Full Terms & Conditions of access and use can be found athttp://www.tandfonline.com/action/journalInformation?journalCode=ghbi20

Download by: [Maxillo Facciali] Date: 23 September 2015, At: 07:32

Historical BiologyAn International Journal of Paleobiology

ISSN: 0891-2963 (Print) 1029-2381 (Online) Journal homepage: http://www.tandfonline.com/loi/ghbi20

Taking up the legacy of Waterhouse Hawkins andOwen: art and science for a new Italian project tobring back dinosaurs to life

Marco Romano, Simone Maganuco, Stefania Nosotti & Fabio Manucci

To cite this article: Marco Romano, Simone Maganuco, Stefania Nosotti & FabioManucci (2015): Taking up the legacy of Waterhouse Hawkins and Owen: art andscience for a new Italian project to bring back dinosaurs to life, Historical Biology, DOI:10.1080/08912963.2015.1089436

To link to this article: http://dx.doi.org/10.1080/08912963.2015.1089436

Published online: 22 Sep 2015.

Submit your article to this journal

View related articles

View Crossmark data

Historical Biology, 2015http://dx.doi.org/10.1080/08912963.2015.1089436

Taking up the legacy of Waterhouse Hawkins and Owen: art and science for a new Italian project to bring back dinosaurs to life

Marco Romanoa,b, Simone Maganucoc, Stefania Nosottic and Fabio Manuccid

aDipartimento di scienze della terra, sapienza, Università di roma, roma, italy; bsam Noble Museum, Norman, oK, Usa; cMuseo di storia Naturale di Milano, Milano, italy; dassociazione Paleontologica Parmense italiana, Parma, italy

ABSTRACTSince their initial formal recognition by Richard Owen in 1842, dinosaurs have always stood out in the collective imagination for their size and unusual appearance. Therefore, these marvellous animals are a source of curiosity and wonder for people of all ages, social and cultural backgrounds. Thanks to improved research techniques, palaeontologists have been able to work reconstructing the most plausible appearance of dinosaurs. Starting with petrified bones, they tried to make a dream come true – to bring the planet’s ancient inhabitants back to life. The new Italian exhibition Dinosaurs in the Flesh: Science and Art bring the Rulers of a Lost World Back to Life reveals the marriage of science and art that brings back to life animals that lived tens or hundreds of millions of years ago. Palaeontologists and artists collaborate on reconstructing the appearance of organisms from the distant past through study of the fossils, often with the aid of new technologies. The new project, which takes up the idea of Waterhouse Hawkins and Owen and their legacy to restore these ancient vertebrates based on solid scientific foundations, represents to date the only way to reanimate these fascinating lost animals.

© 2015 taylor & Francis

KEYWORDSDinosaur exhibitions; palaeoart; owen; Waterhouse Hawkins

ARTICLE HISTORYreceived 2 June 2015 accepted 29 august 2015

CONTACT Marco romano marco.romano@uniroma1.it

Introduction

On New Year’s Eve 1853, twenty-one prominent Victorian scientists dined inside the belly of a giant model of the dinosaur Iguanodon, celebrating the inauguration of the first dinosaur theme park (Figure 1). The exhibition, staged at Sydenham Park, South London, which had become the new location for an enormous glass building known as the Crystal Palace (Figure 1) and designed and built for the Great Exhibition of 1851 (McGowan-Hartmann 2013), was the result of a collab-oration between the sculptor Benjamin Waterhouse Hawkins and the famous British palaeontologist Richard Owen (Marshall 2007; Bramwell and Peck 2008) (Figure 2). It represented the first attempt to reconstruct life-sized models of dinosaurs, a group formally recognised by Owen in 1842, and other extinct ani-mals. Earlier the German palaeontologist Hermann von Meyer (1801–1869) had already recognised this group of animals as distinct and named them ‘Pachypoda’ (literally ‘thick foot’), but his proposal was not accepted (Avanzini et al. 2010).

More than a century and an half has passed since that dinner. Research on dinosaurs has flourished. The two eminent American palaeontologists, Edward Drinker Cope and Othniel Charles Marsh, engaged in the most famous and lively fossil feud (bone war), with a heated rivalry that led to vast collections of Mesozoic and Cenozoic vertebrate fossils from western North America (Wheeler 1960; Colbert 1997; Wallace 2000). Following the evoc-ative account by Colbert (1997), the late nineteenth and early

twentieth century witnessed a ‘Golden Age’ of dinosaur explo-ration in North America, with scientists from the United States National Museum (Smithsonian Institution) in Washington, American Museum of Natural History in New York, Carnegie Museum of Natural History in Pittsburgh, the National Museum of Canada in Ottawa and the Royal Ontario Museum in Toronto collecting, studying and publishing numerous finds. Among the key figures who have worked actively in the Morrison Forma-tion (Upper Jurassic) of the American West were Earl Douglass, Barnum Brown, Charles Whitney Gilmore (Colbert 1997) and Arthur Lakes (Kohl and McIntosh 1997). It followed the so-called ‘Dark Ages’ and the subsequent ‘Renaissance of Dinosaur Studies’ (Colbert 1997), with a couple of decades of forced break due to the Great Depression and World War II.

During the twentieth century, museum exhibition of pictorial and sculptural reconstructions became increasingly common, and in some cases, the example of Hawkins was followed in the construction of prehistoric parks. Among the early exam-ples is that of the Tierpark Hagenbeck, the Zoo of Stellingen (Hamburg, Germania), where an exhibition of life-size models was opened in 1909. The curator was a well-known zoological sculp-tor, Joseph Pallenberg, who updated to the new vision of the period the image of Iguanodon and other dinosaurs (including the North American species). In 1911, the founder of the Zoo, Carl Hagenbeck Jr., opened a second park in Rome, and the newspapers of that year also mention the prehistoric park in Germany, in an Italy still unfamiliar to the Mesozoic world:

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

2 M. ROMaNO eT al.

and described, and much has been learned about the biology of dinosaurs. Along with this dramatic growth in scientific knowl-edge, these amazing animals continue to inspire a sense of awe and wonder in both scientists and the public. The debate on some key issues concerning the distribution, extinction, physiology and general biology of dinosaurs is now more than ever increas-ingly vibrant. The controversy about the origin of birds and their phylogenetic relationships within the theropod clade provides a case in point (e.g. Feduccia 2002, 2012, 2013; Zelenitsky et al. 2012; Clarke 2013; Brusatte et al. 2014; Chiappe et al. 2014; Foth et al. 2014; Han et al. 2014; Smith et al. 2015).

