Evangelos Vlachos

CONICET and Museo Paleontológico Egidio Feruglio

Turtles of the clade Pan-Testudinoidea have a rich fossil record in North America, including the Caribbean, ranging from the late Paleocene to the Holocene. All earlier reports cannot be substantiated herein. The earliest members of this clade probably immigrated in multiple waves from Asia. Current phylogenies of crown Testudinoidea recognize four...

The application of network methodology in anatomical structures offers new insights on the connectivity pattern of skull bones, skeletal elements and their muscles. Anatomical networks helped to improve our understanding of the water-to-land transition and how the pectoral fins were transformed into limbs via their modular disintegration. Here, we...

Background In a recent work I transformed a complex and integrated text like the International Code of Zoological Nomenclature into a network of interconnected parts of text. This new approach allowed understanding that a continuous body of text cannot accurately reflect the true structure of the Code, and provided a scientific methodology to ident...

Turtles are among the most common and easily identifiable elements of the vertebrate fossil record, especially in continental deposits. In Greece, turtle fossils have been identified in at least 68 continental and insular localities, ranging from the Early Miocene to Holocene deposits. The review of the known turtle fossil record from Greece reveal...

The Late Miocene vertebrate fauna from the Entre Ríos Province, Argentina, comes from two units: the marine Paraná Formation and the Lower Member of Ituzaingó Formation (LMIF). This latter includes a lower conglomerate set, with fine quartz gravel, and clay and chalcedony clasts, and fossil remains, traditionally known as ‘Mesopotamiense’ or ‘Congl...

The Late Miocene (Messinian) Gargano “Terre Rosse” in southern Italy has yielded a diverse and peculiar insular vertebrate fauna, comprising an array of endemic taxa, including forms characterized by extreme sizes and unique morphologies. We here present a new testudinid from this insular assemblage. The material comprises few shell remains (both c...

After the end-Triassic extinction, parvipelvian ichthyosaurs diversified and became dominant elements of marine ecosystems worldwide. By the Early Jurassic, they achieved a thunniform body plan that persisted for the last 100 m.y.a of their evolution. Diversification and extinctions of thunniform ichthyosaurs, and their swimming performance, have b...

The use of horns and frills for sexual display and attack has been proposed and demonstrated in extinct taxa on several occasions, with the ceratopsian dinosaurs as the most iconic example. Niolamia argentina is a large meiolaniid turtle from Patagonia, characterized by the presence of extensive frills and massive horns in the skull. Here, Finite E...

Chelydropsis is a genus of the clade Pan-Chelydridae, which existed and was distributed in Eurasia from the Eocene to the end of the Pliocene. The Chelydropsis fossil record of North of the Alps is particularly rich throughout the Miocene. In this study, we describe a new Chelydropsis species from the Late Miocene (Vallesian, MN9) fossiliferous sit...

After the success of the first two editions of the Palaeontological Virtual Congress in 2019 (first PVC) and 2021 (second PVC; Crespo & Manzanares 2019; Crespo & Citton 2021), we have decided to try to replicate the success with a third meeting of the PVC (Fig. 1). The appearance of new applications and technological advances has played a crucial r...

We describe a new species of a giant tortoise of the genus Titanochelon from the locality of Sandelzhausen in south Germany (MN5, Burdigalian/Langhian boundary, Early/Middle Miocene). The material comprises at least two different individuals, one of which is a male individual preserving large parts of the carapace and plastron and several appendicu...

With their particular body plan within amniotes and their amazing fossil record, turtles represent a great interest for both neontologists and paleontologists with a strong anatomical background. The Turtle Evolution Symposia are regular international meetings that gather scientists working with different aspects related to the evolutionary history...

The emergence of new applications and technological advances has opened new and wider pathways for people to ensure effective science communication. This has become more significant after more than two years of difficulties and restrictions due to the COVID-19 epidemiological crisis, in which online platforms gained great relevance and proved key t...

