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Las Hoyas: A Cretaceous WetlandAt human scale, 25 years seems time enough to relate academic and scientific experiences. The particular experiences presented in this book are based, first of all, on the earliest works and research at Las Hoyas led by Prof. José Luis Sanz. This book honors him as a tribute to those early years of Las Hoyas, where, while ad-miring the beauty of its fossils, we all learned the huge diversity of Nature and, at the same time, how to handle a number of enthusiastic students, manage the collection, and ask for funds year after year. The best of what we have all learnt from Prof. Sanz is the importance of a work that is properly done, and this sort of obsession will hound us forever.

Much or little was made in these 25 years, depending on how one looks at it. Either way, this synthesis volume aims to con-vey the story of our experience in the study of Las Hoyas. The present book is aimed to both specialists and informed aficio-nados. It incorporates many international researchers who have largely contributed to the understanding of the palaeobiology and taphonomy of this locality. The volume is organized into four major parts. Part I is devoted to the Las Hoyas environment by providing the geological framework to understand how the interpretation of this fossil Lagerstätte has changed from a deep lake at first to the current hypo thesis of a shallow aquatic environ-ment made of a mosaic of pools, filled by microbial mats, in a wetland-like landscape. Part II unfolds the palaeobiology of the different groups of plants and animals recorded at Las Hoyas by providing an updated revision of their taxonomic diversity. The information provided for each group is presented in a ho-mogeneous manner by developing a summarized description of the specimens, their life styles and ecomorphology, phylogenetic context, preservation, and relevance of the Las Hoyas record within a worldwide or European frame. The processes involved in the fossil preservation are explored in Part III. This part includes not only the study of the preservation of the fossils themselves, but also some chapters devoted to experimental taphonomy and the role of microbial mats in preservation. Part IV incorporates our experience in the fieldwork in establishing how palaeo-ecological restorations need of a previous taphonomic study. The final chapter presents restorations by some brilliant young artists of the landscape and organisms living in the Las Hoyas Wetland during the Early Cretaceous in the framework of their trophic network. An Addendum incorporates taxa discovered or described during the elaboration of the present volume, and a final Appendix presents an updated taxonomic classification of the biota from Las Hoyas.

Las Hoyas: A Cretaceous Wetland

A multidisciplinary synthesis after 25 years of research on an exceptional

fossil Lagerstätte from Spain

Francisco José Poyato-Ariza & Ángela D. Buscalioni

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Verlag Dr. Friedrich Pfeil · München

Las Hoyas: A Cretaceous Wetland

A multidisciplinary synthesis after 25 years of research on an exceptional

fossil Lagerstätte from Spain

Francisco José Poyato-Ariza & Ángela D. Buscalioni

(editors)

Las Hoyas: A Cretaceous Wetland

A multidisciplinary synthesis after 25 years of research on an exceptional fossil Lagerstätte from Spain

Francisco José Poyato-Ariza & Ángela D. Buscalioni (editors)

262 pages, 32.6 × 24.5 cm41 coloured and 1 b&w plates, 64 coloured and 101 b&w figures

(altogether 838 photos and drawings)

ISBN 978-3-89937-153-6

75 Euro plus shipping charges: ˜ 15 to EU countries / ˜ 20 to other countries

Contents

Contributors (authors) (p. 6)Acknowledgements (p. 8)Preface: a bit of history (Francisco José Poyato-Ariza & Ángela D. Buscalioni) (p. 9)I IntroductionI.1 Relevance of Las Hoyas as a Mesozoic Lagerstätte

(Á. D. Buscalioni & F. J. Poyato-Ariza) (p. 13)I.2 Environmental reconstruction: a historical review

(Marian Fregenal-Martínez & Nieves Meléndez) (p. 14)II Fossil RecordII.1 Las Hoyas in the Tree of Life (F. J. Poyato-Ariza,

H. Martín-Abad & Guillermo Navalón-Fernández) (p. 29)II.2 Palynomorphs

(Montserrat de la Fuente & Reinhard Zetter) (p. 31)II.3 Plants and their landscapes (Carles Martín-Closas,

Bernard Gomez & Véronique Daviero-Gomez) (p. 43)II.4 Mollusca (Graciela Delvene & Martin Clive Munt) (p. 57)II.5 Arachnida (Paul A. Selden) (p. 64)

III.5 Exceptional preservation (F. J. Poyato-Ariza & Á. D. Buscalioni) (p. 229)