Obviously, the numerous scientific advances enormously changed even the traditional iconography of the dinosaurs. Robert Bakker himself used his drawing skills to better con-vey those new hypotheses, enriching its articles of dynamic reconstructions (Bakker 1968). During the years, this extreme depiction of their dynamism has been resized and challenged. Beyond the debates, Bakker’s reconstructions immediately hit the imagination of contemporary illustrators, and among them, there was also Giovanni Caselli, one of the first authors (and the first Italian) to devote himself to the new look for dinosaurs (Halstead 1975). Rather than follow the reconstructions by Bakker, these and other works of the period show a blend of traditional and modern elements. There are, however, some exceptions well represented by Gregory Paul, a freelance researcher who has profoundly contributed to the palaeoartistic study, dedicating

In short, visitors in Stellingen are able to see, in one fence, flying reptiles with wings that look like ship sails, birds armed with formi-dable teeth, giant lizards, turtles and toads, which compared with their brethren of our times, make the latters look like miniature toys. (Un serraglio… 1911)

The chance to see similar life-size reproductions from real must certainly be the reason for the success of these parks, built on the sense of wonder. Since the 1930s, theme parks became increas-ingly common and popular (Debus and Debus 2002), although they not always resulted from a collaboration between palae-ontologists and artists. The dinosaurs were already well known to the general public, but towards the middle of the century, a revolution in palaeontology made obsolete the previous inter-pretations of fossils and consequently the reconstructions of the appearance of these ancient inhabitants of the planet.

In the late 1960s, starting with Robert Bakker and John Ostrom, a new generation of palaeontologists began to study dinosaurs from new perspectives (Colbert 1997). In the dec-ades that followed, we witnessed a veritable second golden age for collecting, study and publication of dinosaurs, with leading researchers like Philip Currie, John ‘Jack’ Horner (Figure 3), and the above-mentioned Robert Bakker and John Ostrom. Since then dinosaur research has never stopped, with a growing effort towards the discovery and documentation of new dinosaurs, and the development of new ideas concerning their phylogeny, phys-iology and behaviour. Hundreds of new taxa have been found

Figure 1. the famous dinner of twenty-one prominent Victorian scientists inside the belly of a giant model of the Iguanodon, on New year’s Eve 1853 (above). the crystal Palace from the great Exhibition, installed at sydenham. coloured photomechanical print, later than 1854? (below; from ‘Wellcome library, london’).

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

HISTORIcal BIOlOgy 3

himself to the realisation of skeletal, myological and in vivo reconstructions (Paul 1987). Many other authors have followed the example of Bakker and Paul. Among these, one of the most original and significant is Douglas Henderson, who has fully enhanced the palaeobotanist and landscape study. From his col-laboration with Jack Horner, resulted an innovative vision of the gregarious life and reproduction of dinosaurs, now shown as complex and social animals, in line with studies on Maiasaura (Horner 1984a). In Italy, the new ideas (and images) arrived with some delay, despite the growing popularity of the subject and the birth of the first prehistoric theme parks. Among them was the ‘Parco della Preistoria’ at Rivolta d’Adda (Cremona), founded in 1978, with a series of sculptures by Marcello Brandispada.

Figure 2. Benjamin Waterhouse Hawkins. Photograph by Maull & Polyblank, 1862. (above; from ‘Wellcome library, london’). Portrait of sir richard owen (below; from ‘Wellcome library, london’).

Many of the models are clearly inspired by the work of Zdeněk Burian. In fact, the illustrations of this extremely valued and prolific Czech artist, among the most prominent of the modern era, often became true icons, and since the late 1950s until the 1980s remarkably influenced prehistoric iconography, especially in Europe and Italy. A second theme exhibition was opened in 1978 inside the ‘Parco Natura Viva’, at Bussolengo (Verona). Even in such cases, the sculptures are of classical inspiration, while referring more faithfully to the osteological studies and photos provided in scientific papers. The ‘Dinomania’ was inaugurated in Italy from what has become one of the icons of the Museo di Storia Naturale di Milano: a sculpture of Triceratops, realised by Luciano Menghi, under the supervision by Giovanni Pinna.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

4 M. ROMaNO eT al.

as imagined by Owen and Waterhouse Hawkins for the Crys-tal Palace models? What was the appearance of the skin on the various parts of the body? What type of posture did the animals assume during locomotion or/and at rest? All these questions arise from the creative path of the artist, and the answers, worked out in intense dialogue with palaeontologists, help to further our knowledge on the subject (Figure 4).

In the reconstruction of each hyper-realistic model, a com-plete iconographic study of the dinosaur under consideration is conducted, trying to carefully analyse how the conception and interpretation of the animal have changed over time in terms of overall appearance, colour, posture and ornamentation details (see also Paul 1987; Witton et al. 2014). A classic, highly instruc-tive example is the change of interpretation in the case of the genus Iguanodon (Figure 5), with transition from a squat liz-ard-like quadrupedal reptile, almost monochromatic in colour integument and with a horn on its nose (which proved to be the spike-like ungual phalanx of the thumb), through a tripodal (with the tail used as support in addition to the hind legs) or obligatory bipedal, to the currently accepted interpretation of a quadruped/facultative biped. Drawings by Davide Bonadonna.

To reach the final model, all skeletal materials available for a particular taxon are analysed carefully, to take into account possible intraspecific variability, structures related to the sec-ondary sexual characters or even teratological forms. In this way, the team avoids the issue that peculiar structures, perhaps not representative of the taxon to be reconstructed, will become the new iconographic paradigm. In several cases, given the peculiar nature of the fossil record, it is necessary to retrodeform the analysed bones which, during diagenetic or tectonic processes, have suffered deformation. In this phase, an unavoidable dose of subjectivity is always present and it is attempted to minimise the interpretative part by using, where possible, comparisons with closely related forms to get a plausible result. A procedure of this type was used for the modelling and realisation of the recent model of Styracosaurus (Figure 6), which represents, to date, the most up-to-date and scientifically best-grounded reconstruction of this ceratopsian dinosaur (see Manucci et al. 2014).

