Determining the contents of bronze jars from an Ancient Greek shrine in southern Italy.

In 1954 archaeologists excavated a shrine to an unknown deity at the Greek settlement of Paestum in southern Italy, which was dated to the sixth century BC. Within this shrine they found six bronze hydriai (storage jars) and two amphorae, arranged around a large iron bed. The jars contained a pasty residue with a wax-like aroma. Traces of this substance were also found on the outside of the jars, which were originally sealed with cork, leading the archaeologists to conclude that it was originally a liquid, although possibly a fairly viscous one. It was interpreted that the bed was intended to be the residing place of the unknown deity, with the contents of the hydriai and amphorae were intended as offerings. 

Honey, in the Greek and Roman worlds, was a substance of some significance. As the only practical way to sweeten food and drinks, it was economically important, but it also had spiritual significance, being associated with wisdom and immortality, and therefore a suitable offering to the gods. With this in mind, it seemed highly likely that the original contents of the Paestum hydriai was honey.

With this in mind, the Bee Research Association in London arranged for an analysis of the residue to be carried out. The substance was found to be insoluble in water, but soluble in organic solvents, and to contain trace amounts of Plant and Insect remains, Fungi, and pollen, which led the scientists carrying out the analysis to conclude that it was probably a remnant of the wax which had originally sealed the jars.

In 1970 scientist at the Istituto Centrale del Restauro in Rome carried out an analysis on residues found around the neck and in the bottom of one of the amphorae from Paestum. This was found to comprise a saponifiable substance (substance which will react with an alkali to form a soap), such as a wax, fat, or resin, but contain no detectable sugar or protein (the major components of honey).

The residue was tested again in 1983 by the Laboratory of the Rome Chamber of Commerce, who again found that it was a saponifiable substance insoluble in water but soluble in organic solvents, and did not contain any sugary or starchy substances. On this occasion gas chromatography was also used to analyse the residue, concluding that it was 77.4% palmitic acid, 6.1% oleic acid, 5.2% stearic acid, 1.0% heptadecanoic acid, 1.0% linoleic acid + arachidic acid, 0.4% linoleic acid, and 6.5% unidentified substance. Since triglycerides of palmitic acid are extremely common in nature, the researchers concluded that the container had held animal fat or a vegetable oil.

In 2019, the residue from the Paestum hydriai was loaned to the Ashmolean Museum in Oxford, for an exhibition, 'Last Supper in Pompeii', and permission was obtained to carry out a new analysis of the biomolecular composition of the substance, using modern equipment not available when the previous tests were carried out.

In a paper published in the Journal of the American Chemical Society on 30 July 2025, Luciana da Costa Carvalho and Elisabete Pires of the Mass Spectrometry Research Facility at the University of Oxford, Kelly Domoney of the Ashmolean Museum, Gabriel Zuchtriegel of the Parco Archeologico di Pompei, and James McCullagh, also of the Mass Spectrometry Research Facility at the University of Oxford, present the results of this new analysis, and confidently identify the original material within the Paestum hydriai.

(A) Underground shrine in Paestum. (B) One of the hydriai on display alongside a Perspex box containing the residue at the Ashmolean Museum in 2019. (C) Graphic representation of the arrangement of the bronze jars inside the shrine. (D) Sample from the core of the residue.  Carvalho et al. (2025).

The residue arrived at the Ashmolean Museum in a non-hermetically sealed Perspex box, in which it had apparently been displayed at the Paestum Museum. In order to reduce the chances of modern contamination affecting their results, Carvalho et al. took samples from 40 mm below the surface for analysis, as well as from each distinct colour zones observed on the exterior of the material; black, orange, and green, colours which suggest some sort of interaction with the bronze vessel itself over the past 2500 years. Carvalho et al. also obtained modern beeswax, honey, and honeycombs from locations in Italy and Greece in order to compare these to the residue sample.

Carvalho et al. first used Fourier Transform Infrared Spectroscopy to obtain an overview of the chemical substances present within the sample. This yielded a spectrum almost identical to that of modern beeswax for the interior sample, strongly supporting the idea that this was the original substance. They also compared the spectra of new and artificially aged honeycombs from both Greece and Italy, establishing that there was little difference in these, and that these structures appear to be chemically stable at least 

Fourier Transform Infrared Spectrum  of the core sample of the archaeological residue superimposed on beeswax (a) and honey’s (b). Carvalho et al. (2025).

