Paleoart (and the paleoartist) in the Service of Scientific Research

by Simone Zoccante

Unless otherwise noted, all images © Simone Zoccante

Figure 1: Life reconstruction
of Megantereon cultridens, an
extinct saber-toothed cat
closely related to the betterknown
Smilodon. This species
possessed a rather robust body
structure and was roughly the
size of a large jaguar. Its limbs,
particularly the forelimbs,
were strong and sturdy, while
its tail was relatively short,
similar to that of a lynx.

Figure 1: Life reconstruction of Megantereon cultridens, an extinct saber-toothed cat closely related to the better-known Smilodon. This species possessed a rather robust body structure and was roughly the size of a large jaguar. Its limbs, particularly the forelimbs, were strong and sturdy, while its tail was relatively short, similar to that of a lynx.

 When observing a piece of paleoart, such as a reconstruction of an extinct organism or ecosystem, it’s easy to be captivated by the visual impact of the image. Scenes of a Deinotherium feeding in a Miocene forest, an Allosaurus hiding in vegetation ready to launch its attack, or a woolly rhinoceros wandering through a desolate steppe may seem like the products of an artist’s imagination, but behind each of these images lies a complex process and a demanding craft, where scientific research, in-depth study of a wide variety of sources, comparative anatomy, and artistic interpretation coexist in a delicate balance. 

It is precisely this extensive process of research and source analysis that distinguishes paleoart from other forms of illustration. A wildlife illustrator has the privilege and advantage of being able to observe their subject directly, consult a photographic reference archive, and study its behavior through documentaries and specimens kept in captivity. By contrast, the paleoartist almost always works without any direct contact with the subject they are depicting. The subject may be extinct organisms or environments. The starting point is typically fossil remains which are often incomplete or distorted by taphonomic processes and therefore provide only limited information regarding the anatomy and nature of a particular species. The living appearance and the execution of specific behaviors are thus inferred through a rigorous process of research, deduction, and inference. 

One of the most widespread misunderstandings about paleoart arises from unfamiliarity with the concept of inference. Those unacquainted with paleoart may imagine that a paleontological illustrator’s work consists of “filling in the gaps” in the fossil record with a generous dose of imagination. In reality, reconstruction by inference is an intellectual exercise and the resulting image is never the product of artistic whim. Every element added to the image and every decision made regarding how to reconstruct a specific part of an organism’s body should stem from reasoning based on available data and the study of a range of possibilities consistent with the anatomical, biological, and ecological processes that paleoscientists and paleoartists currently understand. 


When the available evidence is insufficient, the paleoartist’s role is not to invent, but to identify the most plausible solution within that range of possibilities, declaring when possible the degree of uncertainty that accompanies that choice. 



Another common misconception made about paleoart is assuming that, since creating paleoart is an artistic process, art takes precedence over the scientific component, and that being a skilled draftsman is sufficient to plausibly and credibly reconstruct an extinct animal. In reality, the creation of knowledge through study (which ultimately forms the intellectual framework that shapes our vision of an extinct world) is itself part of the artistic research process, just as in broader art, one conducts research on materials or specific concepts to express. In paleoart, art is guided by scientific information. Paleoart serves as a tool for disseminating precise scientific knowledge. Without the appropriate cultural background, even the best draftsman in the world will fail to make plausible and coherent reconstruction choices and will inevitably end up creating copies of what already exists. 


Ultimately, the quality of a paleoartistic work does not depend solely on its aesthetic rendering, but primarily on the solidity of the reconstruction choices made throughout the process. 


CASE STUDY 


At the beginning of 2026, I embarked on a truly stimulating project, one that combined passion, a rewarding commission, and a variety of skills that a professional in my field must have in their toolkit. 


The project for this brief journey into the mind of this paleoartist involved the reconstruction of certain anatomical parts and some behaviors of Megantereon cultridens, a felid with saber-like teeth closely related to the more famous Smilodon, that inhabited Europe during the Pliocene and Pleistocene (3.3–1.8 million years ago). 