The reproductions that make up the exhibition are repre-sented as static models in dynamic postures, which allow viewers to grasp the salient aspects of the dinosaur’s biology immedi-

Despite the bizarre appearance, the model was immediately able to excite audiences in Milan and one of the twentieth century’s most famous Italian writers, Dino Buzzati, made a provocative praise inviting art critics to not ignore the giant prehistoric por-traits.

Concerning the modern view, it has become popular since the 1990s, simultaneously with the discovery of the first skeletons and sites of dinosaur footprints in the peninsula, a true double revolution. New companies became specialised in the creation of life-size models under the supervision of Italian palaeontologists, in particular the Tridimont founded by Carmensilvio Tassinari, and the Geomodel by Mauro Scaggiante, whose work is merged in the exhibition here treated.

In this long history of scientific discoveries and artistic resto-rations, the main questions have remained the same: What did these extinct animals look like in life? Which differences and similarities can be found among the known species? How far can we push biological inferences based on the available, often limited data?

The new Italian project Dinosaurs in the Flesh: Science and Art bring the rulers of a lost World back to life is now trying to address these questions. It was born out of a passion shared by millions of ‘supporters’ and palaeontologists and was entirely developed and curated by professional researchers. The reconstructions of the extinct animals on display are based on the most up-to-date scientific ideas, which have been critically assessed without ever losing sight of the actual fossil record.

The central role of palaeoart

A key objective of the project is to underscore the central role of palaeoart in the process of scientific discovery. Illustrative draw-ings, environmental reconstructions and palaeoart in general are often dismissed as a mere ornamental addition to scientific publications. The time-consuming collaboration by artists and palaeontologists in reconstructing every model contradicts this point of view. When the artist puts her or his brush on canvas, she or he immediately starts posing questions to which science must provide answers: Which are the most likely colours for dinosaurs? Were these animals brightly coloured like extant birds (their living descendants) or were the colours dull and lifeless

Figure 3. Model of Spinosaurus aegyptiacus (length: 15.5 m) predating Mawsonia lavocati, with John ‘Jack’ Horner. Dinosaurs in the Flesh, Florence, 2012. Photo by andrea grigioni.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

HISTORIcal BIOlOgy 5

low 1981; Thulborn 1981, 1982, 1984; Lockley et al. 1994; Irby 1996; Kim and Huh 2010; Day et al. 2002; Mossman et al. 2003). The attitude of the tail is confirmed as straight and aligned with the remainder of the spine, since traces of the tail are relatively uncommon in the context of the entire ichnological record (in many classical representations, the tail – especially in sauropods – is shown as resting on the ground and being dragged during locomotion). The study of theropod and ornithopod footprints with substantial metatarsal impression has shed light on particu-lar behaviours in these dinosaurs, on the modalities of resting and crouching and on the peculiar pubic impressions when an

ately. Much evidence about the dinosaur biology in general, and about their static and dynamic posture in particular, is derived from a branch of palaeontology that only in recent decades has been recognised as important: ichnology. The study of fossil tracks makes it possible to obtain insight regarding behaviour of the living animal that left them, providing essential infor-mation that the individual bones cannot provide. In study after study in the last few decades, ichnological analyses have shed light on dinosaurian postures, both quadrupedal and bipedal; it is possible to obtain an idea of the type of gait and average speed in the various studied groups (e.g. Alexander 1976; Far-

Figure 4. the palaeoart gallery. Dinosaurs in the Flesh, Piacenza, 2011. Photo by Massimo Demma.

Figure 5. change of interpretation in the iconography of Iguanodon, with transition from a squat lizard-like quadrupedal reptile, through a tripodal with the tail used as support in addition to the hind legs or obligatory bipedal, to the currently accepted interpretation of a quadruped/facultative biped. Drawings by Davide Bonadonna.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

6 M. ROMaNO eT al.

Manning et al. 2009; Zheng et al. 2009; Herrero and Farke 2010; Li et al. 2010; Lingham-Soliar and Plodowski 2010; Dal Sasso and Maganuco 2011; Schweitzer 2011; Bell 2012; Davis 2014). It becomes a challenging but fascinating task of the collabora-tion between palaeoartist and palaeontologist to draw on all this physiological and behavioural evidence to generate the models and reconstructions (Figure 3), bearing in mind that, although a hypothesis may sound extravagant or bold, nature always man-ages to surprise us and exceed expectations (see for example, the amazing cranial ornamentation in Dracorex hogwartsia; Bakker et al. 2006; Horner and Goodwin 2009).

The educational aspect and the ‘making of’ section

The word ‘dinosaur’ is commonly used by the public in reference to many kinds of extinct animals without considering their phy-logenetic relationships, their anatomical characteristics and/or the different geological time in which they lived. Many extinct animals – for example, flying reptiles, Mesozoic marine reptiles or colossal land mammals – are not dinosaurs. People identify them as dinosaurs because of their look and size. In fact, when Sir Richard Owen coined the word ‘dinosaur’ in 1842, he specifically explained the English meaning of the word ‘deinos’ as ‘fearfully great’. Now people translate the name as ‘terrible’. A fundamental

animal adopts this posture (e.g. Lockley et al. 1998, 2003; Nicosia et al. 2007; Gierliński et al. 2009; Milner et al. 2009; Wilson et al. 2009). The gregarious behaviour in different groups of dino-saurs has been highlighted by the discovery and study of groups of parallel trackways, with individuals of different ages, moving in herds or in ‘families’ (e.g. Thulborn and Wade 1979; Lockley et al. 1983; Lockley 1994; Cotton et al. 1998; Lockley and Mat-sukawa 1999; Matsukawa et al. 1999; Schumacher 2003; Barco et al. 2006; Lockley et al. 2012). All this evidence demonstrates markedly social behaviour in different groups of dinosaurs, with well-structured relationships and much more complex dynamic than the old stereotype of slow and clumsy ‘big lizards’. As in present-day vertebrates, dinosaurs had well-marked secondary sexual characters, with function of sexual display apparently important during courtship. Most likely, some of these struc-tures was cartilaginous or fleshy in nature, and bright colours probably played a crucial role. Unequivocal evidence of struc-tures linked to sexual dimorphism has not yet been documented; however, there are some cases of exceptional preservation that allow artists to go beyond the simple skeletal reconstruction (e.g. Horner 1984b; Martill 1991; Kellner 1996; Briggs et al. 1997; Benton 1998; Chiappe et al. 1998; Chen et al. 1998; Monasterski 1999; Xu et al. 1999a, 1999b; Martill et al. 2000a, 2000b; Gatesy 2001; Norell et al. 2001; Mayr et al. 2002; Schweitzer et al. 2007;

Figure 6. Model of Styracosaurus albertensis (length: 4.5 m): the new, most updated paradigm by Dinosaurs in the Flesh, rome, 2014–2015. Photo by anna giamborino.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

HISTORIcal BIOlOgy 7

reconstructed and placed in a virtual aquatic environment, where they interact with each other.

