Blaine W. Schubert

Extinction implications of a chenopod browse diet for a giant Pleistocene kangaroo

Kangaroos are the worlds most diverse group of herbivorous marsupials. Following late-Miocene intensification of aridity and seasonality, they radiated across Australia, becoming the continents ecological equivalents of the artiodactyl ungulates elsewhere. Their diversity peaked during the Pleistocene, but by approximately 45,000 years ago, 90% of larger kangaroos were extinct, along with a range of other giant species. Resolving whether climate change or human arrival was the principal extinction cause remains highly contentious. Here we combine craniodental morphology, stable-isotopic, and dental microwear data to reveal that the largest-ever kangaroo, Procoptodon goliah, was a chenopod browse specialist, which may have had a preference for Atriplex (saltbushes), one of a few dicots using the C4 photosynthetic pathway. Furthermore, oxygen isotope signatures of P. goliah tooth enamel show that it drank more in low-rainfall areas than its grazing contemporaries, similar to modern saltbush feeders. Saltbushes and chenopod shrublands in general are poorly flammable, so landscape burning by humans is unlikely to have caused a reduction in fodder driving the species to extinction. Aridity is discounted as a primary cause because P. goliah evolved in response to increased aridity and disappeared during an interval wetter than many it survived earlier. Hunting by humans, who were also bound to water, may have been a more decisive factor in the extinction of this giant marsupial.

Direct comparisons of 2D and 3D dental microwear proxies in extant herbivorous and carnivorous mammals.

The analysis of dental microwear is commonly used by paleontologists and anthropologists to clarify the diets of extinct species, including herbivorous and carnivorous mammals. Currently, there are numerous methods employed to quantify dental microwear, varying in the types of microscopes used, magnifications, and the characterization of wear in both two dimensions and three dimensions. Results from dental microwear studies utilizing different methods are not directly comparable and human quantification of wear features (e.g., pits and scratches) introduces interobserver error, with higher error being produced by less experienced individuals. Dental microwear texture analysis (DMTA), which analyzes microwear features in three dimensions, alleviates some of the problems surrounding two-dimensional microwear methods by reducing observer bias. Here, we assess the accuracy and comparability within and between 2D and 3D dental microwear analyses in herbivorous and carnivorous mammals at the same magnification. Specifically, we compare observer-generated 2D microwear data from photosimulations of the identical scanned areas of DMTA in extant African bovids and carnivorans using a scanning white light confocal microscope at 100x magnification. Using this magnification, dental microwear features quantified in 2D were able to separate grazing and frugivorous bovids using scratch frequency; however, DMTA variables were better able to discriminate between disparate dietary niches in both carnivorous and herbivorous mammals. Further, results demonstrate significant interobserver differences in 2D microwear data, with the microwear index remaining the least variable between experienced observers, consistent with prior research. Overall, our results highlight the importance of reducing observer error and analyzing dental microwear in three dimensions in order to consistently interpret diets accurately.

Carnivoran dental microwear textures: comparability of carnassial facets and functional differentiation of postcanine teeth

No abstract available

Was the Giant Short-Faced Bear a Hyper-Scavenger? A New Approach to the Dietary Study of Ursids Using Dental Microwear Textures

Dramatic environmental changes associated with global cooling since the late Miocene, and the onset of glacial-interglacial cycles in the Pleistocene served as a backdrop to the evolutionary radiation of modern bears (family Ursidae). These environmental changes likely prompted changes in food availability, and triggered dietary adaptations that served as motive forces in ursid evolution. Here, we assess correspondence of dental microwear textures of first and second lower molars with diet in extant ursids. We use the resulting baseline data to evaluate the hypothesis that the Pleistocene giant short-faced bear, Arctodus simus, was a bone consumer and hyper-scavenger at Rancho La Brea, California, USA. Significant variation along the tooth row is consistent with functional differentiation, with the second molar serving as a better dietary recorder than the first. Results evince significant variation among species: carnivorous and omnivorous ursids (Ursus maritimus, U. americanus) have significantly higher and more variable complexity (Asfc) than more herbivorous ones (Ailuropoda melanoleuca, Tremarctos ornatus, U. malayanus), and A. melanoleuca is differentiated from U. maritimus and U. americanus by significantly higher and more variable anisotropy (epLsar) values. Arctodus simus from Rancho La Brea exhibits wear attributes most comparable to its closest living relative (T. ornatus), which is inconsistent with hard-object (e.g., bone) consumption, and the hypothesis that short-faced bears were bone consuming hyper-scavengers across their range.

Implications of diet for the extinction of saber-toothed cats and American lions.

