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Pollen and carbon isotope data from Natural Trap Cave provides insights into the environmental conditions of the Bighorn Basin, in north-central Wyoming, USA during the time intervals 151.1 to 132.2 cal ka BP, 51.8 to 17.4 cal ka BP, and 10.4 cal ka BP to present. Similarities between the buildup of the penultimate and last glacial maxima include abundant Artemisia, Amaranthaceae, Asteraceae, and Poaceae pollen, which, when combined with high δ13C values (>−25.0‰) and low plant discrimination values (18.0–19.0‰) from sedimentary organic matter, indicate steppe-like, cold-dry conditions. Between 151 and 142.5 cal ka BP, abundant Poaceae associated with much lower δ13C values (−28.0‰ to −26.0‰) and higher plant discrimination values (19.0‰–21.0‰) could be indicative of relatively wet summers. Similarly, after 25 cal ka BP increases in Poaceae may indicate wet conditions during the growing season. Holocene conditions differ from the earlier sections of the record, with greater arboreal pollen abundance (i.e., Pinus), consistent with woodland expansion in the region. Carbon isotope data from sedimentary organic matter show the Holocene was more humid than the glacial conditions of the late Pleistocene. This is especially clear at the start of the Holocene when a large negative carbon isotope excursion (CIE

Limited numbers of high-resolution records predate the Last Glacial Maximum (LGM) making it difficult to quantify the impacts of environmental changes prior to peak glaciation. We examined sediments from Last Canyon Cave in the Pryor Mountains of Montana and Wyoming to construct a >45 ka environmental record from pollen and stable isotope analysis. Artemisia pollen was hyper-abundant at the beginning of the record. Carbon isotope values of bulk organic matter (>40 ka) showed little variation (-25.3 ± 0.4‰) and were consistent with a arid C3 environment, similar to today. After 40 cal ka BP, Artemisia pollen decreased as herbaceous taxa increased toward the LGM. A significant decrease in δ13C values from 40–30 cal ka BP (~1.0‰) established a new baseline (-26.6 ± 0.2‰), suggesting cooler, seasonally wetter conditions prior to the LGM. These conditions persisted until variation in δ13C values increased significantly with post-glacial warming, marked by two spikes in values at 14.4 (-25.2‰) and 13.5 cal ka BP (-25.4‰) before δ13C values dropped to their lowest values (-26.9 ± 0.2‰) at the onset of the Younger Dryas (12.8 ka). These results provide insights into late Pleistocene conditions and ecological change in arid intermontane basins of the Rocky Mountains.

Interactions between midlatitude westerlies and the Pamir–Tian Shan mountains significantly impact hydroclimate patterns in Central Asia today, and they played an important role in driving Asian aridification during the Cenozoic. We show that distinct west-east hydroclimate differences were established over Central Asia during the late Oligocene (ca. 25 Ma), as recorded by stable oxygen isotopic values of soil carbonates. Our climate simulations show that these differences are present when relief of the Pamir–Tian Shan is higher than 75% of modern elevation (∼3000 m). Integrated with geological evidence, we suggest that a significant portion of the Pamir–Tian Shan orogen had reached elevations of ∼3 km and acted as a moisture barrier for the westerlies since ca. 25 Ma.

Stable oxygen isotope (δ18O) compositions from vertebrate tooth enamel are widely used as biogeochemical proxies for paleoclimate. However, the utility of enamel oxygen isotope values for environmental reconstruction varies among species. Herein, we evaluate the use of stable oxygen isotope compositions from pronghorn (Antilocapra americana Gray, 1866) enamel for reconstructing paleoclimate seasonality, an elusive but important parameter for understanding past ecosystems. We serially sampled the lower third molars of recent adult pronghorn from Wyoming for δ18O in phosphate (δ18OPO4) and compared patterns to interpolated and measured yearly variation in environmental waters as well as from sagebrush leaves, lakes, and rivers (δ18Ow). As expected, the oxygen isotope compositions of phosphate from pronghorn enamel are enriched in 18O relative to environmental waters. For a more direct comparison, we converted δ18Ow values into expected δ18OPO4* values (δ18OW‐PO4*). Pronghorn δ18OPO4 values from tooth enamel record nearly the full amplitude of seasonal variation from Wyoming δ18OW‐PO4* values. Furthermore, pronghorn enamel δ18OPO4 values are more similar to modeled δ18OW‐PO4* values from plant leaf waters than meteoric waters, suggesting that they obtain much of their water from evaporated plant waters. Collectively, our findings establish that seasonality in source water is reliably reflected in pronghorn enamel, providing the basis for exploring changes in the amplitude of seasonality of ancient climates. As a preliminary test, we sampled historical pronghorn specimens (1720 ± 100 AD), which show a mean decrease (a shift to lower values) of 1–2‰ in δ18OPO4 compared to the modern specimens. They also exhibit an increase in the δ18O amplitude, representing an increase in seasonality. We suggest that the cooler mean annual and summer temperatures typical of the 18th century, as well as enhanced periods of drought, drove differences among the modern and historical pronghorn, further establishing pronghorn enamel as excellent sources of paleoclimate proxy data., We evaluate the use of stable oxygen isotope compositions (δ18O in phosphate; δ18OPO4) from pronghorn (Antilocapra americana Gray, 1866) enamel for reconstructing paleoclimate seasonality by serially sampling lower third molars of recent adults (died 1970, 1972, 2010) from Wyoming. We found that pronghorn δ18OPO4 values from tooth enamel record nearly the full amplitude of seasonal variation from Wyoming. Collectively, our findings establish that seasonality in source water is reliably reflected in pronghorn enamel, providing the basis for exploring changes in the amplitude of seasonality of ancient climates.

Van Valkenburgh et al. challenge the conclusions of a recent study by DeSantis et al. that claimed that sabertooth cats and dire wolves did not compete for similar prey.

Near-modern ecosystems were established as a result of rapid ecological adaptation and climate change in the Late Miocene. On land, Late Miocene aridification spread in tandem with expansion of open habitats including C(4) grassland ecosystems. Proxy records for the central Andes spanning the Late Miocene cooling (LMC) show the reorganization of subtropical ecosystems and hydroclimate in South America between 15 and 35°S. Continental pedogenic carbonates preserved in Neogene basins record a general increase of δ(18)O and δ(13)C values from pre-LMC to post-LMC, most robustly occurring in the subtropics (25 to 30°S), suggesting aridification and a shift toward a more C4-plant-dominated ecosystem. These changes are closely tied to the enhancement of the Hadley circulation and moisture divergence away from the subtropics toward the Intertropical Convergence Zone as revealed by climate model simulations with prescribed sea-surface temperatures (SSTs) reflecting different magnitudes of LMC steepening of equator-to-pole temperature gradient and CO(2) decline.

