- Email: [email protected]
New dinosaur ichnotaxa from the Early Jurassic of the Holy Cross Mountains, Poland
Palaeogeography. Palaeoclimatology, Palaeoecology, 85 (1991): 137-148
137
Elsevier Science Publishers B.V., Amsterdam
New dinosaur ichnotaxa from the Early Jurassic of the Holy Cross Mountains, Poland G e r a r d Gierlifiski
Ul. Jasielska 38/42 m.89, 02-128 Warsaw,Poland (Received September 15, 1988; revised and accepted October 25, 1990) ABSTRACT Gierlifiski, G., 1991. New dinosaur ichnotaxa from the Early Jurassic of the Holy Cross Mountains, Poland. Palaeogeogr., Palaeoclimatol., Palaeoecol., 85: 137-148. The numerous ornithopod tracks discovered during the last four years, in the Holy Cross Mts., allow the identification of two new ichnospecies--Anomoepuspienkovskii ichnosp, nov. and Moyenisauropuskaraszevskiiichnosp, nov. The moyenisauropodid structure indicates that the trackmakers may well have been hitherto unknown forms of the earliest iguanodontids. Newly discovered theropod footprints, Grallator (Grallator)zvierzi ichnosp, nov. and Grallator (Eubrontes) soltykovensis ichnosp, nov., supplement knowledge of the Early Jurassic terrestial fauna of Poland. The majority of the tracks occurring in the Gliniany Las exposure (Upper Hettangian) were left by dinosaurs which occupied the margins of a lagoonal environment. The other footprints, discovered in the Sottyk6w (Lower Hettangian) outcrop, were left by carnosaurs walking across the fluvial plain.
Introduction All the Polish dinosaur footprints were discovered on the northern slope of the Holy Cross Mts. These tracks have been found in the Gliniany Las and Zapni6w exposures of the Przysucha Orebearing Series (Upper Hettangian) and in the Sottyk6w outcrop of the Zagaje Series (Lower Hettangian). The first two footprints were found by Karaszewski in the Gliniany Las quarry, in 1959. He proceeded to describe them respectively as "tetradactyl track" (Karaszewski, 1969) and as Neochirotherium jurassicum (Karaszewski, 1975). Later, these specimens, and other similar footprints collected in 1983-1984, were identified as Moyenisauropus sp. (Gierlifiski and Potemska, 1987). The discovery of two new tracksites at Sottyk6w and Zapni6w (Piefikowski and Gierlifiski, 1987), and the numerous new specimens collected since 1986, have supplemented our knowledge of Hettangian dinosaurs in the Holy Cross Mountains area of Poland. 0031-0182/91/$03.50
In 1986, Anomoepus sp. was found in the uppermost part of the Sottyk6w outcrop (Pieflkowski and Gierlifiski, 1987). Further, Piefikowski and Gierlifiski (1987) described Moyenisauropus sp. from the Zapni6w outcrop. In 1988, Grallator (Eubrontes) soltykovensis ichnosp, nov. was discovered in the lowermost part of the Sottyk6w outcrop. Moreover, the exploration of the Gliniany Las quarry during 1988 and 1989 revealed the theropod footprints Grallator (Eubrontes) sp. (Gierlifiski, 1990) and Grallator (Grallator) zvierzi ichnosp, nov. Newly discovered ornithopod tracks in Gliniany Las allow me to establish two new ichnospecies Anomoepus pienkovskii ichnosp, nov. and Moyenisauropus karaszevskii ichnosp, nov.
lchnosystematic descriptions Suborder THEROPODA Marsh 1881 Ichnofamily GRALLATORIDAE Lull 1904 Ichnogenus Grallator Hitchcock 1858 Subichnogenus Grallator (Grallator) Olsen and Galton 1984
© 1991 - - Elsevier Science Publishers B.V.
138
Grallator (Grallator) zvierzi nov. sp. (Fig. 1) Holotype: Trackway preserved as a natural cast on one layer and as undertracks on another layer. The specimen preserved as a natural cast is owned by the author and the specimen preserved as a couple of undertracks is lodged in MGIW (Museum of the Geological Institute in Warsaw, Poland), as 1560.11.9. Paratype: Single footprint preserved as a natural cast. Original specimen owned by the author and its plaster cast lodged in MGIW, as 1560.11.13. Type horizon: Przysucha Ore-bearing Series (Upper Hettangian). Type locality: Gliniany Las, Holy Cross Mts., Poland. Etymology: Named in honor of Jadwiga Zwierz, head of the Museum of Geological Institute in Warsaw, who supported my research during the last few years. Diagnosis: Bipedal. Tridactyl pes with digits II-IV impressed. In the trackway, pes directed forward. Ratio of the pes length to pace is 1 : 3. The pes is wider than long by an average of 5%. Digit III is longest. The angle between digits II and IV varies from 18° to 37 °. Description: The paratype differs from the holotype by having the third digit relatively shorter in comparison to the lateral digits and by having angles beween the digits about twice as large. Other specimens, not described here (1560.II.14), 1560.11.15, 1560.11.16, in MGIW), demonstrate transitional features between the holotype and paratype; therefore, the length of digit II is 48-62% of the length of digit III, while the length of digit IV is 57-84% of that of digit III. The angles between the digit axes are: I I - I I I = 10-20 °, I l l - I V = 7 - 1 7 °. The pes length varies from 9 cm to 12.6 cm. Comparison: Among the smaller grallatorids with comparably divaricate digits, the most similar ichnites are: Grallator (Grallator) cuneatus and G. (G.) tenuis Hitchcock 1858; Grallator (Grallator) gracilis Hitchcock 1865; Grallator (Grallator) eloensis and G. (G.) variabilis Lapparent and Montenat 1967; several footprints from Lesotho reassigned to Grallator (Grallator) by Olsen and Galton (1984); Grallator (Grallator) sp. Conrad,
G. GIERLINSKI
Lockley and Prince 1987; Grallator (Grallator) ssatoi Zhen, Li, Rao, Mateer and Lockley 1989; Grallator sp. Hunt, Lucas and Kietzke 1989. However, Grallator (Grallator) zvierzi ichnosp, nov. is broader than long, whereas the ichnites mentioned above are always longer than broad. Discussion: Olsen and Galton (1984) considered the names Eubrontes and Anchisauripus as synonymous with Grallator. They used the subichnogeneric designations to denote grallatorid tracks in the size categories. Consequently, the size and morphology of those footprints assigns them to the subichnogenus Grallator ( Grallator). Subichnogenus Grallator (Eubrontes) Olsen and Galton 1984
Grallator (Eubrontes) soltykovensis nov. sp. (Fig.2) Holotype: Single footprint preserved as a natural cast. Original specimen owned by the author and its piaster cast lodged in MGIW, as 1560.11.12. Paratype: Single footprint preserved as a natural cast. Original specimen owned by the author and its plaster cast lodged in MGIW, as 1560.11.10. Type horizon: Zagaje Series (Lower Hettangian). Type locality: Sottykrw, Holy Cross Mts., Poland. Etymology: Named after the type locality, Sottyk6w. Diagnosis: Bipedal. Tridactyl large pes with digits II-IV impressed. Metatarsal-phalangeal pads of digits III and IV made an isolated oblong posterior impression. The third digit is longest. The angle between digits II and IV varies from 50 ° to 60 °. The angle between digits IV and III is always larger than that one between digits III and II. Description: The length of holotype is 22.5cm, whereas the paratype is 28 cm long. Moreover, the paratype has a larger angle between digits III and IV and a smaller angle between digits II and III than in the holotype. The paratype is very deeply impressed, inclined forward and sideward by 6 °, opposite to shallow and equally impressed holotype. The paratype's digits are thickly impressed and the third digit is curved; most probably, this was caused by a non-taxonomic factor, probably resulting from the trackmaker's mode of locomotion and the nature of the sediment.
139
N E W D I N O S A U R I C H N O T A X A F R O M T H E EARLY J U R A S S I C O F T H E HOLY C R O S S M O U N T A I N S , P O L A N D
~." ,v
."~:~.
':,
,
~:-:!~ .,~,.,~t~-~. ..." ;:"~
I'.,::."::.~.~ .....
"z. :".('. ""-':.':-.
".".,.,.~'v'.: ", ~ "'.."':"~.. :
°~o.
, •
.
:
a
10 cm I
I
:i;.......':!'~i!:~'~ ;~]'.. • ...~
~:i~~
~v~,,~ ~: ,
C".
....:......
":~
i~-:., :.,.
"." ".':'~'~ ,
".:.
.~..
_ •
"" ;" ""'
. ::'.
"-~
...~....~-~;.!.....,"~...,"..'.......'~
b
Fig. 1. Grallator (Grallator) zvierzi ichnosp, nov., natural casts from Gliniany Las (Upper Hettangian). a. Trackway designated as holotype, b. Paratype.
140
G. GIERLINSKI
~-:,-.. ;,~.
"~::'P"
.:~,
. :;I
.~ '" " ~
~":'..~
~ 4." ~,
'.:...'~~,~. L- :~.~ x4<.t :~ %:"~'!:":7~.-.';.':'";~'.'.~. :~,~ ;:~ "";:'" ~::~i!& .... $:. ~' ::'~7. "s,~s'~.~' :~' ~.. "..'7~ "!:~,,:.' ..... ;..f;' ..f..i" .~' ~--'-.; : :.,.', ... ":;; ~.:~9. .;....-..~ ..7~ "~.." ' ;.. q.~-"..
.. • "
~t[..-~
.Jcm
, :.:.~ ]'~'::
~7!::?~:~7;;.: ~.:" .".'.:'4' •~"., .. ".:'~"':;;
l:.:
• :: ..u
...~::'~,~ ~".o...'
.'~ "'.~.
