Josep Fortuny

• PhD
• Group Leader at Institut Català de Paleontologia Miquel Crusafont

Meet your gibbon cousin Apes are divided into two groups: larger-bodied apes, or hominoids, such as humans, chimps, and gorillas; and smaller-bodied hylobatids, such as gibbons. These two lineages are thought to have diverged rather cleanly, sharing few similarities after the emergence of crown hominoids. Alba et al. describe a new ape from the Mio...

Macroevolutionary, palaeoecological and biomechanical analyses in deep time offer the possibility to decipher the structural constraints, ecomorphological patterns and evolutionary history of extinct groups. Here, 3D comparative biomechanical analyses of the extinct giant early amphibian group of stereospondyls together with living lissamphibians a...

Understanding cranial sutures and how they relieve and dissipate stress is essential to assess their role in cranial biomechanics and to develop highly accurate predictive models. This involves examining how ontogeny affects cranial sutures, as well as their morphology and function, and how these changes through time may impact essential biomechani...

Therapsids were a dominant component of middle–late Permian terrestrial ecosystems worldwide, eventually giving rise to mammals during the early Mesozoic. However, little is currently known about the time and place of origin of Therapsida. Here we describe a definitive therapsid from the lower–?middle Permian palaeotropics, a partial skeleton of a...

The Early to Middle Triassic is a key time interval in tetrapod evolution. After the end-Permian biotic crisis, harsh environmental conditions due to global warming and aridity persisted during the Early Triassic. This led to an impoverished biodiversity, especially in equatorial Pangea, until the Middle Triassic, when vertebrate ecosystems re-flou...

Inner skull cavities provide key characters to elucidate taxonomic, phylogenetic, and palaeobiological inferences in reptiles. Herein, an integral description of the extinct alligatoroid Diplocynodon tormis is presented based on its holotype, recovered from the Middle Eocene site of “Teso de la Flecha” (Salamanca, Spain). It is an almost complete s...

The phylogenetic relationships of the small-bodied catarrhine Pliobates cataloniae (~11.6 Ma, NE Iberian Peninsula) have been controversial since its original description. However, the recent report of additional dentognathic remains has supported its crouzeliid pliopithecoid status. Based on the available hypodigm, the molar enameledentine junctio...

La Mora is an early−to−middle Anisian (early Middle Triassic) locality in the Catalan Basin, northeastern Spain. Since its discovery by amateur archaeologists looking for medieval pottery shards in 1989, the site has been excavated during four field seasons: 1990, 2008, 2010, and 2023. Stratigraphically, La Mora is situated within the upper Bundsan...

La Mora, an early−to−middle Anisian (early Middle Triassic) site in the Catalan Basin, northeastern Spain, has yielded an abundance of tetrapod fossils and is one of the most important localities from this time period on the Iberian Peninsula. The site was discovered in 1989 and has been excavated during four field seasons: 1990, 2008, 2010, and 20...

The systematic status of the small-bodied catarrhine primate Pliobates cataloniae, from the Miocene (11.6 Ma) of Spain, is controversial because it displays a mosaic of primitive and derived features compared with extant hominoids (apes and humans). Cladistic analyses have recovered Pliobates as either a stem hominoid or as a pliopithecoid stem cat...

Pierolapithecus catalaunicus (~12 million years ago, northeastern Spain) is key to understanding the mosaic nature of hominid (great ape and human) evolution. Notably, its skeleton indicates that an orthograde (upright) body plan preceded suspensory adaptations in hominid evolution. However, there is ongoing debate about this species, partly becaus...

Tetrapod diversity in Permian terrestrial ecosystems of southwestern Europe is poorly recorded by bone specimens, but it is better represented by an important tetrapod ichnological record that is relevant to our understanding of vertebrate communities in the equatorial Pangaea. Herein, two tetrapod ichnoassociations from three new ichnosites, withi...

Dipnoi (lungfish) are one of the three extant sarcopterygian clades and one of the closest to tetrapodomorphs. They became highly diverse during the Devonian, but this diversity subsequently reduced drastically, with only a few taxa known thus far from upper Palaeozoic rocks. Here we report the first discovery of lungfish in terrestrial red-beds (“...

