EAVP_2022_Abstract volume
EAVP_2022_Abstract volume
In collaboration with:
Yurac Diaz M., Salazar C., Meyer C.A., Suárez M.E, Belvedere M. 2022. The dinosaur
ichnological record of northern Chile: a review and its potential development. In:
Belvedere M., Mecozzi B., Amore O., Sardella R (eds.). Abstract book of the XIX Annual
Conference of the European Association of Vertebrate Palaeontologists,
Benevento/Pietraroja, Italy, 27th June-2nd July 2022. PalaeoVertebrata, Special Volume 1-
2022, 224. Doi: 10.18563/pv.eavp2022
ii
Index
Dal Sasso and CELLULAR-LEVEL PRESERVATION IN THE SOFT TISSUES OF THE THEROPOD
1
Maganuco DINOSAUR SCIPIONYX SAMNITICUS: AN OVERVIEW
THE BODY MASS OF PALUDOTONA (LAGOMORPHA, MAMMALIA), THE LAST STEM
Angelone et al. 3
LAGOMORPH (TUSCO-SARDINIA PALAEOBIOPROVINCE, LATE MIOCENE)
INTEREST OF ACCURATE SPECIES IDENTIFICATION IN FOSSIL RECORD, THE
Arbez et al. 5
STORY OF LEMMINGS
PRELIMINARY STUDY OF THE RODENTS ASSEMBLAGES FROM THE LATE
Badea et al. 6
MIOCENE OF MOȘNA 1 (MOLDAVIAN PLATFORM – ROMANIA)
CHALLENGES AND OPPORTUNITIES FOR INTEGRATING TRAIT DATA ACROSS THE
Balk et al. 7
TEMPORAL DIVIDE
iii
NEW VERTEBRATE REMAINS FROM THE JURASSIC-CRETACEOUS TRANSITION OF
Castanera et al. 34
THE EASTERN CAMEROS BASIN (SPAIN)
Chakravorti et DEFORMATION MECHANISM AND RETRODEFORMATION OF INDIAN RHYTIDOSTEID
35
al. TEMNOSPONDYL RESULTS IN A NEW COMBINATION OF BRAZILIAN TAXON
A NEW, HIGH-SPEED PROTOCOL TO RUN MUSCLE-DRIVEN BIOMECHANICAL
Chatar 37
SIMULATIONS THROUGH FINITE ELEMENT ANALYSES
A COMPUTER-AIDED PROTOCOL FOR THE VIRTUAL RECONSTRUCTION OF
Cirilli et al. THEROPOD DINOSAUR SKULLS. TYRANNOSAURUS REX AS AN ICONIC CASE 38
STUDY
Ciudad Real et NEUROANATOMY OF GROSSIUS, AN EARLY PREDATORY LOBE-FINNED FISH
40
al. FROM THE DEVONIAN OF SPAIN
E UNUM PLURIBUS – THE FIRST HIGH-DIVERSITY KOGAIONID (MAMMALIA,
Csiki-Sava et
MULTITUBERCULATA) ASSEMBLAGE FROM THE UPPERMOST CRETACEOUS OF 42
al.
THE TRANSYLVANIAN AREA, ROMANIA
RANGE SIZE AND EXTINCTION RISK OF CUBAN MAMMALS: A CONSERVATION
Dädlow et al. 43
PALEOBIOLOGY PERSPECTIVE
Danisi and
BRINGING THE DEVIL BACK HOME 44
Carotenuto
Darlim and TIME-SCALING OF CROCODYLIAN EVOLUTION AND THE CONTRIBUTION OF TOTAL
45
Rabi EVIDENCE TIP-DATING ANALYSIS
PHYLOGENETIC SIGNAL OF CRANIAL CHARACTERS IN CLADISTIC ANALYSIS OF
Dederichs et al. 46
DIPLODOCOID SAUROPOD DINOSAURS
Delsett et al. THE HYOID ARCH AND FEEDING IN TWO FISH-SHAPED MARINE TETRAPODS 48
Ferreira et al. A GIANT FRESHWATER TURTLE FROM THE PLEISTOCENE OF BRAZILIAN AMAZON 63
Fistola et al. A ROOM OF CIRO'S OWN - BUILDING A HYBRID DIGITAL PALEO SPACE 70
iv
HOW DOES A GIANT INSULAR LIZARD FEED? A 3D FINITE ELEMENT MODEL OF
Fortuny et al. 71
GALLOTIA STEHLINI AND ITS IMPLICATIONS FOR FOSSIL GIANT LIZARDS
García Zamora THICK TAILS! NON-AVIAN DINOSAUR CAUDOFEMORAL MUSCLE ESTIMATION FROM
74
et al. CT-SCAN OF EXTANT TAXA
BIOMECHANICAL AND GROWTH REMODELLING DIFFERENCES BETWEEN MODERN
Genochio et al. 75
HUMANS AND NEANDERTHALS
Ghezzo et al. BIAS IN EXTINCT LION RECORD AFFECTS OUR UNDERSTANDING OF THE SPECIES 77
v
Kynigopoulou THE PHYLOGENY OF THE EARLY PALAEOCENE ORDER TAENIODONTA USING NEW
103
et al. SPECIMENS FROM NEW MEXICO, USA
FIGHTER OR BLUFFER: COMPARISON OF BENDING AND COMPRESSION IN
Lategano et al. MODELS OF THE CAUDAL SPINES OF DACENTRURINE AND STEGOSAURINE 104
STEGOSAURS
SIZE REDUCTION AND ITS MORPHOLOGICAL CONSEQUENCES IN MAMMOTH
Lister 105
MOLARS FROM THE PLEISTOCENE OF BRITAIN
THE MIDDLE PERMIAN TRACKSITE OF GONFARON: A COMPLEX ECOSYSTEM IN
Logghe et al. 106
THE MIDST OF THE END-GUADALUPIAN MASS EXTINCTION
THE FIRST DUCKBILL DINOSAUR (HADROSAURIDAE: LAMBEOSAURINAE) FROM
Longrich AFRICA AND THE ROLE OF OCEANIC DISPERSAL FOR LATE CRETACEOUS AND 108
PALEOGENE VERTEBRATE BIOGEOGRAPHY
NEW INSIGHTS INTO TRICERATOPS LIMB BONE HISTOLOGY:
Lucassen et al. 109
REVEALING SKELETAL UNITY THROUGH PATTERNS IN HISTOVARIABILITY
Macaluso et al. REVIEW AND COMPLETENESS OF THE FOSSIL RECORD OF PALEARCTIC CAUDATA 110
Madurell- HOMOTHERIUM FROM THE LATE EARLY PLEISTOCENE OF PIRRO NORD (APULIA,
111
Malapeira et al. SOUTHEASTERN ITALY)
ISOLATED THEROPOD TEETH FROM THE TENDAGURU FORMATION (UPPER
Malafaia et al. 112
JURASSIC, TANZANIA)
THE BROMACKER LOCALITY: A REFERENCE SITE FOR PALAEOZOIC TETRAPOD
Marchetti et al. 114
ICHNOLOGY
Marcos-
FILLING CRACKS TO IMPROVE THE CONSERVATION OF DINOSAUR EGGS FROM
Fernández et 116
POYOS (UPPER CRETACEOUS. SPAIN)
al.
NEW PRIMATE MATERIAL FROM THE MIDDLE EOCENE FOSSIL SITE OF CAENES
Marigó et al. 117
(DUERO BASIN, SALAMANCA, SPAIN)
vi
Mondanaro et
A NEW METHOD TO MODEL RARE SPECIES BY RELYING ON PHYLOGENY 136
al.
Moya-Costa et THE EARLY- MIDDLE PLEISTOCENE SORICIDAE (MAMMALIA) OF THE IBERIAN
137
al PENINSULA
BITE TRACES, A POWERFUL TOOL IN PALAEOECOLOGY: THE CASE OF THE
Mujal et al. 139
MIDDLE TRIASSIC LOWER KEUPER FOSSIL LAGERSTÄTTEN
PALAEOENVIRONMENTS AND TAPHONOMY OF THE KUPFERZELL VERTEBRATE
Mujal et al. 141
LAGERSTÄTTE (MIDDLE TRIASSIC, S GERMANY)
A RE-EVALUATION OF THE NOTOSUCHIAN CROCODYLIFORM EREMOSUCHUS
Nicholl et al. ELKOHOLICUS FROM THE EARLY EOCENE OF NORTH AFRICA AND THE 143
PHYLOGENETIC RELATIONSHIPS OF SEBECOSUCHIANS
Oliver and
Carro- PHYLOGENETICAL ISSUES WITHIN THE SUPERFAMILY CTENODACTYLOIDEA 144
Rodríguez
ON THE DIAGNOSIS OF IBEROSUCHUS MACRODON (CROCODYLIFORMES,
Ortega et al. 145
NOTOSUCHIA) FROM THE EOCENE OF PORTUGAL
NEW MATERIAL OF THEROPODS (ABELISAUROIDEA?) FROM LO HUECO (LATE
Ortega et al. 147
CRETACEOUS. CUENCA, CENTRAL SPAIN)
LIGHTS AND SHADOWS IN THE EVOLUTIONARY DYNAMICS OF THE PLEISTOCENE
Palombo et al. 148
SARDINIAN MAMMOTH POPULATIONS
Pandolfi and THE ENDEMIC GIRAFFID FROM THE OREOPITHECUS-FAUNAL ASSEMBLAGES OF
149
Rook THE TUSCO-SARDINIAN ARCHIPELAGO (LATE MIOCENE, ITALY)
MORPHOLOGICAL CONVERGENCE IN THE HINDLIMB SKELETON WITHIN
Páramo et al. 150
TITANOSAURIA
Pardo-Pérez et AN ICHTHYOSAUR BREEDING GROUND IN SOUTHERNMOST GONDWANA
151
al. (TYNDALL GLACIER, SOUTHERN CHILE)
Parparousi et LATEST EARLY PLEISTOCENE DAMA-LIKE CERVID REMAINS FROM CAL
153
al. GUARDIOLA AND THE TANGLE OF VILLAFRANCHIAN CERVID TAXONOMY
LIMB DEVELOPMENT IN EARLY TETRAPODS: INSIGHTS FROM
Perez-Ben 154
MACROEVOLUTIONARY PATTERNS OF LIMB PROPORTIONS
A NEW MEMBER OF ERYMNOCHELYINI (TESTUDINES, PLEURODIRA) FROM THE
Pérez-García 155
EARLY PLIOCENE OF CENTRAL AFRICA
Pérez-García
CRANIAL ANATOMY OF THE IBERIAN CENOMANIAN BOTHREMYDID TURTLE
and Martín- 156
ALGORACHELUS PEREGRINA
Jiménez
Pérez-García et THE HOLOTYPE OF HYLAEOCHELYS KAPPA IS NO LONGER THE ONLY KNOWN
157
al. SPECIMEN FOR THIS TURTLE GENUS IN THE JURASSIC RECORD
GIANT SHADOWS IN LATE TRIASSIC SEAS: HISTOLOGICAL ANALYSIS ON PUTATIVE
Perillo et al. 158
AND GENUINE GIANT ICHTHYOSAURS BONES
Pittman et al. FEET REFINE THE ECOLOGY OF EARLY THEROPOD FLYERS 159
vii
STABLE ISOTOPE RECORD OF TRICERATOPS FROM A MASS ACCUMULATION
De Rooij et al. (LANCE FORMATION, WYOMING, USA) PROVIDES INSIGHTS INTO TRICERATOPS 168
BEHAVIOUR AND ECOLOGY
A BASAL ANKYLOPOLLEXIAN DINOSAUR FROM THE LATE JURASSIC OF
Rotatori et al. 170
PORTUGAL AND ITS IMPLICATIONS FOR IGUANODONTIAN DIVERSITY
3D LIFE RECONSTRUCTION OF THE INTRASPECIFICAL VARIABILITY IN THE FOSSIL
Ruiz Pedraza et
GIRAFFID DECENNATHERIUM REX RÍOS, SÁNCHEZ AND MORALES 2017 (LATE 172
al.
MIOCENE, IBERIAN PENINSULA)
THE LARGE, THE SMALL AND THE UNUSUAL – A NEW, MULTISPECIFIC
Rytel et al. 173
TANYSTROPHEID ASSEMBLAGE FROM SOUTHERN POLAND
174
Sacco et al. A NEW DIGITAL TOOL FOR STUDYING SMALL TETRAPOD FOOTPRINTS 174
Surmik et al. NEOTAPHONOMICAL OBSERVATION OF A BONE BED FORMATION IN A RIVER BED 199
viii
LONG BONE HISTOLOGY OF AETOSAURS AND PHYTOSAURS FROM KRASIEJÓW
Teschner et al. (NORIAN) REVEALS STRONG ENVIRONMENTAL INFLUENCE ON THE GROWTH 204
PATTERN
TROPHIC NETWORK RECONSTRUCTION AND ANALYSIS OF THE DIFFERENT
Torchet et al. FOSSILIFEROUS LAYERS OF THE MIDDLE TRIASSIC MONTE SAN GIORGIO WORLD 205
HERITAGE (SWITZERLAND/ITALY)
PALEOECOLOGICAL CHARACTERISTICS OF PLATYOPOSAURUS (AMPHIBIA,
Uliakhin 206
TEMNOSPONDYLI) FROM THE MIDDLE PERMIAN OF EASTERN EUROPE
PAPIONIN AFFINITIES IN THE ENDOCRANIAL ANATOMY OF CERCOPITHECOIDES
Urciuoli et al. 207
WILLIAMSI CALVARIA KA195
SKELETAL STRESS IN 3D SIMULATIONS OF HIGH-SPEED LOCOMOTION IN LARGE
van Bijlert et al. 209
THEROPODS
VIRTUAL ENDOCASTS OF THE BRAIN AND INNER EAR OF LEPTICTIS (LEPTICTIDA:
van Gent et al. 211
MAMMALIA)
van Rijssen et FLY PUPARIA IN A MAMMOTH SKULL FROM THE NETHERLANDS: IMPLICATIONS
212
al. FOR MAMMOTH TAPHONOMY
NETWORK MODELING AND CAPTURE-MARK-RECAPTURE METHODS FOR THE
Varvarà et al. 213
IDENTIFICATION OF SPECIES FACTORIES
THE LATE CAMPANIAN MICROVERTEBRATE ASSEMBLAGE FROM PETREȘTI
Vasile et al. (TRANSYLVANIAN BASIN, ROMANIA), A NEW WINDOW INTO THE EVOLUTION OF 214
EUROPEAN CRETACEOUS INSULAR FAUNAS
A NEW VERTEBRATE ASSEMBLAGE FROM THE LATE PLIOCENE OF MĂRU (DACIAN
Vasile et al. 216
BASIN, SOUTHERN ROMANIA)
SHAKE A LEG: VARIABILITY ON PELVIC MUSCULATURE OF THE TITANOSAURS
Vidal et al. 218
FROM LO HUECO (LATE CRETACEOUS, SPAIN)
QUANTIFYING BONE SURFACE ROUGHNESS IN BRANTA CANADENSIS USING 3D
Vintges et al. DIGITAL MICROSCOPY: IMPLICATIONS FOR TEXTURAL AGEING OF EXTANT AND 220
EXTINCT SPECIES
GLOBAL PHYLOGENY OF DIPLOCYNODONTINAE AND ITS IMPLICATIONS FOR
Walter et al. 221
CROCODYLIAN EVOLUTION
Yurac Diaz et THE DINOSAUR ICHNOLOGICAL RECORD OF NORTHERN CHILE: A REVIEW AND ITS
223
al. POTENTIAL DEVELOPMENT
ix
CELLULAR-LEVEL PRESERVATION IN THE SOFT TISSUES OF THE
THEROPOD DINOSAUR SCIPIONYX SAMNITICUS: AN OVERVIEW
1Sezione di Paleontologia dei Vertebrati, Museo di Storia Naturale di Milano, Corso Venezia 55,
20121, Milano, Italy.
2Associazione Paleontologica Paleoartistica Italiana, Via Nicolò Paganini 19, 43121, Parma, Italy.
In 1998 the baby theropod Scipionyx samniticus, from the Lower Cretaceous of Pietraroia
(Benevento Province, southern Italy), became famous worldwide thanks to its unique preservation
of internal soft tissues. Subsequent detailed studies demonstrated that in this 110-million-year-old
fossil the soft tissues are not simply imprints but are mineralized in three dimensions, and that their
preservation is exceptional even at a cellular, and at some points subcellular, level.
On the other hand, we demonstrated that the remains or imprints purported by some authors to be
of the “diaphragmatic muscles” are, in fact, a calcite nodule of amorphous microstructure,
inconsistent with the preservation of other muscle tissue in this fossil. This evidence denies the
hypothesis of a hepatic-piston assisted breathing mechanism in Scipionyx.
Outstandingly, the degree of preservation of the soft tissues also permitted to map the relative
position of the food remains in the digestive apparatus and thus to reconstruct a feeding
chronology, an insight that is usually impossible to obtain from fossils. Scipionyx’s guts contain
1
bones from a lepidosaurian reptile in the stomach region, lizard-like polygonal squamae in the
duodenum, fish scales in the rectum, and a variety of tiny remains in several points of the intestine.
This is evidence that Scipionyx fed on both lizards and fish, and confirms that in theropod
dinosaurs the partially undigested bones were guided through the entire intestine, eventually being
incorporated within the faeces.
The exceptional preservation of labile soft tissue indicates that, after death, the carcass of this
theropod hatchling was subjected to very little decay and rapid authigenic mineralisation. The
highest taphonomic threshold is represented by substrate microfabrics, that occur where soft
tissues have been phosphatised directly by very small apatite crystallites, replicating details at
subcellular level. The amount and detail of information gained from this single specimen make the
Pietraroja Plattenkalk a unique fossil Lagerstätte. In spite of the more frequent finds in China of
dinosaurs with integumentary structures, the preservation of such a variety of internal soft tissues
revealed by Scipionyx samniticus remains a unique in the fossil record.
2
THE BODY MASS OF PALUDOTONA (LAGOMORPHA, MAMMALIA), THE
LAST STEM LAGOMORPH (TUSCO-SARDINIA PALAEOBIOPROVINCE,
LATE MIOCENE)
1,2,3 4 1,5 6
C. Angelone *, S. Čermák , B. Moncunill-Solé , L. Rook
1Dipartimento di Scienze, Università degli Studi Roma Tre, Largo S. Murialdo 1, 00146 Roma,
Italy.
2Institut Català de Paleontologia Miquel Crusafont (ICP), Carrer de les Columnes s/n, 08193
Cerdanyola del Vallès, Spain.
3Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and
Paleoanthropology, Chinese Academy of Sciences, 142 Xi Zhi Men Wai Da Jie, 100044 Beijing, China.
4Institute of Geology of the Czech Academy of Sciences, Rozvojová 269, CZ-16500 Prague 6,
Czech Republic.
5Departamento de Física e Ciencias da Terra, Universidade da Coruña, Campus da Zapateira,
Rúa da Fraga 10, E-15008 A Coruña, Spain.
6Dipartimento di Scienze della Terra, Paleo[Fab]Lab, Università degli Studi di Firenze, Via La Pira
4, I-50121 Firenze, Italy.
Keywords: Insular endemism, refugium, gigantism, ghost lineage, Mediterranean islands, Turolian
The genus Paludotona (Lagomorpha, Mammalia) is an insular taxon endemic of the Tusco-
Sardinia palaeobioprovince (~8.3-6.7 Ma). Paludotona is the last representative of stem
lagomorphs, a heterogeneous basal group of early lagomorphs, and its last record (6.7 Ma)
postpones of 2.5 Ma the LAD of this stem group. Thus, during the late Miocene, Paludotona can
be considered a living fossil. Since how long Paludotona was thriving in its insular refugium is
unknown: fossils record less than 15% of its temporal distribution range, as there is evidence that
the lineage developed in isolation sensu lato for at least 14 Ma (ghost lineage). Paludotona
acquired a gigantic size with respect to continental ancestors, in accordance with the pattern
followed by small mammals in insular environments. Its body mass (BM) is estimated in ~1100 g,
which is three to five times the BM of MP28-MN1 stem lagomorphs of continental Europe, the
group from which Paludotona likely stemmed. The reasons for such an enormous BM increase are
multiple and interconnected. Some of them likely are synecological factors s.l., lower extrinsic
mortality, changes in size and degree/pattern of fragmentation of the area of the insular
palaeobioprovince, and successive climate changes (which caused a rapid evolutionary pulse
followed by a relative stasis following the pattern of Mein’s biphasic model). Through BM
estimation and morphological observation, we observed that Paludotona probably increased its
3
lifespan (though, at present, this datum is not quantitatively determined), and we suspect a change
in posture and locomotion caused by the noticeable BM increase.
4
INTEREST OF ACCURATE SPECIES IDENTIFICATION IN FOSSIL RECORD,
THE STORY OF LEMMINGS
3 Department of Zoology, Faculty of Science, Charles University, Vinicna 7, 128 44 Praha 2, Czech
Republic.
Taxonomic identification is the cornerstone of all investigations regarding past faunal communities
or ecosystem and, more generally, fossil biodiversity. For ancient time period, systematics is only
based on morphology, due to the absence of preserved DNA in fossil specimens. However,
similarity between closely related species and intra-specific morphological variability make
identification a difficult task, and can lead to spurious classification of fossils. Accurate
identification is crucial in periods of major environmental changes implying important faunal
turnovers and rapid species evolution, such as the climatic cycles during the Pleistocene.
This study proposes a global revision of the status of an emblematic species complex of European
glacial environments, the Lemmings (Lemmus sp., Myopus sp.). Their dental morphological
similarity makes them difficult to identify in the fossil record, and it was considered that the only
genus present in Europe during Pleistocene was the True Lemming (Lemmus sp.). The
reinvestigation of Late Pleistocene fossil specimens using geometric morphometrics demonstrated
that Myopus sp. was also a member of the Pleistocene European faunas. These results question
the identification of other Lemming specimens in Europe, as well as the relevance of the
taxonomic status of previously identified fossil species, such as Lemmus kowalski.
To avoid a typological description and in order to characterize the variability of morphological
characters with statistical confidence, we emphasize the need of using geometric morphometrics,
in taxonomic identification. The re-investigation of fossil specimens using such methods is
undoubtedly necessary to have a more accurate understanding of past faunal communities and
thus environmental reconstructions.
5
PRELIMINARY STUDY OF THE RODENTS ASSEMBLAGES FROM THE LATE
MIOCENE OF MOȘNA 1 (MOLDAVIAN PLATFORM – ROMANIA)
1Faculty of Geography and Geology, “Alexandru Ioan Cuza” University, Bulevard “Carol I”, Nr.11,
707006, Iași, Romania.
Keywords: micromammals, rodents, Late Vallesian, Late Miocene, Moldavian Platform, Romania.
The micromammals assemblage from one Late Miocene locality (Moșna 1) of the Moldavian
Platform present the occurrences of new specimens of rodents. This locality is located south of Iași
county, in the northeast part of Romania. Fossils were collected from at least 2.5 tons of sediment
that was wet sieved on a set of stable sieves. Important fossil fragments have been identified in
these sediments, including micromammals teeth. The rodents assemblage contains 25 identifiable
teeth belonging to eight taxa: Spermophilinus bredai, Progonomys hispanicus, Byzantinia sp.,
Neocricetodon moldavicum, Pliospalax sp., Lophocricetus sp., Glirulus lissiensis, Muscardinus sp.
The age of the assemblage is interpreted as Late Vallesian, Late Miocene. Moreover, Byzantinia ,
Neocricetodon, Lophocricetus, and Pliospalax are for the first time described in the Late Miocene
of Romania. This rodents assemblage represent an important contribution to the study of
micromammals from the Late Miocene of Moldavian Platform and also for interpreting
paleoenvironment.
6
CHALLENGES AND OPPORTUNITIES FOR INTEGRATING TRAIT DATA
ACROSS THE TEMPORAL DIVIDE
1National Ecological Observatory Network, Battelle Memorial Institute, Boulder, CO, United States
of America.
2University of Arizona, Tucson, AZ, United States of America.
3Florida Museum of Natural History, University of Florida, Gainesville, FL, United States of
America.
4University of California, Berkeley, CA, United States of America.
5Howard University, Washington, DC, United States of America.
6University of Oregon, Eugene, OR, United States of America.
Keywords: functional traits, climate change, evolution, ontology, linked open data
Functional traits are the measurable features of organisms that directly interact with the
environment. Quantifying changes in these traits is key to understanding how climate and
anthropogenic change affects species. An historical perspective can illuminate changes in intensity
of change for animal populations. Zooarchaeological data, representing early to modern human
occupation periods, provide a bridge from paleontological and prehuman data, to modern data.
However, paleontological and zooarchaeological trait data are based on skeletal metrics rather
than soft tissues, include additional long-tail data such as contextual and temporal data, and are
often curated in discipline specific data repositories, hindering discoverability of datasets across
the temporal divide. Further, gathering trait data that cuts across time requires much onus on the
researcher to standardize disparate datasets. However, modern, paleontological and
zooarchaeological datasets can be linked based on locality, and zooarchaeological and
paleontological datasets can be temporally linked as well. Preserving the necessary metadata and
aligning discipline-specific standards to make these linkages is a challenge. We present the
Functional Trait Resource for Environmental Studies (FuTRES), a data store that aggregates and
standardizes trait data from paleo, archaeo, and modern specimens, making such data findable
and searchable. We show the utility and potential of FuTRES in accelerating science across time,
and present a tool to help multidisciplinary researchers standardize their datasets for trait based
research.
7
3D CRANIAL RECONSTRUCTIONS OF THE ENIGMATIC
THALATTOSAURIFORMES
3University of Alaska Museum, University of Alaska Fairbanks, 1962 Yukon Drive, AK 99775,
Fairbanks, United States of America.
4Department of Paleobiology, National Museum of Natural History, 10th St. & Constitution Avenue,
DC 20560, Washington DC, United States of America.
5Institute for Photon Science and Synchrotron Radiation (IPS), Karlsruhe Institute of Technology
(KIT), Hermann-von-Helmholtz-Platz 1, 76344, Eggenstein-Leopoldshafen, Germany.
8
We show that the crania of thalattosauroid thalattosauriforms are highly akinetic, with intricate
interlocking and extensive bony contacts between skull elements. These likely helped
thalattosauriform skulls sustain large stresses related to the durophagous aspects in their diets. In
addition, Thalattosaurus displays a set of unique dental characteristic including variation in tooth
orientation, degree of enamel covering and root dimensions. Overall, the morphological variability
of dentigerous elements is much greater than previously assumed as illustrated by a thorough
sampling of North American thalattosauroid thalattosauriforms.
9
REVISING THE NEGLECTED POST-CRANIUM OF JURASSIC
RHYNCHOCEPHALIANS FROM THE SOLNHOFEN ARCHIPELAGO
V. Beccari1*, O. Rauhut1,2,3
The significant diversity of rhynchocephalians during the Mesozoic has been known for over a
century and is getting increasing attention in the last decade. However, most studies of this group
of lepidosaurs still focus mainly on the cranial anatomy, with emphasis on the jaws and dentition.
This has led to a poor understanding of the postcranial anatomy of rhynchocephalians, and the
implications this might have for their taxonomy, phylogeny, and palaeoecology. The high diversity
and exceptional preservation of Jurassic rhynchocephalians from the Solnhofen Archipelago can
provide new information on the postcranium of these animals. Taxa, such as Kallimodon and
Homoeosaurus, have been formerly described, but still lack proper diagnosis, leading to
disagreements over the validity and real diversity of rhynchocephalians during the Late Jurassic of
Europe. Revisions of these taxa are long overdue and can help in identifying new species based
solely on postcranial material, as well as to quantify morphological variation between specimens.
In the last five years, the postcranium of rhynchocephalians has accounted from 0% to 30% of the
total phylogenetic characters. In our preliminary works, we already observed new characters that
may improve and resolve some issues of their evolution and relationships. The postcranial
anatomy can also be used for palaeoecological inferences for the clade. Most of these inferences
are, so far, based on teeth and jaw morphologies. However, the closest relatives of
rhynchocephalians, squamates, show how important postcranial anatomy is, with studies that
correlate the appendicular skeleton proportions and morphology to lifestyle and habitat use.
10
DISSCO PREPARE: TOWARDS A EUROPEAN INFRASTRUCTURE FOR
SCIENTIFIC COLLECTIONS
1Sistema Museale di Ateneo, Università degli Studi di Firenze, via Giorgio La Pira 4, 50121,
Firenze, Italy.
2Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Università degli Studi di
Firenze, via Giorgio La Pira 4, 50121, Firenze, Italy.
3CNR – Istituto di Scienze del Patrimonio Culturale (ISPC), Via Cardinale Guglielmo Sanfelice 8,
80134, Napoli, Italy.
4CNR – Istituto di Geoscienze e Georisorse (IGG), Via G. Moruzzi 1, 56124, Pisa, Italy
5MiC – Istituto Centrale per il Catalogo e la Documentazione, Via di San Michele 18,
00153, Roma, Italy.
6MiC – Parco archeologico di Ostia antica, Viale dei Romagnoli 717, 00119, Roma, Italy.
7CNR – Istituto di Scienze e Tecnologie della Cognizione, Via San Martino delle Battaglia 44,
00161, Roma, Italy.
European Natural Sciences Collections (NSCs) consist of around 1,5 B specimens that are crucial
for answering fundamental scientific questions about ecological, evolutionary, and geological
processes. In the last decades the volume and diversity of information derived from NSCs are
exponentially increasing due to the digital transformation and the progressive use of analytical
instrumentation, remote sensing and molecular approaches. The Distributed System of Scientific
Collections (DiSSCo) is a world-class Research Infrastructure (RI) for NSCs currently in its
preparatory phase. The DiSSCo RI aims to create a new organisational model for collections that
digitally unifies all European natural sciences assets under common access, curation, policies and
11
practices ensuring that all the data are easily Findable, Accessible, Interoperable and Reusable
(FAIR principles). DiSSCo thus represents the largest ever formal agreement between natural
history museums, botanic gardens and collection-holding universities in the world.
The NHM of the University of Florence represents the current Italian National Node of the DiSSCo
Prepare Project and is playing a key role in establishing innovative relationships between its
internal Collection Management System and: i) ArCo, i.e. the Knowledge Graph of the Italian
Cultural Heritage promoted by the Italian Central Institute for Catalogue and Documentation and
the Italian National Research Council, ii) the General Catalogue of Italian Cultural Heritage web
portal created by the ICCD (3 M catalogue records) and, finally iii) the DiSSCo RI. In particular,
ArCo ontology network, adopted by Agenzia per l’Italia Digitale (AgID), allows the representation
on the semantic web of ICCD standards for natural heritage.
12
THE EMERGENCE AND EARLY DIVERSIFICATION OF THE PLACENTAL
MAMMAL BRAIN
O.C. Bertrand1*, S.L. Shelley1,2, T.E. Williamson3, J.R. Wible2, S.G.B. Chester4,5,6, J.J.
Flynn7,8,9,10, L.T. Holbrook11, T.R. Lyson12, J. Meng7, I.M. Miller13,12, H.P. Püschel1, T.
Smith14, M. Spaulding15, Z.J. Tseng16, S.L. Brusatte1,3
12Denver Museum of Nature & Science, 08028, Denver, CO, United States of
America.
13
Sciences, 1000, Brussels, Belgium.
Mammals have the largest brains among vertebrates, and boast a novel cerebral neocortex, center
of sensory integration and complex behavior. The emergence and diversification of the mammalian
brain remains challenging to decipher because fossil specimens that can provide brain endocasts
are relatively rare. We used 34 new computed tomography scans of Paleogene fossils including 17
taxa from the Paleocene of the USA and France alongside previously published endocasts.
Contrary to previous hypotheses that relative brain size increased continuously over time, our
results show that following the end-Cretaceous extinction, body mass increased at a faster rate
leading to a decrease in relative brain size in early Paleocene mammals. Ten million years after
the extinction during the Eocene, greater encephalization independently emerged in several
mammalian lineages. The neocortex and the petrosal lobules were substantially more expanded in
early members of crown placental mammals. Intensified competition in increasingly saturated
ecosystems might have triggered an arms race, in which greater encephalization and behavioral
specialization promoted lineage survival. Indeed, predators and potential prey had low and
indistinguishable phylogenetic encephalization quotients in the Paleocene, but both guilds
encephalized in the Eocene, arming carnivores with significantly larger relative brain size than
herbivores. Additionally, the Eocene experienced intense hyperthermal warming spikes, leading to
higher precipitation and increased seasonality, during which behavioral flexibility and ability to
disperse, linked to higher relative brain size, may have been beneficial. These findings indicate
that increased ecological opportunity and guild interactions both were instrumental during the
establishment of crown mammalian brains.
14
REDESCRIBING LIBYCOSUCHUS BREVIROSTRIS STROMER 1914
(CROCODYLIFORMES, NOTOSUCHIA)
1Departamento de Física e Ciencias da Terra, Universidade da Coruña, Rúa da Fraga 10, 15008,
A Coruña, Spain.
3Museo Paleontológico Egidio Feruglio, CONICET, Av. Fontana 140, U9100, Trelew, Argentina.
The German palaeontologist Prof. Ernst Stromer discovered a striking bizarre vertebrate fauna
during his expeditions into the Western Dessert of Egypt at the beginning of the 20 th century.
Among the species described by him is the crocodyliform Libycosuchus, being one of the few
fossils of Stromer’s collection that was not destroyed during the bombing of Munich in the Second
World War (1944).
Libycosuchus is represented by an almost complete, articulated skull, lower jaws and four
vertebrae found at the Gebel El Dist site (Cenomanian of Bahariya, Egypt). The skull is well
preserved, but damaged at several points (right orbit and postorbital bar, left jugal, right pterygoid
and palatines), which have been reconstructed. In the lower jaw, the anterior ends of both
dentaries are damaged, and the area around the left mandibular fenestra is broken and
reconstructed. We performed a CT-scan of these remains in order to improve the morphological
descriptions and assess the internal anatomy.
When first described (1914), anatomical comparisons of Libycosuchus were limited, and the taxon
was compared to Notosuchus, Alligatorellus, Theriosuchus, goniopholids, Bernissartia and
Crocodylia. This led Stromer to consider Libycosuchus as a “highly-specialized form with ancestral
features” closely related to Notosuchus and Theriosuchus and placed in its own genus and family.
Since then, a vast number of newly discovered taxa has increased the complexity in the
systematics of the Notosuchia, resulting in different interpretations of the position of Libycosuchus.
Our aim is to update comparisons with taxa from different notosuchian clades, and discuss its
phylogenetic relationships.
15
A SINGULAR ASSEMBLAGE OF SMALL AMPHIBIANS AND REPTILES
FROM THE EARLY MIOCENE (RAMBLIAN, MN3) OF TURÓ DE LES
FORQUES 1 (VALLÈS-PENEDÈS BASIN, CATALONIA)
1Institut Català de Paleontologia Miquel Crusafont, Edifici ICTA-ICP, c/ Columnes, s/n, Campus de
la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain.
2School of Earth Sciences, University of Bristol, 24 Tyndall Avenue, BS8 1TQ Bristol, United
Kingdom.
4Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via T. Valperga Caluso 35,
10125 Torino, Italy.
Turó de les Forques 1 is a fossil vertebrate locality located close to Castellbisbal, in the Neogene
Vallès-Penedès Basin, and correlated to zone A of the Ramblian regional age (MN3, Early
Miocene) based on its rodent fauna. Here we report a diverse assemblage of small amphibians
and reptiles that was recovered, together with other microvertebrates, using screen-washing
techniques, and is currently housed at the Institut Català de Paleontologia Miquel Crusafont. The
palaeoherpetofauna includes amphibians, lizards, snakes, and crocodiles, mostly represented by
fragmentary material. Among the amphibians, the identified taxa correspond to alytid and ranid
anurans, and possibly, but not certainly, to an indeterminate urodele. Among lizards, we report the
presence of chameleons, scincoids, lacertids, and anguines. Snakes are represented by
numerous but rather fragmentary vertebrae (mostly condyles), but we recognised at least the
presence of “colubrines”. Finally, crocodilians are represented by scarce teeth and osteoderms,
16
whose morphology is reminiscent of that of Diplocynodon, although we regard this identification as
tentative given the lack of more diagnostic material. Besides a relatively high diversity, the locality
is interesting in providing the earliest evidence of many small amphibian and reptilian taxa typical
of the Vallès-Penedès Basin, and a particular assemblage composition that lacks some typical
elements found in other localities of the same basin, like blanid amphisbaenians and the varanid
lizard Varanus. On the other hand, it includes less common taxa such as chameleons and
scincoids, representing the first record of the former in the basin.
