Person:

Flynn, Lawrence

Past ecological responses of mammals to climate change are recognized in the fossil record by adaptive significance of morphological variations. To understand the role of dietary behavior on functional adaptations of dental morphology in rodent evolution, we examine evolutionary change of tooth shape in late Miocene Siwalik murine rodents, which experienced a dietary shift toward C4 diets during late Miocene ecological change indicated by carbon isotopic evidence. Geometric morphometric analysis in the outline of upper first molars captures dichotomous lineages of Siwalik murines, in agreement with phylogenetic hypotheses of previous studies (two distinct clades: the Karnimata and Progonomys clades), and indicates lineage-specific functional responses to mechanical properties of their diets. Tooth shapes of the two clades are similar at their sympatric origin but deviate from each other with decreasing overlap through time. Shape change in the Karnimata clade is associated with greater efficiency of propalinal chewing for tough diets than in the Progonomys clade. Larger body mass in Karnimata may be related to exploitation of lower-quality food items, such as grasses, than in smaller-bodied Progonomys. The functional and ecophysiological aspects of Karnimata exploiting C4 grasses are concordant with their isotopic dietary preference relative to Progonomys. Lineage-specific selection was differentially greater in Karnimata, and a faster rate of shape change toward derived Karnimata facilitated inclusion of C4 grasses in the diet. Sympatric speciation in these clades is most plausibly explained by interspecific competition on resource utilization between the two, based on comparisons of our results with the carbon isotope data. Interspecific competition with Karnimata may have suppressed morphological innovation of the Progonomys clade. Pairwise analyses of morphological and carbon isotope data can uncover ecological causes of sympatric speciation and define functional adaptations of teeth to resources.

Stable carbon isotope analysis in tooth enamel is a well-established approach to infer C3 and C4 dietary composition in fossil mammals. The bulk of past work has been conducted on large herbivorous mammals. One important finding is that their dietary habits of fossil large mammals track the late Miocene ecological shift from C3 forest and woodland to C4 savannah. However, few studies on carbon isotopes of fossil small mammals exist due to limitations imposed by the size of rodent teeth, and the isotopic ecological and dietary behaviors of small mammals to climate change remain unknown. Here we evaluate the impact of ecological change on small mammals by fine-scale comparisons of carbon isotope ratios (δ13C) with dental morphology of murine rodents, spanning 13.8 to ∼2.0 Ma, across the C3 to C4 vegetation shift in the Miocene Siwalik sequence of Pakistan. We applied in-situ laser ablation GC-IRMS to lower first molars and measured two grazing indices on upper first molars. Murine rodents yield a distinct, but related, record of past ecological conditions from large herbivorous mammals, reflecting available foods in their much smaller home ranges. In general, larger murine species show more positive δ13C values and have higher grazing indices than smaller species inhabiting the same area at any given age. Two clades of murine rodents experienced different rates of morphological change. In the faster-evolving clade, the timing and trend of morphological innovations are closely tied to consumption of C4 diet during the vegetation shift. This study provides quantitative evidence of linkages among diet, niche partitioning, and dental morphology at a more detailed level than previously possible.

The modern Asian monsoonal systems are currently believed to have originated around the end of the Oligocene following a crucial step of uplift of the Tibetan-Himalayan highlands. Although monsoon possibly drove the evolution of many mammal lineages during the Neogene, no evidence thereof has been provided so far. We examined the evolutionary history of a clade of rodents, the Rhizomyinae, in conjunction with our current knowledge of monsoon fluctuations over time. The macroevolutionary dynamics of rhizomyines were analyzed within a well-constrained phylogenetic framework coupled with biogeographic and evolutionary rate studies. The evolutionary novelties developed by these rodents were surveyed in parallel with the fluctuations of the Indian monsoon so as to evaluate synchroneity and postulate causal relationships. We showed the existence of three drops in biodiversity during the evolution of rhizomyines, all of which reflected elevated extinction rates. Our results demonstrated linkage of monsoon variations with the evolution and biogeography of rhizomyines. Paradoxically, the evolution of rhizomyines was accelerated during the phases of weakening of the monsoons, not of strengthening, most probably because at those intervals forest habitats declined, which triggered extinction and progressive specialization toward a burrowing existence.

Hedgehogs (erinaceid insectivores) are a common element in Miocene small mammal faunas of Pakistan, but little material has been formally described. Here, we report on extensive collections from numerous localities across Pakistan, most from the Potwar Plateau, Punjab, and the Sehwan area in Sind. The dominant erinaceid is Galerix, which is also known from Europe, Turkey, and East Africa. We document a new early species of Galerix, G. wesselsae, in sites from Sehwan, the Zinda Pir Dome, the Potwar Plateau, and Banda Daud Shah ranging in age from about 19 to 14 Ma. This species briefly co-occurs with and is then replaced by the smaller Galerix rutlandae, previously based on a few teeth from Daud Khel. G. rutlandae is a common element of Chinji Formation faunas through about 11.5 Ma. Scanty material of an apparently new species of Schizogalerix succeeds Galerix. After 10.2 Ma, no galericines occur in Pakistan, but rare erinaceines are later faunal components.