The use of digital technologies is extended to the prepara-tion of the sculptures, whose base is modelled digitally, and then printed in real size into several mountable parts. The digi-tal maquette, manipulable in every angulation, can be prepared faithfully tracing the proportions of skeletal studies. The art-ist works on the computer, followed step by step by a scientific adviser, through a mutual exchange of comments and visual references. The use of these useful tools is spreading only in the last few years, and its practicality becomes increasingly apparent. Most of the models prepared in Dinosaurs in the Flesh are works by Davide Bonadonna, under the direct supervision of Simone Maganuco.

An entire section of the exhibition is devoted to the main Ital-ian illustrators. The palaeoart gallery presents the different artists and their fundamental contribution to science education (Figure 4), highlighting the different techniques and stylistic solutions followed in creating various illustrations. Prominent Italian pal-aeoartists include Marco Auditore, Davide Bonadonna, Massimo Demma, Fabio Fogliazza, Alberto Gennari, Fabio Manucci, Sante Mazzei, Marzio Mereggia, Lukas Panzarin, Fabio Pastori, Loana Riboli, Emiliano Troco, and Renzo Zanetti, one of the first Ital-ian illustrators to have specialised in this field. Not all of these authors share the same idea of palaeoart and vary in interests and techniques, although in the research of a common scientific substantiation (e.g. Figures 8, 9; Dal Sasso and Maganuco 2011, p. 269–278). Some, as in the case of Marco Auditore and Massimo

aspect of the project and of the exhibit is to try to communicate as simply and clearly as possible the key differences between dinosaurs and other animals often connected to this group. The educational process, while remaining accessible to the general public, is designed to underscore the great complexity of the history of life on Earth with numerous and complex feedback and interrelated processes.

The structure of the exhibit is different from that of other current exhibitions dealing with ancient life. The visitors are not just passive spectators but go on a guided tour behind the scenes. They will witness the suggestive ‘making of ’ the exhibition, a sort of open laboratory, and observe every phase of the development and fabrication of the models (Figure 7). At the same time, visi-tors are introduced to palaeontology as a scientific discipline and to the work of the different kinds of professionals who, called in by palaeontologists, share their passion and – with curiosity and patience – took their competence in new directions to help in the difficult task of fossil interpretation. These professionals, who can help explain lost worlds by translating them in ways that are immediately accessible to everyone, include illustrators, sculptors, craftsmen, special-effects artists and experts in 2D and 3D modelling.

An innovative aspect of this section is represented by virtual animations, a modern approach to representing the animals in their original habitats, which parallels the static reconstruc-tions. In this regard, there are many projects in progress, as the ‘paleo-aquarium’ section (a project that uses cutting-edge 3D technology) that is dedicated to extinct sea animals, virtually

Figure 7. Bringing back to life Stegosaurus: final fleshed-out restoration; cast of the skull; styrofoam maquette; pencil sketch. Photos by andrea leanza, drawing by Davide Bonadonna.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

8 M. ROMaNO eT al.

Concluding remarks

Riding the wave of a never-exhausted, indeed growing enthu-siasm for dinosaurs and taking the rich legacy of almost two hundred years of studies on dinosaurs, the Italian project Dino-saurs in the Flesh has two primary objectives. The first is the close collaboration between professional palaeontologists and prominent palaeoartists. Each new work undertaken, aside from presenting a review and synthesis of previous studies and interpretations, provides the most up-to-date reconstruc-tion, that, as has already occurred, may become the new par-adigm for a particular type of dinosaur (e.g. Jinfengopteryx, Scipionyx, Sinosauropteryx, Spinosaurus, Styracosaurus, Figure 10). The creative process between the scientist and the artist is not merely concerned with the realisation of a model. Each new question generates others, and, as often happens in the process of scientific research, the culmination of a study can be far away from the expected target. This does not exclude that seemingly simple questions of stylistic character will lead to new arguments, even new theories on biological, physiological and behavioural aspects, or the formulation of entirely new hypotheses. Reconstructions and how-to guide by Paul (1987) are now to be renewed in some details, but their construc-tive intent remains admirable, and should be integrated with the contemporary interest for less conventional hypotheses, as it occurred recently with the palaeoart book All Yesterdays (Conway et al. 2012; Witton et al. 2014).

Demma, are more dedicated to traditional scientific illustration, preparing accurate anatomical drawings and illustrations of fossil specimens, fundamental for academic publications (e.g. Nosotti and Pinna 1996, 1999; Nosotti and Rieppel 2002; Nosotti 2007; Maganuco et al. 2009; Dal Sasso and Maganuco 2011; Maganuco et al. 2014).

Even Davide Bonadonna accompanies with his work numer-ous scientific publications, but is more focused on in vivo recon-struction (e.g. Tschopp and Mateus 2012; Maganuco et al. 2014). The same is true for Fabio Fogliazza (e.g. Garassino and Schwei-gert 2006; Nosotti 2007; Nosotti and Teruzzi 2008), illustrator and traditional sculptor who recently almost completely devoted his work to palaeoanthropology (Fogliazza and Peresani 2013): the artistic choices of his portraits, which fully comply with the anatomical knowledge provided by experts and always draw cre-ative inspiration from the scientific data, prove the unique com-municative power of art in summarising research achievements in an intuitive and accessible-to-all way. Lukas Panzarin makes both scientific and in vivo reconstructions, and even co-authored scientific papers thanks to his deep knowledge of some taxa (e.g. McDonald et al. 2010). Others, like Emiliano Troco, turning their interest in a more comprehensive naturalistic vision, also seeking an artistic purpose for the work, while maintaining central the collaboration with a scientific supervisor.