Supplemental Data 1 Detailed Descriptions

The Isle of Crete contains an important endemic fossil fauna that has been studied extensively in the past. An endemic fossil tortoise, Testudo marginata cretensis Bachmayer, Brinkerink & Symeonidis, 1975, had previously been named from Pleistocene cave deposits in the island, presumably being larger than the mainland species and having some import...

Previously, only one small‐sized species of Testudinidae (Chelonoidis gringorum) was named from Lower‐Middle Miocene of Central Patagonia. In this short article, we describe a new large Testudinidae, here named Chelonoidis meridiana sp. nov. This large new species (carapace up to 80 cm) differs from other large species of the southern South America...

Although the existence of fossil turtles in Makrygialos and the surrounding areas, on the west side of the Thermaikos Gulf in Northern Greece, has been known for almost two decades, it has not been studied so far in detail. Over the last 5 years, we have conducted systematic fieldwork and exploration, focusing on the discovery and recovery of fossi...

Archaeological investigations carried out in the cave Zubbio di Cozzo San Pietro, Bagheria, Sicily, revealed the presence of a few skeletal elements of a large-sized tortoise in a funerary area dating to the Copper/Bronze Age. The tortoise has been AMS-dated revealing an age of 12.5 ± 0.5 kyr BP and therefore it pre-dates the funerary activities. T...

Nexus File

The early–middle Miocene continental Cerro Boleadoras Formation (CBF) crops out in the area of Cerro Boleadoras and Cerro Plomo on the western slope of the Meseta del Lago Buenos Aires, northwestern Santa Cruz Province, Argentina. The lower levels of the CBF consist of laterally extensive medium to pebbly sandstone beds with trough cross-bedding, i...

Background Along with the transition to the aquatic environment, cetaceans experienced profound changes in their skeletal anatomy, especially in the skull, including the posterodorsal migration of the external bony nares, the reorganization of skull bones (= telescoping) and the development of an extreme cranial asymmetry (in odontocetes). Telescop...

The genus Testudo was considered, for more than two centuries, to group almost all tortoises that are part of current biodiversity (i.e. the members of Testudinidae), as well as the extinct representatives of that successful lineage. However, only three extant species, as well as scarce late Miocene to late Pliocene extinct ones, are now considered...

From the many post-Oligocene basins in Greece that contain vertebrate fossils, we focus on three major basins with their accompanied minor ones, which present synthetic and complicated processes of geotectonic and stratigraphic evolution. These are the Mesohellenic (MHB) with Ptolemais (PTB), Axios-Thermaikos (ATB) with Mygdonia (MB), and Strymonik...

The sedimentary basins of Greece contain an important fossil record of vertebrates that is known and studied for nearly two centuries. Vertebrate paleontological research in Greece has passed through three main stages: the Early Stage until the end of World War II, the Intermediate Stage until the end of the twentieth century, and the Modern Stage...

This 2-volume set provides a state-of-the-art study of the fossil record and taxonomy of the main vertebrate groups from Greece. Greece stands between 3 continents and its vertebrate fossil record is of great importance for paleontological and evolutionary studies in Europe, Asia and Africa. Fossils from classic, world-famous localities (e.g., Pike...

The extinct herpetofauna of the Chubut Province is one of the most diverse, temporally and spatially extensive, and well-known extinct faunas in Argentina and South America. These fossils help understanding the evolution of the herpetofauna during more than 180 million years, not only in the Patagonian region, but also in a worldwide scale due to t...

The sedimentary basins of Greece contain an important record of fossil vertebrates that has been known and studied for nearly two centuries. Here, we present our collective effort to review and summarize this fossil record. A combination of our original research and previously published records permits the complete reassessment of the identified ve...

The sedimentary basins of Greece contain an important fossil record of vertebrates that is known and studied for nearly two centuries. Vertebrate paleontological research in Greece has passed through three main stages: the Early Stage until the end of World War II, the Intermediate Stage until the end of the twentieth century, and the Modern Stage...

