It is generally accepted that during the Triassic the composition 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 unknown 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 sediments at Lisowice (Lipie Śląskie clay-pit) in southern Poland. 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 carnivorous dinosaurs, pterosaurs, as well as capitosaur and plagiosaur amphibians. Dicynodonts were represented in the Germanic 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 appears, thus, that the fossil record of tetrapod succession in the Late Triassic was strongly controlled by ecological factors and biased by uneven representation of particular environments. The Lisowice assemblage proves that faunas dominated by dicynodonts did not entirely disappear at least until the end of the Triassic.
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 lenticular body of grey mudstone and clay stone in the northern part of the Lipie Śląskie clay-pit, mostly covered with calcareous and pyritic crust or preserved within limestone concretions. The vertebrate 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 reddish mudstones of the Keuper occur, lithostratigraphically correlatable 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 apparently supported two ovules. Unlike S. septentrionalis, as interpreted 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 probable gingkoalean pollen Monosulcites cf. M. minimus Cookson, 1947 (leaves—ZPAL V33/215 similar to Schmeissneria are associated) 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 dipteridacean 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 1980). Taken together, both macrofloral and palynological evidence indicates Rhaetian as the age of the Lisowice site.
Also suggestive of Rhaetian age are conchostracans of unusually 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.
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 similar to that of Ischigualastia (Cox 1965). Articular condyles of the right quadrate (ZPAL V33/84) suggest that the skull was approximately 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, and Placerias. In the Polish form and Stahleckeria, the femur head is directed more ventrally 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 generally 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 boundary. Jachaleria colorata Bonaparte, 1970 from the Los Colorados 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. Dicynodonts 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 the Permian (terminated with Rhachiocephalus). Near the end of the Triassic, dicynodonts achieved large rhino proportions and became the dominant terrestrial herbivores in certain faunas, 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 credibility to the report of their occurrence in the Cretaceous of Australia (Thulborn and Turner 2003).
Large theropod dinosaur.—Bones of a large predatory archosaur were found as four accumulations in a 2-meter thick horizon 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 skeletons of two individuals.
The skull differs from “rauisuchians” in that the frontals (ZPAL V3 3/21) form the anterior part of the supra-temporal fossa. In the braincase (ZPAL V33/15), a sheet-like crista tuberalis is visible, 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, and Dilophosaurus), 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., Dilophosaurus and Allosaurus). Laterally, above the basisphenoid sinus, a pneumatic recess, known also in Coelophysis rhodesiensis (Raath 1984) is visible. C. rhodesiensis, Ceratosaurus, and Allosaurus have a very narrow depression, but in the neotheropods the depression 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 coelophysoids 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 lateral 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 holotype 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., tetanurans, ceratosaurs).
The femur (ZPAL V33/45; Fig. 2A) is massive and elongated. In posterior view, tibiofibularis crest is relatively narrow mediolaterally and its long axis is longitudinal, unlike the condition 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.
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, Europe 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 similar 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 neotheropods (e.g., Coelophysis, Liliensternus, and Lophostropheus) were typical elements of the Norian and Rhaetian dinosaur communities 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), fragments of the maxilla (ZPAL V33/6 and ZPAL V33/7), squamosal with an edge of the closed otic notch (ZPAL V33/4), fragments of the skull roof, and the mandible (ZPAL V33/13). The postcranial materials include a part of the clavicle (ZPAL 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.
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. 2A2) 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 Europe, quite different vertebrate communities inhabited different continental ecosystems in the Late Triassic. Basal sauropodomorphs and coelophysoids in Württemberg (Rauhut and Hungerbühler 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 xerophytic Hirmeriella-dommated 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).
Robert Borzęcki (Warsaw, Poland), Piotr Menducki (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 supported by a research grant from the Polish Ministry of Science and Informatisation No. 1665/P01/2007/32 and National Geographic Polska.
