A Dicynodont-Theropod Association in the Latest Triassic of Poland

Abstract

It is generally accepted that during the Triassic the composi− tion of tetrapod faunas underwent a series of fundamental transformations, mainly as a result of diversification of archosaurs and decline of therapsids (Benton 1994, 2004, 2006). The last herbivorous basal synapsids, dicynodonts, disappeared from the record in the early Norian of the Americas, about 220 Ma (Langer et al. 2007), being un− known from the Late Triassic of Europe. Here, we report a partially articulated skeleton and isolated bones of a giant rhino−size dicynodont in the Upper Triassic fluvial sedi− ments at Lisowice (Lipie Śląskie clay−pit) in southern Po− land. Paleobotanical data indicate an early Rhaetian age for the fauna (Dzik et al. 2008; Niedźwiedzki and Sulej 2008). The dicynodont bones are associated with bones of carnivo− rous dinosaurs, pterosaurs, as well as capitosaur and plagio− saur amphibians. Dicynodonts were represented in the Ger− manic Basin throughout the Late Triassic, as proven by findings of smaller dicynodonts in older deposits in the same area, associated there with temnospondyl amphibians. It ap− pears, thus, that the fossil record of tetrapod succession in the Late Triassic was strongly controlled by ecological fac− tors and biased by uneven representation of particular envi− ronments. The Lisowice assemblage proves that faunas dominated by dicynodonts did not entirely disappear at least until the end of the Triassic.

Full text

A dicynodont−theropod association in the latest Triassic of Poland
JERZY DZIK, TOMASZ SULEJ, and GRZEGORZ NIEDŹWIEDZKI
It is generally accepted that during the Triassic the composi
−
tion of tetrapod faunas underwent a series of fundamental
transformations, mainly as a result of diversification of
archosaurs and decline of therapsids (Benton 1994, 2004,
2006). The last herbivorous basal synapsids, dicynodonts,
disappeared from the record in the early Norian of the
Americas, about 220 Ma (Langer et al. 2007), being un
−
known from the Late Triassic of Europe. Here, we report a
partially articulated skeleton and isolated bones of a giant
rhino−size dicynodont in the Upper Triassic fluvial sedi
−
ments at Lisowice (Lipie Śląskie clay−pit) in southern Po
−
land. Paleobotanical data indicate an early Rhaetian age for
the fauna (Dzik et al. 2008; Niedźwiedzki and Sulej 2008).
The dicynodont bones are associated with bones of carnivo−
rous dinosaurs, pterosaurs, as well as capitosaur and plagio−
saur amphibians. Dicynodonts were represented in the Ger−
manic Basin throughout the Late Triassic, as proven by
findings of smaller dicynodonts in older deposits in the same
area, associated there with temnospondyl amphibians. It ap−
pears, thus, that the fossil record of tetrapod succession in
the Late Triassic was strongly controlled by ecological fac−
tors and biased by uneven representation of particular envi−
ronments. The Lisowice assemblage proves that faunas
dominated by dicynodonts did not entirely disappear at
least until the end of the Triassic.
Introduction
Bone−bearing greenish, reddish, and grey fluvial mudstones and
siltstones, interbedded with cross− or horizontally−stratified grey
−
wacke sandstones (the whole stratigraphic section is aproximately
12 meters thick) are exposed in the Lipie Śląskie clay−pit (Fig. 1)
at Lisowice village, near the town Lubliniec in southern Poland
(Szulc et al. 2006). Well−preserved vertebrate bones occur in a len
−
ticular body of grey mudstone and claystone in the northern part of
the Lipie Śląskie clay−pit, mostly covered with calcareous and py
−
ritic crust or preserved within limestone concretions. The verte
−
brate assemblage is dominated by bones of a giant dicynodont, as
well as large theropod bones. Other vertebrate remains, including
a coelophysoid dinosaur, pterosaur, large capitosaur, and small
plagiosaur, as well as dipnoan teeth, hybodont shark fin spines,
and ganoid skull elements and scales, are rare. About 150 m below
this fossil−bearing greenish−gray sequence, playa and fluvial red
−
dish mudstones of the Keuper occur, lithostratigraphically corre
−
latable with the Late Carnian strata of Krasiejów containing
Paleorhinus and Metoposaurus, located about 25 km to east (Dzik
et al. 2000; Dzik 2001, 2003; Dzik and Sulej 2007; Sulej 2005).