IV PalaeoecologyIV.1 From Taphonomy to Palaeoecology

(A. D. Buscalioni & F. J. Poyato-Ariza) (p. 232)IV.2 The wetland of Las Hoyas (Á. D. Buscalioni,

F. J. Poyato-Ariza, J. Marugán-Lobón, M. Fregenal-Martínez, Óscar Sanisidro, G. Navalón & Carlos de Miguel) (p. 238)

Addendum 1: New taxa and some latest findings (p. 254)Addendum 2: Spinolestes (p. 256)Appendix: Systematic list (p. 258)Added in proof (p. 260)

II.6 Diplopoda (P. A. Selden & William A. Shear) (p. 68)II.7 Insecta (Xavier Delclòs & Carmen Soriano) (p. 70)II.8 Ostracoda (Julio Rodríguez-Lázaro) (p. 89)II.9 Spelaeogriphacea (Damiá Jaume, Eva Pinardo-Moya &

Geoff A. Boxshall) (p. 94)II.10 Decapoda (Alessandro Garassino) (p. 98)II.11 Chondrichthyes (Rodrigo Soler-Gijón, F. J. Poyato-Ariza,

John G. Maisey & Jennifer A. Lane) (p. 103)II.12 Osteichthyan fishes

(F. J. Poyato-Ariza & H. Martín-Abad) (p. 114)II.13 Albanerpetontidae (Susan E. Evans) (p. 133)II.14 Urodela (S. E. Evans) (p. 138)II.15 Salientia (Ana María Báez) (p. 143)II.16 Chelonia (Adán Pérez-García, Marcelo S. de la Fuente &

Francisco Ortega) (p. 151)II.17 Squamata (S. E. Evans & Arnau Bolet) (p. 156)II.18 Crocodylomorpha

(A. D. Buscalioni & Beatriz Chamero) (p. 162)II.19 Pterosauria (Romain Vullo & Jesús Marugán-Lobón) (p. 170)II.20a Dinosauria (Ornithischia) (Mercedes Llandres Serrano,

R. Vullo & F. Ortega) (p. 175)II.20b Dinosauria (non-avian Saurischia) (Fernando Escaso,

F. Ortega & José Luis Sanz) (p. 177)II.21 Aves (J. L. Sanz, B. Chamero, Luis M. Chiappe, J. Marugán-

Lobón, Jingmai K. O’Connor, F. Ortega & F. Escaso) (p. 183)II.22 Feathers (J. Marugán-Lobón & R. Vullo) (p. 190)II.23 Ichnoassemblage (trace fossils) (Jordi M. de Gibert, José J.

Moratalla, M. Gabriela Mángano & Luis A. Buatois) (p. 195)III TaphonomyIII.1 Biostratinomic factors involved in fish preservation

(H. Martín-Abad & F. J. Poyato-Ariza) (p. 202)III.2 Anuran biostratinomy (Álvaro López-García,

H. Martín-Abad & Óscar Cambra-Moo) (p. 211)III.3 Molecular preservation (Derek E. G. Briggs, Neal S. Gupta &

Ó. Cambra-Moo) (p. 216)III.4 Microbial mats and preservation (María del Carmen Guerrero,

Ana Isabel López-Archilla & Miguel Iniesto) (p. 220)

241

pursuing to highlight the interactions between organisms across aquatic, amphibious and terrestrial environments in order to test the ecologic hypothesis of Las Hoyas as a wetland. Grouping species intro ecological categories allows testing whether the expected ecological properties of wetland-type are present in the interactions of the organisms from Las Hoyas. Therefore, plants are organized into two groups, aquatic and terrestrial. In turn, animal categories are based on their life cycles and their dependence on wetland hydrology (van der Valk 2006), so they are grouped into: 1) obligate aquatic: species found always either in the water column or in flooded soils; 2) amphibious: species considered to spend at least part of their life cycle in the water and part in terrestrial environment, with life cycles linked in general to shoreline trophic habits; and 3) terrestrial, being this category divided into facultative and incidental. Facultative applies to species that may be found in wetlands and in other terrestrial environments as well, including those that would prefer terrestrial environments for feeding; incidental applies to species found in wetlands only in a sporadic manner.