Next, Carvalho et al. carried out a Gas Chromatography coupled to Quadrupole Time-Of-Flight Mass Spectrometry analysis of the sample, along with samples of honey, beeswax, and fresh and artificially aged honeycomb. This produced almost no results for the beeswax, suggesting that the bulk components of this material were broken down by the high temperature at which this method operates (over 300°C), but did produce results from the sample, as well as from the honey and honeycomb controls, suggesting that the sample was never pure beeswax.

Electron ionisation chromatograms from the Quadrupole Time-Of-Flight Mass Spectrometry analysis of beeswax (a), the residue core sample (b), honey (c), and honeycomb from Greece, fresh (d) and aged (e). Compounds identified: (1) Hexadecanoic acid, (2) Heneicosane, (3) Octadecanoic acid, (4) Pentacosane, (5) Heptacosane, (6) Nonacosane, and (7) Hentriacontane. Carvalho et al. (2025).

Anion exchange Ion-Chromatography coupled to Mass Spectrometry identified seven hexose sugars, hexadecanoic acid, heneicosane, octadecanoic acid, pentacosane, heptacosane, nonacosane, and hentriacontane within the sample at levels higher than would be expected in beeswax, but lower than would be expected in honey. It also found significant levels of the sulphur amino acid taurine, which was not present in any of the control samples.

These hexose sugars were also recovered from aqueous extracts of honeycomb (i.e. the liquid obtained by soaking honeycomb in water)along with gluconolactone (a derivative of glucose) and galacturonic acid, and low levels of succinic, malic, and citric acids. All of these compounds were yielded at higher levels by the fresh honeycombs than by the artificially aged honeycombs.

Finally Carvalho et al. used a proteomic approach to try to identify specific proteins within the sample, as well as the beeswax and honeycomb controls, which were compared to the UniProt All Proteins database. The reesidue sample produced matches for three proteins derived from the royal jelly produced by the Western Honeybee, Apis mellifera, several Bacteria-derived proteins, and a number of common contaminant proteins, including keratins, caseins, lysyl endopeptidase, and trypsin. Encouraged by this, they then compared the sample to the Bee-specific UniProt Honey database, a search which yielded eight matches, including some associated with the Eastern Honeybee, Apis cerana cerana,

Proteins were also recovered from the modern honeycomb samples, but not the beeswax, indicating that they were derived from the honey portion of the comb. Notably, the royal jelly protein signature from the Greek and Italian honeycombs was quite different, although this was not completely unexpected as the two looked different. Carvalho et al. note that factors such as climate and the floral sources from which nectar is obtained can affect protein expression in Bee products, so this difference does not necessarily mean the Bees were particularly different.

Finally, a sample of the surface residue showing orange, black, and green colouration was subjected to X-ray photoelectron spectroscopy. This determined that the green areas of this residue were composed of 74.98% carbon, 20.78% oxygen, and 4.24% copper, with the copper portion largely dominated by Cu²⁺ ions, while the black areas were composed of 77.96% carbon, 20.12% oxygen, and 1.92% copper, with the copper dominated by Cu⁺ ions, and the orange areas were comprised of 86.50% carbon and 13.50% oxygen. The discoloration in these areas is, therefore, presumed to be derived by interactions between the original substance and the copper portion of the bronze vessel.

Honey is comprised primarily of sugars (typically 79% of the total mass, including 39% fructose), along with water (typically 18% of total mass), acids (0.17-1.17% of total mass) and trace amounts of other substances, such as vitamins, enzymes, flavonoids, and phenolic compounds. Over time this mixture undergoes Maillard reactions ('browning') as the amino acids of the proteins react with the sugars. This will occur more rapidly if the honey is stored at a warmer temperature. Eventually, the honey will take on a dark hue, as the sugars break down into furans and the acid content rises.

Previous studies of the Paestum hydriai residues concluded that this was a wax, most likely used to seal the vessels. The most commonly use wax in the ancient world was beeswax, a substance with quite different properties to honey. Beeswax is typically comprised of about 64% esters, 14% odd medium chain alkanes, 12% free acids, 2% acid polyesters, 1% acid monoesters, 1% free alcohols, and 6% other materials. Beeswax is much more stable than honey, but over time the acid and alcohol contents will increase as the was esters hydrolyse and the shorter chain alkanes are eliminated. 