The fresh−water hybodont sharks elsewhere tend to be especially
abundant in times of global sea level rise. Their presence in
Lisowice and absence in Krasiejów may thus be meaningful.
In these bone−bearing strata numerous well−preserved micro−
and macrofloral remains were found. The dominant plant species
at Lisowice is a conifer similar to Hirmeriella muensteri (Schenk,
1867), as is the case with the Rhaetian and earliest Liassic floras
of the region (Clement−Westerhof and Van Konijnenburg−Van
Cittert 1991; Reymanówna 1992), although its twigs (in ZPAL
V33/212) are more robust than in Jurassic specimens. Abundant
pollen of Classopollis morphology occurring there is elsewhere
associated with this species (Reymanówna 1992). The second
most common plant species is represented by twigs (ZPAL
V33/213) similar to Stachyotaxus septentrionalis (Agardh, 1823)
from the Rhaetian of Greenland and Scania (Harris 1932; Arndt
2002). Associated seed scales (ZPAL V33/214), with no signs of
separate bract and scale being present, have entire margins and ap−
parently supported two ovules. Unlike S. septentrionalis,asinter−
preted by Arndt (2002), the scale apex is blunt, resembling instead
cone scales of the south−Gondwanan Jurassic Mataia (Townrow
1967). Stachyotaxus produced spherical wingless pollen (Harris
1931; Stewart and Rothwell 1993), which could be represented by
Brachysaccus neomundus (Leschik, 1955) at Lisowice. The prob
−
able gingkoalean pollen Monosulcites cf. M. minimus Cookson,
1947 (leaves—ZPAL V33/215 similar to Schmeissneria are asso
−
ciated) and the liverwort spores Ricciisporites cf. R. tuberculatus
Lundblad, 1954 are present at Lisowice; these taxa are known
only from other central European sites believed to correspond
to late Norian and early Rhaetian (Orłowska−Zwolińska 1983;
Schultz and Heunisch 2005). Well preserved cuticule fragments of
typical Rhaetian seed−fern, Lepidopteris cf. L. ottonis were also
described from the Lisowice site (Staneczko 2007). The dipterida
−
cean fern Clathropteris, a member of a generally Jurassic flora of
SE Asian origin, which entered Europe and Greenland in the
Rhaetian (Harris 1937; Lundblad 1950; Mader 1995; Czier 1998),
was reported from a carbonate facies which overlies the clastic
strata in this region (Roemer 1867), although Ash (2005) claimed
presence of Clathropteris in the early Norian of western North
America.
Among probable isoëtalean macrospores from Lisowice,
Trileites cf. T. pinguis (Harris, 1935) is common in the Rhaetian of
central Europe, although it is also known from the late Norian of
Germany and Poland. Horstisporites bertelseni Fuglewicz, 1977
has at least a close relative in the Rhaetian of the core Rødby N. 1
in southern Denmark (Fuglewicz 1977; Fuglewicz and Śnieżek
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Acta Palaeontologica Polonica 53 (4): 733–738, 2008
Brief report
Acta Palaeontol. Pol. 53 (4): 733–738, 2008

1980). Taken together, both macrofloral and palynological evi
−
dence indicates Rhaetian as the age of the Lisowice site.
Also suggestive of Rhaetian age are conchostracans of un
−
usually small size for the Norian of the Germanic Basin (1.5–3.0
mm), similar to Euestheria brodieana (Jones, 1862) from the
Rhaetian of England and United States (Kozur and Weems
2005; Heinz Kozur, personal communication 2007). A more
thorough study of the fossil assemblage is under way and will be
published separately.
Institutional abbreviations.—BGS, British Geological Survey,
Keyworth, Nottinghamshire, UK; BMNH, Museum of Natural
History, London, UK; ZPAL, Institute of Paleobiology of the
Polish Academy of Sciences, Warsaw, Poland.
Description
Dicynodont.—Partially articulated skeleton and numerous bones
of at least two individuals are dispersed in the claystone unit at the
exposure.
The left maxilla (ZPAL V33/85), represented by the postero
−
ventral edge lacking the tusk, displays a short ventral process simi
−
lar to that of Ischigualastia (Cox 1965). Articular condyles of the
right quadrate (ZPAL V33/84) suggest that the skull was approxi
−
mately 47 cm wide (across the quadrates), if its proportions were
similar to Ischigualastia.
Unlike other dicynodonts, the 49 cm long humerus (ZPAL
V33/96; Fig. 2C) lacks the entepicondylar foramen on the distal
head, and the distal end of the deltopectoral crest is bent anteriorly
to become narrow and rounded ventrally. The posterior part of the
humerus head is similar to proportions as that of Ischigualastia.