N}! i;'::: 7 ~:".' :. ~ ts',..
."!
~.::'~'.'~.-':.'~
.'."
:,,.~o
..,sc~e;$'>'~.~<..J
--,~.
N'::':-!
. ....:.,:4
• ',~ ~:.:.....- .~::.:.,~.........:.!.;,:!......¢~7. ..:...:.;'-:...'.!~ ~ ..........~ .:¢,.~.',.'-,~ .:. :., "~:~:'.;,:;.. " •~.~,.:.,..->~. .....~-..,...,:....::::..!i~
-K¢~d."~ "'"
' ~ , 7 ° '~ ' ; . . . .
IV .~.,..W
..
.
•
.,.;° . # , ~ i , . , . : ~ r
'~'~" ,~4.:'.'*, :"-,":.,.;;2.;,~,~ ~ b
Fig.2.
Grallator
(Eubrontes)
soltykovensis
" ...~...~ ~~.~L "." ......~:. ;..~.~;:':;.,.:.. : "":.;':::;.,.,-
ichnosp, nov,, n a t u r a l casts f r o m S o i t y k 6 w ( L o w e r H e t t a n g i a n ) . a. H o l o t y p e . b. P a r a t y p e .
Phalangeal pads are visible on digits II and III (especially in the holotype). In digit II, imprints of two phalangeal pads are present and three phalangeal pads are in digit III. The length of digit II is 54-57% of the length of digit III, while the length of digit IV is 70-76% of that of digit III. Angles between digits are: I I - I I I = 15-25 °, IIIIV = 30-40 o. Comparison: The large size of these footprints combined with their grallatorid structure assigns
them to the ichnotaxon Grallator (Eubrontes), following Olsen and Galton (1984). Grallator (Eubrontes) soltykovensis ichnosp, nov. has highly divaricated digits. Among Liassic, large theropod tracks, this bears a resemblance to: Grallator (Eubrontes) divaricatus (Hitchcock 1865) Lull 1904; Kayentapus and Dilophosauripus Welles 1971; Neotrisauropus, Plastisauropus and Kainotrisauropus, considered to be synonyms of Grallator by Olsen and Galton 1984; Schizograllator Zhen, Li
NEW DINOSAUR ICHNOTAXA FROM THE EARLY JURASSIC OF THE HOLY CROSS MOUNTAINS, POLAND
and Rao 1986. However, Grallator (Eubrontes) divaricatus, Dilophosauripus and Schizograllator differ from G. (E.) soltykovensis by having the third digit relatively shorter in comparison to the lateral digits. On the other hand, G. (E.) soltykovensis differs from Kayentapus, Neotrisauropus, Plastisauropus and Kainotrisauropus, in that digit II originates further posteriorly. Discussion: The numeration of the digits of single tridactyl tracks from the Liassic poses no problems, if the phalangeal pads are visible. The pedal digit II has always two phalangeal pads, while the pedal digit IV has three or four phalangeal pads. Suborder ORNITHOPODAMarsh 1878 Ichnofamily ANOMOEPODIDAELull 1904, emendedherein Included ichnogenera: Anomoepus Hitchcock 1848, Apatichnus Hitchcock 1858, Jialingpus Zhen, Li and Zhen 1983
Occurrence: Lower Jurassic to Upper Jurassic of North America, Europe, southern Africa and southeastern Asia. Emended diagnosis: Normally bipedal, but sometimes also quadrupedal. Pes functionally tridactyl. Hallux rarely impressed. Pedal digits II, III and IV tend to be subequal in length. In the trackway, the pes is directed forward. Manus imprints are four-toed or five-toed. Comparison: Anomoepodidae Lull 1904, emend. herein show the transitional morphology between Atreipodidae Olsen and Baird 1986 and Moyenisauropodidae Ellenberger 1974, emended herein. In Anomoepodidae, the pedal digits II, III and IV tend to be subequal in length (Olsen and Galton 1984), the g r a l l a t o r i d - like pes of the Atreipodidae has digit III clearly as longest, while in the Moyenisauropodidae, the pedal digit IV is always the longest one. Moreover, the manus of the Atreipodidae is tridactyl or tetradactyl, the Anomoepodidae have tetradactyl or pentadactyl manus, whereas the manus of the Moyenisauropodidae is pentadactyl. Ichnogenus Anomoepus Hitchcock 1848, emended herein.
141
1848 AnomoepusHitchcock,p. 220 1858 AnornoepusHitchcock, p. 55 1904 AnomoepusHitchcock. Lull, p. 500 1915 AnomoepusHitchcock. Lull, p. 208 1953 AnomoepusHitcbcock.Lull, p. 192 1971 AnomoepusHitchcock. Haubold, p. 83 1984 AnomoepusHitchcock. Olsen and Galton, p. 99 Type ichnospecies: Anomoepus scambus Hitchcock 1848, p. 222, pl. 13, figs.l-6, from the Portland Formation (Upper Liassic) of Massachusetts, USA. Included ichnospecies: Anomoepus scambus Hitchcock 1848, A. intermedius, At curvatus, A. minimus Hitchcock 1865, A. cuneatus, A. crassus, A. isodactylus Hitchcock 1889, A. gracillimus Hitchcock 1844, A. ranivorus Ellenberger 1970, A. natatilis, A. dodai, A. vermivorous, A. longicauda, A. minor Ellenberger 1974 and A. pienkovskii ichnosp, nov.
Occurrence: East Berlin Formation (Sinemurian) and Portland Formation (Pliensbachian to Toarcian) of Connecticut, Massachusetts and New Jersey, USA. Upper Red Beds (Lower Hettangian) of Lesotho, southern Africa. Zagaje Series (Lower Hettangian) and Przysucha Ore-bearing Series (Upper Hettangian) of the northern slope of the Holy Cross Mts., Poland. Emended diagnosis: Normally bipedal, but sometimes quadrupedal also. Pes with three functional digits. Hallux rarely imprinted. Pedal digits II, III and IV usually tend to be subequal in length, but sometimes digit IV is clearly the longest one. In the trackway, the pes is directed forward or inward. Metatarsal impression occurs when the trackmaker rests. The manus is tetradactyl or pentadactyl. Manual digits II and III are longest. Comparisons: In Anomoepus, the angle between pedal digits III and IV in comparison to that one between digits II and III, is smaller than in Apatichnus. Metatarsal Anomoepus tracks evidently occur as resting traces only, whereas metatarsal tracks of Jialingpus were made during active locomotion. Anomoepus pienkovskii nov. sp. (Fig.3) Holotype: Trackway preserved as a natural cast. Original specimen owned by the author and its plaster cast lodged in MGIW, as 1560.I1.20.
142
G. GIERLINSKI
IV )."
..,/
.....
:
.t
10 cm ~
..:,:....
Fig.3. Holotypeof Anomoepuspienkovskiiichnosp, nov., natural cast of the trackwayfrom GlinianyLas (Upper Hettangian).
Type horizon: Przysucha Ore-bearing Series (Upper Hettangian).
Type locality: Gliniany Las, Holy Cross Mts., Poland.
Etymology: Named in tribute to Dr. Grzegorz Piefikowski, my co-worker in the studies on footprints from the Holy Cross Mts., who has made sedimentological interpretations of the Liassic deposits in Poland. Diagnosis: Quadrupedal in gait. Pes functionally tridactyl with hallux sometimes also imprinted.
The fourth pedal digit is longest. In the trackway, pes and manus are directed forward, with the manus situated anteriorly and slightly outward in comparison to the pes. Ratio of pes length to pes pace 1:2.4. Ratio of manus length to manus pace 1:5.4. The angle between pedal digits II and IV varies from 48 ° to 58 °. Manus tetradactyl, half the size of the pes and wider than long by an average of 33%. Manual digits II and III are longest. Description: Holotype length: pes = 14 cm, m a n u s = 7 cm. The length of pedal digit II averages
143
NEW DINOSAUR ICHNOTAXA FROM THE EARLY JURASSIC OF THE HOLY CROSS MOUNTAINS. POLAND
77% of the length of digit IV, while the length of digit III averages 84% of that of digit IV. Angles between the pedal digits: I - I I = 8 0 °, I I - I I I = 30-32 °, Ill-IV = 20-27 °. The manus with digits IIV impressed. Manual digits II and III are longest followed in length by digits I and IV. Angles between the manual digits: I-II = 54-57 °, II-III = 57-59 °, Ill-IV = 65-75 °. Comparisons: The manus is only half the size of the pes. In the trackway, the manus is situated slightly outward to the pes impression, whereas in other ichnospecies of Anomoepus the manus imprints are only one-third the size of the pes imprints, the manus imprints lying closer to the midline of the trackway. lchnofamily MOYENISAUROPODIDAEEllenberger 1974, emended herein Included ichnogenus: Moyenisauropus Ellenberger 1974, emended herein
Occurrence: Lower Jurassic of southern Africa and Europe. Emended diagnosis: Normally bipedal, but sometimes also quadrupedal. Pes with three functional digits. Hallux rarely imprinted. Pedal digits massive, short and highly divaricated, with imprints of two phalangeal pads on the third digit. The fourth pedal digit is longest. In the trackway, the pes is directed inward. Metatarsal impression occurs when the trackmaker rests. The pentadactyl manus has short and thick digits. Comparison: The pes of Moyenisauropodidae EIlenberger 1974 emend, herein shows two phalangeal pads on digit III, whereas, the pedal digit III of Anomoepodidae has three phalangeal pads and iguanodontid tracks (e.g., Caririchnium Leonardi 1984) have only one phalangeal pad in that digit. The remaining features of Moyenisauropodidae make this ichnofamily closer to iguanodontid footprints than to the ichnofamily Anomoepodidae. The pes and manus imprints of the Iguanodontidae and Moyenisauropodidae differ from those of the Anomoepodidae in having shorter and thicker digits. In the moyenisauropodid and iguanodontid trackway the pes was clearly directed inward,
whereas in the anomoepodid trackway the pes was usually turned forward. Ichnogenus Moyenisauropus emended herein
Ellenberger
1974,
1974 MoyenisauropusEllenberger, p. 25 1984 AnomoepusHitchcock. Olsen and Galton, p. 99 1987 Moyenisauropus Ellenberger. Gierliflski and Potemska, p. 109 Type ichnospecies: Moyenisauropus natator EIlenberger 1974, p. 26, pls. C and D, from the Upper Red Beds (Lower Hettangian) of Lesotho, southern Africa. Included ichnospecies: Moyenisauropus natator, M. levicauda Ellenberger 1974 and M. karaszevskii ichnosp, nov.