The earliest cetaceans were interpreted as semi-aquatic based on the presence of thickened bones and stable oxygen isotopes in tooth enamel. However, the origin of aquatic behaviors in cetacean relatives (e.g., raoellids, anthracotheres) remains unclear. This study reconstructs the origins of aquatic behaviors based on long bone microanatomy and stable oxygen isotopes of tooth enamel in modern and extinct cetartiodactylans. Our findings are congruent with published accounts that microanatomy can be a reliable indicator of aquatic behaviors in taxa that are obligatorily aquatic, and also highlight that some "semi-aquatic" behaviors (fleeing into the water to escape predation) may have a stronger relationship to bone microanatomy than others (herbivory in near-shore aquatic settings). Bone microanatomy is best considered with other lines of information in the land-to-sea transition of cetaceans, such as stable isotopes. This study extends our understanding of the progression of skeletal phenotypes associated with habitat shifts in the relatives of cetaceans.

The tempo-spatial development of the Cenozoic Asian aridification across the Eocene-Oligocene and its controlling factors are important scientific topics in Earth Sciences, which are pertinent to regional and global tectonic and climatic events. However, sedimentary rocks preserving the record of aridification during this time from central Asia (ACA) are rare. Here we present a preliminary analysis of sedimentary facies of the lower Paleogene in the northeastern Tajik Basin, which reveals that: the lower part of the studied section is dominated by shallow marine deposits of the Paratethys, the middle part is typical of alternations of eolian dune and fluvial deposits, the upper part is represented by eolian loess-sandy loess (L&SL) like facies, and the top exhibits alternations of fluvial-lacustrine and loess like facies. Based on a chronological framework derived from preliminary magnetostratigraphy, published U-Pb dating of a volcanic ash, and regional litho-stratigraphic correlations, we determine that eolian and L&SL facies accumulated in the northeastern Tajik Basin during the Late Eocene and through most of the Oligocene. These sedimentary units indicate that semi-arid to arid environments of ACA had developed at least since the late Eocene. This initial aridification is closely linked to the westward retreat of the Paratethys that was likely driven by a combination of tectonic activity and sea level changes.

Plate tectonics and eustatic sea-level changes have fundamental effects on paleoenvironmental conditions and bio-ecological changes. The Paratethys Sea was a large marine seaway that connected the Mediterranean Neotethys Ocean with Central Asia during early Cenozoic time. Withdrawal of the Paratethys from central Asia impacted the distribution and composition of terrestrial faunas in the region and has been largely associated with changes in global sea level and climate such as cooling associated with the Eocene/Oligocene transition (EOT). Whereas the regression has been dated in the Tarim basin (China), the pattern and timing of regression in the Tajik basin, 400 km to the west, remain unresolved, precluding a test of current paleogeographic models. Here we date the Paratethys regression in Tajikistan at ca. 39 million years ago (Ma), which is several million years older than the EOT (at ca. 34 Ma) marking the greenhouse to icehouse climate transition of the Cenozoic. Our data also show a restricted, evaporitic marine environment since the middle–late Eocene and establishment of desert like environments after ca. 39 Ma. The overall stratigraphic record from the Tajik basin and southern Tien Shan points to deposition in a foreland basin setting by ca. 40 Ma in response to active tectonic growth of the Pamir–Tibet Mountains at the same time. Combined with the northwestward younging trend of the regression in the region, the Tajik basin record is consistent with northward growth of the Pamir and suggests significant tectonic control on Paratethys regression and paleoenvironmental changes in Central Asia.

Pinnipeds (seals, sea lions, and walruses) show variation in tooth morphology that relates to ecology. However, crown size and spacing are two aspects of morphology that have not been quantified in prior studies. We measured these characters for nearly all extant pinnipeds and three fossil taxa and then determined the principal sources of variation in tooth size and spacing using principal components (PCAs) and hierarchical cluster analysis (HCA). PCA and HCA showed that species sorted into three groups: taxa with small crowns and large diastemata, taxa with large crowns and small diastemata, and taxa that fell between these two extremes. We then performed discriminant function analysis (DFA) to determine if tooth morphology correlated with foraging strategy or diet. DFA results indicated weak correlation with diet, and stronger correlation with prey capture strategies. Tooth size and spacing were most strongly correlated with the importance of teeth in prey acquisition, with tooth size decreasing and tooth spacing increasing as teeth become less necessary in capturing food items. Taxa which relied on teeth for filtering prey from the water column or processing larger or tougher food items generally had larger crowns and smaller tooth spacing then taxa which swallowed prey whole. We found the fossil taxa Desmatophoca and Enaliarctos were most similar in tooth morphology to extant otariids, suggesting that both taxa were generalist feeders. This study established the relationship between tooth size and feeding behavior, and provides a new tool to explore the paleoecology of fossil pinnipeds and other aquatic tetrapods.

Cope's rule describes the evolutionary trend for animal lineages to increase in body size over time. In this study, we tested the validity of Cope's rule for a marine mammal clade, the Pinnipedimorpha, which includes the extinct Desmatophocidae, and extant Phocidae (earless seals), Otariidae (fur seals and sea lions), and Odobenidae (walruses). We tested for the presence of Cope's rule by compiling a large dataset of body size data for extant and fossil pinnipeds and then examined how body size evolved through time. We found that there was a positive relationship between geologic age and body size. However, this trend is the result of differences between early assemblages of small-bodied pinnipeds (Oligocene to early Miocene) and later assemblages (middle Miocene to Pliocene) for which species exhibited greater size diversity. No significant differences were found between the number of increases or decreases in body size within Pinnipedimorpha or within specific pinniped clades. This suggests that the pinniped body size increase was driven by passive diversification into vacant niche space, with the common ancestor of Pinnipedimorpha occurring near the minimum adult body size possible for a marine mammal. Based upon the above results, the evolutionary history of pinnipeds does not follow Cope's rule.