The saber-toothed cat, Smilodon fatalis, and American lion, Panthera atrox, were among the largest terrestrial carnivores that lived during the Pleistocene, going extinct along with other megafauna ∼12,000 years ago. Previous work suggests that times were difficult at La Brea (California) during the late Pleistocene, as nearly all carnivores have greater incidences of tooth breakage (used to infer greater carcass utilization) compared to today. As Dental Microwear Texture Analysis (DMTA) can differentiate between levels of bone consumption in extant carnivores, we use DMTA to clarify the dietary niches of extinct carnivorans from La Brea. Specifically, we test the hypothesis that times were tough at La Brea with carnivorous taxa utilizing more of the carcasses. Our results show no evidence of bone crushing by P. atrox, with DMTA attributes most similar to the extant cheetah, Acinonyx jubatus, which actively avoids bone. In contrast, S. fatalis has DMTA attributes most similar to the African lion Panthera leo, implying that S. fatalis did not avoid bone to the extent previously suggested by SEM microwear data. DMTA characters most indicative of bone consumption (i.e., complexity and textural fill volume) suggest that carcass utilization by the extinct carnivorans was not necessarily more complete during the Pleistocene at La Brea; thus, times may not have been “tougher” than the present. Additionally, minor to no significant differences in DMTA attributes from older (∼30–35 Ka) to younger (∼11.5 Ka) deposits offer little evidence that declining prey resources were a primary cause of extinction for these large cats.

A Dental Microwear Texture Analysis of the Mio-Pliocene Hyaenids from Langebaanweg, South Africa

Hyaenids reached their peak diversity during the Mio-Pliocene, when an array of carnivorous species emerged alongside dwindling civet-like and mongoose-like insectivorous/omnivorous taxa. Significantly, bone-cracking morphological adaptations were poorly developed in these newly-emerged species. This, their general canid-like morphology, and the absence/rarity of canids in Eurasia and Africa at the time, has led researchers to hypothesise that these carnivorous Mio-Pliocene hyaenas were ecological vicars to modern canids. To shed further light on their diets and foraging strategies, we examine and compare the dental microwear textures of Hyaenictitherium namaquensis, Ikelohyaena abronia, Chasmaporthetes australls, and Hyaenictis hendeyi from the South African Mio-Pliocene site of Langebaanweg with those of the extant feliforms Crocuta crocuta, Acinonyx jubatus, and Panthera leo (caniforms are not included because homologous wear facets are not directly comparable between the suborders). Sample sizes for individual fossil species are small, which limits confidence in assessments of variation between the extinct taxa; however, these Mio-Pliocene hyaenas exhibit surface complexity and textural fill volume values that are considerably lower than those exhibited by the living hyaena, Crocuta crocuta. Dental microwear texture analysis thus supports interpretations of craniodental evidence suggesting low bone consumption in carnivorous Mio-Pliocene hyaenas.

The Largest Known Bear, Arctotherium angustidens, from the Early Pleistocene Pampean Region of Argentina: With a Discussion of Size and Diet Trends in Bears

Abstract The South American giant short-faced bear (Arctotherium angustidens Gervais and Ameghino, 1880) is one of five described Arctotherium species endemic to South America and it is known for being the earliest, largest, and most carnivorous member of the genus. Here we report an extraordinarily large A. angustidens individual exhumed from Ensenadan sediments (early to middle Pleistocene) at Buenos Aires Province, Argentina. Based on overall size, degree of epiphyseal fusion, and pathologies, this bear was an old-aged male that sustained serious injuries during life. Body mass of the bear is estimated and compared to other ursid species based on a series of allometric equations. To our knowledge, this specimen now represents the largest bear ever recorded. In light of this discovery, we discuss the evolution of body size in Arctotherium (from large-to-small) and compare this to bears that exhibited different evolutionary trajectories. We suggest that the larger size and more carnivorous nature of A. angustidens, compared to later members of the genus, may reflect the relative lack of other large carnivores and abundance of herbivores in South America just after the Great American Biotic Interchange.

Helodermatid Lizard from the Mio-Pliocene Oak-Hickory Forest of Tennessee, Eastern USA, and a Review of Monstersaurian Osteoderms