Many of these artists have worked and continue to work in the project, providing personal expressive qualities in artistic reconstructions.

Figure 8. Examples of recent artworks made by italian palaeoartists: a Neanderthal shaman by Fabio Fogliazza, Edaphosaurus by Davide Bonadonna.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

HISTORIcal BIOlOgy 9

sistent and thorough, through the use of a specialised literature, as already it happens in some cases (Antón 2007).

Being a travelling exhibition for all types of public venues, a second key objective is the structuring and implementation of the educational component. The introductory section and the

The proposal for new hypotheses or speculative ideas may prove helpful, if the foundations and evidences are reported and discussed, exploring the different plausible possibilities, and excluding the most blatantly unlikely. In the future, these scientist–artist collaborations hopefully will become more con-

Figure 9. artworks made by italian palaeoartists: Sivatherium by Emiliano troco and a pack of Deinonychus by loana riboli.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

10 M. ROMaNO eT al.

millions of years ago? The legacy of Waterhouse Hawkins and Owen and the later projects they inspired, including Dinosaurs in the Flesh, represents the only way to reanimate these fascinating extinct species as living animals.

AcknowledgementsHans-Dieter Sues is warmly thanked for his comments and corrections that greatly improved an early version of the manuscript. We thank Mau-ro Scaggiante and the staff Geomodel (models Dinosaurs in the Flesh, ex-hibition set-up), Associazione Paleontologica Parmense Italiana A.P.P.I. (management, didactics and special events Dinosaurs in the Flesh), the art-ist Davide Bonadonna (illustrations and digital sculpturing of the models Dinosaurs in the Flesh), and Andrea Pirondini and the Prehistoric Minds (layout, graphic design, Dinosaurs in the Flesh). The Editor and an anony-mous reviewer are thanked for their suggestions and corrections that have improved the manuscript.

Disclosure statementNo potential conflict of interest was reported by the authors.

ReferencesAlexander R. 1976. Estimates of speeds of dinosaurs. Nature. 261:24–130.Antón M. 2007. El secreto de los fósiles. El arte y la ciencia de reconstruir

a nuestros antepasados (y otras criaturas) [The secret of fossil. Art and science to reconstruct our ancestors (and other creatures)]. Madrid: Aguilar; p. 357.

Avanzini M, Bernardi M, Melchiori L, Petti FM. 2010. Le orme dei dinosauri del Castello di San Gottardo a Mezzocorona con cenni della storia del castello [The dinosaur footprints of the Castle of San Gottardo at Mezzocorona with hints of history of the castle]. Mori (TN): La Grafica srl; p. 140.

Bakker RT. 1968. The superiority of dinosaurs. Discovery. 3:11–22.Bakker RT, Sullivan RM, Porter V, Larson P, Saulsbury SJ. 2006. Dracorex

hogwartsia, n. gen., n. sp., a spiked, flat-headed pachycephalosaurid dinosaur from the Upper Cretaceous Hell Creek Formation of South Dakota. In: Lucas SG, Sullivan RM, editors. Late Cretaceous vertebrates from the Western Interior. Albuquerque (NM): NMMNH 35; p. 331–345.

Barco JL, Canudo JI, Ruiz-Omeñaca JI. 2006. New data on Therangospodus oncalensis from the Berriasian Fuentesalvo Tracksite (Villar del Río, Soria, Spain): an example of gregarious behaviour in theropod dinosaurs. Ichnos. 13:237–248.

illustrative panels for visitors have been designed to explain, in simple but scientifically accurate fashion, even the most complex aspects of the biology of dinosaurs and other extinct animals and the history of life on Earth. Popularising science is often neglected but represents a fundamental and important process for a mature and educated society. Italy, and Italian scientists, has played a key role in the early development of geology and palaeontology (e.g. Romano 2013, 2014, 2015a, 2015b; Romano and Cifelli 2015; Romano et al. 2015a, 2015b), and now is finally entering in a consistent way also in dinosaur study, although with some delay if compared to other countries. Dinosaurs in the Flesh seeks to create a stable and lasting bridge between the academic community and the general public. Being conceived as a travelling exhibition, each new instalment of the exhibit is designed to fully enhance the host structure and the permanent collections of the hosting venue, allowing the public to visit nat-ural treasures previously largely unknown to laypersons, and to learn about the work of Italian museums, universities and associations which are committed to sharing the knowledge of the past with general audiences.

In addition, new Italian palaeontologists become involved as the exhibition travels to new locations, collaborating fully as active partners in the project. Dinosaurs in the Flesh could then create a kind of bond for the Italian palaeontological community, with a renaissance of vertebrate palaeontology in our country.

In conclusion, we once again extend our heartfelt thanks to sci-entists for pioneering projects such as the collaboration between Owen and Waterhouse Hawkins. The models of the extinct ani-mals at Crystal Palace have often been unfairly criticised in later publications, which focus on the mistakes in the design of the models and ignore the fact that these reconstructions repre-sented the limited state of scientific knowledge available at that time. Indeed we should be grateful for this effort by Waterhouse Hawkins and Owen, who provided the first reconstructions ‘in the flesh’ of dinosaurs and other ancient vertebrates. Despite the astonishing advances in biotechnology over the past two decades (e.g. Rogaev et al. 2006), to bring non-avian dinosaurs back to life based on fossilised DNA very much remains in the realm of science fiction. Is there any possibility to ever bring back to life dinosaurs and other animals that lived tens or hundreds of

Figure 10. Model of Jinfengopteryx elegans with true plumage, length: 55 cm. By Dinosaurs in the Flesh. By Dinosaurs in the Flesh and Boban Filipović.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

HISTORIcal BIOlOgy 11

Herrero L, Farke AA. 2010. Hadrosaurid dinosaur skin impressions from the Upper Cretaceous Kaiparowits Formation of Southern Utah, USA. PalArch J Vert Pal. 7:1–7.