Fur seals and sea lions (Carnivora: Otariidae) evolved in the North Pacific and later dispersed throughout the Southern Hemisphere. However, the timing and number of dispersals into the Southern Hemisphere still remain poorly understood. To determine the biogeographical patterns of dispersal within fur seals and sea lions, we conducted cladistic analyses using combined evidence incorporating morphological and molecular data. The phylogeny produced in this study was then incorporated into Bayesian biogeographical analyses to reconstruct ancestral points of origin and dispersal patterns for otariid clades. Combined evidence analyses supported Callorhinus as the earliest diverging extant otariid, and a strongly supported northern sea lion clade (Zalophus, Eumetopias, and Proterozetes) as the sister group to a southern clade comprising the remainder of Otariidae. Fossil data constrained the timing and location of this dispersal as occurring between 6 and 7 Mya during a period of unusually cool sea surface temperatures and high productivity in the eastern equatorial Pacific, far older than suggested by prior studies. Our study indicates that the distribution of fur seals and sea lions is tightly linked to sea surface temperature and productivity, and suggests that otariids may be vulnerable to future anthropogenic climate change. © 2014 The Linnean Society of London

Body size plays an important role in pinniped ecology and life history. However, body size data is often absent for historical, archaeological, and fossil specimens. To estimate the body size of pinnipeds (seals, sea lions, and walruses) for today and the past, we used 14 commonly preserved cranial measurements to develop sets of single variable and multivariate predictive equations for pinniped body mass and total length. Principal components analysis (PCA) was used to test whether separate family specific regressions were more appropriate than single predictive equations for Pinnipedia. The influence of phylogeny was tested with phylogenetic independent contrasts (PIC). The accuracy of these regressions was then assessed using a combination of coefficient of determination, percent prediction error, and standard error of estimation. Three different methods of multivariate analysis were examined: bidirectional stepwise model selection using Akaike information criteria; all-subsets model selection using Bayesian information criteria (BIC); and partial least squares regression. The PCA showed clear discrimination between Otariidae (fur seals and sea lions) and Phocidae (earless seals) for the 14 measurements, indicating the need for family-specific regression equations. The PIC analysis found that phylogeny had a minor influence on relationship between morphological variables and body size. The regressions for total length were more accurate than those for body mass, and equations specific to Otariidae were more accurate than those for Phocidae. Of the three multivariate methods, the all-subsets approach required the fewest number of variables to estimate body size accurately. We then used the single variable predictive equations and the all-subsets approach to estimate the body size of two recently extinct pinniped taxa, the Caribbean monk seal (Monachus tropicalis) and the Japanese sea lion (Zalophus japonicus). Body size estimates using single variable regressions generally under or over-estimated body size; however, the all-subset regression produced body size estimates that were close to historically recorded body length for these two species. This indicates that the all-subset regression equations developed in this study can estimate body size accurately.

This study contributes to the uplift history of the Andes, which has received increasing attention in recent years because of its implications for geodynamic models and climate feedbacks. Shortening resulting in crustal thickening and removal of gravitationally unstable mantle lithosphere has been proposed to control deformation and uplift of Cordillera-type orogenic systems such as the Puna Plateau of the central Andes and its eastern margin, the Eastern Cordillera. We present new clumped isotope (Δ47), δ18O, and δ2Η data from carbonate nodules, marlstone, spring deposits, and volcanic ashes from the Puna Plateau and Eastern Cordillera of NW Argentina. When combined with other geological evidence, our data indicate that the Puna Plateau was near its present elevation since at least ~10 Ma, whereas the Eastern Cordillera rose ~1.5 km between ~14 and ~7 Ma. This history of uplift correlates with active shortening in the Eastern Cordillera and with incorporation of a regional foreland into the propagating orogenic wedge. Our study suggests that the elevation of the Puna Plateau changed little during the Miocene-Pliocene, whereas the margin experienced significant uplift associated with active deformation and crustal thickening.

The first appearance and evolution of the crown group Cetacea – Mysticeti (baleen whales) and Odontoceti (toothed whales and dolphins) – from archaeocete ancestors corresponds with major climatic and oceanographic changes shortly before the Eocene–Oligocene boundary (ca. 34.0 Ma). These environmental changes impacted marine productivity and may have sparked the evolution of the distinct bulk-feeding strategy used by mysticete whales today. The movements and feeding habits of living cetaceans can be tracked using the isotopic composition of the structural carbonate within bioapatite of their bones and teeth, which records latitudinal gradients in marine δ13C and δ18O values. Here, we exploit this relationship to determine the past movements and feeding habits of late-surviving archaeocetes (kekenodontids) and early odontocetes and mysticetes sampled from Oligocene fossil sites in the USA (latitude: ~ 30°N) and New Zealand (latitude: ~ 50°S). Bioapatite δ13C and δ18O values for fossilized tooth enamel and bullae from fossil cetaceans were surprisingly similar to those for living cetaceans, which indicates that the feeding habits of these species and the isotopic composition of Oligocene seawater (ca. 28 Ma) were very close to present-day conditions. Values for toothed mysticetes and odontocetes match expectations based on cetacean species feeding at these latitudes today, suggesting these individuals were residents that foraged in close proximity to where their fossils were discovered. In contrast, the extremely low δ13C and δ18O values for some edentulous mysticetes from New Zealand are more similar to values observed in extant mysticetes that seasonally migrate to high latitudes, where they bulk-feed on lipid-rich zooplankton. Enamel δ13C values for kekenodontids showed the greatest range and overlapped with both edentulous mysticetes and resident species; these differences correlate most strongly with tooth type and may reflect an ontogenetic shift in diet associated with the transition from nursing to marine prey. When combined with evidence of increasing morphological disparity, fossil bioapatite δ13C and δ18O values support interpretations that the Oligocene was an interval of heightened taxonomic and ecological diversification within Cetacea.

Uplift of the Central Andes is largely thought to have occurred during the past 10 m.y. based on paleoaltimetry studies from the Altiplano of Bolivia. However, the spatio-temporal uplift history may not be uniform across the Central Andes. We present new stable isotopic results from the Salar de Antofalla, Salina del Fraile, and Arizaro Basin on the Puna Plateau (24°–26°S) of northwestern Argentina. Samples of volcanic glass give δD paleowater values and modeled paleoelevations that indicate an elevated (∼4 km) Puna Plateau since ca. 36 Ma with limited (