Occurrence: Upper Red Beds (Lower Hettangian) of Lesotho, southern Africa. Przysucha Ore-bearing Series (Upper Hettangian) of the northern slope of the Holy Cross Mts., Poland. Emended diagnosis: Normally bipedal, but sometimes also quadrupedal. Pes functionally tridactyl. Hallux rarely imprinted. Pedal digits massive, short and highly divaricated. Two phalangeal pads are present in the pedal digit III. The pedal digit IV is longest. In the trackway, the pes is directed inward. Metatarsal impression occurs when the trackmaker rests. In the pes imprints, which were left during active locomotion, the angle between digits II and IIl is always larger than that one between digits III and IV. The pentadactyl manus has short and thick digits. Manual digits are subequal in length. Comparison: Moyenisauropus is distinguished from other tridactylous ichnogenera from the Liassic in having only two phalangeal pads in the pedal digit IIl. Moreover, in contrast to Anomoepus, the manual digits of Moyenisauropus are subequal in length. In Moyenisauropus the angle between pedal digits II and III in comparison to that one between digits III and IV, is usually larger than in the pes of Anomoepus. Discussion: Ellenberger (1974) erected the ichnogenus Moyenisauropus to comprise the ichno-
144
species M. natator, M. natatilis, M. dodai, M. vermivorous, M. longicauda, M. minor and M. levicauda. In addition Ellenberger proposed their placement into the ichnofamily Moyenisauropodidae. Olsen and Galton (1984) and Haubold (1984, 1986) treated Moyenisauropus as a junior synonym of Anomoepus. However, Gierlifiski and Petemska (1987) proposed to retain Moyenisauropus as a separate ichnogenus in the ichnofamily Anomoepodidae. In fact, Moyenisauropus natatilis, M. dodai, M. vermivorous, M. longicauda and M. minor all bear a resemblance to Amomoepus. In contrast, Moyenisauropus natator, M. levicauda and M. karaszevskii ichnosp, nov. do not correspond in morphology to the Anomoepodidae. Consequently, I propose to transfer Moyenisauropus natatilis, M. dodai, M. vermivorous, M. longicauda and M. minor to the ichnogenus Anomoepus and to retain M. natator, M. levicauda and M. karaszevskii ichnosp, nov. as Moyenisauropus, in their own ichnofamily Moyenisauropodidae.
Moyenisauropus karaszevskii nov. sp. (Fig.4) Holotype: Trackway preserved as a natural cast on one layer and as undertracks on another layer. The specimen preserved as a natural cast is owned by the author and the specimen preserved as a couple of undertracks is lodged in MGIW, as 1560.11.9. Type horizon: Przysucha Ore-bearing Series (Upper Hettangian). Type locality: Gliniany Las, Holy Cross Mts., Poland. Etymology: Named in honor of Dr. Wtadystaw Karaszewski, the discoverer of the first dinosaur footprints in Poland. Diagnosis: Bipedal in gait. Tridactyl pes, with hallux sometimes also imprinted. Two phalangeal pads are present in the pedal digit III. The pedal digit IV is longest. In the trackway, the pes is directed inward. Ratio of pes length to stride 1:3.8. Ratio of pes length to pace 1:2.3. Pace angulation 103°. The angle between pedal digits II and III is always larger than that one between
G. GIERLII~SKI
digits III and IV. The angle between pedal digits II and IV varies from 50 ° to 56 °. Description: The pes length varies from 22 to 26 cm. The length of digit II averages 67% of the length of digit IV, while the length of digit III averages 84% of that of digit IV. The angles between the digit axes are: I - I I = 6 5 °, I I - I I I = 30-36 °, I I I - I V = 17-22 °. The manus imprint associated with the pes has not been found. Comparison: The pedal digit II of this new ichnospecies originates further posteriorly than in the pes of M. natator. Moyenisauropus levicauda differs from M. karaszevskii ichnosp, nov. by having the pedal digit III relatively longer in comparison to the pedal digit IV.
Moyenisauropus sp. (Fig.5) Material: Original specimen owned by the author and its plaster cast lodged in MGIW, as 1560.11.8. It is a single manus imprint preserved as natural cast, from Gliniany Las, Holy Cross Mts., Poland, (Przysucha Ore-beating Series; Lower Hettangian). Description: Pentadactyl manus, wider than long by 48.5% (16.5 cm width and 8.5 cm long). Digits short, massive and subequal in length. The first digit is thinnest; more markedly bent away from the others; ends in a long, straight claw. The tips of the other four digits are rounded. Angles between the digits: I-II--62 °, I I - I I I = 60 °, III-IV = 32 °, IV-V = 42 °. Comparison: The features mentioned above clearly differ from those of the Anomoepodidae and are identical to that of the Moyenisauropus manus. Consequently, in contrast to the manual digits of Anomoepus, digits of this ichnite are subequal in length. Moreover, our specimen has shorter and thicker digits than are typical of the Anomoepodidae. Discussion: Because of the lack of material where the manus would occur together with the pes, we cannot with entire certainty identify it as M. karaszevskii, an ichnospecies created on the basis of the analysis of the pes. According to the structure of the moyenisauropodid manus, the differences between the digits make the numeration of the digits possible. Digits II, III, IV and V are similar in width, whereas
145
N E W D I N O S A U R I C H N O T A X A F R O M T H E EARLY J U R A S S I C O F T H E HOLY C R O S S M O U N T A I N S , P O L A N D
IV • .:;:~!~
~.
~...'.::.
~:"
-:.q :.::..~,..-....~ ,#
,:.~.?".:. .... •:.':."./?~)i~ . . :.~,?q~:,:-- : .: •
:,.,...... " . , : ~
~ ,::::i:!! III.................. :
:{.i....
i:.,...':........... ."-
lOcm ..~::~
"t ..... :
..° ....,
:'-...:
.. ......
"":::~....
)
~
IV
Fig.4. H o l o t y p e o f Moyenisauropus karaszevskii ichnosp, nov., natural cast o f the t r a c k w a y from G l i n i a n y Las ( U p p e r Hettangian).
146
G. GIERLIIqSKI
5 cm
:~.-,..~'?...~ ~,'~," ." : 4 .-". . . .-. ! "., "~: '~.., .~ L~'.:;' .Z:,,:~'.":.-..-':~"" ~,..~,,,~
V ~,~r÷':".~,.-" l ~:~..'.."'" y'::':.:-...".:":.~"'.";';'z.:~ ::'::~ :.'-:....-~?~--"~::";<,;t'.:~,~
~@~
Fig.5. Moyenisauropussp., natural cast of the single manus ichnite from Gliniany Las (Upper Hettangian). a. Whole specimen, b. Pollex with a spike impressed at the end. digit I is thinner and is more markedly bent away from the others. Systematic
comparisons
The grallatorid structure is characterized by the third pedal digit being always longer than the lateral digits (Baird 1957; Lapparent and Montenat, 1967; Olsen 1980). Among Liassic dinosaurs, this bears a resemblance to the pes of the Podokesauridae and of Megalosauridae. Consequently, Grallator (Grallator) might have been imprinted either by podokesaurids (Lull, 1953; Colbert 1963) or by a juvenile megalosaurs, whereas Grallator (Eubrontes) imprints were probably made exclusively by megalosaurs like Dilophosaurus. Anomoepus is conventionally considered as the footprints of Fabrosauridae, previously classed as Hypsilophodontidae (Lull, 1953; Haubold, 1984; Olsen and Baird, 1986). The differences between Anomoepus and the tracks here referred to Moyenisauropodidae have allowed me to ascertain that their trackmakers belong to different dinosaur families.
In the trackways of Moyenisauropus natator described by Ellenberger 1974 and in M. karaszevskfi ichnosp, nov. the pes turned inward, whereas the Anomoepus pes was usually directed forward. The inward rotation of the pes is indicative of large ornithopods such as iguanodontids and hadrosaurids (Lockley 1987). Moreover, the size and structure of Moyenisauropus ichnites suggest that their trackmakers were larger and more massive than were the Fabrosauridae. The pes prints of Anomoepus are 5 cm up to 16 cm long, whereas the pes length of Moyenisauropus varies from 10 cm to 26 cm. The short, thick and highly divaricated pedal digits of Moyenisauropodidae are closest to iguanodontid footprints. Moreover, the moyenisauropodid manus shows less differentiation in the digit lengths than in the manus of Heterodontosauridae or Fabrosauridae. It is further noteworthy that Moyenisauropus manus has a clearly offset, iguanodont-like pollex. Sometimes, a straight spike is impressed at the end of it (Fig.5b), in contrast to the hooked pollex claw of the prosauropod manus, as in Navahopus (Baird 1980).