Toward the end of 2025, I was contacted by Dr. Saverio Bartolini- Lucenti, a researcher at the University of Florence specializing in carnivores. At the time, Saverio was leading a study of Megantereon cultridens focusing on the animal’s brain which was being studied for the first time. He commissioned me to produce a set of scientific illustrations as supporting material for the study, including: 


  • A life reconstruction of Megantereon cultridens (Fig. 1, first image on page); 
  • A reconstruction of its brain; and 
  • A hunting scene illustrating some behaviors and environmental preferences deduced from the study of the brain. 



Figure 2: Reconstruction sequence illustrates the skull, the digital endocast, and the reconstructed brain
of Megantereon. Computed
tomography (CT): Highresolution
CT scanning allows the internal anatomy of fossil specimens to be
visualized non-destructively.
In this study, CT data were
used to digitally reconstruct
the endocranial cavity and
generate a virtual endocast
for neuroanatomical analysis.

Figure 2: Reconstruction sequence illustrates the skull, the digital endocast, and the reconstructed brain of Megantereon. Computed tomography (CT): High-resolution CT scanning allows the internal anatomy of fossil specimens to be visualized non-destructively. In this study, CT data were used to digitally reconstruct the endocranial cavity and generate a virtual endocast for neuroanatomical analysis.


THE THREE LEVELS OF KNOWLEDGE 

For this project, as with every occasion in which I find myself working as a paleoartist, I relied on three levels of classification of the available information and of the choices I would make throughout the reconstruction process. 


1–Observation 

The first level consists of recognizing and recording directly observable data, primarily consisting of the information contained within the fossil record. This includes details such as the shape of the skull, the body plan and skeletal proportions, muscle attachment sites, the configuration of the joints and the osteological correlates that allow us to infer the position of soft tissues such as the eyes, nose, and ears.


Regarding Megantereon cultridens, we are quite fortunate as the species in question has numerous fossil remains, including complete skulls and even an excellent nearly-complete skeleton known as “the Senèze skeleton,” discovered at the eponymous French site and now housed in the Basel Museum in Switzerland. With this abundance of fossil material, we had access to a wealth of information, forming the first level of knowledge. 


2–Inference 

The second level of knowledge consists of inferences—conclusions based on evidence and reasoning—namely all those pieces of information that are not directly observable in the fossil record, but deducible and reconstructable through parallel disciplines such as comparative anatomy, biomechanics, and the phylogenetic placement of the animal under investigation. Typically, most of the reconstruction of soft tissues falls within this second level.


Megantereon cultridens is once again a relatively fortunate case. Making inferences is considerably easier when living relatives or close biological equivalents are still present within modern ecosystems. Although Megantereon is a machairodont felid, belonging to a lineage that is now entirely extinct, it remains a large cat whose anatomical foundation, body plan, and muscle insertions and groups are, overall, quite similar to those of a jaguar or tiger—animals we can observe in nature and for which there are numerous studies on myology (the study of muscles) and biomechanics. If, by contrast, we were to reconstruct an animal with no modern bioecological counterparts, such as Stegosaurus, the inferential process would have to begin from a much more challenging foundation. 

3–Speculation 

Finally, we reach the third level, the most delicate one: speculation. This level is perhaps the most misunderstood aspect of paleoartistic reconstruction, precisely because it is the stage in which creative expression and imagination play the greatest role. This category includes all those characteristics that are not only unobservable in the fossil record, but whose inference also carries a very high degree of uncertainty. Examples include the coloration and chromatic patterns of a given animal, the structure and nature of its integument (such as hair length, the presence of hair or feathers, and scale size), the extent of its labial tissues, the exact shape of the rhinarium or external ear pinna, eye coloration, and, of course, everything related to behavior, vocalizations, and communication signals. 


Such information can only be inferred when exceptionally rare and well-preserved fossil material is available, such as mummified remains or fossils that preserve soft tissues. In most cases, however, these details are not deducible with absolute accuracy. The fact that certain physical and behavioral attributes remain unknown does not mean that all speculation is equally valid. Even at this third level of knowledge, reconstruction must be approached with caution and guided by our understanding of biological patterns and ecological principles. 