Carbon isotope records from organic matter provide a key data set for constraining past fluctuations in the isotopic composition of atmospheric carbon dioxide (δ13Catm), hydrologic conditions, and plant community composition. Here we evaluate spatiotemporal shifts in these three conditions using δ13C values of bulk organic carbon (δ13Cbulk) from Paleocene strata in northwest Washington, U.S.A. and west-central Alberta, Canada. Our data set contains 352 analyses from the Chuckanut and Paskapoo formations. Mean δ13Cbulk values in the two areas are statistically indistinguishable (− 24.5‰ and − 24.4‰ (VPDB), respectively), and exhibit a wide range of values (~ 5–7‰). In both areas sequential vertical samples show large fluctuations in δ13Cbulk values, on average of ~ 1.0‰ ± 1.0 (1σ). There are no distinct up-section trends or structured excursions in the ~ 850 meter stratigraphic section in the Chuckanut Formation nor the ~ 125 meter section in the Paskapoo Formation. Furthermore, δ13Cbulk values are independent of the weight percent organic carbon (%C) and lithology of the sample. The lack of a systematic relationship and correlation between δ13Cbulk and %C values indicates isotopic fractionation during post-mortem processes, such as microbial respiration, exerted minimal influence on the isotopic patterns. The independence of δ13Cbulk values from lithology suggests variability is unrelated to local spatial gradients in vegetation community and water availability across the paleo-landscape. However, the wide range and large fluctuations in δ13Cbulk values is similar to stochastic intra-site variability of plant δ13C values within modern settings. The absence of structured, up-section trends indicates minimal secular variation in the carbon isotopic composition of atmospheric CO2, rainfall, and plant community composition. Moreover, we show that a previous carbon isotope excursion ascribed to the Paleocene–Eocene Thermal Maximum in the Paskapoo Formation is actually a result of aliasing related to sampling density. Finally, the statistically indistinguishable mean δ13Cbulk values in the two regions imply similarly wet and humid climatic conditions when combined with sedimentologic and paleobotanical data. The records support early Paleogene circulation models that show strong orographic precipitation patterns in the Pacific Northwest associated with incipient extensional tectonics in the region and enhanced transport of moisture-laden polar air masses southward into the Canadian continental interior.

The relative concentrations of strontium to calcium (Sr/Ca) and barium to calcium (Ba/Ca) in mammalian bioapatite are common biogeochemical indicators for trophic level and/or dietary preferences in terrestrial foodwebs; however, similar research in marine foodwebs is lacking. This study combined environmental and biological Sr/Ca and Ba/Ca data from both terrestrial and marine settings from 62 published books, reports, and studies along with original data collected from 149 marine mammals (30 species) and 83 prey items (18 species) and found that variations in Sr/Ca and Ba/Ca ratios of biological and environmental samples are appreciably different in terrestrial and marine systems. In terrestrial systems, environmental sources account for most of the variations in Sr/Ca and Ba/Ca ratios. In contrast, environmental sources in marine systems (i.e., seawater) are comparatively invariant, meaning most of the variations in Sr/Ca and Ba/Ca ratios originate from biological processes. Marine consumers, particularly non-mammalian and mammalian vertebrates, show evidence of biopurification of Ca relative to Sr and Ba, similar to what is observed in terrestrial systems; however, unlike terrestrial systems, variations in Sr/Ca and Ba/Ca ratios of environmental sources are overprinted by bioaccumulation of Sr and Ba at the base of marine foodwebs. This demonstrates that in marine systems, spatial or temporal differences may have little to no effect on Sr/Ca and Ba/Ca ratios of marine vertebrates, making Sr/Ca, and to a lesser extent Ba/Ca, potentially useful global proxies for trophic level and dietary preferences of marine vertebrates.

Sr/Ca and Ba/Ca ratios of bone are commonly used as biochemical indicators of trophic level in modern and fossil mammals. Concerns over the effects of diagenesis on Sr/Ca and Ba/Ca ratios of bone led archaeologists and paleontologists to favor tooth enamel, which is less prone to alteration. Sr/Ca and Ba/Ca ratios of bone, enamel, and dentin from three farm-raised steers (Bos taurus) and five wild white-tailed deer (Odocoileus virginianus) from central Missouri were compared. Our results show that changes in diet, discrimination, and growth rate during ontogeny can lead to significant differences in Sr/Ca and Ba/Ca ratios of different bioapatite types as well as significant differences within the same bioapatite forming at different times. Early- and late-forming tooth enamel can have significant differences in Sr/Ca and Ba/Ca ratios equivalent to almost one full trophic step. Although differences between early- and late-forming dentin are typically not significant, dentin Sr/Ca and Ba/Ca ratios are significantly greater than enamel values. This difference in Sr/Ca or Ba/Ca ratios between enamel and dentin from the same tooth can be greater than one full trophic step. These results have profound implications for the use of dental bioapatites in trophic level reconstructions. They highlight the importance of consistency in bioapatite selection, tooth selection, and relative location of sampling within the enamel cap. Furthermore, this expected difference in Sr/Ca and Ba/Ca ratios could be used as another means of checking for diagenetic alteration in ancient samples.

Stable isotope analysis of fossil materials has become an increasingly important method for gathering dietary and environmental information from extinct species in terrestrial and aquatic ecosystems. The benefits of these analyses stem from the geochemical fingerprint that an animal’s environment leaves in its bones, teeth, and tissues. Ongoing study of living mammals has found the stable isotopic composition of several light (hydrogen, carbon, nitrogen, oxygen, and sulfur) and even a few heavy (calcium and strontium) elements to be useful tracers of ecological and physiological information; many of these can be similarly applied to the study of fossil mammals. For instance, the carbon isotopic composition of an animal’s tissues tracks that of the food it eats, whereas the oxygen isotopic compositions of the carbonate and phosphate in an animal’s bones and teeth are primarily controlled by that of the surface water it drinks or the water in the food it ingests. These stable isotope proxies for diet and habitat information are independent of inferences based on morphological characters and thus provide a means of testing ecological interpretations drawn from the fossil record. As such, when well-preserved specimens are available, any dietary study of fossil species should seriously consider including this approach. To illustrate the potential benefits associated with applying these methods to paleontological research, a review of current work on the ecological and evolutionary history of fossil mammals through geochemical analysis is presented. After a brief introduction to issues associated with the preservation of stable isotopic information in soft and mineralized tissues, a series of case studies involving the application of stable isotope analysis to fossil mammal research is discussed. These studies were selected to highlight the versatility of this analytical method to paleontological research and are complemented by a discussion of new techniques and instrumentation in stable isotope analysis (e.g., laser ablation and compound-specific isotope ratio mass spectrometry, and calcium and clumped isotopes), which represent the latest advances in the extension of these geochemical tools to the paleontology of fossil mammals.

