
larga (rapportoL/W= t,43)i moifologica*tt similea Isochirotherium marshalli Peabody,I9<
duhqueunaforma di Isochirotheriumofrrt al. marshalliper il gruPPodigitaleI-M ma di ass
metitarsale V menosuiluppato.
OUTLINE
INDRODUCTIONAND STRATIGRAPHIC
The middle Triassic sedimentary Units of the
Adige Valley and Upper Valle di Non are currently
Geological
for' the National
beirig rrttii.d
CartSgraphy Project (the CARG Project for Thento
and g6lzin olBolzano Province). On the basis of this
researchit has been possibleto recognise,in the interval between the iop of the Verfen Formation
(Spatian) and the base of the Contrin Formation
(Late.Illyrian), at least three terrigenous-cgTglomeratic units and two carbonate tida[ units with tetrapod
footprints (Avanzini 6{ Neri, 1998, Avanzini, 1999;
Avanzini, 2000, Avanzini et al., 2000; Avanzim et al.,
2001). The Upper
-its Anisian carbonate successionis
base by a sequence of siltstones
characterised at
and dark grey clays with coal. This is followed by a
peritid"l ril...ssioh of limestones and yellowish-grey
siltstones.The subtidal layers are of bioturbated bioclastic yellowish wackstone-packstone; the intersupratidal layersare essentiallymlde up of micaceous
siltstones and blackish-grey sandstones.These units
can be related to the M;biac Dark Limestone of the
Eastern Dolomites, Anisian (Illyrian) in age (De
Zanche et al.,1993; Gianolla et al., 1998). A relatively rich ichnoassociationin which various ichnogenera
i^n be recognised,comes from the basal levels6f thit
unit. Amoigst the association, the most common
Maidwell,
ichnogenera appear to be RhynchosAuroides
191 l, Synaptichniun Nopcsa, 1923 and Chirotherium
Kaup, 1835 (Avanzini 6c Mietto in prep.).
A group
of large tracks has been found in the out-of
Morbiai Dark Limestone on the western
crops
side of Adiqe Valley aboveTermeno, a few kilometres
south of B5l zano. Most of these are concentrated on
the same layer which crops out along Rio di Inferno,
at the baseof Monte Roen (Text-fig. 1).
PALEOICHNOLOGICAL DESCRIPTION
The footprints are preservedas natural casts(concave epirelief) at the base of a level of bioclastic yellowish- grey packstone, cropping out in the shapeof
a roof, located a linle more than 2 metres above the
base of Morbiac Dark Limestone (Text-fig. 2). The
underlying layer, fragmented by mud cracks, is made
up of ihi.r- layers of irarls and siltstones with vegetal
remalns.
The main tracked surface is a little larger than 6
mt. This surface bears over thirry prints. The morphology of the tracks is relatively variable and obvibusly tr.p.trds on the deepnessof the impressions in
a substrite that was very heterogeneous.In several
cases,they are preserved'as,tttdeiprints and because
of this many morphological details are difficult to
recognise. Sbmetimes, e"rosion has made both the
prini and underprint visible in the same impression.
i.t caseswhere the print is deeper the digiti appear
M. AVANZINI, G. LEONARDI
42
Abel, 1935
Form-Family CHTnoTHERIIDAE
Ichnogenus IsocutRorHERIUM Haubold, 197 |
ichnosp. n.
IsocHIRoTHERIUMINFERNT
Pl. 1; Text-fig. 3
Deriuatio nominis
localiry of the species.
After Val d'Inferno,
ryPe
/GAMPENPASS BOLZANO
E
J
ci
o
PPIANO
o
G
o
E
j
(9
C)
Irex'.ng
fr:'ffi::
l;F;t5|'';*H}l}i$i5#"t'
juxtaposed and come together in a depression that
iacks morphological,elernents.In casesin which the
pnnts are shallow, the first four digits are distinctly
i.p"rated from the fifth. In intermediate cases the
conn.ction berween the first four digits and the fifth
pads I-IV is visible. In
by the metatarsal-phalangal
-the
print is very superficial, only
some cases,where
the digits I-IV are visible with good morphological
detail.