With regard to Megantereon cultridens and other organisms known only from the fossil record, this level presents the highest degree of uncertainty; however, since numerous living relatives of Megantereon exist, and given that the feliform group to which Megantereon belongs is vast and includes many species, it is possible (by comparing and analyzing modern species as analogs of extinct species) to form a plausible idea about aspects of this animal that are unknown. These include features such as the shape of the nose, the size and position of the external ear pinnae, eye coloration, pupil shape, hair length and coloration, and the patterns present on its coat. Furthermore, this comparative approach can allow speculations to be made regarding aspects of its ethology, such as hunting strategies, preferred habitats, and its trophic position within the predator guild of its ecosystem. 


BRAIN RECONSTRUCTION


At this point theory transitions into practice. In the following sections, I will describe the process I used to complete the commissioned work. 


Although the reconstruction of the brain was the most complex part of the project, it was the part in which I personally carried out the least amount of work. The reconstruction of Megantereon cultridens' brain was primarily conducted by the team of researchers from the University of Florence who, through computed tomography, mapped the endocasts of several Megantereon cultridens skulls, obtaining a fairly detailed approximation of the brain’s morphology as it would have appeared when the animal was alive.† My contribution to the project focused mainly on creating a realistic rendering of the brain and illustrating the position of the encephalon within Megantereon’s cranial cavity (Fig. 2).

 

For the sake of completeness, it is worth mentioning that the study of this extinct felid’s brain, compared with specimens of brains from several extant felid species (including lion, tiger, jaguar, leopard, snow leopard, clouded leopard, puma, wildcat, and Eurasian lynx) revealed that Megantereon cultridens possessed an encephalon characterized by two particularly well-developed regions: the frontal lobe, which is involved, among other functions, in motor dexterity and somatosensory specializations (such as manipulation and tactile perception), and the cerebellum, an area dedicated to spatial coordination, which in this case aligns with a lifestyle of climbing and a preference for environments with numerous obstacles, such as shrubland, wooded areas, and rocky terrains. A notable development of the visual cortex, located in the occipital lobe, suggests also that Megantereon likely possessed highly developed visual capabilities. 


In the dedicated infographic (Fig. 3), we compared the brain of Megantereon cultridens with those of a puma, jaguar, and tiger. These species display similar proportions and similarly developed regions, suggesting potential similarities and affinities in certain behaviors and habitat preferences among the four taxa. 


†  A comprehensive report of the materials and methods used to reconstruct Megantereon’s brain can be found in: Meli, I., Madurell-Malapeira, J., Rook, L., Michaud, M., Zoccante, S., & Bartolini-Lucenti, S. (2026). In the brain of a sabertooth: First neuroanatomical description and ethological insights on Megantereon cultridens (Felidae, Machairodontinae). The Anatomical Record, 1–24. https://doi.org/10.1002/ar.70213 


Figure 3: Comparative infographic of the brain of Megantereon cultridens
alongside those of three extant felids (the puma, jaguar, and tiger).

Figure 3: Comparative infographic of the brain of Megantereon cultridens

alongside those of three extant felids (the puma, jaguar, and tiger). The upper section highlights behavioral traits and inferred abilities associated

with particularly well-developed brain regions, including the frontal lobe and cerebellum.


LIFE RECONSTRUCTION 

This was undoubtedly the most extensive part of the project. It was not my first time working on Megantereon cultridens, in fact, this animal is a subject that I have studied for several years as both a paleoartist and an independent researcher. As a result, the body of knowledge and the amount of information, inferences, and reasoned speculations I had accumulated were quite substantial. This foundation allowed me to navigate confidently through the reconstruction of this animal.


In the following sections, I will discuss the reconstruction from three different perspectives: the head, the whole animal, and its behavior. 


The face of Megantereon 

Following a traditional paleoartistic approach, the reconstruction of the head (Fig. 4) begins with the skull. A fossil skull is not identical to the original bony structure of the living animal. Typically, fossil bone material has been altered by taphonomic processes. Therefore, in order to reconstruct original skeletal morphology, a process of retrodeformation is required. In the case of Megantereon cultridens, this process is relatively straightforward due to the abundance of available skull specimens and their overall similarity to those of extant felids. However, the skull of Megantereon is not exactly like that of a tiger. Even though it might seem obvious or trivial, the primary difference lies in the size of the upper canines. 


