Monday, 30 November 2020

Dating the earliest Myriapods.

Understanding how organisms colonised the land, is crucial to clarify extant biodiversity and biological adaptation. But, evaluating the rate and pattern of land colonisation requires precise dating of the fossil of early land biotas and reconciling them with evolutionary divergence based on morphology and molecular clocks. The frequent striking inconsistency between the ages of fossils and their phylogenies limits our understanding of macroevolution, and it reduces confidence in phylogenetic inference: this is especially the case for the first land Animals. Arthropoda (Insects, Spiders, Centipedes and their allies) were the first, and are the largest group of land Animals in both numbers and biomass. The first fossil land Arthropod-Plant assemblages are rare and only found in Late Silurian (about 425 million years old) to Early Devonian (about 410 million years old) equatorial terrestrial and freshwater sediments within and marginal to the ancient Caledonian mountains which stretched from New York to Germany. The earliest fossil land Arthropods are Myriapods (Millipedes and Centipedes) from the latest Silurian (about 425 million years old) of Scotland and Wales. 

Yet, while Arachnid molecular phylogenies fit the fossils, Myriapod molecular clock phylogenies do not and suggest a late Cambrian (500 million years) monophyletic origin and divergence of the Myriapod classes, which implies not only a marine origin of the classes and possible independent land colonisation events during Myriapod evolution, but also that pre-late Silurian Myriapods are not preserved in the geological record. Though the first land Arthropods were initially very small with thin cuticle, and are difficult to preserve except under unusual conditions, nevertheless, there are enough organic-rich sediments with soft bodied fossils in appropriate environments in Cambrian to early Silurian times, that any existing land Arthropods would be at least occasionally preserved if present. But they are not. Apart from the doubtful Diplopod (Millipede), Casiogrammu ichthyeros, from the Hagshaw Hills, none occur in the Silurian Fish beds of the Midland Valley of Scotland, famed for their diverse freshwater Arthropods and oldest complete Fish fossils (but with no vascular plant remains), and currently assigned on fossils spores to the Wenlock (about 430 million years old). In any case, given that the dating of the Silurian Fish beds is so uncertain, this Diplopod? Could be the same age as or only slightly older than the Kerrera fossils, and similarly possibly aquatic. Pre-late Silurian tracks and trails of supposed land Arthropods are suspect and may represent, as is common today, only temporary excursion onto land. The earliest land Arthropod fossils are Millipedes associated with the first Vascular Plants in the latest Silurian (Pridoli) around 420 million years ago. But, while the earliest fossil Vascular Plant fossils are found at about 425 million years old (Upper Silurian), molecular phylogenies indicate a 515 million years old to 470 million years old (Late Ordovician-Silurian) origin. The precise timing of the appearance of the first land Arthropod and Plant fossils is difficult to determine because of the problem of correlating the deposits of the lakes, river and coasts, in which they occur, with the standard marine-based geological time scale, and there are almost no radiometric dates from associated sediments to help in this.

In a paper published in the journal Historical Biology on 15 May 2020, Michael Brookfield and Elizabeth Catlos of the Department of Geological Sciences at the University of Texas at Austin, and Stephanie Suarez of the Department of Earth and Atmospheric Sciences at the University of Houston, present the latest results in an ongoing study which aims to provide accurate dates for these first land biotas.

Caledonian mountains with cited localities. Ron Blakey in Brookfield et al. (2020).

Brookfield et al. determined the ages of these first land arthropods, with uranium/lead dating of zircons in the earliest Millipede-bearing sediments from three places, in western England (Ludlow) and Scotland (Cowie, Kerrera). These sediments are associated with contemporary explosive volcanic activity, so the youngest concordant zircons give not only the maximum age of the enclosing sediment  (a sediment cannot be older than the youngest thing in it, though it can be younger) but also a good estimate of the actual age of the sediment from the age of the contemporary volcanic rocks Zircons were separated from sediment samples taken either just above and below the arthropod-bearing beds (Kerrera and Cowie) or from heavy mineral separates obtained for other studies (Ludlow). Both the Kerrera and Ludlow dates are new. The Cowie date comes from an earlier study by Stephanie Suarez, Michael Brookfield, Elizabeth Catlos, and Daniel Stöckli, also of the Department of Geological Sciences at the University of Texas at Austin, and is included by Brookfield et al. to show the apparently rapid progressive coevolution of land Arthropods and Floras in semi-arid continental environments during the latest Silurian. The Kerrera Arthropods come from a temporary lake deposit with Anapsid Fish, interbedded with coarse semi-arid intermontane basin sediments, at the base of the Lorne Plateau Lavas. The only previous radiometric ages for Kerrera come from a lava at the top of the Lorne Plateau Lavas, 600 metres above the Kerrera sediments. Their uranium/lead zircon age of 425 ± 0.7 million years is a uranium/lead thermal ionisation mass spectrometry concordia age from two zircons which give individual ages of 425.4 ± 0.8 million years and 424.5 ± 0.8 million year. Zircons, however, are among the first minerals to crystallise from a cooling magma chamber, may be mixed populations from separate batches of magma, and can be several 100 000 years older than the lava eruption at the surface. The youngest concordant zircon gives a maximum age for the eruption of 424.5 ± 0.6 million years. The time taken for 600 metres of lavas to erupt is not known, but from other more recent similar lava piles, which accumulated over a fairly short time geologically, we can make an estimate. The average 1000 metre thick Grand Ronde Basalt lavas of the Miocene Columbia River Plateau were erupted over a 400 000 years period between 16.5 and 16.1 million years ago. A comparable duration is likely for the Lorne Plateau lavas, which seem to have erupted fairly continuously. Of the 52 zircons Brookfield et al. analysed, the youngest concordant ages were 426.5 ± 4.5 million years and 425.4 ± 4.8 million years, which are statistically indistinguishable. The 1 million years between the midpoints of the youngest Kerrera zircon and the youngest top Lorne Lava zircon is thus a reasonable estimate of the time taken for the Lorne Lava pile to accumulate.

Sections at Ludlow, Kerrera, and Cowie with Youngest dated zircons. Brookfield et al. (2020).

The Ludlow Arthropods, accompanied by a fragmentary Cooksonia flora come from one organic-rich siltstone lens filling ripple troughs in fine sandstone just above the Ludlow bone bed lag deposit, a sandstone marking the change from shallow marine to semi-arid continental environments. Brookfield et al. analysed 29 zircons from heavy mineral concentrations, obtained for Conodont analysis, from two samples of the basal Bone Bed near Ludlow. Nineteen of these were within the expected latest Silurian (Pridoli) biostratigraphic age. The youngest dates of 420.0 ± 8.9 million years and 420.3 ± 8.1 million years are, like those from Kerrera, statistically indistinguishable. The Cowie dates of 413.7 ± 4.4 million years and 414.3 ± 7.1 million years which bracket the Millipede-bearing Fish bed, are younger than both the Kerrera and Ludlow.

These ages are consistent with the evolutionary stages shown by the fossil Millipedes and associated Arthropods. 

The Kerrera and Ludlow Myriapods are Kampecarids which show no obvious structural adaptation to land. The Kerrera Kampecarids, Kampecaris obanensis and Archidesmus sp. occur with an early vascular ‘Cooksonia’ flora, Eurypterids and Anapsid fish in a freshwater lake deposit in a semi-arid fluvial environment. Kampecarids also sporadically occur in late Silurian to early Devonian freshwater lake environments throughout Scotland. The Ludlow Millipedes are also accompanied by the early vascular Cooksonia flora, and by a more diverse fauna of Arachnids, a Centipede and Eurypterids, eroded and redeposited either by a coastal storm deposit from a back-barrier environment or by a tsunami from more inland semiarid coastal plain environments. The most diverse Cowie Millipedes belong to the extinct Archipolyploda superorder and consist of Cowiedesmus eroticopodus which derives its name from preserved male gonopods, and Albadesmus almondi and Pneumodesmus newmani which do not have modified appendages preserved. Pneumodesmus newmani, however, has spiracles on the lateral parts of the sternites which are direct evidence of air breathing, and, before our dating, was taken to be the oldest fully terrestrial Animal. The Cowie deposit was laid down in a temporary lake environment related to braided and meandering streams in a semi-arid intermontane basin, but curiously, contains no identifiable Plant remains or spores. There is thus a progressive change in the Arthropod faunas across the Silurian/Devonian (Pridoli/Lochkovian series) boundary around 419 to 421 million years ago; from Kerrera (about 425 million years old, Ludlow series), through Ludlow (about 420 million years old, Pridoli series), to Cowie (about 414 million years old. Lochkovian series), and this increased diversification continues into younger more diverse early Devonian (Emsian) land biotas, like the Rhynie Chert intermontane basin hot springs biota from Aberdeenshire, and the Gaspé, eastern Canada and Alken, Germany, delta marsh biotas. The diverse Rhynie Chert, with an argon⁴⁰/argon³⁹ age of 407.1 ± 2.2 million years, has a diverse flora and fauna dominated by the Vascular Cryptogam Plants, Rhynia, Aglaophyton, and Horneophyton, and Arthropods; Arachnids (Trigonotarbida), Mites (Acariformes), Harvestmen (Opiliones), a possible Millipede (Diplopoda), Centipedes (Chilopoda), and Springtails (Collembola). Neither the Gaspé nor the Alken biotas have been dated radiometrically. But both have somewhat similar biota to Rhynie, despite their very different environment, with Vascular Cryptogams of the slightly more evolved Zosterophyllum flora, accompanied not only by Arachnids (Trigonotarbids), and Centipedes (Chilopoda), but also by freshwater Eurypterids and Anapsid Fish. By the Middle Devonian (about 385 million years ago), complex forests with ten-metre-tall trees and associated diverse arthropod communities had become established, for example at Cairo and Gilboa, New York. Nevertheless, the composition of the land biotas does not differ significantly from upland lake margins to delta front marshes from latest Silurian to mid-Devonian, a rapidly evolving successions of pioneer communities of Arthropods/Vascular Plants seems to have exploded over the Acadian landscape in a very few million years.

The Kampecarid Millipede, Kampecaris obanensis, from the 425-million-year-old Kerrera deposits of Scotland, currently the oldest known Myriapod. British Geological Survey.

From fossil evidence, the time from initial colonisation by Arthropod-Vascular Plant communities to complex forest communities took place over less than 40 million years (425–385 million years ago). An essentially intermontane lake margin Pridoli FW/land biota in the Acadian mountains centred on Scotland evolved rapidly and migrated, undoubtedly via rivers, to colonise floodplain marshes in the adjacent lowlands by the Emsian, as there is little difference in contemporary biotas. Molecular phylogenies of the Myriapods, however, indicate a monophyletic origin and divergence about 100 million years earlier, while the origin of Vascular Plants is similarly placed from 100 to 50 million years prior to their first body fossils. Which is correct?

Though non-Vascular Plants and possibly amphibious Arthropods were around in the Ordovician, the fossil evidence is clear that Vascular Plant/land Arthropod biotas evolved together in the Late Silurian (about 425 million years ago) and not earlier. If the molecular clock timing were correct, then land Arthropods should be found under exceptional circumstances in earlier deposits. These might be meiofaunal. So, such Arthropod should be looked for in silicified pre-Late Silurian organic soils or hot springs deposits like the Devonian Rhynie Chert, though we know of no such deposits or biotas.

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Smelting zinc in medieval China.

Themain difficulty in zinc-metal production is that while zinc oxide is not reduced by carbon below 1000°C, zinc-metal boils at 907°C, becoming vapour that must be condensed to liquid metal before it comes into contact with enough air to oxidise it back to zinc oxide. A great deal of research has been done on zinc smelting in India. It is widely acknowledged that India was the first country in the world to smelt zinc; the earliest archaeological evidence comes from Zawar in Rajasthan and dates as early as AD 1025–1280. Chinese zinc production can be traced back to at least the early Ming Dynasty (early sixteenth century). Chinese and Indian zinc-smelting technologies are quite different in their condensing techniques, furnaces and retorts (the condensing pots). The discovery of zinc-smelting sites in the Upper and Middle Yangtze River region shows that Chinese zinc-smelting technology probably has an independent technical origin and tradition.

According to historical records, brass was used instead of bronze during the Jiajing period of the Ming Dynasty (AD 1522–1566); this significantly reduced the cost of coin casting. The scale of zinc smelting, however, suggests that it was an important element of the economy during the Ming and Qing Dynasties. Interest in ancient Chinese zinc-smelting technology has increased in recent years.

In a paper published in the journal Antiquity on 9 June 2020, Yuniu Li of Sichuan University, Birui Xiao of the Chongqing Cultural Heritage Research Institute, Gill Juleff of the Department of Archaeology at the University of Exeter, Wan Huang of the Sichuan Provincial Cultural Relics and Archaeology Research Institute, and Dadi Li and Jiujiang Bai, also of the Chongqing Cultural Heritage Research Institute, present the results of a study which began in 2004, in which a series of archaeological surveys and excavations of zinc-smelting sites in the Upper and Middle Yangtze River region have been conducted.

Some 40 zinc-related sites have been discovered and recorded. The majority of the sites are located along the banks of the Yangtze River, probably for convenience of transportation. Our study suggests that people in this region mastered the retort method (reduction and condensation) of extracting zinc as early as the middle of the Ming Dynasty (AD 1435–1572).

Locations of the zinc-smelting sites in the Upper and Middle Yangtze River region. Li et al. (2020).

The smelting sites can be identified in three regions: on the banks of the Yangtze River; the Qiyao Mountain region; and the Wu River region in Youyang County. One of the bestpreserved sites is Linjiangerdui in Zhong County. This site was excavated in 2013, 2017 and 2018. As the site lies within what is now a reservoir (as part of the Three Gorges scheme) it is partially submerged and excavation can only be carried out between May and September each year when water levels are low. The Linjiangerdui site lies to the south of the Yangtze River, west of the Shuiyang stream, and east of a Han Dynasty cemetery site (202 BC–220 AD). The site lies on a slope with the elevation varying between 155 m on the north side and 170m on the south; it covers a total area of 15 000 m². Over the three seasons, 22 zinc-smelting furnaces were excavated and recorded, together with a large number of pits and postholes representing places of preparation and storage for ore and coal, over 1000 retorts (both used and unused) and domestic pottery and porcelain.

Site view of Linjiangerdui. Li et al. (2020).

The ore recovered is lead-zinc ore, which possibly comes from the Qiyao Mountain region. A number of samples of unsmelted ore and slag were analysed and 14 radiocarbon assays were taken from charcoal samples. These produced dates between 1408 and 1650 AD (at 95.4% confidence).

Example of zinc-smelting furnace at Linjiangerdui. Li et al. (2020).

Historical documents provide some information on ancient zinc-smelting processes. The Tiangongkaiwu (The products of heaven and man by Song Yingxing, 1587–1666), records a zinc-smelting method of Luganshi (smithsonite) together with a drawing of the process. Many details of the process, including the precise dimensions of the retorts, the smelting process and the structure of the furnace are either inaccurate or missing from this text, however. Li et al.'s fieldwork has revealed new archaeological evidence of the zinc-smelting process in this region. The excavated retorts were ceramic with a condensing zone at the top. The natural distillation effect is caused by the differential temperature gradient between the top and the bottom of the retorts, which causes zinc vapour to rise from the heated base and condense asmetal in the upper condensing zone. Based on the historical texts and radiocarbon results, the zinc-smelting sites in the Upper and Middle Yangtze River region date to the middle of the Ming Dynasty (AD 1435–1572). Linjiangerdui is the earliest and largest ancient zinc-smelting site discovered in China to date and provides valuable data for understanding zinc-smelting technology as well as the social processes enmeshed in zinc smelting in China.

(a) Unused retorts; (b) used retorts; (c) retort pockets; (d) porcelain of the Ming Dynasty. Li et al. (2020).

The zinc-smelting sites in the Upper and Middle Yangtze River region vary in date, scale and technological complexity. While Li et al. have established the chronological sequence of these sites, further work is needed to understand the differences between sites and the reasons for variation. Comparison with other zinc-smelting sites known from the Hunan and Guangxi provinces will offer a macro-perspective on ancient Chinese zinc production, and illuminate the origin, development and spread of this technology.

Illustration of zinc smelting in Tiangongkaiwu. Li et al. (2020).

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Fireball meteor over Germany.

Witnesses across much of Western Europe have reported observing a bright fireball at about 2.30 am local time on slightly after 5.40 pm GMT on Saturday 28 November 2020. The fireball is described as having moved from east to west, and as having been brighter than the Full Moon, as well as having made an audible boom which caused windows to rattle. A fireball is defined as a meteor (shooting star) brighter than the planet Venus. These are typically caused by pieces of rock burning up in the atmosphere, but can be the result of man-made space-junk burning up on re-entry. 

A fireball meteor seen over central Germany on 28 November 2020. Patricia H/American Meteor Society.

The fireball was seen from France, Belgium, Luxembourg, The Netherlands, England, Denmark, Germany, and Switzerland, and has been calculated to move from west to east over central Germany, dissapearing to the northeast of the town of Harzgerode in Saxony-Anhalt. The fireball has been described as being greenish in colour, which tends to be indicative of a body with a high magnesium content.

Heat map of Western Europe showing areas where sightings of the meteor were reported (warmer colours indicate more sightings), and the apparent path of the object (blue arrow). American Meteor Society.

