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  • A Review of The Early Cretaceous Terrestrial Vertebrate Trackbearing Strata of England and Spain Part 1

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    Arturo Palma Ramírez
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    Paleontología

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    A Review of the Early Cretaceous Terrestrial Vertebrate Trackbearing Strata of England and Spain Part 1

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    89 tayangan4 halaman

    A Review of The Early Cretaceous Terrestrial Vertebrate Trackbearing Strata of England and Spain Part 1

    Diunggah oleh

    Arturo Palma Ramírez
    Paleontología

    Hak Cipta:

    © All Rights Reserved
    Unduh sebagai PDF atau baca online dari Scribd
    Tandai sebagai konten tidak pantas
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    cS GK Lan te, Wet, 15, Lonernd Miers Teer eames, Ne Maco Mcam Nt Hin nS Hub No A REVIEW OF THE EARLY CRETACEOUS TERRESTRIAL VERTEBRATE ‘TRACK-BEARING STRATA OF ENGLAND AND SPAIN JOANNA L. WRIGHT, P. M. BARRETT, MARTIN G. LOCKLEY? and E. COOK" "Department of Geology, University of Bristol, Wills Memorial Building, Queen's Road, Bristol, BSB IR), UNITED KINGDOM; ‘Department of Barth Sciences, Univers ‘of Cambridge, Dovning Steet, Cambridge, CB2 3EQ, UNITED KINGDOM; Department of Geology, University of Colorado at Denver, CO 80217-3368, USA ABSTRACT: Vertebrate footprints from the Lower Cretaccous formations of Europe have been known for almost 150 years. Recent discoveries, andl more rigorous study of tracks, have revealed a greater diversity of trackmakers than previously thought, including theropods, sauropods, various omithischians, pterosaurs, birds and turtles. ‘The pattern of the vertebrate trace and body fossil record varies with lithological unit and between regions, Very few dinosaur skeletal remains are known from the Purbeck Limestone Group, although microvertebrate remains are abundant, and the footprint record is one of the _most diverse and best documented in the country. Dinosaur skeletal remains from the Wealden ‘of both the Isle of Wight and the Weald are more abundant, although footprints are an important and diverse component. Vertebrate tracks from Farly Cretaceous rocks of Spain (Oneala and Enciso Groups) are morphologically similar to those from England, though they are somewhat ‘more diverse in terms of named ichnotaxa. The body fossil record from Spain indicates a diverse Early Cretaceous fauna, Early Cretaceous dinosaur-bearing strata of pre-Aptian age are uncommon worldwide. The Purbeck and Wealclen Groups of England and the Oncala and Enciso Groups of Spain therefore provide an important window into this transitional phase in the evolution of dinosaur faunas. ‘These lithological units reveal that a paleofauna of true Cretaceous aspect was already in place at the beginning of the Berriasian, indicating rapid faunal turnover atthe Jurassic-Cretaceous, 143 boundary. INTRODUCTION In 1851, §. H. Beckles published the first report of dinosaur footprints from the Barly Cretaceous sediments of the Wealden Group, southeastern England (Beckles, 1851), and the first description of footprints from the “Purbeck Marble" (= Purbeck Limestone Group) came three years later (Beckles, 1854), §. H, Beckles was the pioneer of dinosaur footprint studies, or “palaeolithichnology" as he termed it, in England. Despite ‘opposition from various parties, who denied that his omithoidichnites" were of organic origin, and inspired by the work of Edward Hitchcock in the United States (e., Hitchcock, 1858), Beckles eventually found, in the Wealden Beds near Hastings, several long trackways (more than 10 footprints) which proved beyond a doubt that these were the fossil footprints of some long extinct animal (Beckles, 18628). Since the mid- nineteenth century, ongoing exploration of the Purbeck Limestone Group, and of the Wealden Group of both the Weald and the Isle of Wight, has continued to reveal new occurrences of dinosaur footprints and trackways. The first long dinosaur trackways from the Purbeck Limestone Group were found in 1960 {(Charig and Newman, 1962); these were subsequently excavated and collected by the British Museum (Natural History). ‘The first Cretaccous dinosaur tracks from Spain were reported in the 1970s (Casanovas and Santafe, 1971, 1974). Since that time dozens of sites have been described (Moratalla, 1993; Moratalla et al, 1992a, b, 1993, 1994, 1995; Moratalla and Sanz, 1997), The majority of these are in the Cameros Basin, La Rioja Province, where a "Ruta de Ichnitas’ or “ichnological trail” and comprehensive guidebook have been created for the benefit of visitors (Moratalla et al., 1988). ‘The purpose of this brief paper is to