The Fourier Transform Infrared Spectrum obtained by Carvalho et al. yielded results very similar to beeswax, suggesting that this may have formed a significant portion of the material from which the residue was derived. However, the Gas Chromatography coupled to Quadrupole Time-Of-Flight Mass Spectrometry analysis carried out suggested that the substance could not be pure beeswax. A study of the proteins present within the sample found several associated with honey production in the Western Honeybee, Apis mellifera, with a subsequent search of the Bee-specific UniProt Honey database yielded proteins associated with the Eastern Honeybee, Apis cerana cerana. Also produced were proteins associated with the wood-decay Fungus Armillaria gallica, and the parasitic Mite Tropilaelaps mercedesae, which targets Honeybees.

Eastern and Western Honeybees are closely related, and even where divergence has occurred, their proteins tend to be very similar. Furthermore, there is ample evidence for the cultivation of the Western Honeybee in ancient Italy, but none for the Eastern Honeybee. For these reasons, Carvalho et al. conclude that the readings suggesting the Eastern Honeybee as a source of proteins are probably erroneous, caused by the software trying to 'best fit' ancient degraded proteins.

The presence of proteins associated with the Mite Tropilaelaps mercedesae is also interesting. This Mite originated in Asia, and has a long historic connection with the Eastern Honeybee, but is generally thought only to have begun to infect Western Honeybees in the past few centuries. Carvalho et al. observe that it would be tempting to interpret this as evidence that the material at Paestum originated in Central Asia, but that a more likely scenario is that the proteins in question are common to a range of Acarid Mites.

Carvalho et al, conclude that a bulk composition similar to beeswax combined with the presence of proteins and other molecules found in honey make it likely that the material within the Paestum hydriai was almost certainly a honeycomb. This matches well with ancient literature, which frequently cites honey and other Bee-products as being suitable offerings for the gods, but contradicts earlier studies which were unable to obtain this result. Carvalho et al. emphasise that this underlines the importance of revisiting samples which have previously been analysed with less modern techniques, thus allowing our imptoving technology to improve our understanding of the past.

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Identifying the skins used to make Scythian leather.

The term 'Scythian' has been used to describe a vast array of nomadic people's living on the Eurasian steppes in the first millennium BC, who played an important role linking the sedentary civilizations of Europe to those of Asia. The term was originally used by Greek writers such as Herodotus to describe the peoples of the Pontic Steppes to the north of the Black Sea, a group best known in the archaeological record for their spectacular elite burials and the highly decorative gold items found within them. The lives of ordinary Scythians, however, are less well understood, as the materials from which they made the majority of their clothing, tools, and weapons, such as wood, bone, leather, and textiles, tend not to preserve well, and often degrade into unphotogenic fragments.

In a paper published in the journal PLoS One on 13 December 2023, Luise Ørsted Brandt of the Globe Institute at the University of Copenhagen, Meaghan Mackie, again of the Globe Institute, and of the Novo Nordisk Foundation Center for Protein Research, also at the University of Copenhagen, Marina Daragan of the Institute of Archaeology of the National Academy of Sciences of Ukraine, Matthew Collins, also of the Globe Institute at the University of Copenhagen, and of the McDonald Institute for Archaeological Research at the University of Cambridge, and Margarita Gleba of the Dipartimento dei Beni Culturali at the Università degli Studi di Padova, present the results of a study of Scythian leather samples from southern Ukraine, which sought to identify the Animals whose skins were used to make the leather.

The leather examined came from 18 burials at 14 different sites in southern Ukraine. Many of the leathers were in an extremely fragmentary state, making it impossible to tell what sort of object they had come from. Some of these leathers may have come from leather clothing, particularly trousers, boots, or vessels of various types, although the majority are thought to have come two iconic Scythian leather objects, quivers, used to hold arrows, and gorytos, which were used to carry both arrows and bows. These items were clearly very important to the Scythians, and are found in almost all burials, as well as being depicted on numerous decorative items. The majority of quivers and gorytos are heavily decomposed when found, but can be identified by the presence of metal arrowheads. Quivers used by elite members of Scythian society were artistic and decorative objects, with the best-preserved examples, such as those from Bulhakovo and Ilyinka, give us some idea of how these items were constructed, but little us understood about the manufacture of the quivers used by ordinary members of Scythian society.

The sites from which leather samples were recovered: (1) Bulhakovo; (2) Ilyinka; (3) Kairy; (4) Kislychevate; (5) Ol’hyne; (6) Orikhove; (7) Otradne; (8) Sadove; (9) Tyahinka; (10) Vil’na Ukraina; (11) Vodoslavka; (12) Vysuns’k; (13) Zelene; (14) Zolota Balka. Marina Daragan in Brandt et al. (2023).

