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Abstract The fossil collections made by early explorers in South America have been fundamental to reveal the past diversity of extinct mammals and unravel their evolutionary history. One important early explorer in South America was the Swiss-Argentine palaeontologist Kaspar Jacob Roth, known as Santiago Roth (1850, Herisau, Switzerland-1924, Buenos Aires, Argentina), who made significant collections of fossil mammals that are housed in museums in Europe and Argentina. The important collections of Roth in Switzerland include iconic Pleistocene megafauna from the Pampean Region (Argentina). The palaeontological significance of the Pampean Region relies on its abundant record of fossil vertebrates that documents diversity dynamics and paleoenvironmental change in southern South America, serving as the basis for the South American biostratigraphical scale of the late Neogene and Quaternary. The South American native ungulates (SANUs) were hoofed placental mammals that radiated in South America. The clades Notoungulata and Litopterna include, among others, the last representatives of SANUs megafauna in the continent. We revise and describe for the first time the SANUs specimens from the Pampean Region of the Roth collections in Switzerland. The collections include two species of notoungulates (Toxodon cf. T. platensis and Mesotherium cristatum) and one litoptern species (Macrauchenia patachonica). The occurrences are restricted to the early and middle Pleistocene (pre-Lujanian Stages/Ages). Although the SANUs diversity in the Roth collections is low in comparison with other groups (e.g., xenarthrans), some of the specimens are very complete, including skulls and postcranial remains. The completeness of the Ma. patachonica material allows an update and reinterpretation of some of the details of the dentition and the postcranial skeleton of this iconic species. In addition to its historical importance, the SANU specimens from the Roth collections provide important information to study the paleobiology and evolution of South American megafauna and evaluate hypotheses about their extinction in the continent.

Roth’s explorations, the resulting collections many now allocated in La Plata, Zurich, Geneva and Copenhagen, and his significant contributions in geological—especially stratigraphic—and paleontological topics, are a paradigmatic case for the global history of paleontology and for the Swiss migration history in Latin America. His work included the discovery of a diverse megafauna from the Pampean region, of sites and strata in Patagonia of paleontological significance, and the recognition of a group of endemic ungulate mammals, Notoungulata. Roth’s discovery of a human skeleton associated with a glyptodont carapace is one of the first reports of the coexistence of humans with the extinct fauna of the South American Quaternary. Roth became a renowned scholar at the Museo de La Plata, which was a leading scientific institution in the nation-making of Argentina, particularly in the expansion of the Patagonian frontier. He also kept strong ties with his native Switzerland, where late in his adult life he obtained some formal training and tried to attract other Swiss nationals to work in natural sciences in Argentina. His biography sheds light about the circumstances of his scientific collection and career in the interstices between amateur and professional science, modernity and imperialism at the turn of the twentieth century. [ABSTRACT FROM AUTHOR]

The article titled "Systematic Revision of Pre- and Post-Santacrucian Species of Protypotherium (Interatheriidae, Notoungulata)" provides a taxonomic revision of the species of Protypotherium, a genus of native ungulates from South America. The study focuses on the Interatheriidae family, which is considered one of the most diverse and abundant clades of South American native ungulates. The authors propose junior synonyms for certain species and conclude that Protypotherium includes several species. The article also presents a comprehensive phylogenetic analysis of Interatheriidae. [Extracted from the article]

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Body mass (BM) is a fundamental variable for many paleobiological investigations that is challenging to accurately infer for species that lack living representatives and/or close morphological analogs. This study explores this issue using notoungulates, a diverse group of extinct South American herbivorous mammals with an extensive fossil record. We use a new dataset of 1,900+ extant mammal species (from ~80,000 specimens) to estimate notoungulate BM based on head-body length and a published dataset of 400+ species (~2,100 specimens) to estimate BM based on occipital condyle width. Condylobasal length, stylopod diameter and circumference, and neck length data are used to explore factors that can confound BM predictions. We estimate the following BM ranges for 10 osteologically well-characterized species and calculate similar ranges for 30 others known from less complete remains: Toxodontia: Thomashuxleya externa (80–120 kg), Homalodotherium cunninghami (250–350 kg), Scarrittia canquelensis (450–550 kg), Adinotherium ovinum (75–90 kg), Nesodon imbricatus (350–400 kg), and Toxodon platensis 1,000–1,200 kg); Typotheria: Interatherium robustum (1.9–2.0 kg), Miocochilius anomopodus (9–14 kg), Protypotherium australe (3.5–4.0 kg), and Pachyrukhos moyani (1.2–1.6 kg). We suggest that species such as these can be used as "calibration points" when inferring BM of species known from more limited remains. Discrepancies between our estimates and previously-published studies are primarily due to the distinctive craniodental morphology of notoungulates and the robust limb bones of toxodontians. There is significant, non-random error correlated with body habitus (i.e., being relatively robust or gracile) in many variables traditionally used to estimate BM, including femur circumference, and new methods are needed to compensate for this. [ABSTRACT FROM AUTHOR]