NEW DINOSAURICHNOTAXAFROM THE EARLYJURASSICOF THE HOLYCROSSMOUNTAINS,POLAND
Consequently, the features described above confirm that Moyenisauropodidae represent footprints of some hitherto unknown early members of the family Iguanodontidae. This contributes to our knowledge of the early evolution of iguanodontids and indicates that this family appeared much earlier than has been indicated by their osteological remains, near the boundary between the Triassic and Jurassic. Paleoenvironmental and paleoecological remarks From knowledge of the profile of the Gliniany Las outcrop (Piefikowski 1985, p. 47) and my investigations performed during the last two years, the footprints were left in certain mudstone layer of lagoonal origin, which were superimposed upon barrier deposits. The tracks occur either as natural casts on the bottom surface of the superimposed sandstone layer or as undertracks preserved on the top of another sandstone layer underlying the lagoonal mudstone. The small and shallow lagoon, with a strong influx of fresh water, was situated between a barrier system separating it from the shallow brackish sea and a deltaic-fluvial area (Piefikowski 1985). It appears likely that the dinosaurs from Gliniany Las found quite favourable living conditions in the barrier - - lagoon transitional environment. Up to this moment, the theropod tracks form 11% of the number of all the footprints discovered in this outcrop. The rest of the footprints belong to ornithopods. In the lowermost part of the Sottyk6w outcrop only the carnosaur tracks, here described as Grallator (Eubrontes) soltykovensis ichnosp, nov., have been found. Those footprints were imprinted into the muddy fluvial plain deposits and preserved at the bottoms of superimposed sandy layers deposited by fluvial processes - presumably crevasse splays (Piefikowski, 1981, unpublished; Piefikowski and Gierlifiski 1987). Acknowledgements I would like to thank my wife for her help in preparing this paper. I am deeply indebted to Professor Dr. William A. S. Sarjeant (University of Saskatchewan, Canada) for his valuable corn-
147
ments and suggestions. I also thank Dr. G. Pieflkowski (Geological Survey of Poland) for his helpful remarks. I am grateful to my friends: A. Piotrowski, Z. W6jcik, T. Jarzabek, W. Knapczyk and P. Kabulski for field assistance. J. Zwierz and P. Nowacki offered their help during my researches. My special thanks go to A. Potemska for her help in preparing this paper, as well as for her collaboration during the last few years in the study of Polish dinosaurs. Her help was invaluable to me, both as co-worker and as a great friend. References Baird, D., 1957. Triassic reptile footprint faunules from Milford, New Jersey. Mus. Comp. Zool. (Harvard Univ.), Bull., 117: 449-520. Baird, D., 1980. A prosauropod dinosaur trackway from the Navajo Sandstone (Lower Jurassic) of Arizona. In: L. L. Jacobs (Editor), Aspect of Vertebrate History. Mus. Northern Ariz. Press, pp. 219-230. Colbert, E. H. 1963. Fossils of the Connecticut Valley: The age of dinosaurs begins. State Geol. Nat. Hist. Surv. Conn. Bull., 96, 31 pp. Conrad, K., Lockley, M. G. and Prince, N. K., 1987. Triassic and Jurassic vertebrate-dominated trace fossil assemblages of the Cimarron valley region: implications for paleoecology and biostratigraphy. In: S. G. Lucas and A. P. Hunt (Editors), New Mexico Geol. Soc. Guidebook, 38th Field Conf. Univ. New Mexico Press, pp. 127-138. Ellenberger, P., 1970. Les niveaux paleontologiques de premi6re apparition des Mammif6res Primordiaux en Afrique du Sud et leur Ichnologie: Establissement de zones stratigraphiques detaill6es dans le Stormberg du Lesotho (Afrique du Sud), (Trias Superieur fi Jurassique). In: S. H. Haughton (Editor), I.U.G.S., 2nd Symposium on Gondwana Stratigraphy and Palaeontology, Pretoria, pp. 343-370. Ellenberger, P., 1974. Contribution ~, la classification des pistes de Vertebres du Trias: Les types du Stormberg d'Afrique du sud (2 6me Partie: Le Stormberg Superieur. I. Le biome de la zone B/1 au niveau de Meyeni: ses bioc6noses). Palaeovertebrata M+me. extraordinaire, 141 pp. Gierlifiski, G. and Potemska, A., 1987. Lower Jurassic dinosaur footprints from Gliniany Las, northern slope of the Holy Cross Mountains, Poland. Neues Jahrb. Geol. Pal~iontol. Abh., 175: 107-120. Gierlifiski, G., 1990. First find of a carnosaur footprint in the Lower Jurassic deposits of Gliniany Las, Holy Cross Mts. Przeg. Geol., 7-8: 315-317. Haubold, H., 1971. Ichnia Amphibiorum et Reptiliorum fossilium. In: O. Kuhn (Editor), Encyclopedia of Palaeoherpetology part 18. Fischer, Stuttgart, 124pp. Haubold, H., 1984. Saurierffihrten, (Neue Brehm-Bficherei, 479) Ziemsen, Wittenberg, 231 pp. Haubold, H., 1986. Archosaur footprints at the terrestial
148 Triassic-Jurassic transition. In: K. Padian (Editor), The Beginnings of the Age of Dinosaurs: Faunal change across the Triassic-Jurassic boundary. Cambridge Univ. Press, pp. 189-201. Hitchcock, C. H., 1889. Recent progress in ichnology. Proc. Boston Soc. Nat. Hist., 24- 117-127. Hitchcock, E., 1844. Report on ichnolithology, or fossil footmarks, with a description of several new species, and the coprolites of birds, from the valley of Connecticut river, and of a supposed footmarks from the valley of Hudson river. Am. J. Sci., 47: 292-322. Hitchcock, E., 1848. An attempt to discriminate and describe the animals that made the fossil footmarks of the United States, and especially New England. Mere. Am. Acad. Arts. Sci., 3(2): 129-256. Hitchcock, E., 1858. Ichnology of New England. A report on the sandstone of the Connecticut Valley, especially its fossil footmarks. White, Boston, 220 pp. Hunt, A. P., Lucas, S. G. and Kietzke, K. K., 1989. Dinosaur footprints from the Redonda Member of the Chinle Formation (Upper Triassic), East-central New Mexico. In: D. D. Gillette and M. G. Lockley (Editors), Dinosaur Tracks and Traces. Cambridge Univ. Press, pp. 277-280. Karaszewski, W., 1969. Tracks of Reptilia in the Lower Liassic of the Swietokrzyskie Mountains, Middle Poland. Kwart. Geol., 13, 1: 115-119. Karaszewski, W., 1975. Footprints of pentadactyl dinosaurs in the Lower Jurassic of Poland. Bull. Acad. Pol. Sci., 23, 2: 133-136. Lapparent, A. F. and Montenat, C., 1967. Les empreintes de pas de reptiles de I'Infralias du Veillon, Vend6e. M6m. Soc. G6ol. Fr., 46, 2, 107, 44 pp. Leonardi, G., 1984. Le impronte fossili di dinosauri. In: J. F. Bonaparte et al., Sulle Orme dei dinosauri. Erizzo, Venezia, 333 pp. Lockley, M. G., 1987. Dinosaur footprints from the Dakota Group of eastern Colorado. Mount. Geol., 24, (4): 107-122. Lull, R. S., 1904. Fossil footmarks of the Jura-Trias of North America. Boston Soc. Nat. Hist., 5: 461-557.
G. GIERLI~SKI Lull, R. S., 1915. Triassic Life of the Connecticut Valley. State Geol. Nat. Hist. Surv. Conn. Bull., 24, 285 pp. Lull, R. S., 1953. Triassic Life of the Connecticut Valley. State Geol. Nat. Hist. Surv. Conn. Bull., 81,336 pp. Olsen, P. E., 1980. Fossil great lakes of the Newark Supergroup in New Jersey. In: W. Manspeizer (Editor), Field Studies in New Jersey Geology and Guide to Field Trips, 52nd Annu. Meet., New York State Geol. Assoc., Newark College of Arts and Sciences, Newark, N.J. pp. 352-398. Olsen, P. E. and Baird, D., 1986. The ichnogenus Atreipus and its significance for Triassic biostratigraphy. In: K. Padian (Editor), The Beginning of the Age of Dinosaurs: Faunal change across the Triassic-Jurassic boundary. Cambridge Univ. Press, pp. 61-87. Olsen, P. E. and Galton, P. M., 1984. A review of the reptile and amphibian assemblages from the Stormberg of southern Africa with special emphasis on the age of the Stormberg. Palaeontol. Aft., 25: 87-I 10. Piefikowski, G., 1981. Sedimentology of the Lower Liassic of the northern slope of the Holy Cross Mountains. Thesis. Library Geol. Dep., Warsaw Univ. 146 pp (unpublished). Piefikowski, G., 1985. Early Liassic trace fossil assemblages from the Holy Cross Mountains, Poland: their distribution in continental and marginal marine environments. In: H. A. Curran, (Editor), Biorganic structures - - their use in interpreting depositional Environments Soc. Econ. Paleontol. Mineral., Spec. Publ., 35: 37-51. Piefikowski, G. and Gierlifiski, G., 1987. New finds of dinosaur footprints in Liassic of the Holy Cross Mountains and its paleoenvironmental background. Przegl. Geol., 4: 199-205. Welles, S. P., 1971. Dinosaur footprints from the Kayenta Formation of Northern Arizona. Plateau, 44: 27-38, Zhen, S., Li, J. and Rao, C., 1986. Dinosaur footprints of Jinning, Yunan. Mem. Beijing Nat. Hist. Mus., 33: 1-18. Zhen, S., Li, J., Rao, C., Mateer, N. J. and Lockley, M. G., 1989. A Review of Dinosaur Footprints in China. In: D. D. Gillette and M. G. Lockley (Editors), Dinosaur Tracks and Traces. Cambridge Univ. Press, pp. 187-197. Zhen, S., Li, J. and Zhen, B., 1983. Dinosaur footprints of Yuechi, Sichuan. Mem. Beijing Nat. Hist. Mus., 25: 1-19.