Starch grains are a ubiquitous component of plants that have been used in tandem with phytoliths, pollen, and macrofossils to reconstruct past floral diversity. This tool has yet to be fully explored for aquatic plants, specifically seagrasses, which lack phytoliths and are rarely preserved as macrofossils or pollen. If starch grains in seagrasses are morphologically distinct, this method has the potential to improve seagrass identification in the fossil record in such cases where its starch is preserved (e.g. scratches and occlusal surfaces of tooth enamel from seagrass consumers). The goals of this study were twofold: (1) to determine if starch is present in seagrass material and (2) to assess how starch grain morphology differs between different seagrasses. This study focused on four abundant and ecologically distinct seagrasses from the Caribbean: Halodule wrightii , Ruppia maritima , Syringodium filiforme , and Thalassia testudinum . Starch grains were observed in all species except S. filiforme . Grains from H. wrightii are typically observed in side-on orientation, are sub-round to angular, and are fairly small (3-19 μm, end-on). Grains of R. maritima are small spherical grains (4–8 μm) that have a centric hilum and a straight extinction cross with a median angle between the arms of 90°. Grains from T. testudinum are large (9–31 μm, end-on), conical in side-on and round/sub-round in end-on orientation, have a slightly eccentric hilum with an obvious particle, and prominent lamellae. Visual assessment and comparative statistics demonstrate that the morphology of starch grains from T. testudinum , R. maritima , and H. wrightii are significantly different. With more extensive research, there is potential for the positive identification of starch grains from an unknown seagrass. The ability to identify seagrass from starch grains could facilitate the identification of seagrasses in the fossil record and supply information on seagrass evolution and distribution, climate effects on seagrass distribution, and the diets of seagrass consumers.

The Eocene greenhouse climate state has been linked to a more vigorous hydrologic cycle at mid- and high latitudes; similar information on precipitation levels at low latitudes is, however, limited. Oxygen isotopic fluxes track moisture fluxes and, thus, the δ(18)O values of ocean surface waters can provide insight into hydrologic cycle changes. The offset between tropical δ(18)O values from sampled Eocene sirenian tooth enamel and modern surface waters is greater than the expected 1.0 per mil increase due to increased continental ice volume. This increased offset could result from suppression of surface-water δ(18)O values by a tropical, annual moisture balance substantially wetter than that of today. Results from an atmospheric general circulation model support this interpretation and suggest that Eocene low latitudes were extremely wet.

The timing of initiation of continent-continent collision between Asia and India is controversial, but this major tectonic event is generally thought to have occurred in the Early Eocene, ca. 50 Ma. New and independent data from strontium isotopes, stable carbon isotopes, microfossil biostratigraphy, and mammal fossils from an Early Eocene marginal marine sequence (Cambay Shale) at the Vastan Lignite Mine of western India indicate that terrestrial faunal exchanges, and therefore continental collision, between Asia and the Indian subcontinent took place before 53.7 Ma. This age coincides with the second Eocene Thermal Maximum (ETM2), a short-lived warming episode that followed the Paleocene-Eocene Thermal Maximum (PETM) ca. 55.5 Ma. Our data also document, for the first time, a clear record of the ETM2 in terrestrial organic material from a low-latitude site, which is represented by a 3‰−4‰ carbon isotope excursion (CIE) in lignite and dispersed organic carbon δ13C values. The magnitude of the CIE at this location closely matches that observed in marine cores from the Arctic Ocean, which supports an interpretation that this hyperthermal event, though of lower magnitude, was similar in character to that of the PETM, being a global phenomenon that affected both terrestrial and marine ecosystems.

Dental morphology changes dramatically across the artiodactyl-cetacean transition, and it is generally assumed that this reflects the evolutionary change from herbivory and omnivory to carnivory. To test hypotheses regarding tooth function and diet, we studied size and position of wear facets on the lower molars and the stable isotopes of enamel samples. We found that nearly all investigated Eocene cetaceans had dental wear different from typical wear in ungulates and isotope values indicating that they hunted similar prey and processed it similarly. The only exception is the protocetid Babiacetus, which probably ate larger prey with harder skeletons. The closest relative of cetaceans, the raoellid artiodactyl Indohyus, had wear facets that resemble those of Eocene cetaceans more than they do facets of basal artiodactyls. This is in spite of Indohyus's tooth crown morphology, which is unlike that of cetaceans, and its herbivorous diet, as indicated by stable isotopes. This implies that the evolution of masticatory function preceded that of crown morphology and diet at the origin of cetaceans.

This study evaluates the effect of climate on facies, grain size, and sedimentation rates using sedimentology, geochronology, and stable isotope geochemistry for Miocene–Pliocene deposits in the Angastaco basin (Eastern Cordillera, northwest Argentina). U-Pb zircon data from ash layers constrain the transition between the finer grained fluvial-lacustrine Palo Pintado Formation and the coarser grained fluvial-alluvial San Felipe Formation to ca. 5.2 Ma and the first deposition of sediment derived from the present-day orographic barrier to ca. 4 Ma. δ 13 C values from pedogenic carbonate nodules range from −15.4‰ to −10.2‰ for the Palo Pintado Formation and from −9.5‰ to −8.2‰ for the San Felipe Formation; this can be best explained by increased, sustained aridity since ca. 5 Ma. The δ 18 O values range from −9.6‰ to −5.9‰ for the Palo Pintado Formation and from −6.1‰ to −5.2‰ for the San Felipe Formation, corroborating this interpretation. The shift toward more arid conditions correlates with a significant increase in grain size but no significant change in sedimentation rate. Because aridity precedes the development of an orographic effect, we interpret the grain size increase in the Angastaco basin since ca. 5 Ma to be a response of the sedimentary system to aridification resulting from regional climate change.

Isotopic studies of palaeoecological and ecological questions often use bone collagen or bioapatite as substrates, but rarely both. Substantial new information can be gained from the incorporation of isotopic values from both the organic and inorganic fractions of bone. Here we show that combining isotopic data from both substrates provides valuable and unique insights into (1) trophic relationships and dietary interactions; (2) differences in digestive physiologies and (3) identification of palaeontological or archaeological remains that lack diagnostic morphological characters. We present a range of new isotopic data collected from modern and fossil mammals, and investigate patterns within several published datasets. We define carbon isotope spacing variables, and then explore four diverse palaeoecological and ecological case studies. Copyright © 2009 John Wiley & Sons, Ltd.