Objects of this size probably enter the Earth's atmosphere several times a year, though unless they do so over populated areas they are unlikely to be noticed. They are officially described as fireballs if they produce a light brighter than the planet Venus. The brightness of a meteor is caused by friction with the Earth's atmosphere, which is typically far greater than that caused by simple falling, due to the initial trajectory of the object. Such objects typically eventually explode in an airburst called by the friction, causing them to vanish as an luminous object. However this is not the end of the story as such explosions result in the production of a number of smaller objects, which fall to the ground under the influence of gravity (which does not cause the luminescence associated with friction-induced heating).

These 'dark objects' do not continue along the path of the original bolide, but neither do they fall directly to the ground, but rather follow a course determined by the atmospheric currents (winds) through which the objects pass. Scientists are able to calculate potential trajectories for hypothetical dark objects derived from meteors using data from weather monitoring services.

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Thousands evacuated as Mount Lewotolo erupts in eastern Indonesia.

The Pusat Vulkanologi dan Mitigasi Bencana Geologi (Center for Vulcanology and Geological Disaster Mitigation) has ordered about 2800 people to evacuate their homes in 28 villages on Lembata Island in the Lesser Sunda Islands of Indonesia, following an eruption on Mount Lewotolo on Sunday 29 November 2020. The eruption produced an ash column about 4 km high, leading to ash falls in several areas, and leading to the closure of Wunopitu Airport. There are no reports of any casualties associated with this eruption, however the evacuations are still ongoing, and there are concerns about the whereabouts of five children from Waienga, who are believed to have fled into a forest in panic after the volcano erupted, and who have not yet been located.

An ash column over Mount Lewotolo on Lembata Island in the Lesser Sunda Islands on 29 November 2020. Muhammad Ilham/Reuters.

Lembata sits on the northern part of the Timor Microplate; a small fragment of crust caught between the Banda Sea Plate to the north and the Australian Plate to the south. Both these other plates are subducting beneath the Timor Plate, and as they sink into the Earth, melted by the friction and the heat of the planets interior. Some of this melted material then rises through the overlying plate, fuelling the volcanoes of Flores, Timor and the neighbouring islands.

The subduction zones beneath the Timor Microplate. Hamson (2004).

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Eomakhaira molossus: A new species of small saber-toothed Sparassodont Marsupial from the Early Oligocene of Chile.

The Sparassodonta, an extinct group of Metatherians (Marsupials and their extinct relatives), were the dominant group of carnivorous mammals in South America from the early Paleocene (Tiupampan South American Land Mammal Age) to the late early Pliocene (Chapadmalalan South American Land Mammal Age), sharing the ecological role of large terrestrial predator with Phorusrhacid Birds and Sebecid Crocodyliforms during the continent’s long Cenozoic isolation. Sparassodonts occupied many of the niches filled by Placental Carnivorans and 'Creodonts' on other continents and often strongly converged with these groups in morphology. Perhaps the best-known example of this phenomenon is the Thylacosmilinae (Thylacosmilidae of most previous authors), a group of Sparassodonts whose striking morphological resemblance to Placental sabertooths literally makes them a textbook example of convergent evolution. 

Thylacosmilines, as traditionally conceived, are characterized by numerous autapomorphies relative to other Sparassodonts, including hypselodont (ever-growing) upper canines, a highly reduced incisor series that may have been essentially nonfunctional, loss of one premolar locus (thought to be the first upper and lower premolar), retention of the deciduous upper third premolar into adulthood, and a highly distinctive basicranium with a compound squamosal/exoccipital bulla, no alisphenoid tympanic process, no external opening for the primary jugular foramen, and large paratympanic spaces. Several of these features appear to be related to a 'sabertooth' mode of life, while others occur in various other groups of Sparassodonts (i.e. some basicranial features are shared with either Hathliacynids or Borhyaenoids), complicating attempts to phylogenetically place Thylacosmilines within Sparassodonta. This may reflect the fact that most well-known Thylacosmilines come from geologically young deposits (middle Miocene to early Pliocene) and hence exhibit high numbers of apomorphies not present in non-Thylacosmilines.

Both the early evolutionary history of Thylacosmilines and the origins of their distinctive saber-toothed morphology remain poorly understood. Phylogenetic analyses indicate that most major Neogene Sparassodont lineages, including Hathliacynids, Borhyaenids, and Thylacosmilines, diverged from their nearest relatives prior to the late middle Eocene, but representatives of these groups are unknown prior to the late Oligocene. The early Oligocene record, in particular, is essential for clarifying whether these long ghost lineages result from poor taxonomic sampling or are an artifact of insufficient sampling of morphological characters in character-taxon matrices.

Unfortunately, the early Oligocene is among the most poorly sampled intervals in the evolutionary history of Sparassodonts. Several authors have remarked on the near-absence of Sparassodont remains from the earliest Oligocene Tinguirirican South American Land Mammal Age. Until now only two specimens have been identified from this interval: a fragmentary upper molar of an extremely small (Pseudonotictis-sized) species, and an isolated premolar of a larger taxon, both from the La Cancha Fauna of Gran Barranca (Chubut, Argentina). A third specimen, a left dentary tentatively assigned to the Borhyaenoid Pharsophorus lacerans, has been described from slightly higher early Oligocene levels (La Cantera) at Gran Barranca. These strata, which postdate the Tinguirirican South American Land Mammal Age and predate the Deseadan South American Land Mammal Age, are informally referred to as the 'Canteran' interval.

In a paper published in the American Museum Novitates on 17 July 2020, Russell Engleman of the Department of Biology at Case Western Reserve University, John Flynn of the Division of Paleontology and Richard Gilder Graduate School at the American Museum of Natural History, André Wyss of the Department of Earth Science at the University of California, Santa Barbara, and Darin Croft of the Department of Anatomy at Case Western Reserve University, describe a new Sparassodont taxon based on a specimen recovered from the upper Cachapoal River drainage in the Andean Main Range of central Chile, approximately 100 km southeast of Santiago.

Middle Eocene to Oligocene South American Land Mammal Ages. The Tinguirirican or 'pre-Deseadan'–aged La Cantera Fauna is represented as a thin bar, as it is thought to represent a very short interval of geologic time (less than 150 000 years). Engleman et al. (2020).

Taxa recovered from Cachapoal are regarded as Tinguirirican in age based on biochronologic evidence, making this the first Oligocene Sparassodont to be described from Chile, and only the second to be described from the many diverse faunas of the Abanico Formation (after the late Eocene Chlorocyon phantasma). Engleman et al.'s specimen, SGOPV 3490, consists of the anterior portion of the skull of a senescent individual and is the most complete early Oligocene Sparassodont known. SGOPV 3490 shows several morphological similarities to Thylacosmiline Sparassodonts, and phylogenetic analyses indicate that it likely represents an early member of the group.

Locations of the Cachapoal locality and the similar-aged (likely coeval) Tinguiririca locality in central Chile. Gray area in inset box represents outcrops of the Abanico Formation. Engleman et al. (2020).

The new species is named Eomakhaira molossus, where 'Eomakhaira' derives from the Greek root 'Eos', meaning 'dawn', and 'makhaira', a type of short sword or large knife (often translated as 'carving knife'), in reference to the bladelike canines of Thylacosmilines, and 'molossus' comes from the Greek 'molossus', a term used to refer to short-snouted, robust-skulled Dog breeds such as mastiffs and bulldogs and refers to the short, robust snout of this species. 

Eomakhaira molossus is described from a single specimen, SGOPV 3490, a partial rostrum of a senescent individual preserving the right maxilla with C-P3, alveoli and partial roots of M1–2, and part of M3; left maxilla with C-P3, anterior root of M1, and M3–4; left and right horizontal rami of the mandible, including both lower canines and most of the postcanine dentition, as well as parts of the coronoid processes; the entire left and parts of the right nasal; parts of the palatine; and the orbital process of the left lacrimal.

This comes from the Abanico Formation at the Cachapoal locality, on the west side of Estero Los Llanos of the upper Río Cachapoal drainage, in the Libertador General Bernardo O’Higgins Region of central Chile. Abanico Formation. Most specimens from Estero Los Llanos were recovered from talus cones at the southeast nose of a roughly north-south running ridge of about 1500 m relief. This ridge roughly parallels the strike of the steeply west-dipping beds. The thickness of the Abanico Formation in the Cachapoal region has not been measured in detail but is on the order of 2000–4000 m. Within this thick succession, the exact horizon that produced SGOPV 3490 is not known, as the specimen was collected from talus. 

Fossils from the Cachapoal locality are likely at least 29.3 ± 0.1 million years old (at least in part), based on an unpublished date for a volcanic tuff that is thought to either correlate with or overlie the fossil-producing horizons at Los Llanos. It must be cautioned, however, that this date is from ~5 km to the south, in the neighboring Las Leñas drainage, and that the units involved have not been traced directly between the two locations due to precipitous intervening topography. The only radioisotopic date for the Cachapoal Valley itself is an argon⁴⁰/argon³⁹ date of 11.1 ± million years reported by Abagael West from levels far above the fossil-producing strata, which does little to precisely constrain the age of the fossils. The presence of the polydolopid Kramadolops and the Archaeohyracid Archaeotypotherium suggest a pre-Deseadan age, and the presence of the Interathere Johnbell hatcheri, otherwise known only from the Tinguirirican type locality, suggests that fossils from the Cachapoal locality are probably similar in age to those of the Tinguiririca Fauna (about 33–32 million years old).

Eomakhaira molossus is considered to be a member of the Borhyaenoidea based on its short, robust rostrum, presence of lingual median canine sulci, extremely small protocone, small and unicuspid talonid on m4. Differs from all other Borhyaenoid Sparassodonts in the following combination of features: small size (smaller than most other Borhyaenoids; length of m1–4 is 37.3 mm, comparable to Fredszalaya hunteri or the extant Dasyuromorphian Sarcophilus harrisii); maxilla very deep and maxillary 'cheeks' absent; mandibular symphysis unfused and anteroposteriorly narrow; two mental foramina present; length/width ratio of palate more than 1.5; palate extending to level of M4; presence of postpalatine tori (shared only with Arminiheringia and possibly Callistoe among Borhyaenoids); absence of postpalatine torus foramen; sphenorbital foramen opening dorsal to M4; large canines; absence of longitudinal striations on the canine roots (shared only with other Thylacosmilines and possibly Lycopsis viverensis); median keel on the labial face of upper canines; medial sulcus on lingual face of upper and lower canines; short lower canine roots; presence of three premolars with no diastemata between them; premolars large and robust but not globular; asymmetric protoconid of P1 (shared only with Arminiheringia and Callistoe); P3 significantly longer than p3 (possibly autapomorphic for this taxon); bulbous roots only on p3; preparacingulum absent; M3 with narrow stylar shelf and prominent ectoflexus; M4 extremely narrow anteroposteriorly (only comparable to Patagosmilus among Borhyaenoids), subequal or greater in width to M3, and with three roots; protocone vestigial (at least on M4); absence of an anteriorly projecting ventral keel of paraconid (which only occurs in proborhyaenids among sparassodonts); protoconid of m4 posteriorly salient; metaconid absent on m4 and probably m2–3; posterolabial cingulid present; talonid of m4 almost absent; and p1–3 short relative to m1–4 (shared with Paraborhyaena among Borhyaenoids with three premolars). Canines more mediolaterally compressed than in Borhyaenoids other than Patagosmilus, Thylacosmilus, and possibly Proborhyaena. P/p3 labiolingually narrower than in Fredszalaya, Plesiofelis, Acrocyon, Arctodictis, Australohyaena, Borhyaena, and Callistoe, but wider than in Prothylacynus and some individuals of Pharsophorus, comparable in relative proportions to Arminiheringia, Paraborhyaena, and Proborhyaena.

SGOPV 3490 is uncharacteristically poorly preserved compared to most fossils described from the Abanico Formation. The specimen was altered syn- or postdepositionally through the actions of heat, fluids, or both, resulting in the thinning or elimination of much of the bone. The density contrast between the volcaniclastic matrix and specimen is low, making it difficult to distinguish rock from bone with the naked eye as well as via computerised tomography imagery. Many portions of the specimen were damaged or destroyed during deposition or diagenesis, leaving many surviving elements isolated but 'floating' in matrix in near-life position, as has been described for some other specimens from the Abanico Formation. For example, the left lower molar row of SGOPV 3490 is preserved in life position, but much of the mandibular ramus is absent labially. Intact toothrows preserved in the absence of bone are occasionally recovered from the Abanico Formation. These specimens may reflect high temperatures or corrosive fluids in the lahar or pyroclastic flow in which the specimens were deposited or subsequent diagenetic processes (the Abanico Formation is locally hydrothermally altered). The specimen also shows clear signs of postmortem crushing and distortion, particularly on the left side, where elements of the skull show signs of breakage and have been displaced anteriorly. By contrast, the right side of the specimen is nearly undistorted; the upper and lower teeth are nearly in occlusion, and the maxilla and mandible show no signs of crushing.

(A), (C) Photographs and (B), (D) computerised tomography segmentation of the holotype of Eomakhaira molossus, a partial skull of a senescent individual preserving the rostrum and the anterior portion of the mandible (SGOPV 3490) in left (A), (B) and right (C), (D) lateral views. In renderings of the computerised tomography segmentation, nasal in orange, facial process of the lacrimal in teal, palatine in blue, all other bones of the cranium (maxilla, jugal, frontal, etc.) in purple, teeth in yellow, and dentary in green. Anterior to left in (A)–(B) and to right in (C)–(D). Scale is 30 mm. Engleman et al. (2020).

Anatomical positions and directions can be difficult to consistently establish and apply to SGOPV 3490. Many landmarks typically used for orientation (e.g. the alveolar border of the postcanine toothrow or the ventral edge of the dentary) sometimes provide conflicting orientations; orienting the skull based on one landmark results in physically impossible orientations for others. Assuming that the fragments of the palate indicate the horizontal plane and that the roots of several postcanine teeth (P2–3, m1–3) approximate the vertical plane, the canines were procumbent and the lower molar rows were inclined to a degree similar to that observed in other Sparassodonts (e.g. Arctodictis, Arminiheringia, Australohyaena, Callistoe, some individuals of Thylacosmilus). Determining the orientation of the rostrum in Eomakhaira more securely would require a more complete or less distorted specimen.

SGOPV 3490 represents a highly senescent individual, as extreme tooth wear obscures much of its dental morphology. The canines are extremely blunt, even compared to many other sparassodonts, with the apices of both the upper and lower canines nearly rounded. Most of the postcanine dentition is also heavily worn. The main cusp of the left p2 is worn nearly flat, and the occlusal morphology of the right M3 is largely obliterated by wear. The entire crown of the right P3 is worn away, and its roots are in direct occlusion with the lower dentition. The occlusal surfaces of the right m1 and the trigonid of right m2 are virtually flat due to wear. Obliteration of the occlusal morphology of m2 indicates that this specimen pertains to a senile individual. Even M4, typically the least worn and last tooth to erupt in Sparassodonts, exhibits well-developed wear facets in SGOPV 3490. The dentition of SGOPV 3490 is obviously heavily worn through use rather than postmortem abrasion, as the posterior face of P3 and the trigonid of m1 almost occlude and have perfectly matching wear facets. Heavy wear obscures some important morphological details and makes it difficult to determine whether certain features typify the species or are only wear related.

(A) Photograph and (B) line drawing of the exposed right upper canine and postcanine dentition of SGOPV 3490, showing their extreme wear. In (B) enamel is denoted in dark grey, dentine in brown, and wear surfaces by light grey. The morphology of these teeth is not entirely visible, as much of the occluded upper and lower jaws of the specimen remain encased in matrix. Scale is 10 mm. Engleman et al. (2020).

The maxilla of SGOPV 3490 is proportionally deeper than in most other sparassodonts, including the robust-skulled Proborhyaenids Arminiheringia and Callistoe and Borhyaenid Arctodictis sinclairi. Only Australohyaena antiquua, Arctodictis munizi, and Thylacosmilus atrox have relatively deeper maxillae among the taxa analyzed. A small portion of the dorsal border of the infraorbital foramen is preserved in SGOPV 3490, indicating that this structure opened dorsal to the P3/M1 embrasure, as in Patagosmilus, Thylacosmilus, and Australohyaena but unlike in: (1) the Eocene taxa Callistoe and Arminiheringia, in which the infraorbital foramen opens above or anterior to the anterior root of P3; (2) Borhyaena and cf. Proborhyaena (MLP 79-XIII-18-1), in which the foramen opens slightly more anteriorly over the posterior root of P3; and (3) species of Arctodictis, in which the foramen is more posterior (above or posterior to the posterior root of M1).

Relative maxilla height and dentary depth (measured at m3–4 embrasure) in sparassodont specimens for which both maxilla and dentary are known, scaled to lower molar row length. Eomakhaira molossus is denoted by a star. Skulls of several taxa are illustrated to highlight variation. Engleman et al. (2020).

Although the alveolar border of the maxilla posterior to the infraorbital foramen is fragmentary, the parts preserved suggest that Eomakhaira lacked maxillary 'cheeks', i.e. protrusions of the maxilla posterior to the infraorbital foramen that extend lateral to the toothrow (best seen in ventral view). Maxillary 'cheeks' are a highly variable feature within Sparassodonta. They are present in Borhyaenids, Prothylacynus, and many Hathliacynids but absent in most species of Lycopsis (except Lycopsis torresi), Acyon, Patagosmilus, and cf. Proborhyaena (MLP 79-XII-18-1). Contrary to some reports, maxillary 'cheeks' appear to be absent in Paraborhyaena and Thylacosmilus (FMNH P14531, MLP 35-X-4-1, MMP 1443). The state in Arminiheringia could not be determined based on available information.