From the many post-Oligocene basins in Greece that contain vertebrate fossils, we focus on three major basins with their accompanied minor ones, which present synthetic and complicated processes of geotectonic and stratigraphic evolution. These are the Mesohellenic (MHB) with Ptolemais (PTB), Axios-Thermaikos (ATB) with Mygdonia (MB), and Strymonik...

Novel phenotypic configurations can profoundly alter the evolutionary trajectories of species. Although innovation can precede lengthy periods of evolutionary stasis, the potential for species to diversify further can be realized via modular changes across distinct levels of hierarchical organization. To test this expectation, we undertook anatomic...

The Late Miocene (MN9) locality of Höwenegg is well known for its diverse faunal assemblage during a key time period of the Neogene. Several articulated and nearly complete skeletons of mammals, fishes and a large amount of turtle specimens have been excavated in the past decades. Four different species of tortoises and freshwater turtles from at l...

The sedimentary basins of Greece contain an important record of fossil vertebrates that has been known and studied for nearly two centuries. Here, we present our collective effort to review and summarize this fossil record. A combination of our original research and previously published records permits the complete reassessment of the identified ve...

In this work we focus on the fossil record of turtles and tortoises from the Chubut Province, in Patagonia, Argentina. This record is the richest, most diverse, the longest and continuous in the country and one of the most important in the continent. In this work, we present and study new fossils from all known clades of turtles from the province,...

Díez Díaz, V.; Belvedere, M.; Böhmer, C.; Bueno, E.; Choiniere, J.; Darlim, G.; Drozdz, D.; Iannucci, A.; Kotthoff, U.; Malafaia, E.; Mallison, H.; Marigó, J.; Miedema, F.;Mujal, E.; Pardo, J.; Perillo, M.; Sciscio, L.; Tajika, A.; Tschopp, E.; van Heteren, A.H.; Vlachos, E. 2021. How to bring taxonomy into the third dimension: developing guideline...

Recently I have developed a new approach to use and understand codes of nomenclature by transforming them into networks. Conceptualizing a code as a network provides more efficient navigation and learning, and is a powerful tool for amendments and revisions. Also, this methodology allows the objective comparison between different codes in terms of...

Turtles are among the most common and easily identifiable elements of the vertebrate fossil record, especially in continental deposits. In Greece, turtle fossils have been identified in at least 68 continental and insular localities, ranging from the early Miocene to Holocene deposits. The review of the known turtle fossil record from Greece reveal...

Stylemys nebrascensis is a testudinid tortoise from North America that lived in subtropical areas during the Paleogene, being one of the anatomically best-known species within extinct turtles. The specimen at the Raymond M. Alf Museum (USA), RAM 8315, corresponds to an almost complete skull with articulated jaws, which were virtually removed — toge...

In this response to Dubois (2020) I clarify some allegations directed personally to me therein and I present my opinion and recommendations on the availability of names created in papers that are later retracted.

The world-famous upper Miocene fossil localities on the Aegean island of Samos in Greece have produced a rich fossil record that sheds light on the evolution of eastern Mediterranean terrestrial faunas over a one-million-year interval of the late Neogene. Fossils have been discovered on Samos since antiquity, although a succession of paleontologica...

This contribution presents a complete revision of the extinct geoemydid Echmatemys from North America, based on new information and a re-evaluation of previously published specimens. Echmatemys is the most diverse and widely distributed geoemydid turtle in the fossil record of North America, known from numerous shells. The different morphotypes are...

This article deals with the fossil tortoises of one of the most iconic fossil localities of the Neogene of the Old World, the upper Miocene locality of Pikermi, near Athens, Greece. We describe the type, previously published, and new material of the fossil tortoises from Pikermi, along with new material from the coeval Azmaka 6 locality in Bulgaria...