Sirenians (sea cows: manatees and dugongs) have been primary consumers in tropical and subtropical shallow-water marine ecosystems for more than 50 m.y. Though fossils of the earliest sirenians have been recovered from the Caribbean, sirenians are thought to have originated in the Tethys-Mediterranean region, which underwent significant climate change over the course of the Cenozoic. Application of stable isotope analysis can provide new insight into how diet and habitat preferences were affected by climate change and how these ecological changes correlate with diversity patterns for this region. Tooth enamel was sampled from 57 specimens of sirenians from the Tethys-Mediterranean and adjacent seas spanning the Eocene to Pliocene. Enamel δ13C values across this time interval were consistently high and indicative of diets composed mostly of marine seagrasses. In contrast, enamel δ18O values showed significant change over time and across the region. Sirenians from southern sites (i.e., Egypt, Libya) had significantly higher mean δ18O values than specimens from most European locations, suggesting variation in salinity within the Tethys-Mediterranean. The range in enamel δ18O values increased from the Paleogene through the Neogene, but enamel δ13C values remained steady, indicating that changes in environmental conditions did not necessarily coincide with major changes in dietary preferences. Declines in sirenian taxonomic diversity in the late Miocene were likely due to the rapid climate and oceanographic changes during the latter half of the Cenozoic and their impacts on available dietary resources. Elevated δ13C and δ18O values in Mediterranean sirenians during the Messinian salinity crisis corroborate the hypothesis of ecophenotypic dwarfing in these animals.

Large-bodied, semiaquatic herbivorous mammals have been a recurring component of most continental ecosystems throughout the Cenozoic. Identification of these species in the fossil record has largely been based on the morphological similarities with present-day hippopotamids, leading to the designation of this pairing of body type and ecological niche as the hippo ecomorph. These morphological characters, however, may not always be diagnostic of aquatic habits. Here, enamel δ13C and δ18O values from living hippopotamuses were examined to define an isotopic signature unique to the hippo ecomorph. Although δ13C values do not support unique foraging habits for this ecomorph, living and fossil hippopotamids typically have low mean δ18O values relative to associated ungulates that fit a linear regression (δ18Ohippopotamids = 0.96 ± 0.09·δ18Ofauna − 1.67 ± 2.97; r2 = 0.886, p < 0.001). Modeling of oxygen fluxes in large mammals suggests that high water-turnover rates or increased water loss through fece...

Although the first ten million years of whale evolution are documented by a remarkable series of fossil skeletons, the link to the ancestor of cetaceans has been missing. It was known that whales are related to even-toed ungulates (artiodactyls), but until now no artiodactyls were morphologically close to early whales. Here we show that the Eocene south Asian raoellid artiodactyls are the sister group to whales. The raoellid Indohyus is similar to whales, and unlike other artiodactyls, in the structure of its ears and premolars, in the density of its limb bones and in the stable-oxygen-isotope composition of its teeth. We also show that a major dietary change occurred during the transition from artiodactyls to whales and that raoellids were aquatic waders. This indicates that aquatic life in this lineage occurred before the origin of the order Cetacea.

Carbon isotope differences (Δ13C) between bioapatite and diet, collagen and diet, and bioapatite and collagen were calculated for four species of sirenians, Dugong dugon (Muller), Trichechus manatus (Linnaeus), Trichechus inunguis (Natterer), and the extinct Hydrodamalis gigas (Zimmerman). Bone and tooth samples were taken from archived materials collected from populations during the mid eighteenth century (H. gigas), between 1978 and 1984 (T. manatus, T. inunguis), and between 1997 and 1999 (D. dugon). Mean Δ13C values were compared with those for terrestrial ungulates, carnivores, and six species of carnivorous marine mammals (cetaceans = 1; pinnipeds = 4; mustelids = 1). Significant differences in mean δ13C values among species for all tissue types were detected that separated species or populations foraging on freshwater plants or attached marine macroalgae (δ13C values −4‰; Δ13Cbioapatite–diet ∼11‰). Likewise, Δ13Cbioapatite–collagen values for freshwater and algal-foraging species (∼7‰) were greater than those for seagrass-foraging species (∼5‰). Variation in Δ13C values calculated between tissues and between tissues and diet among species may relate to the nutritional composition of a species’ diet and the extent and type of microbial fermentation that occurs during digestion of different types of plants. These results highlight the complications that can arise when making dietary interpretations without having first determined species-specific Δ13Ctissue–diet values.

The development of pierce-feeding and loss of oral processing represented major adaptations for underwater feeding in marine mammals. We examined the evolution of pierce-feeding and its association with changes in tooth spacing and tooth size to determine whether pierce-feeding was practiced by the earliest known pinnipeds. Data on crown size and spacing in postcanine dentition were collected and 1) analysed by principal components analysis (PCA) to determine the tooth morphospace of arctoid carnivores, 2) analysed by least squares (LS) regression and phylogenetic independent contrasts (PIC) to determine what morphological variables were associated with increases in tooth spacing, and 3) used to reconstruct the evolution of feeding related traits within a phylogenetic context. The PCA analysis revealed that within arctoid carnivores, the greatest differences in morphospace were associated with pierce-feeding, and the early-diverging seal Enaliarctos was placed within the pinniped morphospace. Increased tooth spacing within Pinnipedia is a result of decreased postcanine crown size. When the evolution of dental characters is reconstructed, 'enaliarctines' were found to represent an intermediate stage in evolution between 'fissiped' and pinniped carnivores. They retained the limited tooth spacing of terrestrial carnivores, possessed postcanine crown lengths intermediate in size between pinnipeds and fissipeds, and possessed reduced heterodonty characteristic of crown pinnipeds. Our study indicated that pierce-feeding evolved early within pinnipeds. This suggested either that pierce-feeding evolved prior to the loss of mastication, or that pierce-feeding evolved at the same time as loss of mastication, and well before simplification of the dentition was completed.

Recent fossil discoveries of early cetaceans and sirenians document the functional transitions that occurred as each group adapted to a completely aquatic existence, but the timing and path of their ecological transition remain uncertain. We analyzed the stable-isotope composition of tooth enamel from several early members of each group to reconstruct the dietary, foraging, and habitat preferences of basal taxa. Carbon isotope (δ13C) values provided evidence of foraging within freshwater, terrestrial, and marine environments, including seagrass beds, whereas oxygen isotope (δ18O) variation was used as a measure of commitment to aquatic habitats. Enamel samples were collected from four regions (south Asia, north and west Africa, and southern Europe) spanning the late early Eocene (ca. 53.5 Ma) to the late Eocene (ca. 36 Ma). Sirenian and cetacean taxa included species that were likely capable of some terrestrial locomotion, and more specialized forms that were morphologically fully aquatic. Cetacean δ13C and δ18O values indicate that some early members of this group (some pakicetids) inhabited freshwater environments, but later members (e.g., remingtonocetids, protocetids, and basilosaurids) moved quickly into estuarine and marine environments. Low δ18O variation confirms that all of these early forms were primarily aquatic, but δ13C and δ18O values for early sirenians indicate an early transition to a marine seagrass diet without any evidence of an intermediate connection to freshwater habitats.