We describe three new Toxodontidae (Notoungulata) remains from Mendoza Province, west-central Argentina. One is a new mandible (IANIGLA-PV 993) collected at Huayquerías del Este, from the Huayquerías Formation (Late Miocene-earliest Pliocene). This finding adds to two other unpublished specimens from the area stored in old collections. MCNAM-PV 113 is a skull fragment from Huayquerías de San Carlos, lacking exact origin data, and MACN PV 14854 corresponds to associated upper and lower teeth from the so-called Bajada Grande locality, in Huayquerías del Este, from levels now assigned to the Bajada Grande Formation (Late Pliocene-Pleistocene). IANIGLA-PV 993 and MCNAM-PV 113 belong to Xotodon major. This species was described in 1914 based on cranial remains likely from levels now assigned to the Tunuyán Formation. Slight differences between IANIGLA-PV 993 and the holotype of X. major are attributable to intraspecific variation and their ontogenetic stage: smoother anteroventral mandibular angle; ascending ramus with blunt crest between the fossae for the zygomaticomandibularis and masseteric muscles; and tooth features (e.g., p4 with deeper grooves and slightly shorter teeth) that relate to a lesser wear degree. MCNAM-PV 113 shows nasals and upper teeth comparable with the holotype, although teeth are slightly less compressed. The temporal gap between levels yielding the holotype and IANIGLA-PV 993 is not reflected by our taxonomical interpretation. Finally, the specimen MACN PV 14854 is identified as Toxodon chapalmalensis, based upon its similarities with specimens from the Chapadmalal area, Buenos Aires Province, showing a correspondence with the Chapadmalalan Stage/Age of the Pampean Region. [ABSTRACT FROM AUTHOR]

Here we describe a new notoungulate taxon, Pascualhyrax irqi sp. nov., from Eocene levels of north-western Argentina. The materials studied include cranial and dental remains and were exhumed in sediments of the Quebrada de los Colorados Formation, corresponding to the late middle Eocene (Bartonian). General traits of the referred specimens (e.g. hypsodont postcanines, nasal bulge) point to a close relationship with different typotherian taxa traditionally included among the paraphyletic 'Archaeohyracidae'. In turn, a combination of characters indicates that the new material represents a new taxon (e.g. smaller size, deep ectoflexus in upper molars in all wear stages, maxillary process extending posterior to the caudal end of the nasals). A phylogenetic analysis was performed, based in a previously published matrix and the addition of new characters, resulting in a close relationship of Pascualhyrax irqi sp. nov. with different 'Archaeohyracidae' of Oligocene radiation. This fact shows the early expression of some characters until now only known for later forms, and developed in the context of a recently acknowledged diversity of hypsodont typotherians in north-western Argentina. Regarding other evolutionary issues, the strong similitude of the juvenile specimens referred to Pascualhyrax irqi sp. nov. with adult individuals of the Neogene clade Pachyrukhinae, indicates possible heterochronic (paedomorphic) traits in the later forms. [ABSTRACT FROM AUTHOR]

Cranial endocasts are one of the most direct tools available to obtain information about the endocranial cavity of fossil mammals, but few anatomical comparisons have analyzed endocranial data within a phylogenetic framework. Our study combines analyses of new digital endocasts from high-resolution X-ray micro-computed tomography (µCT) for two notoungulates, Cochilius sp. and Notostylops murinus, with data from published anatomical descriptions and endocasts (prior µCT digital endocasts and natural and extracted plaster endocasts), resulting in 22 endocranial characters (11 new) scored on 20 notoungulates and five outgroups. These data permitted a comparative study of cranial endocasts of notoungulates, an enigmatic group of extinct South American mammals, shedding light on several important issues concerning their phylogenetic relationships. The new endocranial data were integrated into a craniodental and postcranial matrix, resulting in 259 characters across 65 taxa, the most comprehensive character and taxon sampling to date and the only one including a substantial suite of endocranial characters. We further performed ancestral state reconstructions for endocranial characters using parsimony optimization on the resulting phylogenetic trees. Body mass estimates and endocranial volumes were used to calculate Encephalization Quotients (EQs) and Phylogenetic Encephalization Quotients (PEQs), which then were optimized onto the phylogenies. Endocast data from ancestral character state reconstructions bear on relationships among meridiungulates, revealing that many endocast characters may be synapomorphies for Notoungulata and other clades within the group. Notoungulates, in general, display relatively low EQs when compared with Holarctic herbivorous mammals, potentially due to their long history of evolution in isolation in South America during the Cenozoic. [ABSTRACT FROM AUTHOR]

On the basis of selected osteological markers, Patterson (1936) identified two pathways (here designated A and B) along which the internal carotid artery in notoungulates was said to travel in order to enter the skull and supply the brain. The existence of these pathways, at least as Patterson defined them, has been disputed but no substantive alternatives have been proposed. Using comparative embryological and morphological evidence based on conditions in extant mammals, we find that pathway A (intratympanic) is not supported. Pathway B (enclosed extratympanic) is supportable, but as far as is now known applies only to a small number of notoungulate taxa. On the basis of new evidence, we propose another route, pathway C, briefly mentioned by Scott (1912) but subsequently ignored, that may apply to the majority of notoungulates. In pathway C (unenclosed extratympanic) the internal carotid passed directly into the endocranium via a naturally unossified area of the basicranium, the piriform fenestra, rather than coursing through or alongside the middle ear in a canal. The absence of a separate, bony carotid foramen on the basicranium's ventral surface may explain the hesitancy of previous workers to consider this routing. There is no evidence that the function of the internal carotid artery was supplanted by another vessel (e.g., external carotid artery) in any notoungulate. Conditions in other major clades of South American native ungulates are poorly investigated, but some clearly differed from notoungulates in carotid patterning, pointing to the existence of substantial intertaxon disparities. [ABSTRACT FROM AUTHOR]