- R. Allain , R. Tykoski , N. Aquesbi , N.E. Jalil , M. Monbaron , D. Russell , and P. Taquet 2007. An abelisaurid (Dinosauria: Theropoda) from the Early Jurassic of the High Atlas Mountains, Morocco, and the radiation of ceratosaurs. Journal of Vertebrate Paleontology 27: 610–624. Google Scholar
- S. Arndt 2002. Morphologie und Systematik ausgewahlter Mesozoischer Koniferen. Palaeontographica B 262: 1–23. Google Scholar
- S.R. Ash 2005. Synopsis of the Upper Triassic flora of Petrified Forest National Park and vicinity. Mesa Southwest Museum Bulletin 9: 53–62. Google Scholar
- S. Bandyopadhyay 1988. A Kannemeyeriid dicynodont from the Middle Triassic Yerrapalli formation. Philosophical Transactions of the Royal Society of London. B 320: 185–233. Google Scholar
- M.J. Benton 1994. Late Triassic to Middle Jurassic extinctions among continental tetrapods: testing the pattern. In : N.C. Fraser and H.-D. Sues (eds.), In the Shadow of the Dinosaurs , 366–397. Cambridge University Press, Cambridge. Google Scholar
- M.J. Benton 2006. The origin of the dinosaurs. In : Actos de III Jornadas Internacionales sobre Paleontología de Dinosaurios y su Entorno, 11–19. Colectivo Arqueológico-Paleontológico de Salas, Burgos. Google Scholar
- R.J. Butler , L.B. Porro , and A.B. Heckert 2006. A supposed heterodontosaurid tooth from the Rhaetian of Switzerland and a reassessment of the European Late Triassic record of Ornithischia (Dinosauria). Neues Jahrbuch fur Geologie und Palaontologie Monatschefte 2006: 613–633. Google Scholar
- C.L. Camp and S.P. Welles 1956. The North American genus Placerias. Memoirs of the Univeristy of California. 13: 255–304. Google Scholar
- J.A. Clement-Westerhof and J.H.A. Van Konijnenburg-Van Cittert 1991. New data on the fertile organs leading to a revised concept of the Cheirolepidiaceae. Review of Palaeobotany and Palynology 68: 147–179. Google Scholar
- C.B. Cox 1965. New Triassic dicynodonts from South America, their origins and relationships. Philosophical Transactions of the Royal Society of London B 248: 457–516. Google Scholar
- Z. Czier 1998. Gingko foliage from the Jurassic of the Carpathian Basin. Palaeontology 41: 349–381. Google Scholar
- J. Dzik 2001. A new Paleorhinus fauna in the early Late Triassic of Poland. Journal of Vertebrate Paleontology 21: 625–627. Google Scholar
- J. Dzik 2003. A beaked herbivorous archosaur with dinosaur affinities from the early Late Triassic of Poland. Journal of Vertebrate Paleontology 23: 556–574. Google Scholar
- J. Dzik and T. Sulej 2007. A review of the early Late Triassic Krasiejów biota from Silesia, Poland. Palaeontologia Polonica 64: 3–27. Google Scholar
- J. Dzik , G. Niedźwiedzki , and T. Sulej 2008. Zaskakujące uwieńczenie ery gadów ssakokształtnych. Ewolucja 3: 2–21. Google Scholar
- J. Dzik , T. Sulej , A. Kaim , and R. Niedźwiedzki 2000. Późnotriasowe cmentarzysko kręgowców lądowych w Krasiejowie na Śląsku Opolskim. Przegląd Geologiczny 48: 226–235. Google Scholar
- M.D. Ezcurra and G. Cuny 2007. The coelophysoid Lophostropheus airelensis, gen. nov.: a review of the systematics of “Liliensternus” airelensis from the Triassic—Jurassic boundary outcrops of Normandy (France). Journal of Vertebrate Paleontology 27: 73–86. Google Scholar
- R. Fuglewicz 1977. New species of megaspores from the Trias of Poland. Acta Palaeontologica Polonica 22: 405–431. Google Scholar
- R. Fuglewicz and P. Śnieżek 1980. Upper Triassic megaspores from Lipie Śląskie near Lubliniec. Przegląd Geologiczny 28: 459–461. Google Scholar
- P.M. Galton 1998. Saurischian dinosaurs from the Upper Triassic of England: Camelotia (Prosauropoda, Melanorosauridae) and Avalonianus (Theropoda, ?Carnosauria). Palaeontographica A 250: 155–172. Google Scholar