Thacesof manus prints are rare. Manual prints are
not clearly marked ind have a mostly rounded outlines; in severalcasestracesof digits II-IV are recogpes associationswere
nisable. No certain manus
identified on the main track-bearingsurface.
On an adjacent outcropping surface,other tracks
were found, that can be atributed to the same form;
amongst these a manus pes coypLe ryas identified
(BzTVI/1 - Bolzano Termeno Val d'lnferno, no. 1).
Due to the characteristicsof BzTYII I and the general characteristicsof the tracks found, as well as on the
basis of the relative distance berween them, when
associatedin a trackway, they are attributable to:
Class Rlprlln
SubclassARcHosAURIACope, 1891
z
UJ
lr
o
I
F
I
I
E
j
(,
o
o
(9
F
J
o
oE
?m 8ffi
=lI
9
1 0 1 1
Text-fig . 2 - Stratigraphic section o[ the Anisian successionoutcropp'ing along the Inferno (Hell) River. The
.trlire.ddiaeram on the right shows the track-bearing liyer. I )"dolostone, Zf bioclastic packstone, 3)
midy-limestone, 4) sandstone, 5) sifry-sandstone,
6) siltstone, 7) conglomerate,8) silry-limestone, 9)
red, l0) grey, I l) yellow.
43
ISOCHIROTHERIUM INFERNI IN THE ILLT'RUN OFADIGE VALLEY
Y''/'
i
- % , i >
tL-=-----1
')
Text-fig.3 -Isochirotherium inferni ichnosp. n.,. holoryPe
"ll'*1's*":11'!'3"lai's
:3;ff.jll,ytli"!::
set of right pes and manus
Holoty\e - Bffinll,
(incompl6te)reverseprint (Pl. I ; iextJig. 3).
\rr,
Typelocality- Val d'Inferno (Hell Valler, eastern
sidebf Mt. Roen,Termeno(Bolzano).
F
Text-fig. 4 -Th. main track-bearingslab. In the lower PaT, two
lncomplete footprints are associatedin a trachvay.
Scde = I m.
\
-\
44
Typt leuel
LateAnisian.
M. AVANZINI, G. LEONARDI
Morbiac Dark Limestone, Illyrian,
respectto the midline of 10"
of about I 60".
15" and a Paceangle
Remarks- Due to the greater length of digit II in
respect to IV and the relaiionship between the other
(like the other prints from the same
dieits, BzNIll,
tra'ck-bearingsurface), can be attributed to the ichnosenus Isochi/otheriumformalised by Haubold in 197 | .
Tn. characteristicsof this ichnogenus are ? Pes.witll
longer digits II and III and digi, Y smaller than digit I
GII;II>IV>I>'VI, baseof digiiV located within group
I-M distal phalangesof digit V not very Pronounced,
thai the foJt (l:4.5 ,tP to l:6.8), angle
manus r-il.r
Depository-MuseoThidentinodi ScienzeNaturali,
pace
165", hand and foot divergence is about 20".
of
Tlento (Italy).
ttt. relatively large dimenslons of the. print
I suggesti l"tgJr sized members of the ichnoBzTYII
Material BzTVI l2-3. Tivo isolatedpedal prints
Isochirotherium.
senus
situ.
in
l,
BzTVII
of
slab
(naturalcasts)on the same
c)
Isochirotheriumherculis (Egertor, 1839) (Text-fig.
asnaturalcasts
BzTM 14-29:25footprintspreserved
5. 1), is a large chiroteriid wiih a Pes longer than 30
slab6 m' *ide, in situ (Text-fi8. 4).
on a track-bearing
.-. Digit Il"has almost the samq.length,as digit III
and sorietimes it is even longer. The angle between I
Description Pes:wide and stout with a low
and IV is greater than 50" "id the crosi axis is close
reland
robust
I-IV
digits
ratio
length/width
1t.43),
to 80'. Th"e I-IV group is wider than its length. The
ati.ftly short.The I-IV grouf is 15.5 cm wide (digit
rype speciescomes-from the British Anisian (Tarpoley
III length: 13 cm) and is wider than its length.JJl.,
Sherwood Sandstone Group; Theasise&
Siltrrotr.
IV
and
III
Digits
45".
is
diverqJtt.. of group I-IV
It is a form that, while it is similar to
1997).
