Astragalar Morphology of Selected Giraffidae

The artiodactyl astragalus has been modified to exhibit two trochleae, creating a double pullied structure allowing for significant dorso-plantar motion, and limited mediolateral motion. The astragalus structure is partly influenced by environmental substrates, and correspondingly, morphometric studies can yield paleohabitat information. The present study establishes terminology and describes detailed morphological features on giraffid astragali. Each giraffid astragalus exhibits a unique combination of anatomical characteristics. The giraffid astragalar morphologies reinforce previously established phylogenetic relationships. We find that the enlargement of the navicular head is a feature shared by all giraffids, and that the primitive giraffids possess exceptionally tall astragalar heads in relation to the total astragalar height. The sivatheres and the okapi share a reduced notch on the lateral edge of the astragalus. We find that Samotherium is more primitive in astragalar morphologies than Palaeotragus, which is reinforced by tooth characteristics and ossicone position. Diagnostic anatomical characters on the astragalus allow for giraffid species identifications and a better understanding of Giraffidae.     

On the existence of non‐microbiotherian Australidelphian marsupials (Diprotodontia) in the Eocene of Patagonia

A diverse assemblage of extinct mammals of early–middle Eocene age (Ypresian–Lutetian boundary) come from the Patagonian localities of La Barda and Laguna Fría around Paso del Sapo in northwestern Chubut Province (Argentina). Metatherians are well represented, mostly by dental remains of ‘Didelphimorphia’, Paucituberculata, Sparassodonta, Microbiotheria, and Polydolopimorphia. Here we analyse three calcanea and one astragalus referable to the same, indeterminate taxon, from La Barda, showing the fusion of their ectal and sustentacular facets. This facet arrangement characterizes the Australidelphia, a marsupial clade represented by Microbiotheria from South America and Antarctica, plus all Australasian lineages. Other australidelphian features shown by these tarsals include: in the calcanea, a reduced peroneal process and a tripartite cuboid facet; in the astragalus, a round, very small astragalar head in relation to the body; a convex trochlea with a half for the tibia and half, in slightly lower position, for the fibula. Their size and other anatomical features suggest that the new tarsals cannot be referred to the Microbiotheria. Phylogenetic analysis suggests that the La Barda taxon lies within the Australidelphia, and that it is either closely related or belongs to, the Diprotodontia. The existence of advanced australidelphians in Patagonia adds further evidence of the close relationship between Patagonia, Antarctica, and Australia during the Late Cretaceous – early Palaeogene.     

Molecular phylogenetic relationships of moles,shrew moles,and desmans from the new and old worlds

A Rich variety of anatomical and physiological specializations has enabled members of the family Talpidae (moles, shrew moles, and desmans) to exploit a diverse range of habitats: terrestrial, semi-aquatic, aquatic/fossorial, semi-fossorial, and fossorial. While numerous morphological and biochemical studies pertaining to the origin and radiation of the Talpidae have been completed, phylogenetic hypotheses remain controversial. To address this shortcoming we sequenced the mitochondrial DNA cytochrome b gene (1140bp) from 29 individuals spanning 12 talpid species. Phylogenetic trees incorporating 12 New and Old World genera (18 species; all 3 extant subfamilies) were then constructed using NJ, MP, ML, and NJ-ML (NJ with ML parameters) methods. Our results provide molecular support for a mononphyletic Talpidae, and suggest that the 12 genera are clustered into seven major clades; (1) Asiatic shrew-like moles (Uropsilus), (2) North American aquatic/fossorial moles (Condylura), (3) North American fossorial moles (Parascalops, Scalopus, and Scapanus), (4) North American semi-fossorial shrew moles (Neurotrichus), (5) Japanese semi-fossorial shrew moles (Dymecodon and Urotrichus), (6) European semi-aquatic desmans (Desmana), and (7) Eurasian fossorial moles (Euroscaptor, Mogera, and Talpa). None of these groupings comprised mole species from both continents. In fact, North American moles and shrew moles do not appear to have specific affinities with Asian moles and shrew moles, respectively. Although low bootstrap support was generally found for evolutionary nodes uniting the major talpid clades, all gene trees constructed identified fossorial North American and Eurasian mole lineages as nonmonophyletic groups, suggesting subterranean specializations arose independently at least twice during the evolution of the Talpidae. Additionally, our data set provides molecular support for a basal divergence and long independent history of Uropsilus from the main talpid line, and refutes the traditional taxonomic status and secondarily basal phylogenetic placement of the subfamily Desmaninae within the Talpidae.     