17
EARLY MIOCENE REMAINS OF MELISSIODON FROM MOKRÁ-QUARRY
(MORAVIA, CZECH REPUBLIC) SHED LIGHT ON THE EVOLUTIONARY
HISTORY OF THIS RARE CRICETID GENUS
2Institute of Geology of the Czech Academy of Sciences, Rozvojová 269, 16500 Prague 6, Czech
Republic.
4Masaryk University, Faculty of Science, Department of Geological Sciences, Kotlářská 267/2, 611
37 Brno, Czech Republic.
The extinct genus Melissiodon is a cricetid considered endemic to Europe, known from the early
Oligocene to the early Miocene. Among micromammal remains, its finds are usually very rare, and
most of them are scarce and fragmentary. Currently, two species are considered taxonomically
valid from the early Miocene: Melissiodon schlosseri, which is based on two teeth from the MN2
German locality of Haslach and only reported in two other sites coeval in age (i.e., Ulm-Uniklinik
and La Chaux; Germany and Switzerland respectively); and Melissiodon dominans, found in MN3
and MN4 localities across Europe, even though the scarce and fragmentary remains make some
of these taxonomical assignments dubious. For this reason, Melissiodon dominans has become a
“wastebasket” species. However, the Mokrá-Quarry site provided one of the best-documented
findings of Melissiodon remains in MN4 localities of Europe. These new remains of Melissiodon
have been identified as a new morphotype that clearly differs from Melissiodon dominans by its
unique m1 morphology but still shows some resemblance to Melissiodon schlosseri. Based on
that, the material presented here is referred to Melissiodon aff. schlosseri. We also propose the
hypothesis of an evolutionary line starting from Melissiodon schlosseri that would have reached
18
the late early Miocene. Therefore, at least two different taxa of Melissiodon coexisted before the
genus extinction. Thus, the evolutionary history of the genus is more complex than previously
thought and a complete revision of the type material of Melissiodon dominans and Melissiodon
schlosseri is required.
19
MITOCHONDRIAL PHYLOGENY OF THE EXTINCT PROLAGUS SARDUS
(LAGOMORPHA, MAMMALIA): REIGNITING THE DEBATE ABOUT FAMILY
PROLAGIDAE
2IUCA - Aragosaurus, Universidad de Zaragoza, C/ Pedro Cerbuna 12, 50009 Zaragoza, Spain.
4Grupo Ecología de la resistencia bacteriana, H.C.U. Lozano Blesa, IIS Aragón, Avda. San Juan
Bosco 15, 50009, Zaragoza, Spain.
6Departamento de Física e Ciencias da Terra, Universidade da Coruña, Rúa da Fraga 10, 15008
A Coruña, Spain.
7Dipartimento di Scienze, Università degli Studi Roma Tre, Largo S. Murialdo 1, 00146 Roma,
Italy.
8Institut Català de Paleontologia Miquel Crusafont (ICP), Carrer de les Columnes s/n, 08193
Cerdanyola del Vallès, Spain.
9Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and
Paleoanthropology, Chinese Academy of Sciences, 142 Xi Zhi Men Wai Da Jie, 100044 Beijing, China.
Sardinian Quaternary palaeofauna potentially represents a gold mine for application of ancient
DNA techniques. Dozens of small and large mammals, now extinct, populated the island in the
recent past (sometimes until historical times). Palaeogenomic studies could cast a light on their
phylogeny, often obscure due to the modifications acquired under insular evolution, and provide
valuable data about past faunal diversity and dispersal routes.
The lagomorph Prolagus sardus is an emblematic Sardinian small mammal, and one of the last
20
survivors of the Plio-Pleistocene faunal assemblages. Archaeological evidence, as well as
testimonies from historians, highlight that P. sardus, the last representative of a genus that can be
traced back to ~20 Ma in continental Europe, survived in Sardinia (at least) until the Bronze Age.
According to Western European taxonomic “tradition”, Prolagus is a member of family
Ochotonidae. In this case, Prolagus represents, together with the extant Ochotona, the only
ochotonid to survive in historical times. Other scholars, mainly from Eastern Europe and Asia,
consider the existence of the family Prolagidae to include all the species of Prolagus and possibly
Ptychoprolagus.
A femur of P. sardus from San Lussorio cave (Romana, NW Sardinia, early Neolithic, 4.7-2.4 ky
BC) yielded DNA, providing promising results. Preliminary phylogenetic analyses based on a discrete
mitochondrial DNA dataset show that Prolagus is a sister clade to extant family Ochotonidae. Further
perspectives are: 1. clarify if Prolagus deserves to be included in a separate family (Prolagidae), and 2.
provide the time of the divergence of the two clades.
21
POSTCRANIAL OSTEODERM MORPHOLOGY OF PERMIAN PAREIASAURS
(PARAREPTILIA)
E.I. Boyarinova1,2*, V.K. Golubev1,2, I. Kogan3,4*, M.J. Van den Brandt5
1Borissiak Paleontological Institute of Russian Academy of Sciences, Profsoyuznaya 123, 117647
Moscow, Russia.
2Kazan (Volga region) Federal University, Kremlyovskaya 18, 420008 Kazan, Russia.
3Museum für Naturkunde Chemnitz, Moritzstr. 20, 09111 Chemnitz, Germany.
4TU Bergakademie Freiberg, B.-v.-Cotta-Str. 2, 09599 Freiberg, Germany.
5Evolutionary Studies Institute (ESI) and School of Geosciences, University of the Witwatersrand,
Johannesburg, PO WITS 2050, South Africa.
22
HIGH MORPHOLOGICAL DISPARITY IN A BIZARRE PALEOCENE FAUNA OF
PREDATORY FRESHWATER REPTILES
C. Brownstein1*
1Department of Ecology and Evolutionary Biology, Yale University, New Haven, CT, United States
of America.
The Cretaceous-Paleogene mass extinction was a pivotal event that fundamentally changed global
ecosystems and shaped extant biodiversity. However, the floras and faunas that appeared
immediately after the extinction event were very different from ecosystems found today.
Choristoderes are a phylogenetically problematic, species-poor reptile clade with a fossil record
stretching from the Middle Jurassic to the Miocene. Members of one major choristodere clade, the
Neochoristodera, seem to have their largest sizes and peak diversity right after the asteroid
impact. I describe two new choristoderes from a ~60-million-year-old ecosystem in western North
America based on complete skulls and partially articulated skeletons that deviate dramatically in
morphology. Kosmodraco magnicornis gen. et sp. nov. ossesses an extremely short snout,
extensive cranial ornamentation, and a sacrum with enlarged muscle attachment sites reminiscent
of some giant crocodylians. In contrast, Champsosaurus norelli sp. nov., is a longirostrine taxon
with an unornamented skull. Together with the North American choristodere Kosmodraco
dakotensis comb. nov., K. magnicornis substantiates a new clade of giant brevirostrine
choristoderes endemic to the American West. The new taxa in turn increase Cenozoic
choristodere richness and suggest that choristodere faunas showed trans-Atlantic divergences in
the aftermath of the asteroid impact.
23
INFERENCE OF LIMB PHASE AS A CRUCIAL GAIT PARAMETER FOR THE
PRODUCERS OF ICHNIOTHERIUM TRACKS FROM THE EARLY PERMIAN
TAMBACH FORMATION, GERMANY
2School of Biological and Environmental Sciences, Liverpool John Moores University, Liverpool,
United Kingdom.
3Museum für Naturkunde Berlin, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Berlin,
Germany.
The order of limb movements in symmetric gaits of quadruped tetrapods can be described as
continuous spectrum between pace (limb phase at 0 and 100%) and trot (limb phase at 50%) with
intermediates referred to as sequence walk. Given the availability of relatively long Ichniotherium
sphaerodactylum trackways in larger numbers and of complete skeletons that belonged to a very
likely trackmaker (Orobates pabsti), the well documented trackway sample of the early Permian
Bromacker locality yields the unique possibility to infer the limb phase of the I. sphaerodactylum
trackmakers via two independent approaches: (a) use of the along-track variability in stride length
for calculation of an optimum order of limb movements that minimizes variability under the
assumption of a homogeneous walking style, (b) comparison of the distance between the centers
of hip and shoulder joints in Orobates with the so-called glenoacetabular distance (GAD) in I.
sphaerodactylum of comparable size. The GAD represents a trackway measure that describes the
distance between successive pairs of manus and pes footprints and is supposed to correspond to
the trunk length of a skeleton. Approach (b) resulted in limb phases between 25% (single foot
lateral sequence walk) and 60% (diagonal sequence walk with diagonal couplets) with a maximum
between 40 and 50%. Approach (a) was inconclusive for most of the longer trackways, which were
too homogenous, and only one specimen produced a well-constrained optimum limb phase of 40%
(lateral sequence with diagonal couplets), a result in good agreement with an average limb phase
24
of 41% inferred by means of approach (b). Accordingly, Orobates probably employed a walking
trot or a slightly deviating type of lateral sequence walk.
25
REVISION OF THE CROCODYLIAN TOMISTOMA DOWSONI FROM THE
MIOCENE OF NORTH AFRICA PROVIDES NEW INSIGHTS INTO GAVIALOID
NEUROANATOMY
The interrelationships of the crocodylian species Gavialis gangeticus and Tomistoma schlegelii has
been historically disputed. Whereas molecular analyses indicate a sister taxon relationship
between the two species, morphological datasets typically place Gavialis as the outgroup to all
other extant crocodylians. Recent morphological-based phylogenetic analyses have begun to
resolve this discrepancy, with Gavialis more closely related to Tomistoma than to any other living
species. However, several stratigraphically older fossil taxa are recovered as closer to Gavialis
than Tomistoma, resulting in anomalously early divergence rates. This includes several species
currently referred to Tomistoma. Here we present an anatomical revision of Tomistoma dowsoni
from the Miocene of Egypt, known from a holotypic partial snout and a near-complete referred
skull. This work demonstrates its validity as a distinct species, but, combined with its incorporation
into a phylogenetic analysis, excludes it from the genus Tomistoma, which is restricted to the
extant species. The referred skull is well preserved, allowing enhanced visualization of the
braincase and thus an insight into the neuroanatomy of this species. The inner ear of Tomistoma
dowsoni exhibits an intermediate morphology between Gavialis gangeticus and Tomistoma
schlegelii, a starting point in understanding more about the evolution of ecomorphological features
in gavialoids that might help resolve the group’s interrelationships. Features such as the inner ear,
cerebrum, pituitary fossa and olfactory tracts vary in size amongst crocodylians. By incorporating
such previously inaccessible information from the neuroanatomy of extinct and extant crocodylians
into phylogenetic analyses, we might be able to resolve remaining problems surrounding the
Tomistoma-Gavialis debate.
26
ELLIPTIC FOURIER ANALYSIS VS FIXED LANDMARKS IN THE DENTITION
OF ANCHITHERIUM
2Department of Botany and Geology, Universitat de València, C/Doctor Moliner, 50, 46100,
Burjassot, Spain.
4Sedimentary Geology and Environmental Change, Geosciences Institute (CSIC, UCM), C/ del Dr.
Severo Ochoa, 7, 28040, Madrid, Spain.
Anchitherium is a genus of equid (Equidae) belonging to the extinct subfamily Anchiteriinae. This
subfamily is characterized by brachydont teeth and tridactyl limbs. Nevertheless, the isolated teeth
of Anchitherium are difficult to differentiate because of the morphological similarities between the
molars (M/m) and the premolars (P/p), and between the decidual and the definitive dentition
especially in the upper cheek tooth rows. Therefore, comparisons between species and
paleoecological reconstructions can be biased by this dental element determination problem. In
order to identify the different dental elements, we use geometric morphometric analysis on upper
molars applying Elliptic Fourier Analysis and a set of fixed landmarks. We use isolated and
associated maxilla teeth from the Spanish Middle Miocene sites of Mahou and Los Nogales (MN5,
Madrid) trying to interpret the observed distribution. Our geometric morphometric approaches
consist of the digitization of the different dental pieces using TPSDig2. To do that, we used scaled
photographs of the occlusal surfaces of each dental pieces and transformed them to have the
same orientation. We use the RStudio packages Stereomorph to collect the data and Geomorph
to perform different statistical analyses. The provisional results indicate that these techniques may
be useful not only to distinguish between premolars and molars, but also to help identify each
dental element in anchitherines.
27
THE AXIAL SKELETON OF THE HOLOTYPE OF THE SIMOSAURID
PALUDIDRACO MULTIDENTATUS (EOSAUROPTERYGIA) FROM THE LATE
TRIASSIC OF CENTRAL SPAIN
Simosaurids are Triassic eosauropterygians belonging to the clade Nothosauroidea. Its fossil
record ranges from the Anisian or Ladinian (Middle Triassic) to the Carnian (Late Triassic) of
Europe and the Middle East. Simosauridae was, until recently, a monospecific clade only including
Simosaurus gaillardoti from the Ladinian (Middle Triassic) of France and Germany, due to the
Austrian Ladinian ‘Partanosaurus zitteli’ was considered as a nomen dubium. In 2018, a new
simosaurid, Paludidraco multidentatus, from the Carnian (Late Triassic) fossil site of El Atance
(Guadalajara, Spain) was described. It was interpreted as a bizarre taxon, showing adaptations for
slow movement near the marine bottom of shallow littoral environments, thus, extremely different
from its sister taxon, the active predator S. gaillardoti. Paludidraco multidentatus was briefly
described based on a relatively complete skeleton (the holotype) and an isolated skull. The
anatomy of the vertebrae of this taxon has not been hitherto described. Conversely, the skeleton of
S. gaillardoti was described in detail in the literature, being well-known not only the skull but also
the postcranium, including elements of the axial series from cervical to caudal vertebrae. The
description in detail of the vertebral elements preserved in the holotype of P. multidentatus is
proposed. Most of these elements are articulated, corresponding to the almost complete dorsal
and sacral series but also some cervicals and, probably, the most proximal caudal vertebrae.
Detailed comparisons with the vertebrae of S. gaillardoti are performed, along with those of some
indeterminate simosaurids from other European palaeontological sites.
28
PEER INTO THE PELECANIMIMUS EYE: SCLEROTAL RING
RECONSTRUCTION OF AN EARLY CRETACEOUS ORNITHOMIMOSAUR
FROM SPAIN
2Department of Organismal Biology, University of Chicago, 60637, Chicago, IL, United States of
America.
Keywords: Theropod, Dinosauria, Visual capabilities, CT-Scan, Las Hoyas fossil site
Ornithomimosaurids were a distinctive group of theropod dinosaurs. Some taxa have been
suggested to be omnivores, unlike the vast majority of non-avian theropods. This is the case of
Pelecanimimus polyodon, with slender mandibles with more than 200 teeth. But the dietary
behaviour is not the only questioned issue in their palaeobiology. Diet could be strongly related to
sensory habits such as visual capabilities. Non-avian theropod dinosaurs have been assumed as
essentially diurnal, but recent studies have revealed probable nightlife capabilities for specific
lineages. These studies correlated the relative dimensions of the scleral ring and the orbit with the
probability of nocturnality. The results indicated that some ornithomimosaurs, such as
Ornithomimus, had high probability of nocturnal vision capabilities while others such as
Garudimimus would have had cathemeral activity (day and night habits). To test the vision
capabilities of Pelecanimimus, an accurate reconstruction of its scleral ring was performed.
The reconstruction was made using CT-Scans of the skull of Pelecanimimus. We carried out an
accurate segmentation of the different ossicles of the slightly disarticulated scleral ring separately
based on the tomography sections. Posteriorly, the scleral ring was virtually reconstructed,
reintegrated and manually articulated.
29
With these reconstructions, we will be able to correlate statistically precise measurements of the
scleral ring and orbit with the vision capabilities of Pelecanimimus in future approaches. Resolving
if it was a nocturnal or cathemeral dinosaur will provide insight on the visual capabilities of early
ornithomimosaurs.
30
SPECIFIC SEPARATION OF DICRAEOSAURUS HANSEMANNI AND
DICRAEOSAURUS SATTLERI: OSTEOLOGICAL AND TEMPORAL EVIDENCE
A. Campbell1,2*, D. Schwarz1
Dicraeosaurus is a flagellicaudatan sauropod from the Late Jurassic of the Tendaguru Formation
in Tanzania that was originally described by Werner Janensch in 1914. It is divided in to two
species, D. hansemanni and D. sattleri based largely on the temporal distribution of the known
material and the generally more gracile morphology of D. sattleri. Here we present a new
comparison of the material with the aim of clarifying the differences between the two species, and
determine whether they are sufficient to justify the separation of the genus. The holotype material
for D. hansemanni is mounted at the MfN, Berlin and D. sattleri remains in the collections, all
material, including specimens currently assigned to genus level, was digitised using laser surface
scanning. We found that based on presented analyses and stratigraphy there is sufficient
distinction between the specimens to justify the separation of the two species, with all material
found in the Upper Dinosaur Member having a morphology akin to that of D. sattleri and that from
the Middle Dinosaur Member with D. hansemanni.
31
REVISITING THE RADIATION OF HORSES
Horses were common elements of Cenozoic communities and reached high diversity during the
Neogene. Most research of their diversification has so far focused on the Early Miocene
cladogenesis of the subfamily Equinae (equines, the so-called ‘grazing horses’), interpreting raw
diversity counts at face value. Using occurrence data, diversification models and phylogenetic
methods, we reconstruct speciation and extinction in horses and interpret these trends in the
context of the evolution of body size and relative tooth crown height (hypsodonty). We found that
fast species proliferation (speciation rate) was not restricted to equines but was a shared feature in
both anchitheres and equines, likely stemming from the more variable environments of the
Neogene. The fast early-phase radiation in the subfamily Equinae was likely the result of reduced
extinction rates and higher lineage survival rather than a substantial acceleration in speciation. The
evolution of body size and hypsodonty was not rapidly acquired in regions of the phylogeny with
faster diversification, suggesting a broad-scale decoupling of ecomorphological and taxonomic
diversification. Our phylogenetic modeling approach reveals that major phenotypic trends were not
caused by phyletic progression, derived from sustained and ubiquitous directional selection, but
more likely were the outcome of differential lineage-level survival and multiplication (species
sorting).
32
NEW REMAINS OF NEOTROPICAL BUNODONT LITOPTERNS FROM THE
MIDDLE MIOCENE, LA VENTA FAUNA: PALAEOBIOLOGY AND
SYSTEMATICS
2CR2P, UMR 7207, Muséum National d’Histoire Naturelle, CNRS, Sorbonne Université Rue
Buffon 8,75005, Paris, France.
Litopterna is one the most diverse and long–lived clades of the South American Native Ungulates
(SANUs). Megadolodus and Neodolodus are bunodont litoptern genera recorded in the middle
Miocene tropical fauna of La Venta (Colombia). Both taxa were initially identified as a didolodontid
‘condylarths’, but later transferred to Proteroteriidae, within Litopterna. Previous studies proposed
their inclusion within Proterotheriidae, but possible affinities with early litopterns and didolontids
have not been properly tested in phylogenetic analyses. We report new material of Megadolodus
and Neodolodus and reassess their phylogenetic relationships with Litopterna and Didolodontidae.
In our phylogenetic analyses, we tested two alternative approaches of character construction for
serial characters on the dentition. The reductive coding approach scores characters separately
assuming the observed morphological variations are independent, whereas the composite coding
approach favours the construction of a single character when the observed variation is
hypothesized to be non-independent. Based on the new fossil material, our analyses with either
one or the other coding approach support a close relationship between Megadolodus and
Neodolodus, within Litopterna, and not a close relationship with Didolodontidae. At a less inclusive
level, the relationships of Megadolodinae within Litopterna and its inclusion within
Protherotheriidae vary depending on the coding approach used. However, all our analyses
unambiguously support the monophyly of Megadolodinae as a clade of bunodont Neotropical
litopterns. While the discovery of these new remains enlightens part of the litoptern phylogeny, the
sensitivity of our analyses to coding approaches highlights the importance of critical evaluation of
character construction in morphological phylogenetics.
33
NEW VERTEBRATE REMAINS FROM THE JURASSIC-CRETACEOUS
TRANSITION OF THE EASTERN CAMEROS BASIN (SPAIN)
3Museu de la Conca Dellà. C/ del Museu 4, 25650, Isona i Conca Dellà, Spain.
The eastern Cameros basin is well known by the high number of dinosaur tracksites, although
sites with osteological remains are scarce. One of the units that has yielded fossil bones is the
Matute Alloformation, from where the freshwater aquatic turtle Pleurosternon moncayensis and the
semionotiform actinopterygian Camerichthys lunae have been described. This geological unit is
included within the upper part of the Tera Group and was deposited in shallow carbonate lakes.
The most probable age of the Matute Alloformation is late Tithonian-earliest Berriasian. A new
paleontological site, named Ribota, has been found close to the locality of Ágreda (Soria province,
north-central Spain) in the lower part of this unit. The Ribota site is included in a carbonate-
dominated interval outcropping over a large area of almost 10 hectares. There, three main areas
with 36 small but rich, fossiliferous concentrations of isolated bones have been identified. The
excavation of these areas has allowed to recover more than 75 vertebrate fossils. These mainly
include crocodylomorph remains, such as isolated teeth, skull fragments, vertebrae and
osteoderms, of different sized individuals with possible goniopholidid affinity. Other findings,
although less abundant, includes testudine plates and few isolated cranial and appendicular
pterosaur bones. In addition, cranial fragments and several articulated partial specimens of
indeterminate osteichthyans have been recovered. Further systematic studies of these fossils will
shed new light on these lacustrine vertebrate fossil assemblages from the Jurassic/Cretaceous
transition, a poorly known time period in the Iberian Peninsula and Europe.
34
DEFORMATION MECHANISM AND RETRODEFORMATION OF INDIAN
RHYTIDOSTEID TEMNOSPONDYL RESULTS IN A NEW COMBINATION OF
BRAZILIAN TAXON
1Department of Earth Sciences and Remote Sensing, JIS University, 81, Nilgunj Road, Agarpara,
Kolkata – 700109, West-Bengal, India.
2Universidade Federal de Santa Maria, Av. Roraima nº 1000 Cidade Universitária Bairro - Camobi,
Santa Maria - RS, 97105-900, Brazil.
3Geological Studies Unit, Indian Statistical Institute, 203 Barrackpore Trunk Road, Kolkata 700108,
India.
35
comb. nov., for the Brazilian taxon Sangaia lavinai is erected, based on a detailed phylogenetic
study.
36
A NEW, HIGH-SPEED PROTOCOL TO RUN MUSCLE-DRIVEN
BIOMECHANICAL SIMULATIONS THROUGH FINITE ELEMENT ANALYSES
Finite element analysis (FEA) is a computational method used to predict the behaviour (stresses,
strains, and deformation) of a structure under predefined loading conditions. It can be applied to
biological structures such as bone to study defined biomechanical scenarios. However, as muscle
is an extremely complex structure to model, biologists usually model muscle forces indirectly. In
2007, the Boneload MATLAB routine was developed to distribute muscle forces on a surface
defined by the user. This routine needs to be coupled with a pre-existing FEA software (e.g.,
Strand7) to perform the analyses, and has been widely used ever since. Here, we demonstrate a
new method to run muscle-driven finite element simulations on bone by distributing muscle forces
on their insertion areas within a single computational environment. We test this protocol on three
different biomechanical systems linked to feeding and locomotion. Our automated script meshes
the 3D model, applies the constraints, and distributes muscle forces within a single simulation
software: Metafor. Metafor can perform the entire protocol on high resolution volumetric meshes
(millions of tetrahedra) rapidly, outpacing the processing time of other widely used software by up
to 12 times, all-the-while producing similar and congruent results. Our new protocol thus opens up
the possibility to routinely and rapidly simulate the behaviour of high-precision, muscle-driven FE
models and investigate a wide range of biomechanical scenarios at unprecedented resolution.
37
A COMPUTER-AIDED PROTOCOL FOR THE VIRTUAL RECONSTRUCTION
OF THEROPOD DINOSAUR SKULLS. TYRANNOSAURUS REX AS AN
ICONIC CASE STUDY
2Dipartimento di Scienze della Terra, Università di Firenze, Via G. La Pira 4, 55121, Firenze, Italy.
5Negaunee Integrative Research Center, Field Museum of Natural History, 1400 S DuSable Lake
Shore Drive, 60605 Chicago, United States of America.
7Faculty of Geosciences, Utrecht University, Princetonlaan 8a, 3584 CB Utrecht, the Netherlands.
Taphonomic processes impact the physical preservation of fossil remains and often involve their
deformation. Recently developed R-based virtual reconstruction protocols ‘Retrodeformation’ and
‘Target Deformation’ aim to reduce taphonomic distortion. These protocols have been tested and
applied to Cenozoic fossil mammals and fossil human digital remains. Retrodeformation removes
asymmetric alteration by applying a set of corresponding bilateral landmarks on the model. Target
Deformation uses target specimens of the same species (less distorted, even if incomplete) as a
virtual reconstruction guide, ideally permitting the protocol to reconstruct the original morphology.
The procedure of undistorting the specimen thus follows two steps: 1. Retrodeform by applying
bilateral landmarks to the target specimen; 2. warp the second original specimen onto the
retrodeformed target 3D model. Here, we present the preliminary result of the application of
Retrodeformation and Target Deformation on two iconic Tyrannosaurus rex skulls, FMNH PR2081
(informally known as 'SUE') and RGM.792000 (informally known as 'Trix'). ’SUE’ is based on the
38
published raw CT-scans, which include considerable matrix around and within the skull, whereas
the “Trix” 3D model was obtained by surface scanning. Retrodeformation was run on Trix first, then
Target Deformation methodology was applied on “SUE', using the better-preserved and
retrodeformed “Trix” as the target specimen. The results are extremely positive, despite the matrix
surrounding the skull of “SUE”. The result provides a baseline for deeper applications of the
method to more T. rex specimens and other dinosaur skulls.
39
NEUROANATOMY OF GROSSIUS, AN EARLY PREDATORY LOBE-FINNED
FISH FROM THE DEVONIAN OF SPAIN
Onychodonts (Osteichthyes, Sarcopterygii) were an extinct group of predatory marine fishes from
the Devonian traditionally characterized by kinetic skulls, hypertrophied parasymphisial teeth
inserted in whorls on the dentary symphysis, and a modified ethmosphenoid region of the
neurocranium to accommodate the tusks. The holotype and only specimen of Grossius
aragonensis from the Moyuela Formation (Eifelian, Middle Devonian) of Aragón province (Spain)
represents one of the most completely-preserved onychodont skulls but, despite its three-
dimensional preservation, a detailed study of its internal anatomy was impossible, until now.
The application of computed microtomography techniques has enabled to gain access to
previously unknown cranial anatomical information and to virtually reconstruct the morphology of
the braincase and its internal cavities for the first time. The neurocranium of Grossius shows
differences with those of other onychodonts, such as a more acute angle between the otic-occipital
region and the anterior-posterior axis of the ethmosphenoid than in Onychodus and Qingmenodus.
Comparisons with other osteichthyan neurocrania revealed that onychodonts share features
markedly different from other sarcopterygians such as Dipnorhynchus (dipnoans) or Diplocercides
(coelacanths), like deep, large and ventrally-directed nasal capsules forming a broad angle
between them.
The study of the neuroanatomy of Grossius highlights unique features, as well as new
synapomorphies of Onychodontida. These new anatomical data constitute a rich source of
morphological information for such an enigmatic group of early osteichthyans and will have a
40
broad impact on the study of sarcopterygian interrelationships and character evolution in future
phylogenetic analyses.
41
E UNUM PLURIBUS – THE FIRST HIGH-DIVERSITY KOGAIONID
(MAMMALIA, MULTITUBERCULATA) ASSEMBLAGE FROM THE
UPPERMOST CRETACEOUS OF THE TRANSYLVANIAN AREA, ROMANIA
Z. Csiki-Sava1,2*, R.-C. Lintz1, Ș. Vasile1,2
1Faculty of Geology and Geophysics, University of Bucharest, 1 N.Bălcescu Avenue, 010041
Bucharest, Romania.
2Center for Risk Studies, Space Modeling and Dynamics of Terrestrial and Coastal Systems,
University of Bucharest, 1 Nicolae Bălcescu Ave., 010041 Bucharest, Romania..
Here, we report a remarkable kogaionid locality (Fântânele1 in the northwestern Hațeg Basin) that
is both specimen-rich and taxonomically diverse. It was discovered about 25 years ago, when
isolated teeth from here had been tentatively referred to the Paleocene kogaionid Hainina. During
the past two decades, over 50 isolated teeth have been recovered from this locality. Their survey
suggests that at least three different taxa are represented in the local assemblage. Small-sized
kogaionids dominate, including the probably first definitive, apomorphy-based report of Barbatodon
oardanesis outside its type-locality. These are accompanied by less common medium- and large-
bodied taxa, some of which are possibly related to the recently erected Kogaionon radulescui,
while others are reminiscent of Barbatodon transylvanicus or Litovoi tholocephalos, all taxa known
from other parts of the basin. Despite being taphonomically and lithofacially similar to the
monotaxic oardaensis type-locality, the politaxic Fântânele1 locality documents in premiere the
existence of diversified sympatric kogaionid assemblages in Transylvania.
Work supported by the Romanian Ministry of Research, Innovation and Digitalization, CNCS–
UEFISCDI grant PN-III-P4-ID-PCE-2020-2570, within PNCDI III.
42
RANGE SIZE AND EXTINCTION RISK OF CUBAN MAMMALS: A
CONSERVATION PALEOBIOLOGY PERSPECTIVE
1Ruhr University Bochum, Faculty of Biology and Biotechnology, Theoretical and Applied
Biodiversity Research, Universitätsstr. 150, 44780 Bochum, Germany.
2German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Puschstr. 4, 04103
Leipzig, Germany.
Predicting why and when a species will become extinct is a crucial, yet challenging task in
conservation research. Geohistorical data can provide essential information on how ecosystems
reacted to disturbances, such as climate change and anthropogenic impacts, and have proven
especially useful in understanding extinction risks.
Here, we used a newly assembled dataset of late Quaternary fossil occurrences on Cuba (1,522)
and a new method of range size reconstruction based on co-occurrence analysis to analyze the
relationship between extinction risk and geographic range size for mammals across four orders.
Furthermore, we investigated changes in range size of extant and extinct species through the late
Quaternary.
We found that the probability of being threatened or extinct was negatively and significantly related
to range size for Cuban mammals. Moreover, we detected a significant decrease in range size
since the Late Pleistocene across all species in our dataset. Finally, we recorded significant
differences in range size shifts between taxonomic orders and between extant and extinct species.
In particular, while range contractions were significantly associated with the extinction of mammals
on Cuba, extant species experienced range expansions. We also discussed patterns of changes in
range size in response to human arrival and climatic changes at the end of the Last Glacial
Maximum.
Our results offer an insight into range size shifts of Cuban mammals through the late Quaternary,
and highlight the importance of historical range contractions as predictors of extinction risk.
43
BRINGING THE DEVIL BACK HOME
S. Danisi1*, F. Carotenuto1
1Dipartimento di Scienze della Terra, dell’Ambiente e delle Risorse, Università degli Studi di Napoli
“Federico II”, Via Cinthia, 21, 80126, Monte Sant’Angelo, Napoli.
44
TIME-SCALING OF CROCODYLIAN EVOLUTION AND THE CONTRIBUTION
OF TOTAL EVIDENCE TIP-DATING ANALYSIS
G. Darlim1*, M. Rabi1,2
A manageable number of living species and rich fossil record make Crocodylia, the crown group of
Crocodyliformes, an excellent model clade for divergence dating analyses. Most time-scaling
approaches of crocodylian evolutionary trees rely on often poorly established fossil calibrations in
molecular divergence studies as well as on the a posteriori distribution of time among nodes and
internal branches based on stratigraphic range of fossils and interspecific relationships. Tip-dating
analyses, on the other hand, have the advantage of simultaneously generating a time-scaled
phylogeny using both phenotypic and molecular rates and stratigraphic ages of fossils. In
Crocodylia, only a single total evidence tip-dating analysis have been performed so far,
contributing to the reinterpretation of character evolution and to well-supported divergence age
estimates. Here we present an additional tip-dating study employing a different taxon sampling and
focusing on the origination time of Crocodylia. Our results were overall consistent with the previous
tip-dating analysis as well as with some published molecular clock estimates explicitly following
best practices for fossil node calibrations. Inconsistency with estimates from published a posteriori
methods is apparent on the other hand, as well as sensitivity of tip-dating to sampling, particularly
autapomorphies. Finally, our analyses demonstrate the critical importance of integration of DNA
and morphological data for estimating clade ages when morphology-based relationships among
living taxa are robustly contradicted by genomic data.
45
PHYLOGENETIC SIGNAL OF CRANIAL CHARACTERS IN CLADISTIC
ANALYSIS OF DIPLODOCOID SAUROPOD DINOSAURS
3Department of Vertebrate Paleontology, American Museum of Natural History, 200 Central Park
West, 10024, New York, United States of America.
4GeoBioTec, NOVA School of Science and Technology, Largo da Torre, 2825-149, Caparica,
Portugal.
5The Dinosaur Institute, Natural History Museum of Los Angeles County, 900 W Exposition Blvd,
CA 90007, Los Angeles, United States of America.
Diplodocoidea is a highly diverse clade of sauropodomorph dinosaurs, which has been reassessed
for its systematics in great detail in the past decade. Recent descriptions of new taxa (e.g.
Galeamopus) and the reestablishment of Brontosaurus as a valid genus are just two outcomes of
analyses with large datasets at both specimen- and species-level. However, several taxa included
in these analyses are represented by small-sized individuals, which had raised doubts about the
validity of phylogenetic analyses. Here, we test the impact of ontogeny onto two versions of a
matrix with focus on flagellicaudatan Diplodocoidea with both presence and absence of postcranial
characters. The matrix includes 86 operational taxonomic units, 38 of which preserve cranial
remains at varying ontogenetic stages, and was analyzed with TNT. Using the cranial-only dataset,
the traditionally recognized partition into Apatosaurinae and Diplodocinae was not recovered, and
small-sized individuals appear to cluster at the base of a grade of diplodocid specimens leading
towards a monophyletic Dicraeosauridae. Using the complete dataset, large skulls usually
identified as diplodocines group with the large type specimens of Apatosaurus ajax and A. louisae,
whereas smaller skulls group with the smaller type specimens of Galeamopus pabsti, G. hayi, and
Kaatedocus siberi. Hence, body size, and thus possibly ontogeny, seems to impact tree structure,
with skull-only specimens being particularly prone of being attracted to similarly-sized specimens.
46
It is therefore crucial to identify ontogenetically variable characters in the skull, and avoid scoring
them in juvenile specimens to reduce directed noise in the phylogenetic analysis.
47
THE HYOID ARCH AND FEEDING IN TWO FISH-SHAPED MARINE
TETRAPODS
The hyoid apparatus is essential for underwater feeding in marine tetrapods, but the extent of
convergent adaptations is not well known. This study compares ossified hyoid elements in
ophthalmosaurid ichthyosaurs and odontocetes, two groups with an overall similar body shape
often interpreted to indicate a similar ecology, asking whether they also share a similar feeding
mode.
Three sources of data were used: size, morphology and internal bone microstructure. Size and
morphology data was collected from all ophthalmosaurid specimens (Middle Jurassic - middle
Cretaceous) where at least one hyoid is preserved (n= 13). Inner bone microstructure was mapped
via CT scanning of four ophthalmosaurid hyoid elements from the Late Jurassic Slottsmøya
Member at Spitsbergen and two odontocete specimens (Lagenorhynchus albirostris and
Hyperoodon ampullatus). By comparing the size data from ophthalmosaurids to earlier
ichthyosaurs (Early Triassic – Middle Jurassic), we show that the relative size and shape of the
hyoids did not change. With regard to the inner microstructure of the hyoid elements, the
odontocetes have an overall less compact structure, whereas the ophthalmosaurids possess
cancellous inner cones in an outer, more compact sheath. These are likely mechanical adaptations
to feeding mode. Together with the consistently slender hyoid element, this indicates that
ichthyosaurs never evolved the same type of suction feeding as extant odontocete whales.
This research project is supported by a EAVP Research Grant (awarded 2020 to LL Delsett).