Cenozoic sediments of Florida contain one of the most highly fossiliferous sequences of extinct sirenians in the world. Sirenians first occur in Florida during the Eocene (ca. 40 Ma), have their peak diversity during the late Oligocene-Miocene (including the widespread dugongid Me- taxytherium), and become virtually extinct by the late Miocene (ca. 8 Ma). Thereafter during the Pliocene and Pleistocene, sirenians are represented in Florida by abundant remains of fossil man- atees (Trichechus sp.). Stable isotopic analyses were performed on 100 teeth of fossil sirenians and extant Trichechus manatus from Florida in order to reconstruct diets (as determined from d 13C val- ues) and habitat preferences (as determined from d18O values) and test previous hypotheses based on morphological characters and associated floral and faunal remains. A small sample (n 5 6) of extant Dugong dugon from Australia was also analyzed as an extant model to interpret the ecology of fossil dugongs. A pilot study of captive manatees and their known diet revealed an isotopic enrichment («*) in d13C of 14.0‰, indistinguishable from previously reported «* for extant medium to large terrestrial mammalian herbivores with known diets. The variation in d18OV-SMOW reported here is interpreted to indicate habitat preferences, with depleted tooth enamel values ( 25‰) representing freshwater rivers and springs, whereas enriched values ( 30‰) indicate coastal marine environments. Taken together, the Eocene to late Miocene sirenians (Protosirenidae and Dugongidae) differ significantly in both d 13 Ca ndd18O from Pleistocene and Recent manatees (Trichechidae). In general, Protosiren and the fossil dugongs from Florida have carbon isotopic values that are relatively positive (mean d13C 52 0.9‰) ranging from 24.8‰ to 5.6‰, interpreted to represent a specialized diet of pre- dominantly seagrasses. The oxygen isotopic values (mean d18O 5 29.2‰) are likewise relatively positive, indicating a principally marine habitat preference. These interpretations correlate well with previous hypotheses based on morphology (e.g., degree of rostral deflection) and the known ecology of modern Dugong dugon from the Pacific Ocean. In contrast, the fossil and extant Trichechus teeth from Florida have relatively lower carbon isotopic values (mean d 13C 52 7.2‰) that range from 218.2‰ to 1.7‰, interpreted as a more generalized diet ranging from C3 plants to seagrasses. The relatively lower oxygen isotopic values (mean d18O 5 28.1‰) are interpreted as a more diverse array of freshwater and marine habitat preferences than that of Protosiren and fossil dugongs. This study of Cenozoic sirenians from Florida further demonstrates that stable isotopes can test hy- potheses previously based on morphology and associated floral and faunal remains. All these data sets taken together result in a more insightful approach to reconstructing the paleobiology of this interesting group of ancient aquatic mammalian herbivores.

The Desmostylia, an extinct order of mammals related to sirenians and proboscideans, are known from the late Oligocene to late Miocene of the North Pacific. Though often categorized as marine mammals on the basis of fossil occurrences in nearshore deposits, reconstructions of desmostylian habitat and dietary preferences have been somewhat speculative because morpho- logical and sedimentological information is limited. We analyzed the carbon, oxygen, and stron- tium isotope compositions of enamel from Desmostylus and co-occurring terrestrial and marine taxa from middle Miocene sites in California to address the debate surrounding desmostylian ecol- ogy. The d 13C value of tooth enamel can be used as a proxy for diet. Desmostylus had much higher d13C values than coeval terrestrial or marine mammals, suggesting a unique diet that most likely consisted of aquatic vegetation. Modern aquatic mammals tend to exhibit lower variability in d18O values than terrestrial mammals. Both fossil marine mammals and Desmostylus exhibited low d18O variability, suggesting that Desmostylus spent a large amount of time in water. Finally, the Sr isotope composition of marine organisms reflects that of the ocean and is relatively invariant when com- pared with values for animals from land. Sr isotope values for Desmostylus were similar to those for terrestrial, rather than marine, mammals, suggesting Desmostylus was spending time in estu- arine or freshwater environments. Together, isotopic data suggest that Desmostylus was an aquatic herbivore that spent a considerable portion of its life foraging in estuarine and freshwater ecosys- tems.

Recent research has shown that calcium isotopes are fractionated by metabolic processes, leading to a decrease in 44 Ca/ 40 Ca ratio with increasing trophic level. If so, calcium isotopes could provide information on trophic relationships within foodwebs millions of years older than what we have been able to study thus far with alternative methods (i.e., nitrogen isotopes (δ 15 N), Sr/Ca). To explore whether δ 44 Ca values provided marine trophic level information, we measured the δ 44 Ca composition of tooth enamel and bone from modern marine mammals representing a 2.5 order range in trophic level. Marine mammal enamel δ 44 Ca values clustered into two groups—mammals foraging on vegetation or invertebrates exhibited higher δ 44 Ca values than those foraging on fish or other marine mammals. We next examined whether this correlation was preserved in the fossil record by examining a 15 Ma marine fauna from southern California and observed that the relationship between δ 44 Ca values of specimens followed the same pattern as observed in modern faunas, but the mean δ 44 Ca values were significantly different from modern δ 44 Ca values for mammals of similar trophic level. We conclude that the relative spacing of δ 44 Ca values amongst fossil taxa can serve as a valuable tool for defining trophic level of extinct organisms and can provide critical information on relationships within ancient foodwebs. Copyright  2003 John Wiley & Sons, Ltd.