Herein, we describe Alatochelon myrteum gen. et sp. nov., a large tortoise from the post‐Messinian (lower Pliocene) of the area of Puerto de la Cadena (Region of Murcia), Spain. The new taxon cannot be attributed to Titanochelon, which represented the only lineage of large tortoises previously recognized in the Neogene record of Europe. Alatochelon...

The collections of the Museum of La Plata contain the holotype (MLP 26-400) of a giant fossil tortoise, Chelonoidis australis (Moreno, 1889), from the early Pliocene (Montehermosan SALMA) of the Atlantic coastline between Monte Hermoso and Punta Alta, Buenos Aires Province, Argentina. With this contribution I present my revision of this taxon, and...

Background In order to designate the various concepts of taxa in biology, evolution and paleontology, scientists have developed various rules on how to create unique names for taxa. Different Codes of Nomenclature have been developed for animals, plants, fungi, bacteria etc., with standard sets of Rules that govern the formation, publication and ap...

One of the most iconic tortoise species, coming from the Sivalik Hills in India and probably the largest tortoise that ever lived, has a complicated nomenclatural history. Although initially published as Megalochelys sivalensis, the same authors decided to replace the name a few years later as Colossochelys atlas, starting a discussion on the prope...

The palaeoecology of the basal turtle Proganochelys quenstedti has been open to debate. Whereas it was firstly described as semi-aquatic/aquatic, anatomical and histological data support adaptation to a terrestrial habitat. Our objective is to explore whether the palatal denticles of basal turtles could contain evidence of dental microwear. Two Pro...

I perform the first network analysis on the shell of turtles (Testudinata), using representatives from all major clades of extant and extinct turtles. Their shell is modeled into four networks (bony plates and horny scutes of carapace and plastron). The overall properties of the entire networks, as well as of the various nodes (= plates and scutes)...

Analyses of diversity and extinction are of key importance in many paleontological works. Usually assisted by the construction of diversity curves of taxa at different taxonomical levels (species, genera, families, or clades), diversity analyses require the concentration, analysis, and presentation of a large amount of data. For this reason, the ex...

The abundance and richness of the fossil record place Milia as the richest Pliocene locality in Greece and among the richest in Europe, dating back to a crucial period for the faunal evolution in the Northern Hemisphere. During the initial period of research, the paleontologists were able to confidently identify the majority of the species that liv...

Turtles were always thought to have been unaffected by major extinction events that occurred on Earth, like the Cretaceous–Paleogene one that wiped out the non-avian dinosaurs. This thought was mainly based on genera counts across the K-Pg boundary. The unprecedented growth of the South American turtle fossil record allows for a thorough analysis o...

To describe the various concepts of taxa in biology, evolution, and paleontology, scientists have developed various rules on how to create unique names for taxa. Zoological nomenclature is governed by the International Code of Zoological Nomenclature, a standard set of rules that govern the formation, publication, and application of the nomina of e...

Extinct turtles associated to marine environments in Argentina have a discontinuous record since the Late Jurassic to the Miocene. These turtles adapted to marine environments belong to different clades: the extinct Platychelyidae (Pan-Pleurodira) and Thalassochelyidia (Pan-Cryptodira), and the extant Chelonioidea (formed by two clades: Dermochelyi...

The fossil record of turtles of the clade Testudinoidea is generally poor in South America. Fossil tortoises (Testudinidae) are known from the Late Oligocene to Holocene, but they likely arrived from Africa during the Late Eocene or Early Oligocene. The fossil record of Trachemys (Emydidae) and Rhinoclemys (Geoemydidae) is restricted to the Pleisto...

Recent advances in the fossil record, anatomy, and evolutionary history of South American turtles allow a thorough analysis of their changes in diversity, as well as to identify several major extinction events. The history of turtles in South America starts with stem turtles surviving in the southwestern margin of the Pangea. With the onset of the...