Some members of Lepospondyli, such as Diplocaulus magnicornis, are renowned for their distinctive boomerang-shaped skulls. Computational fluid dynamics (CFD) simulations were conducted to analyze their hydrodynamic performance under various environmental conditions.

Moradisaurine captorhinid eureptiles are one of the best‐known groups of Permian herbivorous tetrapods. Moradisaurines, with several rows of teeth on the maxillae and jaws, first appeared in the Cisuralian and went extinct at the end of the Lopingian; they were especially abundant in the equatorial latitudes of Pangaea. However, the postcranial ske...

A vast diversity of catarrhines primates has been uncovered in the Middle to Late Miocene (12.5-9.6 Ma) of the Vallès-Penedès Basin (northeastern Spain), including several hominid species (Pierolapithecus catalaunicus, Anoiapithecus brevirostris, Dryopithecus fontani, Hispanopithecus laietanus, and Hispanopithecus crusafonti) plus some remains attr...

In the Middle Triassic (Anisian) outcrops of the Montseny area located in the Catalan Coastal Ranges, northeast Iberian Peninsula, several osteological remains providing crucial information of the palaeobiodiversity in equatorial Pangaea have been recovered in the last two decades. The faunal assemblage is consistent with the record from higher lat...

Southwest Asia is at the epicenter of zooarchaeological research on pivotal changes in human history such as animal domestication and the emergence of social complexity. This volume continues the long tradition of the ASWA conference series in publishing new research results in the zooarchaeology of southwest Asia and adjacent areas. The book is or...

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 Per...

An iconic, insular, endemic genus of lizards (Gallotia) is present on the Canary Islands (Spain), comprising gigantic to smaller-sized species. Despite numerous studies on various biological aspects of this genus, the osteological knowledge available is scarce. This makes it difficult to identify to species level the bone remains recovered from bot...

Pliobates cataloniae is a small-bodied catarrhine (~4–5 kg) from the Miocene of NE Iberian Peninsula. Originally this species was recovered by a cladistic analysis as a stem hominoid showing a mixture of primitive (stem catarrhine-like) and derived (modern hominoid-like) craniodental and postcranial features (Alba et al., 2015 ). Later on, this tax...

Tooth crown morphology plays a critical role in primate systematics, notably to make taxonomic assessments and to reconstruct the evolutionary history of hominids and hominins in particular. Compared with the outer enamel surface, which can be affected by wear and various taphonomic processes, the enamel-dentine junction (EDJ) is generally better p...

The Cisuralian–Guadalupian (early–middle Permian) was a period of climate transition between the Carboniferous icehouse conditions to the latest Permian–Early Triassic hothouse. The landmasses had coalesced in the supercontinent Pangaea and the climate was progressively becoming more arid, especially in a belt over the palaeoequator. The deposits o...

Bioerosions produced by the osteophagous diet of animals that fed on dinosaur bones are very scarce in the European fossil record. Herein we present bioerosion on hadrosaurid remains from the Maastrichtian Tremp Formation of the Pyrenean Basin, which is only the second such case recorded from the Iberian-Occitan Plate besides a sauropod from the Ju...

PERMIAN AND TRIASSIC CONTINENTAL PALAEOECOSYSTEMS OF MALLORCA (BALEARIC ISLANDS, WESTERN MEDITERRANEAN): SYNTHESIS AND FUTURE PERSPECTIVES. The present paper offers an exhaustive review of all the previous works on the continental Permian and Triassic of Mallorca, synthesising all the relevant data regarding their geology, palaeontology and age att...

The 1-m-tall dwarf elephant Palaeoloxodon falconeri from the Pleistocene of Sicily (Italy) is an extreme example of insular dwarfism and epitomizes the Island Rule. Based on scaling of life-history (LH) traits with body mass, P. falconeri is widely considered to be ‘r-selected’ by truncation of the growth period, associated with an early onset of r...