The extant venomous Gila monster and beaded lizards, species of Heloderma, live today in southwestern USA and south along the Pacific coastal region into Central America, but their fossil history is poorly understood. Here we report helodermatid osteoderms (dermal ossicles) from the late Miocene-early Pliocene Gray Fossil Site, eastern Tennessee USA. Twenty-three species of mammals are known from the fauna including abundant Tapirus polkensis, as well as fishes, anurans, salamanders, turtles, Alligator, birds, and snakes. Beaded lizards belong to the Monstersauria, a clade that includes Primaderma + Paraderma + Gobiderma + Helodermatidae (Estesia, Eurheloderma, Lowesaurus, and Heloderma). Osteoderms of lizards in this clade are unique within Squamata; they typically are circular to polygonal in outline, domed to flat-domed in cross-section, have a vermiculate surface texture, are not compound structures, and do not have imbricate surfaces as on many scincomorph and anguid lizards. We review and characterize the osteoderms of all members of Monstersauria. Osteoderms from the cranium, body, and limbs of Heloderma characteristically have a ring-extension (bony flange) at least partly surrounding the dome. Its presence appears to be a key character distinct to all species of Heloderma, consequently, we propose the presence of a ring-extension to be an apomorphy. Three osteoderms from the Gray Fossil Site range from 1.5 to 3.0 mm in diameter, have the circular shape of helodermatid osteoderms with a domed apical surface, and have the ring-extensions, permiting generic identification. Macrobotanical remains from the Gray Fossil Site indicate an oak-hickory subtropical forest dominated by Quercus (oak) and Carya (hickory) with some conifer species, an understorey including the climbing vines Sinomenium, Sargentodoxa, and Vitis. Plant and mammal remains indicate a strong Asian influence.

Assessing niche conservatism using a multiproxy approach: dietary ecology of extinct and extant spotted hyenas

Abstract. A central premise of bioclimatic envelope modeling is the assumption of niche conservatism. Whereas such assumptions are testable in modern populations, it is unclear whether niche conservatism holds over deeper time spans and over very large geographic ranges. Hyaenids occupied a diversity of ecological niches over time and space, and until the end-Pleistocene they occurred in Europe and most of Asia, with Asian populations of Crocuta suggested as being genetically distinct from their closest living relatives. Further, little is known regarding whether and how the dietary ecology of extinct populations of Crocuta differed from those of their extant African counterparts. Here, we use a multiproxy approach to assess an assumption of conserved dietary ecology in late Pleistocene extant spotted hyenas via finite element analysis, dental microwear texture analysis, and a novel dental macrowear method (i.e., whether teeth are minimally, moderately, or extremely worn, as defined by degree of dentin exposure) proposed here. Results from finite element simulations of the masticatory apparatus of Chinese and African Crocuta demonstrate lower skull stiffness and higher stress in the orbital region of the former when biting with carnassial teeth, suggesting that Chinese Crocuta could not process prey with the same degree of efficiency as extant Crocuta crocuta. Dental microwear texture data further support this interpretation, as Chinese Crocuta have intermediate and indistinguishable complexity values (indicative of hard-object feeding) between the extant African lion (Panthera leo) and extant hyenas (C. crocuta, Hyaena hyaena, and Parahyaena brunnea), being most similar to the omnivorous P. brunnea. The use of dental macrowear to infer dietary behavior may also be possible in extinct taxa, as evinced by dietary correlations between extant African feliforms and dental macrowear assignments. Collectively, this multiproxy analysis suggests that Chinese Crocuta may have exhibited dietary behavior distinct from that of living C. crocuta, and assumptions of niche conservatism may mask significant dietary variation in species broadly distributed in time and space.

Fossil musk turtles (Kinosternidae, Sternotherus) from the late Miocene–early Pliocene (Hemphillian) of Tennessee and Florida

ABSTRACT The oldest fossil musk turtles, genus Sternotherus, are reported from the Hemphillian of eastern Tennessee and central Florida. Sternotherus palaeodorus, n. sp., is known from five partial shells discovered at the late Miocene—early Pliocene (7–4.5 Ma; late Hemphillian) Gray Fossil Site in Washington County, Tennessee. Sternotherus palaeodorus possesses an enlarged intergular scute, wide first vertebral scute that overlaps peripheral set 1, posteriorly extensive hypoplastron to peripheral 7 contact, and a posteriorly situated inguinal musk duct pore (characteristics more typically seen in Kinosternon). A cladistic analysis recovered S. palaeodorus within crown group Sternotherus in the strict consensus and on the stem of Sternotherus in the majority rule consensus. Sternotherus bonevalleyensis, n. sp., from the Palmetto Fauna (5.5–5 Ma; late Hemphillian) of central Florida was perhaps contemporaneous with S. palaeodorus and is known only from isolated shell fragments. It is morphologically most similar to the Sternotherus minor complex and Sternotherus depressus. Subsequent Blancan fossils from the Suwannee River of Florida represent aff. S. minor peltifer. Additionally, a fragmentary left hyoplastron of cf. Sternotherus from Haile 19A, Alachua County, Florida, could be the oldest record for the genus (ca. 9–8.5 Ma; early Hemphillian). These accounts reveal that Sternotherus was diverse and moderately well distributed geographically by its first known fossil occurrences and support previous hypotheses that the Sternotherus minor complex evolved in the Gulf coastal plain and dispersed throughout that region since at least the latest Miocene-earliest Pliocene.