Horner JR. 1984a. The nesting behavior of dinosaurs. Sci Am. 250:130–137.Horner JR. 1984b. A ‘segmented’ epidermal tail frill in a species of

hadrosaurian dinosaur. J Paleontol. 58:270–271.Horner JR, Goodwin MB. 2009. Extreme cranial ontogeny in the Upper

Cretaceous Dinosaur Pachycephalosaurus. PLoS ONE. 4(10):e7626. doi:10.1371/journal.pone.0007626

Irby GV. 1996. Paleoichnological evidence for running dinosaurs worldwide. Mus North Ariz Bull. 60:109–112.

Kellner AWA. 1996. Fossilized theropod soft tissue. Nature. 379:32.Kim BS, Huh M. 2010. Analysis of the acceleration phase of a theropod

dinosaur based on a Cretaceous trackway from Korea. Palaeogeogr Palaeoclimatol Palaeoecol. 293:1–8.

Kohl MF, McIntosh JS. 1997. Discovering dinosaurs in the old west: the field journals of Arthur Lakes. Washington (DC): Smithsonian Books; p. 198.

Li Q, Gao K-Q, Vinther J, Shawkey MD, Clarke JA, D’Alba L, Meng Q, Briggs DEG, Prum RO. 2010. Plumage color patterns of an extinct dinosaur. Science. 327:1369–1372.

Lingham-Soliar T, Plodowski G. 2010. The integument of Psittacosaurus from Liaoning Province, China: taphonomy, epidermal patterns and color of a ceratopsian dinosaur. Naturwissenschaften. 97:479–486.

Lockley MG. 1994. Dinosaur ontogeny and population structure: interpretations and speculations based on footprints. In: Carpenter K, Hirsch K, Horner J, editors. Dinosaur eggs and babies. Bloomington (IL): Cambridge University Press; p. 347–364.

Lockley MG, Matsukawa M. 1999. Some observations on trackway evidence for gregarious behavior among small bipedal dinosaurs. Palaeogeogr Palaeoclimatol Palaeoecol. 150:25–31.

Lockley MG, Young BH, Carpenter K. 1983. Hadrosaur locomotion and herding behavior: evidence from footprints in the Mesa Verde Formation, Grand Mesa Coal Field, Colorado. Mountain Geol. 20:5–14.

Lockley MG, Hunt AP, Moratalla J, Matsukawa M. 1994. Limping dinosaurs? Trackway evidence for abnormal gaits. Ichnos. 3:193–202.

Lockley MG, Hunt AP, Meyer C, Rainforth EC, Schultz RJ. 1998. A survey of fossil footprint sites at glen canyon national recreation area (western USA): a case study in documentation of trace fossil resources at a national preserve. Ichnos. 5:177–211.

Lockley M, Matsukawa M, Jianjun L. 2003. Crouching theropods in taxonomic jungles: ichnological and ichnotaxonomic investigations of footprints with metatarsal and ischial impressions. Ichnos. 10:169–177.

Lockley MG, Huh M, Gwak S-G, Koo HG, Paik IS. 2012. Multiple tracksites with parallel trackways from the Cretaceous of the Yeosu City Area Korea: implications for gregarious behavior in Ornithopod and Sauropod dinosaurs. Ichnos. 19:105–114.

Maganuco S, Steyer JS, Pasini G, Boulay M, Lorrain S, Bénéteau A, Auditore M. 2009. An exquisite specimen of Edingerella madagascariensis (Temnospondyli) from the Lower Triassic of NW Madagascar; cranial anatomy, phylogeny and restorations. Mem Soc Ital Sci Nat Mus Civ Stor Nat Milano. 34:1–72.

Maganuco S, Pasini G, Auditore M. 2014. A revision of the short-faced stereospondyls Mahavisaurus dentatus and Lyrosaurus australis from the Lower Triassic of Madagascar: cranial anatomy, ontogenetic remarks, palaeoecology and rhytidosteid phylogeny. Mem Soc Ital Sci Nat Mus Civ Stor Nat Milano. 39:1–64.

Manning PL, Morris PM, McMahon A, Jones E, Gize A, Macquaker JHS, Wolff G, Thompson A, Marshall J, Taylor KG, et al. 2009. Mineralized soft-tissue structure and chemistry in a mummified hadrosaur from the Hell Creek Formation, North Dakota (USA). Proc R Soc B: Biol Sci. 276:3429–3437.

Manucci F, Maganuco S, Panzarin L, Bonadonna D. 2014. Improving collaboration between paleontologists and paleoartists: a case study, focused on the ceratopsid dinosaur Styracosaurus. Society of Vertebrate Paleontology – abstract of papers 74th Annual Meeting, Berlin; p. 177.

Marshall NR. 2007. ‘A dim world, where monsters dwell’: the spatial time of the Sydenham Crystal Palace Dinosaur Park. Vic Stud. 49:286–301.

Martill DM. 1991. Organically preserved dinosaur skin: taphonomic and biological implications. Mod Geol. 16:61–68.

Bell PR. 2012. Standardized terminology and potential taxonomic utility for hadrosaurid skin impressions: a case study for Saurolophus from Canada and Mongolia. PLoS ONE. 7:1–12.

Benton MJ. 1998. Dinosaur fossils with soft parts. Trends Ecol Evol. 13:303–304.

Bramwell V, Peck RM. 2008. All in the bones: a biography of Benjamin Waterhouse Hawkins. Philadelphia (PA): The Academy of Natural Sciences of Philadelphia; p. 128.

Briggs DEG, Wilby PR, Perez-Moreno BP, Sanz JL, Fregenal-Martinez M. 1997. The mineralization of dinosaur soft tissue in the Lower Cretaceous of Las Hoyas, Spain. J Geol Soc. 154:587–588.

Brusatte SL, Lloyd GT, Wang SC, Norell MA. 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across dinosaur–bird transition. Curr Biol. 24:2386–2392.

Chen P-J, Dong Z-M, Zhen S-N. 1998. An exceptionally well-preserved theropod dinosaur from the Yixian Formation of China. Nature. 391:147–152.

Chiappe L, Coria R, Dingus L, Jackson F, Chinsamy A, Fox M. 1998. Sauropod dinosaur embryos from the Late Cretaceous of Patagonia. Nature. 396:258–261.