- P.M. Galton 2005. Bones of large dinosaurs (Prosauropoda and Stegosauria) from the Rhaetic Bone Bed (Upper Triassic) of Aust Cliff, southwest England. Revue de Paléobiologie, Genève 24: 51–74. Google Scholar
- P. M. Galton and F. Knoll 2006. A saurischian dinosaur braincase from the Middle Jurassic (Bathonian) near Oxford, England: from the theropod Megalosaurus or the sauropod Cetiosaurus? Geological Magazine 143: 905–921. Google Scholar
- T.M. Harris 1931. Rhaetic floras. Biological Reviews 6: 133–162. Google Scholar
- T.M. Harris 1932. The fossil flora of Scoresby Sound East Greenland. Part 4: Gingkoales, Coniferales, Lycopodiales and isolated fructifications. Meddelelser om Grønland 112: 1–12. Google Scholar
- T.M. Harris 1937. The fossil Flora of Scoresby Sound East Greenland. Part 5. Stratigraphic relations of the plant beds. Meddelelser om Grønland 112: 82–86. Google Scholar
- A.P. Hunt , V.L. Santucci , M.G. Lockley , and T.J. Olsen 1993. Dicynodont trackways from the Holbrook Member of the Moenkopi Formation (Middle Triassic: Early Anisian), of Arizona, USA. New Mexico Museum of Natural History & Science Bulletin 3: 213–218. Google Scholar
- R.B. Irmis , W.G. Parker , S.J. Nesbitt , and J. Liu 2007a. Early ornithischian dinosaurs: the Triassic record. Historical Biology 19: 3–22. Google Scholar
- R.B. Irmis , S.J. Nesbitt , K. Padian , N. Smith , A. Turner , D. Woody , and A. Downs 2007b. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science 317: 358–361. Google Scholar
- H.W. Kozur and R.E. Weems 2005. Conchostracan evidence for a late Rhaetian to early Hettangian age for the CAMP volcanic event in the Newark Supergroup, and a Sevatian (late Norian) age for the immediately underlying beds. Hallesches Jahrbuch für Geowissenschaften B 27: 21–51. Google Scholar
- M.C. Langer 2005. Studies on continental Late Triassic tetrapod biochronology. I. The type locality of Saturnalia tupiniquim and the faunal succession in south Brazil. Journal of South American Earth Sciences 19: 205–218. Google Scholar
- M.C. Langer , A. M. Ribeiro , C.L. Schultz , and J. Ferigolo 2007. The continental tetrapod-bearing Triassic of south Brazil. In : S.G. Lucas and J.A. Spielmann (eds.), The Global Triassic. New Mexico Museum of Natural History and Science Bulletin 41: 201–218. Google Scholar
- M.G. Lockley and A.P. Hunt 1995. Dinosaur Tracks and Other Fossil Footprints of the Western United States. 338 pp. Columbia University Press, New York. Google Scholar
- M.G. Lockley and C.A. Meyer 2000. Dinosaur Tracks and Other Fossil Footprints of Europe. 323 pp. Columbia University Press, New York. Google Scholar
- B. Lundblad 1950. Studies in the Rhaeto-Liassic floras of Sweden 1. Kungliga Svenska Vetenskapsakademiens Handlingar 1: 2–82. Google Scholar
- D. Mader 1995. Taphonomy, Sedimentology and Genesis of Plant Fossil Deposit Types in Lettenkohle (Lower Keuper) and Schilfsandstein (Middle Keuper) in Lower Franconia (Germany). 164 pp. Peter Lang, Frankfurt am Main. Google Scholar
- M. Moser 2003. Plateosaurus engelhardti Meyer, 1837 (Dinosauria: Sauropodomorpha) aus dem Feuerletten (Mittelkeuper; Obertrias) von Bayern. Zitteliana B 24: 1–186. Google Scholar
- G. Niedźwiedzki and T. Sulej 2008. Lipie Śląskie koło Lisowic — okno na późnotriasowy ekosystem lądowy. Przegląd Geologiczny 56: 821–822. Google Scholar
- T. Orłowska-Zwolińska 1983. Palinostratygrafia epikontynentalnych osadów wyzszego triasu w Polsce. Prace Instytutu Geologicznego 104: 1–88. Google Scholar