Notoungulates represent the most taxonomically diverse and temporally and geographically widespread group among South American native ungulates. Here, we analyze anatomical and systematic aspects of proximal tarsal bones recovered from the Lower and Upper Lumbrera formations (middle and late middle Eocene) in northwestern Argentina. We provide detailed descriptions, comparisons, and infer foot stances and range of movements for the taxa implicated. Material studied includes astragali belonging to the oldfieldthomasiid Colbertia lumbrerense (Lower Lumbrera Formation), a set of proximal tarsals referred as Typotheria indet. (Lower Lumbrera Formation), and tarsals (also including navicular and cuboid) of the informal taxon “Campanorco inauguralis” (Upper Lumbrera Formation). The comparison of the tarsals of Colbertia lumbrerense (middle Eocene of Argentina) with Colbertia magellanica (early Eocene of Brazil) reveals several differences including variations on the development and arrangement of articular facets, and the size of the dorsal astragalar foramen in the Argentinean species. The specimen of Typotheria indet. shows morphological affinities with basal interatheriid taxa. However, its larger size contrasting with the overall small body sizes of Eocene interatheriids precludes an indisputable taxonomic assignment. Concerning “Campanorco inauguralis”, our observations indicate that there is no morphological evidence for a close phylogenetic relationship with Mesotheriidae. It presents a “reversed alternating tarsus” condition, which is also observed in Leontiniidae, “Notohippidae”, Toxodontidae, and some typotherians. However, the spectrum of singularities exhibited by this form precludes the assessment of its relationships in the context of the Paleogene radiation of Typotheria and it is necessary to extend the comparison to Eocene notoungulates. Finally, in a morphofunctional context a plantigrade foot posture is inferred for the specimens here reported. These observations have the potential to provide functional proxies for paleoecological reconstructions to be applied to the study of the early radiation of these notoungulate faunas.

Protypotherium (Mammalia, Notoungulata, lnteratheriinae) is a well-known and very diverse genus of extinct native ungulates of South America, widely distributed from southern to middle latitudes of Argentina, Chile, and Bolivia. This genus exhibits distinctive species throughout the Miocene to the beginning of the Pliocene that display an interesting size pattern. The large sample of specimens studied during several years of research allows us to analyze the shape and size of upper and lower molars for nine species of Protypotherium, in order to test the hypothesis of reduction of size ranges preserving a general tooth morphology as a response to climate deterioration, documented in South America by a marked reduction of temperature that occurred during Miocene times. Elliptic Fourier analyses were used to capture the shape of the occlusal morphology and the centroid size was also retained for subsequent analyses. Our results demonstrate that: 1) a similar morphological tooth pattern is observed among all species; 2) tooth shape variation is not associated with a change in size; and 3) there is a tendency to increase the size and decrease the number of species. This striking pattern could be correlated with a global trend to lower temperatures, which indicates a deterioration of paleoenvironmental conditions. A conservative tooth pattern, together with an increase of size and a reduction in number of species were the main evolutionary and ecological tendencies accounted in Protypotherium from the early to late Miocene-beginning of the Pliocene, which were accompanied by a latitudinal displacement from Patagonia to northern areas at the end of the Miocene. [ABSTRACT FROM AUTHOR]

Here we describe a new notoungulate mammal from the early Eocene (Itaboraian SALMA) of Chubut, Argentinian Patagonia, from the localities of Las Flores and Las Violetas Farm, represented by fragments of maxilla and isolated teeth. The specimens were found in the Las Flores Formation, Río Chico Group, and assigned to the Henricosborniidae, a primitive family within the order Notoungulata. Orome deepi gen. et. sp. nov. differs from other henricosborniids in a less developed metacone column in the upper molars, a larger metaloph in the third upper molar and a larger mean size. The morphometric analysis does not show any difference between the individuals of the two studied localities. The new addition to the henricosborniid diversity in Patagonia allows to provide a better understanding of the early radiation of notoungulates in South America.

Abstract Geological studies in the northern sector of the Chaco foreland Basin, Bolivia, yielded new fossils coming from late Oligocene-late Miocene of the Petaca Formation. Few fossil mammals were known from the Subandean Region of Bolivia. We report a partially complete mandible of a hegetotheriid Hegetotheriinae (Notoungulata, Typotheria) from Abapó (Río Grande River). The specimen (YPFB-LIT-PAL-005) is very close in size and dental morphology to the late Oligocene (Deseadan South American Land Mammal Age, SALMA) — Santacrucian (early Miocene) Prohegetotherium schiaffinoi (Kraglievich 1932) from Fray Bentos (Uruguay and Argentina), Salla (Bolivia), Divisadero Largo-Potrerillos and Quebrada Fiera (Argentina). However, mandibular characteristics, as the shape with a marked change in height along its length, increasing towards the back, a prominent masseteric crest, a deep mandibular groove, and a remarkable thickening of the ventral rim of the mandible, indicate differences between this specimen and Prohegetotherium schiaffinoi and the other Hegetotheriinae. The affinity of YPFB-LIT-PAL-005 with P. schiaffinoi suggests a late Oligocene to early Miocene for the upper bearing horizon of the Petaca Formation (in Abapó), an older age than previously assigned to the top of this unit (late Miocene), and confirms the taxon distribution between ∼ 36 ° to ∼ 17 ° south latitude.