The distance between the deltopectoral crest and supinator process
is small and the shaft is only weakly twisted, like the condition in
Ischigualastia. The complete supinator process is prominent and
narrow. The merged capitulum and trochlea of the humerus are
smaller than in Ischigualastia, Placerias (Camp and Welles 1956),
and Stahleckeria.
The femur (ZPAL V33/75; Fig. 2D) is wider (in anterior view)
and generally more massive than in other Triassic dicynodonts,
apparently in connection with its unusually large size (56 cm
length). The lateral edge of the distal head is even more convex in
anterior view than in Wadiasaurus (Bandyopadhyay 1988),
Stahleckeria or Ischigualastia, and it is straight in Placerias
(Camp and Welles 1956) and Jachaleria (Vega−Dias and Szultz
2004). The spherical dorsomedially directed proximal head makes
it similar to Wadiasaurus, Stahleckeria,andPlacerias.InthePol−
ish form and Stahleckeria, the femur head is directed more ven−
trally than reconstructed in Placerias. Its trochanter major is the
longest of all these genera. The tibia (50 cm length) is massive, but
does not differ significantly from Stahleckeria and Placerias.The
fibulae (ZPAL V33/76 and ZPAL V33/77) are generally similar to
Stahleckeria, although the proximal head is narrower and gener
−
ally slender then in this genus. The straight mesial side of the distal
head (in anterior view) is like Placerias (it is strongly concave in
Stahleckeria).
Benton (2006) listed dicynodonts among the groups that
vanished in the mass extinction at the Carnian–Norian bound
−
ary. Jachaleria colorata Bonaparte, 1970 from the Los Colo
−
rados Formation of Argentina and Jachaleria candelariensis
Araújo and Gonzaga, 1980 from the Caturrita Formation of Rio
Grande do Sul, Brazil are considered to be the last dicynodonts
of middle Norian age (Langer 2005; Langer et al. 2007). The
new finding shows that Late Triassic dicynodonts survived at
least to the late Norian and possibly to the Rhaetian. Dicyno
−
donts apparently occurred in southern Poland for the whole Late
Triassic, as indicated by another occurrence under study at
Woźniki, 30 km E of Lisowice, where a medium−sized species
(ZPAL V. 34/1) is associated with a temnospondyl (ZPAL V.
34/28). The unexpectedly large size of the last Polish species
confirms the general evolutionary trend of increasing body size
in the Triassic (Ray 2006), a repetition of a similar succession in
734 ACTA PALAEONTOLOGICA POLONICA 53 (4), 2008
1m
Fig. 1. Exposure of fossiliferous strata of probable Rhaetian or late Norian
age in Lipie Śląskie clay−pit at Lisowice, southern Poland. A–C. Measured
sections at locations shown on blockdiagram (D). E. View of section C
from the SW. Abbreviations: Sl, siltstones; M, mudstones; Sa, sandstones.)

the Permian (terminated with Rhachiocephalus). Near the end
of the Triassic, dicynodonts achieved large rhino proportions
and became the dominant terrestrial herbivores in certain fau−
nas, as suggested by numerical dominance of dicynodont bones
and lack of any other large herbivore at Lisowice. Survival of
the dicynodonts to near the end of the Triassic adds also credi−
bility to the report of their occurrence in the Cretaceous of Aus−
tralia (Thulborn and Turner 2003).
Large theropod dinosaur.—Bones of a large predatory archo
−
saur were found as four accumulations in a 2−meter thick hori
−
zon at the Lisowice clay−pit, one representing the skull bones
and others with postcranial elements. They fit each other in size
and morphology and seem to represent the disarticulated skele
−
tons of two individuals.
The skull differs from “rauisuchians” in that the frontals
(ZPAL V33/21) form the anterior part of the supra−temporal fossa.