Traditional microscopy can sometimes be used to identify leathers, although this is difficult, as the scraping and tanning significantly alter the surface of the material significantly, and leathers from archaeological contexts tend to be further degraded by decay processes. Two samples within Brandt et al.'s study material were preserved with fur on, making it possible to use hair strands to identify the Animals from which the skin had come, but none of the other leathers could be identified by this technique. The first of the two fur samples came from what appears to have been a fur garment from Burial 1 within Kurgan 22 at the Vil’na Ukraina 3 cemetery (an adult woman, apparently of high status buried with jewellery, a mirror and mirror case, and domestic items in the second half of the third century BC), and was identified as having come from an unknown Mustelid, while the second came from a decorative quiver with fur fragments from Burial 3 (a child buried with weapons in the second or early third quarter of the fourth century BC) in Kurgan 4 at the Ilyinka cemetery, and was identified as coming from an unknown Rodent.

In order to identify the remaining leathers (and better identify the furs) Brandt et al. turned to biomolecular techniques. These have become increasingly important in archaeological investigations in recent decades, with DNA analysis allowing not just the identification of Animal and remains to species level, but quite often Human and Animal remains to specific populations, and the illumination of relationships between ancient and modern populations. DNA, however, is seldom recoverable from leather, as it is typically destroyed by the tanning process. Proteomics offers an alternative approach, enabling archaeologists to identify proteins (such as collagen in leather or keratin in hair) from small samples of material, including samples of material, which is likely to be to old, to degraded, or otherwise treated in ways which make the preservation of DNA unlikely.

A selection of the leather object fragments analysed: (1) Ilyinka Kurgan 4 Burial 2; (2) Ilyinka Kurgan 4 Burial 3; (3) Vodoslavka Kurgan 8 Burial 4; (4) Orikhove Kurgan 3 Burial 2; (5) Zelene I Kurgan 2 Burial 3; (6) Kairy V Kurgan 1 Burial 1; (7) Ol’hyne Kurgan 2 Burial 1; (8) Bulhakovo Kurgan 5 Burial 2; (9) Zolota Balka Kurgan 13 Burial 7. The units of the scale bars are cm. Marina Daragan in Brandt et al. (2023).

Forty five samples of leather from the eighteen burials were included in the study. Of these, thirty three samples were identified, sixteen to species level, four to a probable species, seven to family level, one to probable family level, and five to one of two or more species. The majority of the leathers come from domestic Animals, with more than half coming from Sheep and Goats. One sample, from part of a quiver recovered from Burial 2 at Kurgan 3 at Orikove (two adult males buried with a variety of weapons in the first half of the fourth century BC) came from either a Goat or a Reindeer. Another sample, from one of three quiver found in Burial 2 at Kurgan 5 at Bulhakovo (a probable adult male, buried with weapons, jewellery, and domestic items in the second quarter of the fourth century BC), was identified as Cattle leather. Another three samples of leather were identified as either coming from a Bovid or a Cervid, although they were too poorly preserved for any more precise diagnosis.

One of the samples, from a fragment of a leather mirror case found with Burial 1 at Kurgan 6 at Vysuns’k, which comprised two skeletons buried with weapons, jewellery, a mirror, and a Greek kantharos cup, in the second quarter of the fourth century BC, was found to have come from a Red Fox. Another, from a decorative quiver found in Burial 2 of Kurgan 3 at Orikove, came from an unknown Carnivore, probably either a Tiger, Lion, Marten, Wolverine, Otter, or Hyena. 

The piece of fur from a quiver buried with a child, previously identified as an unknown Rodent, was more precisely identified as having come from a Squirrel, although the exact species could not be determined. The fur garment buried with a high status woman, previously thought to be from an unidentified Mustelid, was re-classified as having come from an unknown Felid.

Finally, two samples of leather appear to have been made from Human skin. The first of these comes from one of the three quivers buried with a probable adult male from Kurgan 5 at Bulhakovo, and was identified as definitely Human. The second from Burial 2 of Kurgan 5 at Bulhakovo, in which two skeletons, interpreted as a man and a woman, were buried together in the second quarter of the fourth century BC, with a variety of goods including weapons, domestic items, and jewellery. This fragment was identified as coming from a member of the Family Homininae, which includes Chimpanzees, Bonobos, and Gorillas, as well as Humans, though it is unlikely that any of the other Hominin species were present on the Pontic Steppes in the fourth century BC, so this leather can also be assumed to be of Human origin.