The caudal cranium and occipito-cervical region, although usually overlooked, are informative about the paleobiology of fossil mammals, allowing inquiry into vision and hearing abilities, as well as head and neck postures. Particularly for Pachyrukhinae, some related features remain unexplored. In this contribution, 22 specimens of Paedotherium and Tremacyllus were analyzed in a mainly qualitative and comparative framework. Pachyrukhines are characterized by having large orbits and hearing cavities, moderate to short necks with generalized morphologies, and S-shape postures. These features allow rejecting some types of specialized digging habits, and support the preference of open or low-humidity environments. Paedotherium typicum is distinguished by the presence of laterally oriented eyes, marked vaulted cranium and predominant short extensor and stabilizing neck muscles, and cervico-occipital-hyoid configurations suitable for ventro-flexed resting posture. These features indicate accentuated frontation and panoramic-vision, upward head postures enhancing substrate perception, and the resistance of impacts during leaping-cursorial locomotion. Conversely, P. bonaerense, and to lesser degree Tremacyllus spp., show less frontation and probably adopted more horizontal head postures. More particularly, stronger ventral and lateral neck and head flexors and extrinsic arm musculature are reconstructed for P. bonaerense, compatible with generalist or scratch-digging habits. Its smaller auditory cavities and stronger ear musculature, compared to the contemporary P. typicum and especially Tremacyllus spp., would indicate larger ears and microhabitat segregation. The integrative analysis proposed here and in the accompanying contribution aims to shed light on convergences with extant models, paleobiology, niche partitioning, and external appearance of the latest rodent-like ungulates. [ABSTRACT FROM AUTHOR]

Inquiring into the paleoecology of extinct forms is always a challenge, particularly when the taxa under study correspond to derived ecomorphs of ancient and completely extinct clades. In this contribution, the configuration of the masticatory apparatus and associated features of the Neogene pachyrukhines Paedotherium and Tremacyllus are studied in a detailed, mainly qualitative, comparative analysis of 36 specimens. Tooth morphology and the reconstructed muscular configuration of pachyrukhines indicate an important mediolateral component during chewing, and predominant crushing over grinding, as well as anteroposterior movements for the coupling and action of stronger gnawing incisors. These actions are more compatible with hard and brittle or turgid fruit food consumption than specialized folivorous, and particularly grazing, habits. The infraorbital and palatal foramina morphology and other rostral features indicate increased touch sensibility for object recognition and are congruent with the presence of infoldings of the lips protecting the gingiva during gnawing on hard foods. Additionally, there was a morphological gradient between Tremacyllus and P. bonaerense, from high selection of relatively soft and small food items, to specialized hard item consumption and higher resistance for abrasion and masticatory efforts (e.g., in eventual association with digging habits), respectively. Paedotherium typicum presents intermediate characteristics, with incisors designed for better cropping action or poorer selectivity during feeding. This more profound understanding of the feeding habits of pachyrukhines further allows the suggestion of paleoecological factors that could have contributed to niche segregation between these long-term coexisting rodent-like taxa. [ABSTRACT FROM AUTHOR]

Geological studies in the northern sector of the Chaco foreland Basin, Bolivia, yielded new fossils coming from late Oligocene-late Miocene of the Petaca Formation. Few fossil mammals were known from the Subandean Region of Bolivia. We report a partially complete mandible of a hegetotheriid Hegetotheriinae (Notoungulata, Typotheria) from Abapó (Río Grande River). The specimen (YPFB-LIT-PAL-005) is very close in size and dental morphology to the late Oligocene (Deseadan South American Land Mammal Age, SALMA) -- Santacrucian (early Miocene) Prohegetotherium schiaffinoi (Kraglievich 1932) from Fray Bentos (Uruguay and Argentina), Salla (Bolivia), Divisadero Largo-Potrerillos and Quebrada Fiera (Argentina). However, mandibular characteristics, as the shape with a marked change in height along its length, increasing towards the back, a prominent masseteric crest, a deep mandibular groove, and a remarkable thickening of the ventral rim of the mandible, indicate differences between this specimen and Prohegetotherium schiaffinoi and the other Hegetotheriinae. The affinity of YPFB-LIT-PAL-005 with P. schiaffinoi suggests a late Oligocene to early Miocene for the upper bearing horizon of the Petaca Formation (in Abapó), an older age than previously assigned to the top of this unit (late Miocene), and confirms the taxon distribution between ~ 36 ° to ~ 17 ° south latitude. [ABSTRACT FROM AUTHOR]

Notoungulates were the most diverse group of South American native ungulates during the Cenozoic. They ranged from small rodent-like to heavy rhino-like forms and, despite this morphological diversity, their monophyly is widely accepted. Within Notoungulata, Typotheria and Toxodontia (the two major clades) are well supported, whereas phylogenetic relationships between early diverging lineages (formerly grouped as 'Notioprogonia') are still a matter of debate. Within Toxodontia, 'Notohippidae' sensu Simpson (1967) is a paraphyletic group lacking systematic consensus and represented by small to medium-sized taxa from the late Eocene to early/middle Miocene, but that were especially diverse during the late Oligocene. In this context, we describe a new toxodont (Teushentherium camaronensis gen. et sp. nov.), found at Barrancas Blancas, an Oligocene locality in the south-east of Chubut (Patagonia, Argentina). It is represented by the anterior portion of the skull (palate and anterior snout) and the associated horizontal rami of the mandible. A phylogenetic analysis places the new taxon as closely related to taxa traditionally regarded as 'Notohippidae'. In addition, extensive taxonomic sampling allowed us to assess phylogenetic relationships among other representatives. As expected, only Homalodotheriidae, Leontiniidae and Toxodontidae were recovered as monophyletic groups, whereas neither 'Notohippidae' nor 'Isotemnidae' were supported. We also identified a clade (sister to Homalodotheriidae) for which we propose the name Eutoxodontia. It includes 'Notohippidae' sensu Simpson, Leontiniidae and Toxodontidae, and provides a more accurate definition for middle and later diverging toxodonts, avoiding terminology such as 'advanced' or 'derived'. We hope to contribute to the stabilization of toxodontian taxonomy, to highlight the main gaps in this respect, and to promote future contributions dealing with this intricate issue. [ABSTRACT FROM AUTHOR]