The upper canines of a tiger (Panthera tigris) measure, on average, approximately 7–8 cm from the tip of the tooth to the end of the root, whereas those of Megantereon reached roughly twice that length, measuring around 15–16 cm. The need to accommodate these exceptionally large teeth resulted in major changes to the skull’s proportions, representing the primary factor behind the distinctive differences between the head of Megantereon and that of a tiger. 




Figure 4:  Life reconstruction. Portrait of Megantereon cultridens.

Figure 4: Life reconstruction. Portrait of Megantereon cultridens.


The most notable difference between tiger and Megantereon skulls lies in the elevation of the nasal bone which was influenced by the enormous canine roots extending deep into the animal’s skull. As a consequence, the nasal stop (the “step” between the end of the rostrum and the beginning of the forehead, which is relatively pronounced in many extant felids) would have been much less evident in Megantereon. This difference in proportions also affected the orientation of the orbits and resulted in narrower zygomatic arches. If we could observe the head of a living Megantereon, we would immediately notice that, compared to a tiger, it would have had a less flattened face and a generally more elongated profile. 

A body built for the forest 

As with the reconstruction of the head, the reconstruction of the body must also begin with the available fossils. In this case, we can rely on the remarkable Senèze skeleton, which has a particularly well-preserved postcranial section. From the body plan of Megantereon, we can infer that (as corroborated by the study of its brain) it was likely an animal that preferred wooded environments, whether it be dense shrubs or a true forest. It is highly probable that this animal possessed good climbing abilities and could move effortlessly up trees and rocks. This conclusion is supported by its body proportions, short but strong and robust limbs which are proportionally similar to those of a leopard and jaguar and allow agile movement through thick vegetation or rapid scaling of tree trunks. The forelimbs, slightly shorter than the hindlimbs, suggest that Megantereon was likely an excellent jumper; a very useful trait for a felid hunting by ambush in a forested area.

 

What was the coat Like? 

One particular challenge in reconstructing the body was creating the coat pattern. Although color patterns obviously do not fossilize, for felids it is possible to make a reasonably accurate prediction of the range of colors and patterns that adorned their coats by considering factors such as habitat type, prey size and type, latitude, and climate. Taking phylogenetic implications into account as well, there is a strong possibility that Megantereon had a relatively short coat, with a yellowish or reddish base coloration and a spotted pattern composed of medium to large rosettes or blotches, similar to those we might observe in modern jaguars or clouded leopards. The pattern I created is not simply a “copy-paste” of any modern felid but was developed with careful consideration of the bio-ecological implications of pattern development in feliforms and behavior based on the Turing model or reaction-diffusion system. The resulting interpretation is realized in the two reconstructions presented in this article. 


Reconstructing behaviour and ecology 

In the final stage of the work, I was asked by the research team to depict a potential scene of life for Megantereon cultridens, in which not only could the abilities examined in the study be observed, but a reference to the habitat of this feline would be present. Together with Saverio, the study’s lead researcher, we chose to portray a hunting scene set in a subtropical forest typical of Mediterranean Europe during the transition from the Pliocene to the Pleistocene (Fig. 5). 


The scene depicts two Megantereon individuals bringing down a male Eucladoceros ctenoides, a large extinct deer roughly comparable in size to the modern sambar (Rusa unicolor). This species was likely among the predator’s primary prey, as Megantereon is thought to have preferred hunting medium- to large-sized deers. Along with the scientific basis of the reconstruction, the illustration incorporates a strong narrative component that is intended to convey the dynamics of the hunt. 


At first glance, it might appear to be a cooperative hunting scene, in which the two felids unite their efforts to bring down the prey in a manner reminiscent of modern lionesses hunting on the African savannah. In reality, however, the scene was designed to offer a dual layer of interpretation. The depicted hunt can be interpreted either as an episode of accidental gregariousness or as typical social behavior seen in lions. 


Figure 5: In a forest of Plio-Pleistocene Italy, a large male Eucladoceros ctenoides is attacked by a Megantereon.

Figure 5: In a forest of Plio-Pleistocene Italy, a large male Eucladoceros ctenoides is attacked by a Megantereon. The deer is wounded by the felid but manages to escape; the ungulate runs through the underbrush and crosses a stream, getting wet in the process. However, the Megantereon does not give up and continues chasing it in hopes of delivering a second attack. Believing itself to be nearly safe, the deer is suddenly ambushed by a second Megantereon concealed among the ferns on the opposite bank. The saber-toothed cat leaps onto the deer, driving its claws into the shoulder while using its body weight to bring the animal to the ground before delivering a fatal bite to the throat with its elongated upper canines


What do the details tell us? 