We synthesize geologic observations with new isotopic evidence for the timing and magnitude of uplift for the central Andes between 22°S and 26°S since the Paleocene. To estimate paleoelevations, we used the stable isotopic composition of carbonates and volcanic glass, combined with another paleoelevation indicator for the central Andes: the distribution of evaporites. Paleoelevation reconstruction using clumped isotope paleothermometry failed due to resetting during burial. The Andes at this latitude rose and broadened eastward in three stages during the Cenozoic. The fi rst, in what is broadly termed the “Incaic” orogeny, ended by the late Eocene, when magmatism and deformation had elevated to ≥4 km the bulk (~50%) of what is now the western and central Andes. The second stage witnessed the gradual building of the easternmost Puna and Eastern Cordillera, starting with deformation as early as 38 Ma, to >3 km by no later than 15 Ma. The proximal portions *quadej@email.arizona.edu Quade, J., Dettinger, M.P., Carrapa, B., DeCelles, P., Murray, K.E., Huntington, K.W., Cartwright, A., Canavan, R.R., Gehrels, G., and Clementz, M., 2015, The growth of the central Andes, 22°S–26°S, in DeCelles, P.G., Ducea, M.N., Carrapa, B., and Kapp, P.A., eds., Geodynamics of a Cordilleran Orogenic System: The Central Andes of Argentina and Northern Chile: Geological Society of America Memoir 212, p. 277–308, doi:10.1130/2015.1212(15). For permission to copy, contact editing@geosociety.org. © 2014 The Geological Society of America. All rights reserved. on January 15, 2015 memoirs.gsapubs.org Downloaded from

Microwear, the quantification of microscopic scratches and pits on the occlusal surfaces of tooth enamel, is a commonly used as a paleodietary proxy. For ungulates (hoofed mammals), scratch-dominant microwear distinguishes modern grazers from browsers, presumably as a result of abrasion from grass phytoliths (biogenic silica). However, it is also likely that exogenous grit (i.e., soil, dust) is a contributing factor to these scratch-dominant patterns, which may reflect soil ingestion that varies with feeding height and/or environmental conditions (e.g., dust production in open and/or arid habitats). This study assesses the contribution of exogenous grit to tooth wear by measuring the effects of fine- and medium-grained silica sand on tooth enamel using a novel live animal tooth molding technique. This study therefore constitutes the first controlled feeding experiment using ungulates and the first in vivo experiments using abrasives of different sizes. Four sheep were fed three diet treatments: 1) a mixture of Garrison and Brome hay (control), 2) hay treated with fine-grained silica sand (180–250 µm), and 3) hay treated with medium-grained silica sand (250–425 µm). We found a significant increase in pit features that was correlated with an increase in grain size of grit, corroborating earlier chewing simulation experiments that produced pits through grit-induced abrasion (i.e., the ‘grit effect’). Our results support an interpretation of large silica grains fracturing to create smaller, more abundant angular particles capable of abrasion, with jaw movement defining feature shape (i.e., scratch or pit).

Cope's rule describes the evolutionary trend for animal lineages to increase in body size over time. In this study, we tested the validity of Cope's rule for a marine mammal clade, the Pinnipedimorpha, which includes the extinct Desmatophocidae, and extant Phocidae (earless seals), Otariidae (fur seals and sea lions), and Odobenidae (walruses). We tested for the presence of Cope's rule by compiling a large dataset of body size data for extant and fossil pinnipeds and then examined how body size evolved through time. We found that there was a positive relationship between geologic age and body size. However, this trend is the result of differences between early assemblages of small-bodied pinnipeds (Oligocene to early Miocene) and later assemblages (middle Miocene to Pliocene) for which species exhibited greater size diversity. No significant differences were found between the number of increases or decreases in body size within Pinnipedimorpha or within specific pinniped clades. This suggests that the pinniped body size increase was driven by passive diversification into vacant niche space, with the common ancestor of Pinnipedimorpha occurring near the minimum adult body size possible for a marine mammal. Based upon the above results, the evolutionary history of pinnipeds does not follow Cope's rule.

Cetaceans are obligate aquatic mammals derived from terrestrial artiodactyls. The defining characteristic of cetaceans is a thick and dense lip (pachyosteosclerotic involucrum) of an ear bone (the tympanic). This unique feature is absent in modern terrestrial artiodactyls and is suggested to be important in underwater hearing. Here, we investigate the mineralogical and biochemical properties of the involucrum, as these may hold clues to the aquatic adaptations of cetaceans. We compared bioapatites (enamel, dentine, cementum, and skeletal bone) of cetaceans with those of terrestrial artiodactyls and pachyosteosclerotic ribs of manatees (Sirenia). We investigated organic, carbonate, and mineral composition as well as crystal size and crystallinity index. In all studied variables, bioapatites of the cetacean involucrum were intermediate in composition and structure between those of tooth enamel on the one hand and those of dentine, cementum, and skeletal bone on the other. We also studied the amino acid composition of the cetacean involucrum relative to that of other skeletal bone. The central involucrum had low glycine and hydroxyproline concentrations but high concentrations of nonessential amino acids, unlike most bone samples but similar to the tympanic of hippos and the (pachyosteosclerotic) ribs of manatees. These amino acid results are evidence of rapid bone development. We hypothesize that the mineralogical and amino acid composition of cetacean bullae differs from that of other bone because of (1) functional modifications for underwater sound reception and (2) structural adaptations related to rapid ossification.

Pinnipeds (seals, sea lions, and walruses) show variation in tooth morphology that relates to ecology. However, crown size and spacing are two aspects of morphology that have not been quantified in prior studies. We measured these characters for nearly all extant pinnipeds and three fossil taxa and then determined the principal sources of variation in tooth size and spacing using principal components (PCAs) and hierarchical cluster analysis (HCA). PCA and HCA showed that species sorted into three groups: taxa with small crowns and large diastemata, taxa with large crowns and small diastemata, and taxa that fell between these two extremes. We then performed discriminant function analysis (DFA) to determine if tooth morphology correlated with foraging strategy or diet. DFA results indicated weak correlation with diet, and stronger correlation with prey capture strategies. Tooth size and spacing were most strongly correlated with the importance of teeth in prey acquisition, with tooth size decreasing and tooth spacing increasing as teeth become less necessary in capturing food items. Taxa which relied on teeth for filtering prey from the water column or processing larger or tougher food items generally had larger crowns and smaller tooth spacing then taxa which swallowed prey whole. We found the fossil taxa Desmatophoca and Enaliarctos were most similar in tooth morphology to extant otariids, suggesting that both taxa were generalist feeders. This study established the relationship between tooth size and feeding behavior, and provides a new tool to explore the paleoecology of fossil pinnipeds and other aquatic tetrapods.

Anthracobunidae is an Eocene family of large mammals from south Asia that is commonly considered to be part of the radiation that gave rise to elephants (proboscideans) and sea cows (sirenians). We describe a new collection of anthracobunid fossils from Middle Eocene rocks of Indo-Pakistan that more than doubles the number of known anthracobunid fossils and challenges their putative relationships, instead implying that they are stem perissodactyls. Cranial, dental, and postcranial elements allow a revision of species and the recognition of a new anthracobunid genus. Analyses of stable isotopes and long bone geometry together suggest that most anthracobunids fed on land, but spent a considerable amount of time near water. This new evidence expands our understanding of stem perissodactyl diversity and sheds new light on perissodactyl origins.