Friday, November 20, 2015

The Secret Life of the Middle Holocene Eastern Great Plains Bison

Photograph taken by author at Neal Smith Wildlife Refuge

Imagine yourself wandering through the Iowa tallgrass prairie, you admire the lush tranquility of the big bluestem and compass plant, perhaps you stop to pick a couple of wild blackberries while listening to the call of the red-headed woodpecker. Suddenly, as if out of nowhere, you are confronted by a thunderous sound that seems grow louder and louder and discover the source of the noise appears to be the footsteps of a herd of large, four-legged animals. As these great beasts approach, you see clearer and clearer a herd of bison. Nowadays, this scene seems almost impossible but during the middle Holocene, 7,000 to 8,500 years ago, bison ruled the prairie. This species served as a keystone species: creating necessary disturbances such as grazing and wallowing to continue the success of native prairie plants. While ecologists now know that there were indeed large populations of bison, little is known about the everyday habits of these animals. What did they eat? Where did they drink? How far did they travel? All of these questions become even more pertinent when considering that during the middle Holocene, there was an increase in aridity and drought. Current research suggests that bison respond to a drier climate by increasing their range (Fortin et al. 2003), which might change their diet and water source. So the question remains: how did the bison population in the eastern Great Plains respond to climate changes in the middle Holocene? Answering these questions will not only provide insight into bison behavior but can also reveal the history of Iowa vegetation. Since the bison’s diet consisted of the flora available, the content of their diet should reflect the prairie’s composition.
            A study conducted by Widga et al. (2010) set out to answer where the middle Holocene bison wandered and what they ate. Using fossils of bison from five different populations located in tallgrass prairie and prairie-forest border in Iowa, Minnesota and Nebraska, the researchers used isotopic analysis to determine the diet, water source and seasonal migration patterns of each herd. Isotopes are a type of a specific element containing a different number of neutrons than the typical version of the same element, making it heavier or lighter. Isotopes are useful for ecologists because certain natural processes discriminate between isotopes in their incorporation of elements. Thus, these natural processes experience different isotopic ratios than would normally be expected. Analyzing the ratios of isotopes can lend insight into the historical behaviors of bison by comparing the different isotopic ratios in the fossils of bison compared with the expected ratios in the sources of forage and water.
Map of Locations of Bison Fossils Studied (Widga et al. 2010)