Based on the posterior border of the left nasal, the two naso-frontal sutures of Eomakhaira form an angle of ~100° in dorsal view, greater than the acute-angled naso-frontal sutures of Callistoe, but narrower than those of Paraborhyaena, Patagosmilus, Pharsophorus, and Borhyaenids. An internasal projection of the frontals is absent (i.e., the naso-frontal suture is V-shaped rather than W-shaped). The preserved lateral edge of the left nasal is straight, suggesting it represents the border of the naso-lacrimal suture, based on a similar morphology of this suture in other Sparassodonts. As in Callistoe, a portion of the nasals may have extended onto the lateral surface of the snout, but this cannot be determined with certainty. The nasals are proportionally slender compared to the rest of the skull. Scaling the greatest width of the nasals to the length of M3, the nasals of Eomakhaira are narrower than in most borhyaenoids except Callistoe, Patagosmilus, and a juvenile specimen of Prothylacynus patagonicus (MACN-A 5931), and they are much narrower than the nasals of Paraborhyaena, Arminiheringia, Patagosmilus, and Borhyaenids. The nasals of SGOPV 3490 are only about 25% as wide anteriorly as they are at their widest point. In Borhyaenoids, this figure is typically about 30%, with the exception of Callistoe, in which the nasals vary less in width along their length (though this may be affected by mediolateral compression of the holotype). Nasal bones of the Hathliacynids Sipalocyon and Acyon show less anterior tapering than in borhyaenoids apart from Callistoe, while the proportions of the nasals in Cladosictis more closely resemble those of Borhyaenoids. Interestingly, the Sparassodont UF 27881 from the middle Miocene of Quebrada Honda, Bolivia, originally described as a basal Sparassodont but recovered as a Borhyaenoid in later analyses does not resemble Borhyaenoids in its nasal proportions; rather, it is more similar to a specimen tentatively assigned to the basal Sparassodont Hondadelphys (IGM 250364).

The palate of SGOPV 3490 is fragmented and patchily preserved, but enough is present to establish that Eomakhaira lacked maxillopalatine fenestrae, as in other Sparassodonts. The length/width ratio of the palatal process of the maxilla exceeds 1.5, even taking distortion into account, a value similar to most sparassodonts (with the exception of Thylacosmilines and the Borhyaenids Australohyaena and Arctodictis, in which the process is anteroposteriorly shorter and mediolaterally wider and the ratio is less than 1.5). A pair of palatal pits occurs on the palatal process of the maxilla between M3–4; whether an additional pair was present between M2–3 cannot be determined. Accounting for deformation, anterior displacement of the palatine, and the separation of the maxillary and palatine borders of the minor palatine foramen, the horizontal process of the palatine does not appear to have extended posterior to M4, reminiscent of the condition in Borhyaena, Patagosmilus, and some specimens of Prothylacynus.

Cranium of the holotype of Eomakhaira molossus (SGOPV 3490) in palatal view. Anterior to right. Abbreviations: mpps, medial postpalatine spine; pp, palatal pit; ptor, palatine torus; dental abbreviations as in Materials and Methods. Scale is 30 mm. Engleman et al. (2020).

A pair of low palatine tori are present at the posterior end of the palate. These structures are mediolaterally broad, extending across each palatine bone, but do not contact one another medially. Among Metatherians, Sparassodonts are unusual in the general absence of a palatine torus, a feature also observed in Deltatheroidans, basal Didelphids (Caluromyines and Glironia), and some Dasyuromorphians (i.e. Thylacinus). In all of these taxa, the posterior border of the palate is typically single or double arched in ventral view (depending on the presence or absence of a medial postpalatine spine). Although most Sparassodonts lack a palatine torus, in many (e.g. UF 27881, Cladosictis, Sipalocyon, Borhyaena, Australohyaena, Arctodictis, Thylacosmilus) the posterior border of the palate is slightly thickened. This thickening can be extremely pronounced (e.g. as in species of Arctodictis), but it follows the borders of the choanae rather than forming a straight torus. Other than Eomakhaira, the only Sparassodonts with true palatine tori are the basal taxon Allqokirus and the Proborhyaenids Callistoe and Arminiheringia. However, even in these taxa, the palatine tori do not resemble those of most other Metatherians, wherein a straight palatine torus defines the posterior border of the palate. In Allqokirus, the palatine torus is well developed but is posteriorly concave rather than straight. Callistoe and Arminiheringia resemble Eomakhaira in having palatine tori that are mediolaterally oriented ridges at the back of the palate. Compared to the thickened choanal border of other Sparassodonts, these structures are much more prominent in Eomakhaira, forming distinct processes that do not follow the borders of the choanae. However, unlike in other Metatherians, the palatine tori in Callistoe and Arminiheringia do not form a single, complete torus; rather, they are paired structures that do not meet at the midline (except possibly in Callistoe) and do not constrict the choanae (i.e. in ventral view, the choanae still exhibit the classic 'double arch' pattern typical of Sparassodonts).

Posterior palate of the holotype of Eomakhaira molossus (SGOPV 3490), in oblique anterior view. Anterior to lower left. Shows the paired palatine tori and broken border of the minor palatine foramen. Abbreviations: mpf, minor palatine foramen; ptor, palatine tori. Scale is 10 mm. Engleman et al. (2020).

A small foramen occurs just lateral and slightly dorsal to the lateral edge of each palatine torus. It likely represents the minor palatine foramen based on its position, though it is slightly damaged on both sides of the skull due to anterior displacement of the palatines. In Eomakhaira, the minor palatine foramen is located between the maxilla and the palatine, as in most Metatherians, including many Sparassodonts (including UF 27881, Cladosictis, Arctodictis, Callistoe, Arminiheringia, Patagosmilus, and some but not all specimens of Thylacosmilus). In Callistoe and Arminiheringia, as in Eomakhaira, the minor palatine foramen is dorsal to and slightly tucked under the lateral edge of the palatine torus. The minor palatine foramen is positioned more laterally in Eomakhaira than in other Sparassodonts; in most Sparassodonts, it is closer to the choanae than to the upper teeth, whereas in Eomakhaira, it is closer to the upper dentition. In this respect, Eomakhaira resembles Patagosmilus (but not Thylacosmilus). The minor palatine foramen of Eomakhaira is fairly large, more comparable in size to that of Patagosmilus than Arminiheringia or Callistoe (in which it is smaller). This foramen is clearly not the postpalatine torus foramen present in most groups of New World Metatherians, as the homologous structure is either an open notch or absent in Sparassodonts. The postpalatine torus foramen opens directly posteriorly into the basipharyngeal canal in other Metatherians, whereas the inferred minor palatine foramen of Eomakhaira opens posterolaterally into the orbitotemporal region, similar to the path of the minor palatine foramen of other Sparassodonts. The postpalatine torus foramen appears to be absent in Eomakhaira, as it is in Proborhyaenids (including Thylacosmilines) and Borhyaenids.

Left orbital region of the holotype of Eomakhaira molossus (SGOPV 3490) in oblique posterolateral view. Anterior to left. Abbreviations: laf, lacrimal foramen, mpf, minor palatine foramen; spf, sphenopalatine foramen. Scale is 30 mm. Engleman et al. (2020).

SGOPV 3490 preserves a small portion of the orbital region, primarily on the left side. This consists of part of the ascending process of the palatine posteriorly and several isolated plates of bone separated by distinct gaps and holes that form part of the orbital wall anteriorly. Based on their position and morphology, these bone fragments likely represent parts of the orbital process of the lacrimal and the anterior part of the ascending process of the palatine. No clear suture is visible between the palatine and maxilla in lateral view. Based on size and location, these gaps may represent sutures and foramina that were enlarged postmortem by fragmentation of the fragile surrounding bone. The largest of these openings compares well to the sphenopalatine foramen in location. This foramen opens approximately dorsal to M4, as it does in some other Borhyaenoids (Callistoe, Thylacosmilus).

The anteriormost fragment of the orbital wall appears to represent a small portion of the orbital process of the lacrimal. Parts of the facial and zygomatic processes of the lacrimal may be present but cannot be distinguished from surrounding elements (i.e. maxilla). At the anterior end of the fragment of the orbital process is a small, partially preserved canal, likely the lacrimal foramen. Based on its position and surrounding elements, the lacrimal foramen appears to have opened within the orbit. Most Sparassodonts have a single lacrimal foramen opening inside the orbit. However, in Mayulestes, Allqokirus, Lycopsis padillai, and Arminiheringia there are two lacrimal foramina, and in Callistoe, the lacrimal foramen number is polymorphic. Additionally, in Allqokirus and Mayulestes, one of the two lacrimal foramina is laterally exposed rather than enclosed within the orbit. It is clear that at least one lacrimal foramen that opened within the orbit was present in SGOPV 3490, though the lacrimal foramen count is uncertain due to the limited preservation of this element.

SGOPV 3490 preserves parts of both jugals. On the left side, this element is represented by a fragmentary bone 'floating' near the orbital region. Part of the right jugal also seems to be present, represented by small patches of bone (including parts of the rostrum formed by the jugal in other metatherians) and remnants of a marrow cavity dorsal to the upper molars. The shape and location of the maxillo-jugal suture cannot be determined.

The mandible of Eomakhaira is robust and deep. It is comparatively shallower than the mandible of the Proborhyaenids Callistoe and Arminiheringia and deeper than in Proborhyaena and Paraborhyaena. The dentary is fractured parallel to the long axis of the horizontal ramus of the dentary, and these complementary fractures are displaced and infilled by matrix. The thickness of these matrix-infilled gaps between complementary bone fragments suggests that the dentary of SGOPV 3490 was originally roughly 2+ mm shallower than what is reported by Engleman et al. As in Callistoe, Arminiheringia, and Pharsophorus tenax, the horizontal ramus of Eomakhaira deepens posteriorly, has a ventral border that is curved in lateral view, and terminates anteriorly in a simple curve. Indeed, the horizontal ramus in Eomakhaira strongly resembles that of Arminiheringia (MACN-A 10970). This contrasts with the condition in the Borhyaenids Australohyaena and Arctodictis, the Proborhyaenids Proborhyaena and Paraborhyaena, and MPEF-PV 4170, a specimen assigned to Pharsophorus cf. Pharsophorus lacerans (but not in the holotype of Pharsophorus lacerans, MACN-A 52-391), in which the horizontal ramus is nearly uniform in depth, its ventral margin is flat, and its anterior end forms a distinct 'chin', with a sharp angle between the anterior border of the symphysis and the ventral border of the horizontal ramus. A distinct 'chin' is also present in the basal Sparassodonts Allqokirus and Mayulestes but is absent in most other members of this group in which the anterior border of the dentary is curved. The dentary shows no sign of a genial flange, in contrast to Thylacosmilus, Anachlysictis, or the unnamed Thylacosmilid from La Venta (in which it is present). The deepest point of the horizontal ramus appears to have been below m4 in SGOPV 3490, as in most Sparassodonts.

Mandible of the holotype of Eomakhaira molossus (SGOPV 3490). Right dentary in (A) labial and (C) lingual views. Left dentary in (B) labial and (D) lingual views. Abbreviations: cor, coronoid process of dentary; sulc, lingual sulcus of the lower canine; menf, mental foramina; symph, mandibular symphysis. Scale is 50 mm. Engleman et al. (2020).

The left dentary is displaced slightly anteriorly relative to the right. The anteroventromedial edge of the left dentary is straight in ventral view, and the medial face of this element preserves a small portion of the symphyseal surface. The symphyseal surface does not bear well-developed interdigitating ridges (in contrast to Sparassodonts like Borhyaena, in which such ridges are present). In dorsal and ventral views, the medial face of the right dentary inflects medially near the p2/3 embrasure, a feature denoting the posteriormost extent of the mandibular symphysis in most Mammals, suggesting the symphysis of Eomakhaira extended posteriorly to the level of the p2/p3 embrasure, or at most, just slightly below the anterior root of p3. The symphysis of Eomakhaira is thus rather short compared to closely related Sparassodonts; in these taxa, the mandibular symphysis reaches its midpoint below the: (1) p3 roots (Borhyaena macrodonta, Borhyaena tuberata, and Pharsophorus lacerans, including MPEF-PV 4190, the specimen of Pharsophorus cf. Pharsophorus lacerans from La Cantera), (2) posterior root of p3 (Plesiofelis, Australohyaena and Prothylacynus), (3) p3/m1 embrasure (Callistoe, Paraborhyaena, Proborhyaena, and Arctodictis spp.), or (4) m1 (Arminiheringia). In Acrocyon riggsi and Pharsophorus tenax, the symphysis extends below the anterior root of p3 but further posteriorly than in SGOPV 3490. In Thylacosmilus atrox and Anachlysictis gracilis, the symphysis is much shorter than in all aforementioned taxa, ending below the canines.

Several lines of evidence suggest that the mandibular symphysis of SGOPV 3490 was unfused. The two dentaries are offset anteroposteriorly with respect to one another. The preserved portion of the left symphyseal surface bears a straight ventromedial edge, suggesting a clean break between the two rami, as would be expected if they had been held together by ligaments. This configuration would be unlikely if a fused symphysis were broken postmortem. Had the mandibular symphysis been fused in vivo, one would expect an uneven, jagged break between the two dentaries, or for one or both to be broken immediately posterior to the mandibular symphysis, where the dentary is comparatively the weakest. This is the case in several other Sparassodonts with fused symphyses and broken mandibles (e.g., MACN-A 706, Prothylacynus patagonicus; MLP 85-VII-3-1, Arctodictis sinclairi; UATF-V-000129, Paraborhyaena boliviana).

Each dentary of SGOPV 3490 bears only two mental foramina, one located beneath p2–3 and another below the m1–2 embrasure. The posterior foramen is well defined on both dentaries and opens posteriorly; the anterior foramen is not well preserved on either dentary, but its existence and position can be inferred from a gap in the bone fragments labially beneath the premolar row. The presence of only two mental foramina on each dentary in Eomakhaira is unusual for a Borhyaenoid. Most Hathliacynids, Borhyaenoids, and the basal Sparassodont Stylocynus bear three or more mental foramina on each dentary, and individuals of some species possess as many as five or six (Acrocyon riggsi, Arctodictis sinclairi, Australohyaena antiquua, Borhyaena tuberata, and Lycopsis longirostrus). Additional mental foramina may have been present in SGOPV 3490, but the location of the preserved bone fragments relative to the positions of mental foramina in other sparassodonts (typically between p2 and m1 or m2) make this unlikely.

Enough of the left coronoid process is preserved to indicate that it is tall and well developed. However, due to crushing and distortion, it is not possible to determine the shape of the masseteric fossa, nor can the angle between the anterior border of the coronoid process and the toothrow be securely determined. A small portion of the right ascending ramus (which is not obviously deformed) indicates that the anterior border of the coronoid process is oriented approximately 110°–113° relative to the toothrow, a value typical for Sparassodonts.

Little of the precanine dentition is preserved in SGOPV 3490. A cylindrical fragment of a small tooth appressed to the lingual side of the lower right canine may represent i3, based on the position of this tooth in other Sparassodonts. If this fragment represents part of a lower incisor, then the lower incisors of Eomakhaira molossus were proportionally smaller than those of Arminiheringia auceta, Arctodictis sinclairi, and potentially even Australohyaena antiquua and Paraborhyaena boliviana (scaling by both p3 and m4) but still larger than in MLP 77-VI-13-1, a specimen assigned to Arctodictis sinclairi that has been noted to have relatively small teeth compared with other specimens of this taxon.

Lower right dentition of SGOPV 3490 in (A) labial, (B) occlusal, and (C) lingual views.Anterior to right in (A), (B) and to left in (C). Scale is 30 mm. Engleman et al. (2020).

The most conspicuous feature of the holotype of Eomakhaira is its large, robust canines. These teeth are disproportionately large compared to most Sparassodonts, comparable (in relative size) only to Proborhyaenids (including Thylacosmilines), and the Borhyaenids Australohyaena, Acrocyon, and Arctodictis. The surfaces of the canine roots in SGOPV 3490 are smooth. In most Borhyaenoids, the canine roots bear a series of small longitudinal grooves that sometimes nearly reach the apex (e.g., in Arminiheringia and Proborhyaena). The only Borhyaenoids that do not have these grooves are Thylacosmilines and possibly Lycopsis viverensis.

(A) Photograph and (B), (C) computerised tomography images of the right upper canine of the holotype of Eomakhaira molossus (SGOPV 3490). (A) Lateral view, showing bluntness of canine apex and the absence of enamel, longitudinal ridges, and labial median canine sulcus. (B) Transverse section of canine, slightly below alveolar border (actual point at alveolar border obscured by a crack), showing posterior keel. Anterior root of P1 (aP1) marked to show that the longitudinal ridge is not an artifact of postmortem damage. (C) Transverse section of canine, at level of tooth row, showing presence of median labial keel and lingual median sulcus. Approximate location of sections in (B), (C) denoted by arrows on (A). Anterior to right in all images, and lingual to top in (B), (C). Abbreviations: aP1, anterior root of P1; mk, labial median keel; ms, lingual median sulcus; pr, posterior ridge. Scale is 10 mm (A); 5 mm (B), (C). Engleman et al. (2020).