In the braincase (ZPAL V33/15), a sheet−like crista tuberalis is vis
−
ible, characteristic for theropods (Galton and Knoll 2006). It is
ventrally short, as in the coelophysoids (Raath 1984) whereas in
most theropods (e.g., Piatnitzkysaurus, Sinraptor,andDilopho
−
saurus), the crista tuberalis is very deep (see Rauhut 2004). A
lateroventrally directed bony sheet of the crista ventrolateralis is
developed in the posterior part of basisphenoid. The condition is
not as vertical as in more derived neotheropods (e.g., Dilopho
−
saurus and Allosaurus). Laterally, above the basisphenoid sinus, a
pneumatic recess, known also in Coelophysis rhodesiensis (Raath
1984) is visible. C. rhodesiensis, Ceratosaurus,andAllosaurus
have a very narrow depression, but in the neotheropods the de
−
pression is wide, although not as much as in the Polish form. The
base of the paroccipital process is higher than the dorsal rim of the
occipital condyle, a plesiomorphic condition compared to coelo−
physoids and Dilophosaurus (Raath 1984; Welles 1984).
The teeth (ZPAL V33/50 and ZPAL V33/51) show transverse
enamel wrinkles that sweep ventraly and away from the serrations
on the distal margins of the labial and lingual surfaces.
Like most theropod dinosaurs, the Lisowice form bears a lat−
eral ridge ventral to the tooth margin, which extends throughout
the length of the dentary (Fig. 2B), which makes it similar also
to the “megalosaurid” dentary (cast BMNH R2912, from holo
−
type natural sandstone mold BGS 6532) from the Rhaetian of
England (Galton 1998, 2005).
The location of the humeral greater tubercle relative to the
internal tuberosity is proximal, as in most theropods (Rauhut
2003). The humeral shaft (ZPAL V33/46) is longitudinally
twisted in the Lisowice dinosaur, unlike basal theropods, which
have nearly parallel proximal and distal articular surfaces. They
became increasingly rotated in more derived forms (e.g., teta
−
nurans, ceratosaurs).
The femur (ZPAL V33/45; Fig. 2A) is massive and elon
−
gated. In posterior view, tibiofibularis crest is relatively narrow
mediolaterally and its long axis is longitudinal, unlike the condi
−
tion in neoceratosaurs.
In the partially preserved pubis (ZPAL V33/220), the distal
blade is oriented almost vertically, and the obturator foramen is
open ventrally. In many tetanuran theropods, there is a tendency
towards reduction of the bony plate that encloses the foramen,
and it is not fully enclosed in bone in Allosaurus, Coelurus,
sinraptorids, Siamotyrannus, and carcharodontosaurids (Rauhut
2003). Because both the cranial material and the pelvis show
some dinosaur features we suggest that all these bones represent
a theropod species.
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BRIEF REPORT 735
10 cm
Fig. 2. Selected bones of large tetrapods from the Late Triassic of Lisowice, southern Poland. A, B. Theropod dinosaur. A. Left femur ZPAL V33/45 in lat
−
eral (A
1
) and anterior (A
2
) views. B. Right dentary ZPAL V33/25 in lateral view. C, D. Ischigualastia−like dicynodont. C. Left humerus ZPAL V33/96
in posterior (C
1
) and medial (C
2
) views. D. Left femur ZPAL V33/75 in anterior (D
1
) and medial (D
2
) views.

The new find shows that large theropods inhabited Laurasia as
early as the Late Triassic. Their skull was initially massive and
deep, with shortening haven taken place later, in the Jurassic. The
fossil record of large theropod dinosaurs was previously restricted
to the Early and Middle Jurassic sediments of North America, Eu
−
rope and Asia (Rauhut 2003; Allain et al. 2007). Large, about
40–45 cm long, tridactyl footprints (ZPAL V.33/219) were also
found at Lisowice. Morphology of those footprints are very simi
−
lar to ichnites from the Early and Middle Jurassic (Lockley and
Hunt, 1995; Lockley and Meyer 2000).
Small archosaurs.—Smaller cervical vertebrae with elongated
centra (ZPAL V33/41; ZPAL MB/1) are also known from
Lisowice, similar in morphology to Late Triassic coelophysoid
dinosaurs. In a sandstone intercalation near the top of the section
footprints left by a theropod of body size comparable with that
inferred from those vertebrae and digit proportions suggestive
of a coelophysoid has been found. The coelophysoid neothero
−
pods (e.g., Coelophysis, Liliensternus, and Lophostropheus)
were typical elements of the Norian and Rhaetian dinosaur com
−
munities in Europe (Rauhut and Hungerbühler 1998; Ezcurra
and Cuny 2007).
Isolated wing bones of a pterosaur (extremely elongated and
with a wide medullar cavity), and further small archosaur bones
represent other important elements of the assemblage.