A new species of Megatherium (Mammalia: Xenarthra: Megatheriidae) from the Pleistocene of Sacaco and Tres Ventanas, Peru

Megatherium urbinai sp. nov. is a new megatheriine ground sloth from Pleistocene beds of the Sacaco area and Tres Ventanas Cave, Peru. This new species is referred to the genus Megatherium based primarily on humeral, femoral, and astragalar morphology. It is characterized by various features, such as a strongly reduced metacarpal-carpal-complex (MCC) and a reduced, oval scaphoid facet of the MCC located at the medial extremity of the bone that does not contact without the McII facet. The distal side of the unciform bears an undivided articular surface for the metacarpals III–V. The lateral femoral margin is nearly rectilinear and the tibia and fibula do not fuse distally. The astragalus is oriented more medially, the navicular is in a medial position, the ectocuneiform articulates with MtIII–IV, and the MtIV is more inclined laterally than in M. tarijense and M. americanum . M. urbinai is one of the smallest Megatherium species known along with M. altiplanicum . It has the most marked torsion of the pes for the genus. It extends the palaeogeographic distribution of the genus in South America. It is phylogenetically close to other Andean megatheriines and is less specialised than M. tarijense , and does not belong to the clade constituted by [ M. altiplanicum  +  M. americanum ].     

Stance and gait in the hindlimb of a therocephalian mammal-like reptile

The ilium, femur, tibia, fibula, astragalus and calcaneum of the therocephalian therapsid Regisaurus jacobi Mendrez are described. Functional consideration of the bones and particularly the nature of the hip, knee and ankle joints indicate that this animal was capable of at least two distinct gaits, a primitive, sprawling type and an advanced upright type. The ankle joint had a moveable articulation between the astragalus and the calcaneum analogous to that of the crocodiles. The possibility of such a multi-gaited intermediate stage having occurred in the course of the evolution of the mammalian locomotory pattern is suggested.     

Fossil primates and the evolution of some primate locomotor systems

Of Paleocene primates only Plesiadapis is complete enough to reconstruct locomotor patterns; it was an arboreal scrambler, perhaps functioning like a large squirrel. Eocene lemurs (adapids) show an array of locomotor types much like certain modern Malagasy lemurs. The European Eocene tarsiid Necrolemur and the American Hemiacodon show the beginning of saltatory specializations in possession of elongated calcaneum and astragalus. Although not a direct anthropoid ancestor Necrolemur seems one of the best models for representing the early locomotor type from which higher primates arose. The Oligocene primates of Egypt (among which are the earliest undoubted pongids) are preserved with a forest fauna. Structures of long bones suggest they were arboreal. A considerable number of Miocene ape bones are known and those of Pliopithecus and Dryopithecus indicate similar adaptations. Of African Miocene forms, Dryopithecus major was a large, gorilla-sized animal, and hence perhaps primarily terrestrial. D. africanus was somewhat more arboreally adapted and a partial brachiator. The Italian fossil Oreopithecus, a coal-swamp dweller, shows indications of bipedality in pelvic structure. Ramapithecus, which is presumably ancestral to Australopithecus, shows palatal and facial patterns much like these later hominids, and probably hence had locomotor patterns more like men than like living apes; its lack of the dental specializations of apes strongly supports this suggestion.     

Astragalar morphology of late Eocene anthropoids from the Fayum Depression (Egypt) and the origin of catarrhine primates