We revise the mammal fossils from the Fray Bentos Formation previously assigned to Notoungulata, as well as some new remains recovered from levels cropping out in Corrientes Province (Argentina). In some cases, previous determinations have been completed to the species level, but others have been substantially modified, identifying different families. At present, identified taxa are: "Prohegetotherium" schiaffinoi (Hegetotheriidae), an undetermined interatheriid that is hypothesized as representing the upper teeth of Eopachyrucos ranchoverdensis, and Argyrohyrax proavus (Interatheriidae), cf. Archaeohyrax suniensis (Archaeohyracidae), cf. Mendozahippus fierensis ("Notohippidae"), a Leontiniidae indet., and some undetermined Notoungulata. The presence of Leontiniidae is based on the specimen CTES-PZ 7871, but those previously assigned to this family are herein discarded as such and reinterpreted (notohippid and undetermined taxon). The Mesotheriidae are discarded among the revised materials, but they are present in the Fray Bentos Formation in Corrientes Province with Trachytherus spegazzinianus (= Ameghinotherium curuzucuatiense). The revised sample presents affinities with the fauna from the type locality of the Fray Bentos Formation, in Uruguay, and the Deseadan assemblages from Quebrada Fiera (Mendoza Province, Argentina), Patagonia (Argentina), and Salla (Bolivia). These similarities are important from a paleobiogeographic point of view, especially by the recognition of cf. M. fierensis, a species only known from Mendoza up to now. Only the finding of new material from Fray Bentos Formation in Corrientes Province will allow us to corroborate these tentative determinations and a possible eastward extension of up-to-now exclusive taxa from Quebrada Fiera. [ABSTRACT FROM AUTHOR]

Notoungulates are an extinct clade of South American mammals, comprising a large diversity of body sizes and skeletal morphologies, and including taxa with highly specialized dentitions. The evolutionary history of notoungulates is characterized by numerous dental convergences, such as continuous growth of both molars and incisors, which repeatedly occurred in late-diverging families to counter the effects of abrasion. The main goal of this study is to determine if the acquisition of high-crowned incisors in different notoungulate families was accompanied by significant and repeated changes in their enamel microstructure. More generally, it aims at identifying evolutionary patterns of incisor enamel microstructure in notoungulates. Fifty-eight samples of incisors encompassing 21 genera of notoungulates were sectioned to study the enamel microstructure using a scanning electron microscope. We showed that most Eocene taxa were characterized by an incisor schmelzmuster involving only radial enamel. Interestingly, derived schmelzmusters involving the presence of Hunter-Schreger bands (HSB) and of modified radial enamel occurred in all four late-diverging families, mostly in parallel with morphological specializations, such as crown height increase. Despite a high degree of homoplasy, some characters detected at different levels of enamel complexity (e.g., labial versus lingual sides, upper versus lower incisors) might also be useful for phylogenetic reconstructions. Comparisons with perissodactyls showed that notoungulates paralleled equids in some aspects related to abrasion resistance, in having evolved transverse to oblique HSB combined with modified radial enamel and high-crowned incisors. [ABSTRACT FROM AUTHOR]

A remarkable diversity of plant-eating mammals known as South American native ungulates (SANUs) flourished in South America for most of the Cenozoic. Although some of these species likely filled ecological niches similar to those of modern hoofed mammals, others differed substantially from extant artiodactyls and perissodactyls in their skull and limb anatomy and probably also in their ecology. Notoungulates and litopterns were the longest-lived and most diverse SANU clades and survived into the Quaternary; astrapotheres went extinct in the late Miocene, whereas other SANU groups were restricted to the Paleogene. Neogene notoungulates were quite specialized in craniodental structure, but many were rather unspecialized postcranially; in contrast, litopterns evolved limb specializations early in their history while maintaining more conservative dentitions. In this article, we review the current understanding of SANU evolutionary relationships and paleoecology, provide an updated compilation of genus temporal ranges, and discuss possible directions for future research. ▪ South American native ungulates (SANUs) were a diverse, long-lived, and independent radiation of mammals into varied terrestrial plant-eater niches. ▪ We review origins, evolution, and paleoecology of the major SANU clades: Notoungulata, Litopterna, Astrapotheria, Xenungulata, and Pyrotheria. ▪ At their peak, during the Eocene and Oligocene, more than 40 genera of native ungulates inhabited South America at any one time. ▪ SANUs ranged from <1 kg to several tons and evolved many combinations of diet and locomotor adaptations not seen in living ungulates. [ABSTRACT FROM AUTHOR]