Sarjeant,
so
much
I-III;
to
grouP
inrespect
diverfiedistin.tTy
:o
to be more robust with a slightappear
TVIll, would
that ,Tt. IVIII diverg.ttc.anglEis greaterthan that of
I-IV
wider
lv
sroup.
III-I (25" compar.f, to 20"). The metatargaJ-ph1-_ '
The pes oT ltothirotherium demathieui Haubold,
langeal.pad
y ir well pronounced.The base"f$lgilv
5.2) reaches22 cm in length;jlt.
197| (Text-fig.
alongthe axisof the P{ilt (digit III). On
"more
rsposrttoned
or less as wide as its length. The
srouD I-lV is
dieit I rwo phalarieealpadsarevisible.tices of skin
il "_characterdigir_Y_b.elongargd
of
end
foldr that probabl/ correspond
fiUi"geal
^pg4yto the joints berween
(32"1 and the
low
is
I-IV
qulte
".gL
the
-int.r,
istic
.t.titn)
by
removed
phalang."i p."dt (ghat"_tS
.
to
close
is
7
5"
axis
cross
metatarsalThe
IV.
ahd
iII
II,
are prer.r,n.i on digits
Isochirotherium d.emathieui comes from the
phalangealpads "tE not particularly develoP.^d,ih.
Middle French Thias as does Isochirotherium circad.eb"r.r oTdisits II, III and IV seemto consistof a sinma*ieui Gand, 1979, and Isochirotherium coureli
ele wide ira. The digits end in robust triangular
(Demathieu, 1977) (Text-fig. 5.3), which have pedes
digit
of
claws
The
ihaped clawswith a rolnded tiP.
a lensth that is compirable to the above menwith
of
is
that
longest
the
while
its
base,
at
widest
IV is the
tioned "ttd."ch have an I-IV divergence that is quite
on digi, l_is smallan-drounded.
digit
"(LtIII. The claw
\W:
low (30"; and a crossaxis close to 90". The low I-IV
III:
205mm; 160mm;LI: 75mm,II: 117mm,
of these forms makes their comparison
divergence
to
equal
90o).
13 1lrrln, N: 97 mm,V: 102mm.Cross-oris
improbable.
TYVt
with
with
is-small
V
Digit
dimensions.
small
Manus:of
(Text-fig.,
Isochirotherium soergeli(Haubold, 1967)
a rounded claw and separatedFrom grouP .I-IV
"small
manus"
the
of
teptJs.ntative
first
the
was
5.4),
(which is incomplete).DirrergenceI-IV450", diverBuntsandstein
from
recognised
chirotherii
of
nto,tp
genceI-V: 97".
'')
lscythian) (Haubold, !9O). The dimensions of the
(L:?;\7: 95mm; LI:26mm, II: ?, III: ?, IV: ?,V:
feei are much smaller than in our specimensand vary
22mm).
from 4 rc 15 cm. Digit IV of the foot is much longer
incompletepedesalignedin a PosTra,ckwaysiTwo
than I. Dieit II and III ar. almost the same size.The
sible tr".L*ay (Texr-fig.4). In this casethe distances
Is close to 90" and the external digit has a
cross-axis
a
cln,
60
oblique
would seemto suggesian
Pace9f
angle between 30 and 40".
divergence
in
(sensu
1987)
Leonardi,
angle
positivedivaricatiJi
Stout foot with
Diaqnosis- LarseIsochirotheriltm.
Foot
III>II>IV>I>V.
Digit
trigitt.
robust
short "'"a
digit V is divergent ( 19") to .digir--glgyP I-IV.
Dlvarication I-IV 45", divarication IV-III greater
than III-I (25" 20"). Crossaxis 90". The manus
print.issmalland in front of the foot (manus.digr_tly
rn axrswrth digit III of the Pel. Manus digit V is
small and t.patite from digit group I-IV.
EXPLANATIONOF PLATEI
couple.
n., holotype(BzTVI/1).Manus-Pes
inferniichnosp.
Isochirotherium
, \ l , 1 \ A l { Z l M , ( ; I - E O I , I A R D I ,S O C H I R O T H E R I ( r A l l V F E R h rIlN T H E I I - I - Y R I A No
I ) 1 I.