The Ladinian-Carnian transition in the Tethys domain was accompanied by an important environmental change representing a milestone in the climate evolution of the Triassic. However, estimations on paleodiversity composition and paleoenvironmental conditions across this interval are scarce in marine settings due to the lack of fossil-bearing succes...

During the Early–Middle Triassic, the biosphere was recovering from the most severe mass extinction event of multicellular life, in the Permian–Triassic transition. Continental basins corresponding to present-day Mallorca (Balearic Islands, western Mediterranean) were located in the equatorial region of the supercontinent Pangaea, in the western pe...

Moradisaurine captorhinid eureptiles were a successful group of high-fibre herbivores that lived in the arid low latitudes of Pangaea during the Permian. Here we describe a palaeoassemblage from the Permian of Menorca (Balearic Islands, western Mediterranean), consisting of ichnites of small captorhinomorph eureptiles, probably moradisaurines (Hylo...

The European Buntsandstein facies, of Early to Middle Triassic age, record a key period in tetrapod evolution. After the end-Permian biotic crisis, harsh conditions of global warming and aridity persisted during the Early Triassic, which induced a decrease in biodiversity, especially in equatorial Pangaea. The full recovery of vertebrate ecosystems...

Moradisaurine captorhinid eureptiles were a cosmopolitan and successful group of high-fibre herbivores that lived in the arid low latitudes of Pangaea during the Permian. They are known to be quite diverse in southern North America, with lower-middle Permian fossil deposits that have been intensely sampled over the years. In other areas, such as th...

“Pelycosaur”-grade synapsids were a successful group of terrestrial tetrapods that lived during the Carboniferous and Permian, utilising a wide diversity of ecological niches. They are considered the trackmakers of the ichnogenera Dimetropus and possibly also Dromopus and Tambachichnium, found in upper Palaeozoic deposits of North America, North Af...

Extensive fieldwork at Abocador de Can Mata (north-east Iberian Peninsula) has uncovered a previously unsuspected diversity of catarrhine primates in the middle Miocene (12.5–11.6 Ma) of Europe. However, the distinction of the great ape genera Pierolapithecus and Anoiapithecus from Dryopithecus (supported by craniodental differences) has been dispu...

Pareiasauromorpha is one of the most important tetrapod groups of the Permian. Skeletal evidence suggests a late Kungurian origin in North America, whereas the majority of occurrences come from the Guadalupian and Lopingian of South Africa and Russia. However, Pareiasauromorpha footprints include the ichnogenus Pachypes, which is unknown from strat...

Tetrapod ichnology is a powerful tool to reconstruct the faunal composition of Middle Triassic ecosystems. However, reconstructions based on a single palaeoenvironment provide an incomplete and impoverished picture of the actual palaeodiversity. In this paper, we analyse Middle Triassic tetrapod ichnoassociations from the detrital Muschelkalk facie...

In the last decade, the first ichnoassamblages from the middle Muschelkalk facies (upper Anisian–middle Ladinian) of the Catalan Basin (NE Iberian Peninsula) have been discovered. Herein, the frst xiphosuran trace fossils are described from the locality of Penya Rubí, a newly discovered ichnosite from the Catalan Basin. The fnding opens a window...

Editorial on the Research Topic Evolving Virtual and Computational Paleontology During the last few decades, the development of new technologies and methods, and increases in computational power, are widening Paleontology's research frontiers, moving this discipline-situated halfway between Biology and Geology-toward increasing use of (paleo) bio...

During the Triassic, many groups of predatory marine reptiles appeared and diversified. On Mallorca, fossils of marine reptiles have been found in Muschelkalk (Anisian–Ladinian) and Keuper (Carnian–Norian) facies. Here we describe an anterior caudal vertebra of a basal ichthyosauriform similar to Grippidia (Reptilia: Ichthyosauriformes) from the La...

Quercus. 415. Septiembre 2020. 54-59. La paleopatología estudia las enfermedades que afectaron a los animales de épocas pasadas. Su registro es muy difícil de detectar, pues tienen que haber dejado señales en las estructuras que fosilizan posteriormente, así que está limitada a un reducido número de ejemplares. Sin embargo, el creciente interés por...