Notoungulates represent the most taxonomically diverse and temporally and geographically widespread group among South American native ungulates. Here, we analyze anatomical and systematic aspects of proximal tarsal bones recovered from the Lower and Upper Lumbrera formations (middle and late middle Eocene) in northwestern Argentina. We provide detailed descriptions, comparisons, and infer foot stances and range of movements for the taxa implicated. Material studied includes astragali belonging to the oldfieldthomasiid Colbertia lumbrerense (Lower Lumbrera Formation), a set of proximal tarsals referred as Typotheria indet. (Lower Lumbrera Formation), and tarsals (also including navicular and cuboid) of the informal taxon "Campanorco inauguralis" (Upper Lumbrera Formation). The comparison of the tarsals of Colbertia lumbrerense (middle Eocene of Argentina) with Colbertia magellanica (early Eocene of Brazil) reveals several differences including variations on the development and arrangement of articular facets, and the size of the dorsal astragalar foramen in the Argentinean species. The specimen of Typotheria indet. shows morphological affinities with basal interatheriid taxa. However, its larger size contrasting with the overall small body sizes of Eocene interatheriids precludes an indisputable taxonomic assignment. Concerning "Campanorco inauguralis", our observations indicate that there is no morphological evidence for a close phylogenetic relationship with Mesotheriidae. It presents a "reversed alternating tarsus" condition, which is also observed in Leontiniidae, "Notohippidae", Toxodontidae, and some typotherians. However, the spectrum of singularities exhibited by this form precludes the assessment of its relationships in the context of the Paleogene radiation of Typotheria and it is necessary to extend the comparison to Eocene notoungulates. Finally, in a morphofunctional context a plantigrade foot posture is inferred for the specimens here reported. These observations have the potential to provide functional proxies for paleoecological reconstructions to be applied to the study of the early radiation of these notoungulate faunas. [ABSTRACT FROM AUTHOR]

South American native ungulates include several extinct lineages that evolved within the context of South American Cenozoic geographic isolation. By the late early Miocene Santacrucian Age, the orders Notoungulata, Litopterna, and Astrapotheria were differentiated. Recent ecomorphological studies highlighted complex functional patterns between their habitat/diet and hypsodonty. In order to understand these relationships in an evolutionary context we used geometric morphometrics methods (27 3-D landmarks) to evaluate the morphological integration of two of the primary functional units of the mandible: the alveolar region and the ascending ramus. Principal component analyses (PCA) and two blocks Partial Least Squares analyses (PLS) were used to explore the patterns of variation and covariation between these two modules. The main variation in shape is related to the robusticity and curvature of the mandibular corpus, and the relative size of the angular and coronoid process. The PLS analyses show a very strong morphological integration within the Litopterna, but a pattern of modularity in the Notoungulata. There is a common pattern of covariation in Astrapotheria and Litopterna, which might be due to similar constraints. The different covariation patterns in notoungulates with respect to litopterns are in concordance with both the taxonomic and morphological diversity of the clades, particularly in the typotheres. The modularity present in notoungulates could have allowed specializations in each module of the mandible, related to crown height or muscle attachments, without a high morphological constrain. This could have resulted in their high morphological diversity, in contrast with the more conservative morphology present in litopterns.

The Interatheriinae (Notoungulata, Interatheriidae) from the Collón Curá Formation (Colloncuran South American Land Mammal Age, SALMA) are revised here, based on old and new collections from western Neuquén, Río Negro, and Chubut provinces where this geologic unit crops out. After a detailed study of the holotype of Icochilus endiadys, we conclude that its cranial and dental morphology are diagnostic of the genus Protypotherium, and as a result we include I. endiadys in this genus (P. endiadys comb. nov.). Deciduous dentition and postcranial remains are also ascribed to P. endiadys, which allows us to determine its pattern of dental eruption and describe part of its limbs, expanding its diagnosis. In addition, we describe a new species of Protypotherium, P. colloncurensis sp. nov., which differs from P. endiadys in having larger size, a more robust mandible, strongly imbricate upper molars, a well-developed parastyle on P1, a subcircular and non-overlapping p1, a much reduced p2, and a smaller talonid on p3–4. Based on the revision, we identified only one genus of Interatheriinae in the Collón Curá Formation (i.e., Protypotherium). Protypotherium endiadys extends its distribution from Neuquén to Chubut provinces, including Río Negro; the new species, in turn, was only recognized in Río Negro Province, appearing together with P. endiadys in Estancia El Criado, Comallo, and Chico River localities. Discriminant and cladistic analyses were performed including P. endiadys, the new taxon, and other middle Miocene interatheres, and yielded strong support for our systematic conclusions. A thorough phylogenetic analysis of Protypotherium is provided for the first time. Based on this analysis, Protypotherium and Miocochilius form a monophyletic group supported by four synapomorphies.

Hegetotheriidae is one of the most derived families of South American notoungulates, which are known from the late Oligocene to Pliocene. Several genera of hegetotheriids have been described; most of them have been recently re-examined, but the Miocene taxa Hegetotherium and Pachyrukhos have not been revised for more than a century. Numerous species of both genera were described mainly from the Santa Cruz Formation, Santa Cruz Province, but also from the Colhuehuapian beds in Chubut Province, and the Colloncuran beds in Neuquén and Río Negro provinces, Argentina. Recently, Hegetotherium was recorded in Bolivia and Chile. Sinclair was the first reviewer to postulate some synonymies, in the first decade of the twentieth century. The present contribution provides a re-evaluation of these genera, including the study of previously unpublished materials. We conclude that Hegetotherium mirabile and Pachyrukhos moyani are the only hegetotheres in the Santa Cruz Formation (Santacrucian SALMA) with Hegetotherium novum and Pachyrukhos politus in the Colhuehuapian SALMA. Additionally, H. convexum, H. anceps, H. minum and H. andinum are proposed as synonyms of H. mirabile; H. novum is removed from Prohegetotherium; and Hegetotherium arctum is excluded from Hegetotheriidae. This taxonomic revision led to a new phylogenetic analysis of Hegetotheriidae, based on a previous data matrix, but enlarged and corrected, as some taxonomic issues concerning other genera were revealed. The analysis supports the monophyly of Hegetotheriidae and the subfamily Pachyrukhinae, as well as the monophyly of the genera Hemihegetotherium and Paedotherium, if the proposed removal of Pd. kakai from the latter is admitted. In contrast, Hegetotheriinae and Hegetotherium appear to be paraphyletic. The position of H. novum supports its removal from the genus Prohegetotherium, in which it was recently included. [ABSTRACT FROM AUTHOR]