Las Hoyas networkAccording to these premises, the trophic network of Las Hoyas (Fig. 2) shows the expected mixture of trophic relationships be-tween habitats typical of a wetland, resulting on blurry ecotones, with a high variety of interconnected aquatic-to-terrestrial plants and animals. The trophic web is clearly biased towards the interactions among water-dependant organisms (aquatic plus amphibious). Trophic interactions are dense among the terres-trial vertices, whereas terrestrial organisms have comparatively

fewer interactions with aquatic and amphibious nodes. The number of obligate aquatic plus amphibious vertices exceeds the number of terrestrial vertices, which is more patent when considering animals only (Fig. 2). In terms of diversity (rather than interactions), the terrestrial nodes are bigger because they are taxonomically more diverse. This is due to the substantial diversity of terrestrial coleopterans, neuropterans, and plants. Such richness in terrestrial species is known, nonetheless, from a relatively low abundance in specimens; that is, they are known from one to a few individuals per species, due to taphonomic conditions; these specimens appear in wet facies (Buscalioni & Poyato-Ariza, present volume). Invertebrate stand out in the Las Hoyas food web; the aquatic, amphibious, and terrestrial insect vertices bear the maximal number of trophic connections. In turn, the trophic interactions of vertebrates are biased towards the obligate aquatic and amphibious categories, which forage on micro-invertebrates and small fish. The carnivorous tetrapods from the terrestrial environment are relatively diverse, although, for obvious evolutionary reasons, they lack the great diversity and disparity of birds, mammals, and snakes in Recent wetland ecosystems. Concerning aquatic-amphibious organisms, the trophic connections of seven categories are to be highlighted: plants, plankton, worms, gastropods, insects, crustaceans, and vertebrates (Fig. 3). Plankton-worms and aquatic plants are the principal trophic resources in these realms. The pair plankton-worms receive linkages from fishes, frog larvae, and insects thus becoming a relevant source of matter and energy. In turn, charophytes and aquatics angiosperms are the resources for crustaceans, gastropods, and aquatic insects. In addition,

Plate 1. General view of the regional landscape in the area of La Huér-guina Formation during the late Barremian, some 125 million years ago. It is represented as a patchy mosaic of environments composed by ponds, small lakes, channels, sloughs, and inundated plains. Clouds represent a hu-mid and warm climate. Landscape supervision by M. Fregenal-Martínez (see also Chapter I.2). Artwork by Ó. Sanisidro.

IV.2 The wetland of Las Hoyas Palaeoecology

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Plate 3. Angiosperms from Las Hoyas. A, B. Montsechia vidalii. A. Branched shoot with leaves in opposite decussate phyllotaxis (MCCM-LH-26463a). B. Shoot with fruit attached (MCCM-LH-26811b). C. Ranunculus ferreri, axis showing swollen bodies and slender, dichotomized, branched filaments (MCCM-LH-15416a). D-E. Proteae-phyllum sp., part and counterpart showing a swollen petiole and a reniform lamina with a reticulate venation; specimen photographed in field before labelling. F-G. Cf. Jixia (now Iterophyllum, see Addendum 1, Plate 1A); trilobate lamina showing pinnate venation and tapered laminar base. F. MCCM-LH-26740b. G, H. Specimen MCCM-LH-26873, part and counterpart. I. Cf. Ficophyllum, apical lamina of a simple, oblong pinnately-veined eudicot leaf with festooned brochidodromous major secondary veins. Specimen MCCM-LH-18600. Photos J. A. Gracia, courtesy of the editors (except D, E, and G, photos B. Gomez).

II.3 Plants and their landscapes Fossil Record

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II.12 Osteichthyan fishes Fossil Record

245

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BPlate 4. Life restorations of some terrestrial plants. A. The fern tree Weichselia reticulata, well known for possessing rigid pinnules with thick laminas. Its pinnules and second-order pinnae are arranged in a “butterfly position” that reduces the solar irradiance and excessive heating. B. The cheirolepidacean conifer Frenelopsis. Anatomic supervision by B. Gomez (see also Chapter II.3). Artwork by Ó. Sanisidro.