In recent years photogrammetry has become an essential tool in the study of tetrapod footprints. Morphological analyses of footprints are interpretative; thus, researchers should use as much information as possible in order to eventually provide an objective conclusion. In this regard, photogrammetry is an extremely helpful tool to avoid potential...

The Canary Islands are an Atlantic archipelago known for its high number of endemic species. Among the most known endemic vertebrate species are the giant lizards of the genus Gallotia. We describe the cranial osteology of the first almost complete and articulated fossil skull of the taxon Gallotia auaritae, recovered from the lower-middle Pleistoc...

With the aim of characterising the different stress patterns in several egg types and evaluating the resistivity of the eggshell while impacting in free fall, experimental tests and computational simulations have been performed on eggs of the species Gallus gallus (domestic hen), Struthio camelus (ostrich), and Testudo sp. (tortoise). The different...

Cranial sutures connect adjacent bones of the skull and play an important role in the absorption of stresses that may occur during different activities. The Late Triassic temnospondyl amphibian Metoposaurus krasiejowensis has been extensively studied over the years in terms of skull biomechanics, but without a detailed description of the function o...

First alligatoroid Diplocynodon remains from the Ahníkov/Merkur Mine site (A/MM) are described here. A/MM is the richest fossil site of the Most Basin (Eger Graben, Czech Republic), including more than 150 taxa of fossil vertebrates reported to date. Specifically, more than 300 remains of fossil crocodilians have been collected, but they have never...

The early Miocene deposits of Bohemia (Czech Republic) contain numerous fragmentary crocodylian remains. Despite this abundance, a detailed taxonomical assignment of these remains was impossible due to the absence of diagnostic cranial elements. Here, we report two partially preserved skulls together with some osteoderms and a partially preserved v...

A new actinopterygian, Moradebrichthys vilasecae gen. et sp. nov., from the Middle Triassic (Late Ladinian) of Móra d'Ebre-Camposines (Catalonia, NE Iberian Peninsula) is erected on the basis of several, almost complete, articulated and well-preserved specimens. It is referred to the controversial family Perleididae based on the presence of a combi...

There is considerable debate regarding whether mandibular morphology in ungulates primarily reflects phylogenetic affinities or adaptation to specific diet. In an effort to help resolve this debate, we use three-dimensional finite element analysis (FEA) to assess the biomechanical performance of mandibles in eleven ungulate taxa with well-establish...

Percent of the variance of the PC scores. For the six biting cases (DOCX)

Plots displaying the first two PCs of the different PCAs for N = 10, 25, 50, 75 and 100 for case 2. Lateral biting in the second molar. The species are coloured by order: blue: Perissodactyla and brown Cetartiodactyla. The axes of each pair of PCs are in the same scale. (TIF)

FEA results for the lateral biting cases. Number of mandible mesh elements and statistics: Arithmetic Mean (AM), Mesh-Weighted Arithmetic Mean (MWAM), Percentage Error of the Arithmetic Mean (PEofAM), Median (M), Mesh-Weighted Median (MWM), Percentage Error of the Median (PEofM) and the value quartiles (M25, M50, M75 and M95) according to Marcé-Nog...

Convergence of the R2 values of the PC scores. Each value is the R2 for a different pair of PCAs of the correlation matrix. Each PC was correlated with the equivalent PC of the PCA developed using a larger number of intervals. (DOCX)

Intervals’ method data of Von Mises stress when N = 100 M for the six biting cases. (XLSX)

Plots displaying the first two PCs of the different PCAs for N = 10, 25, 50, 75 and 100 for case 1. Lateral biting in the first molar. The species are coloured by order: blue: Perissodactyla and brown Cetartiodactyla. The axes of each pair of PCs are in the same scale. (TIF)

FEA results for the orthal biting cases. Number of mandible mesh elements and statistics: Arithmetic Mean (AM), Mesh-Weighted Arithmetic Mean (MWAM), Percentage Error of the Arithmetic Mean (PEofAM), Median (M), Mesh-Weighted Median (MWM), Percentage Error of the Median (PEofM) and the value quartiles (M25, M50, M75 and M95) according to Marcé-Nogu...