Chiappe LM, Zhao B, O’Connor JK, Chunling G, Wang X, Habib M, Marugan-Lobon J, Meng Q, Cheng X. 2014. A new specimen of the Early Cretaceous bird Hongshanornis longicresta: insights into the aerodynamics and diet of a basal ornithuromorph. PeerJ. 2:e234.

Clarke JA. 2013. Feathers before flight. Science. 340:690–692.Colbert EH. 1997. North American dinosaur hunters. In: Farlow JO, Brett-

Surman MK, editors. The Dinosaur. Indiana University Press; p. 25–33.Conway J, Kosemen CM, Naish D. 2012. All yesterdays: unique and

speculative views of dinosaurs and other prehistoric animals. Raleigh (NC): Irregular Books; p. 1–100.

Cotton WD, Cotton JE, Hunt AP. 1998. Evidence for social behavior in ornithopod dinosaurs from the Dakota group of northeastern New Mexico, U.S.A. Ichnos. 6:141–149.

Dal Sasso C, Maganuco S. 2011. Scipionyx samniticus (Theropoda: Compsognathidae) from the Lower Cretaceous of Italy: osteology, ontogenetic assessment, phylogeny, soft tissue anatomy, taphonomy and palaeobiology. Mem Soc Ital Sci Nat Mus Civ Stor Nat Milano. 37:1–282.

Davis M. 2014. Census of dinosaur skin reveals lithology may not be the most important factor in increased preservation of hadrosaurid skin. Acta Palaeontol Pol. 59(3):601–605.

Day JJ, Norman DB, Upchurch P, Powell HP. 2002. Biomechanics: dinosaur locomotion from a new trackway. Nature. 415:494–495.

Debus AA, Debus DE. 2002. Paleoimagery: the evolution of dinosaurs in art. Jefferson (NC): McFarland & Company; p. 285.

Farlow JO. 1981. Estimates of dinosaur speeds from a new trackway site in Texas. Nature. 294:747–748.

Feduccia A. 2002. Birds are dinosaurs: simple answer to a complex problem. The Auk. 119:1187–1201.

Feduccia A. 2012. Riddle of the feathered dragons: hidden birds of China. New Haven (CT): Yale University Press; p. 368.

Feduccia A. 2013. Bird origins anew. The Auk. 130:1–12.Fogliazza F, Peresani M. 2013. Amazing Neanderthal. A view from the

country of fashion. Wien: Hugo Obermaier Conference.Foth C, Tischlinger H, Rauhut OWM. 2014. New specimen of Archaeopteryx

provides insights into the evolution of pennaceous feathers. Nature. 511:79–82.

Garassino A, Schweigert G. 2006. The Upper Jurassic Solnhofen decapod crustacean fauna: review of the types from old descriptions. Part I. Infraorders Astacidea, Thalassinidea and Palinura. Mem Soc Ital Sci Nat Mus Civ Stor Nat Milano. 34:1–64.

Gatesy SM. 2001. Skin impressions of Triassic theropods as records of foot movement. Bull Mus Comp Zool. 156:137–149.

Gierliński GD, Lockley MG, Niedzwiedzki G. 2009. A distinctive crouching theropod trace from the Lower Jurassic of Poland. Geol Q. 53:471–476.

Halstead LB. 1975. The evolution and ecology of the dinosaurs. London: Peter Lowe; p. 120.

Han G, Chiappe LM, Ji S, Habib M, Turner AH, Chinsamy A, Liu X, Han L. 2014. A new raptorial dinosaur with exceptionally long feathering provides insights into dromaeosaurid flight performance. Nat Commun. 5:1–9.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15

12 M. ROMaNO eT al.

Romano M. 2014. From petrified snakes, through giant ‘foraminifers’, to extinct cephalopods: the early history of ammonite studies in the Italian peninsula. Hist Biol. 27:214–235.

Romano M. 2015a. ‘Per tremoto o per sostegno manco’: the geology of Dante Alighieri’s Inferno ["For an earthquake or erosion": the geology of Dante Alighieri’s Inferno]. Ital J Geosci. doi:10.3301/IJG.2015.21

Romano M. 2015b. Reviewing the term uniformitarianism in modern Earth sciences. Earth-Sci Rev. 148:65–76.

Romano M, Cifelli R. 2015. Evolving ‘creature’: an avoidable oxymoron. Evol Dev. 17:173–174.

Romano M, Cifelli R, Vai G. 2015a. Natural history: first museologist’s legacy. Nature. 517:271.

Romano M, Cifelli R, Vai G. 2015. The first geological map: an Italian legacy. Ital J Geosci doi:10.3301/IJG.2015.23

Schumacher BA. 2003. An addition to the dinosaur freeway megatracksite, Dakota Group (Upper Cretaceous), Bent County, Colorado. Ichnos. 10:255–262.

Schweitzer MH. 2011. Soft tissue preservation in terrestrial mesozoic vertebrates. Annu Rev Earth Planet Sci. 39:187–216.

Schweitzer MH, Suo Z, Avci R, Asara JM, Allen MA, Arce FT, Horner JR. 2007. Analyses of soft tissue from Tyrannosaurus rex suggest the presence of protein. Science. 316:277–280.

Smith NA, Chiappe LM, Clarke JA, Edwards SV, Nesbitt SJ, Norell MA, Stidham TA, Turner A, van Tuinen M, Vinther J, Xu X. 2015. Rhetoric vs. reality: a commentary on ‘Bird Origins Anew’ by A. Feduccia. The Auk. 132:467–480.

Thulborn RA. 1981. Estimated speed of a giant bipedal dinosaur. Nature. 292:273–274.

Thulborn RA. 1982. Speeds and gaits of dinosaurs. Palaeogeogr Palaeoclimatol Palaeoecol. 38:227–256.

Thulborn RA. 1984. Preferred gaits of bipedal dinosaurs. Alcheringa. 8:243–252.

Thulborn RA, Wade M. 1979. Dinosaur stampede in the Cretaceous of Queensland. Lethaia. 12:275–279.

Tschopp E, Mateus OV. 2012. The skull and neck of a new flagellicaudatan sauropod from the Morrison Formation and its implication for the evolution and ontogeny of diplodocid dinosaurs. J Syst Palaeontol. 1:853–888.

Wallace DR. 2000. The Bonehunters’ revenge: dinosaurs, greed, and the greatest scientific feud of the gilded age. Reprint ed. New York (NY): Mariner Books; p. 384.