The bison populations sampled were selected because there were at least 29 individuals in each herd, and their deaths appeared to occur simultaneously. This study used tooth enamel as the source of isotopes as tooth enamel can easily reveal age allowing for a comparison of behaviors within the individual and between herd mates of the same age and different ages.
            13Carbon isotopes were used to analyze the bison’s’ diet. Tallgrass prairie plants incorporate carbon dioxide during photosynthesis in two distinct ways. The different pathways and the plants that utilize them are referred to as C3 and C4 plants. Most grasses are C4 plants, while woody vegetation is typically C3. The 13C to 12C ratio is higher in C4 plants. Since bison eat these plants and incorporate the carbon throughout their bodies including their teeth, investigating the ratio of 13C to 12C in the bison teeth will lend insight into the types of plants the middle Holocene bison ate. Widga et al. (2010) found that most of the bison sampled had diets higher in C4 plants than C3 plants. In fact, the bison’s’ diets in all but one herd consisted of 60-85% C4 plants. This insight on the bison’s diet enables us to better understand the composition of the tallgrass prairie during the middle Holocene, through the assumption that the bison ate a diet representational of the prairie flora. Therefore, we can conclude that the prairies in northwestern Iowa, northeastern Nebraska and southern Minnesota were roughly composed of 60-85% C4 plants. These findings add detail to a study conducted by Baker et al. (1996), which investigated vegetation composition in northeast Iowa using a variety of methods including pollen fossils, vascular plant macrofossils and isotopic evidence. Their study found that at this time there were more mesic deciduous forest species, C3 plants, than C4 grasses in northeastern Iowa. However using the same methods, Baker et al. (1996) found that 5,500-3,500 years ago (about 2,000 years after the bison investigated by Widga et al. (2010) died) northeast Iowa experienced an ecosystem transformation from forest to prairie. Thus, the bison data reveals that prairie arrived in northwest Iowa before spreading into northeast Iowa.
            Widga et al. (2010) used 18Oxygen as a proxy to determine what types of water sources bison utilized during this period. 18O is heavier than 16O and because of the increase in mass, 18O is less likely to be evaporated than 16O. In the tallgrass prairie, there are a variety of water sources available to bison. Sources such as ephemeral or upland lakes and ponds experience intense evaporation, especially during the summer. At these water sources, there would be a high ratio of 18O to 16O. In contrast, more stable water sources such as large lakes or rivers have low ratios of 18O to 16O. Therefore, bison were more likely to consume 18O in water sources prone to evaporation. The researchers found high concentration of 18O in the teeth sampled, revealing that these bison utilized water sources susceptible to high levels of evaporation. The bison may not have found it necessary to travel outside their normal water sources in the middle Holocene despite the increase in drought conditions.
            The last question investigated by Widga et al. (2010) concerned the movement of bison. Did the bison respond to the increase in temperature with an expansion in movement and home ranges, or as with diet and water sources, did the bison stick with similar patterns before the climate change? To answer this question, researchers compared isotopic ratios of 86Strontium and 87Strontium found in the bison’s teeth to the ratio found in the soil of different geologic areas where the bison may have grazed. Different soil types naturally contain different concentration of 86Sr and 87Sr, so one can differentiate which substrate type a bison was grazing on by the concentration found in the bison’s teeth. If the ratios of 87Sr and 86Sr found in the bison teeth match the soil surface ratio, it would be expected that these areas were utilized by the bison. Surprisingly, few bison of the eastern Great Plains actually traveled across more than one examined area. Further, every herd’s home ranges were smaller than expected, with a radius less than 50 kilometers. These data reveal that the bison did not respond to climatic pressures by searching for literally greener patches. This finding additionally indicates that despite rising temperature, the bison had enough to eat in these prairies.
            It appears from this study that the bison during the middle Holocene adopted a policy of if ain’t (that) broken, don’t fix it. The bison did not change their diet, water source or mobility in response to the increase in temperature and the expected limitation of resources that accompanies low water availability. Surprisingly the habits of the bison from the middle Holocene closely mimic the behaviors of bison today. One explanation for this finding may be that the flora that Bison grazed in the middle Holocene may be surprisingly similar to today’s reconstructed prairies where the bison are currently confined due to the destruction of almost all of C4 tallgrass prairie in Iowa. Importantly, these findings contradict the belief that bison movement and grazing preferences were strongly influenced by the structure of local resources. From Widga et al.’s (2010) findings, we can glean that bison roamed the eastern Great Plains tallgrass prairie year round, creating a constant influence on the behaviors and growth of tallgrass prairie species. While the ecosystem and climate have changed and shifted over thousands of years in the eastern Great Plains, bison activities concerning diet preferences, water source utilization and migration patterns have stayed fairly constant. Hence, at least for the middle Holocene eastern Great Plains bison, old habits truly die hard.

References
Baker, R. G., Bettis III, E.A., Schwert, D.P., Horton, D.G., Chumbley, C.A., Gonzalez,
            L.A. and Reagan, M.K. 1996. Holocene Paleoenvironments of Northeast Iowa.
            Ecological Monographs 66(2): 203-234.

Fortin, D., Fryxell, J.M., O'Brodovich, L. and Frandsen, D. 2003. Foraging ecology of
            bison at the landscape and plant community levels: the applicability of energy
            maximization principles. Oecologia 134: 219–237.

Widga, C., Walker, J.D. and Stockli, L.D. 2010. Middle Holocene bison diet and mobility
            in the eastern Great Plains (USA) based on 13C, 18O and 87Sr/86Sr analyses of
            tooth enamel carbonate. Quaternary Research 73 (2010) 449-460.


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