Excavations conducted by the University of Pennsylvania at Phrygian Gordion have so far only revealed wall paintings from one small building. This structure was prominently located between two large Middle Phrygian megara at the outer court of the citadel and dates to the beginning of the Achaemenid period (ca. 500–490 BCE). The building was unique regarding both its decoration and architectural features. It was a semi-subterranean structure that consisted of one main chamber measuring about 3.50 × 4.75 m with a narrow antechamber in front. The mudbrick walls, once decorated with painted plaster, had collapsed during the Persian period, probably because the walls had been robbed of their supporting timber beams. Thousands of painted plaster fragments were recovered during the excavations of the late 1950s. This paper discusses the artistic style, techniques, and iconography of these paintings from an Anatolian perspective, and takes into account contemporary Etruscan wall paintings.

An expansion of the use of molecular data in the 21st C has accompanied a greatly increased number of amphibian and reptile species descriptions. Some of the newly discovered or recognised diversity has been considered ‘cryptic’. Islands and island endemic taxa represent worthwhile systems to address questions about cryptic species. Being comprised of multiple islands of partly Gondwanan origin, the Seychelles Archipelago offers the opportunity to address crypsis for a biota comprised of both deep-time vicariant and more recent overwater dispersed forms. We focus on the history of discovery, taxonomy, inter- and intraspecific variation, and phylogeography. Most recent studies investigating intraspecific variation in Seychelles herpetofauna have identified (often similarly) geographically structured, genetically divergent populations, and we note the potential of Seychelles herpetofauna in understanding lineage crypsis. We attempt to assess why some lineages have remained cryptic for so long; whether recently discovered lineages are truly cryptic and/or distinct taxa; and if ecology (vagility) and/or length of time on the islands can explain lineage crypsis. Patchy data and/or lack of sufficient study currently limit possible interpretations. Detailed ecological and environmental data, population genetic assessments, reconstruction of phylogeographic histories, and integration of phenotypic trait data will all aid future assessments.

The origin and early evolution of snakes has long been studied, but little research has focused on soft-tissue organs such as the brain. I report data from dissections and 3D reconstructions of the endocasts of diverse species, including the Cretaceous stem snake Dinilysia patagonica in order to provide a comparative evolutionary framework for the snake brain. Snakes are a special case among reptiles because the braincase almost entirely encloses the whole brain, so endocasts provide realistic representations of brain size and shape. Diversity of brain gross anatomy among snakes is remarkable, encompassing two major cerebrotypes occurring in surface-dwelling and burrowing species. The repeated acquisition of the burrowing cerebrotype in different and phylogenetically distant snake clades suggests that brain gross anatomy is surprisingly evolutionary labile in snakes. Brain gross anatomy and other features such as body size and the absence of any unequivocal osteological feature related to burrowing is interpreted as evidence that D. patagonica was surface-dwelling, and that at least some of the early history of snakes occurred above ground.

Sea-serpent sightings were popular subjects of nineteenth century fictional tales. One of the most famous sightings, the 1817 appearance in the harbor of Gloucester (Massachusetts), generated a report published by the Linnean Society of New England. In 1869, ED Cope introduced a new reptilian order, Pythonomorpha, comprising large Upper Cretaceous marine lizards (mosasaurs) that he thought rather well captured in historical depictions of sea-serpents. The name Pythonomorpha emphasized the many striking features that Cope found mosasaurs to share with snakes. Cope’s Pythonomorpha was resurrected in the late 1990s, as a clade including mosasaurs plus snakes. This was supported by the placement of mid-Cretaceous marine snakes with well-developed hindlimbs as evolutionarily intermediate between mosasauroids and snakes. Critics pointed to features indicating that those fossil snakes are instead evolutionarily advanced, which would imply that hindlimbs of these fossil snakes re-developed from rudiments such as occur in pythons. Recent molecular developmental studies confirmed that the embryonic limb bud of the python hindlimb conserves the genetic program to generate a complete limb.

We give a review of all published Palaeogene snake taxa from all localities worldwide. Several conceptual and material advances in the past two decades—a focus on apomo+P31rphies, greater attention to variation, quantification of morphology, and new fossil discoveries—have vivified the fossil record. Particularly noteworthy have been new fossils from Gondwanan continents and complete, articulated skeletons. Species known only from vertebrae are unlikely to be placed precisely phylogenetically, but a high number of vertebrae is a strong indication that cranial remains are present, which in turn allow more precise phylogenetic placement. Extrapolations of snake palaeodiversity are of the same order of magnitude as rough calculations of cumulative lineage diversity in the Palaeogene, raising the prospect that palaeontological morphospecies may more closely approximate biological species than is commonly conceived. As their interrelationships become better known, Palaeogene fossils will increasingly help elucidate the early evolution of snakes.

This brief chapter introduces the book. The rationale, scope, and coverage are summarized, including mention of topics that are not covered. Aspects of debate, disagreement, and consensus in the field are summarized before the chapter is concluded with a look to future potential progress.

The Cretaceous fossil record of snakes demonstrates the origin and evolution of the snake body and the early ecological and biogeographic history of the clade during the first 80 million years of their history. Consisting primarily of isolated vertebrae as well as a small number of mostly complete specimens, the record shows the elongate body of snakes evolved no earlier than approximately 100 million years ago. Stem snakes are present in terrestrial and marine palaeoenvironments throughout the Late Cretaceous of northern and southern continents. Conversely, the oldest records of living clades are restricted to the Campanian of South America, Africa, and possibly North America, which requires episodes of dispersal, or unrealistically long ghost lineages extending back to tectonically-mediated vicariance, to explain geographic distributions. Most Maastrichtian-aged stem and primary living clades extend into the Paleogene, indicating that the K-Pg extinction event had little visible effect on the evolution of snakes, whereas major diversifications of crown clades are constrained from middle Paleogene to Neogene.