137
Elsevier Science Publishers B.V., Amsterdam
New dinosaur ichnotaxa from the Early Jurassic of the Holy Cross Mountains, Poland G e r a r d Gierlifiski
Ul. Jasielska 38/42 m.89, 02-128 Warsaw,Poland (Received September 15, 1988; revised and accepted October 25, 1990) ABSTRACT Gierlifiski, G., 1991. New dinosaur ichnotaxa from the Early Jurassic of the Holy Cross Mountains, Poland. Palaeogeogr., Palaeoclimatol., Palaeoecol., 85: 137-148. The numerous ornithopod tracks discovered during the last four years, in the Holy Cross Mts., allow the identification of two new ichnospecies--Anomoepuspienkovskii ichnosp, nov. and Moyenisauropuskaraszevskiiichnosp, nov. The moyenisauropodid structure indicates that the trackmakers may well have been hitherto unknown forms of the earliest iguanodontids. Newly discovered theropod footprints, Grallator (Grallator)zvierzi ichnosp, nov. and Grallator (Eubrontes) soltykovensis ichnosp, nov., supplement knowledge of the Early Jurassic terrestial fauna of Poland. The majority of the tracks occurring in the Gliniany Las exposure (Upper Hettangian) were left by dinosaurs which occupied the margins of a lagoonal environment. The other footprints, discovered in the Sottyk6w (Lower Hettangian) outcrop, were left by carnosaurs walking across the fluvial plain.
Introduction All the Polish dinosaur footprints were discovered on the northern slope of the Holy Cross Mts. These tracks have been found in the Gliniany Las and Zapni6w exposures of the Przysucha Orebearing Series (Upper Hettangian) and in the Sottyk6w outcrop of the Zagaje Series (Lower Hettangian). The first two footprints were found by Karaszewski in the Gliniany Las quarry, in 1959. He proceeded to describe them respectively as "tetradactyl track" (Karaszewski, 1969) and as Neochirotherium jurassicum (Karaszewski, 1975). Later, these specimens, and other similar footprints collected in 1983-1984, were identified as Moyenisauropus sp. (Gierlifiski and Potemska, 1987). The discovery of two new tracksites at Sottyk6w and Zapni6w (Piefikowski and Gierlifiski, 1987), and the numerous new specimens collected since 1986, have supplemented our knowledge of Hettangian dinosaurs in the Holy Cross Mountains area of Poland. 0031-0182/91/$03.50
In 1986, Anomoepus sp. was found in the uppermost part of the Sottyk6w outcrop (Pieflkowski and Gierlifiski, 1987). Further, Piefikowski and Gierlifiski (1987) described Moyenisauropus sp. from the Zapni6w outcrop. In 1988, Grallator (Eubrontes) soltykovensis ichnosp, nov. was discovered in the lowermost part of the Sottyk6w outcrop. Moreover, the exploration of the Gliniany Las quarry during 1988 and 1989 revealed the theropod footprints Grallator (Eubrontes) sp. (Gierlifiski, 1990) and Grallator (Grallator) zvierzi ichnosp, nov. Newly discovered ornithopod tracks in Gliniany Las allow me to establish two new ichnospecies Anomoepus pienkovskii ichnosp, nov. and Moyenisauropus karaszevskii ichnosp, nov.
lchnosystematic descriptions Suborder THEROPODA Marsh 1881 Ichnofamily GRALLATORIDAE Lull 1904 Ichnogenus Grallator Hitchcock 1858 Subichnogenus Grallator (Grallator) Olsen and Galton 1984
© 1991 - - Elsevier Science Publishers B.V.
138
Grallator (Grallator) zvierzi nov. sp. (Fig. 1) Holotype: Trackway preserved as a natural cast on one layer and as undertracks on another layer. The specimen preserved as a natural cast is owned by the author and the specimen preserved as a couple of undertracks is lodged in MGIW (Museum of the Geological Institute in Warsaw, Poland), as 1560.11.9. Paratype: Single footprint preserved as a natural cast. Original specimen owned by the author and its plaster cast lodged in MGIW, as 1560.11.13. Type horizon: Przysucha Ore-bearing Series (Upper Hettangian). Type locality: Gliniany Las, Holy Cross Mts., Poland. Etymology: Named in honor of Jadwiga Zwierz, head of the Museum of Geological Institute in Warsaw, who supported my research during the last few years. Diagnosis: Bipedal. Tridactyl pes with digits II-IV impressed. In the trackway, pes directed forward. Ratio of the pes length to pace is 1 : 3. The pes is wider than long by an average of 5%. Digit III is longest. The angle between digits II and IV varies from 18° to 37 °. Description: The paratype differs from the holotype by having the third digit relatively shorter in comparison to the lateral digits and by having angles beween the digits about twice as large. Other specimens, not described here (1560.II.14), 1560.11.15, 1560.11.16, in MGIW), demonstrate transitional features between the holotype and paratype; therefore, the length of digit II is 48-62% of the length of digit III, while the length of digit IV is 57-84% of that of digit III. The angles between the digit axes are: I I - I I I = 10-20 °, I l l - I V = 7 - 1 7 °. The pes length varies from 9 cm to 12.6 cm. Comparison: Among the smaller grallatorids with comparably divaricate digits, the most similar ichnites are: Grallator (Grallator) cuneatus and G. (G.) tenuis Hitchcock 1858; Grallator (Grallator) gracilis Hitchcock 1865; Grallator (Grallator) eloensis and G. (G.) variabilis Lapparent and Montenat 1967; several footprints from Lesotho reassigned to Grallator (Grallator) by Olsen and Galton (1984); Grallator (Grallator) sp. Conrad,
G. GIERLINSKI
Lockley and Prince 1987; Grallator (Grallator) ssatoi Zhen, Li, Rao, Mateer and Lockley 1989; Grallator sp. Hunt, Lucas and Kietzke 1989. However, Grallator (Grallator) zvierzi ichnosp, nov. is broader than long, whereas the ichnites mentioned above are always longer than broad. Discussion: Olsen and Galton (1984) considered the names Eubrontes and Anchisauripus as synonymous with Grallator. They used the subichnogeneric designations to denote grallatorid tracks in the size categories. Consequently, the size and morphology of those footprints assigns them to the subichnogenus Grallator ( Grallator). Subichnogenus Grallator (Eubrontes) Olsen and Galton 1984
Grallator (Eubrontes) soltykovensis nov. sp. (Fig.2) Holotype: Single footprint preserved as a natural cast. Original specimen owned by the author and its piaster cast lodged in MGIW, as 1560.11.12. Paratype: Single footprint preserved as a natural cast. Original specimen owned by the author and its plaster cast lodged in MGIW, as 1560.11.10. Type horizon: Zagaje Series (Lower Hettangian). Type locality: Sottykrw, Holy Cross Mts., Poland. Etymology: Named after the type locality, Sottyk6w. Diagnosis: Bipedal. Tridactyl large pes with digits II-IV impressed. Metatarsal-phalangeal pads of digits III and IV made an isolated oblong posterior impression. The third digit is longest. The angle between digits II and IV varies from 50 ° to 60 °. The angle between digits IV and III is always larger than that one between digits III and II. Description: The length of holotype is 22.5cm, whereas the paratype is 28 cm long. Moreover, the paratype has a larger angle between digits III and IV and a smaller angle between digits II and III than in the holotype. The paratype is very deeply impressed, inclined forward and sideward by 6 °, opposite to shallow and equally impressed holotype. The paratype's digits are thickly impressed and the third digit is curved; most probably, this was caused by a non-taxonomic factor, probably resulting from the trackmaker's mode of locomotion and the nature of the sediment.
139
N E W D I N O S A U R I C H N O T A X A F R O M T H E EARLY J U R A S S I C O F T H E HOLY C R O S S M O U N T A I N S , P O L A N D
~." ,v
."~:~.
':,
,
~:-:!~ .,~,.,~t~-~. ..." ;:"~
I'.,::."::.~.~ .....
"z. :".('. ""-':.':-.
".".,.,.~'v'.: ", ~ "'.."':"~.. :
°~o.
, •
.
:
a
10 cm I
I
:i;.......':!'~i!:~'~ ;~]'.. • ...~
~:i~~
~v~,,~ ~: ,
C".
....:......
":~
i~-:., :.,.
"." ".':'~'~ ,
".:.
.~..
_ •
"" ;" ""'
. ::'.
"-~
...~....~-~;.!.....,"~...,"..'.......'~
b
Fig. 1. Grallator (Grallator) zvierzi ichnosp, nov., natural casts from Gliniany Las (Upper Hettangian). a. Trackway designated as holotype, b. Paratype.
140
G. GIERLINSKI
~-:,-.. ;,~.
"~::'P"
.:~,
. :;I
.~ '" " ~
~":'..~
~ 4." ~,
'.:...'~~,~. L- :~.~ x4<.t :~ %:"~'!:":7~.-.';.':'";~'.'.~. :~,~ ;:~ "";:'" ~::~i!& .... $:. ~' ::'~7. "s,~s'~.~' :~' ~.. "..'7~ "!:~,,:.' ..... ;..f;' ..f..i" .~' ~--'-.; : :.,.', ... ":;; ~.:~9. .;....-..~ ..7~ "~.." ' ;.. q.~-"..
.. • "
~t[..-~
.Jcm
, :.:.~ ]'~'::
~7!::?~:~7;;.: ~.:" .".'.:'4' •~"., .. ".:'~"':;;
l:.:
• :: ..u
...~::'~,~ ~".o...'
.'~ "'.~.
N}! i;'::: 7 ~:".' :. ~ ts',..
."!
~.::'~'.'~.-':.'~
.'."