M. AVANZINI,
46
G. LEONARDI
Text-fig. 5 - 1) Isochirotherium herculis, 2) I. dtmathieu|
[ coureli,4) I. soeygeli
?)
i
i)i;,!;i'!::ff;:;,'J,
rc
197 |
Isochirotherium hessebergenseHaubold,
(Text-fig. 5.5) has pes dimeniions !h1t Iaqge berween
15 and 18 cm, the group I-IV is slightly longer than
it is wide. The digital divergence I-IV is very small
(29"). Digit I is longer than IV. These are all characteristic t[at make iivery different from our material.
Isocbirotherium lomasi (Baird, 1954) (Text-fig.
5.6), from the British Anisian has feet close to 18 cm
lone. Group I-IV is longer than its width and in gen.ruI"i, has . -ot. slende-rmorpholory than our fJrm.
Isochirotherium coltoni (Peabody, 1948) (Text-fig.
5.7) of the Scythian-Anisian bound ary (Lower
Moenkopi), has digit IV shorter than III and more or
less the lame dimtnsions of II. The foot is much
smaller than TVI/ 1, and measures about 12 cm.
Group I-IV is relatively long and the metatarsal-phalangeil pad of V is relatively t-41.
In ]sochirotherium marshalli (Peabody, 1948)
(Text-fig,.5.8) the foot is characterisedby rather short
disits. The base of disits II-III is closed in a
-Et"t"rsal-phalangeal F"d, as also occurs in
Chirotherium bartFi Kaup, 1835. The edge of digits I-
marshalli.Scalebar = l0
cm. lAfter Haubold,
(1971a)modifiedl.
IV is indented berween the base of I and IV. Digits I
I-IV are
and IV divergence is 40". The claws of digits
"
in betwe.n" the robust triangular claws of
Chirotherium barthi and the spatulf shaped claws of
Chirotherium rex Peabody, 1948. The cliw of digit I
is smaller than the others. The phalangealportion of
digit V seems much smaller in respect to the large
ovoidal shaped metatarsal-phalangealpad. This phalaneeal pad-is relatively lari.t
"of than in other chirotheChirotherium rex and
riaris *ith the .*..plion
Chirotherium moquinensePeabody, 1948, where it is
more angled with an elongated shape.
The itack BzTYll l, isielativeli comparable with
f. marshalli, with regards the parameters relative to
digits I-IV. Although-it is knowh that the inter-digital
"tigt..-1y u?rydeien{i"g on the substrare,the it.t;
enie in both forms of an accentuated divergence of
digits III-ry in respectto the others appearspeculiar.
H6wever, the wide metatarsal-phalangealpaa V that
characterises^L marshalli, is much less pronounced in
TVI/ l. Similar differences can also be found in the
morphology and robustness of the claws, which, in
ISOCHIROTHEzuUMINFERNI IN THE ILLYRIANOFADIGE VALLEY
rextng
-7-
47
'iiftri::i:ifii4i#i*#l*
?:jr,,#r;t
;;X;:;p,X:!,1:iL::
H,.'f,li::
;ffi ;
posed the use of the Landmark anafusismethod for
,lt. comparison of tetrapo4 ichnofauna. This application seems to have potential and appears ro provide
another instrument for the comparison of ichnospecies "qd ichnogenera along side traditional
morphometric methods.
The seven fixed points of the print (Text-fig. 6)
(Karl 6c Haubold, 1998), are connected togetheiby a
rex'ng6-tr#Xif,3 ';ii:'*:
f.:u+n+,f:J2:(y#re!r;
the Termeno specimen are in general slender and
more triangular (similar to Chirotherium barthi).
Comparison with other representatives of this
group does_not give risg to any substantial morphological conformiry and for these there are nor enough
coincidirg morphological parametersfor an attribution to be proposed. Futhermore, f. inferni is shorter
and wider than all the other forms referable to the
Isochirotherium ichnogenus, with a L/\f ratio that
appearsto be peculiar (1 .43) and which would place
it outside the field of variabiliry for the ichnogenus
(Tab. I ).
