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Faunal and Environmental Change in the Late Miocene Siwaliks of Northern Pakistan
- John C. Barry, Michèle E. Morgan, Lawrence J. Flynn, David Pilbeam, Anna K. Behrensmeyer, S. Mahmood Raza, Imran A. Khan, Catherine Badgley, Jason Hicks, Jay Kelley
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- Journal:
- Paleobiology / Volume 28 / Issue S2 / Spring 2002
- Published online by Cambridge University Press:
- 14 July 2015, pp. 1-71
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The Siwalik formations of northern Pakistan consist of deposits of ancient rivers that existed throughout the early Miocene through the late Pliocene. The formations are highly fossiliferous with a diverse array of terrestrial and freshwater vertebrates, which in combination with exceptional lateral exposure and good chronostratigraphic control allows a more detailed and temporally resolved study of the sediments and faunas than is typical in terrestrial deposits. Consequently the Siwaliks provide an opportunity to document temporal differences in species richness, turnover, and ecological structure in a terrestrial setting, and to investigate how such differences are related to changes in the fluvial system, vegetation, and climate. Here we focus on the interval between 10.7 and 5.7 Ma, a time of significant local tectonic and global climatic change. It is also the interval with the best temporal calibration of Siwalik faunas and most comprehensive data on species occurrences. A methodological focus of this paper is on controlling sampling biases that confound biological and ecological signals. Such biases include uneven sampling through time, differential preservation of larger animals and more durable skeletal elements, errors in age-dating imposed by uncertainties in correlation and paleomagnetic timescale calibrations, and uneven taxonomic treatment across groups. We attempt to control for them primarily by using a relative-abundance model to estimate limits for the first and last appearances from the occurrence data. This model also incorporates uncertainties in age estimates. Because of sampling limitations inherent in the terrestrial fossil record, our 100-Kyr temporal resolution may approach the finest possible level of resolution for studies of vertebrate faunal changes over periods of millions of years.
Approximately 40,000 specimens from surface and screenwash collections made at 555 localities form the basis of our study. Sixty percent of the localities have maximum and minimum age estimates differing by 100 Kyr or less, 82% by 200 Kyr or less. The fossils represent 115 mammalian species or lineages of ten orders: Insectivora, Scandentia, Primates, Tubulidentata, Proboscidea, Pholidota, Lagomorpha, Perissodactyla, Artiodactyla, and Rodentia. Important taxa omitted from this study include Carnivora, Elephantoidea, and Rhinocerotidae. Because different collecting methods were used for large and small species, they are treated separately in analyses. Small species include insectivores, tree shrews, rodents, lagomorphs, and small primates. They generally weigh less than 5 kg.
The sediments of the study interval were deposited by coexisting fluvial systems, with the larger emergent Nagri system being displaced between 10.1 and 9.0 Ma by an interfan Dhok Pathan system. In comparison to Nagri floodplains, Dhok Pathan floodplains were less well drained, with smaller rivers having more seasonally variable flow and more frequent avulsions. Paleosol sequences indicate reorganization of topography and drainage accompanying a transition to a more seasonal climate. A few paleosols may have formed under waterlogged, grassy woodlands, but most formed under drier conditions and more closed vegetation.
The oxygen isotopic record also indicates significant change in the patterns of precipitation beginning at 9.2 Ma, in what may have been a shift to a drier and more seasonal climate. The carbon isotope record demonstrates that after 8.1 Ma significant amounts of C4 grasses began to appear and that by 6.8 Ma floodplain habitats included extensive C4 grasslands. Plant communities with predominantly C3 plants were greatly diminished after 7.0 Ma, and those with predominantly C4 plants, which would have been open woodlands or grassy woodlands, appeared as early as 7.4 Ma.
Inferred first and last appearances show a constant, low level of faunal turnover throughout the interval 10.7–5.7-Ma, with three short periods of elevated turnover at 10.3, 7.8, and 7.3–7.0 Ma. The three pulses account for nearly 44% of all turnover. Throughout the late Miocene, species richness declined steadily, and diversity and richness indices together with data on body size imply that community ecological structure changed abruptly just after 10 Ma, and then again at 7.8 Ma. Between 10 and 7.8 Ma the large-mammal assemblages were strongly dominated by equids, with more balanced faunas before and after. The pattern of appearance and disappearance is selective with respect to inferred habits of the animals. Species appearing after 9.0 Ma are grazers or typical of more open habitats, whereas many species that disappear can be linked to more closed vegetation. We presume exceptions to this pattern were animals of the mixed C3/C4 communities or the wetter parts of the floodplain that did not persist into the latest Miocene. The pace of extinction accelerates once there is C4 vegetation on the floodplain.
The 10.3 Ma event primarily comprises disappearance of taxa that were both common and of long duration. The event does not correlate to any obvious local environmental or climatic event, and the pattern of species disappearance and appearance suggests that biotic interactions may have been more important than environmental change.