48
AEOLOSAURINE TRAITS IN VERTEBRAL CAUDAL SERIES FROM
BELLEVUE LOCALITY (FRANCE): MORPHOFUNCTIONAL HOMOPLASY OR
EVIDENCE FOR SOUTH AMERICAN TITANOSAURS IN THE LATE
CRETACEOUS OF EUROPE?
1
Dynamics of Nature, Museum für Naturkunde - Leibniz Institute for Evolution and
Biodiversity Science, Invalidenstraße 43, 10115 Berlin, Germany.
2
Institut Català de Paleontologia Miquel Crusafont – ICTA-ICP, Edifici Z, C/ de les
Columnes s/n. Campus Universitat Autònoma de Barcelona, 08193 Cerdanyola del
Vallès, Spain.
3
Museu de la Conca Dellà, c/Museu 4, E-25650 Isona, Lleida, Spain.
4
Musée des Dinosaures, 11260 Espéraza, France.
Systematics and palaeobiogeography have always been controversial topics when dealing with
Late Cretaceous European titanosaurs. Thanks to thorough works carried out in the last decades,
the taxonomic diversity of this group of sauropods has significantly increased, but we are still far
from perceiving the real image. The early Maastrichtian Bellevue site (Aude, France) is well-known
for its dinosaur fauna, especially the titanosaur taxa, with Ampelosaurus atacis as the only species
described to date. Among the sauropod material, interesting features were identified in a sequence
of anterior caudal vertebrae associated with an incomplete sacrum: the caudal centra present an
anterior margin tilted anteriorly, the neural arches are positioned on the anterior margin of the
centrum, and the prezygapophyses are notably elongated. Besides these characters, other isolated
anterior-middle caudal vertebrae show a neural spine, in which the distal tip is tilted anteriorly.
These features are commonly recognized as synapomorphies of the South American clade
Aeolosaurini, and some of them were reported in a caudal series from Lo Hueco site (Spain). With
the current data at hand, two main hypotheses emerge: 1) aeolosaurines were present in Europe
during the early Maastrichtian, or 2) the acquisition of such particular caudal features would reflect
some kind of morphofunctional homoplasy between distant-related titanosaur taxa. The ongoing
49
study will shed light on the question as well as on a better comprehension of the complex
evolutionary history of this clade of sauropods.
50
SMALL BRAINS PREDISPOSED LATE QUATERNARY MAMMALS TO EXTINCTION
3The Steinhardt Museum of Natural History, Tel Aviv University, Tel Aviv, 6997801, Israel.
During the Late Quaternary, many of the world’s largest terrestrial mammals went extinct. These
extinctions are often attributed to either human hunting, climate change, or both. However, many
large mammals similar in size to those that went extinct are still alive today, implying that body size
was not the only predictor of extinction risk. We hypothesized that extant species have significantly
larger brains than those that went extinct during the Late Quaternary which allowed them to cope
with the changing environment and/or novel predators (humans). We created a dataset that
included brain sizes of 291extant mammals and 50 that went extinct during the Late Quaternary.
We performed logistic and mixed effect models, controlling for phylogeny and body mass to test if
extant mammals have larger brains than extinct ones and if this was correlated with extinction
status. We found that extant species have brains on average 53% larger than those that went
extinct, and 83% larger when a single regression line is fitted. We also found that brain size was a
significant predictor of extinction status. We propose that small-brained species were predisposed
to extinction during the Late Quaternary whereas their large-brained extant counterparts were
better suited to cope with changing environments and human hunters. We propose that
possessing a large brain was a crucial, yet so far, neglected characteristic of surviving megafauna
species.
51
TETRAPOD ICHNOASSOCIATIONS, BIOSTRATIGRAPHY AND
PALEOENVIRONMENTAL RECONSTRUCTION OF THE PERMIAN
TERRESTRIAL SUCCESSION OF CASTELLAR DE N'HUG (CATALAN
PYRENEES, SOUTHERN EUROPE)
4School of Earth Sciences, University of Bristol, Wills Memorial Building, Queens Road, Bristol
BS8 1RJ, United Kingdom.
5Istituto Nazionale di Geofisica e Vulcanologia, Via di Vigna Murata 605, I-00143 Roma, Italy.
The early–middle Permian in the Western peri-Tethys was marked by an aridification process, with
environments changing from wet- to dry-dominant conditions. The evolution of tetrapod continental
faunas in Southern Europe is unclear due to the remains scarcity, especially bones and teeth.
However, tetrapod ichnites are better known from European Permian deposits, thus being a
powerful tool to reconstruct ecosystems. The discovery of three new ichnosites from the lower and
middle Permian volcanosedimentary terrestrial successions from the Catalan Pyrenees (NE
Spain), together with stratigraphic and facies analyses, allows a better understanding of these
ecosystems. Tetrapod ichnofossils were discovered in red-beds from the Lower Red Unit and
Upper Red Unit, corresponding to alluvial deposits and building an almost 1000 m thick continuous
succession, the longest of the Pyrenean Basin. The stratigraphic succession also includes
volcaniclastic deposits, sometimes reworked by fluvial processes. The volcaniclastic deposits
notably diminish towards the upper part of the succession, which is dominated by floodplain
52
deposits with desiccation cracks. The identified tetrapod ichnofossils correspond to the
ichnogenera Batrachichnus, Dromopus, Varanopus, Hyloidichnus, Brontopus, Pachypes and
Charachichnos. 3D photogrammetric models of selected footprints and trackways were generated,
complementing morphological descriptions and biomechanical analyses. The composition of the
ichnoassociations changes along the succession, as well as the accompanying invertebrate fauna,
showing features of the Dromopus biochron in the volcaniclastic and fluvial deposits and features
of the Brontopus sub-biochron in the floodplain deposits. The biostratigraphic and
magnetostratiphic results of this study suggest a late Cisuralian to early–middle Guadalupian age.
53
SYNCHROTRON SCANNING REVEALS THE UNIQUE ANATOMY OF A
LATEST CRETACEOUS PADDLEFISH FROM TANIS (ND, USA)
1
Department of Organismal Biology, Uppsala University, Norbyvägen 18A, 75236
Uppsala, Sweden.
The Cretaceous-Palaeogene event deposit Tanis (North Dakota) yields fossils from yet
undescribed Acipenseriformes. Propagation Phase-Contrast Micro-Computed Tomography was
applied to study anatomical features preserved in a partial paddlefish from this assemblage.
Comparison with the two extant paddlefishes Polyodon and Psephurus, and with fossil forms
including the Maastrichtian Paleopsephurus shows that The Tanis paddlefish represents a new
taxon.
The elongate cranial roof appears intermediate between the more robust Polyodon skull and the
more slender Psephurus skull. It is distinctly curved anteroposteriorly, as is the parasphenoid,
which trends parallel with the cranial roof up to its anterior margin. This differs from the condition in
the extant forms, especially Polyodon, in which the parasphenoid is straight and converges on the
skull roof anteriorly. The postorbital foramen features a unique interfenestral strut. The
posttemporal widens posteriorly, whereas it narrows in both Polyodon and Psephurus. The parietal
is significantly longer and intermediate in width relative to both extant genera. Its centre of
ossification is positioned much further forward than in Polyodon. The gill rakers appear
comparatively short and unlike those of the modern paddlefishes.
The cause of death was likely burial in the massive seiche wave and/or suffocation with impact
spherules. The spherules are almost exclusively encountered between the second and the third gill
arch, possibly indicating the main respiratory water flow path; since the gill arches are obscured by
a gill cover that cannot be removed without damaging the specimen, such details would remain
invisible without the use of synchrotron microtomography.
54
A STYRACOSTERNAN SACRUM FROM A NEW FOSSIL LOCALITY FROM
THE ARCILLAS DE MORELLA FORMATION IN MORELLA (SPAIN)
2 Real Academia de Ciencias Exactas, Físicas y Naturales, c/ de Valverde, 24, 28004 Madrid,
Spain.
An isolated styracosternan ornithopod sacrum from the new Barremian fossil locality of Fábrica
Giner (Morella, Castellón, Spain) is described. This partial sacrum comprises six fused sacral
centra including the badly preserved posterior end of a most anterior sacral vertebra. Additionally,
some portions of the sacral ribs of the posteriormost sacral centra are preserved. The
posteriormost sacral centrum is transversely broad and has a wider than deeper concave posterior
articular facet. Its ventral surface is slightly eroded but seems to be flat. The base of the sacral ribs
is positioned at the posterior half of the centrum. The two preceding sacral vertebrae are
transversely broad and have flat ventral surfaces. In these vertebrae the sacral ribs are
intervertebrally positioned. The most anteriorly positioned sacral centrum is spool-shaped and
bears a well-defined ventral keel whereas the succeeding sacral centrum is not transversely
constricted and present a modest ventral keel anteriorly placed.
To date, the styracosternan sacral material from the Arcillas de Morella Formation have been
ascribed to Iguanodon bernissartensis and to Morelladon beltrani. The analyzed partial sacrum
clearly differs from those of the I. bernissartensis by the absence of a ventral sulcus in the most
posterior sacral vertebrae. Comparing it with those attributed to Morelladon, both differences (e.g.
un-keeled penultimate sacral vertebrae, third last sacral centrum transversely broad) and
similarities can be observed. Although some of these differences can be part of the intraspecific
variability of Morelladon, it has been preferred to provisionally assign this new sacrum to an
undetermined styracosternan ornithopod.
55
A NEW GNATHOSAURINE (ARCHAEOPTERODACTYLOIDEA, PTEROSAURIA)
FROM THE LATE JURASSIC OF PORTUGAL
1 Museu da Lourinhã, Rua João Luís de Moura 95, 2530-158 Lourinhã, Portugal.
The known global distribution and diversity of pterosaurs demonstrates their success as a group,
and yet their relatively sparse fossil record and often incomplete preservation (outside of
Lagerstätten environments) pose a challenge for further understanding their palaeobiology, when
compared with other vertebrates. Such is the case for the Lourinhã Formation of Portugal, which is
largely recognized for its rich and taxonomically diverse fossil representation of dinosaurs, fishes,
amphibians, marine reptiles, crocodilians, and eggs. Despite this abundance, up to now, pterosaur
material recovered from this deposit has been restricted to scant and often fragmentary isolated
bones and teeth, hindering any confident taxonomic assignments. Recently, a remarkably-sized
dentated rostrum fragment and associated partial cervical vertebrae (ML2554) of a pterosaur were
discovered from the Late Jurassic (Kimmeridgian-Tithonian) of Praia do Caniçal, Lourinhã. The
specimen exhibits features such as a spatulated anterior expansion of the rostrum, a robust comb-
like dentition, and pronounced rims of the tooth alveoli, indicating gnathosaurine affinities. The
presence of this taxon adds yet another ecological niche element to the fluvio-deltaic lagoonal
environment that has been suggested to be representative of Lourinhã Formation in the Late
Jurassic, and contributes to the diversity and distribution of gnathosaurines worldwide.
56
Furthermore, based on its unique tooth and dentary morphology, ML2554 clearly represents a new
taxon, the first named pterosaur species from Portugal.
57
FIRST APPROACH TO LIFE HISTORY OF THE ISLANDER PROLAGUS SARDUS
(LAGOMORPHA) BY STUDYING FEMORAL BONE HISTOLOGY
1Departamento de Física e Ciencias da Terra, Universidade da Coruña, Rúa da Fraga 10, 15008
A Coruña, Spain.
2Dipartimento di Scienze, Università degli Studi Roma Tre, Largo S. Murialdo 1, 00146 Roma,
Italy.
3Institut Català de Paleontologia Miquel Crusafont (ICP), Carrer de les Columnes s/n, 08193
Cerdanyola del Vallès, Spain.
4Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology and
Paleoanthropology, Chinese Academy of Sciences, 142 Xi Zhi Men Wai Da Jie, 100044 Beijing, China.
5CAS Center for Excellence in Life and Paleoenvironment, 100044 Beijing, China.
6College of Earth and Planetary Sciences, University of Chinese Academy of Sciences, 100049
Beijing, China.
Islands hold an outstanding potential for studying evolution due to their geographical isolation and
singular ecological pressures (resource limitation, low extrinsic mortality and high intraspecific
competition). In small mammals, the evolution in insular regimes entails later maturation and larger
body sizes. We decided to evaluate the life history strategy of the islander Prolagus sardus
(Ochotonidae) assessing the femoral bone histology of an ontogenetic series (N = 11), recovered from
the Late Pleistocene Dragonara Cave faunal assemblage (22.39-21.94 ka, Sardinia). Bones were
categorized into 3 ontogenetic groups (juvenile, young adult, and adult) according to the epiphyseal
plates fusion, and bone thin sections were examined under polarized microscopy. The osteohistological
analysis revealed rapid growth rates in juveniles, characterized by fibrolamellar bone, whereas a
decrease in growth rate (parallel fibered and lamellar bone) with secondary remodeling (Harvesian
systems) was noted in adults. Young individuals showed smaller medullar area, because of its smaller
size (p<0.05). LAGs were identified in the outer cortex of juvenile to adult specimens. In young adults
(with at least one unfused epiphysis) a range of 1-3 LAGs was determined, pointing to a delayed
achievement of skeletal maturity. In the larger specimen 8 LAGs were identified, hinting an
58
approximated longevity of 8 years. Considering the size of the Dragonara population (700-750 g), the
observed lifespan is longer than the expected, suggesting a shift towards the slow end of the life history
continuum. This study contributes to understanding the eco-evolutionary responses of small mammals
to insular regimes.
59
SMALL-MAMMAL ASSEMBLAGES FROM THE LATE PLEISTOCENE IN
NORTHERN IBERIA (SOUTHWESTERN EURASIA): ENVIRONMENTAL
CLUES FOR ASCERTAINING NEANDERTHALS’ ECOLOGY
1
Grupo de I+D+i EVOADAPTA (Evolución Humana y Adaptaciones Económicas y Ecológicas
durante la Prehistoria). Dpto. Ciencias Históricas. Universidad de Cantabria. Av. Los Castros 44,
39005, Santander, Spain.
60
NEW SPECIMEN OF DRYOMORPHAN (ORNITHISCHIA, IGUANODONTIA)
REMAINS FROM THE UPPER JURASSIC OF PORTUGAL
3Museu da Lourinhã, Rua João Luis de Moura 95, 2530-158 Lourinhã, Portugal.
4CI2Paleo/Sociedade de História Natural (SHN), Travessa Florêncio Augusto Chagas, nº8B, 2560-
230 Torres Vedras, Portugal.
Upper Jurassic Lourinhã Fm. has yielded a diverse vertebrate fauna dated to the
Kimmeridgian/Tithonian interval. Extremely common are saurischian dinosaurs, although recent
work made by the working team highlighted an over-looked diversity and abundance of
ornithischian and ornithopod dinosaurs. Fieldwork activities of Museu da Lourinhã have unearthed
ML 2700, a partially articulated left hind limb and associated carpal elements. The specimen is
fractured and distorted, however several elements are identifiable and provide a useful taxonomic
signal. The hind limb of ML 2700 comprises a partial tibia, an almost complete fibula, a complete
metatarsal (MT)-I, a complete MT-II, partial MT-III and several phalanges, which articulate with one
another, including three pedal claws. The carpal elements include an isolated partial metacarpal,
and an ungual phalanx. The diagnostic characters of ML 2700, include: a rounded and well
distinguishable cnemial crest that projects cranio-laterally; fibular condyle that projects laterally and
forms a 90° structure together with the caudal condyle; symmetric margins of the proximal end of
fibula and enlarged distal one; reduced splint-like MT-I; MT-II overlaps MT-III dorsally; extremely
61
shortened pedal phalanx III-3 and claw-like pedal claws with well-developed lateral and medial
flanges. Furthermore, the manual ungual is claw-like and strongly arched. This combination of
characters indicates a basal iguanodontian affinity for ML 2700, and it is not consistent with two
taxa identified in Lourinhã Fm, Eousdryosaurus nanohallucis and Draconyx loureiroi. Phylogenetic
analyses, employing Maximum Parsimony and Bayesian Inference, confirmed these results,
recovering ML 2700 at the base of Dryomorpha.
62
A GIANT FRESHWATER TURTLE FROM THE PLEISTOCENE OF BRAZILIAN
AMAZON
1Senckenberg Centre for Human Evolution and Palaeoenvironment at the Eberhard Karls
Universität Tübingen, Hölderlinstr. 12, 72074, Tübingen, Germany.
Large reptiles are common among the Neogene fauna of the Amazon region but disappear after
the Miocene. The giant podocnemidids, including the largest freshwater turtle, Stupendemys
geographicus, are all closely related to the extant Peltocephalus dumerilianus, also from the
Amazon. Here, we present an isolated 25 cm long dentary from the Rio Madeira Formation,
northwestern Brazil, well beyond the 16 cm of the largest known dentary of Stupendemys. The
morphology of the symphysis and extension of the lingual ridge suggest a closer affinity to
Peltocephalus among extant taxa. We took a set of linear measurements from 42 specimens
sampling all the eight extant podocnemidid species, S. geographicus and performed a Principal
Component Analysis (PCA) to further assess its affinities. The PCA results confirm its similarity to
Peltocephalus and S. geographicus, but also support its distinctiveness. Carapace maximum
length estimates (as a proxy for body size) show great uncertainty due to the lack of other giant
specimens with associated lower jaw and carapace, ranging from 2114 to 697 mm, with a median
at 1406 mm (p = 0.0162, R² = 0.8893). Radiocarbon (14C) dating using three samples with
preserved bioapatite provided an age between 14,290 ±45 and 9,060 ±50 cal BP, potentially
63
placing this gigantic turtle within the set of megafauna extinctions coincident with the human
occupation of the Amazon. The paleontological and stratigraphical context of the region is still
poorly understood, and we hope our results provide an exciting trigger to expand research about
the Quaternary of the Amazon.
64
EARLY PLEISTOCENE HIPPOS FROM BARRANC DE LA BOELLA
(TARRAGONA, SPAIN): PHENOTYPIC AFFINITIES AND PALAEOECOLOGY
4Universitat Rovira i Virgili. Departament d’Història i Història de l’Art. Avinguda de Catalunya 35,
43002, Tarragona, Spain.
The study of European hippopotamus (recorded throughout the Quaternary from the Early to the
Late Pleistocene) presents several unresolved questions and a lack of consensus among
specialists. The number of identified taxa, their geographical and chronological distribution, and
the palaeobiological changes in their populations are still hotly debated. This work presents
comparative analyses using descriptive anatomy, linear biometry and geometric morphometrics of
hippo specimens unearthed in the archaeo-palaeontological outcrops of Barranc de la Boella
(Francolí River basin, Spain), in the context of the record of the genus Hippopotamus in Europe
and its putative migrations from Africa. The deposits from these localities attest the presence of
hominids and hippos during the Early-Middle Pleistocene Transition (EMPT), a period of great
climatic shifts. The evaluation of the studied specimens supports the consideration of
Hippopotamus antiquus as the only taxon recorded in Europe from the Early Pleistocene (ca. 2.1
65
Ma) to the mid-Middle Pleistocene (ca. 0.4 Ma), when the current species of common hippo
(Hippopotamus amphibius) dispersed from Africa. Even so, a more detailed understanding of the
large intraspecific variability reported in the genus Hippopotamus is needed in order to
characterise the variation in its populations across different chronologies, geographic locations and
environmental conditions. The detailed review of the distribution of H. antiquus at the EMPT raises
the possibility that the Iberian and Italic peninsulas acted as climatic refugia for its populations
since ca. 0.94 Ma when a substantial increase in global ice volume accompanied the increasing
severity and duration of cold stages.
66
FIRST APPROACH TO THE PALAEOENVIRONMENTAL AND
BIOCHRONOLOGICAL CONTEXT OF THE NEW QUATERNARY SITE OF
CUEVA DE LLAMAZARES (LEÓN, SPAIN) THROUGH THE STUDY OF
MICROVERTEBRATES
Cueva de Llamazares is a karstic cavity located at 1400 m above sea level, in the
southern part of the Cantabrian Mountains (León, Spain). Initial geochemical evidence
and the estimated chronological range for the formation of the cavity and the sedimentary
deposits suggest a good record of the changes in the climate and landscape. Among the
study materials of the cave, a series of sedimentary levels containing skeletal elements
of different vertebrates stand out. In this work, we focus on the study of the 264
microvertebrate specimens collected during the 2020 field campaigns. The work carried
out has generated a digital catalogue; taxa from orders Rodentia, Eulipotyphla,
Chiroptera, Anura and Squamata were identified. The microvertebrate association has
been characterised by calculating the minimum number of individuals, considering the
environmental range for different biological, physical and chemical parameters and
reviewing the biochronology of each taxon. As a result, the presence of this association
reveals climatic and environmental conditions similar to those present nowadays in the
area of Cueva de Llamazares and that the chronological framework of the deposits is
between the Upper Pleistocene and the Holocene. Although this work has limitations,
67
such as the lack of precise geophysical dating of the sedimentary deposits, it opens up a
research path for new studies. The study of the faunal associations from Cueva de
Llamazares and other caves in the north of León will help to gain a deeper understanding
of the ecosystems, faunas and human occupations in the past of this territory.
68
PERISSODACTYLA DIVERSITY IN THE NEW EXCAVATIONS AT THE UPPER
MIOCENE LOCALITY OF PIKERMI (ATTICA, GREECE)
P.V. Filis1*, I.X. Giaourtsakis2, S.D. Sklavounou1, S.J. Roussiakis1, G.E. Theodorou1
Pikermi is a renowned reference locality for the study of late Miocene vertebrates, its long history
of excavation and research activity dating back 1835. Since 2008, a new series of recurring and
stratigraphically controlled excavations has been conducted under the auspices of the National
and Kapodistrian University of Athens and the municipality of Rafina-Pikermi (NKUA-SARG Project
70/3/12977), revealing several new sites numbered PV1 to PV4 (“PV”=“Pikermi Valley”).
Perissodactyls exhibit a notable degree of diversity and population density at the new sites. Equids
are the most common representatives of the group and account for a large portion of the
mammalian remains found. They are commonly attributed to two hipparionine species, the slender
Cremohipparion mediterraneum and the more robust Hippotherium brachypus. The potential
occurrence of additional hipparionine species has been frequently discussed, although based on
limited and debatable evidence; the ongoing study combining both old and new material may offer
valuable insights on this matter. Rhinocerotids, although less common, are rather diverse and
represented by two tandem-horned species, Dihoplus pikermiensis and Miodiceros neumayri, as
well as by the relatively scarce hornless species Acerorhinus neleus. Chalicotheriids are the rarest
among the locality’s perissodactyls. At the new excavations, the typical Pikermian schizotheriine
chalicothere Ancylotherium pentelicum is hitherto represented only by a few postcranial elements.
Based on a handful of specimens, the rare occurrence of a chalicotheriine chalicothere (formerly
referred to as Chalicotherium goldfussi) is currently known from old collections and the Pikermi-
Chomateri site, but remains thus far absent from the new PV-sites.
69
A ROOM OF CIRO'S OWN - BUILDING A HYBRID DIGITAL PALEO SPACE
R. Fistola1, F. Fabbri2, A. Rastelli1*, I. Zingariello1, G. Pepe Muzzurru1, M.Y. Messina1, A.
Giordano2, C. Argenio3, F.O. Amore3
1Department of Engineering, AURUS Research Group, Università degli Studi del Sannio, Piazza
Roma, 21, 82100 Benevento, Italy.
3Department of Science and Tecnology, Università degli Studi del Sannio, Via de Sanctis, 82100
Benevento, Italy.
The developed project, using the potential of an immersive and interactive experience, has been
envisaged for the paleontological site of Pietraroja, a unique paleontological site in the world that
can significantly contribute to the education, and development of the surrounding area, and the
promotion of the paleontological heritage.
This technology, which allows a reconstruction of the environmental context, is useful in
creating an immersive environment by using integrated digital tools such as interactive devices,
projection mapping, and web technology.
The integrated use of different digital tools and software allows transferring the
knowledge content through an immediate, real-time, and non-conventional perception to the user.
To create the CIRO digital model it is necessary to obtain complete information on the anatomical
structure of Scipionyx samniticus and, through an accurate 3D modeling, generate the digital
model of the dinosaur. The same approach it's used to recreate the landscape and the vegetation
species. The complete reconstruction process is constituted by steps that allow, starting from the
3d modelling process, to create a web based virtual live environment, that it could be streamed in
different amplitude and context: from a personal mobile device to an immersive multi
projection system. The aim is to build an immersive digital space in which the visitor will perceive
the original environment of the Scipionyx samniticus and will interact with the Ciro virtual
avatar. Furthermore, the application, relying on web technology, it's accessible from all kind of
device, each feature will be available according with the device capability.
70
HOW DOES A GIANT INSULAR LIZARD FEED? A 3D FINITE ELEMENT
MODEL OF GALLOTIA STEHLINI AND ITS IMPLICATIONS FOR FOSSIL
GIANT LIZARDS
The endemic giant insular lizard Gallotia stehlini (Lacertidae, Gallotiinae) lives in Gran Canaria
(Canary Archipelago, Spain). Different extant and extinct Gallotia taxa are known from each
Canary Island, with Gallotia stehlini the largest living taxon. Pleistocene fossil occurrences of this
gigantic species are also known, being surpassed in size only by the extinct Gallotia goliath. To
understand how the gigantic size of G. stehlini affects its functional cranial structures, here we
present a high-resolution 3D Finite Element model of its skull. Different scenarios of symmetrical
biting, bite position, and muscle actions have been tested in order to investigate the feeding
behavior of this taxon. Our preliminary results suggest that the palate, and particularly the
articulations of the epipterygoid with the pterygoid as well as the quadrate, plays a key role in
cranial stress patterns, but possibly also on bite power and protection of the braincase. Our results
shed light on the feeding ecology of this gigantic reptile, enabling a better understanding of the
effects of increasing cranial bones size, muscle volume and insertion attachment area. Additional
analyses will be required to assess the role of other structures such as the protractor musculature
and cranial sutures. Overall, our analysis opens the door to future inferences on the cranial
mechanics and the paleoecology of the extinct giant Gallotia goliath.
71
THE POSTERIOR BRAINCASE OF A SMALL TYRANNOSAURID
1Department, Leiden University, Sylviusweg 72, 2333 BE, Leiden, the Netherlands.
2Faculty of Geosciences, Utrecht University, Princetonlaan 8a, 3584 CB, Utrecht, the Netherlands.
4Department of Organismal Biology, Uppsala University, Norbyvägen 18A, 752 36, Uppsala,
Sweden.
6Museum of Evolution, Uppsala University, Norbyvägen 16, 752 36, Uppsala, Sweden.
7European Synchrotron Radiation Facility, 71 Avenue des Martyrs, 38000, Grenoble, France.
In 1988, Nanotyrannus lancensis was described as a new tyrannosaurid taxon. However, the
validity of this enigmatic species has been controversial since. Today, Nanotyrannus lancensis is
generally considered to represent either a separate taxon or a juvenile Tyrannosaurus rex.
Although numerous diagnostic apomorphies have been proposed to distinguish Nanotyrannus
from Tyrannosaurus, many of these have been demonstrated to be ambiguous or susceptible to
ontogenetic variability. As such, distinguishing true taxonomic diversity from ontogenetic, sexual,
and intraspecific variability represents a notorious challenge in vertebrate palaeontology that is
exemplified by the case of Nanotyrannus.
A partial posterior braincase representing putative Nanotyrannus lancensis (BHI-6430) was
scanned at the European Synchrotron Radiation Facility (Grenoble, France) to reveal its internal
osteology. Additionally, the distribution, shapes, and dimensions of osteological correlates of
neural and sensory structures were visualised, including the endosseous labyrinths. This allowed
us to non-destructively visualise the sutures within the braincase as well as create three-
dimensional visualisations of the endocast and endosseous labyrinths. Additionally, the
endosseous labyrinths of Arcovenator escotae were visualised for comparative reference.
The referred methods and materials will allow for interpretation and differentiation between
ontogenetic and functional conditions in the posterior braincase of Nanotyrannus lancensis. The
states of the sutural openings have already indicated that BHI-6430 represents a juvenile. In
72
addition, the neural and sensory structures preserved in the specimen will aid in illuminating its
ecomorphological affinity. Reflected on the diagnostic cranial anatomy of Tyrannosaurus rex, these
insights will provide new information on the taxonomic identity and palaeobiology of Nanotyrannus
lancensis.
73
THICK TAILS! NON-AVIAN DINOSAUR CAUDOFEMORAL MUSCLE
ESTIMATION FROM CT-SCAN OF EXTANT TAXA
2Department of Organismal Biology, University of Chicago, 1027 E 57th St, Chicago, Illinois 60637,
United States of America.
74
BIOMECHANICAL AND GROWTH REMODELLING DIFFERENCES BETWEEN
MODERN HUMANS AND NEANDERTHALS
L. Genochio1,2*, L. Albessard-Ball1, F. Landi3, A. Profico1, V. Doronichev4, L. V.
Golovanova4, B. Maureille5, and P. O’Higgins1,2
1Department of Archaeology, University of York, Wentworth way, YO10 5DD, York, United
Kingdom.
2Hull York Medical School, University of York, University Rd, Heslington, YO10 5DD, York, United
Kingdom.
3Institute of Medical and Biomedical Education, St George's University, Crammer Terrace, SW17,
London, United Kingdom.
4Laboratory of Prehistory, St. Petersburg, Russia.
5PACEA, University of Bordeaux Montaigne, 8 Esplanade des Antilles, 33607 Pessac cedex,
Bordeaux, France.
Modern human and Neanderthal faces present a clear divergence in their post-natal ontogenetic
trajectories. Differences in patterns of bone growth remodelling have also been found in these two
species with modern humans presenting bone resorption on the anterior subnasal surface, while
Neanderthals present extensive bone deposition. These differences have been hypothesized to be
related to genetic signals that differ between these species, or to differences in the forces arising
from differences in food processing during post-natal development. Here we test the mechanical
hypothesis.
Using geometric morphometrics and finite element analysis, we assessed the association between
masticatory loadings, facial strains and facial remodelling during postnatal growth and
development in modern humans and Neanderthals.
During biting simulations, strain contour plots show that, in both species, the highest strains are
localised on the anterior maxilla, orbital, and anterior subnasal surfaces and decrease between
infancy and adulthood. However, strains show differences in distribution and magnitude among
age stages and species. Further FEA shows that biting performance varies with age and differs
between species.
These findings confirm that differences exist in loading history throughout post-natal ontogeny
between Neanderthals and modern humans and these partially reflect differences in remodelling
maps. Therefore, we cannot exclude the possibility that differences in the ontogeny of masticatory
75
system performance and load resistance underlie the differences in the distribution of craniofacial
growth remodeling features. Further work is needed to more directly compare cumulative strains
arising from diverse masticatory system loads in both taxa and facial growth remodeling maps.
76
BIAS IN EXTINCT LION RECORD AFFECTS OUR UNDERSTANDING OF THE
SPECIES
2La Brea Tar Pits and Museum, Natural History Museums of Los Angeles County, 5801 Wilshire
Blvd, 90036, Los Angeles, CA, United States of America.
3Department of Earth Science, University of Oregon, 310G Cascade Hall, 97403, Eugene, OR,
United States of America.
Eurasian and American lions faced extinction at the end of the late Pleistocene. Both disappeared
right before the Younger Dryas (12.9 ka BP), concomitant with the extinction of other megafauna in
the Northern Hemisphere. Because these two species shared morphologies and adaptations,
doubts about their taxonomy persisted until genetic analyses demonstrated that indeed, these
species were distinct.
Currently, most of the defining characteristics of the two species are accepted by the scientific
community, but some of them were built on assumptions that are no longer valid. For example, in
comparison to its closest relative in Eurasia, the American lion record is generally poor. Many
specimens lack precise locality data or even a representation of the specimen, and only a few
have been directly dated. Therefore, any attempt at comparison should be taken with caution.
In the present contribution, analogies and differences observed in the datasets of these two extinct
species were described. Data were analyzed with a particular focus on how the biased record is
affecting our understanding of their distribution over North America and Eurasia, interaction with
other species, and resilience to changing climatic conditions.
We noticed that the estimated distribution of Eurasian lion was less purely defined after the Last
Glacial Maximum, where occurrences associated with any absolute ages are only one fifth of older
periods. The American species has only four localities reporting any absolute dates and it is
frequently reported as a single recovery, forcing a range estimation averaged through multiple
local climatic conditions and preventing correlations with other species.
77
STUDY OF THE LATE MIOCENE RHINOCEROTIDAE FROM PIKERMI AND
SAMOS (GREECE) AT THE COLLECTIONS OF THE NATURAL HISTORY
MUSEUM OF VIENNA: PRELIMINARY RESULTS
I.X. Giaourtsakis1*
The celebrated Pikermi and Samos localities have been extensively excavated and studied since
the 19th century, serving as reference faunas for the systematic, biostratigraphic and biogeographic
assessment of Eurasian Late Miocene mammals. The Natural History Museum of Vienna hosts
significant material from both localities. The Pikermi collection (acquired 1854-1863) comprises 82
rhinocerotid specimens, 89% identifiable at species level. The Samos collection (acquired in 1911)
comprises 76 rhinocerotid specimens, 88% identifiable at species level. The majority of the
material from both localities consists of postcranial elements (87% in Pikermi and 89% in Samos),
clearly suggesting an origin from dedicated excavations and not selective purchases, which are
biased towards well-preserved craniodental specimens. The most notable craniodental specimens
include a relatively well-preserved cranium of Dihoplus pikermiensis from Pikermi; two fairly
complete crania of Chilotherium schlosseri, a partial juvenile cranium of Miodiceros neumayri, and
an exceptionally well-preserved juvenile maxilla of Dihoplus pikermiensis from Samos. The Pikermi
collection comprises only the two tandem-horned species Dihoplus pikermiensis and Miodiceros
neumayri; the rare hornless rhinocerotid Acerorhinus neleus is not represented. The Samos
collection includes both aforementioned horned taxa and the hornless Chilotherium schlosseri. The
marked differences observed in the relative distribution and abundance of both horned and
hornless rhinocerotid taxa among the Turolian localities of Greece seem to have been influenced
by environmentally controlled provincial differences. The relatively slenderer and lower-crowned
Dihoplus and Acerorhinus seem to have preferred more closed and temperate niches, whereas the
more robust and higher-crowned Miodiceros and Chilotherium have favored more open habitats.
78
CLIMATIC NICHES AND DISPERSAL OF MIOCENE HOMINIDS
3Laboratory IPGP, Université Paris Cité, CNRS, Université de Paris, Paris, France.
4Laboratory LSCE, Institut Pierre Simon Laplace (IPSL), UMR 8212, Université Paris-Saclay,
91191, Gif-sur-Yvette, France.
Several scenarios for the dispersal of Miocene hominids from Africa to Eurasia still coexist. The
reason why it remains impossible to decide between these scenarios, and why some fundamental
questions about the biogeographic history of Miocene hominids are so difficult to address, lies in a
fragmented fossil record and uncertain dating. In order to overcome these problems and to
measure the potential effect of climate on the dispersal of Miocene hominids, we use ecological
niche models (ENM) based on palaeoclimatic reconstructions of Early, Middle and Late Miocene.
Using ENM, we demonstrate that the Eurasian climate during the Lower Miocene was not
compatible with the climatic conditions required for the installation of hominids living in East Africa
at the same time. Consequently, while the connection between Africa and Eurasia was active
since the Gomphoterium Datum Event (PDE, ca. 19 myr), the presence of hominids in Eurasia
appears unlikely during Early Miocene. In the Middle Miocene, the long-term cooling following the
Middle Miocene Climatic Optimum profoundly modified the Earth's climate, notably in Eurasia.
South Asia and Southeast Asia, as well as the Mediterranean coastline, became suitable for the
presence of Late Miocene East African hominids. Finally, we provide a case of ecological niche
displacement. The estimated climatic niches for European hominids in the Middle Miocene are
79
distinctly different from those of their Lower Miocene ancestors from East Africa. These arising
taxa found in Europe did not simply 'track' their climatic niche, but adapted to the new climatic
conditions of the European continent.
80
A REVIEW OF THE URSUS ETRUSCUS REMAINS FROM THE LATE
VILLAFRANCHIAN OF EUROPE WITH EMPHASIS ON NEW FINDS
1Institute of Plant and Animal Ecology, Ural Branch, Russian Academy of Sciences, 8th March
Street 202, 620144 Yekaterinburg, Russian Federation.