:,,.~o
..,sc~e;$'>'~.~<..J
--,~.
N'::':-!
. ....:.,:4
• ',~ ~:.:.....- .~::.:.,~.........:.!.;,:!......¢~7. ..:...:.;'-:...'.!~ ~ ..........~ .:¢,.~.',.'-,~ .:. :., "~:~:'.;,:;.. " •~.~,.:.,..->~. .....~-..,...,:....::::..!i~
-K¢~d."~ "'"
' ~ , 7 ° '~ ' ; . . . .
IV .~.,..W
..
.
•
.,.;° . # , ~ i , . , . : ~ r
'~'~" ,~4.:'.'*, :"-,":.,.;;2.;,~,~ ~ b
Fig.2.
Grallator
(Eubrontes)
soltykovensis
" ...~...~ ~~.~L "." ......~:. ;..~.~;:':;.,.:.. : "":.;':::;.,.,-
ichnosp, nov,, n a t u r a l casts f r o m S o i t y k 6 w ( L o w e r H e t t a n g i a n ) . a. H o l o t y p e . b. P a r a t y p e .
Phalangeal pads are visible on digits II and III (especially in the holotype). In digit II, imprints of two phalangeal pads are present and three phalangeal pads are in digit III. The length of digit II is 54-57% of the length of digit III, while the length of digit IV is 70-76% of that of digit III. Angles between digits are: I I - I I I = 15-25 °, IIIIV = 30-40 o. Comparison: The large size of these footprints combined with their grallatorid structure assigns
them to the ichnotaxon Grallator (Eubrontes), following Olsen and Galton (1984). Grallator (Eubrontes) soltykovensis ichnosp, nov. has highly divaricated digits. Among Liassic, large theropod tracks, this bears a resemblance to: Grallator (Eubrontes) divaricatus (Hitchcock 1865) Lull 1904; Kayentapus and Dilophosauripus Welles 1971; Neotrisauropus, Plastisauropus and Kainotrisauropus, considered to be synonyms of Grallator by Olsen and Galton 1984; Schizograllator Zhen, Li
NEW DINOSAUR ICHNOTAXA FROM THE EARLY JURASSIC OF THE HOLY CROSS MOUNTAINS, POLAND
and Rao 1986. However, Grallator (Eubrontes) divaricatus, Dilophosauripus and Schizograllator differ from G. (E.) soltykovensis by having the third digit relatively shorter in comparison to the lateral digits. On the other hand, G. (E.) soltykovensis differs from Kayentapus, Neotrisauropus, Plastisauropus and Kainotrisauropus, in that digit II originates further posteriorly. Discussion: The numeration of the digits of single tridactyl tracks from the Liassic poses no problems, if the phalangeal pads are visible. The pedal digit II has always two phalangeal pads, while the pedal digit IV has three or four phalangeal pads. Suborder ORNITHOPODAMarsh 1878 Ichnofamily ANOMOEPODIDAELull 1904, emendedherein Included ichnogenera: Anomoepus Hitchcock 1848, Apatichnus Hitchcock 1858, Jialingpus Zhen, Li and Zhen 1983
Occurrence: Lower Jurassic to Upper Jurassic of North America, Europe, southern Africa and southeastern Asia. Emended diagnosis: Normally bipedal, but sometimes also quadrupedal. Pes functionally tridactyl. Hallux rarely impressed. Pedal digits II, III and IV tend to be subequal in length. In the trackway, the pes is directed forward. Manus imprints are four-toed or five-toed. Comparison: Anomoepodidae Lull 1904, emend. herein show the transitional morphology between Atreipodidae Olsen and Baird 1986 and Moyenisauropodidae Ellenberger 1974, emended herein. In Anomoepodidae, the pedal digits II, III and IV tend to be subequal in length (Olsen and Galton 1984), the g r a l l a t o r i d - like pes of the Atreipodidae has digit III clearly as longest, while in the Moyenisauropodidae, the pedal digit IV is always the longest one. Moreover, the manus of the Atreipodidae is tridactyl or tetradactyl, the Anomoepodidae have tetradactyl or pentadactyl manus, whereas the manus of the Moyenisauropodidae is pentadactyl. Ichnogenus Anomoepus Hitchcock 1848, emended herein.
141
1848 AnomoepusHitchcock,p. 220 1858 AnornoepusHitchcock, p. 55 1904 AnomoepusHitchcock. Lull, p. 500 1915 AnomoepusHitchcock. Lull, p. 208 1953 AnomoepusHitcbcock.Lull, p. 192 1971 AnomoepusHitchcock. Haubold, p. 83 1984 AnomoepusHitchcock. Olsen and Galton, p. 99 Type ichnospecies: Anomoepus scambus Hitchcock 1848, p. 222, pl. 13, figs.l-6, from the Portland Formation (Upper Liassic) of Massachusetts, USA. Included ichnospecies: Anomoepus scambus Hitchcock 1848, A. intermedius, At curvatus, A. minimus Hitchcock 1865, A. cuneatus, A. crassus, A. isodactylus Hitchcock 1889, A. gracillimus Hitchcock 1844, A. ranivorus Ellenberger 1970, A. natatilis, A. dodai, A. vermivorous, A. longicauda, A. minor Ellenberger 1974 and A. pienkovskii ichnosp, nov.
Occurrence: East Berlin Formation (Sinemurian) and Portland Formation (Pliensbachian to Toarcian) of Connecticut, Massachusetts and New Jersey, USA. Upper Red Beds (Lower Hettangian) of Lesotho, southern Africa. Zagaje Series (Lower Hettangian) and Przysucha Ore-bearing Series (Upper Hettangian) of the northern slope of the Holy Cross Mts., Poland. Emended diagnosis: Normally bipedal, but sometimes quadrupedal also. Pes with three functional digits. Hallux rarely imprinted. Pedal digits II, III and IV usually tend to be subequal in length, but sometimes digit IV is clearly the longest one. In the trackway, the pes is directed forward or inward. Metatarsal impression occurs when the trackmaker rests. The manus is tetradactyl or pentadactyl. Manual digits II and III are longest. Comparisons: In Anomoepus, the angle between pedal digits III and IV in comparison to that one between digits II and III, is smaller than in Apatichnus. Metatarsal Anomoepus tracks evidently occur as resting traces only, whereas metatarsal tracks of Jialingpus were made during active locomotion. Anomoepus pienkovskii nov. sp. (Fig.3) Holotype: Trackway preserved as a natural cast. Original specimen owned by the author and its plaster cast lodged in MGIW, as 1560.I1.20.
142
G. GIERLINSKI
IV )."
..,/
.....
:
.t
10 cm ~
..:,:....
Fig.3. Holotypeof Anomoepuspienkovskiiichnosp, nov., natural cast of the trackwayfrom GlinianyLas (Upper Hettangian).
Type horizon: Przysucha Ore-bearing Series (Upper Hettangian).
Type locality: Gliniany Las, Holy Cross Mts., Poland.
Etymology: Named in tribute to Dr. Grzegorz Piefikowski, my co-worker in the studies on footprints from the Holy Cross Mts., who has made sedimentological interpretations of the Liassic deposits in Poland. Diagnosis: Quadrupedal in gait. Pes functionally tridactyl with hallux sometimes also imprinted.
The fourth pedal digit is longest. In the trackway, pes and manus are directed forward, with the manus situated anteriorly and slightly outward in comparison to the pes. Ratio of pes length to pes pace 1:2.4. Ratio of manus length to manus pace 1:5.4. The angle between pedal digits II and IV varies from 48 ° to 58 °. Manus tetradactyl, half the size of the pes and wider than long by an average of 33%. Manual digits II and III are longest. Description: Holotype length: pes = 14 cm, m a n u s = 7 cm. The length of pedal digit II averages
143
NEW DINOSAUR ICHNOTAXA FROM THE EARLY JURASSIC OF THE HOLY CROSS MOUNTAINS. POLAND
77% of the length of digit IV, while the length of digit III averages 84% of that of digit IV. Angles between the pedal digits: I - I I = 8 0 °, I I - I I I = 30-32 °, Ill-IV = 20-27 °. The manus with digits IIV impressed. Manual digits II and III are longest followed in length by digits I and IV. Angles between the manual digits: I-II = 54-57 °, II-III = 57-59 °, Ill-IV = 65-75 °. Comparisons: The manus is only half the size of the pes. In the trackway, the manus is situated slightly outward to the pes impression, whereas in other ichnospecies of Anomoepus the manus imprints are only one-third the size of the pes imprints, the manus imprints lying closer to the midline of the trackway. lchnofamily MOYENISAUROPODIDAEEllenberger 1974, emended herein Included ichnogenus: Moyenisauropus Ellenberger 1974, emended herein
Occurrence: Lower Jurassic of southern Africa and Europe. Emended diagnosis: Normally bipedal, but sometimes also quadrupedal. Pes with three functional digits. Hallux rarely imprinted. Pedal digits massive, short and highly divaricated, with imprints of two phalangeal pads on the third digit. The fourth pedal digit is longest. In the trackway, the pes is directed inward. Metatarsal impression occurs when the trackmaker rests. The pentadactyl manus has short and thick digits. Comparison: The pes of Moyenisauropodidae EIlenberger 1974 emend, herein shows two phalangeal pads on digit III, whereas, the pedal digit III of Anomoepodidae has three phalangeal pads and iguanodontid tracks (e.g., Caririchnium Leonardi 1984) have only one phalangeal pad in that digit. The remaining features of Moyenisauropodidae make this ichnofamily closer to iguanodontid footprints than to the ichnofamily Anomoepodidae. The pes and manus imprints of the Iguanodontidae and Moyenisauropodidae differ from those of the Anomoepodidae in having shorter and thicker digits. In the moyenisauropodid and iguanodontid trackway the pes was clearly directed inward,
whereas in the anomoepodid trackway the pes was usually turned forward. Ichnogenus Moyenisauropus emended herein
Ellenberger
1974,
1974 MoyenisauropusEllenberger, p. 25 1984 AnomoepusHitchcock. Olsen and Galton, p. 99 1987 Moyenisauropus Ellenberger. Gierliflski and Potemska, p. 109 Type ichnospecies: Moyenisauropus natator EIlenberger 1974, p. 26, pls. C and D, from the Upper Red Beds (Lower Hettangian) of Lesotho, southern Africa. Included ichnospecies: Moyenisauropus natator, M. levicauda Ellenberger 1974 and M. karaszevskii ichnosp, nov.