There is no simple recipe for turning skin into leather; the skins of different Animals need to be treated in different ways to achieve a leather of acceptable quality. The fact that the Scythians were using leather derived from numerous different Animals implies that they had a sophisticated understanding of the leathermaking process, and were likely selecting leather from different Animals for different purposes, just as modern leatherworkers do.

The majority of the skins used for leather by the Scythians appear to have come from domestic Animals which would have been herded on the Steppes by the pastoralist Scythians, particularly Goats and Sheep, although at least one of the leathers in Brandt et al.'s study was derived from a Cow, and Horse leathers have been recovered from burials in the Tuva Region of Russia (although this is a long way from the Pontic Steppes). Such Animals are also frequently depicted in Scythian goldware, and bones of Goats and Sheep have been found within the Kurgans of Scythians, interpreted as the remains of funeral feasts.

Scythian gold pectoral from Tovsta Mohyla, Ukraine, depicting a number of domestic Animals. Brandt et al. (2023).

Several of the leathers, and in particular the furs, in Brandt et al.'s study derive from wild Animals, which appear likely to have been hunted for their skins, including a Red Fox, and unknown Cat, and a Squirrel. None of these have been previously identified in the Ukrainian Scythian archaeological record, but are consistent with the types of furs found in Scythian setting across the wider Eurasian area.

The discovery of Human skin being used to make leather by the Scythians is new, and significant. The Greek historian Herodotus, who wrote extensively on the Scythians, certainly described this practice, however Herodotus is known to have embellished his stories somewhat, leaving modern historians unclear as to what can be taken as fact and what is fiction. On this occasion, Brandt et al.'s work appears to confirm that Herodotus was telling the truth. 

Both the direct presence of leather in Scythian archaeological sites, and iconography produced by the Scythians themselves depicting garments, suggests that these people made extensive use of leather to make vessels, mirror cases, quivers, shoes, garments such as trousers and coats, and the lining for metal armour such as greaves. The Scythians are also known to have made extensive use of scale armour, in which metal scales were sewn onto a leather base. It is presumed that the Scythians made leather themselves, as described by Herodotus, although no direct evidence for this has been found on the Pontic Steppes (the such evidence has been found for the nomads of Kazakhstan and Eastern Tibet, whose life-styles are not thought to have been dissimilar to those of the Pontic Scythians). Notably, the Tovsta Mohyla pectoral appears to show two Scythian men engaged in either skinning a Sheep or production of a garment made of sheepskin.

Depictions of Scythian warriors wearing decorated sleeved leather garments: (1)–(2) Gilded silver bowl from Haimanova Mohyla, north chamber. (3)–(4) Golden cone from Perederiyiva Mohyla, Ukraine. Brandt et al. (2023).

However, many of the embossed decorations on quivers from southern Ukraine have a very Hellenic feel to them, suggesting that the Scythians of this region were obtaining materials by trade with the Pontic Greeks, and that some of the goods obtained in this way were either quivers, or materials used in the making of quivers. It is also possible that those Scythians in contact with Greeks adopted some of their decorative styles. Given that high status objects buried with members of the Scythian elite are often decorated with Greek mythological and decorative motifs, either seems plausible.

Some of the fragments of quivers had traces of a red pigment, which was found to be cinnabar, a naturally occurqring form of mercury sulphide. This pigment is known to have been used by the Scythians for a range of decorative purposes, making its presence on quivers a probable indicator of Scythian manufacture.

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Ancient collagen provides insights into the relationships of the South American ‘Ungulates’.

South America was an isolated continent for much of the Cainozoic,  connected only to Antarctica during the earlier part of the period and not forming a land bridge to North America until the Pliocene. This enabled the continent to develop a unique fauna and flora unlike that of other parts of the world, including creatures such as giant ‘Terror Birds’, Sloths, large carnivorous Marsupials and a number of now extinct groups of large herbivorous Mammals (‘Ungulates’); the Xenungulata, Notoungulata, Litopterna, Pyrotheria and Astrapotheria. The relationships of these extinct groups has remained controversial, with some palaeontologists suggesting they are related to other ‘Ungulates’ - Artiodactyls (Deer, Antelopes, Camels, Pigs etc.) plus Perissodactyls (Horses, Tapirs and Rhinoceroses) – while others suggest they are more closely related to the South American Xenarthrans (Sloths, Armadillos etc.).