Since the pioneering works of Björn Kurtén from the middle of XX century, the Eurasian Early
Pleistocene species Ursus etruscus is considered the putative ancestor of both cave and brown
bears. However, in the last two decades the remains of the late Early Pleistocene has been in a
hotly phylogenetic and taxonomic debate and several authors included it alternatively in the brown
or cave bear lineage.
Here we provide a complete synthesis of the latest published remains from U. etruscus from
Taurida cave, the Orce sites, Dmanisi, Pietrafitta and Pirro Nord comparing it with the classical
material of Saint Vallier, Olivola, Upper Valdarno and Kuruksai.
Our results support the distinction of two different morphotypes of U. etruscus throughout its long
chronologically record, as stated by previous authors: an early forms from the Middle Villafranchian
(e.g. Saint-Vallier or Kuruksay) and late forms from Olivola, Dmanisi, Taurida cave, Pietraffita,
Upper Valdarno or Venta Micena. Later on, at the end of the Early Pleistocene and roughly coeval
with the climatic shifts associated with the ‘Early-Middle Pleistocene Transition’ the earliest large
and stout specimens of speleoid bears (Ursus deningeri) were recorded in Europe at sites such as
Le Vallonnet, Vallparadís Section or Untermassfeld as exemplified by their more derived
characters an overall larger dimensions.
81
UNRAVELLING THE LOCOMOTION OF REPTILES: CONTRIBUTION OF
FEMORAL MICROANATOMY AND PALEOBIOLOGICAL IMPLICATIONS
1Centre de recherche en paléontologie – Paris, UMR 7207, Sorbonne Université, Muséum national
d’histoire naturelle, Centre national de la recherche scientifique, 8 rue Buffon, 75005 Paris,
France.
3Structure and Motion Laboratory, Department of Comparative Biomedical Sciences, The Royal
Veterinary College, Hatfield, United Kingdom.
The water-to-land transition represents a key event in the evolution of tetrapod vertebrates. The
mechanical constraints inherent to this new environment led amniotes to develop novel locomotor
strategies that granted them access to new ecological niches, thus contributing to the evolutionary
success of the clade. Today, amniotes show great locomotor and postural diversity, especially in
reptiles, whose extant representatives include erect bipeds (birds), "semi-erect" quadrupeds
(crocodilians) and sprawling quadrupeds (lepidosaurs and turtles). But the various events that led
to such diversity remain enigmatic and the locomotion and posture of many extinct species, such
as the Triassic Euparkeria capensis (archosauriform) and Marasuchus lilloensis (dinosauriform),
raise questions. The study of reptile microanatomy can help address these uncertainties. Indeed,
this locomotor and postural diversity is accompanied by great microanatomical disparity. We show
how microanatomical parameters measured in cross-section, such as bone compactness and the
position of the medullo-cortical transition, can be related to locomotion and posture in reptiles.
Using statistical methods accounting for phylogeny (phylogenetic flexible discriminant analyses),
we develop several inferential models from a sample of femur cross-sections belonging to 51
reptilian species. We use these models to infer the locomotor mode and posture of 7 extinct reptile
taxa. Our study contributes to the understanding of the evolution of locomotion in various early
reptile species. Our models and methods could be used by palaeontologists to infer locomotion,
including posture, in other extinct reptilian taxa, especially when considered in combination with
other lines of evidence.
82
EVOLUTION OF CRANIOCERVICAL MUSCULATURE IN ABELISAURIDAE
(DINOSAURIA: THEROPODA)
3Área Laboratorio e Investigación, Museo Municipal “Ernesto Bachmann”, Dr. Natali s/n,
Q8311AZA Villa El Chocón, Neuquén, Argentina.
5Department of Organismal Biology, University of Chicago, 1027 E 57th St, Chicago, Illinois
60637, United States of America.
Abelisaurids were dinosaurs that diversified specially in Gondwana during the Cretaceous. They
were medium to large sized theropods, with reduced forelimbs, highly modified cervical vertebrae
and sometimes had horned skulls. Their fossil record is abundant and well-represented making
possible to assess the evolution of craniocervical musculature in Abelisauridae.
We have mapped eight characters, corresponding either to correlates of musculature or to the
motion capacities (moment arms, range of motion...) in recent phylogenetic hypotheses. Five of
these characters are related to cervical ribs and vertebrae and three more to the skull. Characters
were optimized following the principle of maximum parsimony considering Ceratosaurus as the
taxon with the simplesiomorphic condition. These conditions are basal, derived or unknown.
First derived characters occur in Carnotaurinae: (i) epipophyses taller than neural spine, (ii)
paraoccipital processes inclined less than 45 degrees (dorsal view) that provide a big moment arm
for dorsiflexion and dorso-lateroflexion, (iii) absence of ventral keel in posterior cervical and
anterior dorsal vertebrae which implies a smaller insertion for neck ventriflexors, and (iv) the
presence of aliform process in cervical ribs. Range of motion for torsion movements increased due
to (v) wider prezygapophyseal facet joints.
83
These derived characters were present in Brachyrostra and there were indications of (i) widening of
neck base, producing greater levers for lateroflexion and (ii) subhorizontal paraoccipital processes,
granting greater moment arms. Finally, Furileusauria evolved (i) nuchal crests taller than wide.
Craniocervical region evolution enabled abelisaurids greater moment arms for torsion, dorsiflexive
and lateroflexive movements than their earlier branching relatives.
84
NOT ALL ARE CROCODILE BITES: SHELL PITTING ON A NEOCHELYS
TURTLE FROM THE EOCENE OF ZAMORA (SPAIN)
1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Avda. Esparta s/n, 28232, Las Rozas,
Madrid, Spain.
2Colección de Vertebrados Fósiles de la Cuenca del Duero. Departamento de Geología, Facultad
de Ciencias, Universidad de Salamanca, Salamanca, Spain.
Shell pitting is one of the most common abnormal condition identified in extinct turtles and, at the
same time, one of the less understood. References to shell pitting in extant turtles do not usually
include morphological descriptions, illustrations, or data concerning causal agents, etiology or
pathogenesis. In the case of the fossil record, available interpretations on shell pitting are generally
speculative or poorly justified. In this sense, in the literature regarding extinct turtles, pre-mortem
bite marks are the most frequently proposed causative agent to explain shell pitting. As a
consequence, other potential causative agents (e.g., parasitism or bacterial and fungal infections)
have rarely been contemplated.
The anomalies on the shell of an individual determined as Neochelys sp. (Pleurodira,
Podocnemididae) from the middle Eocene (Lutetian) of Corrales del Vino (Zamora, Spain) are
discussed in detail. The anomalies of the specimen were interpreted in a previous study as
traumatic injuries result from a crocodile attack. The lesions produced by the predator were
interpreted as non-fatal for the turtle specimen. The re-study of these anomalies through physical
examination and differential diagnosis procedures, allows proposing alternative causal agents.
This type of analysis indicates an increase in the complexity in the interpretation of the shell pits in
fossil remains of turtles than that which can be generally deduced from the available literature.
85
THE IMPACT OF GIANT GROWTH: A COMPARATIVE FINITE ELEMENT
STUDY OF EXTANT AND EXTINCT INSULAR DORMICE
2Hull York Medical School, PalaeoHub, University of York, Wentworth Way, York, YO10 5DD,
United Kingdom.
Keywords: Finite Element Analysis, Insular Gigantism, Gliridae, Functional Morphology, Island
Evolution
Rodents on islands often drastically adjust their body size and shape as a response to their
isolated environment. Small mammals, like dormice (Gliridae), appear to increase in size
significantly on various Mediterranean islands. The largest extant dormouse population (Eliomys
quercinus ophiusae) is present on the Balearic island of Formentera, Spain. Although significantly
larger than its mainland relatives, these insular giants are relatively small in comparison to fossil
dormice on islands like Mallorca (Hypnomys spp.) and Sicily (Leithia sp.). The mandibular shape of
these dormice vary significantly, and statistical shape analyses indicate only part of these
morphological variations can be explained by allometry. Variation in preferred diet and foraging
techniques are expected between insular populations and could influence skull morphology.
MicroCT data of both giant and normal sized dormouse mandibles used in this study allowed for
evaluating the effects of shape change on the biomechanical performance within the masticatory
apparatus. Finite Element Analyses specified which mandibular morphologies are better adapted
to either chewing or gnawing at the incisors. Our results indicate the mandibular shape to have a
direct effect on the masticatory apparatus, predicting faunivory in some giants and herbivory in
others.
86
FINDING THE TRACEMAKER OF LARGE-SCALE CISURALIAN
VERTEBRATE BURROWS
2Institut für Geologische Wissenschaften, Freie Universität Berlin, Malteserstraße 74-100, 12249,
Berlin, Germany.
3Institut für Biologie, Humboldt-Universität zu Berlin, Invalidenstraße 42, 10115, Berlin, Germany.
Burrowing is one of the most widely distributed behaviors among extant terrestrial vertebrates,
yielding several ecological benefits, such as providing microhabitats with stable environmental
conditions for aestivation and breeding, or subterranean foraging. In the fossil record fossorial
lifestyles are best documented by the presence of burrowing structures. Evidence for fossorial
lifestyles among vertebrates ranges as far back as the Devonian. However, large scale tetrapod
burrows, especially in the Cisuralian are scarce, and information about possible producers are
practically non-existent. The Cisuralian Bromacker locality near Tambach-Dietharz (Thuringia,
central Germany) with its exceptional preservation of both trace- and body fossils, presents unique
insights into terrestrial ecosystems of the early Permian. Among the trace fossils are one of the
oldest known large-scale vertebrate burrows (width up to 50 cm). The observed structures,
previously informally named “Megatambichnus”, include helical shafts, horizontal tunnels, and
terminal chambers. The infilling is almost identical to the host rock and burrow walls are often not
clearly visible. These burrows show extensive scratch marks, including bilaterally-arranged sets of
up to 5 scratches. We try to find possible tracemaker candidates for those burrows, by (1)
comparing scratch set and burrow measurements with body measurements of the taxa found in
the Bromacker locality, and (2) discussing possible fossorial lifestyles in those taxa based on
comparisons with osteological adaptations for fossoriality found in extant and extinct fossorial
animals.
87
MOSASAUR FEEDING ECOLOGY FROM THE BEARPAW FORMATION,
ALBERTA, CANADA: PART II
F.M. Holwerda1*
1Royal Tyrrell Museum of Palaeontology, T0J0Y0 Drumheller, Alberta, Canada.
The Campanian Bearpaw Formation of southern Alberta, Canada, holds very well preserved
specimens of mosasaurs (Mosasaurus missouriensis, Prognathodon overtoni, and Plioplatecarpus
primaevus) and well-preserved specimens of elasmosaurs, turtles, Enchodus, lobsters, cuttlefish,
and bivalves, representing a relatively complete ecosystem north of the Western Interior Seaway.
Here, the ecosystem interrelationships of all these animals, and particularly those of the most
common mosasaurs, are studied via two-dimensional dental microwear analysis, Energy-
dispersive X-ray spectroscopy, and isotope analysis. Teeth of all vertebrates, as well as turtle
carapace, and lobster, cuttlefish and bivalve shells were selected for analysis from the collections
of the Royal Tyrrell Museum of Palaeontology in Drumheller,
The 2D dental microwear analysis reveals a generalist tooth wear for Mosasaurus (equal amounts
of pits, gouges, and scratches), a more heavy wear for Prognathodon (more gouges and pits) and
a variety of both softer wear (fine scratches) and moderate wear (gouges and pits) for
Plioplatecarpus. Coupled with EDX Sr/Ca and Ba/Ca measurements, Mosasaurus pairs with
piscivorous Plioplatecarpus, elasmosaurs and sharks, while Prognathodon pairs with durophagous
sawfish.
Carbon and Oxygen isotope analysis shows, interestingly, the smallest range for Mosasaurus,
indicating a less wide array of prey than its microwear and EDX results predicted. Prognathodon
shows the widest range. Plioplatecarpus, elasmosaurs, and Enchodus show similar δ13C values;
sharks and sawfish less negative values, and lobsters and cuttlefish more negative values.
Prognathodon shows the highest range and slightly higher measurements of δ18O, indicating
perhaps different ingested salinity levels, and therefore a different foraging area for this mosasaur.
88
EARLIEST EVIDENCE FOR FRUGIVORY AND POTENTIAL SEED
DISPERSAL BY BIRDS
1Department of Earth Sciences, University of Oxford, Oxford, OX1 3AN, United Kingdom.
2Zoology Division, School of Environmental and Rural Sciences, University of New England,
Armidale, NSW 2351, Australia.
3Institute of Geology and Paleontology, Linyi University, Linyi, Shandong, 276000, China.
4Field Museum of Natural History, Chicago, Illinois 60605, United States of America.
5Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology
and Paleoanthropology, Chinese Academy of Sciences, 10010 Beijing, China.
6Chinese Academy of Sciences Center for Excellence in Life and Paleoenvironment, 10010
Beijing, China.
7Australian Centre for Neutron Scattering, Australian Nuclear Science and Technology
Organisation, Sydney, Australia.
The Early Cretaceous diversification of birds was a major event in the history of terrestrial
ecosystems, occurring during the earliest phase of the Cretaceous Terrestrial Revolution.
Frugivorous birds play an important role in seed dispersal today, and may have done so since their
origins. However, evidence of this has been lacking. Jeholornis is one of the earliest-diverging
birds, only slightly more crownward than Archaeopteryx, but its cranial anatomy has been poorly
understood, obscuring diet-related functional interpretations. Originally hypothesised to be
granivorous based on seeds preserved as gut contents, this interpretation has become
controversial. We conducted high-resolution synchrotron tomography on an exquisitely preserved
new skull of Jeholornis, revealing remarkable cranial plesiomorphies combined with a specialised
rostrum. We use this to provide a near-complete cranial reconstruction of Jeholornis, and exclude
89
the possibility that Jeholornis was granivorous, based on morphometric analyses of the mandible
(3D) and cranium (2D), and comparisons with the 3D alimentary contents of extant birds. We show
that Jeholornis was at least seasonally frugivorous, providing the earliest evidence for fruit
consumption in birds, and indicating that seed dispersal was more likely to be present from early in
the avian radiation. As highly-mobile seed dispersers, early frugivorous birds could expand the
scope for biotic dispersal in plants, and may explain, in part, the subsequent evolutionary
expansion of fruits, indicating a potential role of bird-plant interactions in the Cretaceous Terrestrial
Revolution.
90
MEDIUM AND SMALL-SIZED SUINES (SUIDAE, MAMMALIA) FROM THE
LATE MIOCENE TO EARLY PLIOCENE OF EUROPE
A. Iannucci1*
1Department of Earth Sciences (PaleoFactory lab.), Sapienza University of Rome, Piazzale Aldo
Moro 5, I-00185 Rome, Italy.
Suids (Suidae, Mammalia), or simply the pigs, are a primitive-looking family of even-toed hoofed
mammals widespread across the Old World. Most, if not all, living species belong to Suinae, a
subfamily that originated during the late Miocene and became dominant soon thereafter. The late
Miocene to early Pliocene is therefore a crucial period for investigating the dynamic of the success
of Suinae, which occurred within a broader context of renewal of mammalian paleocommunities.
Nonetheless, the paucity of complete cranial remains and the overall plesiomorphic anatomy of
suids pose challenges in elucidating their evolution. Here, I present the results of an ongoing
research focused on small and medium-sized species of European Suinae, including the
description of new material as well as the revision of historical collections. Three good species are
recognized during the late Miocene to early Pliocene of continental Europe, with no convincing
evidence of chronological overlap between them: Propotamochoerus palaeochoerus,
Propotamochoerus provincialis, and Sus arvernensis. Propotamochoerus palaeochoerus was a
typical element of early Vallesian (MN 9) ecosystems of Europe, but it disappeared after the so-
called (mid-)Vallesian Crisis (~9.7 Ma), along with many other species. Propotamochoerus
provincialis is recorded in Turolian (MN 11-MN 13) assemblages, which document the presence of
faunas adapted to open environments. The early Pliocene (Ruscinian) witnessed a return to more
humid condition and the concurrent spread of Sus arvernensis. During the considered period,
European suids experienced a significant decline in diversity, especially at the expense of non-
Suinae.
91
UPDATING THE THEROPOD RECORD FROM THE EASTERN CAMEROS
BASIN: SPINOSAURID REMAINS OF THE LOWER CRETACEOUS ENCISO
GROUP (LA RIOJA, SPAIN)
2Centro de Interpretación Paleontológica de La Rioja, Calle Mayor 10, 26525, Igea, Spain.
5Universidad Nacional de Río Negro-IIPG, Avenida Roca 1242, 8332, General Roca, Argentina.
The La Rioja sector of the Cameros Basin (northern Spain) is world-renowned for the abundance
and good quality of preservation of its Lower Cretaceous dinosaur tracksites. In addition,
explorations and digs carried out in recent years have led to new finds of theropod skeletal
material. Nearly all theropod fossils recovered in Eastern Cameros so far come from the Enciso
Group of near Igea village, which is dated as late Barremian-Aptian. These remains consist of
three partial skeletons that can be confidently assigned to Spinosauridae, most likely to
Baryonychinae, from Costarrey-1, Marino-2 and Virgen del Villar-1 sites. The specimen recovered
from Costarrey-1 consists of some caudal vertebrae, ribs and fragments of the pelvic girdle; and
that from Virgen del Villar-1 mostly comprises appendicular bones from the left hindlimb. The
Marino-2 fossil is the most complete specimen. It preserves several cranial bones and elements
from the axial skeleton, pelvic girdle and hindlimbs. Aside from these remains, more fragmentary
material found in the Enciso Group in Igea consists of a partial maxilla and isolated teeth. Studies
based on the maxilla and teeth suggest that they belong to a baryonychine spinosaurid different
92
from Baryonyx. Furthermore, two tooth morphotypes have been distinguished among the sample,
which could indicate dental variation or the presence of two different species. Therefore, future
research may lead to the description of new spinosaurid taxa and to better understanding of the
diversity of Spinosauridae not only in the Cameros Basin, but also in Europe as a whole.
93
INVERTEBRATE BIOEROSION ON MESOZOIC MARINE REPTILE FOSSILS
1Earth Sciences, University College London, Gower Street, WC1E 6BT, London, United Kingdom.
2Natural History Museum, Cromwell Road, SW7 5BD, London, United Kingdom.
Invertebrate assemblages have been recovered in association with whale carcasses in both the
present day and in the fossil record. These whalefall communities are highly specialized for
exploiting the nutrient influx produced by a large vertebrate fall. A few instances of organisms
occupying this type of environment have been described from Mesozoic marine reptiles. Although
it is possible that marine reptile falls provided a similar environment to whalefalls for specialist
invertebrate taxa, there has been little focus on these Mesozoic faunas. Several whalefall taxa
likely originated in the Cretaceous, raising questions of how the community of invertebrates
associated with a vertebrate fall changed across the Jurassic-Cretaceous and Cretaceous-
Paleogene boundaries. In this study, marine reptile fossils from three major UK collections were
examined for traces of bioerosion. A variety of borings and surface traces were consistently
identified on these fossils. Echinoid and gastropod grazing traces are present through the Jurassic
and Cretaceous, and microborings emerge in more oxygenated Jurassic formations, while bivalve
borings are more common in the Cretaceous specimens. This preliminary identification of
ichnotaxa associated with Mesozoic marine reptiles lays the groundwork for describing and
identifying taxonomic assemblages associated with marine reptile dead falls more widely. This
study and those that follow will provide a snapshot of the invertebrate taxa present at a Mesozoic
marine reptile fall and inform us on the ecological niche occupied by these specialist taxa through
deep time.
94
TRACKWAY-DATA-BASED RECONSTRUCTION OF LOCOMOTION
EVOLUTION IN EARLY TETRAPODS
1Museum für Naturkunde Berlin, Leibniz-Institut für Evolutions- und Biodiversitätsforschung, Berlin,
Germany.
Considering the large anatomical range of known late Paleozoic tetrapod postcrania, some of the
variability among contemporaneous tetrapod tracks can be attributed to evolutionary changes in
trackmaker locomotion style close to the origin of amniotes. In a study including Carboniferous
(10%; 19 out of 186) and early Permian trackways, we tried to infer steps of locomotion evolution
close to the origin of amniotes from trackway measurements by means of ancestral state
reconstruction. Due to the imbalanced sample, imperfect track-trackmaker correlation and certain
unresolved controversies about tetrapod phylogeny, many of our preliminary results regarding the
timing, correlation and presumable convergence of locomotion evolution steps were not
conclusive. With the inclusion of additional Carboniferous records of the ichnotaxa Hylopus,
Notalacerta, Dromopus and Varanopus, we gain a better data coverage for the earlier part of the
trackmaker tree, while consideration of several alternative trackmaker phylogenies provides an
indication for the robustness of our conclusions about character evolution. Furthermore, the ages
assigned to certain track assemblages and node ages of our trackmaker trees have been updated
according to recently published data. Our new results suggest that some functionally relevant
features, such as the appearance of narrow gauges, large body sizes and gaits without ground
contact of the belly or tail, are seemingly not as clearly linked to evolutionary change within the
series of common amniote ancestors as we had assumed before. Instead, group-specific patterns
and convergent evolution in diadectomorph, synapsid, parareptile and eureptile track producers
contributed greatly to the observed variability.
95
THE “GHOST” THEROPODS OF THE BAHRIYA OASIS AND THEIR BEARING
ON CRETACEOUS AFRICAN THEROPOD FAUNAS
M. Kellermann1*, O. Rauhut1,2,3
The Cenomanian of North Africa is famous for its large theropod taxa. Several of these, such as
the iconic Spinosaurus and Carcharodontosaurus, were first described from the Egyptian Bahariya
Formation. In recent years, new material from the presumably roughly contemporaneous
Moroccan Kem Kem Group was claimed to be conspecific with some of the Egyptian taxa. Taxa
claimed to be present in both units include Deltadromeus, Carcharodontosaurus,
Sigilmassasaurus, and Spinosaurus. However, no detailed comparisons were presented between
the Moroccan and Egyptian fossils, as the latter were destroyed during World War II. A detailed
reexamination of the North African theropods, with an emphasis on the carcharodontosaurids and
the Moroccan Deltadromeus, reveals that the two faunas are not as similar as previously thought.
The Egyptian carcharodontosaurid is recognized as a basal carcharodontosaur distinct from the
Moroccan neotype of Carcharodontosaurus saharicus, with the latter being recovered within the
traditionally South American clade Giganotosaurinae. Deltadromeus is here recognized as distinct
from any of the Egyptian theropod specimens. Instead, it is recovered as a sister taxon to the
South American Gualicho and both were found to be basal ornithomimosaurs. These results
emphasize the connection between Moroccan and South American theropods while calling
previous comparisons between Egyptian and Moroccan theropods into question. This is consistent
with the paleogeographical situation in the early Late Cretaceous of Gondwana, as Morocco and
Egypt were separated from one another by the Trans Saharan Seaway, while Northwest Africa
remained connected to South America until about 100 million years ago.
96
NEW TETRAPOD TRACKSITE WITH EXTENSIVE TRACKWAYS FROM THE
MOENKOPI FORMATION (MIDDLE TRIASSIC, ANISIAN) OF
NORTHEASTERN ARIZONA: IMPLICATIONS FOR ICHNOTAXONOMY,
ARCHOSAUR COMMUNITIES AND PALEOECOLOGY
H. Klein1*, S.G. Lucas2, E.J. Kappus3, J. Cancellare2, J.N. Lallensack4, P. May2, J. Lang5,
T.J. Olson6
2New Mexico Museum of Natural History, 1801 Mountain Rd. NW, Albuquerque, NM 87104,
United States of America.
3Southwest University at El Paso, 1414 Geronimo Dr, El Paso, 79925 United States of America.
4School of Natural Sciences and Psychology, Liverpool John Moores University, James Parsons
Building, Bryon Street, Liverpool L3 3AF, United Kingdom.
The Moenkopi Formation in the North American Southwest yielded a diverse tetrapod fauna with
body and ichnofossils, being a famous reference for Triassic vertebrate paleontology and
ichnology since the mid-twentieth century. Here we document a recently discovered large tracksite
in the Holbrook Member (Anisian) of northeastern Arizona comprising the most extensive Middle
Triassic footprint material from North America, about 50 trackways and 5000 imprints on a surface
of 75 m2. The site is about 5 m above the base of the Holbrook Member and 18 m below the
overlying Upper Triassic (Carnian) Shinarump Formation of the Chinle Group. The footprint
assemblage consists of Chirotherium barthii, C. rex, Isochirotherium marshalli, Synaptichnium
diabloense, S. pseudosuchoides and Rotodactylus cursorius. Synaptichnium and Rotodactylus are
most abundant at the site. The presence of different size-classes of all chirothere taxa suggests a
mixed archosaur community with individuals of different age. The assemblage confirms the
Holbrook Member to belong to the Chirotherium barthii biochron, and the lack of Chirotherium
sickleri is further evidence of paleobiogeographic peculiarities in the composition, if compared to
97
European early Anisian ichnoassociations. Remarkable is the lack of non-archosaur footprints, for
example Rhynchosauroides and therapsid tracks. Associated invertebrate traces are Arenicolites,
Gordia, Scoyenia and Taenidium, indicating the Scoyenia ichnofacies and an intermittently
subaerial/subaqueous setting on a nonmarine riverine floodplain environment. The track-bearing
stratum is a 0.2-0.4 m thick bed of sandstone with climbing ripple laminae. This thin, tabular and
extensive bed with ripples is interpreted as a sheetflood (unchannelized flow) deposit on the
floodplain.
98
ORAL PROCESSING IN EARLIEST TERRESTRIAL HERBIVORES
RECONSTRUCTED THROUGH TOOTH WEAR AND 3D MODELING
Diadectids are among the earliest clearly identified tetrapod herbivores in the fossil record. This
diagnosis is mainly based on their suspected capability for efficient oral processing of plant fodder
using strongly widened posterior (molariform) teeth and heavy jaws. While coeval localities
preserve mostly faunivorous tetrapods, diadectids are the most abundant vertebrate fossils at the
Cisuralian (lower Permian) Bromacker Locality (Thuringia, central Germany), with Bromacker
representing the earliest example of our modern herbivore-dominated ecosystem structure.
However, evidence for oral processing and thus herbivory in diadectids has never been
demonstrated beyond morphological descriptions. Here we present a hypothesis for the movement
of diadectid jaws in oral processing based on an analysis of attrition marks and wear facets
evaluated using scanning electron microscopy (SEM) on both molariform and incisiform teeth as
well as 3D reconstruction of skull bones and teeth. While the main orientation of attrition marks is
focused in the orthal direction, a minor component of tooth wear supports a fore-aft movement
(proal or palinal). Retrodeformed 3D skull reconstructions of the two exceptionally preserved
Bromacker species Diadectes absitus and Orobates pabsti allow for more detailed reconstruction
of the jaw movements. These results suggest that the origin of efficient oral processing might have
to be predated at least 30 Ma, close to the origin of high fiber herbivory itself and outside the
mammalian lineage. Further studies on metabolic costs might explain why oral processing is not
more widespread among modern herbivorous reptiles and instead mostly restricted to mammals
among tetrapods today.
99
DEFINING AGE AT MATURITY IN THE INSULAR GIANT LEPORID
NURALAGUS REX FROM BONE HISTOLOGY
1Institut Català de Paleontologia Miquel Crusafont, Edifici Z, C/ de Les Columnes s/n, Universitat
Autònoma de Barcelona, 08193 Cerdanyola del Vallès, Spain.
2Institució Catalana de Recerca i Estudis Avançats, Pg. Lluís Companys 23, 08010 Barcelona,
Spain.
Insular dwarfs consistently show a much later age at maturity than predicted from body size
scaling. Little, however, is known about the life history traits that evolve associated to insular
gigantism. Some theoretical works have predicted an increase in age at maturity with increasing
body size in insular giants, but whether these taxa mature later than predicted from body size
scaling remains unknown. To answer this question, we analyzed the bone histology of the tibiae
(n=13) and femora (n=12) of the Menorcan giant leporid Nuralagus rex (Pliocene, Spain). We also
studied in-vivo labelled bones of European brown hare (Lepus europaeus) to calibrate bone tissue
types and bone growth marks with sexual maturity in a close-related extant species. Our results
show that a slow-growing lamellar bone appears in N. rex around the second year of life. The
External Fundamental System, however, is only deposited some years later, which suggests that
the extinct giant leporid grew slower and for a longer time as compared to extant taxa. Indeed,
growth curves obtained from measurements of bone growth marks show that females of N. rex
achieved sexual maturity around the age of four. In an allometric context, this extinct taxon (body
mass of around 8 kg) is an outlier that reached sexual maturity extremely late for leporids, and
even late for slow-developing bovids of the same body mass. This is the first evidence that giant
insular mammals mature later than predicted from body size scaling, mirroring the pattern of
delayed maturation in insular dwarfs.
100
DEEP TIME INVENTION AND HYDRODYNAMIC CONVERGENCES THROUGH
AMNIOTE FLIPPER EVOLUTION
A. Krahl1*, I. Werneburg1,2
2Senckenberg Centre for Human Evolution and Palaeoenvironment an der Eberhard Karls
Universität Tübingen, Sigwartstraße 10, 72076 Tübingen, Germany.
Plesiosauria (Sauropterygia) are secondarily aquatic diapsids. From the Late Triassic to the end of
the Cretaceous, plesiosaurs inhabited the oceans globally. A special feature of plesiosaurus are
their four hydrofoil-like flippers. It has been suggested that plesiosaurs swam by rowing,
combinations of rowing and underwater flying (like Otariinae, Carettochelys insculpta), or by
underwater flying (like Spheniscidae, Chelonioidea, and possibly Cetacea). Rowers use water
drag, while underwater fliers are energy efficiently using lift but at the cost of manoeuvrability. It is
largely accepted that plesiosaurs were underwater fliers. Recently, it has been suggested that for
efficient use of lift during underwater flight, plesiosaurs were able to twist their flippers along the
long axis actively with a special muscular mechanism. Here, the convergent evolution of flipper
twisting in the foreflippers of a plesiosaur and its functional analogues were investigated by
employing anatomical network analysis (AnNA). As functional analogues, taxa with substantial lift-
based propulsion phases were chosen: Caretta caretta (Chelonioidea, Testudines), Spheniscus
demersus (Spheniscidae), Megaptera novaeangliae (Cetacea), Zalophus californianus (Otariinae),
and C. insculpta (Carettochelyidae, Testudines). The AnNA is based on literature data. Bone to
bone and muscle to bone contacts were coded in N x N data matrices. Morphofunctional modules
were detected by running “igraph” in “R” and using a walktrap algorithm. Foreflipper AnNA
revealed that C. eurymerus, C. caretta, and Z. californianus convergently evolved myological
mechanisms for active foreflipper leading and trailing edge twisting. S. demersus actively twists the
leading edge, and C. insculpta and M. novaeangliae cannot actively twist their foreflippers.
101
EVOLUTIONARY HISTORY OF BIRD ECOMORPHOLOGY: USING 3D SCANS
TO ESTIMATE ANCESTRAL STATES OF THE GALLOANSERAN QUADRATE
1 Department of Earth Sciences, University of Cambridge, Downing St, Cambridge CB2 3EQ,
England, United Kingdom.
2 Department of Earth Sciences, University of Oxford, 3 S Parks Rd, Oxford OX1 3AN, England,
United Kingdom.
3 Museum of Zoology, University of Cambridge, Downing St, Cambridge CB2 3EJ, England, United
Kingdom.
In birds and other reptiles, the quadrate acts as a hinge connecting the lower jaw and the cranium,
and plays an important role in avian cranial kinesis. Though previous studies have qualitatively
described its substantial morphological variability, none have tried to quantify evolutionary changes
in its shape. Here, we investigate shape changes of the quadrate in Galloanserae, a major clade of
extant birds uniting relatives of living chickens and ducks. We quantified morphological variation in
the quadrate across 50 extant galloanseran taxa covering all major extant subclades using three-
dimensional geometric morphometrics, and performed ancestral shape reconstructions in the
context of an up-to-date neornithine phylogeny. We find that ancestral state reconstructions based
only on modern quadrates failed to recover plesiomorphic features observed in stem groups, and
instead merely represented the average shape derived from the extant dataset. Therefore, we
strongly encourage adding fossil taxa into three-dimensional geometric morphometric datasets for
ancestral state reconstructions when possible. After incorporating several early fossil galloanseran
quadrates into our analyses (Asteriornis, Presbyornis, and Conflicto), our reconstructed ancestral
galloaneran quadrate approximates aspects of quadrate morphology observed in galliforms,
indicating that the ancestor of Galloanserae might have resembled extant galliforms in its quadrate
morphology and aspects of its jaw biomechanics. The quadrate morphology of Pelagornithidae—
an enigmatic clade of fossil birds potentially allied with Galloanserae—does not closely resemble
our ancestral reconstructions, suggesting that pelagornithids may have exhibited divergent
quadrate mechanics unseen in any other members of total-clade Galloanserae.
102
THE PHYLOGENY OF THE EARLY PALAEOCENE ORDER TAENIODONTA
USING NEW SPECIMENS FROM NEW MEXICO, USA
1School of Geosciences, University of Edinburgh, James Hutton Rd, King's Buildings, EH9 3FE,
Edinburgh, U.K.
2New Mexico Museum of Natural History & Science, 1801 Mountain Rd NW, NM 87104,
Albuquerque, New Mexico, United States of America.
3Carnegie Museum of Natural History, 5800 Baum Blvd., PA 15206-3706, Pittsburgh,
Pennsylvania, United States of America.
After the Cretaceous-Palaeogene (K-Pg) mass extinction mammals thrived in the Cenozoic.
However, the phylogenetic affinities of early Palaeogene ‘archaic’ mammals that lived immediately
after the extinction remain unresolved. Taeniodonta is a group of puzzling ‘archaic’ mammals that
appeared in the early Palaeocene of North America. They are arranged into two subgroups; the
Conoryctidae and Stylinodontidae and are characterised by their extreme degree of dental wear,
indicating an abrasive diet, which led to hypsodonty in the most derived species. Due, in part, to
their worn teeth and their rarity in the fossil record, the position of taeniondonts in the mammalian
phylogenetic tree remains unresolved. New fossils from San Juan basin, New Mexico, USA,
including unworn teeth of four genera and postcranial elements of an early taeniodont, Conoryctes,
shed light on their dental and postcranial anatomy. Both in the forelimb and hind limp of
Conoryctes, there are anatomical adaptations towards fossoriality. Using these specimens, we
scored taeniodonts and other Palaeogene mammals into a phylogenetic data matrix (620
characters, 135 taxa). We then conducted a phylogenetic analysis using parsimony. Our results
show that Taeniodonta is a monophyletic group within Eutheria. We also found that Onychodectes
is basal to the two subgroups previously proposed. Based on the new postcranial fossils and
revised phylogeny, we concluded that digging behaviours were likely ancestral for taeniodonts.
Therefore, a more fossorial mode of life may have been beneficial for their surviving and thriving in
the wake of the K-Pg extinction.
103
FIGHTER OR BLUFFER: COMPARISON OF BENDING AND COMPRESSION
IN MODELS OF THE CAUDAL SPINES OF DACENTRURINE AND
STEGOSAURINE STEGOSAURS
1Dipartimento di scienze della Terra, Università degli studi di Torino, Via Valperga Caluso, 35,
10125 Torino, Italy.
3 Department of Aerospace Science and Technology, Politecnico di Milano, Milano, Italy, via La
Masa 34, 20156, Milano, Italy.
Stegosaurian dinosaurs have a double row of parasagittal osteoderms and a ubiquitous and iconic
set of two pairs of spines at the end of the tail, often referred to as thagomizer. Most derived
stegosaurian taxa are members of two sister clades: Stegosaurinae and Dacentrurinae.
Stegosaurinae includes Stegosaurus stenops, with large dorsal plates; and Dacentrurinae is
represented by Miragaia longicollum and Miragaia longispinus, both characterized by the largest
caudal spines, up to one meter long. The cross-section of the thagomizer spines is different
between these taxa, as the stegosaurine spines are elliptical in cross-section while dacentrurine
ones have a thick sigmoid-like section. These two different morphologies have been compared
and used to estimate the forces withstandable by the spines, under compression and bending
loadings, using a rhomboid shape as a proxy for dacentrurine spines. One estimate was carried
out by attributing the same length and proportions to the two spines and a second one was carried
out by attributing the spines' real length. The two models tested showed that stegosaurine spines
have better performances in withstanding a higher force in bending and compression than
dacentrurine ones. This difference can be explained by the spines of stegosaurine stegosaurs
being better suited for combat than dacentrurine ones, while the latter may have been better suited
for other functions, such as display or intimidation.