The phylogenetic relationships of the late Eocene anthropoids Catopithecus browni and Proteopithecus sylviae are currently a matter of debate, with opinion divided as to whether these taxa are stem or crown anthropoids. The phylogenetic position of Catopithecus is of particular interest, for, unlike the highly generalized genus Proteopithecus, this taxon shares apomorphic dental and postcranial features with more derived undoubted catarrhines that appear in the same region 1-2 Ma later. If these apomorphies are homologous and Catopithecus is a stem catarrhine, the unique combination of plesiomorphic and apomorphic features preserved in this anthropoid would have important implications for our understanding of the crown anthropoid morphotype and the pattern of morphological character transformations that occurred during the early phases of stem catarrhine evolution.Well-preserved astragali referrable to Proteopithecus, Catopithecus, and the undoubted early Oligocene stem catarrhine Aegyptopithecus have provided additional morphological evidence that allows us to further evaluate competing hypotheses of interrelationships among Eocene-Oligocene Afro-Arabian anthropoids. Qualitative observations and multivariate morphometric analyses reveal that the astragalar morphology of Proteopithecus is very similar to that of early Oligocene parapithecids and living and extinct small-bodied platyrrhines, and strengthens the hypothesis that the morphological pattern shared by these taxa is primitive within crown Anthropoidea. In contrast, Catopithecus departs markedly from the predicted crown anthropoid astragalar morphotype and shares a number of apomorphic features (e.g., deep cotylar fossa, laterally projecting fibular facet, trochlear asymmetry, mediolaterally wide astragalar head) with Aegyptopithecus and Miocene-Recent catarrhines. The evidence from the astragalus complements other independent data from the dentition, humerus and femur of Catopithecus that support this taxon's stem catarrhine status, and we continue to maintain that oligopithecines are stem catarrhines that constitute the sister group of a clade containing propliopithecines and Miocene-Recent catarrhines.     

The morphology of the cerebellar nuclei of Galago and Tupaia

The cerebellar nuclei of the lesser bushbaby (Galago senegalensis) and the tree shrew (Tupaia glis) were studied. The cerebellar nuclear grey of Galago is divisible into a medial nucleus, a nucleus interpositus anterior, a nucleus interpositus posterior, and a lateral nucleus. The lateral nucleus is slightly concave medially suggestive of a primitive hilus. The interpositus nucleus is divided into anterior and posterior portions by a delicate lamina of fibers. The medial cerebellar nucleus is an irregular mass of cells located dorsal to the fourth ventricle. The cerebellar nuclear grey of Tupaia is also divisible into a medial nucleus, a nucleus interpositus anterior, a nucleus interpositus posterior, and a lateral cerebellar nucleus. The medial cerebellar nucleus is located dorsal to the fourth ventricle. The nucleus interpositus anterior and nucleus interpositus posterior are joined together and with the lateral nucleus in the caudo-ventral region. The NIA and NIP have an anterior-posterior relationship to each other and the lateral nucleus has no apparent undulations suggestive of early sacculations. The configuration of the cerebellar nuclei of Tupaia more closely resembles the more primitive patterns of the rat, hedgehog, and mole than those of Galago or other primates.     

Additions to the knowledge ofUrumaquia robusta (Xenarthra, Phyllophaga, Megatheriidae) from the Urumaco Formation (Late Miocene), Estado Falcön, Venezuela

We describe new remains ofUrumaquia robusta Carlini, Brandoni &; Sänchez, 2006, which significantly improve the knowledge of the species and allow a more precise discussion about the relationships with the rest of the South American Megatheriinae. The new remains (ulna, phalanx, femora, patella, tibia, astragalus, navicular, metatarsals IV–V) described here suggest close affinities of this taxon with the species ofPyramiodontherium and withMegatheriops rectidens, based on morphological similarities of the ulna (length, gracility), femur (general morphology, shape of the medial margin), and astragalus (angle between the discoidal and odontoid facets). In fact, the shape of femoral medial margin is a shared derived character amongUrumaquia, Pyramiodontherium andMegatheriops. However, and despite the increasing information concerning Tertiary species, their phylogenetic relationships are only partially understood.     

Predicting euarchontan body mass: A comparison of tarsal and dental variables

Objective: Multiple meaningful ecological characterizations of a species revolve around body mass. Because body mass cannot be directly measured in extinct taxa, reliable body mass predictors are needed. Many published body mass prediction equations rely on dental dimensions, but certain skeletal dimensions may have a more direct and consistent relationship with body mass. We seek to evaluate the reliability of prediction equations for inferring euarchontan body mass based on measurements of the articular facet areas of the astragalus and calcaneus. Methods: Surface areas of five astragalar facets (n = 217 specimens) and two calcaneal facets (n = 163) were measured. Separate ordinary least squares and multiple regression equations are presented for different levels of taxonomic inclusivity, and the reliability of each equation is evaluated with the coefficient of determination, standard error of the estimate, mean prediction error, and the prediction sum of squares statistic. We compare prediction errors to published prediction equations that utilize dental and/or tarsal measures. Finally, we examine the effects of taxonomically specific regressions and apply our equations to a diverse set of non‐primates. Results: Our results reveal that predictions based on facet areas are more reliable than most linear dental or tarsal predictors. Multivariate approaches are often better than univariate methods, but require more information (making them less useful for fragmentary fossils). While some taxonomically specific regressions improve predictive ability, this is not true for all primate groups. Conclusions: Among individual facets, the ectal and fibular facets of the astragalus and the calcaneal cuboid facet are the best body mass predictors. Since these facets have primarily concave curvature and scale with positive allometry relative to body mass, it appears that candidate skeletal proxies for body mass can be identified based on their curvature and scaling coefficients. Am J Phys Anthropol 157:472–506, 2015. © 2015 Wiley Periodicals, Inc.     