In a paper published in the Proceedings of the Royal Society: Series B, Biological Sciences on 1 April 2015, Michael Buckley of the Faculty of Life Sciences at the Manchester Institute of Biotechnology presents a study of evolutionary relationships within Mammals based upon collagen protein sequence analysis, which includes two South American ‘Ungulates’ the Litoptern Macrauchenia patachonica and the Notoungulate Toxodon platensis.

The Litoptern Macrauchenia patachonica (larger skeleton at rear). Ryan Somma/Wikimedia Commons.

The last two decades has seen a revolution in taxonomic classification based upon the use of DNA analysis. Loosely speaking, the DNA of animals and plants contains large areas of non-coding DNA, which do not apparently serve any function and therefore are not subject to evolutionary pressures. The mutation rate in these areas of DNA is therefore thought to be a steadily paced ‘random walk’, enabling DNA-based phylogenies to be build up which are not subject to convergent evolution (structures being subject to similar evolutionary pressures and therefore ending up looking the same), as is the case with phytogenic trees based upon morphological comparisons.

DNA-based studies have led to Therian (Placental) Mammals being divided into four main groups, the Euarchontoglires (Primates, Rodents and Lagomorphs), the Laurasiatherians (Bats, Carnivores, Perissodactyls etc.), the Afrotherians (Elephants, Elephant Shrews, Hyraxes etc.) and the Xenarthrans (Sloths, Armadillos etc.), with the Euarchontoglires and Laurasiatherians being considered sister-groups, sometimes referred to collectively as the Boreoeutheria and different studies resulting in either the Afrotherians or Xenarthrans being the closest relatives of this combined group.

DNA sequencing techniques have also been applied to the remains of a variety of ancient organisms, with positive results being obtained from material as old as the early Pleistocene in cool regions (claims of DNA recovery from older material, particularly Mesozoic Dinosaurs, are at best fanciful). However much of South America has a tropical or subtropical climate, severely limiting the potential of DNA recovery from many of its unique fossil groups somewhat unlikely.

Collagen is one of the toughest and most widely used proteins in vertebrate bodies, being found in cartilage, ligaments, tendons and, significantly, in bone. The mineral structure of vertebrate bone is essentially laid down over a framework of collagen, encasing this protein in a mineral matrix which can preserve it from environmental damage long after the death of the animal. Significantly collagen has a somewhat flexible composition, and away from binding sites can contain any amino acids without apparent harm. This means that like DNA, the amino acid structure of collagen is subject to an evolutionary random walk, which can be used to trace evolutionary relationships within vertebrate groups.

Specimen of the Notoungulate Toxodon platensis on display at the Bernardino Rivadavia Natural Sciences Museum. Wikimedia Commons.

Buckley extracted collagen samples from two specimens each of the Litoptern Macrauchenia patachonica and the Notoungulate Toxodon platensis from Pleistocene sites in Buenos Aires, Argentina, and used them to build a collagen proteomic family tree which also included representatives of the Euarchontoglires, Laurasiatherians, Afrotherians, Xenarthrans, Marsupials and Monotremes.

His results recovered the existence of the Boreoeutheria, comprising the Euarchontoglires and Laurasiatherians. The sister group to the Boreoeutheria was the Xenarthrans, with the Afrotherans being the sister group to all other placental Mammals. Within the studied animals, the Litoptern Macrauchenia patachonica and the Notoungulate Toxodon platensis were the closest relatives of eachother, and both were more closely related to the Perissodactyls than any other group.

Phylogenetic analyses of Toxodon and Macrauchenia collagen sequences matched by LC–MS/MS rooted to the duck-billed platypus (Ornithorhynchus), showing maximum-likelihood analysis using PhyML with 10 000 bootstraps (less than 50 not shown except for Toxodon and Macrauchenia). Buckley (2015).

This study suggests that at least two of the extinct South American Ungulate groups, the Litopterns and the Notoungulates, were closely related to the Perissodactyls, leading to a distinct possibility that the other three groups, the Xenungulates, Pyrotherians and Astrapotherians were also. This also suggests that the ancestors of these animals were able to make the crossing between Laurasia (Eurasia plus South America) some-time in the early Cainozoic or possibly the End Cretaceous, suggesting that a land bridge between North and South America must have existed at this time.

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A volcanically preserved Rhinoceros skull from the Late Miocene of Turkey.

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