The first thing a viewer may notice is that two subjects are wet, while one is dry. Specifically, the deer’s lower body (feet, legs, and belly) is soaked, as if it had crossed a stream to escape predators. The Megantereon in the background has wet paws, shoulders, and belly, suggesting it had chased the deer through the stream to catch it. The Megantereon in the foreground, which has grabbed its prey, is completely dry, indicating, that instead of being in water, it waited, camouflaged among the vegetation on dry ground, for the deer. 


One interpretation concerns a potential cooperative behavior for the depicted felid species. Modern felids are mostly solitary as adults (outside of mating season), with the sole exception being the lion (Panthera leo) which lives in family groups composed of dozens of individuals. Although the lion is the only “social felid,” episodes of group living (i.e., two or three individuals, often siblings, spending a period, more or less long, of their lives together) have also been reported in other large felids such as the cheetah (Acinonyx jubatus), the puma (Puma concolor), and the jaguar (Panthera onca). In these three species, coalitions have been documented, where some individuals (often male siblings) spend several years of their lives together, often during the early stages of adulthood, after abandoning life with their mother. For Megantereon, however, there is no particularly solid evidence supporting social or group behaviors. Nevertheless, observations on traditionally solitary felids support the idea that cooperative behavior and coalition formation might not be far-fetched. 


An alternative interpretation is that the simultaneous attack is entirely coincidental. If Megantereon was indeed a strictly solitary predator, then the unfortunate Eucladoceros happened to encounter two lone hunters at exactly the wrong place and the wrong time, thus having a particularly bad day.


IN CONCLUSION


Although there may still be much to say about each part already addressed in this article, what has been written represents a solid compromise in terms of storytelling regarding the process and mindset to adopt when reconstructing an extinct species. More than simply illustrating an animal, paleoart is an exercise in integrating scientific evidence, comparative anatomy, and ecology with informed inference and speculation into a coherent and plausible reconstruction. From my perspective, I hope my work sparks curiosity and desire to delve deeper into learning more about Earth’s past—a body of knowledge that deserves to become part of the cultural repetoire of any naturalist or illustrator interested in paleoart.

Visual abstract of the study: “In the brain of a sabertooth: First neuroanatomical description
and ethological insights on Megantereon cultridens (Felidae, Machairodontinae)” by I.
Meli, J. Madurell-Malapeira, L. Rook, M. Michaud, S. Zoccante, and S. Bartolini-Lucenti.
This infographic visually summarizes the main concepts and results of the research.
This type of illustration is a standard practice in paleoart applied to scientific studies.

Visual abstract of the study: “In the brain of a sabertooth: First neuroanatomical description and ethological insights on Megantereon cultridens (Felidae, Machairodontinae)” by I. Meli, J. Madurell-Malapeira, L. Rook, M. Michaud, S. Zoccante, and S. Bartolini-Lucenti. This infographic visually summarizes the main concepts and results of the research. This type of illustration is a standard practice in paleoart applied to scientific studies. 



ABOUT THE AUTHOR 
Simone Zoccante is an Italian, freelance natural history illustrator, information designer, and paleoartist specializing in biology, zoology, paleontology, and anatomy. 
https://www.simonezoccante.com/

ABOUT THE AUTHOR 

Simone Zoccante is an Italian, freelance natural history illustrator, information designer, and paleoartist specializing in biology, zoology, paleontology, and anatomy. His work consists of creating visual representations of concepts, notions, and discoveries related to the world of natural sciences. A significant portion of his professial work reflects his focus and passion for paleoart—his self-identified specialty. He collaborates with publishers, museums, universities, and scientific institutions, creating illustrations, infographics, and visual communication projects for research, education, and exhibitions. His work has received international recognition, including the Italian Paleoart Prize (2024) and the prestigious John Lanzendorf-National Geographic Paleoart Prize (2025, 2-D Art category). Simone is also a writer and host of Diorama Nature, a web project dedicated to natural history. 

www.simonezoccante.com ∙ www.youtube.com/@dioramanature