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Monography on Las Hoyas exceptional deposit

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 Las Hoyas:  A Cretaceous Wetland
9-2016

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Las Hoyas:
At 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
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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 A Cretaceous Wetland
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. A multidisciplinary synthesis after
Much or little was made in these 25 years, depending on how
one looks at it. Either way, this synthesis volume aims to con-
25 years of research on an exceptional
vey the story of our experience in the study of Las Hoyas. The fossil Lagerstätte from Spain
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
Francisco José Poyato-Ariza &
by providing the geological framework to understand how the Ángela D. Buscalioni
interpretation of this fossil Lagerstätte has changed from a deep
lake at first to the current hypo­thesis of a shallow aquatic environ- (editors)

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different groups of plants and animals recorded at Las Hoyas

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 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  cm
41 coloured and 1 b&w plates, 64 coloured and 101 b&w figures
(altogether 838 photos and drawings)
plus shipping charges: ˜ 15 to EU countries / ˜ 20 to other countries
Contributors (authors)  (p. 6)
Acknowledgements  (p. 8)
Preface: a bit of history  II.20b Dinosauria (non-avian Saurischia)  (Fernando Escaso,
(Francisco José Poyato-Ariza & Ángela D. Buscalioni)  (p. 9)
I Introduction
I.1 Relevance of Las Hoyas as a Mesozoic Lagerstätte  Lobón, Jingmai K. O’Connor, F. Ortega & F. Escaso)  (p. 183)
(Á. D. Buscalioni & F. J. Poyato-Ariza)  (p. 13)
I.2 Environmental reconstruction: a historical review  II.23 Ichnoassemblage (trace fossils)  (Jordi M. de Gibert, José J.
(Marian Fregenal-Martínez & Nieves Meléndez)  (p. 14)
II Fossil Record
II.1 Las Hoyas in the Tree of Life  (F. J. Poyato-Ariza,
H. Martín-Abad & Guillermo Navalón-Fernández)  (p. 29)
II.2 Palynomorphs  III.2 Anuran biostratinomy  (Álvaro López-García,
(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)
II.3 Plants and their landscapes
B This deep crest is slightly curved posteriorly. Such a condition
1 cm
1 cm
A D
G Tapejara (preserved parts
1 cm
F C, D. Concornis lacustris, holotype, MCCM-LH-1184. Prepared specimen (C) and fluorescence induced by means of ultraviolet light (D). Notice the preservation of
H apj
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).
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Contents
Fossil Record II.19 Pterosauria
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
5 mm
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).
is unusual among tapejarid pterosaurs.
Phylogenetic position
All 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).
1 cm
Four clades are generally recognized within the Pterodactylo-
idea: Ornithocheiroidea, Ctenochasma-
1 cm
toidea, Dsungaripteroidea, and
Azhdarchoidea (Fig. 4).
E C
5 mm
5 cm
A scd
pf scp l of the jugal; aprj, anterior
sq
po
naof
o to species found in wetlands only in a sporadic manner.
1 cm
j
ltf
1 cm
q
m temporal fenestra; m, max-
rap
I frontal; po, postorbital;
d Sanz et al. 2001). Photos G. F. Kurtz.
C rd, right dentary; rm, right
D interactions among water-dependant organisms (aquatic plus
scd
53
1 cm
A B
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,
hy lq rap
lsa
pt
ec
o
lj
lsq rm
lpo ltf
aprj
B in A. C. Reconstruction of
rd
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)
F. Ortega & José Luis Sanz)  (p. 177)
II.21 Aves  (J. L. Sanz, B. Chamero, Luis M. Chiappe, J. Marugán-
II.22 Feathers  (J. Marugán-Lobón & R. Vullo)  (p. 190) III.5 Exceptional preservation 
(F. J. Poyato-Ariza & Á. D. Buscalioni)  (p. 229)
Moratalla, M. Gabriela Mángano & Luis A. Buatois)  (p. 195) IV Palaeoecology
III Taphonomy IV.1 From Taphonomy to Palaeoecology 
III.1 Biostratinomic factors involved in fish preservation  (A. D. Buscalioni & F. J. Poyato-Ariza)  (p. 232)
(H. Martín-Abad & F. J. Poyato-Ariza)  (p. 202) IV.2 The wetland of Las Hoyas (Á. D. Buscalioni,
F. J. Poyato-Ariza, J. Marugán-Lobón, M. Fregenal-Martínez,
H. Martín-Abad & Óscar Cambra-Moo)  (p. 211) Óscar Sanisidro, G. Navalón & Carlos de Miguel)  (p. 238)
III.3 Molecular preservation  (Derek E. G. Briggs, Neal S. Gupta & Addendum 1:  New taxa and some latest findings  (p. 254)
Ó. Cambra-Moo)  (p. 216) Addendum 2:  Spinolestes  (p. 256)
III.4 Microbial mats and preservation (María del Carmen Guerrero, Appendix:  Systematic list  (p. 258)
Ana Isabel López-Archilla & Miguel Iniesto)  (p. 220) Added in proof  (p. 260)
Fossil Record II.12Palaeoecology
Osteichthyan fishes IV.2 The wetland
Fossil
of Record
Las Hoyas IV.2 The wetland of Las Hoyas
Fossil Record Palaeoecology II.21 Aves Palaeoecology
II.21 Aves IV.2 The wetland of Fossil
Las Hoyas
Record IV.2 The wetland of Las Hoyas Palaeoecology
1 cm
A 1 cm
A 5 cm
Fig. 2. Or nithocheirid
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.
C 1 cm
B 1 cm
A
B 1 cm
D 1 cm
C 5 mm
lm
Fig. 3. Skull of Europejara Plate 1. General view of