104
SIZE REDUCTION AND ITS MORPHOLOGICAL CONSEQUENCES IN
MAMMOTH MOLARS FROM THE PLEISTOCENE OF BRITAIN
A.M. Lister1*
The late Middle to Late Pleistocene saw the transition from Mammuthus trogontherii (‘steppe
mammoth’) to M. primigenius (‘woolly mammoth’) in Europe, with increase in lamellar number in
the molars, and reduction in lamellar and enamel thickness. There were also, in each species,
episodes of significant size reduction, particularly well-documented in Britain. With a metric dataset
of molars from over 40 localities I explore the pattern of size reduction and the consequences on
molar morphology, a complicating factor in tracing the evolutionary transition. Shortening of molars
increases lamellar packing but its effect on the number of lamellae appears variable across
populations. The interplay of size reduction and additional developmental modifications in the
transition between M. trogontherii and M. primigenius is explored. The sporadic occurrence of
analogously small-sized populations in permanently continental areas (mainland Europe and
Siberia) indicates that this is not always technically an ‘island’ phenomenon, but the exploration of
developmental mechanisms in all these instances is relevant to the interpretation of size-related
changes in the island syndrome.
105
THE MIDDLE PERMIAN TRACKSITE OF GONFARON: A COMPLEX
ECOSYSTEM IN THE MIDST OF THE END-GUADALUPIAN MASS
EXTINCTION
The Permian outcrops of Provence yield one of the most extensive records of continental
ichnofossils, reaching from the Kungurian through the Capitanian (early-middle Permian). The
Pélitique Formation of the Gonfaron site (Provence, France) has recently been revised and is
marked by a diverse vertebrate ichnofauna: Batrachichnus salamandroides
(temnospondyls/lepospondyls), Capitosauroides talus comb. nov. (therocephalian therapsids),
Dicynodontipus isp. (cynodont therapsid), Varanopus isp. (bolosaurian parareptiles), Hyloidichnus
bifurcatus (captorhinomorph eureptiles) and Rhynchosauroides isp. (neodiapsid eureptiles). This
ichnoassociation has been dated to the late Capitanian and assigned to the newly introduced
Dicynodontipus sub-biochron. It is also the earliest evidence of the post-dinocephalian extinction
recovery at low latitudes of Pangaea. Moreover, the Gonfaron site is an excellent site for the study
106
of Permian ecosystems: it yielded two rare full-body impressions of Hexapoda that testify certain
ecological strategies, such as mimetics and jumping adaptation, possibly indicating high trophic
pressure in harsh palaeoenvironmental conditions. The trophic pressure was investigated through
the analysis of co-occurrences of fossil traces (arthropod and tetrapod tracks and trackways),
which show that the association of tetrapod and hexapod imprints was relatively common. Also,
the good preservation of the tetrapod tracks of the Pélitique Formation allowed the first
synapomorphy-based correlation between Hyloidichnus tracks and captorhinomorph trackmakers
and a precise reconstruction of the locomotion of captorhinomorph eureptiles. Further studies and
new excavations on the site are planned to provide more data/material and strengthen our
knowledge of this crucial period in the evolution of tetrapods and hexapods.
107
THE FIRST DUCKBILL DINOSAUR (HADROSAURIDAE: LAMBEOSAURINAE)
FROM AFRICA AND THE ROLE OF OCEANIC DISPERSAL FOR LATE
CRETACEOUS AND PALEOGENE VERTEBRATE BIOGEOGRAPHY
N.R. Longrich1*
1Biology and Biochemistry, University of Bath, Claverton Down, BA1 5HY, Bath, United Kingdom.
Following the breakup of Pangaea in the early Jurassic, continents became increasingly
fragmented over the course of the Cretaceous and isolated by a combination of continental drift
and high sea levels, leading to distinct and endemic dinosaurian communities in Northern,
Laurasian continents and the southern, Gondwanan continents. Late Cretaceous Laurasian
communities were dominated by hadrosaurids, ceratopsians, and tyrannosaurs; Gondwanan
communities by titanosaurians and abelisaurs. The presence of hadrosaurids, a lineage endemic
to North America, in the Maastrichtian of Africa raises questions about processes driving
biogeography in the late Cretaceous and early Cenozoic. Similar patterns are seen in other
dinosaur lineages including hadrosaurids and titanosaurians, and in Paleogene mammals. Long-
range oceanic dispersal, via swimming, drifting, or rafting appears to play an infrequent but central
role in structuring dinosaurian communities. Low-probability, high impact “Black Swan” events,
including oceanic dispersal, mass extinction, and complex adaptations, all play a major role in
driving evolution over long timescales. Microevolutionary processes observable on historical
timescales are necessary but insufficient to explain macroevolutionary patterns: macroevolution is
not just lots of microevolution.
108
NEW INSIGHTS INTO TRICERATOPS LIMB BONE HISTOLOGY:
REVEALING SKELETAL UNITY THROUGH PATTERNS IN
HISTOVARIABILITY
2Institute of Biology Leiden, Leiden University, Sylviusweg 72, 2333 BE Leiden, the Netherlands.
3Faculty of Geosciences, Utrecht University, Princetonlaan 8a, 3584 CB Utrecht, the Netherlands.
Annual field expeditions from 2013-2019 to eastern Wyoming, USA by Naturalis Biodiversity
Center yielded a monospecific bonebed comprising disarticulated remains of at least four
individuals of Triceratops (Dinosauria: Ceratopsia). Hitherto, little research has been done on
Triceratops growth. The abundance of post-cranial elements provides an opportunity to study
Triceratops osteohistology in detail. Drill core samples were taken from multiple hindlimb elements
(tibiae and femora) which were subsequently processed into thin sections and studied using
polarizing light microscopy. All elements show laminar parallel-fibred bone and a lack of cyclical
growth marks, a combination of histological traits similar to that seen in (derived) sauropod
dinosaurs. However, there is significant variation in the degree of bone remodelling, despite minor
variation in element size. Semi-quantification of remodelling patterns allows for the assessment of
inter-elemental variation between the different limb bones and histovariability within Triceratops.
The skeletal disarticulation of the site poses challenges when reconstructing the skeletons. The
assessment of histovariability between different elements helps to reconstruct the individual
skeletal unity of the disarticulated remains from the mass death assemblage. Elements from the
same individual are connected through similarities in histological patterns. This study provides a
first detailed analysis of Triceratops limb bone histology and allows for the opportunity to compare
its histology with that of other ceratopsian dinosaurs. Comparisons with the closely related
members of the Chasmosaurinae and more distantly related Centrosaurinae improve our
understanding of ceratopsian growth and development in general.
109
REVIEW AND COMPLETENESS OF THE FOSSIL RECORD OF PALEARCTIC
CAUDATA
1Dipartimento di Scienze della Terra, Università degli Studi di Torino, Via Valperga Caluso 35,
10125, Turin, Italy.
2Department of Earth Sciences, University College London, WC1E 6BT, London, United Kingdom.
3 Research Department of Cell and Developmental Biology, University College London, WC1E
6BT, London, United Kingdom.
The absence of an up-to-date, taxonomically and spatiotemporally inclusive dataset of the fossil
record of Palearctic caudates limits our ability to synthesize information, precluding an evaluation
of the macroevolutionary and biogeographical history of this group. Here, we provide a critical
reappraisal of the published fossil occurrences of caudates from the Palearctic and quantitatively
evaluate the quality of the group’s fossil record. By contrast with the depauperate nature of extant
biodiversity (especially compared to that of the Nearctic), Palearctic Caudata were much more
diverse in the past, with more than sixty extinct species currently recognized. Stem-Urodela were
already widespread during the Middle Jurassic in Eurasia. Cryptobranchoidea, the earliest-
branching sub-order of Caudata, appear to have radiated in Asia, whereas the origin of the other
sub-order of Caudata, Salamandroidea, is unclear. Not all time bins are equally represented in the
record, which is probably a combination of real biological signals and several sampling biases. The
highest apparent diversity of extinct taxa occurs during the Miocene, with a drastic reduction during
the Pliocene, reflecting a common trend in small ‘herps’ in general. We suggest that this pattern is
partly influenced by a sampling bias, with continental Pliocene outcrops less sampled than the
Miocene ones (for geological or anthropic reasons). Most of the Neogene and Quaternary taxa are
known from isolated vertebrae, and the lack of articulated skeletons from these time bins
constitutes a problem for the understanding of their taxonomic identity and the phylogenetic
relationships among the taxa.
110
HOMOTHERIUM FROM THE LATE EARLY PLEISTOCENE OF PIRRO NORD
(APULIA, SOUTHEASTERN ITALY)
Homotherium was one of the most widely distributed felids, both geographically and
chronologically. Records of this genus were found from the early Pliocene to the latest Late
Pleistocene in Eurasia, Africa and America. Late Pleistocene Eurasian and American fossils,
usually considered as representing two different species and well-known thanks to several
complete specimens from Friesenhahn cave in Texas (USA), were recently ascribed to a single
species thanks to ancient DNA evidence. On the contrary, Pliocene and Early Pleistocene
Homotherium from Africa and Eurasia are ascribed to multiple species and remain poorly known,
particularly their postcranial anatomy. In this regard, up to now only one complete skeleton from
Senèze (France) and the collection of isolated finds from the Incarcal Complex (Iberia)
characterize the early forms of Homotherium.
Here we describe Homotherium collection from a single fissure filling (DE11-1) from the Pirro Nord
area in Gargano (southern Italy). It is one of the most diverse Old-World assemblage of
Homotherium composed by an almost complete skeleton of an adult female, a partial skeleton of
adult male, and remains of a juvenile and other 4 individuals, for a total number of 474 specimens.
The adult female skeleton from Pirro Nord is a lioness-sized animal with a slender, long-legged
appearance adapted to prey in open-environments, with powerful cervical area and reduced
lumbar region. In the lumbar region, the female displays several pathologies on the neural arches.
Even other specimens of the sample show other pathologies probably consequence of traumas in
tibias, pelvis and metapodials.
111
ISOLATED THEROPOD TEETH FROM THE TENDAGURU FORMATION
(UPPER JURASSIC, TANZANIA)
1Instituto Dom Luiz, Faculdade de Ciências da Universidade de Lisboa, Campo Grande, 1749-016
Lisboa, Portugal.
The Oxfordian-Tithonian Tendaguru Formation (Tanzania) is still the most relevant reference for
the study of Late Jurassic dinosaur faunas from Gondwana. This unit has yielded an abundant
fossil record that includes sauropods, stegosaurs, ornithopods and theropods. The record of
theropods consists mainly of fragmentary materials, including a large collection of isolated teeth.
The description mainly of postcranial remains allowed the identification of a diverse fauna that
includes at least four ceratosaurian and three tetanuran taxa, some of which also identified based
on isolated teeth. However, the full diversity of tooth morphotypes represented in these levels had
not been previously evaluated. A collection of 167 theropod teeth from the Tendaguru Formation
was analyzed, using a combination of morphometric and cladistic tools. The identified tooth
morphotypes are tentatively assigned to six taxa, including a non-abelisauroid ceratosaur, two
indeterminate abelisauroids, a megalosaurid, an early branching allosauroid, and a
carcharodontosaurid. No specimen could be clearly related to abelisaurids, which were previously
identified based on postcranial materials. This clade might be represented by indeterminate
abelisauroid teeth or by specimens identified as carcharodontosaurids, given the similarity of their
tooth morphology. As with non-dental materials, the greatest abundance and diversity of tooth
morphologies has been identified in the Middle and Upper Dinosaur Members. However, the
distribution of the tooth morphotypes allowed to expand the record of megalosaurids to the Lower
112
Dinosaur Member, increasing the diversity of theropods in this basal unit of the Tengaduru
Formation.
This study was funded by CEECIND/01770/2018, FCT- UIDB/50019/2020 –IDL, and DE-TAF-
2515.
113
THE BROMACKER LOCALITY: A REFERENCE SITE FOR PALAEOZOIC
TETRAPOD ICHNOLOGY
2Institut für Geologische Wissenschaften, Freie Universität Berlin, Malteserstraße 74-100, 12249,
Berlin, Germany.
3Institut für Biologie, Humboldt-Universität zu Berlin, Invalidenstraße 43, 10115, Berlin, Germany.
The Bromacker locality of central Germany (Tambach Formation, Cisuralian) is known since the
late 19th century for its extraordinary fossil footprint content, especially with regards to abundance
and preservation. The project “Opening science: new ways of knowledge transfer using the
example of the research project Bromacker”, funded by the Federal Ministry of Research and
Education, yields new preliminary results. Ichnotaxonomic studies evidence a more diverse
ichnoassociation than previously thought, belonging to the Dromopus biochron. The occurrence of
long trackways allows for studies on track taphonomy and on locomotion/posture of the
trackmakers. Observed features, such as swimming traces, body/tail/digit drag traces and
body/scaly skin impressions, are key for the inference of the trackmakers’ palaeoecology. The
occurrence of complete, articulated autopodia at the same site allows for a thorough track-
trackmaker correlation of the main producer groups. The correlation between footprints and
autopodia evidences features in the skeletal record that are potentially useful for studies on
phylogeny and biomechanics. The vertebrate footprint association was investigated through
different census methods and our results indicate a dominance of diadectomorph tracks, which is
114
consistent with the proportions observed in the skeletal record of the same site. At least two
distinct types of vertebrate burrows have been identified, and a study on the producers of these
structures and their meaning under an evolutionary perspective has been undertaken. The relative
abundance of tracks attributed to herbivores and of vertebrate burrows is currently investigated in
the context of depositional environment, the evolution of terrestrial Cisuralian biota and climate
change.
115
FILLING CRACKS TO IMPROVE THE CONSERVATION OF DINOSAUR
EGGS FROM POYOS (UPPER CRETACEOUS. SPAIN)
2Grupo de Biología Evolutiva UNED, C/ Esparta s/n, 28232, Las Rozas, Madrid,
España.
In the paleontological site of Poyos (Upper Cretaceous. Guadalajara, Central Spain), a large number
of dinosaur eggs distributed along different stratigraphic levels has been recorded. The matrix
surrounding and filling these eggs has between 34% and 76% of phyllosilicates, of which 10% are
smectites, highly expansive clays that, in terms of conservation, are an important risk factor for
deterioration).
The eggs are preserved in volume, showing a surface formed by a succession of shell fragments
that the clay filler holds in position. However, this clay dehydrates after its extraction, losing volume
and originating cracks that can break the structure that supports the egg, causing it to collapse. To
avoid this, it is necessary to fill these cracks using materials compatible with the clays and
analytically discernible (for example in radiological explorations).
For general consolidation of these eggs, it is used Paraloid B72 dissolved in acetone (sometimes by
papering the surface with the adhesive embedded in Japanese paper). To prevent the treatment of
the cracks from interfering with the general consolidation, it is filled with a water and alcohol soluble
resin such as EVA Art and Polyvinyl alcohol, both of which have been tested. On the cracks, the
consolidant was first applied without any type of thickener and once dry, the resin was introduced
with glass microspheres which, due to their homogeneity and size, can be easily injected until the
crack is filled. This treatment, together with the papering of the surface, improves conservation by
eliminating areas of weakness.
116
NEW PRIMATE MATERIAL FROM THE MIDDLE EOCENE FOSSIL SITE OF
CAENES (DUERO BASIN, SALAMANCA, SPAIN)
Caenes (MP15-16) was excavated in the 1980s-1990s, and yielded interesting primate remains
still not studied in detail. This locality is found int the Western Iberian Bioprovince, a part of Iberia
separated from the Pyrenean basins in some time intervals during the Eocene, that presents
endemic faunas. Past revisions of the material showed that an Anchomomys-like primate was
present in the locality, represented by two complete and one fragmentary upper molars, and two
mandible fragments, one showing a complete P/4 and the other presenting an almost complete
P/4, a parcial M/1 and a complete M/2, and both with the rest of the alveoli from the lower C
onwards. A new revision of the material recovered from this site has allowed us to identify more
primate remains. On the one hand, a calcaneus that we attribute to the anchomomyin primate
mentioned above, with an elongared distal part but smaller than the calcaneus of A. frontanyensis,
the only anchomomyin calcaneus described so far. On the other hand, we have identified dental
remains of a much smaller primate, represented by a mandible fragment with M/2 and M/3. Its
morphology suggests that it is an omomyiform with Pseudoloris affinities, smaller than P.
pyrenaicus, similar in size to P. parvulus and P. cuestai, presenting morphological differences with
all of them. Further studies will elucidate the taxonomy of these primates, their phylogenetic
affinities and the locomotor behavior interpreted from the anchomomyin calcaneus.
117
THE ORIGIN OF AMNIOTA IN ITS PHYLOGENETIC CONTEXT
D. Marjanović1*
1Evolutionary Morphology, Dynamics of Nature, Museum für Naturkunde – Leibniz Institute for
Evolutionary and Biodiversity Research, Invalidenstraße 43, 10115 Berlin, Germany
Ever since the first phylogenetic analyses of early limbed vertebrates in the mid-1990s, there have
been exactly two types of such analyses: (1) from the origin of limbs to that of Amniota, but barely
reaching into Amniota; (2) covering amniote phylogeny and barely reaching beyond Amniota. This
makes it impossible to research whether the diadectomorphs or certain “microsaurs” are amniotes.
Moreover, research on amniote-internal problems like the positions of Varanopidae and the
“parareptiles” may be misdirected if too distant outgroups are used (potentially leading to
misrooted, distorted trees); which taxa are best suited as outgroups for amniote phylogeny
depends on the abovementioned phylogenetic questions. I have updated and enlarged a dataset
of early limbed vertebrates by adding amniotes and other taxa along with numerous characters,
and doubled it to account for uncertainty over the "microsaurian" (etc.) tabular-or-supratemporal.
Both matrices place Diadectomorpha and Varanopidae in Pan-Mammalia; the “parareptiles” and
others are scattered. All “microsaurs” are found as stem-amphibians; moving any into Amniota
requires about 30 extra steps. Brouffia, the only pan-amniote reported to retain dorsal dermal
scales, appears to lie on the pan-amniote stem. Redescriptions of taxa and character complexes
will be necessary, though not sufficient, to improve our understanding of amniote origins. My
results also underscore the importance of “microsaurs” as close amniote relatives, and the
closeness of the origins of Amniota and Lissamphibia. Extant amphibians may be better model
organisms for amniotes like ourselves than used to be thought: phylogenetics using fossil data is
important even for biomedical research.
118
NEW INSIGHTS INTO THE HIPPOPOTAMID (MAMMALIA,
HIPPOPOTAMIDAE) FROM THE CASINO BASIN (TUSCANY, ITALY)
1
Department of Earth Sciences, GeoBioTec, School of Science and Technology, FCT-NOVA,
Universidade NOVA de Lisboa, Campus de Caparica, 2829-516 Caparica, Portugal.
3Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via G. La Pira 4, 50121
Firenze, Italy.
Fossil remains from the Casino basin (Tuscany) are known since the beginning of the 19 th century
thanks to the 1879 notes by D. Pantanelli. Among the others, the mammal assemblage includes
Mesopithecus pentelicus, Tapirus arvernensis, Propotamochoerus provincialis and scarce and
poorly preserved remains of a hippopotamid, consisting of a mandibular symphysis fragment, an
apical fragment of a lower canine, some isolated lower incisors, a fragmented second lower
premolar, and a second lower molar. These specimens were initially referred to Hippopotamus
hipponensis (a pigmy species described by A. Gaudry in 1876 from the Early Pliocene locality of
Pont-De-Duvivier, Algeria) and later ascribed to the new species Hip. pantanellii (now
Hexaprotodon? pantanellii) by L. Joleaud in 1920. However, this attribution has been disputed
during the past years. The hippopotamid remains from the Casino basin are here revised in order
to clarify their systematic position and to infer paleobiogeographic and evolutionary patterns within
the Mediterranean fossil record of Hippopotamidae. The canine displays a narrow longitudinal
groove on the mesial side and the enamel is finely striated and crenulated. The lower molar lacks
accessory cuspids and displays a well-developed trefoil wear pattern. The morphology of the
remains collected from the Casino basin more closely resembles the African than the Asian
hippopotamids’ lineage. Hippopotamids’ dispersal in Tuscany probably involved either the Iberian
Peninsula or southern Italy, where the presence of Archaeopotamus crusafonti (Spain) and Hex.?
siculus (Sicily) is well documented, in contrast to Eastern Europe where no Late Miocene site
reports the occurrence of hippopotamids.
119
TETRACONODONTINE SUID REMAINS FROM CA L’ALMIRALL (MN6;
VALLÈS-PENEDÈS BASIN, NE IBERIAN PENINSULA)
The suid dental remains from the late Aragonian (Middle Miocene) site of Ca l’Almirall located in
the Vallès-Penedès Basin (NE Iberian Peninsula), are revised from a taxonomic and
chronostratigraphic perspective by taking into account recent advances in tetraconodontine
taxonomy. The Ca l’Almirall material, including a mandibular fragment and a few isolated upper
and lower cheek teeth, are housed in the Institut Català de Paleontologia Miquel Crusafont in
Sabadell, Spain. They were first attributed to Hyotherium soemmeringi, then to Conohyus
steinheimensis, and most recently to Versoporcus steinheimensis. However, they were never
described in detail and only the mandibular fragment had been figured thus far. Our taxonomic
revision of the material indicates that, except for a listriodontine incisor fragment, most of the
material belongs to the tetraconodontine genus Versoporcus, differing from both Conohyus
simorrensis and Parachelauastochoerus valentini in premolar morphology. Mostly based on size,
we attribute the Ca l’Almirall material to Versoporcus grivensis instead of V. steinheimensis. On
biostratigraphic grounds, the site of Ca l’Almirall is tentatively correlated with the early MN6 (as
conceived in Western Europe), with an estimated age of ~14.0–13.5 Ma that is roughly coeval with
the oldest records of both species of Versoporcus. These species considerably overlap in
chronostratigraphic range and have been considered synonyms by some authors, although based
on size we provisionally support their distinction.
120
CHARTING EVOLUTION, NEW METHODS TO MAP MORPHOLOGICAL
CHANGE UPON THREE-DIMENSIONAL DIGITAL MODELS
1Dipartimento di Scienze della Terra, dell’Ambiente e delle Risorse, Università degli Studi di Napoli
Federico II, Naples, Italy.
121
PALAEOBIOLOGY OF PANTHERA SPELAEA (GOLDFUSS, 1810) AND
IMPLICATIONS ON POCALA CAVE (TRIESTE, ITALY) PALAEOECOLOGY
1Department of Cultural Heritage, University of Bologna, Ravenna Campus, Via Degli Ariani 1,
48121, Ravenna, Italy.
2Centro di Ateneo per i Musei (CAM), University of Padova, Via Giotto 1, 35121, Padova, Italy.
In Italy, cave lions (Panthera spelaea) are well represented by several fossil specimens but
research mainly focused on their anatomy and taxonomy. In this study, lion remains from Pocala
Cave (Trieste, Italy) preserved in the Museum of Geology and Paleontology of the University of
Padova were redescribed. Morphometric analysis confirmed that these lions belong to the chrono-
subspecies Panthera spelaea spelaea, most individuals are interpreted as adult males, and
several young individuals are present. Taphonomic analysis shows low intensity water flow, few
signs of scavenging, and cut-marks on a lioness jaw, probably of anthropogenic origin. Analysis of
87Sr/86Sr isotopes indicates that the same individual was not native of the Pocala Cave area.
Those results confirm that exploitation of large carnivores was not unusual among Pleistocene
humans, and provide an insight into the complex palaeoecology of Pocala Cave. The cave was
used as a shelter by Pleistocene humans for collecting resources, and rarely by scavengers for
consuming their preys. Furthermore, lions entered the cave to hunt cave bears during their
hibernation, and Pocala Cave was probably used by lionesses as a refuge to give birth and take
care of cubs, as suggested by the numerous remains of young individuals at different growth
stages, providing the first possible evidence of a lion “den”.
122
DINOSAUR TRACKS, SEDIMENTARY ENVIRONMENTS AND
PALAEOCLIMATE OF THE LATE CRETACEOUS EL MOLINO FORMATION IN
BOLIVIA (MAASTRICHTIAN, BOLIVIA; TORO TORO NATIONAL PARK)
3Universidade Federal do Rio de Janeiro, 21.910-200 Av. Athos da Silveira Ramos 273, Instituto
de Geociencias. CCMN, Rio de Janeiro, RJ, Brazil.
4Dipartimento di Scienze della Terra, Università degli Studi di Firenze, Via La Pira, 4, I 50121
Firenze, Italy.
6Dinosaur Trackers Research Group, University of Colorado Denver, PO Box 173364, Denver, CO
80217-3364, United States of America.
7Faculdad de Ciencias, Escuela de Geología, Universidad Mayor, Manuel Montt 367, Providencia,
Santiago de Chile, Chile.
9Facultad de Ciencias Geologicas, Carrera de Ingeneria, Universidad Mayor de San Andrés L Paz,
Campus Universitario Cota Cota, Calle 27, Bolivia.
The El Molino Formation in the Toro Toro syncline (Department of Potosi, Bolivia) has long been
known for its Cretaceous dinosaur tracks. The description of a trackway, Ligabueichnium
bolivianum, attributed either to a ceratopsian or to an ankylosaur comes from the Toro Toro area.
123
The type trackway consists of undertracks with no anatomical details that can be seen under
closer inspection. Five larger surfaces with trackways of theropods and sauropods have been
figured in previous papers which are all situated close to the village center of Toro Toro. Recently,
we assign the sauropod trackways to the ichnotaxon Calorckosauripus lazari, attributed to a non-
derived titanosaur. Several sites, documented in 2019, are situated in the middle member of the El
Molino Formation (Maastrichtian), and 15 different track levels have been recorded. The tracks
occur as negative epichnia on fine-grained sandstones and limestones. In some sites, such as Las
Golondrinas, positive hypichnia of theropods are present and sauropod footprints can be seen in
cross-section. Today, more than 28 track-bearing areas are known inside and close to the National
park Toro Toro, recording the presence of titanosaurid sauropods, different types of theropods,
and ankylosaurs. The Toro Toro area is of key interest as it records dinosaur track levels that are
close to the K/Pg boundary. The project’s main focus is a multidisciplinary attempt to integrate
stratigraphy, climate proxies (e.g., clay minerals, isotopes), sedimentary environments and
ichnoassemblages in the context of the Latest Cretaceous in order to verify or falsify the rapid
decline of dinosaur diversity.
124
NOTOCHORD PIT SIZE AS A POTENTIAL PROXY FOR EMBRYONIC STAGE
IN ICHTHYOSAURS
Recently four different embryonic developmental stages have been proposed in the parvipelvian
ichthyosaur Stenopterygius based on relative cranial ossification. Stenopterygius has the largest
prenatal record of any ichthyosaur by far, including cranial and postcranial material in many
embryos of gravid females. Other ichthyosaurs have a much more limited prenatal record, often
lacking cranial material. This means that determining the prenatal stage in these embryos (relative
to Stenopterygius) is difficult. The most common prenatal elements to preserve are vertebral
centra. In Stenopterygius we noticed that the notochord pit is relatively larger in size in early
development and smaller perinatally. Per embryonic stage preserved in Stenopterygius, we
determined the relative notochord pit size of the vertebral centra. The relative notochord pit size
ranged from ~ 50% in stage 1 to 40% in stage 2, 25% in stage 3, and 15% in stage 4. At every
stage there were clear differences between the more anterior and more posterior vertebrae.
However, preflexural vertebrae in every stage had roughly the same relative notochord pit size.
We then examined gravid specimens of Mixosaurus, Ichthyosaurus, and Leptonectes with both
cranial material and vertebral centra preserved. Comparable stages relative to Stenopterygius
were hypothesized on the basis of cranial ossification. In all cases, the relative notochord pit size
was similar to the size of hypothesized stage based on cranial ossification. This means that
relative notochord pit size can be used as a proxy for embryonic stage in ichthyosaurs in the
absence of cranial material.
125
THE MIDDLE EOCENE VERTEBRATES FROM PONTILS (EBRO BASIN, NE
SPAIN)
5Departamento de Física e Ciencias da Terra, Universidade da Coruña, Rúa da Fraga 10, 15008
A Coruña, Spain.
6School of Earth Sciences, University of Bristol, 24 Tyndall Avenue, BS8 1TQ Bristol, UNITED
KINGDOM.
The Pontils locality (Ebro Basin, NE Spain) corresponds to a continental-to-marine sequence with
several vertebrate-bearing levels. Previous works in this site documented the presence of scarce
vertebrate remains, but only rodents were described in detail. Recent fieldwork carried out in this
site has allowed recognizing seven levels yielding significant vertebrate remains. Among
mammals, primates are particularly diverse and include abundant remains of a small, still
undetermined omomyiform and scarce teeth of Pseudoloris, Necrolemur and an undetermined
anchomomyin. Rodents are present in all the sampled levels and comprise additional material of
126
some species already described from this section (Pseudoltinomys cosetanus, Sciuroides cf.
romani, Paradelomys sp.) and other forms unknown from this site until now (Pseudoltinomys sp.,
Pairomys cf. ibericus). Metatherian and eulipotyphlans are identified in several levels, although the
material is scarce and fragmentary; similarly, chiropterans are only represented by two teeth from
levels PO-22 and 33. Perissodactyls are diverse in PO-39 and include cf. Palaeotherium sp., cf.
Anchilophus (Paranchilophus) sp., cf. Anchilophus (Anchilophus) sp. and “Pachynolophinae”
indet., whereas artiodactyl remains have been found in PO-39 (Dacrytheriinae indet.) and PO-40
(Hyperdichobuninae indet.). Regarding the herpetofauna, amphibians are scarce, squamates are
relatively diverse (including gekkotans, lacertids, iguanids, anguids and snakes), and
crocodyliforms are represented by very abundant isolated teeth in nearly all levels. The
ichthyofauna found in the upper levels (PO-39 and PO-40) includes chondrichthyans (three sharks
and three rays) and actinopterygians. Summarizing, the assemblage from Pontils represents one
of the most diverse Bartonian vertebrate faunas from the Iberian Peninsula.
127
ON THE AGE OF THE PONTILS FOSSIL SITE, AN EXCEPTIONAL CASE OF
CORRELATION BETWEEN MARINE AND CONTINENTAL SCALES
3Natural History Museum of Denmark, University of Copenhagen, Øster Voldgade 5-7, 1350
Copenhagen K, Denmark.
The fossil site of Pontils (Ebro Basin, NE Spain) is placed in a sequence that records the change
from continental to marine conditions, thus representing an excellent opportunity for the correlation
of biostratigraphic schemes based on marine and continental fossils. This locality has been known
since the 1980s and was traditionally assigned to the Bartonian, on the basis of scarce mammal
remains recovered from the lowest levels of the sections and, particularly, of marine fossils
(foraminifera and nannoplankton) found in the overlaying marine sediments. However, some
recent works have proposed an older, Lutetian age, for this site. Two field campaigns carried out
by our team in the Pontils section have led to the recovery of abundant vertebrate remains from
seven different levels, including chondrichthyans, actinopterygians, amphibians, crocodilians,
squamates, metatherians, eulipotyphlans, chiropterans, rodents, artiodactyls, perissodactyls and
primates, together with other non-vertebrate fossils. Among the latter, larger benthic foraminifera
are particularly interesting, since the assemblage of Nummulites garnieri sturi, N. garganicus, and
128
Assilina (Operculina) schwageri allows assigning the site to the zone SBZ17 (Bartonian). Previous
analyses of paleomagnetism determined that this site corresponds to a reversed-polarity interval,
and therefore it can be confidently correlated to chron C18r (40-41 Ma). The rodent assemblage,
including Pseudoltinomys cosetanus, Pseudoltinomys sp., Sciuroides cf. romani, Paradelomys sp.
and Pairomys cf. ibericus, also supports a Bartonian age for this site. Therefore, our results solve
the debate about the age of the locality, which can be definitely assigned to the Bartonian.
129
HINTS OF INCREASED LONGEVITY IN EXTINCT INSULAR DWARF DEER
J.J. Miszkiewicz1,2, A.A.E. van der Geer3*
1School of Archaeology and Anthropology, Australian National University, Canberra, Australian
Capital Territory, Australia.
2School of Social Science, University of Queensland, Brisbane, Queensland, Australia.
3Vertebrate Evolution, Development and Ecology, Naturalis Biodiversity Center, Leiden, the
Netherlands.
Dwarfed insular mammals evolve a suite of morphological and physiological adaptations triggered
by a range of variables associated with geographical isolation. One of such variables is longevity
which can extend in relation to body size reduction in ecological contexts of predator and
competitor release, or reduced resource availability. Longevity in extreme cases of dwarfism can
be reconstructed using palaeohistological approaches. Here, we studied the histology of seven rib
fragments of the extraordinary, endemic dwarf deer Candiacervus (size 2) of Crete (Greece),
dated to the Pleistocene. We tested whether its rib bone matrix shows evidence for extensive bone
remodelling typically seen in larger and long-living mammals. We found that all ribs had well
remodelled Haversian bone, which ranged from moderate to strong degrees of remodelling.
Widespread Haversian bone with superimposed and remodelled secondary osteons implies
experiences of several cortical bone resorption and re-deposition events. We also found some of
the smaller rib cross-sections showed relatively higher osteon densities, along with instances of
cortical bone porosity similar to that expected in elderly animals. These rib histology results agree
with prior literature discussing longevity, and bone histology, in dwarf ruminants, and suggest this
dwarf deer might have lived longer than what would be expected for its size. We highlight the
usefulness of ribs when studying histology in palaeontological specimens.
130
LO HUECO QUARRY (UPPER CRETACEOUS. CUENCA, CENTRAL SPAIN), A
PUZZLE OF ARTICULATED TITANOSAURIAN TAILS
1Instituto Dom Luiz, Faculdade de Ciência da Universidade de Lisboa, Campo Grande, 1749-016
Lisbon, Portugal.
3Dinosaur Institute, Natural History Museum of Los Angeles, 900 Exposition Blvd, 90007 Los
Angeles, California, United States of America.
4Facultad de Bellas Artes, Universidad Complutense de Madrid, Calle Pintor el Greco 2, 28040,
Madrid, Spain.
6Department of Organismal Biology, University of Chicago, 1027 E 57th St, Chicago, Illinois
60637, United States of America.
Titanosaurian record from the Upper Cretaceous of Europe is abundant. This record is mainly
represented by bones recovered from localities in which the degree of skeletal articulation or
association is low, being difficult to establish the relationship between bones and where more than
one titanosaur may be represented. Recently, partially articulated titanosaurian specimens have
been found in the Ibero-Armorican domain, highlighting the fossil-site of Lo Hueco (Cuenca,
Spain). Lo Hueco is a multitaxic bonebed with more than 10,000 collected fossils of which nearly
half are titanosaurian remains. These specimens are key to understand the evolutionary history of
European titanosaurs and to study their intraspecific variability and paleobiology, using a set of
new methodologies. More than 10 individuals with fully to partially articulated or associated
elements were found in this fossil-site, including several partial individuals preserving series of
articulated caudal vertebrae. Analyses on these specimens are particularly important to
understand positional, intra-, and interspecific morphological variability of caudal vertebrae, which
can shed light to the systematics of European titanosaurs. Caudal vertebrae are source of several
131
morphological characters in datasets, and particularly diagnostic for titanosaurian subclades.
Accessing positional variability can be relevant in the way we codify and score these
morphological characters. To test this assumption, we provide a discussion on several caudal
vertebrae anatomical characters such as the orientation of the neural spine, the ventral and
longitudinal hollow of the centrum, the posterior condyle morphology, or the presence of
postzygodiapophyseal lamina.
132
CALCANEAL MORPHOLOGY IS A RELIABLE PROXY FOR THE
LOCOMOTOR BEHAVIOUR OF EXTINCT PRIMATES
3Institute of Ecology and Evolution, University of Bern, Baltzerstrasse 6, 3012, Bern, Switzerland.
4Life Sciences Department, Vertebrates Division, Natural History Museum, SW7 5BD, London,
United Kingdom.