review the dinosaur Jchnofauna from the Early Cretaceous of England and Spain, and to combine these data with information obtained from the skeletal record and from paleoenvironmental reconstructions, The Early Cretaceous was a time of enormous change in dinosaur communities, during which the _sauropod-dominated communities prevalent in the Late Jurassic were replaced by the ornithischian-dominated communities that prevailed in the late Farly and Late Cretaceous (Bakker, 1978). Early Cretaceous dinosaur-bearing strata of pre-Aptian age are very uncommon (Weishampel, 1992); the Purbeck and Wealden Groups of England and the Oncala and Encisco Groups of Spain provide a rare ‘window into this transitional phase, Additional data on non- dinosaurian trackways (pterosaurs, crocodilians, turtles, birds) are also provided. GEOLOGICAL OVERVIEW ‘The Purbeck and Wealden Groups of England were deposited during the Barly Cretaceous over a period of approximately 20 million years. The track-bearing strata of the Purbeck Limestone Group are Berriasian in age, while the overlying Wealden Group of the Weald and the Isle of Wight was deposited between the late Berriasian and the lowermost Aptian (Fig. 1). These sediments record a general regression from the marginal marine setting of the Purbeck Limestone Group to the low-lying alluvial floodplains of the Wealden, The Vectis Formation (Upper Wealden: Barremian) of the Isle of Wight shows signs of inereasing marine influence, heralding the transgression of the overlying Greensand Group. The Wealden Group represents the ‘youngest nonmarine Mesozoic sediments in England, ‘The Oncala-Eneiso Groups of Spain also span at least the Berriasian-Barremian interval, and may extend from near the 144 Kimmeridgian-Tithonian interval up into the Albian (Fig, 1),a total of a8 much as 50 million years, though this dating is somewhat uncertain. Most of the tracksites discussed here are confined to the Bersiasian-Barremian interval, thus facilitating comparison between Spain and England. The track-bearing strata from Spain were deposited under fluvial and lacustrine conditions (Alonso and Mas, 1998; Moralalla, 1993). (El-Shahat and West, 1983). Footprints are not known from the Lower Purbeck (Ensom, 1995) Most of the Early Cretaceous strata of the Purbeck Group were deposited in brackish water, in a subchumid and warm- temperate climate. Freshwater and fully marine environments are also represented, but such strata are rare (Arkell, 1947; Allen and Keith, 1965) The well-lithfied limestones in which footprints are ‘most commonly preserved were deposited on intertidal or supra- tidal flats which underwent early cementation in meteoric water, increasing the preservation potential ofthe footprints (West and El-Shahat, 1985; El-Shahat and West, 1983). ‘THE PURBECK LIMESTONE GROUP Sedimentology and Stratigraphy ‘The marginal marine Purbeck Limestone Group crops out in ‘asmall area on the south coast of England (Fig, 2) and consists of ‘icritic limestones and marls, The Purbeck Limestone Group is divided into two formations, the lower Lulworth Formation and the upper Dutlston Formation (Clements, 1993), but also into three traditional informal sections, the Lower, Middle and Upper Purbeck Beds (Fig. 1). The highest concentration of footprints in the Purbeck Limestone Group is found in the Intermarine Member ofthe Middle Purbeck Beds (Ensom, 1995; Wright, 1996), ‘The Intermarine Member combines a high proportion of well- lthified limestones, in which footprints have a high preservation potential, with a very shallow water to emergent depositional environment (El-Shahat and West, 1983), in which dinosaur footprints would have been easily produced. Members of the ‘Midille Puzbeck in which footprints are not found do not meet these criteria, Dinosaur footprints are much rarer in the Upper Purbeck (Ensom, 1995; Wright, 1996). Where present, they occur in similar lithologies to the footprints from the Middle Purbeck ‘Terrestrial Vertebrate Skeletal Remains ‘An incredibly diverse skeletal assemblage has been recorded from the Purbeck Limestone Group, including several dinosaur taxa and many other terrestrial tetrapods (EI-Shahat and West, 1983; Ensom, 1988; Ensom et al, 1991, 1994; Howse and Milner, 1995), Lissamphibians are represented by four taxa, including a frog, an albanerpetontid, and two salamanders (McGowan and Ensom, 1997). A number of small reptile remains have also been found, indicating the presence of two sphenodontids, an anguiomorph lizard, and numerous crocodilians and turtles. Dinosaur remains are extremely rare, though several taxa were present, including an indeterminate theropod, an indeterminate nodosaurid, and Iguanodon hoggi (Weishampel, 1992). Several genera of pterodactyloid—pterosaurs —_(Gnathosaurus, Plataleorhynchus, and *Ornithocheirus") have been identified on the basis of fragmentary material All major groups of mid-Mesozoie mammals are represented in the Purbeck Limestone Group, ENGLAND Generated Genoralzod FAW Sratgrapty Statgrapty 5 eS oO ALBIAN 7 omg =a Tio Base = r | faery | [thee 9 Lowe APTIAN JE se oC a BARREMIAN mn BARREMIAN g | = h ENCISO [ 3| wautenvan || 2) see OY hn) Havrerian 12 te Ly ronnoisnress — EEE 3 | G 3 URBION & | VALANGINIAN || 2 vy VALANGINIAN, ° Hastings There Woks o | BERAIASIAN || meang ina Ww af FaenniEe [x] BERRIASIAN : Cow 7 | "urbe a] ue NIAN 3) tHonan Paper — | payne 3) ceed fl] THO g 3 _ ¢}——_ : 2 KIMMERIDGIAN 3 | immeriocian’ | [IGURET, Comparative staligraphy ofthe vertebrate rack bearing Barly Cretaceous rods of Bngland and Spain. 145 — Tease Isle of Purbeck — 30%, FE] over Wealden FIGURE2, Simplified geology ofthe vertob including triconodonts, docodants, symmetrodonts, cladotheres, rmultituberculates, and tribosphenids (Ensom and Sigogneau- Russell, 1998). Despite this diversity, however, articulated skeletal remains have never been recovered from the Purbeck Limestone Group, and larger bone elements (> 25 mm in length) are rare. Indeed, the most abundant skeletal elements from the Purbeck are isolated teeth. There seems to be a considerable amount of size and resistance sorting, but no detailed research has yet been carried out to investigate this. Recently, a number of eggshell fragments were recovered from the Cherty Freshwater Member (Ensom, 1997) which oceurs in the lower part of the Middle Purbeck. Four different types of eggshell were identified; two of these have been tentatively associated with sauropods (Ensom, 1997). The most abundant type of eggshell at the site is thought to be crocodilian, suggestion which accords well with the abundant crocodile teeth found throughout the Purbeck Limestone Group. The other ‘eggshell types, which are much less common and all thought to bbe of dinosaurian origin, are suggested to have been transported in from elsewhere (Ensom, 1997). This suggestion may seem at ‘odds with the fragile nature of eggshells, but recent work (Tokaryk and Storer, 1991) has shown that eggshell may be transported considerable distances with minimal abrasion. ‘Terrestrial Vertebrate Tracks Teidactyl footprints found in the Purbeck Limestone Group have traditionally been attributed to Iguanodon (Ensom, 1995). It thas been recognized for many years, however, that some of these teidactyl footprints were made by large theropods (Wright, 1996). ‘This situation has been complicated by the use of invalid criteria in the differentiation of theropod and iguanodontian tracks, and some trackways originally ascribed to theropods were probably made by iguanodontians (Fig. 3). Revision of these trackway has confirmed that both theropod and iguanodontian footprints are present throughout the track-bearing strata of the Purbeck Limestone Group (Wright, 1996). In addition, quadrupedal ‘guanodontian trackways (Fig. 3) have recently been recognized in this unit (Wright, in press). (One of the most important footprint finds made in the area came in the summer of 1986, when a trackway was uncovered at Sunnydown Farm Quarry, in the lower part of the Middle twock-bearing Early Cretaceous rocks of Ragland, ‘Purbeck (Lulworth Formation), near Langton Matravers (Ensom, 1987a,b). This trackway was unusual in thatthe footprint casts of which it was composed were exceptionally large and tetradactyl (Fig. 3), and the trackway was that of a habitual quadruped, rather than a biped. The footprints were provisionally attributed to a sauropod (Ensom, 19872, 6), although subsequent examination indicates that they are more likely to be nodosaurid in origin (Wright, 1996). More recently (january, 1997), extremely large sauropod footprints have been discovered; these also occur in the Middle Purbeck, but higher in the sequence (Durlston Formation) (Wright, unpublished data). Pterosaur tracks (Purbeckopus) were frst found inthe Purbeck Group in 1936, but they were not recognized as tracks of any sort until they were briefly described by Delair (1963). Subsequently referred toas exocodilian (Lockley, 1989, 1991), their eal affinities were only realized recently (Wright et al, 1997) (Fig. 3). The Purbeck Limestone Group is the only lithological unit in Britain from which pterosaur tracks are currently known. ‘THE WEALDEN GROUP The Wealden Group crops out in two main areas in the UK-the Weald and the Isle of Wight (Fig. 2). Although the sediments in these two areas bear some relation to one another and can be at least partially correlated (Fig. 1), they were deposited in two different sub-basins, the Wessex sub-basin (where the Purbeck Group was also deposited) and the Weald

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