Functional-adaptive anatomy of the forelimb in the Didelphidae, and the paleobiology of the Paleocene marsupials Mayulestes ferox and Pucadelphys andinus

An attempt to determine the locomotor activities of Mayulestes ferox (Borhyaenoidea) and Pucadelphys andinus (Didelphoidea) from the early Paleocene site of Tiupampa (Bolivia) is presented. The functional anatomy of the forelimbs of these South American marsupials is compared to that of some living didelphids: Caluromys philander, Micoureus demerarae, Marmosa murina, Didelphis marsupialis, Monodelphis brevicaudata and Metachirus nudicaudatus. Deductions from bone morphology to myology and locomotor behavior in the fossils are inferred from the comparisons with living forms. Some features of the postcranial skeleton, indicative of arboreal adaptations, are found in the extinct marsupials: anteriorly projected acromion, hemispherical head of the humerus, extended humeral lateral epicondylar ridge, medially protruding humeral entepicondyle, proximal ulnar posterior convexity, and deep flexor fossa on the medial side of the ulna. But other features are related to a more terrestrial pattern: the well-developed tubercles of the humeral head, the elongated olecranon process of the ulna, and the oval shape of the radial head. Mayulestes had clear arboreal abilities, but, as a predaceous mammal, probably hunted on the ground. Pucadelphys was less specialized, close to the living Monodelphis, a terrestrial insectivorous form with some skeletal features related to arboreal locomotion that are probably plesiomorphic for marsupials.     

Effects of iron deficiency on the growth and photosynthesis of three bloom‐forming cyanobacterial species isolated from Lake Taihu

Cyanobacterial blooms are found in many freshwater ecosystems around the world, but the effect of environmental factors on their growth and the proportion of species still require more investigation. In this study, the physiological responses of bloom‐forming cyanobacteria M icrocystis aeruginosa FACHB912, M icrocystis flos‐aquae FACHB1028 and P seudanabaena sp. FACHB1282 to iron deficiency were investigated. Their specific growth rates were found to decrease as the available iron concentration decreased. At low available iron concentrations of 1 × 10^?7 M (pFe 21.3) and 5 × 10^?8 M (pFe 21.6), M . aeruginosa had the lowest specific growth rate among three studied species. The cell sizes of M . flos‐aquae and Pseudanabaena sp. were significantly smaller under the lowest iron concentration. The chlorophyll a content of the three species decreased at the lowest iron concentration. The maximal relative electron transport rate, photosynthetic efficiency, and light‐saturation parameter of M . aeruginosa were lower than the other two cyanobacteria at pFe 21.3. Therefore, M . aeruginosa was the least able to adapt to iron deficiency. Under iron deficiency, the functional absorption cross‐section of PSII and electron transport rate on the acceptor side of PSII decreased in M . aeruginosa, while the connectivity factor between individual photosynthetic units increased in M . flos‐aquae, and the electron transport rate on the acceptor side of PSII and between PSII and PSI decreased in P seudanabaena sp. The ability to store iron was highest in M . flos‐aquae, followed by P seudanabaena sp. and M . aeruginosa. Thus, these results provide necessary information for detecting the role of iron in the succession of cyanobacterial species in Lake Taihu, the third largest freshwater lake in China, because all three species were isolated from this lake.     