The upper and lower canines of Eomakhaira bear a well-developed median sulcus lingually, making them somewhat reniform in cross section. By contrast, no sulcus occurs on the labial side of lower canine or the exposed labial surface of the upper canine. A shallow labial sulcus is present on intralveolar portions of the upper canine but is visible only on computerised tomography scans. Although median canine sulci have been considered a synapomorphy of Proborhyaenids, this feature is widely distributed among Sparassodonts, with lingual sulci also occurring in the Borhyaenids Australohyaena antiquua, Arctodictis sinclairi, Arctodictis munizi, and Borhyaena macrodonta, the basal Borhyaenoid Pharsophorus lacerans (as seen in the holotype MACN-A 32-391, YPM-VPPU 20551, and MPEF-PV 4190, a specimen from La Cantera assigned to Pharsophorus cf. Pharsophorus lacerans), and an indeterminate sparassodont from the Fitzcarrald Arch. William Sinclair in 1930 and Larry Marshall in 1978 reported lingual sulci in Acrocyon riggsi, but Engleman et al. could not verify this observation in photographs of this taxon. In all Sparassodonts in which median sulci occur, the labial ones are less prominent than the lingual ones (e.g. Proborhyaenids, Arctodictis munizi) or are absent (all other taxa). Although the extraalveolar portion of the upper canine of Thylacosmilus lacks a median sulcus, a shallow median sulcus is present on the intralveolar portion of this tooth (FMNH P14344, FMNH P14531). The lower canines of Thylacosmilus bear median sulci labially and lingually.

In cross section, the lingual side of the upper canines of SGOPV 3490 is slightly flatter than the labial side but not as flat as in Thylacosmilus atrox. The labial surface of the right upper canine of Eomakhaira is slightly keeled, similar to but less pronounced than in Thylacosmilus. This feature is not evident on the more poorly preserved left upper canine. A small carina occurs on the posterior edge of the upper canine, slightly more marked on the right tooth than the left. Carinae appear to be absent in the upper canines of Arminiheringia auceta, cf. Proborhyaena (MLP 79-XII-18-1), and other Borhyaenids we have observed. By contrast, the upper canines of Thylacosmilines, like those of most saber-toothed mammals, bear well-defined carinae that lend the tooth a knifelike appearance. In Patagosmilus, the upper canine is blunt anteriorly but bears a sharp posterior carina, similar to the condition in Eomakhaira. The anterior and posterior faces of the upper canines form well-defined carinae in Thylacosmilus, the posterior of which is much sharper.

The upper canines of Eomakhaira are mediolaterally compressed compared to non-Thylacosmiline Borhyaenoids. The length/width ratio of the upper canines ranges from 1.50–1.77, depending on orientation of the skull during measurement. The lower end of this range likely underestimates the degree of mediolateral compression of the tooth, however, as it requires an anatomically unlikely orientation of the specimen (one where the palate is highly inclined and the tooth roots are far from vertical). More reasonable orientations of the specimen yield higher estimates. The most reasonable orientations of SGOPV 3490 produce canine length/width ratios of 1.65–1.70. Orientation aside, the upper canines of Eomakhaira are clearly more mediolaterally compressed than in most other Borhyaenoids, including Pharsophorus, Prothylacynus, all Borhyaenids (Acrocyon, Arctodictis, Australohyaena, and Borhyaena), IGM 251108 (the putative Thylacosmiline from La Venta), most non-Thylacosmiline Proborhyaenids (Arminiheringia, Paraborhyaena, Callistoe), and the indeterminate Proborhyaenid from the Tremembé Formation. However, they are less compressed than in Patagosmilus, Thylacosmilus, and cf. Proborhyaena (MLP 79-XII-18-1). In terms of nonSparassodont carnivores, the canine proportions of Eomakhaira are comparable to the machaeroidine Oxyaenid 'Creodont' Machaeroides eothen, the Nimravid Carnivoramorphans Dinictis felina and Nimravus brachyops, and the machairodontine Felid Carnivoran Pseudaelurus quadridentatus. The resemblance to the latter three placental taxa is noteworthy, as each is among the least-specialized members of their respective clades in terms of machairodonty.

Computerised tomography imaging indicates that the canine roots of SGOPV 3490 were closed at the time of death. The canine roots are closed in adulthood in Sparassodonts except in non-Thylacosmiline Proborhyaenids, in which the upper and lower canines are hypselodont and their roots remain open throughout life. In Thylacosmilines, only the upper canines are hypselodont. However, the closed roots of the lower canines may be a secondary reversal from a hypselodont ancestral condition. Canine pulp cavities in SGOPV 3490 are relatively narrow and lack a well-developed opening. Additionally, the canine roots taper slightly apically rather than being uniformly wide with outwardly flaring edges as is typically observed in open-rooted taxa. Nevertheless, several features suggest that the canines of Eomakhaira were open rooted earlier in ontogeny, closing only in extreme senescence, as in the Proborhyaenid Proborhyaena. First, the canines of SGOPV 3490 lack any trace of enamel, being composed solely of dentine. This contrasts with the typical condition in Sparassodonts with nonhypselodont canines (e.g. Borhyaenids), where some enamel occurs on nonoccluding surfaces near the apex of the tooth (such as the lateral side of the upper canines), even in old individuals. The hypselodont canines of Patagosmilus and Thylacosmilus have enamel, but only in a band covering the labial surface of these teeth rather than a simple cap. Postmortem damage and/or unusual wear are unlikely to account for the complete absence of enamel on all four canines of SGOPV 3940, particularly considering that enamel occurs on the right P1 and p2, located only a few millimeters posterior to the canines. A similar condition (enamel absent on the canines but present on the postcanines) is also observed in the Proborhyaenids Proborhyaena and Callistoe. The absence of enamel in Eomakhaira is consistent with this taxon having had open-rooted canines until near the end of its lifespan.

The upper canines of SGOPV 3490 are tall given their high degree of wear. The upper canine roots extend almost to the dorsal border of the maxilla within their alveoli (measuring over 46 mm in total length), indicating that the height of the exposed portion (23 mm) is natural and not the result of the canine slipping ventrally from the alveolus. Scaled to the anteroposterior length of the tooth, the exposed height of the upper canine in SGOPV 3490 exceeds that of most other Borhyaenoids and far exceeds that of other Sparassodonts with a comparable degree of canine wear. Scaled to M3 length, the upper canine is longer than in almost any other taxon, except those with hypselodont upper canines.

The pulp cavities of the upper and lower canines appear to extend to their apices. The cavity is difficult to discern in the right upper canine, but an area of seemingly less dense, nodule-containing material runs the entire length of the tooth. The pulp cavities of the upper canines are much smaller than those of the lower canines. A pulp cavity extending to the apex of the tooth would be expected if these teeth were worn but hypselodont, as is seen in other Mammals with hypselodont caniniforms (i.e. Choloepus), though this condition can also result from extreme wear in some Sparassodonts without hypselodont canines (e.g. the holotype of Pharsophorus lacerans, MACN-A 52-391).

The left lower canine, which has the best-preserved root among the four canines, may not have a fully closed root. Although its pulp cavity is very narrow along most of its length, a deep depression occurs near its base, where the edges of the root flare outward in cross section. This depression connects with the pulp cavity of the tooth. This is reminiscent of the condition in non-Thylacosmiline Proborhyaenids, all of which have hypselodont lower canines. The lack of a basal tip on the left lower canine of SGOPV 3490 cannot be ascribed to damage, as the base of the alveolus is intact and the edges of the apical depression are smooth and rounded rather than jagged, as they would be if broken. Given the advanced ontogenetic age of SGOPV 3490, details of the root of the lower left canine suggest that it had been open for much of the Animal’s life and had only very recently closed. The conditions of the roots of the upper canines are uncertain, as their bases cannot be confidently distinguished from the surrounding matrix and bone. Together, the observations above suggest that the canines of Eomakhaira were hypselodont throughout most of their ontogeny, with roots closing and tooth growth ceasing only in extremely senescent individuals.

Morphology of the left lower canine root in the holotype of Eomakhaira molossus (SGOPV 3490). (A) Computerised tomography reconstruction in oblique lateral view, showing prominent depression on proximal end of canine and its connection to the pulp cavity. Connection to pulp cavity has been artificially darkened for contrast. (B) Oblique lateral computerised tomography image slice of SGOPV 3490, showing flared, smooth distal end of left lower canine (denoted by arrow). Scale is 10 mm. Engleman et al. (2020).

The roots of the canines in Eomakhaira, particularly the lower canines, are shorter and less curved than those of other Sparassodonts. The upper canine roots end above P3 in Eomakhaira but are still relatively large above P3 in Callistoe and Arminiheringia (based on coronal cross sections), suggesting that the roots extended further posteriorly in those taxa. A condition similar to that of Callistoe and Arminiheringia occurs in Australohyaena (UNPSJB PV 113). The lower canines of SGOPV 3490 are emplaced nearly subvertically within the alveoli, similar to what has been described for most Borhyaenoids, with the notable exception of Arminiheringia, in which the lower canines are procumbent. The lower canine roots are much shorter and more vertical than their upper counterparts and probably end at the level of the p1–2 embrasure. However, they certainly do not extend beyond the anterior root of p2. This differs from Callistoe, Proborhyaena, and Arctodictis, in which the lower canine roots reach the level of p3, and from Arminiheringia, in which they reach the molar row. The lower canine roots are more curved in Callistoe and Arctodictis than in Eomakhaira.

SGOPV 3490 possesses the typical metatherian postcanine dental formula of three premolars and four molars. In the dentary, the alveolar margin of the postcanine dentition is higher lingually than labially, a feature noted in other Sparassodonts. The postcanine toothrow of Eomakhaira is straight in occlusal view, as in most Sparassodonts. In the Thylacosmilines Patagosmilus and Thylacosmilus, the postcanine toothrow is strongly curved and concave medially, but this does not appear to be the case for the more basal Thylacosmiline Anachlysictis. In Arctodictis and Australohyaena, the postcanine toothrow is neither straight nor curved. Instead, the long axes of the molar and premolar rows are offset from one another rather than aligned, a feature often associated with an obliquely oriented P3. This condition is more pronounced in Arctodictis munizi and Australohyaena antiquua than in Arctodictis sinclairi. The condition in the Borhyaenid Acrocyon riggsi is unclear; in the holotype (FMNH P13433), the toothrow is straight, but P3 is slightly oblique. This unusual configuration may be an artifact of poor restoration of the holotype skull. FMNH P13433 is also unusual in having postcanine toothrows that do not diverge posteriorly to form a triangular palate. Posteriorly diverging toothrows characterise every Sparassodont for which the shape of the palate can be determined except Hondadelphys (thought be a basal member of the clade), an observation that supports the interpretation of William Sinclair that the straight postcanine toothrows of FMNH P13433 are an artifact. The long axis of P3 in Eomakhaira is parallel to the long axis of the postcanine toothrow, as in Pharsophorus, Borhyaena, and Proborhyaenids, rather than being obliquely oriented as in Australohyaena and Arctodictis.

As in other Sparassodonts, the upper and lower premolars of Eomakhaira increase in size from P1 to P3 and p1 to p3. However, the relative disparity in sizes among the premolars differs between the upper and lower toothrows. P3 is much larger than P1and P2, which are of similar size. By contrast, p1 is distinctly smaller than p2 and p3, which are of similar size. The first upper and lower premolars are both very small. In lateral view, the dorsal alveolar border of the lower premolars slopes anterodorsally-posteroventrally from p1 to p3, a common pattern also seen in Prothylacynus patagonicus, Pharsophorus cf. Pharsophorus lacerans (MPEF-PV 4170), Borhyaena macrodonta, Arctodictis sinclairi, Australohyaena antiquua, Callistoe vincei, and Proborhyaena gigantea. The premolar row is relatively short in Eomakhaira compared to other Sparassodonts.

The right P1 and both p1s are oriented obliquely relative to the remainder of the toothrow (about 35° anterolabially-posterolingually for both loci), but the left P1 is nearly parallel to the toothrow. Given that the left maxilla is poorly preserved, the anterior root of its P1 is damaged, and natural bilateral asymmetry of tooth orientation has never been documented in Sparassodonta, the orientation of this tooth almost certainly reflects postmortem deformation.

The P1 of Eomakhaira is asymmetric in lateral view, with its main cusp located over the anterior root rather than equidistant between the anterior and posterior roots, as in most Sparassodonts. Similarly asymmetric premolars occur in Allqokirus and Mayulestes but not in Hathliacynids, Borhyaenids, or most basal Borhyaenoids. They do occur in the Proborhyaenids Callistoe and Arminiheringia, the only other Proborhyaenids for which P1 is known. P/p1 is thought to have been lost in Thylacosmilines, but the putative basal Thylacosmiline from La Venta retains P/p1 (as evidenced by the presence of the roots of these teeth in IGM 251108). What remains of p1 in SGOPV 3490 suggests that its apex was more centrally positioned than that of P1, as in other Sparassodonts.

The crowns and roots of P/p2 of Eomakhaira are aligned with the rest of the postcanine toothrow, as in Pharsophorus (though possibly not as in MPEF-PV 4170 from La Cantera, which more closely resembles Arctodictis sinclairi in this respect), Callistoe, Arminiheringia auceta, and Borhyaena, rather than oblique to the toothrow, as in Proborhyaena, Paraborhyaena, Arminiheringia contigua (MACN-A 10317), Arctodictis, Acrocyon, and Australohyaena.

The robust P/p3 bear large, stout roots. Nevertheless, these teeth are proportionally narrower labiolingually than the more bulbous teeth of Borhyaenids (P3 length/width ratio 1.75 in Eomakhaira vs. an average of 1.43 in Borhyaenids) and those of species of Pharsophorus (Pharsophorus lacerans, YPM-VPPU 20551; Pharsophorus tenax, AC 3192). Borhyaenid taxa closely resembling Eomakhaira in other respects (e.g. Arctodictis and Australohyaena) also have the most bulbous P/p3s. The P3 of Eomakhaira is more elongate than that of Callistoe vincei, less elongate than that of cf. Proborhyaena (MLP 79-XII-18-1), but comparable to those of specimens of Arminiheringia. Like P3, the p3 of Eomakhaira is proportionally narrower labiolingually than in most Borhyaenids, the basal Borhyaenoid Plesiofelis, and the Proborhyaenids Proborhyaena and Arminiheringia. The length/width ratio of this tooth is comparable to that in the Proborhyaenids Paraborhyaena and Callistoe, but is less narrow than its counterparts in the basal Borhyaenoids Prothylacynus and Pharsophorus.

The P3 of Eomakhaira is about 13%–19% longer than p3. In most Sparassodonts resembling Eomakhaira (Borhyaenids, Proborhyaenids, and Pharsophorus), P3 and p3 are of similar length. The only Borhyaenoid potentially resembling Eomakhaira in this respect is Proborhyaena gigantea. The P3 of MLP 79-XII-18-1, assigned to Proborhyaena by Mariano Bond and Pascual Rosendo but referred to as cf. Proborhyaena by Engleman et al., is extremely large (nearly 30 mm long, judging from its preserved roots). This is considerably longer than the p3 of the largest known specimen of Proborhyaena gigantea (AMNH 29576, where this tooth is ~24 mm long). If MLP 79-XII-18-1 pertains to Proborhyaena, it implies the P3 of this taxon was more than 50% longer than p3, a more extreme size disparity than in Eomakhaira. The tooth at the P3 locus in Thylacosmilus is also much longer than its p3, but since the upper tooth represents dP3 rather than P3, direct comparisons with other Sparassodonts are not possible. No other Thylacosmilines are known from associated upper and lower dentitions.

The p3 of Eomakhaira appears to have been inclined posteriorly. This is common among Borhyaenoids, occurring also in the basal forms Plesiofelis schlosseri and Pharsophorus lacerans; the Borhyaenids Australohyaena antiquua, Arctodictis sinclairi, and Borhyaena macrodonta; and the Proborhyaenid Proborhyaena gigantea. Most of the enamel is missing from p3 in SGOPV 3490, with only a small patch preserved near the apex of the left p3. It is not clear whether this paucity of enamel represents a normal feature of Eomakhaira or is an artifact of preservation, because in most other Sparassodonts (e.g. Pharsophorus, Borhyaenids) the enamel of p3 extends inferiorly to the same level as in the other postcanine teeth, although in Proborhyaena (AMNH 29576, MACN-A 52-382) and Callistoe (PVL 4187), the enamel of p3 is restricted to the apex of this tooth.

The roots of the lower postcanines of many Borhyaenoids (Borhyaenids, Proborhyaenids, and closely related taxa) are often robust and 'bulbous'. The degree to which this condition is expressed is variable, ranging from taxa with bulbous roots only on p3 (e.g. Borhyaena) to taxa in which the roots of all premolars are bulbous, but the roots of the molars are not (e.g. Thylacosmilus) to those having bulbous roots on all lower premolars and some molars (e.g. Australohyaena antiquua and most Proborhyaenids). The roots of p3 in Eomakhaira are extremely robust and nearly in contact, resulting in little interradicular space. Whether this condition qualifies as bulbous is uncertain, as previous studies have typically defined roots as bulbous when they are wider than the crown in occlusal view, but the crown of p3 is incompletely preserved in SGOPV 3490. Nevertheless, the morphology of the roots of p3 in SGOPV 3490 closely resembles that of Sparassodonts considered to have bulbous roots, such as Arctodictis sinclairi, in which the p2–m2 roots are so swollen that the interradicular space is nearly eliminated. The crowns of right p2 and m1 are preserved in SGOPV 3490, but their roots are not wider than their crowns, suggesting that these roots are not bulbous if prior definitions are strictly applied. The roots of right p2 are comparatively more robust than the roots of the molars but still much less bulbous than in Arctodictis sinclairi. SGOPV 3490 resembles Thylacosmilus in that the roots of the lower premolars are more robust than those of the molars, though in contrast to Eomakhaira the premolar roots of Thylacosmilus are bulbous.