olcadesorum from Las the regional landscape
Hoyas (holotype, MCCM-
pursuing to highlight the interactions between organisms across fewer interactions with aquatic and amphibious nodes. The in the area of La Huér-
LH-9413) (after Vullo et al. aquatic, amphibious and terrestrial environments in order to test number of obligate aquatic plus amphibious vertices exceeds guina Formation during
the ecologic hypothesis of Las Hoyas as a wetland. Grouping the number of terrestrial vertices, which is more patent when
2012). A. Acid-prepared the late Barremian, some
ld
counter-slab. B. Interpre- G 5 mm E 1 cm D 1 cm
species intro ecological categories allows testing whether the considering animals only (Fig. 2). In terms of diversity (rather 125 million years ago. It
tative drawing of the skull expected ecological properties of wetland-type are present in than interactions), the terrestrial nodes are bigger because they is represented as a patchy
the interactions of the organisms from Las Hoyas. Therefore, are taxonomically more diverse. This is due to the substantial mosaic of environments
the skull based in part on plants are organized into two groups, aquatic and terrestrial. In diversity of terrestrial coleopterans, neuropterans, and plants. composed by ponds, small
turn, animal categories are based on their life cycles and their Such richness in terrestrial species is known, nonetheless, from lakes, channels, sloughs,
in grey). D. Life restoration dependence on wetland hydrology (van der Valk 2006), so they a relatively low abundance in specimens; that is, they are known and inundated plains.
of the head of Europejara are grouped into: 1) obligate aquatic: species found always from one to a few individuals per species, due to taphonomic Clouds represent a hu-
olcadesorum. Abbrevia- either in the water column or in flooded soils; 2) amphibious: conditions; these specimens appear in wet facies (Buscalioni mid and warm climate.
tions: apj, anterior process species considered to spend at least part of their life cycle in the & Poyato-Ariza, present volume). Landscape supervision by
M. Fregenal-Martínez (see
E 1 cm water and part in terrestrial environment, with life cycles linked Invertebrate stand out in the Las Hoyas food web; the
process of the right jugal; in general to shoreline trophic habits; and 3) terrestrial, being aquatic, amphibious, and terrestrial insect vertices bear the also Chapter I.2). Artwork
d, dentary; ec, ectopte- H 2 cm
by Ó. Sanisidro.
this category divided into facultative and incidental. Facultative maximal number of trophic connections. In turn, the trophic
rygoid; hy, hyoids; j, ju-
applies to species that may be found in wetlands and in other interactions of vertebrates are biased towards the obligate
gal; l, lachrymal; ld, left
terrestrial environments as well, including those that would aquatic and amphibious categories, which forage on micro-
dentary; lj, left jugal; lm,
left maxilla; lpo, left post-
prefer terrestrial environments for feeding; incidental applies invertebrates and small fish. The carnivorous tetrapods from
the terrestrial environment are relatively diverse, although, for
orbital; lq, left quadrate;
obvious evolutionary reasons, they lack the great diversity and
lsa, left surangular; lsp,
left squamosal; ltf, lower F 1 cm G 1 cm
Las Hoyas network disparity of birds, mammals, and snakes in Recent wetland
B ecosystems. Concerning aquatic-amphibious organisms, the
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). trophic connections of seven categories are to be highlighted:
illa; naof, nasoantorbital Plate 4. Life restorations of some terrestrial plants. A. The fern tree Weichselia reticulata, well known for possessing rigid pinnules with thick laminas. Its pinnules According to these premises, the trophic network of Las Hoyas
fenestra; o, orbit; pf, post- and second-order pinnae are arranged in a “butterfly position” that reduces the solar irradiance and excessive heating. B. The cheirolepidacean conifer Frenelopsis. (Fig. 2) shows the expected mixture of trophic relationships be- plants, plankton, worms, gastropods, insects, crustaceans,
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
and vertebrates (Fig. 3). Plankton-worms and aquatic plants
Anatomic supervision by B. Gomez (see also Chapter II.3). Artwork by Ó. Sanisidro. Hoyas fossil pellet, MCCM-LH-11386. Prepared (resin transferred) material (F), and scheme representing at least three recognizable specimens (G, modified from tween habitats typical of a wetland, resulting on blurry ecotones,
pt, pterygoid; q, quadrate; with a high variety of interconnected aquatic-to-terrestrial plants are the principal trophic resources in these realms. The pair
rap, retroarticular process; and animals. The trophic web is clearly biased towards the plankton-worms receive linkages from fishes, frog larvae, and
insects thus becoming a relevant source of matter and energy.
maxilla; scd, sagittal crest In turn, charophytes and aquatics angiosperms are the resources
amphibious). Trophic interactions are dense among the terres-
of the dentary; scp, scle- trial vertices, whereas terrestrial organisms have comparatively for crustaceans, gastropods, and aquatic insects. In addition,
ral plates; sq, squamosal. F 1 cm I 1 cm
171 125 245 185 241