A new species of Pseudoloris (Omomyidae,Primates) from the middle Eocene of Sant Jaume de Frontanyà (Eastern Pyrenees,Spain)

In this article we describe a new species of Pseudoloris (Omomyidae, Primates) from the Robiacian (middle Eocene) locality of Sant Jaume de Frontanyà (Eastern Pyrenees, Spain). Pseudoloris pyrenaicus is characterized by its medium size, thickened paracristid, absence of a distinct paraconid, and well‐developed buccal cingulid in the lower molars, large hypoconulid in the M₃, paraconule and hypocone reduced in the M¹ and M² and absent in the M³. The material from Sant Jaume de Frontanyà constitutes the most abundant sample of the genus Pseudoloris found until now in the Iberian Peninsula. Almost all the dental elements have been recovered, including those teeth hardly known for other species of the genus, such as lower and upper incisors. The new species shows intermediate features between Pseudoloris isabenae from Capella and Pseudoloris parvulus, present in different Spanish and French sites. Therefore, we consider that Pseudoloris pyrenaicus is an intermediate form between these two species. Am J Phys Anthropol 143:92–99, 2010. © 2010 Wiley‐Liss, Inc.     

The impact of local population genetic background on the spread of the selfish element Medea‐1 in red flour beetles

Selfish genetic elements have been found in the genomes of many species, yet our understanding of their evolutionary dynamics is only partially understood. A number of distinct selfish Medea elements are naturally present in many populations of the red flour beetle (Tribolium castaneum). Although these Medea elements are predicted by models to increase in frequency within populations because any offspring of a Medea‐bearing mother that do not inherit at least one Medea allele will die, experiments demonstrating an increase in a naturally occurring Medea element are lacking. Our survey of the specific Medea element, M¹, in the United States showed that it had a patchy geographic distribution. From the survey, it could not be determined if this distribution was caused by a slow process of M¹ colonization of discrete populations or if some populations lacked M¹ because they had genetic factors conferring resistance to the Medea mechanism. We show that populations with naturally low to intermediate M¹ frequencies likely represent transient states during the process of Medea spread. Furthermore, we find no evidence that genetic factors are excluding M¹ from US populations where the element is not presently found. We also show how a known suppressor of Medea can impair the increase of M¹ in populations and discuss the implications of our findings for pest‐management applications of Medea elements.     

New chiropterans from the middle Eocene of Shanghuang (Jiangsu Province,Coastal China): new insight into the dawn horseshoe bats (Rhinolophidae) in Asia

Until recently, the fossil record of Paleogene bats in Asia primarily included extinct families (i.e. ‘Eochiroptera’) from the early Eocene of Vastan in India and from the middle‐late Eocene of the Liguanqiao and Yuanqu basins in central China. Here, we describe a new fauna of Chiroptera from the middle Eocene Shanghuang fissure fillings of China. The fauna includes abundant material referred to a new rhinolophid (Protorhinolophus shanghuangensis gen. and sp. n.), one specimen of a possible rhinopomatid and several indeterminate rhinolophoids. This new bat assemblage constitutes the earliest record of extant families of microbats in Asia. Because it lacks representatives of ‘Eochiroptera’, this Shanghuang bat fauna indicates significant turnover in Asian bat communities. The dental pattern of P. shanghuangensis shows a mosaic of primitive and derived features (‘Eochiroptera’ vs Rhinolophidae dental characteristics), suggesting that this taxon occupies a basal position among the Rhinolophidae. Rhinolophids were already well diversified at the end of the late Eocene in Europe. Interestingly, many dental characteristics of Protorhinolophus are also found in a primitive rhinolophoid taxon, Vaylatsia, from the middle Eocene to late Oligocene of Europe, supporting a close relationship between these taxa. These affinities testify to the widespread Eurasian distribution of rhinolophoids during the Eocene and are consistent with a westward dispersal of the group from eastern Asia to Europe owing to the greater antiquity of Protorhinolophus.     

The distribution and spread of naturally occurring Medea selfish genetic elements in the United States

Selfish genetic elements (SGEs) are DNA sequences that are transmitted to viable offspring in greater than Mendelian frequencies. Medea SGEs occur naturally in some populations of red flour beetle (Tribolium castaneum) and are expected to increase in frequency within populations and spread among populations. The large‐scale U.S. distributions of Medea‐4 (M⁴) had been mapped based on samples from 1993 to 1995. We sampled beetles in 2011–2014 and show that the distribution of M⁴ in the United States is dynamic and has shifted southward. By using a genetic marker of Medea‐1 (M¹), we found five unique geographic clusters with high and low M¹ frequencies in a pattern not predicted by microsatellite‐based analysis of population structure. Our results indicate the absence of rigid barriers to Medea spread in the United States, so assessment of what factors have limited its current distribution requires further investigation. There is great interest in using synthetic SGEs, including synthetic Medea, to alter or suppress pest populations, but there is concern about unpredicted spread of these SGEs and potential for populations to become resistant to them. The finding of patchy distributions of Medea elements suggests that released synthetic SGEs cannot always be expected to spread uniformly, especially in target species with limited dispersal.     