IV.2 The wetland of Las Hoyas Palaeoecology

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Fig. 2. Ornithocheirid teeth from Las Hoyas (af-ter Vullo et al. 2009a). A. MCCM-LH-17264. B. MCCM-LH-21451. The grey area (drawing) repre-sents the enamelled part of the crown.

rd

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Fig. 3. Skull of Europejara olcadesorum from Las Hoyas (holotype, MCCM-LH-9413) (after Vullo et al. 2012). A. Acid-prepared counter-slab. B. Interpre-tative drawing of the skull in A. C. Reconstruction of the skull based in part on Tapejara (preserved parts in grey). D. Life restoration of the head of Europejara olcadesorum. Abbrevia-tions: apj, anterior process of the jugal; aprj, anterior process of the right jugal; d, dentary; ec, ecto pte-rygoid; hy, hyoids; j, ju-gal; l, lachrymal; ld, left dentary; lj, left jugal; lm, left maxilla; lpo, left post-orbital; lq, left quadrate; lsa, left surangular; lsp, left squamosal; ltf, lower temporal fenestra; m, max-illa; naof, nasoantorbital fenestra; o, orbit; pf, post-frontal; po, postorbital; pt, pterygoid; q, quadrate; rap, retroarticular process; rd, right dentary; rm, right maxilla; scd, sagittal crest of the dentary; scp, scle-ral plates; sq, squamosal.

The small teeth MCCM-LH-15448 and MCCM-LH-28572 from Las Hoyas are very similar to those of Haopterus (Wang & Lü 2001, Wang & Zhou 2006). This ornithocheiroid genus of unclear affinities was originally described as belonging to the Pterodactylidae (Wang & Lü 2001) and later considered to be the basalmost member of the Ornithocheiridae (Unwin 2003). Finally, the most recent studies conclude that Haopterus is either a basal Istiodactylidae (Lü et al. 2008, 2010) or the sister-group of this clade (Witton 2012). MCCM-LH-15448 and MCCM-LH-28572 are here tentatively assigned to this family. Considering it is closely related to Haopterus, the istiodactylid from Las Hoyas would correspond to a form basal to all other istiodactylids (i. e., Istiodactylus, Liaoxipterus, Longshengpterus, Nurhachius, and possibly Hongshanopterus). The Istiodactylidae may represent a basal group within the clade Ornithocheiroidea. About the second ornithocheiroid taxon present at Las Hoyas,

is present on the whole surface of the crown. It consists of slight, irregular, longitudinal folds. Some very slight growth folds, concave toward the apex, are present near the base. A poorly marked distal carina is present on the enamelled part of the crown. The mesial edge is rounded and devoid of a carina. The apex shows a small wear facet oriented distally. As suggested by its morphology, this tooth probably occupied an anterior position in the jaw. The holotype (MCCM-LH-9413) and only known specimen of Europejara olcadesorum corresponds to an incomplete skull and lower jaw, partially articulated, and preserved in a slab and counter-slab (Fig. 3, Plate 1). The skull is about 240 mm in pre-served length. The skull is crushed and flattened dorsoventrally, while the mandible is preserved in lateral view. The dorsal part and the rostral end are missing. The palate, exposed in dorsal view, shows two narrow and elongate choanae which are sepa-rated by a thin vomer. The crushed posterior part displays some bones of the left side of the skull (e. g., the postorbital process of the jugal, the postorbital, the quadrate, and the squamosal). The upper and lower jaws are edentulous. The dentary shows a very thin cortical bone and a typical trabecular structure. The rostral end of the mandible is slightly turned ventrally (~ 10 degrees). Europejara olcadesorum is mainly characterized by a well-developed bony sagittal crest in the anterior portion of the dentary (at least twice the height of the dentary ramus). This deep crest is slightly curved posteriorly. Such a condition is unusual among tapejarid pterosaurs.

Phylogenetic positionAll the forms that have been found at Las Hoyas belong to the suborder Pterodactyloidea (short-tailed pterosaurs). This group first appeared in the Late Jurassic and became largely dominant during the whole of the Cretaceous (Lü et al. 2010). Four clades are generally recognized within the Pterodactylo-

idea: Ornithocheiroidea, Ctenochasma-toidea, Dsungaripteroidea, and

Azhdarchoidea (Fig. 4).

II.19 Pterosauria Fossil Record

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Plate 1. Birds from Las Hoyas. A, B. Iberomesornis romerali, holotype, MCCM-LH-22. Prepared specimen (A) and fluorescence induced by means of ultraviolet light (B). C, D. Concornis lacustris, holotype, MCCM-LH-1184. Prepared specimen (C) and fluorescence induced by means of ultraviolet light (D). Notice the preservation of feathers on the left wing lost in the preparation process. E. Eoalulavis hoyasi, MCCM-LH-13500a,b. Original specimen (slab before transfer preparation). F, G. Las Hoyas fossil pellet, MCCM-LH-11386. Prepared (resin transferred) material (F), and scheme representing at least three recognizable specimens (G, modified from Sanz et al. 2001). Photos G. F. Kurtz.

II.21 Aves Fossil Record