\
)
I
I
J
,,LANDMARK
ANALYSIS'
The differentiation of tetrapod ichnofauna is difficult and comparison between the various forms is
sometimes complex. Th e Landmark analysrrmerhod
was proposed for the first time for tlie study of
dinosaur skeletons (Chapman, 1990; Chapman 6(
Brett-Surnam, 1990). This method involves-theconstruction of polygons, the vertexesof which are fixed
by particular anatomic poles. These polygons can be
easily compared between themselves to evaluate
analogies and incongruities between different specimens. Recently Karl and Haubold ( 1998) also pro-
\I
-l
3
l
Y
,r"
------.-
.r//
---,-----'
t'
L -)--->
Text-fig. 8 -The orientation of /sochirotherium inferni footprints
'Dark
and trackways at the base of Morbiac
Limestone from Val d'lnferno, indicares a possible
preferred direction that is parallel to a river-or tidal
channel. The arrow shows the prevalenr
currenr
directions. (N= 25, Interval: l0').
M. AVANZINI, G. LEONARDI
48
taxon
I. inferni
I. hirculis
I. herculis
I. marshalli
I. soergeli
I. hessbergense
I. lomasi
I. demathieui
I. coltoni
I. coureli
L jenense
L/\7 ratio
r.43
r.6
r.75
r.g
r.7
r.76
r.9
r.6
r.76
2
r.7
Thb. I - Length/\7idth ratio of some Isochirotherium ichnosPecles.
polygon (A - G) that characterisesthe morpholory of
ih.' t-rack. Tb. comparison berween different Polygons is made by superimposing the constructlons creIted using the t"-i fixed comiron scale(G - F) (Karl
6c Haubold, 1998).
The comparison of polygons relative to various
speciesof Isochirotheriri _(Hi"hgld, 1967; Haubold,
tgZla; Haubold 1971b; Haubold, 1984; Gand, 1979;
Praszynski, 2000) shows the similariry be_rweenthe
formi almost immediately (Text-fig. 7). Using this
method TVI/ 1 superimPoses well over the general
morpholopy for I. marshilli. The polygon relative to ^L
*orihotli i; dmost completely contaiied in the Polygon of TVI/ I ; in particular the geometric concordance
point A^- D seemsto-be particularly t4i!5.*..n
cant. Tha sfight discordance relative to point E (the
middle mediil point of digit I\0 is caused by the
ts
greater robusttt.tt of TM/ I digit IV. Moreover, this
iobustness is evidenced by the gEneral morpholory of
the polygon which has substandally larger dimensions
.,r.. tho-'ughthey are geometric illy.ottipatible.
Howeier, thLre "i. accentuaied differences with
other forms of Isochirotherium.
L coltoni, the most similar in form notwithstandi.g
the much smaller size, has vertexesA and B slightly
translated in respectto TVI/ I and evidencesthe greater
lengths of digit it in.respectto III.and IV. /. soergelihas
dielt II prop6rtion.lly shorter and digit III longer than
ffil l.-1. herculis,the most robust of the group, has an
evident translation of all the points towards th-emiddle
point of dieit V (E), with a-print that is much wider
and with dfiits IV and V more developed.
In concllsion it can be affirmed that, amongst the
species of the ichnogenera
-also Isochirotherium morphousing Landmark_ analysis
logical comparison,
would seem t.r suggest, for the tracks at Termero,
attribution to a tttf,Et short robust and wide form of
chirotheriid, morphologically similar to Isochirotherium marshalli .The hlpothesis that they may rePresentlarger specimensthan those documented yP.tg
now (Halbold, l97la; Haubold, l97lb; Haubold,
1984; Peabody, 1948) relative to the ichnospecies./.
marshalli doei not seem to be confirmed due to the
difference in the metatarsal-phalangealpad V which,
t6 b. peculiar rn L
becauseof its robustnessappears
^
marshalli and due to the l^essrobust ind different
shape of the claws. The Termeno specimens would
theiefore represent a form of Isochiiotherium similar
rc I. marshilli wirh regards group
-with I-IV but.generally
a less develoPed
shorter and more robust
metatarsal pad V.
CONCLUSIONS
The discov ery of prints attributable to large chirotheriids is not common in our country.
Two prints that are generic4ly attributable to
Chirotheiiidae have been foirnd in the continental sediments of the Late Permian that outcrop in the
Bletterbachgorge(Conti et al., 1977; Conti et a1.,2000).