Oblique lateral computerised tomography image slice of SGOPV 3490 along the postcanine tooth row, showing position of lower molars and depth of horizontal ramus. Note how worn surfaces of P3 and m1 closely match one another. Scale is 30 mm. Engleman et al. (2020).

Left M3–4 and the partial crown of right M3 are the best preserved of the heavily worn upper molars. The left M3 appears to have been displaced posterolingually relative to the left M4. In most sparassodonts, the distal tip of the M3 postmetacrista contacts the anterior end of the M4 preparacrista, whereas in SGOPV 3490, the metastylar corner of M3 is located labial to the end of the M4 preparacrista. Nevertheless, the length of the M3 can be roughly estimated based on preserved parts of this tooth. The M3 is so heavily worn that it is essentially pyramidal in shape, bearing only a single, poorly distinguished cusp. This cusp is located near the labial edge of the tooth, suggesting that the stylar shelf was extremely narrow. Based on comparisons with other Sparassodonts, the main cusp represents either the metacone (which is typically the tallest upper molar cusp in Sparassodonts) or the remnants of a single, completely connate, merged paracone and metacone. In most sparassodonts that have a very small paracone (e.g. Borhyaena, Arctodictis, Australohyaena, Prothylacynus, and the proborhyaenid from the Tremembé Formation), this cusp is typically half (or less) the height of the metacone. However, in Callistoe, Arminiheringia, and Patagosmilus, the paracone is very tall (despite its small base), often nearly as high as the metacone (or only slightly lower). The condition in Proborhyaena is ambiguous; the relative heights of the paracone and metacone in the molars of AMNH 29576 are obscured by wear and damage.

Evidence of an anterolabial cingulum, paracone, or stylar cusps on M3 of SGOPV 3490 is lacking. Whether these structures were once present but obliterated by heavy wear cannot be determined. The ectoflexus on M3 is more than 10% the labiolingual width of the tooth, qualifying it as 'deep'. Among short-snouted Borhyaenoids, a deep ectoflexus on M3 occurs in Prothylacynus patagonicus, Proborhyaena gigantea, and Callistoe vincei. By contrast, upper molar ectoflexi are shallow or absent in Pharsophorus tenax and all Borhyaenids (Borhyaena spp., Arctodictis spp., Acrocyon riggsi, Australohyaena antiquua). Although the M3 ectoflexus of Patagosmilus goini is classified by Engleman et al. as 'shallow', it is deeper than in Borhyaenids and Pharsophorus tenax (close to the threshold between 'deep' and 'shallow') and contrasts with the condition in Thylacosmilus atrox and an indeterminate Colhuehuapian Thylacosmiline, in which the ectoflexus is extremely shallow or absent. computerised tomography images show that the M3 roots are extremely splayed in SGOPV 3490. A similar condition occurs in Proborhyaena, Paraborhyaena, and MLP 79-XII-1-1. Other Sparassodonts may also exhibit this condition, but this is difficult to assess without isolated molars or computerised tomography imaging data. A 2007 study led by Francisco Goin considered anterolabially-posterolingually narrow molar roots to characterise Thylacosmilinae. This condition cannot be scored for Eomakhaira due to distortion of these roots in SGOPV 3490.

Left M3–4 of SGOPV 3490 in (A) labial, (B) occlusal, and (C) lingual views; (C) rotated upside down for easier comparison with (A), (B). Anterior to left in all computerised tomography segmentation images. Scale is 3 mm. Engleman et al. (2020).

The M3 protocone of SGOPV 3490 appears to have been extremely small. This inference is based on: (1) the position of the lingual root M3, which is mostly superior to the trigon rather than directly above it, leaving little space for a talon (unlike the condition in sparassodonts with a larger protocone); and (2) the presence of a small but poorly preserved protocone on right M3. On the left M3, the protocone is entirely missing and the lingual face is dominated by a nearly vertical surface. It is unclear whether this feature formed in vivo or resulted from postmortem damage. The flat lingual face of the left M3 does not appear to be due to carnassial rotation, as carnassial rotation in other Sparassodonts produces wear facets that parallel the preparacrista and postmetacrista, whereas in SGOPV 3490, this feature is oblique to these crests. Significantly, this feature does not occur on the left M3, suggesting that its presence on the right tooth is a preservational artifact.

A small, freshly broken area of enamel on the posterolabial face of the left M3 of SGOPV 3490 shows that the enamel is remarkably thin (about 0.06 mm). The enamel seems to be of similar thickness on the lower teeth, but this is uncertain given the lack of other clean breaks and sufficient density contrast and resolution for distinguishing thin enamel from dentine in computerised tomography scans. In sparassodonts, extremely thin molar enamel has been reported in a proborhyaenid (0.17 mm in MLP 79-XII-19-1, cf. Proborhyaena) and large Hathliacynids (0.07–0.10 mm in Acyon and Cladosictis). Enamel is generally thicker in Borhyaenids and basal Borhyaenoids (about 0.23 mm, Prothylacynus patagonicus; about 0.34 mm, Arctodictis sinclari). Postcanine enamel thickness is unknown in Thylacosmilines, but their canine enamel has been reported to be extremely thin. Although the enamel thickness in Eomakhaira is more similar to that in other Proborhyaenids than in other Borhyaenoids, allometry may play a role, given that Eomakhaira is much smaller than both Prothylacynus and Arctodictis.

The best-preserved upper molar in SGOPV 3490, left M4, is very short anteroposteriorly. In this respect, Eomakhaira more closely resembles Patagosmilus (where the M4 is also very narrow) than other short-snouted Borhyaenoids (e.g. Prothylacynus, Pharsophorus, Arminiheringia, Borhyaena, Arctodictis, and Thylacosmilus) in which the M4 is more robust and less anteroposteriorly narrow. The preparacrista is parallel to the axis of the greatest width of the tooth. The M4 crown is oriented obliquely (anterolabially-posterolingually) to the rest of the toothrow. An oblique M4 occurs in many Sparassodonts (Cladosictis patagonica, Acyon myctoderos, Prothylacynus patagonicus, Callistoe vincei, Arminiheringia auceta, Arctodictis sinclairi, Arctodictis munizi, Thylacosmilus atrox) but can be variable (e.g., M4 is oriented obliquely in some but not all individuals of Arctodictis sinclairi and Acyon mycteros). The M4 of SGOPV 3490 is almost as wide or wider labiolingually than M3, even accounting for damage to the latter tooth, a pattern otherwise seen only in Pharsophorus tenax, Arctodictis sinclairi, Patagosmilus goini, and possibly Borhyaena macrodonta among short-snouted Borhyaenoids.

The simple M4 crown of SGOPV 3490 consists of two poorly distinguished trigon cusps, a paracone and stylar cusp B, in addition to an extremely small protocone. There is no metacone. Both the paracone and stylar cusp B are nearly subsumed within the extremely well-developed preparacrista, though this may be exaggerated by wear. There is no anterior cingulum (preparacingulum) on M4. A labial cingulum, which occurs in some Sparassodonts with simplified M4s (e.g. Prothylacynus patagonicus, MACN-A 707), is also absent in Eomakhaira. No vestigial postparacrista occurs posterior to the paracone, the presence of which has been described for Patagosmilus and Arctodictis.

The M4 protocone of Eomakhaira is tiny, barely a swelling of enamel on the lingual side of the paracone. This differs from the condition in Callistoe and Patagosmilus, in which the vestigial protocone is larger and more distinct from the trigon, as well as from Borhyaena macrodonta (MACN-A 32–390) and Pharsophorus tenax (AC 3192), in which the vestigial protocone retains a small basin. The M4 of Proborhyaena is unknown, and the M4 of Paraborhyaena could not be examined firsthand.

Computerised tomography images of the left M4 of SGOPV 3490 show that this tooth has three roots despite its highly simplified morphology.The apices of the roots are located labially, lingually, and posteriorly, with the lingual and posterior roots merging basally, resulting in only two roots at the level of the crown. By contrast, most sparassodonts with a highly simplified ('linear') M4 are considered to have only two roots (but see below), including nearly all short-snouted Borhyaenoids for which the M4 is known (i.e. Acrocyon riggsi, Arctodictis spp., Borhyaena spp., Callistoe vincei, Paraborhyaena boliviana, Patagosmilus goini, Pharsophorus tenax, Prothylacynus patagonicus, and Thylacosmilus atrox). Australohyaena antiquua, the only short-snouted Borhyaenoid with a definitively three-rooted M4, is deeply nested within a clade otherwise characterised by double-rooted M4s. This observation, combined with the fact that the third root in M4 of SGOPV 3490 could be identified only through computerised tomography imagery, raises the question of whether some Sparassodonts currently identified as having a two-rooted M4 might instead have a three-rooted M4.

Cross-sectional computerised tomography image of left M4 of SGOPV 3490 in (A) oblique occlusal and (B) posterior views. Shows presence of three roots. Scale bars is 5 mm. Engleman et al. (2020).

Despite the senescence of SGOPV 3490, there is no evidence of carnassial rotation like that seen in some Sparassodonts. Carnassial rotation results in a unique wear pattern of completely flat wear facets with exposed dentine on the posterolingual faces of M1–3 (extending from the metastyle to the protocone) and the anterolingual face of M4. Such wear facets are not seen in SGOPV 3490. Although the posterolingual corners of the left and right M3s are poorly preserved, these wear facets are clearly absent on the anterolingual face of M4. At the same time, the crowns of right M3 and left M3 and M4 appear to have been medially canted, a feature usually considered indicative of carnassial rotation. This canting is clearly not an artifact of postmortem distortion, as the crowns are slanted in opposite, complimentary directions on the left and right sides. Medially canted molars occur in all other borhyaenoids and hathliacynids, including many species that show no signs of carnassial rotation, even in the oldest individuals (see discussion). This demonstrates that molar canting occurs independently of, and is thus not necessarily indicative of, carnassial rotation. Whereas the posterior upper molars of Eomakhaira are medially canted, the posterior lower molars are laterally canted. Laterally canted posterior lower molars occur in other sparassodonts, such as Australohyaena, Arminiheringia, and Arctodictis. Labial canting of the lower molars is possibly correlated with the lingual canting of the upper molars in many Sparassodonts.

The lower molars of SGOPV 3490 are not strongly imbricated, at least not to the degree seen in the Borhyaenids Australohyaena, Acrocyon, and Arctodictis, or the Thylacosmiline Thylacosmilus. The m3–4 are slightly angled relative to the long axis of the toothrow, comparable in degree of imbrication to that seen in Arminiheringia and Proborhyaena but more imbricated than in Paraborhyaena, Pharsophorus (specifically the holotype, MACN-A 52-391), and possibly Borhyaena (based on MACN-A 52-366, assigned to Borhyaena macrodonta). In lateral view, the alveolar border of the lower molars, as approximated by the bases of the crowns (in the absence of most of the alveolar bone in this region), rises posteriorly at an angle of about 5°–6° relative to horizontal. Several factors suggest this is real rather than taphonomic. First, when the specimen is positioned such that the alveolar border is horizontal, the nasals point anterodorsally, a biologically unreasonable orientation. Second, the right P3/m1 are preserved in occlusion, and the alveolus of the right P3 is partially preserved, indicating that the right dentary has not moved relative to the cranium. Finally, a lower toothrow that rises posteriorly occurs in a few other Sparassodonts, including Arminiheringia auceta, Paraborhyaena boliviana, Arctodictis sinclairi, Australohyaena antiquua, and possibly Acrocyon riggsi. In those taxa, the alveolar border of the lower molars is angled at about 8° relative to horizontal, similar to the inferred angle in SGOPV 3490.

Little can be said about m1–2 of SGOPV 3490. As mentioned above, the occlusal morphology of right m1–2 has been obliterated by wear, whereas on the left side, m1 is missing its crown, and no trace of m2 is preserved, possibly due to greater distortion of the left side of the skull. The posterior lobes of the crowns of m1–2 are not lower than the anterior lobes, a condition that occurs to a variable degree in all borhyaenids, including Prothylacynus, Plesiofelis, Pharsophorus (including Pharsophorus lacerans and Pharsophorus tenax but not MPEF-PV 4170, the specimen from La Cantera assigned to Pharsophorus), Proborhyaena, and Thylacosmilus. In this respect, SGOPV 3490 resembles Arminiheringia. A small posterolabial cingulid occurs on the labial side of m1 in SGOPV. The presence or absence of a posterolabial cingulid cannot be evaluated in m2 or m3, but it is absent on m4. The presence of a labial postcingulid on m1 is not unexpected given its distribution among sparassodonts. Posterolabial cingulids are absent in the basal Borhyaenoids Lycopsis, Pseudothylacynus, and Prothylacynus but are present in cf. Nemolestes (AMNH 29433), Plesiofelis, Pharsophorus (both Pharsophorus lacerans and Pharsophorus tenax), all Borhyaenids (Acrocyon spp., Australohyaena antiquua, Arctodictis spp., and Borhyaena spp.), and the Proborhyaenid Callistoe. This feature could not be scored for Arminiheringia, Proborhyaena, or Paraborhyaena. The condition in Thylacosmilines is not entirely clear. Photographs of the holotype of Anachlysictis gracilis published by Francisco Goin in 1997 appear to show a posterolabial cingulid, whereas figures of Thylacosmilus show a structure that could be a posterolabial cingulid. Most phylogenies of Sparassodonts imply at least six independent losses of the posterolabial cingulid (in Hondadelphys, Stylocynus, Lycopsis, Prothylacynus, and at least twice among Hathliacynids), or repeated loss and reacquisition of the posterolabial cingulid within Sparassodonta.

The right m4 is the best-preserved lower molar in SGOPV 3940 and the only tooth in which crown morphology has not been largely obliterated by wear. Nevertheless, its paracristid still is highly worn, comparable to the degree of wear in the holotype of Angelocabrerus daptes (MMP 967M) and a specimen assigned to Pharsophorus cf. Pharsophorus lacerans (MPEF-PV 4190). As in all borhyaenoids, the m4 of SGOPV 3490 is characterized by two main cusps, a tall protoconid and a slightly shorter paraconid. The m4 paraconid of SGOPV 3940 lacks an anteriorly projecting ventral keel, unlike most Sparassodonts but as in Proborhyaenids and possibly Thylacosmilus. The protoconid is tall relative to the anteroposterior length of the tooth (i.e. the height is greater than 90% the length of the tooth) and is wider at its midpoint that at its base, as in other Sparassodonts. The m4 of SGOPV 3490 closely resembles that of Proborhyaena and Paraborhyaena in having a posteriorly salient protoconid at the posterior end of the tooth and a barely discernable talonid. The latter feature contrasts with m1–3, each of which shows evidence of a talonid that is very small and worn but nevertheless slightly larger.

The metaconid is clearly absent on the m4 of SGOPV 3490. Assessing whether a metaconid was present or absent on m2–3 is more difficult due to the worn and highly fragmented preservation of these teeth, but this cusp appears to be absent on at least m3. In Borhyaenoids with a metaconid (e.g. Borhyaenids, Pharsophorus), this cusp often occurs as a small, low protuberance at the posterolingual corner of the tooth. In SGOPV 3490, on the other hand, the posterolingual surface of m3–4 is smooth, and the base of the protoconid extends to the lingual margin of the tooth; there is no evidence that a distinct metaconid was present. In fact, the posterior margin of m3 is very similar to the holotype of Arminiheringia auceta (MACN-A 10970), consisting of a cuspless ridge that is oriented dorsolingually-ventrolabially.

The anterior root of the posterior lower molars (primarily m3–4) in SGOPV 3490 is much larger and more robust than the posterior one. This condition is also seen in several other Proborhyaenids and Borhyaenids, including Borhyaena macrodonta, Borhyaena tuberata, Arctodictis sinclairi, Arctodictis munizi, Acrocyon riggsi, Acrocyon sectorius, MLP 88-V-10-4 (the Proborhyaenid from Antofagasta de la Sierra), Arminiheringia auceta, Paraborhyaena boliviana, Proborhyaena gigantea, and Thylacosmilus atrox. However, this condition is not present in the Borhyaenid Australohyaena antiquua, the Proborhyaenid Callistoe vincei, and all species of non-Proborhyaenid, non-Borhyaenid Sparassodonts in which the state of the roots of the lower molars could be determined (e.g. Hondadelphys, Hathliacynids, Pharsophorus spp., Prothylacynus patagonicus). Disparity in size between the anterior and posterior roots varies among taxa. In Arctodictis sinclairi and Arminiheringia auceta, only the roots of m3–4 are unequal in size, whereas in Arctodictis munizi, Proborhyaena gigantea, Paraborhyaena boliviana, and Thylacosmilus atrox, it is the roots of m2–4 that are unequal. In SGOPV 3490, the anterior roots of m2–4 are larger than the posterior ones, but the disparity is much less in m2 than in m3–4. Molar roots differ in size among other groups of carnivorous Mammals. The roots of m2–3 are similar in length and robustness in the extant bone-cracking Sarcophilus harrisii, while the anterior root of m4 is slightly more robust than, but the same length as, the posterior one. This size disparity is also present in the lower carnassial (m1) of some carnivorans that are not specialized bone-crackers, including some species of Barbourofelids, Felids, and Cryptoprocta.

An equal-weights analysis produced 12 most-parsimonious trees, each 1624 steps in length, with a consistency index of 0.302 and a retention index of 0.663. The implied-weights analysis with k = 3 produced a single most-parsimonious tree with a best score of 179.21909. The implied-weights analysis with k = 12 produced a single most-parsimonious tree with a best score of 72.67280. Since the topologies of the most-parsimonious trees, in the three analyses are nearly identical, they are discussed together below.