Plots displaying the first two PCs of the different PCAs for N = 10, 25, 50, 75 and 100 for case 3. Lateral biting in the third molar. The species are coloured by order: blue: Perissodactyla and brown Cetartiodactyla. The axes of each pair of PCs are in the same scale. (TIF)

Plots displaying the first two PCs of the different PCAs for N = 10, 25, 50, 75 and 100 for case 5. Orthal biting in the second molar. The species are coloured by order: blue: Perissodactyla and brown Cetartiodactyla. The axes of each pair of PCs are in the same scale. (TIF)

Plots displaying the first two PCs of the different PCAs for N = 10, 25, 50, 75 and 100 for case 6. Orthal biting in the third molar. The species are coloured by order: blue: Perissodactyla and brown Cetartiodactyla. The axes of each pair of PCs are in the same scale. (TIF)

Plots displaying the first two PCs of the different PCAs for N = 10, 25, 50, 75 and 100 for case 4. Orthal biting in the first molar. The species are coloured by order: blue: Perissodactyla and brown Cetartiodactyla. The axes of each pair of PCs are in the same scale. (TIF)

Temnospondyls from the Middle–Late Triassic of Madagascar are problematic and scarce: ‘Metoposaurus hoffmani’ was erected on the basis of poor material, and this taxon has never been revised. Other remains were also reported and assigned to Temnospondyli indet., but they have never been described, nor figured. Here, we (re)describe in detail this h...

The pterosaur record from the Iberian Peninsula is mostly scarce and undefined, but in the last few years some new taxa have been described from different Lower Cretaceous sites of Spain. Here we describe a new genus and species of toothed pterodactyloid pterosaur from the Barremian of the Iberian Peninsula, Iberodactylus andreui gen. et sp. nov.,...

Ronchi et al. (in press) comment on the stratigraphic, sedimentological, mineralogical and palaeontological analyses performed in the recently described Coll de Terrers Permian–Triassic terrestrial succession from the Catalan Pyrenees by Mujal et al. (2017a). The comment debates our interpretation of a succession of red-beds as Permian (Upper Red U...

Worm lizards, or amphisbaenians, of the genus Blanus are found in various countries around the Mediterranean Sea. In addition to four extinct species, seven extant taxa are currently recognized. Here, we present the first comparative analysis of the cranial osteology of Blanus including all extant species. The results of this analysis show a homoge...

Flying squirrels are the only group of gliding mammals with a remarkable diversity and wide geographical range. However, their evolutionary story is not well known. Thus far, identification of extinct flying squirrels has been exclusively based on dental features, which, contrary to certain postcranial characters, are not unique to them. Therefore,...

The morphology of the mandibular sutures in the Late Triassic temnospondyl Metoposaurus krasiejowensis has been examined in order to determine their role in mandible biomechanics. Until now, no histological studies of mandibular sutures in extinct vertebrates were performed, in contrast to cranial sutures. As a consequence, mandibular suture interp...

Tetrapod ichnites are essential to untangle trackmakers locomotion and behaviour, as they are in situ and in vivo evidences of the trackmaker. They also provide information on the palaeoenvironmental affinities of the trackmakers. All these features can be studied in detail in the Early–Middle Triassic Buntsandstein fluvial successions from the Cat...

Locomotionis one of the basic needs of all animals. Nature has created a variety of locomotion mechanisms to adapt to different environments, to increase chances of survival in competition with other animalsliving in the same area, and to allow exploring otherarea in search for food and shelter[1, 2]. This paper focuses on savannah monitor (Varanus...

Background Trematosaurines are a widespread group of early tetrapods (Temnospondyli, Stereospondyli) known from all continents except South America and Antarctica. They radiated rapidly during the Early Triassic just after the End Permian mass extinction and are of interest to understand the recovery of the ecosystems just after extinction. Tremato...

Objectives: High-resolution imaging of fossils with X-ray computed microtomography (lCT) has become a very powerful tool in paleontological research. However, fossilized bone, embedding matrix, and dental tissues do not always provide a distinct structural signal with X-rays. We demonstrate the benefits of high-resolution neutron radiation in three...