Occurrence: Upper Red Beds (Lower Hettangian) of Lesotho, southern Africa. Przysucha Ore-bearing Series (Upper Hettangian) of the northern slope of the Holy Cross Mts., Poland. Emended diagnosis: Normally bipedal, but sometimes also quadrupedal. Pes functionally tridactyl. Hallux rarely imprinted. Pedal digits massive, short and highly divaricated. Two phalangeal pads are present in the pedal digit III. The pedal digit IV is longest. In the trackway, the pes is directed inward. Metatarsal impression occurs when the trackmaker rests. In the pes imprints, which were left during active locomotion, the angle between digits II and IIl is always larger than that one between digits III and IV. The pentadactyl manus has short and thick digits. Manual digits are subequal in length. Comparison: Moyenisauropus is distinguished from other tridactylous ichnogenera from the Liassic in having only two phalangeal pads in the pedal digit IIl. Moreover, in contrast to Anomoepus, the manual digits of Moyenisauropus are subequal in length. In Moyenisauropus the angle between pedal digits II and III in comparison to that one between digits III and IV, is usually larger than in the pes of Anomoepus. Discussion: Ellenberger (1974) erected the ichnogenus Moyenisauropus to comprise the ichno-
144
species M. natator, M. natatilis, M. dodai, M. vermivorous, M. longicauda, M. minor and M. levicauda. In addition Ellenberger proposed their placement into the ichnofamily Moyenisauropodidae. Olsen and Galton (1984) and Haubold (1984, 1986) treated Moyenisauropus as a junior synonym of Anomoepus. However, Gierlifiski and Petemska (1987) proposed to retain Moyenisauropus as a separate ichnogenus in the ichnofamily Anomoepodidae. In fact, Moyenisauropus natatilis, M. dodai, M. vermivorous, M. longicauda and M. minor all bear a resemblance to Amomoepus. In contrast, Moyenisauropus natator, M. levicauda and M. karaszevskii ichnosp, nov. do not correspond in morphology to the Anomoepodidae. Consequently, I propose to transfer Moyenisauropus natatilis, M. dodai, M. vermivorous, M. longicauda and M. minor to the ichnogenus Anomoepus and to retain M. natator, M. levicauda and M. karaszevskii ichnosp, nov. as Moyenisauropus, in their own ichnofamily Moyenisauropodidae.
Moyenisauropus karaszevskii nov. sp. (Fig.4) Holotype: Trackway preserved as a natural cast on one layer and as undertracks on another layer. The specimen preserved as a natural cast is owned by the author and the specimen preserved as a couple of undertracks is lodged in MGIW, as 1560.11.9. Type horizon: Przysucha Ore-bearing Series (Upper Hettangian). Type locality: Gliniany Las, Holy Cross Mts., Poland. Etymology: Named in honor of Dr. Wtadystaw Karaszewski, the discoverer of the first dinosaur footprints in Poland. Diagnosis: Bipedal in gait. Tridactyl pes, with hallux sometimes also imprinted. Two phalangeal pads are present in the pedal digit III. The pedal digit IV is longest. In the trackway, the pes is directed inward. Ratio of pes length to stride 1:3.8. Ratio of pes length to pace 1:2.3. Pace angulation 103°. The angle between pedal digits II and III is always larger than that one between
G. GIERLII~SKI
digits III and IV. The angle between pedal digits II and IV varies from 50 ° to 56 °. Description: The pes length varies from 22 to 26 cm. The length of digit II averages 67% of the length of digit IV, while the length of digit III averages 84% of that of digit IV. The angles between the digit axes are: I - I I = 6 5 °, I I - I I I = 30-36 °, I I I - I V = 17-22 °. The manus imprint associated with the pes has not been found. Comparison: The pedal digit II of this new ichnospecies originates further posteriorly than in the pes of M. natator. Moyenisauropus levicauda differs from M. karaszevskii ichnosp, nov. by having the pedal digit III relatively longer in comparison to the pedal digit IV.
Moyenisauropus sp. (Fig.5) Material: Original specimen owned by the author and its plaster cast lodged in MGIW, as 1560.11.8. It is a single manus imprint preserved as natural cast, from Gliniany Las, Holy Cross Mts., Poland, (Przysucha Ore-beating Series; Lower Hettangian). Description: Pentadactyl manus, wider than long by 48.5% (16.5 cm width and 8.5 cm long). Digits short, massive and subequal in length. The first digit is thinnest; more markedly bent away from the others; ends in a long, straight claw. The tips of the other four digits are rounded. Angles between the digits: I-II--62 °, I I - I I I = 60 °, III-IV = 32 °, IV-V = 42 °. Comparison: The features mentioned above clearly differ from those of the Anomoepodidae and are identical to that of the Moyenisauropus manus. Consequently, in contrast to the manual digits of Anomoepus, digits of this ichnite are subequal in length. Moreover, our specimen has shorter and thicker digits than are typical of the Anomoepodidae. Discussion: Because of the lack of material where the manus would occur together with the pes, we cannot with entire certainty identify it as M. karaszevskii, an ichnospecies created on the basis of the analysis of the pes. According to the structure of the moyenisauropodid manus, the differences between the digits make the numeration of the digits possible. Digits II, III, IV and V are similar in width, whereas
145
N E W D I N O S A U R I C H N O T A X A F R O M T H E EARLY J U R A S S I C O F T H E HOLY C R O S S M O U N T A I N S , P O L A N D
IV • .:;:~!~
~.
~...'.::.
~:"
-:.q :.::..~,..-....~ ,#
,:.~.?".:. .... •:.':."./?~)i~ . . :.~,?q~:,:-- : .: •
:,.,...... " . , : ~
~ ,::::i:!! III.................. :
:{.i....
i:.,...':........... ."-
lOcm ..~::~
"t ..... :
..° ....,
:'-...:
.. ......
"":::~....
)
~
IV
Fig.4. H o l o t y p e o f Moyenisauropus karaszevskii ichnosp, nov., natural cast o f the t r a c k w a y from G l i n i a n y Las ( U p p e r Hettangian).
146
G. GIERLIIqSKI
5 cm
:~.-,..~'?...~ ~,'~," ." : 4 .-". . . .-. ! "., "~: '~.., .~ L~'.:;' .Z:,,:~'.":.-..-':~"" ~,..~,,,~
V ~,~r÷':".~,.-" l ~:~..'.."'" y'::':.:-...".:":.~"'.";';'z.:~ ::'::~ :.'-:....-~?~--"~::";<,;t'.:~,~
~@~
Fig.5. Moyenisauropussp., natural cast of the single manus ichnite from Gliniany Las (Upper Hettangian). a. Whole specimen, b. Pollex with a spike impressed at the end. digit I is thinner and is more markedly bent away from the others. Systematic
comparisons
The grallatorid structure is characterized by the third pedal digit being always longer than the lateral digits (Baird 1957; Lapparent and Montenat, 1967; Olsen 1980). Among Liassic dinosaurs, this bears a resemblance to the pes of the Podokesauridae and of Megalosauridae. Consequently, Grallator (Grallator) might have been imprinted either by podokesaurids (Lull, 1953; Colbert 1963) or by a juvenile megalosaurs, whereas Grallator (Eubrontes) imprints were probably made exclusively by megalosaurs like Dilophosaurus. Anomoepus is conventionally considered as the footprints of Fabrosauridae, previously classed as Hypsilophodontidae (Lull, 1953; Haubold, 1984; Olsen and Baird, 1986). The differences between Anomoepus and the tracks here referred to Moyenisauropodidae have allowed me to ascertain that their trackmakers belong to different dinosaur families.
In the trackways of Moyenisauropus natator described by Ellenberger 1974 and in M. karaszevskfi ichnosp, nov. the pes turned inward, whereas the Anomoepus pes was usually directed forward. The inward rotation of the pes is indicative of large ornithopods such as iguanodontids and hadrosaurids (Lockley 1987). Moreover, the size and structure of Moyenisauropus ichnites suggest that their trackmakers were larger and more massive than were the Fabrosauridae. The pes prints of Anomoepus are 5 cm up to 16 cm long, whereas the pes length of Moyenisauropus varies from 10 cm to 26 cm. The short, thick and highly divaricated pedal digits of Moyenisauropodidae are closest to iguanodontid footprints. Moreover, the moyenisauropodid manus shows less differentiation in the digit lengths than in the manus of Heterodontosauridae or Fabrosauridae. It is further noteworthy that Moyenisauropus manus has a clearly offset, iguanodont-like pollex. Sometimes, a straight spike is impressed at the end of it (Fig.5b), in contrast to the hooked pollex claw of the prosauropod manus, as in Navahopus (Baird 1980).