Results of parsimony phylogenetic analysis under equal weights, showing the strict consensus of 12 most parsimonious trees. Eomakhaira molossus in bold. Numbers to upper left of each node represent Bremer supports/decay indices, numbers to lower left of each node represent bootstrap values. Support values not given for Dasyuromorphia, as this node was constrained a priori. Engleman et al. (2020).

In all three analyses, Eomakhaira is recovered as the basalmost member of Thylacosmilinae (basal to a clade of Patagosmilus + Thylacosmilus) within Proborhyaenidae (as defined above). Eomakhaira is recovered as a Thylacosmiline in every equal-weights most-parsimonious tree, despite several characters coded as uncertainties (which results in TNT considering all possible coded character states when determining the most-parsimonious tree). Recovery of Thylacosmilinae within Proborhyaenidae is not due solely to the inclusion of Eomakhaira, as Thylacosmilinae is nested within Proborhyaenidae even when Eomakhaira is excluded from the analysis. Among Proborhyaenids, the clade of Paraborhyaena + Proborhyaena and Callistoe vincei (as a distinct branch) represent successive outgroups to Thylacosmilinae. Proborhyaenidae (including Thylacosmilinae) has a high bootstrap support value (71) and a high Bremer support value (4) in the equal-weights analysis; in both implied-weights analyses (76 in the analysis where k = 3, 71 where k = 12), it has a high bootstrap support value. Proborhyaenidae is recovered as the sister group of Borhyaenidae, and the clade of Proborhyaenidae + Borhyaenidae has high bootstrap support values in all three analyses (84 under equal weights, 92 in the implied-weights analysis where k = 3, and 89 in the implied-weights analysis where k = 12).

Eomakhaira could only be coded for about 19.5% of the characters in this analysis (78 of 400 characters), and accurately coding some characters was hindered by the extreme dental wear and challenging preservation of the specimen. Eomakhaira is almost certainly a member of the clade composed of Borhyaenidae, Proborhyaenidae (including Thylacosmilinae), and their closest relatives (i.e. Pharsophorus) based on the unambiguous synapomorphies preserved (e.g. lingual median sulci on the canines, absence of the postpalatine torus foramen). It is highly likely that it represents a proborhyaenid, though there is a small chance it could instead represent a Borhyaenid or a Pharsophorus-like borhyaenoid. All most-parsimonious trees place it with Thylacosmilinae rather than among other proborhyaenids. Excluding Eomakhaira from Thylacosmilinae requires two additional steps to the most-parsimonious trees and results in the new taxon occupying various more basal positions within Proborhyaenidae (as the most basal Proborhyaenid, as sister to Proborhyaena + Paraborhyaena, etc.). Constraining Eomakhaira to be outside of Proborhyaenidae also requires two steps more than the most-parsimonious trees, placing Eomakhaira as the nearest outgroup to Proborhyaenidae. Constraining Eomakhaira as a Borhyaenid requires five steps more than the most-parsimonious trees, and recovers Eomakhaira as the basalmost member of this group. Constraining Eomakhaira to be outside the clade of Borhyaenidae + Proborhyaenidae also requires five steps more than the most-parsimonious trees, and results in Eomakhaira being recovered as the sister taxon of that clade. In all these analyses, Patagosmilus + Thylacosmilus remain nested within Proborhyaenidae. Constraining Proborhyaenidae (excluding Thylacosmilinae) and Thylacosmilinae to each be monophyletic (leaving Eomakhaira as a floating taxon) requires three steps more than the most-parsimonious trees and recovers Eomakhaira as either the basalmost 'Proborhyaenid' or the basalmost Thylacosmiline in different most-parsimonious trees.

Results of parsimony phylogenetic analysis under implied weights with concavity constant k = 3, showing the single recovered most parsimonious tree. Eomakhaira molossus in bold. Numbers to lower left represent bootstrap values. Support values not given for Dasyuromorphia, as this node was constrained a priori. Engleman et al. (2020).

Deltatheroida and Holoclemensia are recovered as the most basal taxa in this analysis, outside of Theria. The placement of Deltatheroida may reflect the selection of Vincelestes as the outgroup, as Deltatheroidans and other carnivorous Mammals frequently exhibit secondarily simplified dentitions that superficially resemble the nontribosphenic dentition of Vincelestes. Within Marsupialiformes, sparassodonts are recovered crownward of Kokopellia, Asiatherium, Pediomyidae, Alphadon, Stagodontidae, and Mimoperadectes (in the equal-weights analysis), sister to the Pucadelphyidae (Pucadelphys + Andinodelphys) within Pucadelphyida. Pucadelphyida is recovered either as sister to a clade of Herpetotherium + crown group Marsupialia (in the equal-weights analysis and implied-weights analysis with k = 12) or Herpetotherium is recovered as sister to Pucadelphyida and this clade is recovered as sister to crown-group Marsupialia (in the implied-weights analysis with k = 3).

Mayulestes, Allqokirus, and Patene are recovered as the earliest-diverging branches of Sparassodonta, with Mayulestes and Allqokirus as sister taxa and Patene apical to the clade of Mayulestes + Allqokirus but basal to the remainder of the group. This result resembles that of of a recent study by a group led by Caio César Rangel, who recovered Mayulestes, Allqokirus, and Patene as successively diverging lineages at the base of Sparassodonta, but it contrasts with that of a study led by Christian de Muizon, which recovered these three taxa as a monophyletic clade. Englman et al. agree with the assessment by Rangel et al. that the recovery of Mayulestidae in Muizon et al. is due to the latter study optimizing sparassodont or pucadelphyidan symplesiomorphies as apomorphies of Mayulestes, Allqokirus, and Patene.

Results of parsimony phylogenetic analysis under implied weights with concavity constant k = 12, showing the single recovered most parsimonious tree. Eomakhaira molossus in bold. Numbers to lower left represent bootstrap values. Support values not given for Dasyuromorphia, as this node was constrained a priori. Engleman et al. (2020).

In all three of Engleman et al.'s analyses, Hondadelphys and Stylocynus form a well-supported clade (Bremer support of 5 and bootstrap support of 71 in the equal-weights analysis, bootstrap support of 59 in the implied-weights analysis with k = 3, bootstrap support of 70 in the implied-weights analysis with k = 12) that is apical to Mayulestes, Allqokirus, and Patene but basal to Hathliacynidae + Borhyaenoidea. This contrasts with previous analyses, in which these two taxa were recovered as successively diverging branches just outside of Hathliacynidae + Borhyaenoidea. If the grouping of Hondadelphys + Stylocynus is upheld in future analyses, Hondadelphidae might be the appropriate name for this clade. Hondadelphys and Stylocynus are united by several apomorphies, including a metacone positioned lingual to the paracone, well-separated paracone and metacone, large protocone, absence of StB, talonid wider than the trigonid, relatively short protoconid, and transverse posthypocristid. Several of these traits are often considered metatherian symplesiomorphies, but in the current analysis, the character states of Hondadelphys and Stylocynus appear to be synapomorphic reversals within Sparassodonta. This is supported by the observation that these features are not observed in any Palaeogene Sparassodont (including additional taxa not considered in the present phylogenetic analysis, e.g. Patene coluapiensis, Procladosictis anomala). Some of these features may be functionally linked (i.e. width of the talonid and size of the protocone) or correlated with omnivory, but others are not or are unique to Hondadelphys + Stylocynus within Sparassodonta. A clade of Hondadelphys + Stylocynus is also more congruent with the stratigraphic record of these taxa than previous phylogenetic analyses. Hondadelphys and Stylocynus date to the middle and late Miocene, respectively, and previous phylogenies that recovered them as successively diverging branches basal to Hathliacynidae + Borhyaenoidea imply a separate ghost lineage for each that extends back to the middle Eocene. By contrast, the present phylogeny requires only a single ghost lineage extending into the Paleogene (representing the common ancestor of the Hondadelphys + Stylocynus clade).

Relationships within Hathliacynidae are slightly better resolved in this analysis than in previous studies. The group is recovered as monophyletic with two major subclades in all analyses: one that includes the large-bodied Hathliacynids Cladosictis and Acyon, and the other that consists a polytomy of Sallacyon, Notogale, and Sipalocyon. In the implied-weights analyses, the polytomy is resolved, with Sallacyon as the sister to Notogale + Sipalocyon. There has been little consensus regarding relationships within Hathliacynidae, as noted previously. Some studies have even failed to recover a monophyletic Hathliacynidae.

The Borhyaenoid Lycopsis is not unambiguously recovered as monophyletic. In some most-parsimonious trees of the equal-weights analysis, Lycopsis longirostrus is sister to remaining species of Lycopsis (resulting in a monophyletic Lycopsis). In others, Lycopsis longirostrus is sister to all other Borhyaenoids (including other species of Lycopsis, which form their own clade distinct from Lycopsis longirostrus and all other Borhyaenoids), resulting in a paraphyletic Lycopsis. Thus, Lycopsis longirostrus forms part of a basal Borhyaenoid polytomy in the consensus tree. In the implied-weights analysis with k = 3, the four species of Lycopsis form a series of successively diverging branches at the base of Borhyaenoidea. Lycopsis is also recovered as paraphyletic in the implied-weights analysis with k = 12; Lycopsis longirostrus is at the base of Borhyaenoidea, and the remaining species of Lycopsis form a monophyletic clade. The paraphyly of Lycopsis is not surprising, as this taxon is not currently diagnosed by any synapomorphies uniquely shared by its four species (Lycopsis torresi, Lycopsis longirostrus, Lycopsis padillai, and Lycopsis viverensis) to the exclusion of other Sparassodonts. Instead, Lycopsis has been diagnosed by general Borhyaenoid apomorphies and a combination of features that are plesiomorphic relative to later-diverging Borhyaenoids (e.g. Prothylacynus, Borhyaenidae, Proborhyaenidae). Even in cases where Lycopsis has been recovered as monophyletic, its monophyly is only supported by two characters: an infraorbital foramen located over the anterior root of P3 and p3 with an anterior edge more convex than the posterior edge. Neither of these character states are unique to Lycopsis spp. among sparassodonts nor definitely present in all four species referred to this genus (the former cannot be coded in Lycopsis padillai and Lycopsis torresi, and the latter cannot be coded in Lycopsis padillai and Lycopsis viverensis). Therefore, Lycopsis sensu lato may represent a grade of basal Borhyaenoids rather than a monophyletic group.

Compared to other Borhyaenoids, Eomakhaira molossus is notable for its small size. Regression equations of the lower dentition from Troy Myers and Cynthia Gordon yield body mass estimates of 9.5–10 kg for Eomakhaira molossus, comparable to a male Tasmanian Devil (Sarcophilus harrisii). The holotype specimen, SGOPV 3490, is comparable in size to a skull of Sarcophilus, though SGOPV 3490 is deeper and narrower. This unusual shape is unlikely to be attributable to postmortem compression. While the skull has been subjected to shear deformation, there are no signs of significant dorsoventral or mediolateral crushing.

Eomakhaira is by far the smallest known Palaeogene Proborhyaenid, its estimated body mass being only about 40% of that of the next smallest, Callistoe vincei (about 23 kg). Such a small Proborhyaenid is quite unexpected during the Oligocene, given that non-Thylacosmiline Proborhyaenids reached their largest size during this interval. The two previously described Oligocene Proborhyaenids, Proborhyaena and Paraborhyaena, both likely exceeded 50 kg in body mass. The small size of Eomakhaira indicates that proborhyaenids exhibited significantly more ecomorphological diversity than previously recognised during the Oligocene. This conclusion is corroborated by an unnamed Proborhyaenid from the late Oligocene (Deseadan South American Land Mammal Age) locality of Taubaté that is comparable in size to Arminiheringia auceta, making it much larger than Eomakhaira but still smaller than Paraborhyaena and Proborhyaena (probably about 30–40 kg based on comparison with Arminiheringia auceta). There are two possible explanations for the high ecomorphological diversity of Proborhyaenids during the Oligocene. First, Oligocene Proborhyaenids could represent holdovers from a much older unsampled Eocene radiation, of which only a few lineages (such as those that gave rise to Proborhyaena + Paraborhyaena and Thylacosmilinae) survived into the Oligocene. Alternatively, the high Oligocene diversity of Proborhyaenidae might reflect an Oligocene radiation related to faunal turnover at the Eocene-Oligocene transition, of which gigantism and small size/incipient machairodonty were but two evolutionary experiments. South American Metatherians underwent a large faunal turnover during the Oligocene, likely in response to climatic change. This turnover (termed the Bisagra Patagónica) has been associated with the Eocene-Oligocene transition by previous authors and has been considered analogous to the Grande Coupure in Europe and the Mongollian Remodelling in Asia. However, the peak of the turnover does not correlate with the Eocene-Oligocene transition (and the faunal events associated with it on other continents) but instead occurs nearly 4 million years later, during the 'middle' Oligocene (about 30.6–30.7 million years old; 'Canteran'). Regardless of its exact correlation with the Eocene-Oligocene transition, this Oligocene turnover resulted in the extinction of many formerly dominant Metatherian groups and subsequent radiation of the survivors. Among Sparassodonts, these changes included the appearance of Hathliacynids and Borhyaenids in the late Oligocene. The extinction of non-Thylacosmiline Proborhyaenids at the end of the Oligocene has been considered an example of the long-term decline of a previously dominant group stemming from climatic changes during the Oligocene. However, given that gigantism and machairodonty do not occur in middle Eocene Proborhyaenids such as Callistoe or Arminiheringia, Proborhyaenids may have radiated in response to the environmental changes and faunal turnover during the Oligocene in a manner similar to Hathliacynids and Borhyaenids. The fossil record of Proborhyaenids is too incomplete at present to favor either hypothesis, though they both represent avenues for future research.

Size comparison of representative Palaeogene Proborhyaenids. From largest to smallest, Proborhyaena gigantea (in blue), the largest known Proborhyaenid (scaled after AMNH 29576, the largest specimen of this taxon); Callistoe vincei (in green), the smallest named Proborhyaenid prior to this study (scaled after the holotype specimen, PVL 4187); Eomakhaira molossus (in red), scaled after SGOPV 3490. Homo sapiens (170 cm tall) to right for comparison. Scale bar is 1 m. Engleman et al. (2020).

The small size of Eomakhaira raises the possibility that an isolated m4 from the late Eocene (Mustersan) of Antofagasta de la Sierra in northwestern Argentina (MLP 88-V-10-4) belongs to the same taxon or a closely related form. Originally assigned to Arminiheringia, later authors have referred this specimen to Callistoe based on its smaller size and reduced talonid. Although MLP 88-V-10-4 is larger than the corresponding tooth of the holotype of Eomakhaira (m4 length is 14.0 mm in MLP 88-V-10-4 versus 12.0 mm in SGOPV 3490), it is smaller than the corresponding tooth in the holotype of Callistoe (length 17 mm). MLP 88-V-10-4 also more closely resembles SGOPV 3490 in the more posterior position of its protoconid and near absence of a talonid; in Callistoe, the protoconid is positioned more anteriorly and is less salient posteriorly, and the talonid is larger. Furthermore, MLP 88-V-10-4 lacks an anterolabial cingulid, similar to Proborhyaena, Paraborhyaena, and Thylacosmilus (but apparently not the Thylacosmiline Anachlysictis). This suggests that MLP 88-V-10-4 cannot be assigned to either of the currently recognised genera of Eocene Proborhyaenids, Callistoe or Arminiheringia, in which an anterolabial cingulid is present on m4. While the size and morphological resemblance support a potential close relationship with Eomakhaira, this is not definitive, as the state of the anterolabial cingulid cannot be determined in SGOPV 3490.

The small size of Eomakhaira relative to other Proborhyaenids suggests that Thylacosmilines were ancestrally characterised by small body size compared to other members of this group. Eomakhaira is small (9.5–10 kg) not only relative to Palaeogene Proborhyaenids but also to many late Cainozoic Thylacosmilines. The Proborhyaenid taxa stemward of the base of Thylacosmilinae are all relatively large (over 20 kg), with the smallest of these species, Callistoe, being two and a half times larger than Eomakhaira. The only members of Proborhyaenidae (including Thylacosmilines) comparable to Eomakhaira in size are other geologically old Thylacosmilines such as the Colhuehuapian Thylacosmiline from Gran Barranca and the possible plesiomorphic Thylacosmiline from La Venta (the latter being substantially smaller than Eomakhaira). The slightly larger Patagosmilus goini is estimated to have been less than 20 kg. If MLP 88-V-10-4 from the late Eocene of Antofagasta de la Sierra also pertains to Eomakhaira or a closely related form, it would further support the idea that Thylacosmilines were ancestrally characterised by small body size, as MLP 88-V-10-4 is slightly larger than SGOPV 3490, potentially implying a decrease in size across the Eocene-Oligocene boundary. The large body size and highly specialised morphology of non-Thylacosmiline Proborhyaenids has previously been used to argue against a close relationship between Thylacosmilines and Proborhyaenids sensu stricto, as large, morphologically specialised carnivorous Mammal clades generally exist for short timespans (under 10 million years) rather than continue to diversify for millions of years. Ancestrally small body size in Thylacosmilines may have circumvented this pattern and allowed for a later, secondary acquisition of large body size in this clade later in the Cainozoic.