The calcaneus plays a critical role in efficient foot movement required to navigate differing
substrate conditions such as of the arboreal milieu. Therefore, we quantitatively studied the
functional morphology of this bone in a large sample of extant primates using a novel 3D
geometric morphometrics approach that combines anatomical landmarks along with curve and
surface semilandmarks. To do so, we assessed the entire morphology of the bone as well as its
different facets independently, such as the subtalar and transverse tarsal joints since they
constitute functionally relevant regions within the foot. Allometric effects, phylogenetic signal and
covariation between calcaneal shape and percentages of locomotor behaviour compiled from
field studies were examined. Finally, representatives of the main Paleogene primate groups
(plesiadapiforms, adapoids, omomyoids and stem anthropoids) were included in an exploratory
analysis to assess their locomotor adaptations. Our results indicate significant allometry in the
distal calcaneal region, with functionally informative departures from the allometric regression.
Phylogenetic signal was found to be moderately low, thus hinting at an important functional
signal. Calcaneal shape is strongly correlated with locomotor behaviour, either when the entire
shape of the bone or its different facets are examined. This highlights the potential of this bone to
extract reliable information regarding the locomotor behaviour of taxa for which only fragmentary
remains are available. Lastly, the analysis including extinct species indicates that early primates
displayed a wide range of calcaneal morphologies, suggesting that they engaged in a diverse
array of locomotor behaviours early on in their evolutionary history.
133
PRIMARY OSTEOARTHRITIS RECORDS IN THE EXTINCT LAGOMORPH
PROLAGUS SARDUS AND ITS LIFE HISTORY IMPLICATIONS
1Departamento de Física e Ciencias da Terra, Universidade da Coruña, Rúa da Fraga 10, 15008
A Coruña, Spain.
2Dipartimento di Scienze, Università degli Studi Roma Tre, Largo S. Murialdo 1, 00146 Roma,
Italy.
3School of Veterinary Medicine, University of California, 1 Garrod Dr, 95616 Davis CA, United
States of America.
4Institut Català de Paleontologia Miquel Crusafont (ICP), Carrer de les Columnes s/n, 08193
Cerdanyola del Vallès, Spain.
5Key Laboratory of Vertebrate Evolution and Human Origins, Institute of Vertebrate Paleontology
and Paleoanthropology, Chinese Academy of Sciences, 142 Xi Zhi Men Wai Da Jie, 100044
Beijing, China.
The study of diseases, both human and nonhuman, in antiquity may discern novel biological details
about extinct taxa. Current work in animal paleopathology has been limited to descriptions of
exceptional abnormalities without quantitative reports at population level. In the present study, we
examined 246 femora of Prolagus sardus (Ochotonidae, Lagomorpha) for evidence of primary
osteoarthritis (pOA). The source of the specimens was from the renowned Dragonara cave, located in
the north-western Sardinia, and dated back to the Late Pleistocene (22.390-21.910 ka cal BP). For
each specimen, we established the age category and reconstructed the body mass, as well as
evaluated hip and knee joints for pOA using high resolution radiographic and histological modalities.
The prevalence of pOA in P. Sardus from Dragonara was 24.9%, particularly affecting hip joint, and its
occurrence was associated with older age categories, but not with larger sizes. Morphologic features of
thickened subchondral bones, irregular bone contours, partial collapse of the femoral neck and
evidence of irregular bone composition, suggestive of adoptive modeling, were recognized in pOA-
affected specimens. In femoral condyles, denser subchondral bone was suggestive of sclerosis.
Prolagus sardus exhibited a pOA prevalence similar to that of domestic present-day rabbits, even
134
though it was a wild species and subjected to natural selection. A longer lifespan may be the main
evolutionary trigger of the observed results, although other causes cannot be excluded. This study is
framed within a comprehensive investigation which aims to unraveling the eco-evolutionary adaptations
of extinct lagomorphs to insular regimes.
135
A NEW METHOD TO MODEL RARE SPECIES BY RELYING ON PHYLOGENY
2Department of Biosciences and Territory, University of Molise, c.da Fonte Lappone, 86090
Pesche, Italy
3DiSTAR, Università degli Studi di Napoli Federico II, Via Cinthia 21, 80126, Naples, Italy
Species distribution models (SDMs) are powerful tools with which to understand how
environmental variation influences species geographical distribution. Although SDMs are widely
used in ecology and applied conservation biology, they consistently lose accuracy precisely when
they are needed the most, that is with rare species, originating the so-called rare-species modeling
paradox. We overcome this issue presenting a new algorithm, ENphylo, developed for the R
environment, which couples Environmental Niche Factor Analysis (ENFA) and phylogenetic
imputation to model rare species. By using the fossil record of two large mammal species
(Rangifer tarandus, a cold-adapted species, and Sus scrofa, a warm-adapted species) that lived
during the late Pleistocene as the source data to sample from, we demonstrate ENphylo provides
high predictive accuracy under different model evaluation metrics, when just 10 fossil occurrences
are randomly drawn from their respective fossil records. ENphylo significantly outperforms ENFA
and provides a fast and accurate solution to perform rare species distribution modeling, which will
help predict their spatial distribution in the light of climate change, and to delineate how extinct
species reacted to past climatic variation.
136
THE EARLY- MIDDLE PLEISTOCENE SORICIDAE (MAMMALIA) OF THE
IBERIAN PENINSULA
4Archaeological Studies Program, University of the Philippines-Diliman, Albert Hall, Lakandula St.,
Quezon City 1101, Philippines.
6National Museum of the Philippines, P. Burgos Drive, City of Manila 1000, Philippines.
Soricids are small mammals that commonly appear in Pleistocene sites of the Iberian Peninsula.
The soricids with a post-subchron-Jaramillo Early-Middle Pleistocene age currently cited in the
Iberian Peninsula are: Sorex minutus, Sorex gr. araneus-coronatus, Dolinasorex glyphodon,
Neomys cf. fodiens and Crocidura russula. Most of the records are indeterminate species due to
the scant and fragmentary nature of the fossils. However, in other parts of Europe, there are up to
ten different species of soricids in the same level.
The archaeo-paleontological locality of Gran Dolina (Burgos, Spain) is a cave infill dated to the
post-Jaramillo Early-Middle Pleistocene where D. glyphodon was first discovered. We present a
review and taxonomic attribution of the remaining soricid species from the whole stratigraphic
sequence of Gran Dolina. Two-hundred fragmentary mandibles from levels TD4 to TD10
(recovered with washing-sieving methods during the 1991-2017 field campaigns), were
photographed, described, measured and compared with data from other European localities.
Nine taxa were identified in the Gran Dolina stratigraphic sequence: S. minutus, Sorex gr.
runtonensis-subaraneus, Sorex (Drepanasorex) gr. margaritodon-savini, D. glyphodon, Asoriculus
137
gibberodon, Neomys cf. newtoni in the Early Pleistocene levels, and S. minutus, S. gr.
runtonensis-subaraneus, Neomys cf. fodiens, Neomyni cf. Macroneomys and C. kornfeldi in the
Middle Pleistocene levels. The biometric data shows extreme size values for the Gran Dolina taxa
when compared to their counterparts in other regions of Europe, increasing the variability of these
species.
Acknowledgements: Fellowships FPU14/ 05528(MECD) and Margarita Salas (MU and the
European Union-NextGenerationEU). Projects CGL2015-65387-C3-2-P(MINECO/FEDER),
PGC2018-093925-B-C33(MCIU/AEI/FEDER, UE) and RTI2018‐093419‐B‐I00(AEI/FEDER, UE).
138
BITE TRACES, A POWERFUL TOOL IN PALAEOECOLOGY: THE CASE OF
THE MIDDLE TRIASSIC LOWER KEUPER FOSSIL LAGERSTÄTTEN
4Palaeontology, Institut für Biologie, Universität Hohenheim, Wollgrasweg 23, 70599, Stuttgart,
Germany.
Trace fossils, or ichnofossils, are important elements to understand the relationships and
interactions of trace makers with the environment and/or with other organisms. In this regard, bite
traces (commonly also known as “bite marks”) provide direct evidence on the feeding habits and
potentially the ecological role of the bite maker, even in the cases in which it is not confidently
identified. Bite traces are especially relevant for palaeoecological studies when coupled with other
analyses, such as on taxonomy and taphonomy (including sedimentology). Here, we present a
study on the bite traces present on tetrapod bones from the Lower Keuper fossil lagerstätten
(Middle Triassic, S Germany). We identified nearly 200 bones and teeth with at least one bite trace
(usually being more). We created a workflow for the documentation of bite traces, which includes:
(1) morphological descriptions (ichnotaxonomy), (2) associations between bite trace morphotype,
(3) location on the bones, (4) orientation of the traces on the bones. Such database allowed
performing statistical analyses, creating “biting spaces” according to the bitten taxon, bone type
and bone region. Derived interpretations on feeding behaviour were contrasted and confirmed with
analyses on the tooth macroscopic wear and microanatomy of the bite maker, which was
confidently identified coupling tooth and bite trace morphologies. In the Lower Keuper ecosystems,
the pseudosuchian archosaur Batrachotomus was the top predator, showing scavenging,
predatory and cannibalistic behaviours and mostly feeding on the giant temnospondyl
139
Mastodonsaurus. Comparisons with the bite trace record suggest a conserved feeding ecology
among pseudosuchians, including present-day crocodilians.
140
PALAEOENVIRONMENTS AND TAPHONOMY OF THE KUPFERZELL
VERTEBRATE LAGERSTÄTTE (MIDDLE TRIASSIC, S GERMANY)
3Palaeontology, Institut für Biologie, Universität Hohenheim, Wollgrasweg 23, 70599, Stuttgart,
Germany.
The Middle Triassic epoch witnessed a flourishing of ecosystems, with the radiation of several
vertebrate lineages, eventually dominated by archosaurs. Fossil sites in terrestrial settings from
this time interval, even if known from all continents, still hold unresolved questions regarding the
evolution of faunas and their significance in the recovery during the aftermath of the end-Permian
mass extinction and the evolution of some tetrapod groups. In order to fill this void of knowledge,
we are carrying out studies on the sedimentology and taphonomy of the vertebrate fossil
lagerstätten from the Lower Keuper (Ladinian, S Germany). In the present work, we focus on the
singular Kupferzell locality, which delivered ~30,000 vertebrate remains in a 3-month-long salvage
excavation in 1977 during road construction. We reconstructed the stratigraphic succession of the
main excavation site with the aim of unravelling the palaeoenvironmental evolution of relatively
complex ecosystems, which included different top predators. The bulk of the fossil material
corresponds to two temnospondyl amphibians (Gerrothorax, ~70%; Mastodonsaurus, ~30%), with
the archosaur Batrachotomus, ranging first among the other tetrapod remains, and a relatively high
diversity of fishes. The sequence consists of: basal coaly mudstones, massive siltstones, green
siliciclastic mudstones, yellow-brown carbonate claystones-marlstones, and massive dolostones.
They represent the succession of relatively similar water bodies, some with characean meadows
and ostracodes, from a lacustrine setting with recurrent marine influence. Drought periods marked
the remodeling of environments and development of different faunal assemblages. The complexity
141
of vertebrate communities suggests recovery of ecosystems, with faunas adapted to changing
environments.
142
A RE-EVALUATION OF THE NOTOSUCHIAN CROCODYLIFORM EREMOSUCHUS
ELKOHOLICUS FROM THE EARLY EOCENE OF NORTH AFRICA AND THE
PHYLOGENETIC RELATIONSHIPS OF SEBECOSUCHIANS
1Department of Earth Sciences, University College London, Gower Street, WC1E 6BT, London,
United Kingdom.
Notosuchian systematics have been highly debated in the past few decades, particularly regarding
the placement of sebecid sebecosuchians, the only putative notosuchian taxa to have survived the
Cretaceous/Paleogene mass extinction event, 66 Ma. Reconciliation of any conflicting views on
notosuchian relationships through standardized approaches to character construction and
improved taxon and character sampling is therefore required. Here, we redescribe the type
material of Eremosuchus elkoholicus, a largely neglected species from the early Eocene of
Algeria, as well as undescribed remains from the same locality that potentially represent a juvenile
of this species. Both specimens are incorporated into one of the largest notosuchian-focused
character-taxon matrices yet to be compiled, comprising 453 characters and 124 taxa. Particular
focus is placed on increasing the sampling of sebecosuchians, especially the frequently neglected
fragmentary remains from Europe and North Africa. Continuous characters relating to the mandible
are implemented for the first time in a notosuchian-relevant dataset, increasing the sampling of
mandibular characters by ~20%. All discrete mandibular characters are reassessed to improve
consistency of scoring. Analyses are run under multiple weighting schemes including equal
weighting and extended implied weighting. In all treatments, Eremosuchus is recovered within
Sebecidae, as the sister taxon to Lorosuchus nodosus, from the Paleocene of Argentina. Given
the sparsity of sebecosuchian remains in Northern Africa during this time interval, this taxon
provides an important datapoint in elucidating the dispersal and radiation of the clade, which is
particularly pertinent considering the ongoing fragmentation of Gondwana during the Late
Cretaceous and Paleogene.
143
PHYLOGENETICAL ISSUES WITHIN THE SUPERFAMILY
CTENODACTYLOIDEA
2Association Mujeres con los Pies en la Tierra, C/ Jose Antonio Novais, 12, 28040 Madrid, Spain.
4Sedimentary Geology and Environmental Change, Geosciences Institute (CSIC, UCM), C/ del Dr.
Severo Ochoa, 7, 28040, Madrid, Spain.
During the last decades, there have been different phylogenetic approaches regarding the
Ctenodactyloidea superfamily. Several molecular phylogenies pointed out the paraphyly of this
group and propose the “Ctenohystrica” clade. Several morphological phylogenies have proposed
different cladistic positions for the different genera included in this superfamily.
For the first time, we proposed the most complete morphological phylogeny referring to this
superfamily, trying to clarify the position and relationships between and within the different genera.
To do that, we performed a parsimony phylogenetic analysis with more than 40 species and 63
characters related to the dental and cranial remains. These remains comprise a sample that
comes from five geographical regions (East Asia, Central Asia, Western Asia, Europe and Africa)
and recorded from the Eocene to the Holocene.
Our results show ctenodactylids are recorded for the first time in East Asia during the Eocene and
since that date, the species flourished and spread out through Western Asia, Europe and Africa
and became one of the dominant rodents groups of the Oligocene. In relation to the morphological
trends of this group, these species show an increase in the hypsodont pattern coinciding with a
climatic change during the Oligocene-Miocene transition. Concerning the phylogenetic position of
the species, we have clarified the relation between the European and African ctenodactylids and
the relation between the extant ctenodactylids and extinct distylomyids.
144
ON THE DIAGNOSIS OF IBEROSUCHUS MACRODON
(CROCODYLIFORMES, NOTOSUCHIA) FROM THE EOCENE OF PORTUGAL
2Instituto Dom Luiz, Faculdade de Ciências da Universidade de Lisboa, Campo Grande, 1749-016
Lisboa, Portugal.
4Área de Zoología, Universidad Nacional de San Luis. Chacabuco 917, 5700 San Luis Argentina.
Laurasian notosuchians are a singularity within the record of crocodyliforms and, particularly,
medium-large notosuchian ziphodonts outside Gondwana are restricted to the Eocene of the
Iberian Peninsula and Southern France. The best-known set of these notosuchians is composed
by remains from the Eocene of Portugal, Spain and France that are usually related to the
Portuguese form Iberosuchus macrodon, although they are not synchronous and have a relatively
wide distribution in time (Lutetian to Bartonian)
Since the holotype of Iberosuchus macrodon is restricted to the anterior portion of a rostrum from
the Eocene of Vale Furado (middle Eocene, Lutetian. Portugal) and no other material is known
from the Portuguese record, the assignment of other specimens from Spain and Southern France,
and thus the characterization of this crocodile from attributed material, should be taken with
caution.
In fact, in the holotype of Iberosuchus macrodon only a series of characters that are common to
many notosuchians (and particularly to the baurusuchids) are identified. The only recognized
automorphic character on the rostrum of specimens attributed to Iberosuchus outside Portugal is a
large premaxillo-maxillary palatal fenestra associated with a complex palatal fossa. This character
could not be confirmed directly in the holotype of Iberosuchus because most of this structure is
internal or hidden by a plaster filling placed during preparation.
145
A CT-Scan of the Iberosuchus holotype has been carried out to describe in detail its rostral
anatomy, both to evaluate its diagnostic capacity, and to compare it with those of specimens from
the Spanish record.
146
NEW MATERIAL OF THEROPODS (ABELISAUROIDEA?) FROM LO HUECO
(LATE CRETACEOUS. CUENCA, CENTRAL SPAIN)
2Instituto Dom Luiz, Faculdade de Ciências da Universidade de Lisboa, Campo Grande, 1749-016
Lisboa, Portugal.
3Museo Carmen Funes, Av. Córdoba 55 (8318), Plaza Huincul, Neuquén, Argentina.
To date, theropod diversity from the Late Cretaceous of the Iberian Peninsula is still far from being
well established, mainly because these dinosaurs are represented by very isolated remains. Up to
now, the known diversity is composed of several taxa attributed to different groups of coelurosaurs
and by medium-sized primitive forms whose characters are congruent with abelisauroid
ceratosaurs. The Lo Hueco fossil-site (Campanian-Maastrichtian of Cuenca, central Spain) has
provided an abundant record of titanosaur sauropods, crocodyliforms and testudines, in which the
presence of ornithischian and theropod dinosaurs is scarcer. However, among the latter, teeth,
and postcranial material of coelurosaurs are common in the fossil-site, but remains of non-
coelurosaurian forms are rarer and so far, only a few teeth and one metatarsal have been
previously described.
A discussion about a theropod posterior caudal vertebra from Lo Hueco is proposed. This vertebra
presents some singular characters, some of them shared with members of Abelisauroidea.
Centrum is amphicoelous and does not present transverse processes nor pleurocoels. The neural
spine is extremely thin and, in lateral view, is relatively tall, triangular, and caudally positioned.
Prezygapophyses are relatively short, postzygapophyses are very reduced and a shallow
longitudinal groove is present along the ventral face of the centrum. This combination of features is
compatible with those of some basal abelisauroids, providing new information for the discussion on
the relationships of the non-coelurosaurian theropods from the Late Cretaceous of the south of the
Ibero-Armorican domain.
147
LIGHTS AND SHADOWS IN THE EVOLUTIONARY DYNAMICS OF THE
PLEISTOCENE SARDINIAN MAMMOTH POPULATIONS
1CNR-IGAG, Dept. of Earth Sciences, Sapienza University of Rome, P.le Aldo Moro 5, 00185
Roma, Italy.
2Dept. of Veterinary Medicine, University of Sassari, via Vienna 2, 07100 Sassari, Italy.
The fossil record of the Sardinian proboscideans mainly consists of isolated remains found in sites
ranging in age from the late Middle to the Late Pleistocene(an incomplete skeleton from Guardia
Pisano, isolated remains from different localities, and some footprints from Funtana Morimenta)
that have been ascribed to a single species of endemic, moderately reduced-in-size mammoth,
Mammuthus lamarmorai. The paucity of remains, the different sizes of molariform teeth from
different localities, the lack of dental remains at Guardia Pisano and the uncertainties about the
chronology of some remains hamper any attempt to infer whether one or more species, originated
by an anagenetic evolutionary process or by multiple arrivals from the mainland, inhabited the
island. Therefore, the continental ancestor/ancestors of Sardinian mammoth populations, the time
and number of dispersals of the ancestral taxon/taxa, the actual number of mammoth species that
inhabited Sardinia, as well as the persistence through time of these populations still remain
unsolved issues, due to the low number of findings and the imperfect chronological constraint of
some remains.
This research aims to put out and discuss the main issues related to the actual number of
mammoth species that inhabited Sardinia, the time and number of dispersals of the ancestral
taxon/taxa, the morphological and dimensional differences shown by tooth remains, the dynamics
of Sardinian mammoth populations and their persistence through time, and the possible causes of
the populations' decline and disappearance, as well as preliminary data on the histological
characteristics of a Sardinian mammoth long bone.
148
THE ENDEMIC GIRAFFID FROM THE OREOPITHECUS-FAUNAL
ASSEMBLAGES OF THE TUSCO-SARDINIAN ARCHIPELAGO (LATE
MIOCENE, ITALY)
L. Pandolfi1*, L. Rook1
1Dipartimento di Scienze della Terra, Paleo[Fab]Lab, Università degli Studi di Firenze, Via G. La
Pira 4, 50121 Firenze, Italy.
Over the past decades, numerous studies have been aiming to reconstruct the evolution of
endemic mammals from the Late Neogene of the Tyrrhenian area in the Western Mediterranean
Basin. Despite these efforts, however, no conclusive evidence has been provided yet to
confidently address the debate about the origin of some of the species characterising the
Oreopithecus-faunal assemblages of the Tusco-Sardinian archipelago. An emblematic case is the
enigmatic giraffid-like Umbrotherium azzarolii. The occurrence of this mammal has been so far
documented by the single type specimen from Casteani (V1 faunal assemblage) in Tuscany, and
by the sample from the slightly younger locality of Fiume Santo (V2 faunal assemblage) in
Sardinia. Despite the presence of teeth, an exhaustive diagnosis of Umbrotherium was not firmly
established and its phylogenetic position remained unresolved. Albeit no new specimens, nor new
localities yielding these taxa have been recently discovered, recent archival research at the
Natural History Museum of Basel and the Department of Earth Sciences of the University of
Florence (under the PalAss grant n. PA-SB202103) enabled to re-discover unpublished remains of
giraffids, including an almost complete mandible, several isolated teeth and other cranial remains
from the localities of Botro della Canonica and Serrazzano (Pisa). The new material shed lights on
the morphological and morphometric variability of Umbrotherium, enabling a comparison between
specimens collected from different Tusco-Sardinian Miocene localities spanning from V1 to V2
Oreopithecus-faunal zone. Phylogenetic affinities are investigated to detect a possible ancestor of
this endemic taxon and reconstruct its dispersal route.
149
MORPHOLOGICAL CONVERGENCE IN THE HINDLIMB SKELETON WITHIN
TITANOSAURIA
3Instituto Dom Luiz, Faculdade de Ciência da Universidade de Lisboa, Campo Grande, 1749-016
Lisbon, Portugal.
4Dinosaur Institute, Natural History Museum of Los Angeles, 900 Exposition Blvd, 90007 Los
Angeles, California, USA.
This work is being funded by an ERG-2020 EAVP research grant, PID2019-111488RB-I00 and
CEECIND/00726/2017.
150
AN ICHTHYOSAUR BREEDING GROUND IN SOUTHERNMOST GONDWANA
(TYNDALL GLACIER, SOUTHERN CHILE)
3Fundación de Historia Natural Félix de Azara, Universidad Maimónides, Hidalgo 775, C1405,
Buenos Aires, Argentina.
4Museo de Historia Natural de Río Seco, Juan Williams 012812, 62000000, Río Seco, Punta
Arenas, Chile.
5Escuela de Geología, Universidad Católica de Temuco, Manuel Montt 56, 4780000, Temuco,
Chile.
Since 2004, paleontological expeditions have been conducted to the Tyndall Glacier in Torres del
Paine National Park, Chilean Patagonia, resulting in the discovery of almost a hundred ichthyosaur
skeletons to date. The ichthyosaurs are exposed in the rocks as a consequence of the ongoing
melting of the glacier caused by climate change. Most of the ichthyosaurs in this locality are
complete and articulated. Invertebrates such as ammonites and belemnites, as well as different
types of fishes, have been found associated with the ichthyosaurs. According to a previously
developed hypothesis, this area was used as a breeding ground for ichthyosaurs in southernmost
Gondwana during the Early Cretaceous. To date, eight years since the hypothesis was published,
we have yet to find marine reptiles other than ichthyosaurs, most finds corresponding to neonates
and small juveniles. Pregnant females have also been recorded. These data, in addition to the
abundance of food and the apparent absence of predators, allow us to support the idea of a
breeding ground. The discovery of glendonites, correlated with cold to freezing water conditions,
suggest that the site may have only been inhabited during summer months. Regarding taphonomy,
151
many of the skeletons have skulls that penetrate the sedimentary layers, indicating headfirst arrival
at the seafloor. Several skulls show breakage consistent with high speed collision with the bottom.
The frequency and depth of embedded skulls suggest relatively soft sediments at the sea floor.
This research was funded by Agencia Nacional de Investigación y Desarrollo (ANID), project
PAI77200036 awarded to JPP.
152
LATEST EARLY PLEISTOCENE DAMA-LIKE CERVID REMAINS FROM CAL
GUARDIOLA AND THE TANGLE OF VILLAFRANCHIAN CERVID TAXONOMY
3Dipartimento di Fisica e Geologia, Università degli Studi di Perugia, Via A. Pascoli, 06123
Perugia, Italy.
The taxonomy and phylogeny of the small to mid-sized Plio-Pleistocene cervids of Europe,
commonly known as Dama-like deer, have been a matter of vast dispute, as specimens referred to
this group have been attributed to multiple genera such as: Dama, Pseudodama, Axis, Euraxis,
Rusa and Metacervocerus. Here we present a preliminary study of 305 cervid remains from the
site of Cal Guardiola (CGR), consisting of 157 craniodental and 148 postcranial elements. The site
of CGR, along with the nearby site of Vallparadís Estació (EVT), are correlated along the
Vallparadís Section, cropping out in the western bank of the Torrent de Vallparadís (Terrassa,
Vallès Penedès Basin, Iberian Península). The CGR section consists of two units separated by an
erosional surface, and it hosts seven fossiliferous layers spanning in age from 1.2 to 0.86 Ma. Both
quantitative and qualitative morphological comparisons of the studied cervid material from CGR
allow us to attribute it to the Plio-Pleistocene Dama-like group of deer. The antler morphology from
the CGR cervid, in particular, resembles those from Capena, Pietrafitta, Redicicoli and Collecurti
(Italy), Le Vallonnet (France) and Untermassfeld (Germany) which have been attributed to a
variety of taxa that we can group in two clusters/taxa/species: Pseudodama farnetensis and
Pseudodama vallonnetensis. Our results support an attribution of the CGR specimens to
Pseudodama vallonnetensis. The present study helps clarifying/to clarify the taxonomic status of
European Plio-Pleistocene Dama-like deer as well as to re-evaluate the geographical and
chronological distribution of this group in the Mediterranean area.
153
LIMB DEVELOPMENT IN EARLY TETRAPODS: INSIGHTS FROM
MACROEVOLUTIONARY PATTERNS OF LIMB PROPORTIONS
C. Perez-Ben1*
1Evolutionary Morphology, Museum für Naturkunde Berlin, Invalidenstr. 43, 10115, Berlin,
Germany.
154
A NEW MEMBER OF ERYMNOCHELYINI (TESTUDINES, PLEURODIRA)
FROM THE EARLY PLIOCENE OF CENTRAL AFRICA
A. Pérez-García1*
1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Avda. Esparta s/n, 28232 Las Rozas
Madrid, Spain.
The pleurodiran turtle Erymnochelys madagascariensis (the ‘Madagascan big-headed turtle’) is the
only representative of Erymnochelyini (Pelomedusoides, Podocnemididae) that is part of the
current biodiversity. Erymnochelys was traditionally used as a wastebasket taxon, but the extant
species from Madagascar is currently considered as the only valid representative of the genus. In
the early 1990s, the remains of several groups of turtles were reported in some Early Pliocene
(Zanclean) outcrops of the Sinda beds, in the Sinda-Mohari region of the Democratic Republic of
Congo. Among them, several postcranial elements were attributed to Erymnochelys sp.
Unpublished cranial remains from that locality, exclusively corresponding to the anterior area of the
skull, were subsequently cited in papers published over the subsequent two decades. These
specimens were recognized, without justification, as compatible with the extant Erymnochelys
madagascariensis. Knowledge about the extinct representatives of the Erymnochelyini lineage has
increased remarkably in recent years. In fact, most of them correspond to genera defined in the
21st century. In this sense, the attribution of the taxon from the Sinda-Mohari region to the genus
Erymnochelys has been recently indicated, without justification, as incorrect. The study of
abundant and well-preserved cranial material from this Pliocene locality, not only corresponding to
the anterior region of the skull but also to elements that make it possible to know most of the
cranial anatomy, allows me to justify its attribution to a new taxon. It corresponds to the only extinct
member of Erymnochelyini represented in the Neogene record by more than one skull.
155
CRANIAL ANATOMY OF THE IBERIAN CENOMANIAN BOTHREMYDID
TURTLE ALGORACHELUS PEREGRINA
A. Pérez-García1*, M. Martín-Jiménez1
1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Avda. Esparta s/n, 28232 Las Rozas
Madrid, Spain.
The European bothremydid turtle Algorachelus peregrina (Bothremydini) is one of the oldest
representatives of the crown Pleurodira known in Laurasia. The species has been identified in
several Iberian Cenomanian outcrops, both in Portugal and in Spain. A partial shell of a single
individual has been found in most of these sites. By contrast, hundreds of remains, attributable to
dozens of individuals, have been identified at its type locality: the uppermost middle or lowermost
upper Cenomanian site of Algora (Guadalajara Province, Central Spain). In fact, Algora have
provided the largest concentration of vertebrate macroremains for the Cenomanian of south-
western Europe so far known, this turtle being the most abundant taxon there. Numerous plates of
Algorachelus peregrina from Algora have been analyzed so far, in addition to several complete or
almost complete shells, not only providing detailed information about its shell anatomy, but also
about its intraspecific variability. Several appendicular elements are also known. However, the
cranial information so far available are exclusively based on a partial skull. A complete and very
well-preserved skull from Algora is here presented. Both the previously known partial skull and the
new complete one have been scanned. The three-dimensional reconstruction of each skull, as well
as that of each of the bones that compose them, has been carried out. Thus, the detailed anatomy
of the skull of Algorachelus peregrina can be characterized. This allows us to noticeably improve
the encoding of the species in the data matrices and, therefore, to provide new information about
its phylogenetic position.
156
THE HOLOTYPE OF HYLAEOCHELYS KAPPA IS NO LONGER THE ONLY
KNOWN SPECIMEN FOR THIS TURTLE GENUS IN THE JURASSIC RECORD
1Grupo de Biología Evolutiva, Facultad de Ciencias, UNED, Avda. Esparta s/n, 28232 Las Rozas
Madrid, Spain.
Turtle remains are very abundant in the Upper Jurassic (Oxfordian to Tithonian) record of the
Lusitanian Basin (West-Central Portugal). The best represented lineage, especially considering its
diversity, is that of the aquatic turtles Thalassochelydia. Thus, several members of the coastal
marine clade Plesiochelyidae have been identified there, through relatively numerous finds since
the 19th century. However, a member of Thalassochelydia not attributable to Plesiochelyidae was
recently identified in the Lusitanian Basin. It corresponds to a representative of Hylaeochelys.
Previously, the presence of this freshwater genus had been exclusively confirmed for the British
Lower Cretaceous record. In fact, Hylaeochelys corresponds to the only representative of
Thalassochelydia found in post-Jurassic levels. Hylaeochelys is known in the Portuguese record
by a single specimen, corresponding to a partial articulated shell. It displays several character
states that allowed its attribution to a different species: Hylaeochelys kappa. Therefore, knowledge
about the Hylaeochelys taxon represented in the Jurassic levels is currently very limited. New data
are provided here by the study of two new specimens of this genus from the Upper Jurassic levels
of the Lusitanian Basin. Each of them comes from a different locality that are not the type locality
of Hylaeochelys kappa. One of these individuals corresponds to a partial articulated shell, which
preserves some regions not represented in the holotype of that species. The other is a
disarticulated partial shell, which provides data on the variability of the genus in Portugal.
157
GIANT SHADOWS IN LATE TRIASSIC SEAS: HISTOLOGICAL ANALYSIS ON
PUTATIVE AND GENUINE GIANT ICHTHYOSAURS BONES
Large to very large unidentified fossil bone shafts from European Late Triassic deposits have been
puzzling the paleontological community since the second half of the 19 th century. Over the
centuries, differing hypothesis have been proposed regarding the nature of these fossils:
amphibian, ichthyosaurian and dinosaurian/archosaurian. In this study a comprehensive
histological description and comparative analysis was conducted to test what we coin as the
“Huene-Lomax hypothesis"; i.e. an ichthyosaurian affinities for large bone fragments of uncertain
origin recovered from different Rhaetian fossil localities across Europe. Resulting histological
comparison of the material in exam, including the Norian holotype of Shastasaurus sikanniensis,
found a common combination of unusual histological features (e.g. Parallel-Woven complex rich in
Interstitial Structural Fibers and preferential secondary remodeling through “double-zoned”
osteons) shared with bonafide giant ichthyosaurs. Our study is therefore unable to find histological
evidence to reject the Huene-Lomax hypothesis, further supporting the widespread presence of
large sized ichthyosaurs at the Tr-Jr boundary. Finally, the aferomentioned shared histological
features, suggests the presence of a yet-to-discover biological or phylogenetic signal possibly
related to the aquisition of large sizes in gravity-free aquatic environment or to specific niches
occupation in late Triassic ichthyosaurs.
158
FEET REFINE THE ECOLOGY OF EARLY THEROPOD FLYERS
M. Pittman1,2,3*, P.R. Bell4, C.V. Miller5, N.J. Enriquez4, X.L. Wang6,7, X.T. Zheng6,7, L.R.
Tsang4,8, Y.T. Tse1, M. Landes9 & T.G. Kaye3
1School of Life Sciences, The Chinese University of Hong Kong, Shatin, Hong Kong SAR, China.
2Department of Earth Sciences, University College London, Gower Street, London, WC1E 6BT,
United Kingdom.
3Foundation for Scientific Advancement, Sierra Vista, Arizona, 85650, United States of America.
4School of Environmental and Rural Science, University of New England, Armidale, NSW 2351,
Australia
5Department of Earth Sciences, The University of Hong Kong, Pokfulam, Hong Kong SAR, China
6Institute of Geology and Paleontology, Linyi University, Linyi City, Shandong, 276005, China.
The feet of modern birds are closely correlated to their ecology. Their toe pads, foot scales, joints
and claws reflect their cursoriality, feeding mode and grasping ability. To reveal these ecological
characteristics from the feet of early theropod flyers, we present equivalent fossil evidence in the
form of soft tissues surrounding the bones. The ecology of the earliest theropod flyers has been
studied using their anatomy, diet and locomotor capabilities as well as the environments and
climates they lived in. Our results are interpreted in the context of these existing lines of evidence.
The earliest theropod flyers including Anchiornis and Archaeopteryx from the Middle-Late Jurassic
had feet indicating a more ground-dwelling lifestyle. In the Early Cretaceous aerial lifestyles
diversified, including generalists such as Confuciusornis and specialists such as the climbing
Fortunguavis. Some early birds had complex ecologies that were seemingly unique among
modern birds e.g., the Berlin Archaeopteryx and the Early Cretaceous Sapeornis. Unexpectedly,
the non-avialan flyer Microraptor was recovered with a more specialised raptorial lifestyle featuring
159
hawk-like characteristics rare among theropod flyers of the time. This suggests that non-avialan
flyers like Microraptor were specialists, similar to certain birds in modern ecosystems.
160
PALEONEUROANATOMY OF THE CENOMANIAN CROCODYLOMORPH,
PORTUGALOSUCHUS
5Department of Vertebrate Zoology, Saint Petersburg State University, Universitetskaya nab. 7-9,
199034, St. Petersburg, Russian Federation.
161
crocodylians. Despite all this, in order to better constrain these results, additional phylogenetic
analyses, including this new morphological character coding, together with DNA data should be
performed.
162
DENTAL VARIATION IN HYPNOMYS MAHONENSIS BATE, 1918 (GLIRIDAE,
RODENTIA, MAMMALIA) FROM THE NEWLY REDISCOVERED TYPE-
LOCALITY
²ICREA, Institut Català de Paleoecologia Humana i Evolució Social (IPHES), Universitat Rovira i
Virgili, Carrer Marcel·lí Domingo s/n (Edifici W3), Campus Sescelades, 43007 Tarragona, Spain.
Punta Esquitxador 17 is a small coastal site of karstic origin, formed by very hard red silt, rich in
remains of small vertebrates. This is the same site from which Dorothea M. A. Bate recovered and
described, in 1918, Hypnomys mahonensis. The limited number of specimens of the type series (a
left maxilla, a right mandible and an M1) has, historically, limited the adequate knowledge of this
species. The recovery of new remains has allowed us to describe the different dental morphotypes
characteristic of H. mahonensis and their comparison with other species of the genus. Overall, H.
mahonensis has simpler tooth patterns than Hypnomys morpheus, with isolated ridges in upper
M1. Moreover, H. mahonensis displays a small transverse ridge connecting the anteroloph and
protoloph. This is a derived feature absent in H. morpheus. Two hypotheses have been put
forward to explain the origin of H. mahonensis: 1) is considered the final representative of a
phyletic lineage originating in Menorca; 2) H. mahonensis evolved from the arrival to Menorca of a
late representative of the existing phylogenetic line in Mallorca (H. morpheus). This study also
made possible to confirm the taxonomic validity of H. mahonensis and its geographical and
temporal distribution.