Wheeler WH. 1960. The uintatheres and the Cope-Marsh War. Science. 131:1171–1176.

Wilson JA, Marsicano CA, Smith RMH. 2009. Dynamic locomotor capabilities revealed by early dinosaur trackmakers from Southern Africa. PLoS ONE. 4:e7331. doi:10.1371/journal.pone.0007331

Witton MP, Naish D, Conway J. 2014. State of the Palaeoart. Palaeontologia Electronica; vol. 17: 5E; p. 10. Available from: http://palaeo-electronica.org/content/2014/768-contributions-by-amateur-paleontologists

Xu X, Zhi-Lu T, Xiao-Lin W. 1999a. A therizinosauroid dinosaur with integumentary structures from China. Nature. 399:350–354.

Xu X, Xiao-Lin W, Xiao-Chun W. 1999b. A dromaeosaurid dinosaur with a filamentous integument from the Yixian Formation of China. Nature. 401:262–266.

Un serraglio preistorico.1911. La Lettura [A prehistoric fence]. 3:282–285.Zelenitsky DK, Therrien F, Erickson GM, DeBuhr CL, Kobayashi Y, Eberth

DA, Hadfield F. 2012. Feathered non-avian dinosaurs from North America Provide insight into wing origins. Science. 338:510–514.

Zheng X-T, You H-L, Xu X, Dong Z-M. 2009. An Early Cretaceous heterodontosaurid dinosaur with filamentous integumentary structures. Nature. 458:333–336.

Martill DM, Batten DJ, Loydell DK. 2000a. A new specimen of the thyreophoran dinosaur cf. Scelidosaurus with soft tissue preservation. Palaeontology. 43:549–559.

Martill DM, Frey E, Sues H-D, Cruickshank ARI. 2000b. Skeletal remains of a small theropod dinosaur with associated soft structures from the Lower Cretaceous Santana Formation of northeastern Brazil. Can J Earth Sci. 37:891–900.

Matsukawa M, Lockley MG, Hunt AP. 1999. Three age groups of ornithopods inferred from footprints in the mid-Cretaceous Dakota Group, eastern Colorado, North America. Palaeogeogr Palaeoclimatol Palaeoecol. 147:39–51.

Mayr G, Peters DS, Plodowski G, Vogel O. 2002. Bristle-like integumentary structures at the tail of the horned dinosaur Psittacosaurus. Naturwissenschaften. 89:361–365.

McDonald AT, Kirkland JI, DeBlieux DD, Madsen SK, Cavin J, Milner ARC, Panzarin L. 2010. New basal iguanodonts from the Cedar Mountain Formation of Utah and the evolution of thumb-spiked dinosaurs. PLoS ONE. 5(11):e14075. doi:10.1371/journal.pone.0014075

McGowan-Hartmann J. 2013. Shadow of the dragon: the convergence of myth and science in nineteenth century paleontological imagery. J Soc Hist. 47:47–70.

Milner ARC, Harris JD, Lockley MG, Kirkland JI, Matthews NA. 2009. Bird-like anatomy, posture, and behavior revealed by an Early Jurassic theropod dinosaur resting trace. PLoS ONE. 4:e4591. doi:10.1371/journal.pone.0004591

Monasterski R. 1999. Getting under a dinosaur’s skin. Science. 155:38.Mossman DJ, Bruning R, Powell HP. 2003. Anatomy of a jurassic theropod

trackway from Ardley, Oxfordshire, U.K. Ichnos. 10:195–207.Nicosia U, Petti FM, Perugini G, Porchetti S, Sacchi E, Conti MA, Mariotti

N, Zarattini A. 2007. Dinosaur tracks as paleogeographic constraints: new scenarios for the Cretaceous geography of the periadriatic region. Ichnos. 14:69–90.

Norell MA, Makovicky PJ, Currie PJ. 2001. The beaks of ostrich dinosaurs. Nature. 412:873–874.

Nosotti S. 2007. Tanystropheus longobardicus (Reptilia, Protorosauria): re-interpretation of the anatomy based on new specimens from the Middle Triassic of Besano (Lombardy, Northern Italy). Mem Soc Ital Sci Nat Mus Civ Stor Nat Milano. 35: 1–88.

Nosotti S, Pinna G. 1996. Osteology of the skull of Cyamodus kuhnschnyderi Nosotti & Pinna 1993 (Reptilia, Placodontia). Pal Lomb Soc Ital Sci Nat Mus Civ Stor Nat Milano. 6:3–42.

Nosotti S, Pinna G. 1999. Skull anatomy of Protenodontosaurus italicus Pinna 1990 (Reptilia, Placodontia). Pal Lomb Soc Ital Sci Nat Mus Civ Stor Nat Milano. 11:3–17.

Nosotti S, Rieppel O. 2002. The braincase of Placodus Agassiz, 1833 (Reptilia, Placodontia). Mem Soc Ital Sci Nat Mus Civ Stor Nat Milano. 31:3–18.

Nosotti S, Teruzzi G. 2008. I rettili di Besano-Monte San Giorgio [The reptiles of Besano-Monte San Giorgio]. Nat Soc Ital Sci Nat Mus Civ Stor Nat Milano. 98:1–99.

Paul GS. 1987. The science and art of restoring the life appearance of dinosaurs and their relatives: a rigorous how-to guide. In: Czerkas SJ, Olson EC, editors. Dinosaurs past and present. Seattle (WA): University of Washington Press; vol II; p. 5–49.

Rogaev EI, Moliaka YK, Malyarchuk BA, Kondrashov FA, Derenko MV, Chumakov I, Grigorenko AP. 2006. Complete mitochondrial genome and phylogeny of pleistocene mammoth Mammuthus primigenius. PLoS Biol. 4(3):e73. doi:10.1371/journal.pbio.0040073

Romano M. 2013. ‘The vain speculation disillusioned by the sense’: the Italian painter Agostino Scilla (1629–1700) called ‘The discoloured’, and the correct interpretation of fossils as ‘lithified organisms’ that once lived in the sea. Hist Biol. 26:631–651.

Dow

nloa

ded

by [

Max

illo

Facc

iali]

at 0

7:32

23

Sept

embe

r 20

15