Temnospondyls.—A large cyclotosaur is represented in the
Lisowice assemblage by the exoccipital (ZPAL V33/5), frag
−
ments of the maxilla (ZPAL V33/6 and ZPAL V33/7), squa
−
mosal with an edge of the closed otic notch (ZPAL V33/4), frag
−
ments of the skull roof, and the mandible (ZPAL V33/13). The
postcranial materials include a part of the clavicle (ZPAL
736 ACTA PALAEONTOLOGICA POLONICA 53 (4), 2008
1m
Fig. 3. Preliminary skeletal reconstructions of the basal theropod (A) and the Ischigualastia−like dicynodont (B) from the Late Triassic of Lisowice. Bones
represented in the collection are stippled.

V33/11), and a complete femur (ZPAL V33/1). Judging from
the mandible, this was one of the largest cyclotosaurs, with a
skull length of 75 cm, larger than its relative from the late
Carnian Krasiejów fauna in the area (Sulej and Majer 2005).
This material and undescribed fragments from the Rhaetian of
Germany (Schoch and Wild 1999) are the last capitosaurids in
the Germanic Basin.
Plagiosaurids are represented by the exoccipital (ZPAL
V33/193), dermal scutes (ZPAL V33/192) and a characteristic
vertebral centrum (ZPAL V33/194) with large parapophyses at
both ends.
Discussion
The composition of the Lisowice fossil assemblage is a curious
mixture of advanced elements (gymnosperm flora, dinosaurs, and
pterosaurs) with forms not expected to occur in latest Triassic
strata (giant dicynodonts and capitosaurs). Partial articulation of
the dicynodont (Fig. 2A
2
) skeleton and excellent preservation of
all fossils excludes redeposition of the assemblage. Regardless of
whether the strata are late Norian or early Rhaetian in age, the
dicynodont from Lisowice is the last unquestionable dicynodont
in the world with reliable stratigraphic origin and the only one
known from the Late Triassic of Europe.
Dicynodonts were living in the region also much earlier in
the Late Triassic, at the same time as the aetosaur−rauisuchian
dominated assemblage of Krasiejów (Dzik and Sulej 2007), as
proven by their occurrence at Woźniki, approximately 60 km
eastward and inland from Lisowice. It appears that in central Eu−
rope, quite different vertebrate communities inhabited different
continental ecosystems in the Late Triassic. Basal sauropodo−
morphs and coelophysoids in Württemberg (Rauhut and Hun−
gerbühler 1998; Moser 2003) were coeval with dicynodonts and
large and small theropods in Poland, as well as with aetosaurs,
the possible ornithischian relative dinosauromorph Silesaurus,
and rauisuchians (Dzik and Sulej 2007). They lived at the same
time in different environments, as indicated by associated floral
assemblages.
It is hard to decide with the available evidence whether the
Lisowice giant dicynodont was living close to water reservoirs
at the site where its remains are preserved, or was transported
from a dry nearby region. The alluvial sedimentary environment
of Lisowice does not preclude transport of the dicynodont and
theropod carcasses from dry inland environments with xero
−
phytic Hirmeriella−dominated flora, the source of large charcoal
stumps associated with bones.
The new unexpected discoveries of dicynodonts in the Late
Triassic of Poland nicely shows how misleading any local, or
even regional, faunal succession may be, if used to generalize
about the pattern of extinction (see also Irmis et al. 2007b).
Acknowledgements.—Robert Borzęcki (Warsaw, Poland), Piotr Men
−
ducki (Ostrowiec Świętokrzyski, Poland), and Marek Błyszcz (Lisowice,
Poland) made us aware of vertebrate bones occurring at Lisowice−Lipie
Śląskie. The excavations by Institute of Paleobiology PAN were sup
−
ported by a research grant from the Polish Ministry of Science and Infor
−
matisation No. 1665/P01/2007/32 and National Geographic Polska.
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Jerzy Dzik [dzik@twarda.pan.pl], Instytut Paleobiologii PAN, Twarda 51/55, PL−00−818 Warszawa, Poland and Instytut Zoologii Uniwersytetu
Warszawskiego, ul. Banacha 2, PL−02−097 Warszawa, Poland;
Tomasz Sulej [sulej@twarda.pan.pl], Instytut Paleobiologii PAN, ul. Twarda 51/55, PL−00−818 Warszawa, Poland;
Grzegorz Niedźwiedzki [gniedzwiedzki@biol.uw.edu.pl], Instytut Zoologii Uniwersytetu Warszawskiego, ul. Banacha 2, PL−02−079 Warszawa, Poland.
738 ACTA PALAEONTOLOGICA POLONICA 53 (4), 2008