Our knowledge of Neogene chelonian diversity in northern Greece is increased with the present description of a new species of Mauremys (Testudines, Geoemydidae) from the late Miocene to Pliocene of three localities in central Macedonia (Gefira‐2, Nea Silata, Allatini). This new species, Mauremys aristotelica sp. nov., is characterized by the presen...

This work is meant as popular and scientific communication work to trigger a wider audience about Zoological Nomenclature. In the World of Ice and Fire, a Maester is looking to forge his weirwood link in his chain by studying animals in nature. He soon realizes that there is no system to allow clear communication between scholars and Maesters acro...

1) Background: Over the last decades, important fossil records of Late Pliocene mammals and reptiles have been unearthed in Milia, Grevena (N Greece). This assemblage shows a remarkable composition and diversity, including the partial skeletons of mastodons that reached record-breaking sizes, abundant rhinos-the first occurrence of this species in...

One of the named species from the South American clade Chelonoidis and the only one from Uruguay is Testudinites sellovii Weiss, 1830. The material consists of isolated plates that come mainly from the plastron, and originates from the Pleistocene deposits of Paso del Catalán, near river Queguay in the area of Paysandú (NW Uruguay). In this note, w...

The mining activities in the Middle Pleistocene lacustrine basin of Megalopolis (Peloponnesus, Greece) have exposed expanded sections of lacustrine sediments. In particular, the northernmost mine of Kyparíssia has yielded numerous vertebrate fossils, recovered during field surveys and small-scale rescue excavations. The stratified specimens indicat...

Despite their global distribution, substantial diversity, and rich fossil record, the evolutionary history of Testudinidae (tortoises) is poorly understood. We here present the first total evidence analysis of Pan-Testudinidae combining previous and novel morphological datasets with fossil data and published molecular information. Parsimony analysi...

Chelonoidis gringorum (Simpson, 1942) is a medium-sized testudinid taxon known mainly from the Early-Middle Miocene of Patagonia. Most fossil specimens come from the deposits in the area of Trelew – Gaiman – Dolavon (Chubut Province, Argentina). Besides the type material (a partial shell), several specimens have been referred to Ch. gringorum over...

Testudinidae (tortoises) is an extant clade of terrestrial turtles of worldwide distribution and with a rich fossil record that provides an exceptional context for studying their evolutionary history. Because of the lack of global phylogenetic analyses integrating extinct taxa, our current knowledge of the relationships of the total clade of Testud...

The presence of remains of a giant tortoise in the lower Pleistocene site of Fonelas P-1 (Guadix Basin, Betic Ranges; Granada, southeastern Spain) is reported and analyzed herein for the first time. This finding represents the youngest evidence of a large tortoise in continental Europe, dating the age of extinction of this successful lineage as sev...

Fossil amphibians and reptiles from the earliest late Miocene (early Tortonian, MN 9) of Plakias (Crete, Greece) are described in this paper. Most of the material is fragmentary, precluding precise taxonomic assignment. Nevertheless, the herpetofauna of Plakias is here shown to be diverse, comprising at least six different taxa: an alytid anuran, a...

The information about the European giant fossil tortoises has been greatly increased in recent years, based on the description of new material and the revision of previously published specimens. A genus with a wide paleobiogeographic and stratigraphic distribution, Titanochelon, has recently been described, containing all large testudinids from the...

The use of fossils in tree calibrations has been rigorously debated, mainly as the result of the uncertainty of the phylogenetic position and age of extinct taxa. Also, the addition of fossil information can be done with various methods, reaching sometimes significantly different results. Here we present our methodological comparative study in whic...

Synchrotron radiation micro-X-Ray Fluorescence (-XRF), X-ray photoelectron (XPS) and X-ray Absorption Fine Structure (XAFS) spectroscopies are applied for the study of paleontological findings. More specifically the costal plate of a gigantic terrestrial turtle Titanochelon bacharidisi and a fossilized coprolite of the cave spotted hyena Crocuta c...