Anatomy and relationships of Pachyrhachis problematicus,a primitive snake with hindlimbs

The anatomy of Pachyrhachis problematicus, an elongate, limb-reduced squamate from the Upper Cretaceous of Israel, is described and evaluated in detail. Previously considered a snake-like ''lizard'' of uncertain affinities, it is here shown to be the most primitive snake, and the sister-group to all other snakes. Pachyrhachis exhibits numerous derived characters uniting it with modern snakes (scolecophidians and alethinophidians): e.g. mobile premaxilla-maxilla articulation, braincase enclosed by frontals and parietals, sagittal parietal crest, absence of tympanic recess, single postdentary bone, over 140 presacral vertebrae, and complete loss of shoulder girdle and forelimb. However, it is more primitive than all modern snakes in retaining some strikingly primitive (lizard-like) features: presence of a jugal, squamosal, normal sacral attachment, and well-developed hindlimb composed of femur, tibia, fibula, and tarsals. Pachyrhachis provides additional support for the hypothesis that snakes are most closely related to Cretaceous marine lizards (mosasauroids). Almost all of the derived characters proposed to unite snakes and mosasauroids are highly developed in Pachyrhachis: the mobile mandibular symphysis, intramandibular joint, long and recurved pterygoid teeth, quadrate suspended by the supratemporal, loosely united pelvic elements (ilium, ischium, and pubis), and separate astragalus and calcaneum.     

Effect of HgCl2 on acetylcholine,carbachol, and glutamate currents ofAplysia neurons

Summary 1. Using conventional two-microelectrode voltage-clamp techniques we studied the effects of inorganic mercury (HgCl₂) on acetylcholine-, carbachol-, and glutamate-activated currents onAplysia neurons. Hg²⁺ was applied with microperfusion.2. Acetylcholine and carbachol activated an inward, sodium-dependent current in the anterior neurons of the pleural ganglion. The medial neurons gave a biphasic current to acetylcholine and carbachol, which was outward at resting membrane potential. The faster component was Cl^– dependent and reversed at about –60 mV, while the slower component was K⁺ dependent and reversed at greater than –80 mV.3. Hg²⁺ (0.1–10 µM) caused a dramatic increase in the acetylcholine- and carbachol-induced inward current in anterior neurons and the fast Cl^– current in medial neurons. With only a 1-min preapplication of Hg²⁺, the acetylcholine- or carbachol-activated sodium or chloride currents were increased to 300% and the effect was only partly reversible. The threshold concentration was 0.1 µM Hg²⁺.4. Contrary to the effects on sodium and chloride currents, concentrations of 0.1–10 µM Hg²⁺ caused a complete and irreversible blockade of K⁺-dependent acetylcholine and carbachol currents. The block of the potassium current was relatively fast and increased with time. The concentration of HgCl₂ that gave a half-maximal blockade of the carbachol-activated potassium current was 0.89 µM. The chloride-dependent current elicited by glutamate on medial neurons was increased by HgCl₂ as well.5. These results suggest that actions at agonist-activated channels must be considered as contributing to mercury neurotoxicity. It is possible that the toxic actions of Hg²⁺ on synaptic transmission at both pre- and postsynaptic sites are important factors in the mechanism of Hg²⁺ toxicity.     

The first described Arsinoitherium from the upper Eocene Aydim Formation of Oman: Biogeographic implications

A new fossiliferous locality is discovered from the upper Eocene Aydim Formation, in Dhofar, Southern Sultanate of Oman. A left ulna of Arsinoitherium is described, and cranial and postcranial specimens found in close proximity are referred to the same taxon. The locality is promising for the recovery of additional fossil specimens. Moreover, the presence of Arsinoitherium in Oman is of biogeographic significance; as the Red Sea did not exist during the late Eocene, these large-bodied animals were able to freely travel between what is now the Arabian Peninsula and continental Africa.