The 7.8 Ma event is characterized solely by appearances, and that at 7.3 Ma by a combination of appearances and disappearances. These two latest Miocene events include more taxa that were shorter ranging and less common, a difference of mode that developed between approximately 9.0 and 8.5 Ma when many short-ranging and rare species began to make appearances. Both events also show a close temporal correlation to changes in floodplain deposition and vegetation. The 7.8 Ma event follows the widespread appearance of C4 vegetation and is coincident with the shift from equid-dominated to more evenly balanced large-mammal assemblages. The 7.3 to 7.0 Ma event starts with the first occurrence of C4-dominated floras and ends with the last occurrence of C3-dominated vegetation. Absence of a consistent relationship between depositional facies and the composition of faunal assemblages leads us to reject fluvial system dynamics as a major cause of faunal change. The close correlation of latest Miocene species turnover and ecological change to expansion of C4 plants on the floodplain, in association with oxygen isotopic and sedimentological evidence for increasingly drier and more seasonal climates, causes us to favor explanations based on climatic change for both latest Miocene pulses.
The Siwalik record supports neither “coordinated stasis” nor “turnover pulse” evolutionary models. The brief, irregularly spaced pulses of high turnover are characteristic of both the stasis and pulse models, but the high level of background turnover that eliminates 65–70% of the initial species shows there is no stasis in the Siwalik record. In addition, the steadily declining species richness and abrupt, uncoordinated changes in diversity do not fit either model.
Corneal Limbal Foreign Bodies
- Abdul-Jabbar Ghauri, Imran J. Khan
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- Journal:
- Canadian Journal of Emergency Medicine / Volume 13 / Issue 4 / July 2011
- Published online by Cambridge University Press:
- 11 May 2015, pp. 277-278
- Print publication:
- July 2011
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Controls on fluvial systems in the Siwalik Neogene and Wyoming Paleogene
- Brian J. Willis, Anna Kay Behrensmeyer, Thomas M. Bown, Mary Kraus, John S. Bridge, Imran Khan
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- Journal:
- The Paleontological Society Special Publications / Volume 6 / 1992
- Published online by Cambridge University Press:
- 26 July 2017, p. 315
- Print publication:
- 1992
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The 3-km thick Neogene Siwalik Group (Himalayan foredeep in northern Pakistan) and the 2-km thick Paleogene Fort Union/Willwood Formations (Bighorn Basin, Wyoming) both preserve long records of fluvial deposition adjacent to rising mountain belts. Depositional environments and associated habitats change with spatially varying physiography and deposition by river systems that may differ greatly in size, sediment loads, depositional rates, drainage of adjacent floodplains, and taphonomy of organic remains. At times, some environments may not be preserved; for example, avulsion of channels to low areas removes more deposits of channel-distal environments as avulsions increase relative to net sediment aggradation rates. Recognition of such large-scale biases is important because they represent time scales over which long term paleoecological change is reconstructed, and requires knowledge of how drainage systems changed in time and space within these evolving basins.
The Siwalik Group was deposited by large rivers that filled a basin extending at least 1000 km along its axis and 150–250 km away from the mountain front. Despite the scale of these rivers relative to Siwalik exposures, transitions between different fluvial systems have been recognized. For example, a 1-km thick sequence bridging the boundary between Chinji and Nagri formations records displacement of a smaller river system (width < 2 km; depth 5-10 m; discharge 1000-1500 m3/s) by a larger system (width <5 km; depth 15-30 m; discharge at least 5,000-10,000 m3/s), with an associated upsection increase (30 to 70%) in the proportion of channel sandstones, increased mean sediment accumulation rates (150 to 300 m/my), decrease in poorly drained floodplain deposits and well developed paleosols, marked decrease in abundance of faunal remains, and a major change in faunal composition. Stratigraphically higher (Dhok Pathan Fm.), there is a lateral transition between deposits of dissimilar, coeval river systems with corresponding differences in local paleoenvironments and vertebrate taphonomy. Although upsection changes in environments and vertebrate faunas may generally reflect extrabasinal controls such as tectonism and climate change, our studies emphasize the importance of recognizing deposits from different contemporaneous river systems before inferring such large-scale controls on paleoenvironmental change through time.
The Bighorn Basin is an intermountain foreland basin extending 200 km along its axis and about 80 km across. A large portion of this basin is exposed, and thus it is possible to reconstruct the distribution of river systems and the spatial paleoenvironments in more detail than in the Siwaliks. The Bighorn Basin was traversed along its axis by an early Eocene, north-south flowing river that was joined by smaller rivers flowing transverse to the axis. The proportion of channel sandstones decreases upsection (50 to 25%) from the Fort Union to the Willwood Fm. The proportion of channel sandstones and the abundance of well developed paleosols decrease with increasing net sediment aggradation rates. Although channel deposits are concentrated along the basin axis in a more complex way in some stratigraphic intervals, it is unclear to what extent these changes reflect deposition by different rivers versus extrinsically controlled changes within individual river systems.