163
NEW THEROPOD REMAINS FROM THE LATE JURASSIC CAÑADÓN
CALCÁREO FORMATION OF CHUBUT, ARGENTINA
2CONICET, Museo Paleontológico Egidio Feruglio, Fontana 140, 9100 Trelew, Argentina.
Late Jurassic theropod dinosaurs from the former continents of Gondwana are still very poorly
known. From South America, only two taxa have been named so far, the poorly known early
branching tetanuran Pandoravenator, and the enigmatic Chilesaurus. Whereas Pandoravenator is
so far only known from the very fragmentary holotype, which does not allow to establish its
phylogenetic position confidently, Chilesaurus shows a unique combination of characters that has
even led to suggestions that this taxon is not a theropod at all. Recent fieldwork in the Oxfordian-
Kimmeridgian Cañadón Calcáreo Formation of Chubut, Argentina, have led to the discovery of
several new theropod specimens. One represents a second specimen of Pandoravenator, and
provides additional evidence on the anatomy of this taxon, including the first cranial remains. This
new material helps to establish Pandoravenator as an early branching coelurosaur, representing
the first representative of this clade known from the Jurassic of the southern hemisphere. A new
taxon of small theropod dinosaur is represented by at least three partial, articulated skeletons,
which together represent most parts of its anatomy. The pelvis resembles that of Chilesaurus in
being opisthopubic, with a relatively low, simple ilium. Likewise, the fore- and hindlimbs also
resemble those of Chilesaurus, although the foot is more typical theropodan in that the first
metatarsal does not reach the ankle joint. The skull is unusual in being rather short and broad, with
an extremely short snout and an apparently herbivorous dentition, whereas the cervical vertebrae
closely resemble those of elaphrosaurine noasaurids.
164
BALEEN WHALES IN MATERA BASIN: PALEOENVIRONMENTAL AND
BIOSTRATIGRAPHIC RECONSTRUCTION AS REVEALED BY COCCOLITH
ASSEMBLAGES
1Dipartimento di Scienze e Tecnologia, Università degli Studi del Sannio, Via De Santis, 82100,
Benevento, Italy.
2Museo Nazionale di Matera “Domenico Ridola”, Via Ridola, 75100, Matera, Italy
Blue whales (Balaenoptera musculus) are the largest living mammals and the largest known
animals ever appeared on the Earth to date. Balaenoptera musculus plays a pivotal role in modern
ocean ecology as it is a major consumer. On the other hand, ecological and oceanographic
feedback processes may influence baleen whales size, thriving and migration paths. Recently, a
Mysticete partial skeleton (sharing multiple traits with extant Balaenoptera musculus) was retrieved
near Matera, buried into the early Pliocene to middle Pleistocene foredeep deposits of the
Bradanic Trough, southern Italy. The fossil was previously dated at 1.5 – 1.3 Ma.
Here, we attempt at reconstructing the ecosystem in which this huge mammal died, and was
successively buried, using coccoliths. These have been widely used for biostratigraphic purposes
in the central Mediterranean, with several recognizable event. Analysis of the calcareous
nannoplankton fraction throughout the sedimentary succession of the Matera Basin may be useful
to corroborate radiometric age data. Besides, coccolithophore association analysis can yield a
broad range of information about the paleoecological/paleoceanographic environment in which this
Pleistocene analogue of modern days blue whale dwelled. The aim of this work is to shed light on
the timing of the onset of whale gigantism in the Mediterranean and its relationships with
paleoecological and paleoceanographic changes in the study area.
165
BODY SIZE IN THE INSULAR MURID MIKROTIA FROM GARGANO (LATE
MIOCENE, ITALY)
Body size is one of the main determining factors in a mammal’s life strategy. Nowhere is this
clearer than on islands, where mammals tend to change in body size compared to their mainland
ancestors. This study focuses on such an island mammal, the giant rat Mikrotia (Muridae) of the
Gargano palaeoisland of the Italian Peninsula. While the cranial and dental elements of Mikrotia
have been studied elaborately, the postcranial skeleton was largely unexplored. In order to
determine the body weight and locomotory habits of Mikrotia, we studied limb bone features and
calculated indices of three species of Mikrotia from San Giovannino, one of the youngest fissures
of the palaeoisland. Our results were then compared to those of extant rodents with known body
masses and locomotion. The distal joint widths of femur and humerus are the most accurate for
estimating weight, the femur length tends to overestimate body mass. Based on the distal joints,
we estimate the body mass of the smallest Mikrotia species at 140 g, for the middle-sized species
at 630 g and for the largest species at 1700 g. Body mass estimates based on limb bones of
Mikrotia are much higher than those from earlier studies based on the lower incisor.
166
THE FIRST SIVATHERIINE (ARTIODACTYLA, GIRAFFIDAE) FROM THE
MIDDLE MIOCENE OF PORTUGAL
The revision of the giraffid material stored at the Museu Geológico (Lisbon, Portugal) using current
analytical tools and methods leads us to the first identification in Portugal of a member of the
Samotheriinae-Sivatheriinae clade. The material consists of a metacarpal from Casais da Formiga,
Azambuja (MG 5733, later Middle Miocene, MN 7: 13-12.5 Ma). Although previously tentatively
assigned to Palaeotragus sp. due to size, several morphological features point to the specimen
belonging to the widely distributed Samotheriinae-Sivatheriinae clade, whose earliest member
recorded in the Iberian Peninsula is the Vallesian Decennatherium (MN 9-10). Giraffid metapodials
have a high diagnostic value, and according to its morphology MG 5733 is most similar to
Decennatherium (especially the proximal epiphysis and palmar diaphysis), which was present
during the Vallesian (MN 9-10) in the Calatayud-Teruel, Duero, and Tagus basins. It is especially
similar to D. pachecoi (MNCN-42769) from Los Valles de Fuentidueña, with the only relevant
difference being a more dorsoventrally compressed proximal epiphysis in MG 5733. When
compared to Palaeotragus, the condyles in P. rouenii are less developed and distally the diaphysis
does not widen. The central trough is also deeper in P. rouenii and the proximal articular surface is
on the same plane. A PCA analysis reveals this specimen as metrically falling between the ranges
of variability of Decennatherium pachecoi and Palaeotragus rouenii. This is the first report of
Sivatheriinae in Portugal.
167
STABLE ISOTOPE RECORD OF TRICERATOPS FROM A MASS
ACCUMULATION (LANCE FORMATION, WYOMING, USA) PROVIDES
INSIGHTS INTO TRICERATOPS BEHAVIOUR AND ECOLOGY
2Department of Earth Sciences, Faculty of Science, Vrije Universiteit, De Boelelaan 1085, 1081
HV Amsterdam, the Netherlands.
3Institute of Biology, Faculty of Science, Leiden University, Sylviusweg 72, 2333 BE, Leiden, the
Netherlands.
4Faculty of Geosciences, Utrecht University, Princetonlaan 8A, 3584 CB Utrecht, the Netherlands.
Our understanding of Late Cretaceous dinosaur ecosystems from North America has considerably
improved through stable isotope analyses on fossil bones and teeth. Oxygen and carbon isotopic
compositions of structurally-bound carbonate in fossil teeth are commonly used to detect
(temporal) variations in ingested water and food sources, which are in turn related to shifts in
environmental and climatic conditions. So far, these reconstructions are based on spatiotemporally
diverse datasets. Here, we present oxygen (δ18O) and carbon (δ13C) isotopic records from a large
well-constrained monospecific Triceratops bonebed from the Lance Formation (Wyoming, USA),
allowing to elucidate the palaeoecology of these large herbivores in detail, as well as their habitat
use, diet and possible migration. Depending on tooth size, three to eight incremental samples were
taken from seven Triceratops teeth, revealing relatively low intra-tooth δ18O variation (average
1.3 ‰), comparable to contemporaneous hadrosaurs and modern mammals. Ambient water
temperature reconstructions based on associated gar scale δ18O follow modern freshwater fish
and comply with earlier temperature-proxies for the Late Cretaceous. Average δ13C values (-
5.2 ‰) are higher than in modern C3 plant grazers, but support the previously stated hypothesis of
different metabolic fractionation in herbivorous dinosaurs. The combined δ18O and δ13C
Triceratops isotope signatures indicate a living environment intermediate between inland forests
168
and coastal floodplains, which was previously believed to strictly adhere to hadrosaur and
ceratopsian niche partitioning, respectively. Our dataset provides an accurate palaeoecological
reconstruction for Triceratops, and highlights the need for spatiotemporally well-constrained fossil
remains in isotope analyses and large-scale isoscape reconstructions.
169
A BASAL ANKYLOPOLLEXIAN DINOSAUR FROM THE LATE JURASSIC OF
PORTUGAL AND ITS IMPLICATIONS FOR IGUANODONTIAN DIVERSITY
2Museu da Lourinhã, Rua João Luis de Moura 95, 2530-158 Lourinhã, Portugal.
3CI2Paleo/Sociedade de História Natural (SHN), Travessa Florêncio Augusto Chagas, nº8B, 2560-
230 Torres Vedras, Portugal.
5Operational Directorate Earth and History of Life, Royal Belgian Institute of Natural Sciences,
Brussels, Belgium.
Historically, Portugal has been a hot spot for Late Jurassic dinosaur diversity and several
institutions have been conducting fossil collections over the past few decades. Here we re-
describe known specimens of basal ankylopollexian iguanodontians hosted at the Sociedade de
História Natural (SHN) and Museu da Lourinhã (ML), and present novel interpretations. The
material has been recovered from the Upper Jurassic Lourinhã Formation from various localities
and has been figured and described already in peer-reviewed publications or other academic
works. The specimens include: SHN.(JJS).015, associated scapula, humerus, femur and
calcaneum; ML 434 isolated femur (previously regarded as Draconyx loureiroi); SHN.073 isolated
femur; ML 2055 associated femur and partial tibia. These specimens share a bowed femoral shaft,
high crest on the cranial surface of the shaft and pendant 4th trochanter of the femur with an
extensive lamina projecting dorsally, and the difference in size suggests they represent different
ontogenetic stages of the same species. With the abovementioned features, SHN.(JJS).015
presents a combination of characters which distinguish it from Draconyx loureiroi and other Late
Jurassic ankylopollexians, such as: straight scapular blade, glenoid and deltoid fossa not strongly
170
separated. We modified published data matrices to explore the systematic affinities of this
specimen. Employing maximum parsimony, we recover SHN.(JJS).015 at the base of
Ankylopollexia. These results increase the diversity of Late Jurassic iguanodontians present in the
Late Jurassic of Portugal, possibly reporting the first ontogenetic series for ankylopollexians in
Europe. Further investigation on histology is needed to elucidate growth patterns in this clade.
171
3D LIFE RECONSTRUCTION OF THE INTRASPECIFICAL VARIABILITY IN
THE FOSSIL GIRAFFID DECENNATHERIUM REX RÍOS, SÁNCHEZ AND
MORALES 2017 (LATE MIOCENE, IBERIAN PENINSULA)
The recovery of hundreds of skeletal remains of the extinct giraffid Decennatherium rex
Ríos, Sánchez and Morales, 2017 from the Late Miocene deposits of the site Batallones-
4 and Batallones-10 (MN 10, Cerro de los Batallones, Madrid Basin) sheds light on the
complex intraspecific variability of the fossil giraffid. Here we reconstruct for the first time
a hypothetical herd including all the morphological variability recovered in the Batallones
sample. Using Zbrush, which allows for digital 3D modelling, sculpture and painting we
were able to show through a paleoartistic approach the wide range of ontogenetic and
sexual variability present in this taxon. The results include an accurate representation of
its paleohabitat and cranial and postcranial morphological variability with several scenes
including females, juveniles and older males. Fur colour was inferred using knowledge of
the ecosystem and actualism principles.
172
THE LARGE, THE SMALL AND THE UNUSUAL – A NEW, MULTISPECIFIC
TANYSTROPHEID ASSEMBLAGE FROM SOUTHERN POLAND
1Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, 00-818, Warsaw, Poland.
2Institute of Evolutionary Biology, Faculty of Biology, University of Warsaw, Żwirki i Wigury 101,
02-089, Warsaw, Poland.
The study was financially supported by the National Science Centre (NCN) grants
2017/27/B/NZ8/01543, 2020/39/O/NZ8/02301 (awarded to T. Sulej) and 2019/35/N/NZ8/03806
(awarded to Ł. Czepiński).
173
A NEW DIGITAL TOOL FOR STUDYING SMALL TETRAPOD FOOTPRINTS
Since the early 2000s, digital photogrammetry has become increasingly popular in ichnology. Such
approach has enabled the digitisation of footprints and the objectivisation of data collected in the
field. In particular, the opportunity to analyse the footprints through colour-coded images
represents a powerful tool to highlight the morphological characters preserved in the tracks. This
approach has been used in the recent study of almost all medium- and large-sized tetrapod
footprints. In contrast, photogrammetry or other 3D application for centimetric and millimetric
footprints is sparse and limited. In this contribution, we introduce the use of optical scanners with
blue light technology in ichnology. This tool allows high-resolution scans with the production of
closed double precision polygonal mesh. It also offers multiple advantages in addition to the very
high-resolution model, with a significant reduction in data acquisition and processing time
compared to photogrammetry.
Here we present preliminary results of the application of this new technology to several very small
Permian footprints from the Museum of Nature South Tyrol (Bolzano, Italy) collection. Three-
dimensional data were acquired using both scanners and traditional photogrammetry and the
results were compared highlighting the huge resolution improvement obtained using scanners. We
developed a new extensive workflow compatible with file type generated by the scanner's
proprietary software to produce colour-coded image (e.g., Digital Elevation Model). The obtained
174
digital elevation models, allow to ensure a new level of detail in the interpretation of these small
footprints, which have so far only been studied with bare eye.
175
A BETA TAXONOMY APPROACH TO LATE JURASSIC AND EARLY
CRETACEOUS DINOSAUR ASSEMBLAGES
2 Museu da Lourinhã, Rua João Luis de Moura 95, 2530-158 Lourinhã, Portugal
3 Department of Aerospace Science and Technology, Politecnico di Milano, via La Masa 34, 20156
Milano, Italy
Although dinosaurs achieved worldwide distribution during the Mesozoic Era, their biogeographic
distribution patterns are subject of high debate. A major geological influence on dinosaur
distribution is the progressive separation between Gondwanan and Laurasian landmasses, leading
to a faunal segregation by mid/Late Cretaceous. To elucidate the tempo and modo of such
segregation, we adopted a Beta Taxonomy approach by comparing the similarity of dinosaur
occurrences of various geological formations around the globe. Jaccard, Dice, and Simpson
similarity indices were employed for the following time-bins: Kimmeridgian-Tithonian, Berriasian-
Valanginian, Barremian, and Aptian-Cenomanian. Based on each of these indices, we performed:
1) a set of cluster analyses (UPGMA, neighbor joining, K-means), 2) Principal Coordinate Analysis,
and 3) Correspondence Analysis. Results derived from the Jaccard index presented the highest
statistical support, appearing to be more robust when compared to the other two indices. Absence
of faunal segregation between Laurasia/Gondwana was identified during Kimmeridgian-Tithonian
interval, although we found evidence of Asiatic provincialism. This distribution remains unchanged
during Berriasian-Valanginian, even though sampling bias in this interval partially affected our
results. Similarly, Barremian sampling bias hampered the possibility to discriminate Laurasian and
176
Gondwanan faunas, despite having higher statistical support in our analyses than the other time-
bins. Finally, the Aptian-Cenomanian interval presented a clear faunal segregation between
Laurasia and Gondwana, suggesting that a major faunal re-organization occurred no later than the
Barremian. Further investigation, using clade-specific historical biogeography techniques, is
required to clarify whether dispersal or vicariance events shaped this faunistic re-arrangement.
177
SPHEROLITH VARIABILITY OF MEGALOOLITHID AND FUSIOOLITHID
DINOSAUR EGGSHELLS
The oofamilies Megaloolithidae and Fusioolithidae have a rich fossil record composed of clutches,
eggs and eggshells, most of which come from the Campanian-Maastrichtian of South America,
Europe and Asia, and have been attributed to sauropods as their producers. Overall,
Megaloolithidae and Fusioolithidae present similar eggshell macro and microstructure features and
are mainly differentiated by the degree of fusion of their eggshell units (spheroliths). Many of these
ootaxa present a relatively wide range of variability in thickness and/or ornamentation. These
features are linked to the height, width, and thus the shape of the spheroliths that form the
eggshell. Therefore, the morphological variability of spheroliths of megaloolithid and fusioolithid
specimens has been assessed. The length and width, along with the morphology of the accretion
lines of each spherolith of each sample, has been registered and analysed to search for
differences between ootaxa. The height of the spheroliths between the analysed ootaxa ranges
from 0.9 to 2.9mm, whereas the width ranges from 0.1 to 1.1mm. Generally, the range of variability
in width is greater than that of spherolith height within each ootaxon, with few exceptions. These
results have been compared with dinosaur eggshells from the Poyos fossil site (Upper Cretaceous,
Villalba de la Sierra Formation, Guadalajara, Spain). The morphotypes from Poyos have the
lowest values for both spherolith height and spherolith width among the analysed ootaxa.
178
2D OPC: A NEW METHOD TO DESCRIBE DENTAL COMPLEXITY OF
HERBIVORE MAMMALS
1Departamento Ciencias de la Vida, GloCEE Global Change Ecology and Evolution Research
Group, Universidad de Alcalá, 28805, Madrid, Spain.
Dental morphology is a major aspect of ecological and evolutionary studies of both extant and
fossil mammalian species. Previously two-dimensional (2D) methods quantified the resistance of
enamel through the cutting area as the ratio between enamel length and tooth area. Other
approach uses the fractal index as an estimate of enamel folding. However, these variables give
no information about local variations of important enamel structures within a tooth. In recent years,
a series of novel methods have been proposed to analyze three-dimensional (3D) dental models.
However, 3D data poses its own limitations: the initiatives to make it widely accessible and freely
available are still limited and 3D scanned data needs computationally expensive pre-processing
and costly software to optimize raw mesh/voxel information. On the other hand, 2D images are
broadly prevalent in the literature as a main source of information for systematic studies, is
available for nearly every species, and requires little preprocessing. With this in mind, we created a
novel workflow using 2D image analysis techniques with unprecedented resolution over individual
tooth’s structures by combining multiple methodological tools included in the image analysis
software ‘Fiji’. A general case study is proposed using two independent clades within the Family
Rhinocerotidae containing species regarded as hypsodont. To test the robustness of this tool
under different practical scenarios, additional samples have been evaluated. These include
differences in image size, rotation, or differences in dental wear.
179
THE ACEDERAL CAVE: A NEW QUATERNARY LOCALITY WITH FOSSIL
LYNXES IN ARAGÓN (NORTHEASTERN SPAIN)
1Department of Earth Sciences, Faculty of Science, University of Zaragoza. Pedro Cerbuna 12,
50009, Zaragoza, Spain.
8Departament d'Història i Història de l’Art. Universitat Rovira i Virgili, Tarragona, Av. Catalunya 32,
43002, Tarragona, Spain.
The Iberian lynx (Lynx pardinus) is an endemic wild cat, now restricted to the South of the Iberian
Peninsula, and the most threatened carnivore in Europe. The Acederal cave, located in the
municipality of Aldehuela de Liestos (Zaragoza, Aragón, Spain), has been found to contain an
important accumulation of lynx remains (at least two almost-complete skulls and abundant
postcranial remains), preliminary assigned to Lynx pardinus. The cavity has a moderate inwards
development, and, at the bottom of a 26-m-deep sinkhole, there is an area where bones of large
mammals, that appear half-buried in the clay infill, were accumulated. Despite a precise
chronological framework is not yet available, the facies, the conservation state, and the faunal
association (including Equus sp., Cervidae indet., Caprinae indet., Felis sp., Martes sp., Vulpes
180
vulpes, Oryctolagus cuniculus, Rhinolophus sp. and Corvidae indet.) would be consistent with a
Late glacial to early Holocene accumulation, but a more recent chronology cannot be ruled out.
The lynx fossil record in the Quaternary of Aragón is limited to the sites of La Puebla de Valverde
(Lower Pleistocene, Villafranchian) and the Cueva de Chaves (Upper Pleistocene, Magdalenian).
Albeit the study of the site is still at an early stage (further studies are scheduled, including ancient
DNA analysis and direct datings), this locality has the potential to contribute to the
phylogeographical knowledge of this species in a significant way.
181
REDESCRIPTION OF THE NEOTYPE OF PLATEOSAURUS AS FOUNDATION
FOR FUTURE STUDIES
J. Schaeffer1*, R. Schoch1,2
The taxonomy of Plateosaurus, the fifth dinosaur genus named, has always been majorly
problematic. In the early 1900’s more than 20 species were erected within three different genera
that are currently all synonymised with Plateosaurus trossingensis. However, there is an enormous
amount of morphological variability within the P. trossingensis specimens in the collection of the
Staatliches Museum für Naturkunde Stuttgart (SMNS) in Germany, as well as with specimens from
the same locality in other collections and from other localities. In the most recent literature, it has
been proposed that the variability is consequential of developmental plasticity, however the
explanation is evidently multifaceted, with further investigation of material from all horizons and
localities essential to resolve this question. In 2019, SMNS 13200 was designated as the neotype
of Plateosaurus after the holotype was deemed undiagnostic – however, no modern description of
a relatively complete, mature specimen of P. trossingensis is currently available. The most recent
description was made by von Huene in 1926, written in German. Here we redescribe the neotype
of Plateosaurus trossingensis and present an updated phylogeny. The results will form the basis of
several other associated projects currently underway, such as a large-scale 3D geometric
morphometric study of Plateosaurus material from across all known localities and collections,
description of pathologies, both in the neotype and other undescribed specimens, as well as
fieldwork at the historical Trossingen site to collect more stratigraphic and sedimentological data
and to excavate additional Plateosaurus specimens from the underrepresented beds.
182
INFERENCE OF SAUROPOD POPULATION ECOLOGY FROM TRACKWAYS?
– ASSESSMENT OF THE SWISS JURA TRACK SITES
3 Dipartimento di Scienze della Terra, Università di Firenze, via G. La Pira 4, 55121, Firenze, Italy.
Studying population ecology in the vertebrate fossil record is a difficult task because of drawbacks
such as taphonomic biases. Fossil tracks, conversely, are the in situ evidence left by animals to
provide data for the likely natural grouping and relative abundance. This raises the question: can
we infer potential changes in populations repeatedly traversing an area, through time, from their
trackway record alone? This question seems impossible to answer in the affirmative as continental
ichnology is fraught with inherent complexities (e.g., small surface areas and time-restricted track
sites, preservation bias, etc.). Uniquely, the Swiss Jura track sites can provide insight into
sauropod populations from their trackways. This is because of the nature of the track sites, their
distribution in space and time, the easy stratigraphic correlation between sites, and the repetitive
use of the track site areas. We document temporal patterns in abundance and distribution of
sauropod trackways and by inference sauropod dinosaurs using the Jura coastal platform in the
Late Kimmeridgian. We deduce population sizes, grouped walking directions, and the presence of
juveniles, subadults, and adults. Overall, the Swiss Jura coastal platform acts as a means to
estimate population sizes, stage classes, and gregarious movement of sauropod herds through
time. Moreover, the rare occurrence of tracks attributed to juveniles at different stratigraphic
heights (i.e., different times), and at different sites, provides an exceptional glimpse into the
sauropod populations. Although preliminary, this approach brings into question seasonal habitat
use, nesting behavior, dispersal, parental care, and predation in the Jura area.
183
MESOZOIC HEAVY MEAL: TELEOSAUROID REMAINS IN A REGURGITALITE
FROM THE UPPER JURASSIC OF NORTHEASTERN ITALY
1Department of Chemical and Geological Sciences, University of Modena and Reggio Emilia, Via
Campi 103, 41125, Modena, Italy.
2Department of Earth and Planetary Sciences, Yale University, 210 Whitney Avenue, New Haven,
Connecticut, 06511, United States.
3Department of Natural Sciences, National Museum of Scotland, EH1 1JF Chambers Street,
Edinburgh, United Kingdom.
4Department of Earth and Environmental Sciences, Pavia University, via Ferrata 1, I-27100, Pavia,
Italy.
5Department of Geosciences, University of Padova, Via Gradenigo 6, I-35131, Padova, Italy.
The Upper Jurassic record of marine reptiles from the western Tethys is poorly documented,
together with information on their trophic interactions. A fragmentary set of small crocodylomorph
bones from the Rosso Ammonitico Veronese Fm. (RAV) of Ponte Serra (northeastern Italy) was
officially described in 2021. The age of the specimen was restricted to the late Kimmeridgian-
earliest Tithonian thanks to calcareous nannofossil analysis. The fossil is tentatively assigned to
the teleosauroid subfamily Aeolodontinae based on the presence and morphology (size, thickness
and ornamentation) of osteoderms, its stratigraphic position, and pelagic nature of the RAV
deposit. The peculiar clustering of axial, pelvic and dermoskeletal elements suggests its
interpretation as a bromalite, a fossilized item that was processed by the digestive system of a
predator or scavenger. This hypothesis is supported by UV fluorescence, SEM microstructural
analysis of the bone and EDS geochemical analysis of the surrounding matrix. Limited bone
etching and a dearth of background phosphorus in the surrounding matrix further distinguish the
specimen as a regurgitalite, an egested fossilized mass that experienced limited gastric processing
by the producer. Data on phosphorous weight percentage from this specimen closely match the
array of other regurgitalites, while substantially deviating from the geochemical composition of
coprolites. The specimen represents the first occurrence of Aeolodontinae in Italy as well as its
most southward record in the western Tethys ocean. The fossil marks the first record of a marine
184
crocodylomorph preserved in a regurgitalite, bringing new insights into the taphonomy and
paleoecology of the Upper Jurassic mesopelagic realm.
185
REDISCOVERED THALATTOSUCHIAN AND ICHTHYOPTERYGIAN
MATERIAL FROM THE MIDDLE-UPPER JURASSIC OF NORTHERN ITALY
1Department of Chemical and Geological Sciences, University of Modena and Reggio Emilia, Via
Campi 103, 41125, Modena, Italy.
2Department of Palaeontology, Staatliches Museum für Naturkunde, Rosenstein 1, 70191,
Stuttgart, Germany.
3Department of Natural Sciences, National Museum of Scotland, EH1 1JF Chambers Street,
Edinburgh, United Kingdom.
4Department of Geosciences, University of Padova, Via Gradenigo 6, 35131, Padova, Italy.
The mesopelagic depositional setting of the Rosso Ammonitico Veronese of northeastern Italy
(RAV, Bajocian-Tithonian) is known since the XVIII century for the very rare, fragmentary, and
badly preserved record of metriorhynchids, ichthyosaurs and pliosaurs. Preliminary information is
presented here for two “new” specimens from the Altopiano d’Asiago area rediscovered during
collections survey. The first, briefly reported in a museum short note, was discovered in the ‘80s
and housed in the collection of Museum of Geology and Paleontology of Padova. After a quick
evaluation the specimen is assigned to Metriorynchoidea thanks to unequivocal large impressions
of the prefrontals in the matrix. The specimen is represented by impressions and fragments of
middle and posterior portions of the skull together with mandibular rami, one tooth, cervical
vertebrae and ribs on a slab of limestone from the RAV lower member (Bajocian-Bathonian). The
second specimen was found in a RAV boulder used in a dam in the ‘90s and later housed in
Chioggia Civic Museum; shortly after restoration the specimen was interpreted as an ichthyosaur.
The specimen is represented by mandibular elements in ventral view, posterior portions of the
skull and some semi articulated vertebrae. A rich invertebrate fossil fauna is also found associated
with the skeleton. The boulder appears to come from the upper member of the RAV
(Kimmeridgian-Tithonian). While the poor preservation challenges any specific taxonomic
attribution, the two specimens enrich the record of marine vertebrates from the RAV Fm., bringing
valuable paleoecological and taphonomical insight on this pelagic depositional environment.
186
FUNCTIONAL MORPHOMETRY OF MAMMALIAN LIMB BONES:
CARNIVORES AND UNGULATES COMPARED THROUGHOUT THE
CENOZOIC
2 Clinic for Zoo Animals, Exotic Pets and Wildlife, Vetsuisse Faculty, University of Zurich,
Winterthurerstr. 260, 8057, Zurich, Switzerland.
Mammals show a broad range of long bones morphologies linked to their ecological diversity.
Even though mammalian morphologies were well established near the beginning of the Cenozoic,
modern taxa exhibit different adaptations within and between clades. For example, Artiodactyla
and Perissodactyla have both evolved elongated limbs in proportion to their large body mass. In
contrast, predatory Carnivora do not show particularly large body sizes and their limb proportions
can vary a lot between taxa due to specific hunting behaviours. The hypothesis of coordinated
diversification events between these clades has rarely been tested, and it is still not clear why and
when predators and prey evolved specific locomotory adaptations. Here, we assessed
morphological variation of limb proportions in a rich sample of extant and fossil Carnivora,
Artiodactyla, and Perissodactyla in order to examine hypothesized associations with ecological
adaptations and diversification through time. The phylogenetic relationships among species were
incorporated into the analysis to allow the use of comparative methods. Results show that limb
bone proportions change in relation to life habit with amphibious species having relatively shorter
limbs than ground-dwellers. Also, major climatic events clearly appeared to have had a temporal
impact on patterns of morphological diversification, expressed as disparity, in several clades and
ecological groups. Ground-dwelling Carnivora and Artiodactyla showed increased morphological
disparity during the coldest Cenozoic phases. Our analysis provided evidence that patterns of
disparity were often synchronous among large mammalian clades although we did not find
evidence that a predator-prey “arms race” was responsible for these changes.
187
DENTAL MESOWEAR ANALYSIS AND MASS ESTIMATION OF GAZELLA
BOUVRAINAE (MAMMALIA, BOVIDAE)
Gazella bouvrainae Kostopoulos & Athanassiou, 1997, is an Early Pleistocene gazelle species,
typical of the Middle Villafranchian faunas of Greece. This taxon is absent from Western Europe,
but constitutes a rather common find in Greece, being the most common species of the genus
Gazella. The material used for this study includes numerous relatively well-preserved dental
elements from three Lower Pleistocene Greek mainland localities, Karnezeika, Sésklo and
Gerakarou, as well as post-cranial elements from Karnezeika. Phylogenetically, G. bouvrainae is
closely related to contemporaneous Asian gazelle taxa, while its palaeoecological profile is yet not
fully investigated. The main aim of this study is to partially fill this gap by analysing its dental wear
pattern, utilizing the extended mesowear technique. Moreover, we attempt to estimate its average
body mass, based on the metric characters of the radius and the femur. Mesowear analysis of G.
bouvrainae molars showed a mixed to browsing diet, similar to most extant gazelles and the partly
contemporary, European species G. borbonica Depèret, 1884. As far as its body mass is
concerned, it has been estimated to 19–20 kg, comparable with the extant Thompson’s gazelle,
Eudorcas thompsonii (Günther, 1884).
188
A NEW EARLY MIOCENE ELASMOTHERIINE FROM SWITZERLAND:
PHYLOGENETIC AND PALAEOBIOGEOGRAPHICAL INSIGHTS
The study of fossil remains from Benken (early Miocene; Zurich Canton, Switzerland) unveils an
unexpected rhinocerotid taxonomic richness, with seven species identified. They are most likely
originating from two distinct levels referable to the MN4, with 6 and 3 species recorded,
respectively. Interestingly, two skulls and several isolated remains (permanent and decidual teeth,
postcrania) are referable to Victoriaceros, which supports the first occurrence of the genus outside
of Africa. These new remains support a new species. The main diagnostic characters are a skull
with a concave dorsal profile, a sole nasal horn, and a U-shaped nasal notch. The third upper
molar (M3) has pinched hypocone and a strongly-developed antecrochet. A parsimony analysis,
including 36 taxa and 282 morpho-anatomical characters allows for retrieving the monophyly of
Victoriaceros, as a sister group to the Elasmotheriina sensu stricto. From a palaeobiogeographical
perspective and thanks to ancestral character reconstruction, the topology retrieved reveals that
the Victoriaceros clade likely diversified in Africa. This is compatible with the first spread of
Eurasian rhinos toward Africa during the earliest Miocene. The unexpected presence of a new
species of Victoriaceros in Benken both considerably expands the palaeobiogeographical range of
the genus and documents the oldest occurrence of rhinocerotids of African ancestry in Eurasia.
189
BOVID DIVERSITY AT THE LATE MIOCENE FAUNA OF PIKERMI (ATTICA,
GREECE) AS REVEALED BY NEW EXCAVATION DATA
S.D. Sklavounou1*, S.J. Roussiakis1, D.S. Kostopoulos2, P.V. Filis1, I.X. Giaourtsakis3,
G.E. Theodorou1
Bovids emerged during the Neogene and continued to disperse and radiate almost on a global
scale, constantly adapting to ongoing ecological conditions. A significant diversity of bovid species
has been revealed at Pikermi -one of the richest fossil-bearing localities of the late Miocene of
Europe, which has been extensively excavated and studied since the 19th century. Since 2008, a
series of recent and stratigraphically calibrated excavations (NKUA-SARG Project 70/3/12977) has
taken place, primarily at the new sites PV1 (“PV”=“Pikermi Valley”) and PV3 (which correlates with
the historical excavations). The abundant bovid material that has been extracted carries much
potential to provide a thorough assessment of the locality’s bovid systematics, paleoecology and
biostratigraphy. The studied specimens consist of numerous craniodental elements, representing
eight different bovid species: Tragoportax amalthea, Miotragocerus valenciennesi, Gazella
capricornis, Oioceros rothii, Palaeoreas lindermayeri, Palaeoryx pallasi, Protragelaphus skouzesi
and Sporadotragus sp. Preliminary examination of the material provides several new insights.
Among them is the presence of a Sporadotragus species, which does not correspond to
Sporadotragus parvidens known hitherto from the historical excavations. Further, the absence of
Protoryx carolinae and Prostrepsicerus rotundicornis thus far from the studied sample is also
notable, since they are considered to be typical species of the Pikermian fauna. The strong
occurrence of Protragelaphus skouzesi at the new sites is also noteworthy. Additionally, the cranial
dimensions of Palaeoreas lindermayeri specimens seem to exhibit inter-site differences. All these
points may indicate variations (with a possible temporal component) in the bovid diversity among
the different sites at Pikermi.
190
NEW INFORMATION ON THE ARCHOSAURS FROM THE LATE TRIASSIC
FISSURE FILLS LOCALITY OF PANT-Y-FFYNNON, WALES
1Department of Earth Sciences, Natural History Museum, Cromwell Road, London SW7 5BD,
United Kingdom.
4National Museums Scotland, Chambers Street, Edinburgh EH1 1JF, United Kingdom.
The Late Triassic to Early Jurassic fissure fill localities of the Bristol Channel area preserve a
diverse fauna of mostly small-bodied vertebrates, which has provided important insights into the
early evolution of major tetrapod groups such as mammaliaforms, rhynchocephalians,
crocodylomorphs, and dinosaurs. The Late Triassic site at Pant-y-ffynnon yields a particularly rich,
but poorly understood assemblage of archosaurs, including the recently named theropod dinosaur
Pendraig milnerae, the cursorial crocodylomorph Terrestrisuchus gracilis, the small
sauropodomorph Pantydraco caducus, and the enigmatic pseudosuchian Aenigmaspina
pantyffonnonensis. Ongoing research has revealed several new insights into this fauna. Pendraig
is identified as a small-sized non-coelophysid coelophysoid dinosaur. A revision of Aenigmaspina,
characterised by unique, bifurcating osteoderms and conspicuously T-shaped neural spines on the
cervical and anterior dorsal vertebrae, recovers this genus as the sister taxon of Erpetosuchidae +
Aetosauriformes in a new phylogenetic analysis. Finally, CT-scanning has elucidated the
braincase anatomy of Terrestrisuchus gracilis, highlighting several features likely plesiomorphic to
Crocodylomorpha, such as a quadrate that is not fused to the braincase. Together, these studies
reveal the significance of the fissure fills fauna from the Bristol Channel? Of Pant-y-ffynnon? for
our understanding of early archosaur evolution and diversity.