The Tertiary sediments of northern Pakistan are an exceptional record of terrestrial sedimentation and represent most of Neogene time. Foremost is the Siwalik Group of the Potwar Plateau, for which multiple, superposed fossil levels span ∼18–6 Ma. Well-developed magnetostratigraphic control provides secure dating so that Siwalik fossil horizons may be interpolated into a time scale with resolution to 100,000 years. We describe the geographic setting of the Potwar, give an overview of the temporal distribution of faunas, and discuss changes in paleohabitat and paleoecology with coinciding faunal change, as seen from the Siwalik viewpoint. The long Siwalik biostratigraphy of many successive assemblages with its resolved time scale may be compared directly with other well-dated sequences. Immigrant arrival and timing of faunal change may be traced. The basins of the Iberian Peninsula show somewhat different timing of introduction of hipparionine horses, and faunal turnover in the Siwaliks clearly precedes the Vallesian crisis in Spain. In contrast to the increasingly seasonal precipitation of the late Miocene Potwar, the paleohabitat of coeval North China appears to have been moist and equable, with high diversity faunas. Continued development and comparison of resolved Neogene records allow increasing resolution of the patterns of faunal change on regional to global levels.

We discuss the history of exploration of Red Clay vertebrate fossils of the Loess Plateau, and record observations on “Hipparion Red Clay” localities of Shanxi and Gansu provinces. Red clay is widespread across the Loess Plateau, but misleading as a descriptive term because many deposits are neither red nor dominated by clay-size sediment. Many red clay sequences contain paleosols, but also water-laid deposits. Our survey includes well-known localities of Yushe Basin, Renjiagou and Leijiahe near Lingtai, and sites of Qingyang and Qin’an. We discuss fossils found at Renjiagou, a new discovery of micromammals (Pliopentalagus) from Yucun, and Pliocene burrows observed at Hu Jia Cun, both latter localities northeast of Lingtai. Observations are consistent with a high rate of supply of clay sized particles, likely air-fall origin, throughout the Loess Plateau during the late Miocene. In some areas where fluvial or lake processes dominate, red clay particles are replaced in part or completely by water-laid coarse-grained deposits. We saw no evidence for a dry Late Miocene-Early Pliocene, but rather hypothesize well-watered habitat of high productivity. Locally diverse vertebrate fossils attest to high terrestrial biomass for Hipparion faunas.

The genus Sinolagomys is an early representative of the Family Ochotonidae, appearing first in the late early Oligocene of Central Asia. It is known in China from Shargaltein Tal, Taben Buluk, Ulan Tatal and northern Junggaria, and a few specimens from Tatal Gol, Mongolia have been published. For most occurrences the genus is not represented by many specimens. Extensive studies in the Valley of Lakes, Central Mongolia, have produced a large number of sinolagomyin fossils spanning at least ten million years, and belonging to five species: Sinolagomys kansuensis, S. major, S. gracilis, S. ulungurensis and S. badamae sp. nov. Descriptions of these are given, as well as definition of the new species. Sinolagomyins flourished during the late Oligocene and early Miocene and came to occupy vast territories from China through Mongolia and Kazakhstan. The evolution of this ochotonid group is characterized by increasing taxonomic diversity and progressive development of rootless cheek teeth.

The consensus view among geologists is that the Lanzhou Basin sequence of sediments contains the Oligocene/Miocene boundary, but where this occurs is unknown. Indeed, this is the crux of the issues concerning mid-Tertiary paleontology in Asia: what fauna corresponds with this time? What turnover events or distinctive taxa signal the beginning of the Miocene epoch? Once known, this will be of broad usefulness because vertebrate-bearing deposits are widespread in Asia and can be used to date basin sediments. Lanzhou and other basins permit a means of correlating to the time scale by using fauna and paleomagnetic data locally, and ultimately radiometric data from distant localities, to develop a precise biochronology. Herein, we summarize faunal constraints, primarily those of small mammal taxa, on correlation of the Lanzhou magnetozone sequence to the magnetic time scale. We conclude that the lower part of the Xianshuihe Formation contains theOligocene/Miocene boundary, and that the top of the formation, both in the south and north parts of the basin, is middle Miocene in age. Rodents from lower white sand units of the Xianshuihe Formation, correlate to the Xiejia assemblage of the Xining Basin, Qinghai Province, and are early, but not earliest, Miocene age. The implication for rodent faunas of the epoch boundary is that they retained a mainly "Oligocene" composition, and that other presumed late Oligocene assemblages may be, in fact, early Miocene in age. The Lanzhou appearance of Proboscidea is 19-18 Ma, as expected.

Faunal assemblages consist of immigrants, endemics and long-term residents; changes in proportions of these categories through time reflect general aspects of faunal stability and turnover. To study stability and change in the long biostratigraphic succession of Miocene terrestrial mammalian faunas recorded mainly in the Potwar Plateau Siwaliks of Pakistan, we distinguish long-term residents and assess new appearances as endemic vs. immigrant components. Our data represent the biogeography of the northern Indian subcontinent, but because neighboring areas of similar age are not as well known, virtually all species are found only there, and the proportion of the fauna that is immigrant is not apparent. We use preceding faunal assemblages to judge whether individual species might have originated there. We find many long-term residents in the Siwaliks, and of new species, most are immigrants and fewer are likely to have originated endemically. Siwalik rodents and artiodactyls show high rates of immigration for both during the early middle Miocene, followed by declining immigration but continuing residents. In the late middle Miocene both groups show stability, with a high proportion of long-term residents, and high endemism among rodents. In contrast, late Miocene rodent and artiodactyl patterns differ. Rodents decline in diversity but show increased immigration and new long-term residents; artiodactyls show turnover but then recover with many persisting residents.