- M.A. Raath 1984. The Theropod Syntarsus and Its Bearing on the Origin of Birds. In : M.K. Hecht , J.H. Ostrom , G. Viohl , and P. Wellnhofer (eds.), The Beginnings of Birds , 219–228. Fruende des Jura-Museums, Eichstatt. Google Scholar
- O.W.M. Rauhut 2003. The interrelationships and evolution of basal thero-pod dinosaurs. Special Papers in Palaeontology 69: 1–213. Google Scholar
- O.W.M. Rauhut 2004. Braincase structure of the Middle Jurassic theropod dinosaur Piatnitzkysaurus. Canadian Journal of Earth Sciences 41: 1109–1122. Google Scholar
- O.W.M. Rauhut and A. Hungerbühler 1998. A review of European Triassic theropods. Gaia 15: 75–88. Google Scholar
- S. Ray 2006. Functional and evolutionary aspects of the postcranial anatomy of dicynodonts (Synapsida, Therapsida). Palaeontology 49: 1263–1286. Google Scholar
- M. Reymanówna 1992. Two conifers from the Liassic flora of Odrowaz in Poland. In : J. Kovar-Eder (ed.), Palaeovegetational Development in Europe and Regions Relevant to its Palaeofloristic Evolution, Proceedings of the Pan-European Palaeobotanical Conference, Vienna, 19–23 September 1991, 307–311. Museum of Natural History, Vienna. Google Scholar
- F. Roemer 1867. Neuere Beobachtungen über die Gliederung des Keupers und der ihn zunnächst überlagernden Abtheilung der Juraformation in Oberschlesien und in den angrenzenden Theilen von Polen. Zeitschrift der Deutschen Geologischen Gesselschaft 14: 255–269. Google Scholar
- R. Schoch and R. Wild 1999. Die Wirbeltier-Fauna im Keuper von Süddeutschland. In : N. Hauschke and V. Wilde (ed.), Trias, 395–408. Verlag Dr. Friedrich Pfeil, München. Google Scholar
- E. Schultz and C. Heunisch 2005. Palynostratigraphische Gliederungsmöglichkeiten des deutschen Keupers. In : G. Beutler (ed.), Stratigraphie von Deutschland. IV Keuper. Courier Forschungsinstitut Senckenberg 253: 43–49. Google Scholar
- K. Staneczko 2007. Nowe dane paleobotaniczne na temat górnego triasu z Lipia Śląskiego koło Lublińca (południowa Polska). Geo-Sympozjum Mlodych Badaczy Silesia 2007: 155–168. Google Scholar
- W.N. Stewart and G.W. Rothwell 1993. Paleobotany and the Evolution of Plants. 535 pp. Cambridge University Press, Cambridge. Google Scholar
- T. Sulej 2005. A new rauisuchian reptile (Diapsida: Archosauria) from the Late Triassic of Poland. Journal of Vertebrate Paleontology 25: 75–83. Google Scholar
- T. Sulej and D. Majer 2005. The temnospondyl amphibian Cyclotosaurus from the Late Triassic of Poland. Palaeontology 48: 157–170. Google Scholar
- J. szulc , M. Gradziński , A. Lewandowska , and C. Heunisch 2006. The Upper Triassic crenogenic limestones in Upper Silesia (southern Poland) and their paleoenvironmental context. In : A.M. Alonso-Zarza and L.H. Tanner (eds.), Paleoenvironmental Record and Applications of Calcretes and Palustrine Carbonates: Geological Society of American Special Paper 416: 133–151. Google Scholar
- T. Thulborn and S. Turner 2003. The last dicynodont: An Australian Cretaceous relict. Proceedings of the Royal Society of London: Biological Sciences 270: 985–993. Google Scholar
- J.A. Townrow 1967. On Rissikia and Mataia podocarpaceous conifers from the Lower Mesozoic of southern lands. Papers and Proceedings of the Royal Society of Tasmania 101: 103–136. Google Scholar
- S.P. Welles 1984. Dilophosaurus wetherilli (Dinosauria: Theropoda) osteology and comparisons. Palaeontographica, Abteilung A: Paläozoologie, Stratigraphie 185: 85–180. Google Scholar
- C. Vega-Dias and C.L. Schultz 2004. Postcranial material of Jachaleria candelariensis Araujo and Gonzaga 1980 (Therapsida, Dicynodontia), Upper Triassic of Rio Grande do Sul, Brazil. PaleoBios 24: 7–31. Google Scholar