Very lar[e ipecimens attributable to Brachychirotherium Betirlen, 1950, have recently been found in a
Thiassiclayer of the Montemarcello Formation, q.oP^be (Sirigu & Nicosia, 1997).
^ping out iouth of Spezia
T.rcks that can
seneriZally attributed to chiisolated and still the sub-otly
rotheriids, that are
ject of study, have b9.-. documented il Upper Tiiassic
(Upp.r
^th. Carnian) tidal flat formations in various areas
Dolomites (Mietto, 2000; Leonardi 1987;
of
Conti et al., 2000).
In Middle Triassic layers only tracks attributed by
Brandner (1973) to Chirotherium sp. cf. Chirotherium
rex have been documented in the Upper Anisian
(Richthofen Conglomerate) of the Braies Dolomites.
There is more-documentation, not yet published
relative to the presenceof large Chirotheria in an area
close to the o.rf describedin ihit report. Tlacks generically attributable to Chirotheria have been found in
various stratigraphical units and at various localities.
A few kilometres north, in the area of Bad-Gfrill
(Tesimo), in a stratigraphical level corresPonding to
that of Termeno, traikJ attributable to Chirotherium
sp. cf . Chirotherium rex and a form similar to Chirotherium barthi were found (Avanzini 6{ Mietto in
prep.).
^
The ichnofauna of Termeno therefore rePresents'
from an ichnological viewpoint, the first documentation of the pr.r.ilce of large continental vertebratesin
the areas ilot. to the Dolomites during the Late
Anisian (Illyrian).
The constant presenceof continental tetrapods at
severalstratigraphical levels, which ongoing rese_arch
also evidencEsin the Morbiac Formation,- confirms
the persiste.nceof continental or at least marginalmanne environments, widely subaerial, at the culmination of sedimentation of terrigenous units (.f.
Richthofen Conglomerate) and "t1h. beginning of
the more marine conditions that led to the formation
of the first carbonate platforms (Contrin F-.) near
the Anisian-Ladinian bound ary.
An analysis of the prevalent direction measured
along digit ill of the prints on the track-bearinglevel,
ISOCHIROTHEzuUM INFERNI IN THE ILLYRIANOFADIGE VALLEY
shows a relatively well represented concentration in
the direction NE-S\7 (N40"-60"). This direction
could be related to the complex topography
that
^environwould have characterised the d.poritiorial
ment of this geographic sector during the course of
the Late Anisian. Groove casts, veqetal remains and
dragging grooves found in the imirediate overlyi.g
levels would seem to indicate the persistencein the
environment of the basal part of the Morbiac
Formation, of current flow and rransporr directions
mainly in a NE-S\7 direction (Text-fig. 8). Therefore
it may be probable that the animals walked parallel to
the bodies of water that constrained their movemenr
and distribution (Fuglewicz et al., 1990; Ptaszynksi,
2000).
ACKNO\TLEDGEMENTS
(Dijon),M. Lockl.y(Denver)and
Thanksto G. Demathieu
U. Nicosia (Rom"), f_ota critical reviewof the manuscriptand to
P.Mietto, (Padova),for useful comments.
Apart from the authors, the following personstook part in
field **.yr and research:P. Bonvecchio,T.Conci, S. Fabris,S.
Mattei, M. Moser, R. Seppi (Museo Ti'identino di Scienze
Naturali).
'We
would like to thank B. Baumgarten,Museo di Scienze
Naturali Alto Adiee - Naturmuseum Sudtirol, V. Mair and A.
Sapelza,Ufficio Giologia e Prove mareriali - Amt ftir Geologie
und BaustoffprtifungoFProvinceof BolzanolBozen,who .tr.oIragedthe study.The Eeld work wassupportedby CARG grant for
BolzanoI Bozen Province.
REFERENCES
ABEL,O., 1935, YorzeitlicheLebensspuren.
xv + 644 pp., G.
Fischer,Jena.
AvnNZINt,M., 1999, New Anisian vertebratetracks from the
Southern Alps (V"l dAdige and Valle di Non - Italy): Riv.
"E.CafFr",20:
Mus. Civ. Sc.Nat.
17-21.