Several systematic revisions of the Hegetotheriidae have been published in the last twenty years, but some important taxonomic work remains limited to unpublished studies. In one such thesis, the late Oligocene genera Prosotherium and Propachyrucos were revised, so that Prosotherium garzoni and Propachyrucos smithwoodwardi were considered the only valid species for each genus. Later authors loosely accepted these informal revisions, but recent phylogenetic coding has yet to accurately reflect these revisions, with character states often at odds with the known material for these genera. This study re-examines most of the fossil material assigned to Propachyrucos and Prosotherium , including both published and unpublished fossils, and contributes to a better understanding of the systematics, taxonomic diversity and early morphological evolution of the pachyrukhines. We conclude that Prosotherium garzoni and Propachyrucos smithwoodwardi are indeed the only valid species for each genus, confirming the influential but unpublished revisions that have become widely accepted. The known material for these species is clarified to aid future phylogenetic studies. At the same time, Propachyrucos aequilatus and Pr. robustus are herein considered nomina dubia. Additionally, Miocene species originally ascribed to Propachyrucos , along with unpublished material loosely assigned to the same genus, are here synonymized with other genera. Finally, several fossils previously recognized as Prosotherium sp. are here recognized as P. garzoni. [ABSTRACT FROM AUTHOR]

Here we describe a new notoungulate mammal from the early Eocene (Itaboraian SALMA) of Chubut, Argentinian Patagonia, from the localities of Las Flores and Las Violetas Farm, represented by fragments of maxilla and isolated teeth. The specimens were found in the Las Flores Formation, Río Chico Group, and assigned to the Henricosborniidae, a primitive family within the order Notoungulata. Orome deepi gen. et. sp. nov. differs from other henricosborniids in a less developed metacone column in the upper molars, a larger metaloph in the third upper molar and a larger mean size. The morphometric analysis does not show any difference between the individuals of the two studied localities. The new addition to the henricosborniid diversity in Patagonia allows to provide a better understanding of the early radiation of notoungulates in South America. [ABSTRACT FROM AUTHOR]

Here, we describe the first skeletal remains of Notostylops recovered from middle Eocene levels of the Sarmiento Formation, Patagonia, Argentina. The remains include two teeth of Notostylops murinus, the axis, vertebral bodies, a rib, a left humerus, both radii, two metapodials, two phalanges, the pelvis, a right femur, a right calcaneus and several broken bones. Radial bones are not fused to ulnas, and are shorter than the humerus, very generalized, with an oval head, a marked neck and a radial tubercle. The humerus and the femur show pronounced insertion structures. Our analysis suggests that the appendicular skeleton of Notostylops is too generalized and shares several features with that of terrestrial rodents as Sciuridae. Unlike the appendicular skeletons of cursorial or saltatorial mammals, which restrict mobility, the skeleton of Notostylops indicates the ability to make a variety of different movements, as would be expected for terrestrial, fossorial or arboreal mammals. This skeleton gives new information about the locomotor behaviour of notoungulates, particularly in their basal forms. The results will also allow the identification of isolated notoungulate bones and raise questions about the previous taxonomic assignment of postcrania to Pleurostylodon. [ABSTRACT FROM AUTHOR]

Paedotherium was the last representative of the pachyrukhines (Notoungulata, Hegetotheriidae), a small rabbit- or caviomorph rodent-like clade, with hypsodont and simplified dentition. In contrast to the Pliocene Paedotherium species, the Miocene species are less known and mainly represented by teeth. Different systematic studies of Paedotherium based on the same set of subtle trait variations of the cheek teeth have arrived at different proposals. With the aim of testing the validity and analysing the morphological variation of Paedotherium species, and assessing the definition and diagnostic value of teeth traits, we performed a dual approach: a traditional qualitative study and exploratory geometric morphometric analyses of the upper and lower cheek teeth (both, partial and complete series). In addition, the configurations of cheek tooth shape were used as landmark characters, in combination with traditional characters, to perform a phylogenetic analysis. We propose the validity of the late Miocene P. borrelloi Zetti, and the assignment of north-western Argentinian representatives of Paedotherium to P. aff. minor. Paedotherium borrelloi is morphologically more similar and phylogenetically more closely related to the Pliocene P. bonaerense than to other Paedotherium species. Our results suggest that during the late Miocene the coexistence of two Paedotherium lineages was already established, whose main morphological differences persisted or even intensified in Pliocene representatives, supporting a more complex evolutionary scenario than previously believed. [ABSTRACT FROM AUTHOR]