Several features suggest that Borhyaenids and non-Thylacosmiline Proborhyaenids used p3, the largest and most robust premolar in all Sparassodonts, to crack bones. These include an interlocking or fused mandibular symphysis, deep dentary, bulbous premolars (especially P/p3) with long roots, enamel microfractures, and high estimated bite force. Contrary to a possible first impression of this robust-skulled new taxon, Eomakhaira does not seem to have been specialised for bone cracking. Previous authors have noted that mandible depth in bone-cracking carnivorous Mammals is often that same below the primary carnassial and the bone-cracking premolar, resulting in the ventral border of the horizontal ramus of the dentary to appear straight in lateral view. More specifically, a 2010 study headed by Paul Palmqvist considered a uniformly deep mandible with a straight ventral border between the bone-cracking premolar and primary carnassial to distinguish obligate scavenging Hyaenids (Hyaena hyaena and Parahyaena brunnea) from facultative scavengers that also hunt for prey (i.e. Crocuta crocuta, wherein the dentary is deeper beneath the carnassial than the bone-cracking premolar). This observation may not be broadly applicable, however, given that it is only based on extant Hyaenids; extinct hyaenid species may have differed in their dietary habits despite similar mandible morphology. Moreover, since Hyaena and Parahyaena are considered sister taxa among extant Hyaenids, it is not clear whether their similar mandible shape reflects functional morphology or shared ancestry. In extant Metatherians, a uniformly deep dentary with a straight ventral border between the primary crushing tooth and carnassial is present not only in bone-cracking Sarcophilus but also in non-bone-cracking Dasyurus. Thus, this feature cannot be used to securely distinguish obligate bone-cracking scavengers from facultative bone-crackers.

In Sparassodonts, a deep mandible with a straight ventral border and roughly uniform depth between p3 and m4 occurs in Australohyaena, Proborhyaena, Paraborhyaena, and Arctodictis, and these taxa have all been considered to be bone-crackers. By contrast, the dentary is much shallower under p3 than under m4 in Eomakhaira, and the ventral border of the mandible is curved in lateral view. The symphysis of Eomakhaira is also much less extensive than in presumed bone-cracking Sparassodonts. In most such Sparassodonts, the symphysis extends posteriorly to at least the main bone-cracking premolar, often to the p3/m1 embrasure and sometimes to the posterior root of m1 (in Arminiheringia auceta). In the extant bone-cracking Dasyuromorphian Sarcophilus, the symphysis extends to the midpoint of m1, which is functionally analogous to the p3 of Sparassodonts. In Eomakhaira, the symphysis ends approximately at the level of the p2/p3 embrasure. Also, the roof of the skull is not vaulted at the level of the primary bone-cracking teeth in Eomakhaira, in contrast to the typical condition in other bone-cracking Mammals, including Australohyaena, in which the nasals are vaulted.

Several metrics have been used to examine dietary habits in carnivorous Metatherians, primarily degree of specialisations for bone-cracking (durophagy) and/or a carnivorous diet (i.e. hypercarnivory, mesocarnivory, etc.). Five of these parameters (premolar shape, relative premolar length, relative premolar size, relative trigonid length, and relative grinding area) can be applied to Eomakhaira. Premolar shape of the putative bone-cracking tooth (width/length of p3) reflects the robustness of the last lower premolar; it is 0.54 in Eomakhaira, slightly below the threshold separating bone-cracking from non-bone-cracking forms (bone-crackers over 0.58). Relative premolar size (width of p3/cube root of body mass in kg) is 1.32, which is substantially lower than the threshold between bone-cracking and non-bone-cracking taxa (bone-crackers over 2.6). The relative premolar length (length of p3/length of m4) is 0.63, below the threshold of hypercarnivores (hypercarnivores over 0.7). However, extant Metatherian hypercarnivores such as Sarcophilus harrisii and Dasyurus maculatus, as well as many fossil Sparassodonts exhibiting other specialisations considered related to hypercarnivory (Callistoe, Arminiheringia, Australohyaena), also fall below this threshold, suggesting that this parameter is not applicable across Metatherians.

With regard to the molars, the relative trigonid length (m4 trigonid length/total m4 length) of Eomakhaira is 0.91, within the range of specialized ('Catlike') hypercarnivores and comparable to some of the most specialised carnivores within Sparassodonta, including the Borhyaenoid Angelocabrerus; the Borhyaenids Australohyaena, Arctodictis, and Acrocyon; the Proborhyaenids Proborhyaena and Paraborhyaena; and the Thylacosmiline Thylacosmilus. Previous authors have considered a relative trigonid length of of greater than 0.9 to indicate Catlike hypercarnivory and a relative trigonid length of 0.8–0.9 to indicate hypercarnivory with bone-crushing specializations. However, only three extant bone-crushing taxa (Crocuta crocuta, Hyaena hyaena, and Parahyaena brunnea; all Placental Hyaenids) were included in the comparative dataset of those studies. Hyaena and Parahyaena are unusual among extant large-bodied hypercarnivores in having small but functional talonids on m1, whereas Crocuta more closely resembles Felids in its longer trigonid and a near-vestigial talonid (and has a relative trigonid length over 0.9). This difference may be related to the substantial amount of fruit consumed by Parahyaena and Hyaena (about 12%–20% of diet by volume) but not by Crocuta. Molar trigonid length is unlikely to be correlated with bone-cracking habits in Hyaenids, as Hyenas primarily employ premolars in this function. Additionally, the relative trigonid lengths of the extant non-bone-cracking hypercarnivorous Canids Cuon, Lycaon, and Speothos are only 0.72–0.74. The narrow range of relative trigonid lengths in living bone-cracking Carnivorans (0.8–0.9) most likely reflects their low diversity (4 species, the three Hyaenids and Sarcophilus harrisii) and phylogenetic signal rather than any unique functional association; three of the four extant bone-crackers (Crocuta crocuta, Hyaena hyaena, and Parahyaena brunnea) belong to the same clade, and this range is within the variation seen in Mammalian hypercarnivores more generally.

Relative grinding area has been assessed via two methods in Sparassodonts, one using only m4 and the other assessing the entire lower molar row. Although the latter method is optimal, m1–3 in the holotype of Eomakhaira are too heavily worn to calculate relative grinding area in this manner. The m4 of Eomakhaira most closely resembles that of Proborhyaena, Paraborhyaena, Arctodictis, and Thylacosmilus, all of which were characterised as lacking a functional talonid and therefore considered to have an relative grinding area of 0. Evaluated together, the short mandibular symphysis, shallow dentary below p3, labiolingually narrow premolars that are small in proportion to those of bone-cracking taxa, long trigonid, and very small talonid of Eomakhaira suggest it was hypercarnivorous but not clearly specialized for bone crushing.

In 1969 James Mellet described an unusual phenomenon in the Hyaenodont Hyaenodon in which the upper molars become progressively more medially oriented throughout ontogeny (carnassial rotation). Mellet also reported the occurrence of carnassial rotation in the Hyaenodont Hemipsalodon, the Oxyaenid Patriofelis, and 'an unnamed Pliocene Marsupial saber-tooth'; almost certainly referring to Thylacosmilus atrox, given that no other well-preserved Thylacosmilines were known at the time). Carnassial rotation has been documented in extinct Carnivorans, including the Barbourofelid Barbourofelis and the Nimravid Hoplophoneus. In 1978 Larry Marshall reported carnassial rotation in several Sparassodonts, most prominently Arminiheringia auceta, but also to a lesser degree in Acrocyon, Arctodictis, and Borhyaena. In 1983 Mariano Bond and Pascual Rosendo described carnassial rotation in a senescent specimen they assigned to Proborhyaena (MLP 79-XII-18-1). Carnassial rotation in the Placental Hyaenodon and the Metatherian Arminiheringia produces a very distinct form of wear in which the entire posterolingual face of the upper molars is worn flat and the dentine is exposed, even on the talon. This produces a sharp, flat edge with vertical wear facets that roughly parallels the main shearing blade of the tooth (typically the postmetacrista of upper molars) but relatively little apical wear. In the holotype of Arminiheringia auceta (MACN-A 10970/10972), in vivo wear had progressed to the point that the pulp cavities of M1–3 were exposed. Additionally, in the holotype of Arminiheringia auceta, a second flat wear facet is present on the anterolingual face of M1–4, roughly parallel to the preparacrista. These facets are not present in Hyaenodon.

Aside from Arminiheringia, the only other Sparassodonts for which true carnassial rotation appears to be present are the Proborhyaenid Callistoe vincei and Thylacosmiline Thylacosmilus atrox. The holotype of Callistoe vincei (PVL 4187) exhibits the same vertically flat wear facets seen in Arminiheringia on the entire posterolingual faces of M2–3 and anterolingual faces of M2–4 (M1 not being preserved in this specimen), including the talons. In Thylacosmilus, FMNH P14531 exhibits vertically flat wear facets on the posterolingual face of M3 and anterolingual faces of M4 (as determined from a cast of this specimen). Francisco Goin and Pascual Rosendo found no evidence of ontogenetic carnassial rotation in Thylacosmilus, perhaps because they examined ontogenetically younger specimens (with less-worn teeth) than FMNH P14531, which pertains to an older individual with a highly worn dentition. Unlike Larry Marshall, Engleman et al. did not observe carnassial rotation in any Borhyaenid. The dentitions of Borhyaenids are not characterised by flat wear facets across the entire lingual faces of the teeth; rather, wear facets are predominantly apical and follow the major cusps and crests of the crowns, as in most other faunivorous Metatherians (e.g. Didelphis). Mariano Bond and Pascual Rosendo reported carnassial rotation in cf. Proborhyaena (MLP 79-XII-18-1) based on lingual canting of the upper molars and labial canting of the lower molars. However, as discussed below, neither of these features is necessarily indicative of carnassial rotation. The molars of MLP 79-XII-18-1 are too damaged to determine whether flat wear facets similar to those seen in Arminiheringia and Thylacosmilus were present. Carnassial rotation does not characterise all Proborhyaenids despite its presence in Callistoe, Arminiheringia, and Thylacosmilus. Upper molars of Proborhyaena gigantea (AMNH 29576) clearly do not exhibit carnassial rotation. Wear facets on the upper molars of AMNH 29576 are restricted to their occlusal faces, as in most other Sparassodonts.

Although only a handful of Sparassodonts exhibit true carnassial rotation, many Sparassodonts do exhibit upper molars that are medially canted. This condition can be identified by measuring the angle between the bases of the crowns of the posterior upper molars (M3–4) and the horizontal plane in posterior (distal) view; Engleman et al. consider an angle greater than 35° indicative of canted molars. In general, the posteriormost upper molars (M3–4) show the highest degree of canting, whereas more anterior molars (e.g., M1) are less canted and often comparable in orientation to the canines and premolars. In taxa with canted upper molars, the paracone and metacone are typically canted medially, and the occlusal faces of the upper molars are partially or completely obscured when the palate is in ventral view, sometimes to the point that the protocone is hidden by the trigon. In effect, this means that the upper molars are not oriented in occlusal view when the palate is viewed ventrally, something that can readily be seen when Sparassodont skulls are figured in ventral view. In addition to SGOPV 3490, molars canted at angles greater than 35° are present in the Hathliacynids Acyon myctoderos, Cladosictis patagonica, and Sipalocyon gracilis, and in the Borhyaenoids Acrocyon riggsi, Arctodictis sinclairi, Prothylacynus patagonicus, Pharsophorus tenax, and Arminiheringia sp. By contrast, the angle between the palate and the crowns of M3–4 is 10°–25° in Allqokirus australis, Patene simpsoni, Patene coluapiensis, Hondadelphys fieldsi, and UF 27881 (as indicated by the alveoli of M2–3). Both states contrast with the condition in Metatherians such as Didelphis, where the crown bases lie nearly level with the palate (angle below 10°). The upper molars are also medially canted in Australohyaena antiquua, Borhyaena spp., Callistoe vincei, cf. Proborhyaena, Paraborhyaena boliviana, and Thylacosmilus atrox, but the precise angle cannot be determined from available photographs and published illustrations (which typically do not show the palate in posterior view). All undoubted members of Hathliacynidae + Borhyaenoidea that could be observed directly exhibit strongly medially canted upper molars.

Although the upper molars of many Sparassodonts are medially canted, they do not appear to have undergone carnassial rotation as seen in Hyaenodon and Arminiheringia. Few Sparassodonts exhibit the distinctive wear pattern observed in Arminiheringia, even in older individuals, suggesting that molar orientation changed little after eruption in those taxa. This interpretation is supported by MACN-A 5931 (Prothylacynus patagonicus) and MLP 82-V-1-1 (an undescribed specimen referred to Arminiheringia sp.), both subadults with M/m4 still erupting and little to no wear on M3. In these individuals, M3 is already medially canted at an angle comparable to that seen in adult specimens, and M4 is erupting in a canted orientation. Additionally, MLP 82-V-1-1 bears flat wear facets on the posterolingual faces of M1–2 similar those of the (adult) holotype of Arminiheringia, indicating that carnassial rotation can be identified early in ontogeny in Sparassodonts for which this condition is present. Thus, the upper molars of most Sparassodonts appear to have erupted in a canted position rather than having rotated into that position later in life.

The morpho-functional significance of medially canted upper molars in Sparassodonts is unclear, given that it appears to be uncorrelated to carnassial rotation. James Mellett suggested that carnassial rotation, as occurs in Hyaenodon, maintained precise occlusion between the carnassials despite wear and prolonged the functional lifespan of the dentition in this taxon. According to Mellett, a carnivorous Mammal like Hyaenodon, with multiple shearing teeth, anisognathus lower jaws, and fused mandibular symphysis, cannot compensate for tooth wear via lateral motion of the lower jaw (as occurs in forms with unfused symphyses, such as most extant carnivorans), and carnassial rotation provided an analogous functional solution. However, medially canted upper molars occur in many Sparassodonts with a ligamentous symphysis (e.g., Acyon, Cladosictis, Sipalocyon), which provides some symphyseal flexibility. The same is true for Barbourofelis and Nimravid Carnivorans. Although Jon Baskin suggested that the long vertical symphysis of Barbourofelis would have been functionally equivalent to a fused symphysis, Harold Bryant and Anthony Russell suggested that this taxon was capable of lateral lower jaw mobility. Furthermore, Engleman et al. note that the upper molars of immature specimens of Arminiheringia sp. and Prothylacynus patagonicus are medially canted, M4 erupts in a canted orientation, and the angle between the base of the molar and the palate in these specimens is comparable to that of adult Sparassodonts (including other specimens of the same taxa). This indicates that medial canting of the upper molars was not attained gradually over an Animal’s lifespan, as might be expected if canting maintained precise occlusion during wear, since M3–4 are already canted while they erupt. Finally, the lower molars of many Sparassodonts are laterally canted (e.g. Australohyaena antiquua), a condition also present to a lesser degree in Dasyurus. The lower molars of Hyaenodon are not canted, suggesting that the functional explanation for carnassial rotation in Hyaenodon by Mellett may not be applicable to Sparassodonts (except possibly to Arminiheringia, Callistoe, and Thylacosmilus). Bryant and Russell suggested that canting in Dinictis and carnassial rotation in Hoplophoneus (medial canting and carnassial rotation do not cooccur in these taxa) counter heavy wear stemming from the dental morphology and more posterior location of the carnassials of these taxa compared to other Carnivorans. Such reasoning would not apply to most Metatherian Carnivores (except Thylacosmilus and possibly Patagosmilus), whose primary carnassials (m4) are located at the anteroposterior midpoint of the mandible, as in Carnivorans.

The Mammaliaform Morganucodon exhibits medially canted posterior upper molars and uncanted anterior molars, similar to what is described here for Sparassodonts. Morganucodon also resembles many Sparassodonts in having prominent pits on the maxilla that receive the main cusp of the lower molars and posterior upper molars positioned lingual to the lateral edge of the maxilla (i.e. 'maxillary cheeks'), which suggest analogous jaw mechanics. These features have been argued to minimize 'roll' during mastication in Morganucodon, allowing precise occlusion despite mainly orthal jaw movements. However, canting of the upper molars in Morganucodon is interpreted to have increased tooth wear rates, which runs counter to the view that upper molar canting prolongs the functional lifespan of the dentition.

Alternatively, canting of the posterior molars in Sparassodonts may be related to their extremely tall lower molar protoconids compared to those of other carnivorous Metatherians. Canting may be a way to accommodate these tall teeth when the mouth is closed. The embrasure pits on the palate of many Sparassodonts have been suggested to represent a similar adaptation to accommodate the extremely tall protoconids.

Ever since the first well-preserved specimens of Thylacosmilines were described, the manner in which these animals acquired their distinctive, highly specialised machairodont morphology has been debated, as has their relationship to other Sparassodonts. Based on various similarities William Berryman Scott tentatively linked Thylacosmilus and the Eocene Proborhyaenid Arminiheringia, but this idea was questioned based on substantial morphological and temporal differences between Thylacosmilus and non-Thylacosmiline Proborhyaenids. Early workers were hampered in their attempts to link Thylacosmilines to other Sparassodonts because the only well-known member of the group was Thylacosmilus atrox, its geologically youngest and most autapomorphic member, which shared few features with other taxa known at the time. Only much later did earlier-diverging forms with less extreme machairodont specializations come to light. Francisco Goin described two taxa from the middle Miocene locality of La Venta, Colombia, (Laventan South American Land Mammal Age) exhibiting less machairodont specialisations than the Mio-Pliocene Thylacosmilus: the Thylacosmiline Anachlysictis gracilis and a second taxon, questionably assigned to the group (IGM 251108). More recently, even older Thylacosmiline remains have been described from the early Miocene (Colhuehuapian South American Land Mammal Age) and early middle Miocene (Colloncuran South American Land Mammal Age) of Patagonia. Paradoxically, these older Patagonian taxa appear to be more closely related to Thylacosmilus than to the taxa from La Venta, even though Thylacosmilus and the La Venta taxa are closer in age; this implies an older (likely pre-Miocene) origin of the clade.