-, 2000, Synaptichniumtracks with skin impressionsfrom the
Anisian (niiddle Ti'iassic)of the Southern^Alps(Val di Non,
Italy): Ichnos, 7 (4): 243-251.
-, CeoLoNI, P., Coxrl, M.A., LEONARDI,
G., Mnxxt, R.,
Mnnrott-t, N., Mtgrro, P., Ntcosn, LJ., SANTI,G. 6{
SppzznuoNTE,M., 2001, Tetrapod footprints as key elements of the Permo-Thiassic
continental biochronology:
"Natura
Bresciana",Ann. Mus. Civ. Sc. Nat., Bresc'ia,
Monografia 25: 89-107.
-s Dnr-m VECCtttR,F.M., DE Znxcup, V., GIRNOLLA,P.,
MIerro, P.,PRETo,N. & RocHr, G., 2000, Aspetti stratigrafici relativi alla presenzadi tetrapodi nelle piattaforme
carbonatichemesozoichedel Sudalpino. In Cherchi, A. 6{
Corradini, C. (eds.),Crisi biologiche,radiazioniadattivee
dinamica delle piattaformecarbonatiche:convegnodi fine
progettonazionaledi ricercaCOFIN 97: CollanaStudi Acc.
Naz. SS.LL.AA.:15-20.
- & Nr.nt,C., 1998,Improntedi tetrapodida sedimentianisici deltaValledi Non (tentino occidentale- Italia):nora preliminare:Ann. Mus. Civ. St. Nat. Ferrara,l: 5-19.
BRtRo,D., 1954, Chirotheriumlulli, a Pseudosuchian
Reprile
from New Jersey:Bulletin of the Museum of Comparitive
Zoology,Harvard College,I I I (4)t 163-192.
BnnNDNER,R., 1973, Tetrapodenfdhrtenaus der unreren
Mitteltrias der Stidalpen: Festschr.Heissel, Veroff. Univ.
Innsbruck,86: 57-7 | .
49
CunpvAN, R. E., 1990, Shapeanalysisin the study of dinosaur
morphology. In Carpenter, K. 6{ Currie, P.J. (eds.),
Dinosaur Systematics.Approaches and Perspectives.
CambridgeUniversiryPress:2l-42.
- & Bnerr-SURNAM,
M. K., 1990, Morphometricobservations
on hadrosauridornitopods. In Carpenter,K. & Currie, P.J.
(eds), Dinosaur Systematics.Approachesand Perspectives.
CambridgeUniversiryPress:163-177.
Coxrr, M.A., LEONARDT,
G., MnRrorrl, N. 6{ Nlcosn, U.,
"Val
1977, Tetrapod fooryrints of the
GardenaSandstone"
(North Italy).Their paleontological,
stratigraphicand paleoenvironmentalmeaning:Palaeontographia
Italica,70 (N.S.
4 0 ) : 9 1p p . , l l p l s .
-s -s Mlnrro, P.& Ntcostn, U., 2000, Orme di tetrapodinon
dinosauriani del Paleozoicoe Mesozoico in Italia. In
Leonardi,G. & Mietto, P.(eds.),I dinosauriitaliani.Le orme
giurassiche
dei Lavini di Marco (Tientino) e eli altri resti fosiili italiani. AccademiaEditoriale,Pisa- Roria: 495 pp.
De ZnNCHE,V., GIRNOLLA,
P.,MtEtro, P.,StoRpRES,
C. & VRtt-,
P.R.,1993,Ti'iassicsequencestratigraphyin the Dolomites
(ltaly):Mem. Sci.geol.,45: l-27.
DevnrutEu, G., I 977, La palichnologiedesVertibris. Ddveloppementrdcentet role dansla stratigraphiedu Ti'ias:Bulletin
d u B . R . G . M . ,4 ( 3 ) : 2 6 9 - 2 7 8 .
EcEnroN, 1839,On two Castsin Sandstoneof impressionof the
Hind Foot of a gigantic Cheirotherium,from^the New Red
Sandstoneof Clielhire: London Edimb. Philos. Magazine,
3rd series,14 (75): l5l-158.
Fuct-Ewrcz,R., PrnszyNsrcr,
T., 6( RDZANEK,
K., 1990, Lower
tiassic footprints from the Swietokrzyskie(Holy Cross)
Mountains,Poland:Acta Palaeont.Pol., 35: 109-164.