Abstract Old and new specimens from the early Miocene middle member of the Mariño Formation (Mendoza, Argentina) allow for a reinterpretation of a controversial Mesotheriidae taxon. Earlier known remains come from the Divisadero Largo locality and were, depending on the specimen, either previously described as Trachytherus? mendocensis or referred to a coeval taxon from Chile, Altitypotherium chucalensis. Based on new specimens, here described in detail for the first time, we propose that all mesotheriid material from Divisadero Largo, as well as new material from the Potrerillos locality belong to a new mesotheriid taxon. Recognition of the same taxon from these two localities establishes a sound biostratigraphic correlation. The combination of morphological characters leads to the proposal of a new generic name, Rusconitherium , for Trachytherus? mendocensis , establishing Rusconitherium mendocense New genus and New Combination. This nomenclatural act is due to the fact that T.? mendocensis , formerly thought as a Trachytheriinae from the Paleogene Divisadero Largo Formation, actually comes from the Mariño Formation and belongs to the same taxon as the new material. The missing holotype of T.? mendocensis prompts us to propose a neotype and a new diagnosis for this species, which better characterize the Santacrucian mesotheriids from Mendoza. A phylogenetic analysis does not support a close relationship between this species and Altitypotherium (or any other mesotheriine genus) counter to a previous hypothesis. An unnamed new taxon from the Chinches Formation (San Juan Province) could belong to Rusconitherium , but its full description is pending, as well as a thorough revision of most Mesotheriinae members. Sediments at Potrerillos reveal two paleoenvironmental scenarios: dune fields, influenced by volcanic events, and capillary water that provided humid conditions encouraging vegetation and fauna. Highlights • A new mesotheriid genus from the Miocene Mariño Formation, NW Argentina, is described. • A new taxonomic combination and a neotype are proposed. • Sedimentary and paleoenvironmental approaches are provided for the Potrerillos locality. [ABSTRACT FROM AUTHOR]

A detailed anatomical analysis is here presented focused on a notoungulate skull recovered from sediments of the lower part of the Quebrada de los Colorados Formation (LC I; late middle Eocene), cropping out in Salta Province, Argentina. The material was identified as a Toxodontia, although it does not exhibit information useful enough for its generic or even familiar assignment. The description was mainly focused on the basicranium and the auditory region (especially the petrosal and auditory ossicles) given the better preservation of these parts in the specimen. Additionally, a phylogenetic analysis was conducted based on previous studies dealing with a wide sample of South American Native Ungulates. This specimen exhibits some traits traditionally mentioned for the suborder Toxodontia, such as the horizontal bullar septum, and others recently regarded as synapomorphies of Notoungulata, including the lateral location of the tensor tympani fossa and the expanded medial margin of the petrosal. However, it also shows some unexpected features for that suborder of notoungulates, such as the presence of a strongly curved promontorium. The study of this skull increases our knowledge of the auditory anatomy of Eocene Toxodontia and yields a good opportunity to test current phylogenetic hypothesis, mostly based on Neogene representatives. [ABSTRACT FROM AUTHOR]

South American native ungulates include several extinct lineages that evolved within the context of South American Cenozoic geographic isolation. By the late early Miocene Santacrucian Age, the orders Notoungulata, Litopterna, and Astrapotheria were differentiated. Recent ecomorphological studies highlighted complex functional patterns between their habitat/diet and hypsodonty. In order to understand these relationships in an evolutionary context we used geometric morphometrics methods (27 3-D landmarks) to evaluate the morphological integration of two of the primary functional units of the mandible: the alveolar region and the ascending ramus. Principal component analyses (PCA) and two blocks Partial Least Squares analyses (PLS) were used to explore the patterns of variation and covariation between these two modules. The main variation in shape is related to the robusticity and curvature of the mandibular corpus, and the relative size of the angular and coronoid process. The PLS analyses show a very strong morphological integration within the Litopterna, but a pattern of modularity in the Notoungulata. There is a common pattern of covariation in Astrapotheria and Litopterna, which might be due to similar constraints. The different covariation patterns in notoungulates with respect to litopterns are in concordance with both the taxonomic and morphological diversity of the clades, particularly in the typotheres. The modularity present in notoungulates could have allowed specializations in each module of the mandible, related to crown height or muscle attachments, without a high morphological constrain. This could have resulted in their high morphological diversity, in contrast with the more conservative morphology present in litopterns.

The Interatheriinae (Notoungulata, Interatheriidae) from the Collón Curá Formation (Colloncuran South American Land Mammal Age, SALMA) are revised here, based on old and new collections from western Neuquén, Río Negro, and Chubut provinces where this geologic unit crops out. After a detailed study of the holotype of Icochilus endiadys, we conclude that its cranial and dental morphology are diagnostic of the genus Protypotherium, and as a result we include I. endiadys in this genus (P. endiadys comb. nov.). Deciduous dentition and postcranial remains are also ascribed to P. endiadys, which allows us to determine its pattern of dental eruption and describe part of its limbs, expanding its diagnosis. In addition, we describe a new species of Protypotherium, P. colloncurensis sp. nov., which differs from P. endiadys in having larger size, a more robust mandible, strongly imbricate upper molars, a well-developed parastyle on P1, a subcircular and non-overlapping p1, a much reduced p2, and a smaller talonid on p3-4. Based on the revision, we identified only one genus of Interatheriinae in the Collón Curá Formation (i.e., Protypotherium). Protypotherium endiadys extends its distribution from Neuquén to Chubut provinces, including Río Negro; the new species, in turn, was only recognized in Río Negro Province, appearing together with P. endiadys in Estancia El Criado, Comallo, and Chico River localities. Discriminant and cladistic analyses were performed including P. endiadys, the new taxon, and other middle Miocene interatheres, and yielded strong support for our systematic conclusions. A thorough phylogenetic analysis of Protypotherium is provided for the first time. Based on this analysis, Protypotherium and Miocochilius form a monophyletic group supported by four synapomorphies. [ABSTRACT FROM AUTHOR]