Purpose Xiphosurids (horseshoe crabs) are aquatic chelicerate arthropods commonly related to marine environments. Although today only four marine species exist, in the geological record they were much more diverse (especially during Carboniferous and Triassic periods), and even inhabited non-marine settings. Here we analyze an exceptional xiphosuri...

Finite Element Analysis (FEA) is a useful method for understanding form and function. However, modelling of fossil taxa invariably involves assumptions as a result of preservation-induced loss of information in the fossil record. To test the validity of predictions from FEA, given such assumptions, these results could be compared to independent lin...

This paper discusses the taxonomic affinities of three isolated teeth discovered in the Buntsandstein facies of the Catalan Coastal Ranges and central-eastern Pyrenean basins that crop out in Catalonia, northeastern Spain. The tooth crowns are blade-like, labiolingually compressed, distally recurved, and proportionally apicobasally tall, in which t...

Developmental changes in salamander skulls, before and after metamorphosis, affect the feeding capabilities of these animals. How changes in cranial morphology and tissue properties affect the function of the skull are key to decipher the early evolutionary history of the crown-group of salamanders. Here, 3D cranial biomechanics of the adult Salama...

Figure S1. Plant fossil distribution in Coll de Sas locality. Note the high lateral thickness variation of the succession. See details of the plant taxonomic groups for each level (including genera and species) in Text S1.

Figure S2. (A-F) Representative in situ orthogonal demagnetization diagrams from the studied Coll de Sas (CS) and les Esglésies (ES) sections. Samples follow a stepwise alternating field (AF) demagnetization protocol after a single heating step to 150ºC. The natural remanent magnetization (NRM) intensity and some demagnetization steps are indicated...

Figure S3. Stereographic projections of the L (A) and ChRM (B) computed components before (in situ) and after bedding correction (tilt corrected) for Les Esglésies section. Open and closed symbols indicate projections onto the upper and lower hemisphere respectively. Mean direction and statistics are given. N = number of samples; Dec = declination;...

The Carboniferous–Permian terrestrial successions record a global climatic shift from icehouse to hothouse conditions. Our multidisciplinary study documents an aridification trend throughout the ~ 1000 m thick composite terrestrial succession of the western Catalan Pyrenees (NE Iberian Peninsula), representing this time period. The detailed stratig...

The most severe biotic crisis on Earth history occurred during the Permian–Triassic (PT) transition around 252 Ma. Whereas in the marine realm such extinction event is well-constrained, in terrestrial settings it is still poorly known, mainly due to the lack of suitable complete sections. This is utterly the case along the Western Tethys region, lo...

Figure S1. The lower URU. A. Overview of the mudstones with fine-grained sandstone channels. B, C. Lateral accretions within sandstone channels. D. Small carbonate edaphic nodules. E. Vertical invertebrate burrow. F. Root trace fossils (whitish) in a medium- to coarse-grained sandstone.

Figure S2. Lower/upper URU boundary. A. Overview of the transition with three levels of nodules (white dashed lines); note the colour change between the lower and upper URU mudstones. B, C. Close view of the nodular levels (C squared in B), including septarian nodules (N) and root traces (Rt). D. Example of septarian nodules squared in C.

Figure S5. Tetrapod footprints of Morphotype I from cycle 38. A. General view of the trackway with the different manus-pes sets (RM-RP and LM-LP). B. Trackway in cross section. C. Well-preserved manus and partial pes, including 3D model and ichnites outline (as in Fig. 6B).

Figure S6. Tetrapod footprints of Morphotype II from cycle 39. A. Manus-pes set. B, C. Pes impressions.

Figure S7. Tetrapod footprints of Morphotype III from cycle 27. A-F. Footprints in surface G. H, I. Footprints from surface J. Note the different shapes of the heel (A-F, H, I) and the rough alignments of the footprints, possibly from the same trackways (G, J).

Figure S8. Slab with tetrapod footprints from Morphotype IV, close to cycle 49 (specimen IPS-99097). Squares are those of Fig. 6.