GnNo, G., 1979, Description de deux nouvelles traces
d'Isochirotheriumobservdesdans le grbs du Ti'ias moyen de
Bourgogne:Bulletin de la Socidti I'Hirtoire Nat,rr.ll. d,t
Creusot, 37: 13.
GTRNoLLR,
P., De ZnNcuE, V. & Mrerro, P., 1998, Ti'iassic
SequenceStratigraphy in the Southern Alpr (Northern
Ita[y). Definition oT sequences
and basin evolution. In De
Graciansky,P. C., Hardenbol,J., J".quin, T. &Vail, P. R.
(eds.), Mesozoic and Cenozoic SequenceStratigraphyof
EuropeanBasins:SEPM sp. pubbl. 60: 721-747.
HnueoLD,H.,1967, Eine PseudosuchierFdhrtenfauna
ausdem
Buntsandstein Stidthtiringens: Hall. Jahrbuch Mitteldt.
Erdg.,8 (1966): 12-48.
-, l97la, Ichnia Amphibiorum et Reptiliorum fossilium.In
Kuhn, O. (.d.), Handbuch der Palieoherpetologie,
18: l124, G. Fischer.,Stuttgart.
-,
197lb, Die tetrapodenfhhrtendes Buntsandsteins
in der
'Westdeutsdeutschen Demokratische Republik und in
chland um ihre aequivalent-ein der gesamten tias:
Palaontologische
Abhandlungen,4, 3; 395-660.
-,
1984, Saurierfdhrten. Die Neue Brehm-Bucherei,A.
Ziemsen,'W'ittenberg.
Lutherstadt:231 pp.
KARL,K., & HnueoLD, H., 1998, Bracbychirotherium
atrsdem
Coburger sandstein (Mittlerer Keuper, Karn/Nor) in
Nordbiyern: Hallesches
Jahrb.Geowiss.,820' 33-58.
KnuR J.1., 1835, Uber TierFirhrtenbei Hildburghausen:
N. Jb.
Min. Geol.Pal. 1835:327-328.
LEoNRRol,G., (.d.) 1988, Glossaryand Manual of Tetrapod
Footprint Palaeoichnology.Departamento Nacional de
Produgio Mineral, Brasflia:ll7 pp.
MntnvELL, F.T., l9l l, Notes on footprints from the Keuper of
RuncornHill: Proc.Geol. Soc.Liverpool,I l: 140-152.
Mterro, P., 2000, La SezioneGeologicadel Museo della Val
Fiorentina.In Museo di Selvadi Cidore. Storia,Archeologia
e Geologiadella Val Fiorentina,Cortina d'Ampezzo: 7 -29.
M. AVANZINI, G. LEONARDI
50
Nopcse, F., 1923,Die Familiender Reptilien:Fortscht Geol.
Pdeont.,2: l-210, Berlin.
from
Pnnnooy,F.E., 1948, Reptile and 4pphibian Thachn'ays
the l.ower Thiassic'Moenkopiformation of fuizona and
295468.
Utah: Bull. Dept. Geol. Sci.,-2718:
nel terriSnrcu, I. 6cNrcosu' IJ., 1994,Pistedi rettili triassici
'Giovanni
torio della Spezia:Mern. Accad. Lunig. Sc.
Capellini", 64-652251-256.
Ilraszn rsKy,T, 2000, lower Thiassicvertebratefootprints from
\7iory Holy ClossMountains,Poland:Acta Paleont.Pol.,
45Q):r5r-r94.
TnneslsE,G. 6c SenlEAI{T,'WA.S,1997,The Tiacksof Thiassic
FossilEvidencefrom Nonh-\flest England.The
Vertebrates.
SationeryOffice,Iondon:204 pP.
(nanusnipt rccciacdI"h 30, 2001
acccpudJanury I 7, 2002)
Marco AveNzNl
Museo Thidentino di ScienzeNanrrdi
Via Calepina 14,38100 Thento,Itdy
e-m-ai[ aaataini@ntn. trr.it
Gitrseppe LroNanol
Museo Thidentino di ScienzeNaturali
Present address:Via Modielieni 2
80070 Monterusciello (Nafili), Italy