191
COMPARATIVE ANATOMY OF THE PASSERINE CARPOMETACARPUS
SUPPORTS THE PRESENCE OF CROWN SUBOSCINES IN THE OLIGOCENE
OF EUROPE
1Department of Earth Sciences, University of Cambridge, CB2 3EQ, Cambridge, United Kingdom.
2Australian Museum Research Institute, Australian Museum, NSW 2010, Sydney, Australia.
4Department of Earth Sciences, University of Oxford, OX1 3AN, Oxford, United Kingdom.
Among crown birds, the major clade Passeriformes (passerines) exhibits unparalleled diversity,
comprising over 6,000 living species and accounting for over 60% of extant bird species. However,
the early evolutionary history of this key living group remains enigmatic due to a relative lack of
research attention on passerine comparative morphology. This has hampered attempts to interpret
the clade’s fossil record, obscuring patterns of morphological evolution across one of the most
diverse clades of extant vertebrates. Numerous potentially important crown passeriform fossils,
including some of the oldest crown passerines, have proven challenging to place in a precise
phylogenetic context. This is due in part to a paucity of phylogenetically informative characters
from across the passerine skeleton, as well as inadequate characterisation of variation in key
skeletal elements that preserve readily in the fossil record. Here, we begin to address this gap in
our understanding of passerine evolution by presenting a comparative framework for a
morphologically variable, functionally important, and frequently-fossilised skeletal component of
the wing: the carpometacarpus. We sampled >70% of extant family-level passerine clades (132
extant species) as well as several fossils from the Oligocene of Europe, and scored them for 54
phylogenetically informative carpometacarpus characters optimised on a recently published
phylogenomic scaffold. We document a considerable amount of previously undescribed
morphological variation among passerine carpometacarpi, and, despite widespread homoplasy,
192
our results support the presence of representatives of both crown group oscines and crown group
suboscines in Europe during the Oligocene.
193
DIVERSIFICATION “EARLY BURSTS” AND SMALLER TAXA DON’T
DIVERSIFY QUICKER DURING THE RISE OF ARCHOSAURS
1Department of Geology, University of Liège, Bât. B18 Quartier Agora, Allée du Six Août 14, 4000
Liège, Belgium.
The archosauromorph radiation, beginning after the End-Permian extinctions and progressing
during the Triassic, was a spectacular evolutionary radiation that ultimately begat the dinosaurs
and pterosaurs, which would dominate the Mesozoic land and skies, and even the stunning
diversity of modern birds. However, phylogenetic relationships underlying the Triassic radiation are
–like of many fossil clades– greatly uncertain, relying as they do on morphology from an
incomplete fossil record. Thus, methods of estimating diversity dynamics that rely on phylogeny
should best be compared and complemented with phylogenetic topology-free methods for a better
understanding of this radiation. Here, we employ, for the first time in this group, the occurrence-
based Bayesian approach PyRate to estimate extinction and origination rates for
archosauromorphs from the late Permian to Early Jurassic. We use chronostratigraphic uncertainty
for most species rather than actual occurrences, due to abundant true singletons and test for
correlations of diversification rate with body size using femoral length as proxy. We find a latest
Permian diversification peak, meaning that the initial radiation was coeval with major extinctions,
and possibly evidencing morphologically cryptic early diversification. Subsequently, net
diversification rates gradually decline, reaching zero in the early Norian, potentially indicating
niche-filling. Individual clades show similar patterns of initially high diversification dropping off
through time, indicating “early bursts” of diversification following development of ecomorphological
novelty. We find no significant correlation of diversification with body size, but the relationships are
194
generally positive, contradicting hypotheses that larger-bodied clades are more prone to extinction
or less likely to diversify.
195
THE RECORD OF ALEPHIS TIGNERESI FROM CAMP DELS NINOTS (LATE
PLIOCENE, IBERIAN PENINSULA)
4Universitat Rovira i Virgili (URV), Facultat de Lletres, Av. Catalunya 35, 43002 Tarragona, Spain.
Camp dels Ninots (Caldes de Malavella, NE Spain) is one of the most remarkable Konservat-
Lagerstätte of the European Pliocene. The site is located in la Selva depression, within the Catalan
Volcanic Complex. The intense volcanic activity that characterized this area during the Pliocene
led to the creation of the Camp dels Ninots maar lake (CN), and the related fossiliferous site
chronologically constrained at ~3.1-3 Ma (MN15–16 transition, late Ruscinian). The particular
anoxic condition of the lake bottom waters allowed the preservation of a rich vertebrate
assemblage with several skeletons in anatomical connection including: bony fishes, amphibians,
reptiles and mammals. The latter group is particularly interesting for the presence of large
ungulates such as the bovid Alephis tigneresi, the rhinoceros Stephanorhinus cf. jeanvireti and the
tapir Tapirus arvernenis. Here we present the preliminary study performed on the Alephis records
which stands out as one of the most abundant collections of Late Pliocene bovid in Europe. This
genus, closely related with the basal bovine Parabos, populated the Mediterranean area until the
earliest stages of Ruscinian. The relatively low-crowned teeth and the extremely slender limbs,
evidence how this animal was adapted to thrive in humid, closed habitats, as the one inferred for
CN site. The progressive harshening of the climatic conditions and the shrinking of the forested
areas, started at the end of Pliocene, ultimately led to the extinction of these animals; replaced by
Leptobos as the main representative of the large bovine guild in the Early Pleistocene of Europe.
196
MIDDLE MIOCENE METATHERIANS FROM JUAN GUERRA (SAN MARTIN,
PERU) EXPAND THE RECORD OF SMALL MAMMALS IN THE PROTO
AMAZONIA
197
13Seção de Paleontologia, Museu de Ciências Naturais, Secretaria do Meio Ambiente e
Infraestrutura, Avenida Dr. Salvador França, 1427, 90690-000, Porto Alegre, Brazil.
Currently, marsupials (modern members of Metatheria), are widely distributed in tropical and
subtropical areas of South America, but poorly represented in their fossil record. Except for the
species-rich fossiliferous localities of La Venta (Colombia), additional Miocene metatherians from
tropical-equatorial South America are sparsely reported in Bolivia, Brazil, Peru, and Venezuela.
Here, we introduce new metatherian remains recovered in the late middle Miocene TAR-31
locality, Peru, assigned to the early Laventan South American Land Mammal Age (~13 Ma) by
mammalian biostratigraphy. U-Pb analysis made on detrital zircon grains from TAR-31 gave a
maximum depositional age of 17.4 ± 0.12 Ma. Three metatherian taxa are recognized at TAR-31:
the didelphid Thylamys cf. T. colombianus, plus the paucituberculatans Palaeothentes sp.
(palaeothentid) and the new abderitid Pitheculites ipururensis nov. sp. This assemblage, along
with a cebid primate at TAR-31, suggests predominantly humid and warm tropical conditions, with
the occurrence of both forests and drier habitats in the surroundings. Comparisons with other
Miocene metatherian assemblages at low and mid latitudes of South America clearly confirm
closest relationships between TAR-31 and La Venta (previously observed on monkeys and
rodents), as well as, to a lesser extent, with Acre River local fauna and Madre de Dios (MD-67)
and in Brazil and Peru, respectively. The current results further the hypothesis of Western
Amazonia as a single and consistent biogeographical region for land mammals over middle
Miocene times, at the western edge of the Pebas Mega-Wetland System, and highlight the role of
the Amazonian region concerning marsupial Neogene radiation.
198
NEOTAPHONOMICAL OBSERVATION OF A BONE BED FORMATION IN A
RIVER BED
2Polish Tourist and Sightseeing Society – Branch in Ojców, Ojców 15, 32-047 Ojców, Poland.
This research project is supported by the National Science Centre, Poland (no.
2019/32/C/NZ4/00150).
199
MALIGNANT NEOPLASM IN METOPOSAURUS VERTEBRA
1Institute of Earth Sciences, Faculty of Natural Sciences, University of Silesia, Bedzinska 60, 41-
200 Sosnowiec, Poland.
3Institute of Paleobiology, Polish Academy of Sciences, Twarda 51/55, 00-818 Warsaw, Poland
5Carnegie Museum of Natural History, 4400 Forbes Ave, Pittsburgh, PA, 15213, United States of
America.
Each case of cancer in the fossil record is interesting for general biology as well as oncology,
providing – from an evolutionary perspective – precious information on somatic vulnerability to
neoplasms in extinct animals. On the other hand, the documentation of skeletal pathologies in
amphibians is scarce, both in herpetological and paleontological literature. Here we demonstrate
the first case of the development of advanced spinal osteosarcoma in the temnospondyl
Metoposaurus krasiejowensis from the Late Triassic of Poland. Massive, irregular outgrowths on
the post-cervical/anterio-dorsal vertebra (intercentrum) of the metoposaur spread in multiple
directions, with the lateral domination. The location of the bone mass surrounding the affected
vertebra and clear boundary between normal and altered bone suggest that the neoplasm
originated from the outer layer of the periosteum. It is consistent with one of the most common
primary neoplasms affecting the musculoskeletal system called osteosarcoma. The neoplasm
started on the cortical surface of the bone, affected the entire periosteal domain and penetrated
inside the vertebral intercentrum towards the endosteal domain, indicating parostotic variant of
osteosarcoma. Spinal osteosarcomas are rare and aggressive neoplasms in humans, making up
only 0.6-3.2% of all osteosarcomas. Their appearance in Metoposaurus a distant relative of
modern amphibians imply that at large, extinct genera were more vulnerable to cancer than much
smaller, recent species, believed to be cancer-resistant.
200
This research project is supported by the National Science Centre, Poland, grant no.
2019/32/C/NZ4/00150.
201
DO CONICAL AND SABERTOOTHED CATS REPRESENT AN EXCEPTION TO
CRANIOFACIAL EVOLUTIONARY ALLOMETRY?
D. Tamagnini1,2*, M. Michaud3, C. Meloro4, P. Raia5, L. Soibelzon6,7, P. S. Tambusso8,9,
L. Varela8,9, L. Maiorano1,2
1Department of Biology and Biotechnologies “Charles Darwin”, University of Rome “La Sapienza”,
00185, Rome, Italy.
2Museum of Zoology, Sapienza Museum Centre, University of Rome “La Sapienza”, 00185, Rome,
Italy.
3Department of African Zoology, Royal Museum for Central Africa, 3080, Leuvensesteenweg 13,
Tervuren, Belgium.
4 Research Centre in Evolutionary Anthropology and Palaeoecology, School of Biological and
Environmental Sciences, Liverpool John Moores University, L3 3AF, Liverpool, United Kingdom.
5Dipartimento di Scienze della Terra, dell’Ambiente e delle Risorse, University of Naples Federico
II, 80126, Napoli, Italy.
6División Paleontología Vertebrados, Museo de La Plata, Facultad de Ciencias Naturales y Museo,
Universidad Nacional de La Plata, B1900, Paseo del Bosque s/n, La Plata, Argentina.
7Consejo Nacional de Investigaciones Científicas y Tecnicas (CONICET), C1425FQB, Godoy Cruz
2290, CABA, Argentina.
8Departamento de Paleontología, Facultad de Ciencias, Universidad de la República, 11400, Iguá
4225, Montevideo, Uruguay.
9Servicio Académico Universitario y Centro de Estudios Paleontológicos (SAUCE-P), Universidad
de la República, 91500, Santa Isabel s/n, Sauce, Departamento de Canelones, Uruguay.
202
felines. By contrast, Machairodontinae fail to support CREA. The adoption of different landmark
configurations, phylogenetic hypotheses, and corrections for phylogenetic effect have a limited
impact on CREA pattern recognition within felids. Our findings suggest that Machairodontinae
constitute one of the first well-supported exceptions to this biological rule currently known. We
hypothesize the acquisition of specific cranial features resulting from extreme ecological
specialisation - such as sabertoothed upper canines - to represent a preferential way to escape
from common evolutionary patterns of morphological variation such as CREA.
203
LONG BONE HISTOLOGY OF AETOSAURS AND PHYTOSAURS FROM
KRASIEJÓW (NORIAN) REVEALS STRONG ENVIRONMENTAL INFLUENCE
ON THE GROWTH PATTERN
Palaeohistology is a good source of information regarding the animals age, metabolism or mode of
life. The Norian Krasiejów locality offers an unique possibility to study multiple skeletal elements of
amphibians and reptiles. We tested the influence of external conditions on the growth pattern of
taxa originating from the same locality but belonging to different phylogenetic groups vs. the same
taxa but from geographically distinct localities. We studied the growth pattern of the phytosaur
Parasuchus cf. arenaceus and the aetosaur Stagonolepis olenkae femora and humeri. For
comparison, we sampled a phytosaur Nicrosaurus sp. femur from the Norian Heslach (S Germany)
locality. All samples show a lamellar-zonal bone, consisting mostly of low-organized parallel-fibred
tissue, and showing a low to moderate vascular density. Beside the Stagonolepis femora, all
bones are stratified with zones and annuli. None of the Krasiejów taxa preserve lines of arrested
growth (LAGs) and only the Nicrosaurus femur displays one LAG. Instead, the Krasiejów taxa
show the deposition of rest lines and sub-cycles throughout the cortex. A change in tissue from
fast-growing inner cortex to slower-growing outer cortex was previously described for numerous
phytosaurs and aetosaurs from North and South America, however, it was not observed in the taxa
studied herein. Parasuchus and Stagonolepis, despite their occupation of different habitats and not
being phylogenetically closely related, show a very similar growth pattern, which varies from their
close relatives originating outside Krasiejów. This implies, that the local environmental conditions
strongly influenced their growth pattern, and potentially overprinted their genetic precondition.
204
TROPHIC NETWORK RECONSTRUCTION AND ANALYSIS OF THE
DIFFERENT FOSSILIFEROUS LAYERS OF THE MIDDLE TRIASSIC MONTE
SAN GIORGIO WORLD HERITAGE (SWITZERLAND/ITALY)
2Department of Geology and Geophysics, Yale University, CT 06520 New Haven, United States of
America.
Since its discovery, the Middle Triassic Monte San Giorgio (MSG) lagerstätte near Meride (Canton
Ticino, Switzerland) has revealed an enormous quantity of exceptionally preserved fossils. Distinct
changes in faunal composition have been identified in the MSG lagerstätte, illustrating significant
ecological changes of this extinct ecosystem over a time-lapse of 3 Ma. However, there has never
been a detailed study on the network of trophic interactions among the organisms of the MSG
Faunas. Here we present the most complete database on MSG fossils with specimens from
museums in Zurich and Lugano in Switzerland and Milano in Italy. These initial results will be
expanded to reconstruct a succession of 8 trophic networks for 8 distinct fossil layers of the MSG
lagerstätte. We are using well-documented feeding behaviours and interactions of several
hundreds of different taxa collected and a new Rstudio package called PFIM (Paleo Food Web
Inference Model). The aim of this project is twofold. Firstly, our study will shed a new light on the
ecological changes of a Middle Triassic marine fauna set between two of the deadliest mass
extinction events in Earth history. Secondly, we will add a new element to the growing list of fossil
faunas studied for trophic network reconstruction. This will open new research opportunities for the
understanding of extinct ecosystem complexities and organizations and how these evolve through
deep time.
205
PALEOECOLOGICAL CHARACTERISTICS OF PLATYOPOSAURUS
(AMPHIBIA, TEMNOSPONDYLI) FROM THE MIDDLE PERMIAN OF EASTERN
EUROPE
A. Uliakhin1,2*
Archegosaurid temnospondyls of the genus Platyoposaurus were large (total reconstructed skull
length up to 0.7 m; body length about 3.5 m) gavial-like amphibians that were widely distributed in
the Middle Permian throughout today’s Eastern Europe. The genus Platyoposaurus includes four
described species (P. rickardi, P. stuckenbergi, P. watsoni, P. vjuschkovi), now confidently
identified from 18 localities. The paleoecological characteristics of Platyoposaurus were
identified and refined based on morphological, paleohistological, coprological, lithological facies,
and taphonomic analyses. Through this study it was possible to establish that Platioposaurus were
actively swimming and specialized piscivorous predators, with a more likely semi-aquatic type of
ecological adaptation. Platyoposaurus habitats included rivers, lakes and lagoons, in which respect
it resembled Archegosaurus, and may indicate a considerable environmental plasticity of members
of Archegosauridae generally. For comparison, the fossils of other archegosaurids have been
found in the following types of deposits: Archegosaurus in lacustrine and fluvial, Bashkirosaurus in
fluvial, Collidosuchus in fluvial, Prionosuchus in lacustrine. For Platyoposaurus a change of
paleobitopes in the process of individual development with relatively passive hydrodynamics (lakes
and lagoons) to rivers with more active hydrodynamics while maintaining their former habitats was
established, as evidenced by the discovered morphological transformations of skeleton elements.
Platyoposaurus actively competed for food resources with Melosauridae in those paleobiotopes in
which they could live together. Thus the use of different methods in the study showed the
effectiveness of complex approach for more accurate determination of paleoecological
characteristics and in particular the type of ecological adaptation.
206
PAPIONIN AFFINITIES IN THE ENDOCRANIAL ANATOMY OF
CERCOPITHECOIDES WILLIAMSI CALVARIA KA195
Cercopithecoids are relatively common in the fossil Plio-Pleistocene record of Africa and are found
in most South African hominin-bearing localities. Due to their usefulness in paleoenvironmental
reconstruction and biostratigraphy, many research efforts have been dedicated to them. However,
the taxonomic attribution of several specimens remains problematic. For example, the KA195
calvaria from Kromdraai was initially designated as the holotype of the papionin Parapapio
coronatus and subsequently assigned to the colobine Cercopithecoides williamsi. Subsequent
studies have reported similarities with either Cercopithecoides kimeui, a very large extinct colobine
from East Africa, or Theropithecus. To investigate the taxonomic affinities of KA195, we analyzed
its endocast and inner ear morphology, both known as good proxies for phylogenetic relatedness.
Then, we compared this specimen to a sample of Plio-Pleistocene and extant cercopithecoids. In
addition to cranial capacity and sulcal pattern assessment, endocast and bony labyrinth
morphology were inspected using a deformation-based 3DGM approach, allowing direct
207
comparison between continuous surfaces. Results show that the morphoarchitecture of the
endocast, and the shape of the semicircular canals contrast with those of fossil colobines and
highlight similarities with papionins, especially Theropithecus. Yet, despite the similarities, a
reassignment of KA195 to Theropithecus would be premature without an inspection of the
endocranial morphology of C. kimeui, currently unavailable. Additional 3DGM analyses, including
the external cranial morphology, are required and should assess if the morphological affinities with
Therepithecus are be homoplastic, potentially related to the increased terrestriality inferred for
fossil colobines and/or to their large body mass.
208
SKELETAL STRESS IN 3D SIMULATIONS OF HIGH-SPEED LOCOMOTION IN
LARGE THEROPODS
P.A. van Bijlert1,2,3*, K.T. Bates4, A.J.K. van Soest3, A.S. Schulp1,2
Non-avian dinosaurs experienced uniquely high skeletal loads due to their large body sizes.
Megatheropods (>1000 kg) may have been particularly limited by stress, because their bodies
were supported by two legs. Multibody dynamic physics simulations represent an ideal method to
test this phenomenon. However, previous studies have simulated maximal speed of
Tyrannosaurus rex while limiting movements to the midsagittal plane. This constrains a 3D model
to move in 2D, which can be physically interpreted as walking between two walls – a necessary
simplification due to computational costs. This earlier work suggested that speeds exceeding 5.5
m s-1 (at 7207 kg body mass) may have resulted in skeletal injuries.
We have built on this work, by combining it with a novel, computationally efficient optimization
method (direct collocation). This sophisticated workflow enables us to simulate full-3D periodic
movements of Tyrannosaurus. Our model has 23 degrees of freedom (versus 11 in previous
models), and weighs 8158 kg. Despite being nearly 1000 kg heavier, our 3D simulations display
substantially lower femoral peak-stresses than found in 2D. Furthermore, because our model
included the gastralia, knee penetration into the belly segment was only preventable in 3D. These
findings suggest that this added model complexity may have relevant biological interpretations:
megatheropods may have been capable of stress-avoidant locomotion at higher speeds. We are
currently developing a model of an emu (Dromaius novaehollandiae) to validate these new
209
predictions. Our results demonstrate the power of applying novel computational methods to the
fossil record.
210
VIRTUAL ENDOCASTS OF THE BRAIN AND INNER EAR OF LEPTICTIS
(LEPTICTIDA: MAMMALIA)
The genus Leptictis is an enigmatic small-sized insectivorous mammal from the middle Eocene to
late Oligocene of North America. This taxon has been challenging to place in the mammal tree,
being considered basal to Placentalia, or closely related to specific mammalian crown groups.
Endocranial data provide a wealth of phylogenetic, neurosensory, and behavioural information.
Published studies on the natural brain endocasts of Leptictis were limited to anatomical
descriptions, and the proportion of different brain regions could not be measured to inform us
about the behaviours and senses of this extinct mammal. For this study, we scanned and
segmented the brain and inner ear endocasts for two specimens of Leptictis from the AMNH
collections in the USA. Our results show that Leptictis had an EQ range of 0.43 – 0.69, similar to
contemporaneous rodents such as Paramys and Ischyromys. The olfactory bulbs represented
9.1% of the total endocast volume, which is in the range of Hyopsodus, indicating that Leptictis
likely relied heavily on olfaction to find its prey. The petrosal lobules were small, representing 1.1%
of the total endocast volume similar to the fossorial rodent Pseudotomus. The neocortical surface
area is relatively small compared to contemporaneous crown placental mammals (12.2% of the
total surface area), which is similar to the condition seen in the stem taxon Hyopsodus. Leptictis
was relatively agile, with an agility score of 4.04. A mosaic of plesiomorphic and derived traits
support the hypothesis that Leptictis was likely a stem placental mammal closely related to crown
groups.
211
FLY PUPARIA IN A MAMMOTH SKULL FROM THE NETHERLANDS:
IMPLICATIONS FOR MAMMOTH TAPHONOMY
1Department of Earth Sciences, Utrecht University, P.O. Box 80115, 3508 TC Utrecht, the
Netherlands.
2Department of Earth Sciences, Uppsala Universitet, Villavägen 16, 752 36, Uppsala, Sweden.
A fragmented mammoth skull of Weichselian age, recovered at a dredging site at Lent in the
Netherlands, has been found to contain fly puparia. The puparia belong to necrophagous flies,
which lay their eggs in carcasses. These fly puparia can inform on the palaeoenvironment and
offer insights into the preservation and taphonomy of fossil mammoth material from Lent. The
uncovered skull contained puparia from a single blowfly species: Protophormia terraenovae. The
presence of a single species allowed for a tentative estimation of the season of death, namely
spring. This is because in warmer seasons the presence of more blowfly species is common. The
minimum temperature for colonisation of a carcass by P. terraenovae is 9 °C. This would rule out
winter for colonisation, but not necessarily for the death of the mammoth, as colonisation could
occur after rapid thawing. The colonisation of the carcass by blowflies not only informs on the
taphonomy of the mammoth, the presence of blowfly puparia in the mammoth remains also further
constrains the age estimate of this individual mammoth fossil. Temperature reconstructions of
Weichselian summers do not frequently show excursions above 9 °C, which only occur in the
beginning of the Weichselian or during interstadials. The mammoth thus must have lived, and
more importantly died, during one of the somewhat warmer periods of the Weichselian.
212
NETWORK MODELING AND CAPTURE-MARK-RECAPTURE METHODS FOR
THE IDENTIFICATION OF SPECIES FACTORIES
1Department of Computer Science, University of Helsinki, Pietari Kalmin katu 5, 00560 Helsinki,
Finland
Species factories are defined as times and places in the fossil record where and when an
exceptionally large number of new species occur. While several tailored solutions for the
mammalian record have been proposed, how to identify species factories computationally in a
standardized way is still an open question. To quantify what is exceptional, we first need to
quantify what is regular. One of the main challenges in this identification process is to account for
sampling unevenness, which depends on several methodological decisions, including the scale of
the analysis (aggregation radius).
Here we use Capture-Mark-Recapture methods (CMR) with spatial aggregation guided by network
modelling, to estimate the sampling probabilities for the species in the NOW database. Since the
mammalian record is sparse and most localities include only a few species, we couple CMR with
tailored spatial aggregation approaches to estimate the sampling probabilities. We then use these
sampling probabilities to quantify background speciation rates and assess what rates are
abnormal. We represented aggregated fossil data as a bipartite network and used community
detection to evaluate how the choice of an aggregation radius impacts the modular structure.
While estimating sampling probabilities allows the adjustment for sampling unevenness so that the
difference in findings can be compared across locations and cannot be due to differences in
sampling. We have identified as species factories the locations with origination rate in the highest
5% per time unit. To characterize those locations we looked for paleoecological patterns in these
places that may be lacking elsewhere.
213
THE LATE CAMPANIAN MICROVERTEBRATE ASSEMBLAGE FROM
PETREȘTI (TRANSYLVANIAN BASIN, ROMANIA), A NEW WINDOW INTO
THE EVOLUTION OF EUROPEAN CRETACEOUS INSULAR FAUNAS
1Faculty of Geology and Geophysics, University of Bucharest, 1 Nicolae Bălcescu Avenue, 010041
Bucharest, Romania.
2Center for Risk Studies, Space Modelling and Dynamics of Terrestrial and Coastal Systems,
University of Bucharest, 1 Nicolae Bălcescu Avenue, 010041 Bucharest, Romania.
3Department of Natural Sciences, Transylvanian Museum Society, 2-4 Napoca Street, 400009
Cluj-Napoca, Romania.
4Department of Geology, “Al. I. Cuza” University of Iași, 20A Carol I Blv., 700505 Iași, Romania.
The latest Cretaceous continental vertebrate faunas of Romania are well known for their unique
features, derived from prolonged evolution in isolation on an insular landmass, the so-called
`Hațeg Island`. The discovery of new vertebrate assemblages, of Campanian–Maastrichtian age,
at Petrești, in the SW Transylvanian Basin, offers the opportunity to take a look at the composition
of insular vertebrate faunas of the region at an older age than other better known records of the
Hațeg Basin. On-going research on the palynomorph, calcareous nannoplankton, and foraminifera
assemblages of the Petrești section more confidently support a late Campanian age (Upper
Cretaceous) for the fossiliferous bonebed that yielded the largest part of the vertebrate remains.
Continued screen-washing of the bonebed sediment from the grey deposits from the base of the
Sebeș Formation, overlying the marine-transitional Bozeș Formation, have yielded numerous
microvertebrate remains, belonging to: lepisosteid fish; albanerpetontid and anuran amphibians;
crocodyliforms (teeth of various morphologies); rhabdodontid, ankylosaurid, and theropod
214
dinosaurs; and multituberculate mammals. This contribution presents the new microvertebrate
material found at Petrești, adding up to the previous, shorter taxonomic list, and compares the new
additions to the later, Maastrichtian faunas of Transylvania, aiming to better understand their
evolution within their insular environment.
Work supported by the Romanian Ministry of Research, Innovation and Digitalization, CNCS–
UEFISCDI grant PN-III-P4-ID-PCE-2020-2570, within PNCDI III.
215
A NEW VERTEBRATE ASSEMBLAGE FROM THE LATE PLIOCENE OF
MĂRU (DACIAN BASIN, SOUTHERN ROMANIA)
2“Emil Racoviță” Institute of Speleology, Romanian Academy, 13-15 Calea 13 Septembrie, 050711
Bucharest, Romania.
7Museu Valencià d’Història Natural, L’Hort de Feliu, BOX 8460, 46018 Alginet, Spain.
Pliocene vertebrate assemblages from the Dacian Basin (southern Romania) are poorly known,
and generally represented by isolated large mammal remains. So far, non-mammalian material
was reported from Podari (freshwater fishes), Berești, and Mălușteni. The latter two localities
yielded reptile and fish remains collected and only preliminarily described the 1930s.
216
Recent investigations in the north-western Dacian Basin led to the discovery of new outcrops in
Măru (Gorj County). Mollusc-bearing mudstones were screen-washed, producing rare rodent
molars, and numerous teeth and postcranial bones of freshwater fish. The rodent material is
tentatively assigned to the arvicolid Mimomys hajnackensis, indicative for biozone MN16a (early
late Pliocene). The fish specimens are assigned to Leuciscus sp., Rutilus sp., Scardinius sp.,
Chondrostoma sp., Abramis sp., Barbus sp., Carassius sp., Tinca sp., another indeterminate
cyprinid, Silurus sp., Esox sp., and an indeterminate percid. The taxonomic composition of the fish
community indicates a lacustrine environment. The fossil assemblage from Măru adds important
information on the composition of late Pliocene freshwater fish assemblages of the Dacian Basin,
previously known only from Podari (Tinca sp., Scardinius sp., Esox sp., Silurus sp.), and provides
new data that will be useful in further assessment of connections to neighbouring basins
comprising similar assemblages.
Work supported by: CNCS – UEFISCDI grants PN-III-P4-ID-PCE-2020-2282 (Ș.V.), and PN-III-P1-
1.1-TE-2021-0664 (Ș.V.), within PNCDI III; grant 0121U110402 (O.K.); requalification of the
Spanish university system from the Ministry of Universities of the Government of Spain, financed
by the European Union, Next Generation EU, grant ZA21-044 (V.D.C.).
217
SHAKE A LEG: VARIABILITY ON PELVIC MUSCULATURE OF THE
TITANOSAURS FROM LO HUECO (LATE CRETACEOUS, SPAIN)
2 Department of Organismal Biology, University of Chicago, 1027 E 57th St, Chicago, Illinois
60637, United States of America.
3Instituto Dom Luiz, Faculdade de Ciência da Universidade de Lisboa, Campo Grande, 1749-016
Lisbon, Portugal.
4Dinosaur Institute, Natural History Museum of Los Angeles, 900 Exposition Blvd, 90007 Los
Angeles, California, United States of America.
5Facultad de Bellas Artes, Universidad Complutense de Madrid, Calle Pintor el Greco 2, 28040,
Madrid, Spain.
Titanosaurs from Lo Hueco fossil-site are among the better preserved and articulated specimens
of the Late Cretaceous of Europe. Preliminary analyses on these titanosaurian specimens have
revealed a high degree of disparity in the caudal axial skeleton, with up to four different
morphotypes in the assemblage. Most of these partial skeletons preserve articulated or closely
associated caudal series and pelvic girdles, making them prime specimens to evaluate their
locomotor capabilities.
Several pelvic girdles and associated femora, tibiae and fibulae were scanned using
stereophotogrammetry. These elements were virtually assembled in osteological neutral pose. We
used the extant phylogenetic bracket to map the attachments of pelvic musculature in the girdle,
limb (when preserved) and their corresponding caudal series of each of the four putative
titanosaurian morphotypes. Then, we compared the proportions and development of the different
218
osteological correlates, as well as disposition of muscle lines of action, to assess potential
locomotor capabilities in each morphotype.
Main differences among the morphotypes were found in the proportions and orientation of the ilium
preacetabular lobe (origin sites for M. iliotibialis 1. M. iliotrochantericus caudalis and M.
iliofemoralis externus). These differences imply changes in the lines action of these muscles and
their volumes. This showcases different capabilities in femoral protraction, abduction and hip
flexion that could be related with minor variations in pelvic width and femoral morphology among
the different morphotypes. This suggests that the four putative titanosaurian morphotypes had
different ecomorphologies and might have occupied different niches.
219
QUANTIFYING BONE SURFACE ROUGHNESS IN BRANTA CANADENSIS
USING 3D DIGITAL MICROSCOPY: IMPLICATIONS FOR TEXTURAL AGEING
OF EXTANT AND EXTINCT SPECIES
Keywords: Textural ageing, surface texture, digital microscopy, skeletal ontogeny, histology
Ontogenetic ageing is applied in the fields of taxonomy, morphology, ecology, archaeozoology and
palaeontology and plays a crucial role when studying the osteology and ontogeny of species.
Textural analysis is a non-destructive ontogenetic ageing method that uses bone surface textures
to determine the relative age of skeletal remains. Variations in bone surface textures between
individuals result from a decrease in penetrating blood vessels through the bone surface during the
growth of an animal. Textural ageing patterns have been observed independently in numerous
species (e.g., Branta canadensis, Ardea cinerea, Pygoscelis antarctica and centrosaurine
dinosaurs), but previous research has exclusively investigated textural ageing patterns through
qualitative, subjective, and unsystematic visual inspection.
Here, we apply 3D digital microscopy (Keyence VHX-6000) to quantify bone surface roughness
and investigate textural ageing patterns conforming the roughness parameters ISO-25178 (3D
areal surface texture) and ISO-4287 (2D line roughness). Measuring the skeletal remains of extant
bird taxa was prioritised over the skeletal remains of fossil taxa, due to potential interference from
taphonomic signals. Therefore, Branta canadensis longbones were analysed. 2D line roughness
analyses on Branta canadensis long bones were able to retrieve numerical surface roughness
parameters conforming to the bone texture patterns observed during visual inspection. Similarly,
Triceratops fossil remains of varying states of preservation returned comparable parameters.
Future research will focus on osteohistological analyses to link the observed patterns in textural
ageing and aims to better constrain taphonomic effects in the analysis of fossil material.
220
GLOBAL PHYLOGENY OF DIPLOCYNODONTINAE AND ITS IMPLICATIONS
FOR CROCODYLIAN EVOLUTION
J. D. Walter1,4*, J. E. Martin2, M. Delfino1,3, M. Rabi4,5
1 Dipartimento di Scienze della Terra, Università di Torino, Via Valperga Caluso 35, I-10125
Torino, Italy.
2 Centre National de la Recherche Scientifique UMR 5276, LGLTPE, Université de Lyon, ENS de
Lyon, Université Lyon 1, 46 Allée d’Italie, F-69342 Lyon Cedex 07, France.
Diplocynodontinae represents the only crocodylian lineage that survived the Eocene-Oligocene
transition in Europe and is known for its long endemic evolution in the continent. Current
phylogenies place Diplocynodontinae as an early diverging member of Alligatoroidea, a group that
otherwise originated in North America during the Late Cretaceous. This phylogenetic position is
incongruent with the spatiotemporal distribution of fossils and leaves the origin of
Diplocynodontinae enigmatic. In this study, we critically reassess the evolutionary relationships of
Diplocynodontinae through first-hand revision of key taxa and homologies placing, according to
previous phylogenies, the group into Alligatoroidea. We find that several synapomorphies
supporting the alligatoroid position represent irreproducible or poorly defined character states.
Parsimony analyses of a revised dataset results in several most parsimonious trees with an
alternative position for Diplocynodontinae, along the stem lineage of crown-group crocodylians.
The traditional basal alligatoroid position is persisting in the remaining trees but the
synapomorphies supporting this placement are poorly known for outgroup taxa. A stem-
crocodylian position for Diplocynodontinae is more congruent with stratigraphy and European
vertebrate paleobiogeographical patterns, and would clarify the origin of the clade as a result of an
221
early Paleogene dispersal from North America. Furthermore, phylogenetic patterns of extinction
across the Eocene-Oligocene transition likely need to be reconsidered together with our current
hypotheses of early alligatoroid morphology.
222
THE DINOSAUR ICHNOLOGICAL RECORD OF NORTHERN CHILE: A
REVIEW AND ITS POTENTIAL DEVELOPMENT
6Dipartimento di Scienze della Terra, Università degli Studi di Firenze, 50121 Firenze, Italy.
Chile's ichnological record has great potential but, apart from a few mentions in the literature, its
study has been neglected. Rich track bearing levels, between Late Jurassic (Oxfordian) to Early
Cretaceous in age, are known especially, but not only, from the Tarapacá to Atacama regions in
northern Chile. These track-bearing sequences were deposited in various depositional
environments (e.g., littoral, fluvial, etc.) providing an excellent base for detailed studies of the
relationship between ichnofauna and their palaeoenvironment. Fossil tracksites on the western
margin of Gondwana at these palaeolatitues have not been fully investigated. However, they
represent a unique record and the finding new tracksites will increase our knowledge on dinosaur
diversity correlated to the different palaeoenvironments.
In addition to the many known tracksites (e.g. Quebrada Chacarilla, Quebrada Arca, Quebrada La
Descubridora), especially in the Tarapacá, Antofagasta and Atacama regions, the potential for new
relevant discoveries is very high. This is because of several not-yet-validated reports of dinosaur
footprints from those regions and a large number of potential outcrops in barely explored areas, no
doubt vastly outnumbering known sites. This potential is going to be exploited in the near future
through a Ph.D. project. Stay tuned for more new and exciting news.
223