Venom, a specialized form of poison, is actively injected by the venomous organism into its target animal to facilitate several quotidian functions. Over a hundred convergent origins of this remarkable functional trait, along with intricate mechanisms of venom delivery, have been documented across animals. Pinpointing the emergence of venom in squamate reptiles has important implications for understanding the evolutionary history of snakes, but it has been challenging. Several competing hypotheses have been put forth to explain the evolutionary origin of squamate venom, including assertions of single, dual and multiple origins. In this chapter, in addition to a summary of this ongoing dialogue, we provide an overview of ecology, composition, delivery mechanisms, and evolutionary models that explain the possible origin and diversification of venom in squamate reptiles.

This chapter assesses morphological characters proposed to support the Pythonomorph Hypothesis— a purported close relationship between snakes and mosasaurians. With an emphasis on early diverging (non-mosasaurid) mosasaurians and mosaurids, new morphological data (including from high-resolution CT) for well-preserved dolichosaurid and mosasaurid fossils are presented. Details of the skull and mandibles are interpreted as supporting the monophyly of Mosasauria as the proximal outgroup of Varanoidea, to the exclusion of snakes. However, mosasaurians do deviate from the typical varanoid condition in aspects of their infraorbital foramen, ventral part of the lacrimal and its relationship with the prefrontal, anterior ramus of the ectopterygoid and its contact with the maxilla and jugal, lack of plicidentine, and (at least in early diverging mosasaurians) anguinoidean tooth replacement. We consider most characters previously reported as supporting the Pythonomorph Hypothesis to be problematic, because of incomplete fossil preparation, artefacts of taphonomy, limited comparisons, misinterpretations of anatomy, incomplete taxon sampling, or inadequate character formulation and/or scoring.

Mosasaurian phylogenetics has been one of the most controversial topics in squamate systematics, with various studies and authors arguing in favor of a varanoid affinity (the Varanoid Hypothesis), a snake affinity (the Pythonomorph and Ophidiomorph Hypotheses) or only distant affinities to these lineages (the Stem-scleroglossan Hypothesis). We review the classification history of mosasaurians over the past two centuries, focusing on non-mosasaurid mosasaurians (dolichosaurs and aigialosaurs). A reappraisal is provided based on a new phylogenetic analysis. Our results clearly support the Varanoid Hypothesis. The Pythonomorph and Ophidiomorph Hypotheses are reviewed, and characters traditionally inferred to support these hypotheses are discussed and reinterpreted. Taxonomic sampling and fossil completeness likely play a major role—our (hopefully improved) phylogenetic hypothesis being based on denser taxon sampling and more complete character scoring resulting from additional studies, including the application of modern imaging techniques. Based on current data, our interpretation is that a particularly close phylogenetic relationship between mosasaurians and snakes can be rejected.

Squamata, the group that comprises lizards, snakes, and amphisbaenians, is the largest and most diverse major group of living reptiles. Although most recent estimates place their divergence from Rhynchocephalia in the early Triassic, the first unequivocal records of squamates currently date from the Middle Jurassic, and the first crown squamates, representing Scincoidea, Anguimorpha, and possibly Gekkota, are known from the Late Jurassic. The record then improves substantially in the Early Cretaceous, with squamate fossils recorded from most major continents and showing evidence of expansion into a diversity of specialist niches. Despite recent claims to the contrary, the earliest unequivocal snake fossils also date from this period, with fragmentary remains from the Aptian-Cenomanian of North Africa and North America, followed by a more substantial global record from the Cenomanian onwards. The re-interpretation of the Middle Jurassic to Early Cretaceous parviraptorids as snakes was based on a misinterpretation of the original material, as shown by new associated specimens from the Middle Jurassic of Scotland.

This chapter is concerned with Alexander in Egypt in both life and legend. Subjects discussed include his foundation of Alexandria, which became a new capital for Egypt on the Mediterranean coast, his expedition through the Libyan desert to Siwah, where the oracle’s recognition of the conqueror as son of Zeus-Ammon resonated in both Greek and Egyptian cultic terms, his acceptance as the pharaoh of Egypt, and finally, after his death in Babylon, his return for burial to Egypt, where his embalmed corpse and tomb in Alexandria became the centre of Ptolemaic ruler cult, a focal point for later visits of Roman emperors, and where the question of its actual location remains a source of continuing fascination and debate. In the accounts of classical historians, Alexander in Egypt is already variously presented; the historiography is as important as the history. From the start some specifically local legendary elements may be seen and over time the Romance or Legend of Alexander in its many different forms overshadows and surpasses any strictly historical account.

In this chapter I uncover the place of animals within imperial discourses. This repertoire of representations served to denigrate Burmese populations as being too close to animals but did so ambivalently. Imperial texts also revealed their authors’ barely sublimated desires for the colonized Other and their own close, emotional connections with non-human creatures.

This chapter examines how both Pindar and Aeschylus invoke snakes as an example of how bodies, no less than voices, are rendered multiform through the reenactment of choral performance.