NEW DINOSAURICHNOTAXAFROM THE EARLYJURASSICOF THE HOLYCROSSMOUNTAINS,POLAND
Consequently, the features described above confirm that Moyenisauropodidae represent footprints of some hitherto unknown early members of the family Iguanodontidae. This contributes to our knowledge of the early evolution of iguanodontids and indicates that this family appeared much earlier than has been indicated by their osteological remains, near the boundary between the Triassic and Jurassic. Paleoenvironmental and paleoecological remarks From knowledge of the profile of the Gliniany Las outcrop (Piefikowski 1985, p. 47) and my investigations performed during the last two years, the footprints were left in certain mudstone layer of lagoonal origin, which were superimposed upon barrier deposits. The tracks occur either as natural casts on the bottom surface of the superimposed sandstone layer or as undertracks preserved on the top of another sandstone layer underlying the lagoonal mudstone. The small and shallow lagoon, with a strong influx of fresh water, was situated between a barrier system separating it from the shallow brackish sea and a deltaic-fluvial area (Piefikowski 1985). It appears likely that the dinosaurs from Gliniany Las found quite favourable living conditions in the barrier - - lagoon transitional environment. Up to this moment, the theropod tracks form 11% of the number of all the footprints discovered in this outcrop. The rest of the footprints belong to ornithopods. In the lowermost part of the Sottyk6w outcrop only the carnosaur tracks, here described as Grallator (Eubrontes) soltykovensis ichnosp, nov., have been found. Those footprints were imprinted into the muddy fluvial plain deposits and preserved at the bottoms of superimposed sandy layers deposited by fluvial processes - presumably crevasse splays (Piefikowski, 1981, unpublished; Piefikowski and Gierlifiski 1987). Acknowledgements I would like to thank my wife for her help in preparing this paper. I am deeply indebted to Professor Dr. William A. S. Sarjeant (University of Saskatchewan, Canada) for his valuable corn-
147
ments and suggestions. I also thank Dr. G. Pieflkowski (Geological Survey of Poland) for his helpful remarks. I am grateful to my friends: A. Piotrowski, Z. W6jcik, T. Jarzabek, W. Knapczyk and P. Kabulski for field assistance. J. Zwierz and P. Nowacki offered their help during my researches. My special thanks go to A. Potemska for her help in preparing this paper, as well as for her collaboration during the last few years in the study of Polish dinosaurs. Her help was invaluable to me, both as co-worker and as a great friend. References Baird, D., 1957. Triassic reptile footprint faunules from Milford, New Jersey. Mus. Comp. Zool. (Harvard Univ.), Bull., 117: 449-520. Baird, D., 1980. A prosauropod dinosaur trackway from the Navajo Sandstone (Lower Jurassic) of Arizona. In: L. L. Jacobs (Editor), Aspect of Vertebrate History. Mus. Northern Ariz. Press, pp. 219-230. Colbert, E. H. 1963. Fossils of the Connecticut Valley: The age of dinosaurs begins. State Geol. Nat. Hist. Surv. Conn. Bull., 96, 31 pp. Conrad, K., Lockley, M. G. and Prince, N. K., 1987. Triassic and Jurassic vertebrate-dominated trace fossil assemblages of the Cimarron valley region: implications for paleoecology and biostratigraphy. In: S. G. Lucas and A. P. Hunt (Editors), New Mexico Geol. Soc. Guidebook, 38th Field Conf. Univ. New Mexico Press, pp. 127-138. Ellenberger, P., 1970. Les niveaux paleontologiques de premi6re apparition des Mammif6res Primordiaux en Afrique du Sud et leur Ichnologie: Establissement de zones stratigraphiques detaill6es dans le Stormberg du Lesotho (Afrique du Sud), (Trias Superieur fi Jurassique). In: S. H. Haughton (Editor), I.U.G.S., 2nd Symposium on Gondwana Stratigraphy and Palaeontology, Pretoria, pp. 343-370. Ellenberger, P., 1974. Contribution ~, la classification des pistes de Vertebres du Trias: Les types du Stormberg d'Afrique du sud (2 6me Partie: Le Stormberg Superieur. I. Le biome de la zone B/1 au niveau de Meyeni: ses bioc6noses). Palaeovertebrata M+me. extraordinaire, 141 pp. Gierlifiski, G. and Potemska, A., 1987. Lower Jurassic dinosaur footprints from Gliniany Las, northern slope of the Holy Cross Mountains, Poland. Neues Jahrb. Geol. Pal~iontol. Abh., 175: 107-120. Gierlifiski, G., 1990. First find of a carnosaur footprint in the Lower Jurassic deposits of Gliniany Las, Holy Cross Mts. Przeg. Geol., 7-8: 315-317. Haubold, H., 1971. Ichnia Amphibiorum et Reptiliorum fossilium. In: O. Kuhn (Editor), Encyclopedia of Palaeoherpetology part 18. Fischer, Stuttgart, 124pp. Haubold, H., 1984. Saurierffihrten, (Neue Brehm-Bficherei, 479) Ziemsen, Wittenberg, 231 pp. Haubold, H., 1986. Archosaur footprints at the terrestial
148 Triassic-Jurassic transition. In: K. Padian (Editor), The Beginnings of the Age of Dinosaurs: Faunal change across the Triassic-Jurassic boundary. Cambridge Univ. Press, pp. 189-201. Hitchcock, C. H., 1889. Recent progress in ichnology. Proc. Boston Soc. Nat. Hist., 24- 117-127. Hitchcock, E., 1844. Report on ichnolithology, or fossil footmarks, with a description of several new species, and the coprolites of birds, from the valley of Connecticut river, and of a supposed footmarks from the valley of Hudson river. Am. J. Sci., 47: 292-322. Hitchcock, E., 1848. An attempt to discriminate and describe the animals that made the fossil footmarks of the United States, and especially New England. Mere. Am. Acad. Arts. Sci., 3(2): 129-256. Hitchcock, E., 1858. Ichnology of New England. A report on the sandstone of the Connecticut Valley, especially its fossil footmarks. White, Boston, 220 pp. Hunt, A. P., Lucas, S. G. and Kietzke, K. K., 1989. Dinosaur footprints from the Redonda Member of the Chinle Formation (Upper Triassic), East-central New Mexico. In: D. D. Gillette and M. G. Lockley (Editors), Dinosaur Tracks and Traces. Cambridge Univ. Press, pp. 277-280. Karaszewski, W., 1969. Tracks of Reptilia in the Lower Liassic of the Swietokrzyskie Mountains, Middle Poland. Kwart. Geol., 13, 1: 115-119. Karaszewski, W., 1975. Footprints of pentadactyl dinosaurs in the Lower Jurassic of Poland. Bull. Acad. Pol. Sci., 23, 2: 133-136. Lapparent, A. F. and Montenat, C., 1967. Les empreintes de pas de reptiles de I'Infralias du Veillon, Vend6e. M6m. Soc. G6ol. Fr., 46, 2, 107, 44 pp. Leonardi, G., 1984. Le impronte fossili di dinosauri. In: J. F. Bonaparte et al., Sulle Orme dei dinosauri. Erizzo, Venezia, 333 pp. Lockley, M. G., 1987. Dinosaur footprints from the Dakota Group of eastern Colorado. Mount. Geol., 24, (4): 107-122. Lull, R. S., 1904. Fossil footmarks of the Jura-Trias of North America. Boston Soc. Nat. Hist., 5: 461-557.
G. GIERLI~SKI Lull, R. S., 1915. Triassic Life of the Connecticut Valley. State Geol. Nat. Hist. Surv. Conn. Bull., 24, 285 pp. Lull, R. S., 1953. Triassic Life of the Connecticut Valley. State Geol. Nat. Hist. Surv. Conn. Bull., 81,336 pp. Olsen, P. E., 1980. Fossil great lakes of the Newark Supergroup in New Jersey. In: W. Manspeizer (Editor), Field Studies in New Jersey Geology and Guide to Field Trips, 52nd Annu. Meet., New York State Geol. Assoc., Newark College of Arts and Sciences, Newark, N.J. pp. 352-398. Olsen, P. E. and Baird, D., 1986. The ichnogenus Atreipus and its significance for Triassic biostratigraphy. In: K. Padian (Editor), The Beginning of the Age of Dinosaurs: Faunal change across the Triassic-Jurassic boundary. Cambridge Univ. Press, pp. 61-87. Olsen, P. E. and Galton, P. M., 1984. A review of the reptile and amphibian assemblages from the Stormberg of southern Africa with special emphasis on the age of the Stormberg. Palaeontol. Aft., 25: 87-I 10. Piefikowski, G., 1981. Sedimentology of the Lower Liassic of the northern slope of the Holy Cross Mountains. Thesis. Library Geol. Dep., Warsaw Univ. 146 pp (unpublished). Piefikowski, G., 1985. Early Liassic trace fossil assemblages from the Holy Cross Mountains, Poland: their distribution in continental and marginal marine environments. In: H. A. Curran, (Editor), Biorganic structures - - their use in interpreting depositional Environments Soc. Econ. Paleontol. Mineral., Spec. Publ., 35: 37-51. Piefikowski, G. and Gierlifiski, G., 1987. New finds of dinosaur footprints in Liassic of the Holy Cross Mountains and its paleoenvironmental background. Przegl. Geol., 4: 199-205. Welles, S. P., 1971. Dinosaur footprints from the Kayenta Formation of Northern Arizona. Plateau, 44: 27-38, Zhen, S., Li, J. and Rao, C., 1986. Dinosaur footprints of Jinning, Yunan. Mem. Beijing Nat. Hist. Mus., 33: 1-18. Zhen, S., Li, J., Rao, C., Mateer, N. J. and Lockley, M. G., 1989. A Review of Dinosaur Footprints in China. In: D. D. Gillette and M. G. Lockley (Editors), Dinosaur Tracks and Traces. Cambridge Univ. Press, pp. 187-197. Zhen, S., Li, J. and Zhen, B., 1983. Dinosaur footprints of Yuechi, Sichuan. Mem. Beijing Nat. Hist. Mus., 25: 1-19.