New material of fossil amphibians and reptiles from the earliest late Miocene (early Tortonian, MN 9) of Plakias, Crete, Greece, are described. Despite the fragmentary nature of the fossil specimens, the herpetofauna of Plakias is shown to comprise at least six different taxa: one alytid anuran, one crocodylian, two turtles (one pan-­‐trionychid an...

Ever since the times of Linnaeus, the use of Latinized Greek names for naming the present and past diversity of our planet has been a common practice. This contribution focuses on the use of Greek words in forming taxon names, as exemplified by names from chelonian literature. The current problems of the guidelines of the International Code of Zool...

The late Pliocene (early Villafranchian, MN 16a) locality of Milia (Grevena, Greece) has yielded numerous remains of mammals, such as zygodons, mastodons, rhinocerotids, hipparions, bovids, cervids, suids, carnivorans and tapirs. Several specimens of chelonians have also been discovered and comprise a small testudinid, a geoemydid and a giant torto...

Soft-shelled turtles (Pan-Trionychidae) are not included in the present-day chelonian fauna of Greece and have been unknown in the Greek fossil record up to now. Here, we report the first fossil occurrence of a soft-shelled turtle from Greece, originating from the Pliocene Gefira Member (Angelochori Formation), in the lower Axios valley. The corres...

During recent years, our knowledge of fossil turtles from northern Greece has improved, as a result of both the discovery and description of new specimens and the revision of previously published material. The sites of Allatini and Pylea are among the classic Latest Miocene–Earliest Pliocene localities of northern Greece and have produced notable c...

Background. Large-sized testudinids had a long evolutionary history in Europe during the last 50 million years before becoming extinct in the beginning of the Pleistocene. Despite a 150-year long history in research and a decent fossil record, the European large testudinids have received limited attention so far. Methods. New excavations, descripti...

Background. Regardless using a rank-based or a phylogenetic nomenclature code, the use of Latinized binomens to describe the extant and extinct species has been essential. Ever since the times of Linnaeus, the use of Latinized Greek names has been a common practice both for neontologists and paleontologists. Methods. I critically analyzed the most...

The chelonian fauna of Sicily is currently limited to a single autochthonous tortoise, Testudo hermanni hermanni, whose shell length reaches approximately 20 cm. Even if the Balkan subspecies of the Hermann's tortoise, Testudo hermanni boettgeri, is significantly larger (up to 35 cm), the Sicilian Pleistocene fossil record testifies for the presenc...

Nel maggio 2014, durante lo scavo archeologico di una sepoltura multipla all'interno della grotta Zubbio di Cozzo San Pietro (PA), attribuito, secondo la cultura materiale, al Bronzo Antico (metà III mill. a.C.), sono stati rinvenuti reperti faunistici di grandi e piccoli vertebrati. Particolarmente interessanti sono i resti di una testuggine di di...

Background. Regardless using a rank-based or a phylogenetic nomenclature code, the use of Latinized binomens to describe the extant and extinct species has been essential. Ever since the times of Linnaeus, the use of Latinized Greek names has been a common practice both for neontologists and paleontologists. Methods. I critically analyzed the most...

Background. Large-sized testudinids had a long evolutionary history in Europe during the last 50 million years before becoming extinct in the beginning of the Pleistocene. Despite a 150-year long history in research and a decent fossil record, the European large testudinids have received limited attention so far. Methods. New excavations, descripti...

Testudinidae are terrestrial turtles currently living on several continents, including Europe. Fossil testudinids are known in Europe since the Eocene, with several taxa of medium size (from 0.3 to 0.7 m) recognized in the Paleogene record. The size of several European Neogene taxa was larger (between 1 and 2 m). These large testudinids were abunda...

Fossil testudinids are known in Europe since the Eocene, with several taxa of medium size (from more than 0.3 m to less than 0.7 m) recognized in the Palaeogene record, most of them being poorly known. The size of several European Neogene taxa was larger (between 1 and 2 m). These large testudinids were relatively abundant and diverse, ranging from...