Engleman et al.'s phylogenetic analysis recovers the early Oligocene Eomakhaira as the basalmost member of a clade that includes the Thylacosmilines Patagosmilus and Thylacosmilus, which collectively are nested within Proborhyaenidae. The idea of a close relationship between Thylacosmilines and Proborhyaenids sensu stricto is not novel. Several studies have recovered Proborhyaenids sensu stricto and Thylacosmilines as sister groups, and several others have recovered Thylacosmilinae within a paraphyletic Proborhyaenidae. Leonardo Carneiro recovered a paraphyletic Proborhyaenidae, with Paraborhyaena as sister group to Thylacosmilinae, but also found Borhyaenidae nested within this group, with Callistoe and Arminiheringia recovered as basal to a clade composed of Borhyaenidae + (Paraborhyaena + Thylacosmilinae). Support for a close relationship between Thylacosmilinae and Proborhyaenidae sensu stricto has not been universal in previous studies, and placements for Thylacosmilinae outside of Proborhyaenidae have also been suggested. Recovery of Eomakhaira as an Oligocene Thylacosmiline not only breaks up the long ghost lineage between Thylacosmilines and non-Thylacosmiline Proborhyaenids but also corroborates the nesting of Thylacosmilines within Proborhyaenidae, given that Eomakhaira exhibits a combination of derived features occurring in other Thylacosmilines and plesiomorphic features retained in non-Thylacosmiline Proborhyaenids.

Eomakhaira resembles non-Thylacosmiline Proborhyaenids and differs from other Thylacosmilines in retaining lingual sulci on the upper canines, three premolars, replacement of dP3, and absence of a genial flange. Its P1 is asymmetric, which is likely ancestral for Proborhyaenids given its presence in Callistoe and Arminiheringia; the tooth is unknown in Proborhyaena and Paraborhyaena and is interpreted as absent/lost in other Thylacosmilines. SGOPV 3490 lacks enamel on the labial surface of the canines, which suggests that enamel, if present in Eomakhaira, was a simple cap lost through wear as in non-Thylacosmiline Proborhyaenids rather than a persistant band extending to the base of the tooth as in Patagosmilus and Thylacosmilus. Finally, Eomakhaira may have had open-rooted (hypselodont) lower canines, as in non-Thylacosmiline Proborhyaenids but unlike Thylacosmilines. Its lower molars lack an anteriorly projecting ventral keel on the paraconid, as in non-Thylacosmiline Proborhyaenids (present in Anachlysictis but evidently absent in Thylacosmilus among Thylacosmilines).

On the other hand, Eomakhaira has canines that lack longitudinal grooves, as other Thylacosmilines (and, possibly, Lycopsis viverensis). The upper canines are narrow labiolingually compared to other Sparassodonts, have a well-developed median keel, and lack a labial median sulcus. The maxilla of Eomakhaira is deep, as in other Thylacosmilines (a condition acquired independently in Borhyaenids), rather than shallow as in Proborhyaenids. The dentary of Eomakhaira is intermediate in depth between those of other Thylacosmilines and non-Thylacosmiline Proborhyaenids, shallower than in the Eocene Proborhyaenids Callistoe and Arminiheringia (though possibly not the Oligocene Paraborhyaena and Proborhyaena) but deeper than in the Thylacosmilines Anachlysictis and Thylacosmilus. The infraorbital foramen of Eomakhaira is located at the P3/M1 embrasure, a position more similar to Thylacosmilines than most non-Thylacosmiline Proborhyaenids (except possibly Proborhyaena), in which it is located more anteriorly. The mandibular symphysis of Eomakhaira also more closely resembles that of Thylacosmilines than non-Thylacosmiline Proborhyaenids. In most Borhyaenoids (especially non-Thylacosmiline Proborhyaenids), the symphysis is broad anteroposteriorly and typically fused in adults. In Eomakhaira, the symphysis is unfused and much narrower, probably ending at the p2/3 embrasure (but perhaps as far posterior as the anterior root of p3). P3 of Eomakhaira is much longer than p3, and the lower premolar row is relatively short. Both conditions are reminiscent of other Thylacosmilines, in which dP3 is much longer than p3, and the upper and lower premolar rows are relatively short (though in Patagosmilus and Thylacosmilus, there is a large diastema between the lower canine and premolars due to a more posterior position of the postcanine teeth). The M4 of Eomakhaira resembles that of Patagosmilus more than that of any other Borhyaenoid examined in being gracile, anteroposteriorly narrow and labiolingually very wide compared with M1–3 and in having a vestigial protocone. Compared with M4 of Eomakhaira and Patagosmilus, the M4 of Thylacosmilus is anteroposteriorly longer than labiolingually wide (resulting in a robust tooth), due partly to the loss of its talon. Some features common to both Eomakhaira and later Thylacosmilines may also occur in Proborhyaena gigantea, especially if MLP 79-XII-18-1 pertains to that species, and potentially represent synapomorphies at a deeper node within Proborhyaenidae; these include a shallow dentary, more posteriorly positioned infraorbital foramen, labiolingually narrow upper canines, and a P3 that is much larger than p3.

Among Eutherians, saber teeth have originated independently three or four times: in Oxyaenid 'Creodonts' (Machaeroidinae), in Nimravid and Barbourofelid Carnivoramorphans (if these are not sister taxa that were ancestrally machairodont, and in Felid Carnivorans (Machairodontinae). Among Metatherians, they are known to have evolved only once, in Sparassodonts (suggestions that the extant Didelphid Monodelphis dimidiata exhibits a saber-toothed morphology have not been supported by later analyses). This disparity between Eutherians and Metatherians is curious given the broad range of predatory Metatherians, which include Deltatheroidans, Stagodontids, the Marsupialiform Anatoliadelphys, other groups of Sparassodonts (i.e. Hathliacynids), Didelphids (i.e. Sparassocynins, Didelphins, and related forms), Dasyurids, Thylacinids, and Thylacoleonids. One potential reason for the rarity of Metatherian sabertooth lineages may be related to how their distinctive upper canines may have functioned. It has been argued that saber teeth would have required a considerable learning period to be used effectively. Functional modeling of saber bites has shown that an imprecise bite can cause the canines to snag, can easily be be too shallow to be lethal, or can penetrate too deeply to be extracted from the prey. Elongate, labiolingually narrow upper canines are also vulnerable to breakage when subjected to sudden, unpredictable loads such as those produced by struggling prey. Many Eutherian sabertooths exhibited prolonged retention of the deciduous canines, which are also large and machairodont in these lineages. It has been hypothesised that this extended retention time resulted in a longer 'training' period, in which breakage would not have had permanent consequences (due to their eventual replacement).

Unlike Eutherians, Metatherians (including Sparassodonts) lack deciduous canines and therefore have only one tooth generation at the canine locus, precluding 'training' canines. At the same time, Thylacosmiline Sparassodonts are the only sabertooths with hypselodont canines, and this may be a different means to achieve the same end: a tooth that would eventually be replaced in the event of breakage (albeit by growth at the base rather than by wholesale replacement). The results of Engleman et al.'s phylogenetic analysis suggest that open-rooted canines were not a novel innovation of Thylacosmilines but a plesiomorphic feature inherited from non-saber-toothed ancestors (i.e. an apomorphy for Proborhyaenids). This suggests that the absence of deciduous canines may have constrained the evolution of saber teeth in Metatherians and that hypselodonty is a prerequisite for evolving saber teeth in this clade. This may be one reason why saber-toothed canines only evolved once in metatherians rather than repeatedly, as in Eutherians. Deciduous canines are widespread within Eutheria, whereas hypselodont canines in Metatherians appear to have a much narrower distribution. Therefore, developing saber teeth in Metatherians would require not only selection for machairodonty but also the prerequisite of an uncommon morphology.

Open-rooted, ever-growing (hypselodont) canines in Proborhyaenids (including Thylacosmilines) are frequently considered a unique feature within Metatheria. However, they may be more widely distributed than generally realised. Hypselodont canines have been reported in Peramelemorphians, and computerised tomography scans of adult Peramelemorphians show open-rooted canines with non-tapering roots and open pulp cavities. A computerised tomography scan of the nonSparassodont Pucadelphyidan Pucadelphys andinus, interpreted as a sexually mature male, appears to show open-rooted upper canines. The canines of didelphids and dasyuromorphians are reported to be hypselodont by several authors, but Engleman et al. were not able to confirm these observations. Among the aforementioned studies, that of Amelia Chemisquy and Francisco Prevosti is the only one that provides imagery to support their claim of hypselodonty in Monodelphis dimidiata. However, their radiographs are shown only in palatal view, making it difficult to determine whether the canine pulp cavities are actually open (or merely show the apical foramen). In one of the radiographs said to demonstrate an open root, the canine root is closed. Other computerised tomography scans and X-rays of adult Didelphids and Dasyuromorphians show nonhypselodont canines. None of the canines of Didelphids and Dasyuromorphians we have observed exhibit the features associated with hypselodonty that have been described previously in Proborhyaenids, such as absence of enamel, remodeling of the apex with well-developed compensatory wear facets, and great height of the tooth despite heavy wear.

Lower canines of non-Thylacosmiline Proborhyaenids. (A)–(B) Proborhyaena gigantea (MACN-A 52-382) in (A) labial and (B) lingual views. (C) Arminiheringia auceta (MACN-A 10970) in labial view. (D) Paraborhyaena boliviana (UATF-V-000129; left canine, reversed) in labial view. The upper canines of Arminiheringia show a comparable morphology (apices of the upper canines are unknown for Proborhyaena and Paraborhyaena). Scale bars are 10 mm. Engleman et al. (2020).

The development of hypselodont, open-rooted canines in Proborhyaenids may be the result of paedomorphosis. The canine roots of extant Marsupials remain open much longer during ontogeny than those of most Placentals, and the canine roots of juvenile non-Proborhyaenid Sparassodonts remain open relatively late in ontogeny. Combined with the observations above, the presence of closed roots in senescent Proborhyaenids, such as SGOPV 3490 and MLP 79-XII-18-1, suggests that canine hypselodonty was achieved by delaying root closure until extremely late in ontogeny. Similar evolutionary transitions from closed-rooted to fully hypselodont teeth via postponement of root formation have been observed in other Mammals, including Notoungulates. It has also been suggested that paedomorphosis played a role in other aspects of Thylacosmiline evolution, such as the retention of dP3 as a functional element in the adult dentition.

Other features in Thylacosmilines typically related to machairodonty, such as those connected with a wide gape, appear to have originated prior to the group’s origin (i.e., among non-Thylacosmiline Proborhyaenids). In Callistoe vincei, the combined heights of the upper and lower canines are comparable to the height of only the upper canine of Thylacosmilus. Thus, Callistoe and Thylacosmilus would have required comparable gapes to achieve clearance between the canines.

The transverse processes of the atlas of Callistoe and Thylacosmilus are longer (anteroposteriorly) than wide (transversely), and extend far posterior to the caudal facets. This contrasts with the condition in Prothylacynus, Borhyaena, and Arctodictis, in which these processes are subequal in length and width and extend only slightly posterior to the posterior border of the caudal facets. Similar distinctions in the morphology of the axial transverse processes are seen in comparisons between saber-toothed and non-saber-toothed Felids, respectively. Elongate transverse processes in saber-toothed Felids increased the area of origin for the obliquus capitis cranialis and obliquus capitis caudalis, which are thought to be the primary muscles involved in the saber bite in saber-toothed Mammals. The transverse processes of the atlas in Thylacosmilus resemble those of saber-toothed Felids. The transverse processes of Callistoe extend further posteriorly than those of most Sparassodonts but less posteriorly than in Thylacosmilus, suggestive of large obliquus capitis cranialis and obliquus capitis caudalis muscles in this non-saber-toothed taxon. This suggests that specialisation of the neck musclature occurred in non-Thylacosmiline Proborhyaenids prior to the evolution of saber teeth in this clade.

The Oligocene Paraborhyaena, which is recovered as more closely related to Thylacosmilus than to Callistoe in Engleman et al.'s phylogenetic analyses, shows additional similarities to Thylacosmilus in the posterior part of the skull not present in the geologically older Callistoe. (The posterior cranium is unknown in other Proborhyaenids, such as Proborhyaena and Patagosmilus). As in Thylacosmilus, the braincase of Paraborhyaena is short anteroposteriorly, and the nuchal crest is nearly vertical, exposing the occipital condyles in dorsal view. A shortened temporalis fossa (covarying with an anteroposteriorly short braincase) and vertical occiput, features common among saber-toothed Mammals, have been considered to reflect either a wide gape or mechanical compensation of a reduced temporalis muscle lever arm created by a small coronoid process. In Proborhyaena, Eomakhaira, and Anachlysictis (in which the temporal and occipital regions of the skull are unknown, but these taxa are phylogenetically bracketed by Paraborhyaena and Thylacosmilus), the coronoid process is large and dorsoventrally tall, contradicting the latter hypotheses. This suggests that Proborhyaenids differed from Placental sabertooths in acquiring a short braincase and vertical occiput early in their history (i.e. prior to appearance of saber teeth), whereas these features generally appeared after the acquisition of saber teeth in Placentals.

Eomakhaira is considered a saber-toothed Sparassodont here because it exhibits a degree of machairodont specialisation comparable to that of early-diverging members of other saber-toothed clades, e.g. Machaeroides, Dinictis, Nimravus, and Pseudaelurus. Accordingly, saber-toothed Sparassodonts (Thylacosmilinae) now have a documented biochron extending from the early Oligocene (32–33 million years ago) to the early Pliocene (3 million years ago), a duration of almost 30 million years, far longer than that of any placental sabertooth clade. Saber-toothed Oxyaenid 'Creodonts' (Machaeroidinae) span approximately 11 million years (52.8–41.5 million years ago), a minimum estimate given the clade’s poor fossil record (no more than six species and fewer than 10 specimens). Securely dated and identified Nimravids are known from the late Eocene (about 37.8 million years ago) to the end of the Oligocene (23 million years ago), a temporal range of 14.8 million years. Isolated upper canine fragments from Asia may push the first appearance datum of this clade back into the late middle Eocene (about 42 million years ago), extending its temporal range to approximately 19 million years. However, given that these specimens consist of isolated fragments of upper canine saber teeth, they may pertain to another saber-toothed clade such as machaeroidine 'Creodonts'. Barbourofelid Carnivoramorphans first appear in the early Miocene (20–19 million years ago) and are last recorded in the late Miocene (6 million years ago), a range of 13–14 million years. Finally, machairodontine Felid Carnivorans are recorded as early as the middle Miocene (16 million years ago) based on the first appearance of Pseudaelurus sensu stricto and last appear during the end-Pleistocene extinctions (about 0.01 million years ago), a temporal range of roughly 16 million years. Excluding Eomakhaira, the biochron of Thylacosmilines spans at least 16 million years, based on a Patagosmilus-like upper molar from the early Miocene of Patagonia (Colhuehuapian South American Land Mammal Age, 20.2–20.0 million years ago). Thus, even excluding Eomakhaira, the temporal range of Thylacosmilines exceeds that of Nimravids and Barbourofelids and is comparable to that of Machairodontines.

Temporal durations of major lineages of Mammalian saber-toothed carnivores, with metatherian lineage in grey and placental lineages in black. A representative skull of each clade is depicted to the right; from top to bottom: Thylacosmilus atrox (Thylacosmilinae), Machaeroides eothen (Machaeroidinae), Hoplophoneus primaevus (Nimravidae), Barbourofelis fricki (Barbourofelidae) and Smilodon fatalis (Machairodontinae).Tick mark on Thylacosmilinae record represents the oldest occurrence of this clade (about 20.2 million years old) prior to the discovery of Eomakhaira. Abbreviations: Plio., Pliocene; Ple., Pleistocene. Engleman et al. (2020).

Sabertooth clades are generally characterised by high rates of extinction and turnover relative to other carnivorous Mammal. This has been suggested to be related to their inferred specialised hunting behavior and comparatively narrow prey base (based on functional morphology and palaeoecological data) and their status as hypercarnivorous apex predators, which would make them more vulnerable to ecological distruptions and environmental perturbations. In this respect, the long stratigraphic range of thylacosmilines compared to placental saber-toothed clades is noteworthy considering the many major faunal/climatic changes faced by the former in South America, including the Bisagra Patagónica during the Oligocene, the faunal turnover at the Oligocene-Miocene boundary (which, notably, marked the end of all non-Thylacosmiline Proborhyaenids), the Middle Miocene Climatic Optimum and subsequent climatic deterioration, and the expansion of grasslands during the late Miocene. The shorter stratigraphic ranges of Placental sabertooths relative to Thylacosmilines do not appear to be attributable to competitive interactions between Placental sabertooth clades in North America, Eurasia, and Africa compared to the relative isolation of Thylacosmilines in South America, as placental sabertooth clades mostly do not overlap in space and time (e.g. North America’s 'Cat gap'). The large disparity between the temporal ranges of thylacosmiline metatherians and placental sabertooth clades suggests dissimilar ecological requirements, with thylacosmilines perhaps having broader dietary and/or habitat preferences. Thylacosmiline Sparassodonts, a distinctive, diverse, and temporally long-lived lineage of machairodont Mammals, hardly represent 'inferior' or 'ineffective' imitations of their Placental analogs as frequently claimed. The discovery of Eomakhaira clarifies the evolution of machairodonty both within Sparassodonta and in Mammals generally and speaks to the enduring utility of the South American fossil record in elucidating the splendid variety of morphological diversity among Mammals and nature’s fantastic capacity for convergence.

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