Terrestrial feeding in aquatic turtles: environment-dependent feeding behavior modulation and the evolution of terrestrial feeding in Emydidae

Evolutionary transitions between aquatic and terrestrial environments are common in vertebrate evolution. These transitions require major changes in most physiological functions, including feeding. Emydid turtles are ancestrally aquatic, with most species naturally feeding only in water, but some terrestrial species can modulate their feeding behavior appropriately for both media. In addition, many aquatic species can be induced to feed terrestrially. A comparison of feeding in both aquatic and terrestrial environments presents an excellent opportunity to investigate the evolution of terrestrial feeding from aquatic feeding, as well as a system within which to develop methods for studying major evolutionary transitions between environments. Individuals from eight species of emydid turtles (six aquatic, two terrestrial) were filmed while feeding underwater and on land. Bite kinematics were analyzed to determine whether aquatic turtles modulated their feeding behavior in a consistent and appropriate manner between environments. Aquatic turtles showed consistent changes between environments, taking longer bites and using more extensive motions of the jaw and hyoid when feeding on land. However, these motions differ from those shown by species that naturally feed in both environments and mostly do not seem to be appropriate for terrestrial feeding. For example, more extensive motions of the hyoid are only effective during underwater suction feeding. Emydids evolving to feed on land probably would have needed to evolve or learn to overcome many, but not all, aspects of the intrinsic emydid response to terrestrial feeding. Studies that investigate major evolutionary transitions must determine what responses to the new environment are shown by naïve individuals in order to fully understand the evolutionary patterns and processes associated with these transitions.

A VEGETATION RECONSTRUCTION OF UNION COUNTY, PENNSYLVANIA DURING THE EARLY HISTORICAL PERIOD (1755-1855)

This thesis assesses relationships between vegetation and topography and the impact of human tree-cutting on the vegetation of Union County during the early historical era (1755-1855). I use early warrant maps and forestry maps from the Pennsylvania historical archives and a warrantee map from the Union County courthouse depicting the distribution of witness trees and non-tree surveyed markers (posts and stones) in early European settlement land surveys to reconstruct the vegetation and compare vegetation by broad scale (mountains and valleys) and local scale (topographic classes with mountains and valleys) topography. I calculated marker density based on 2 km x 2 km grid cells to assess tree-cutting impacts. Valleys were mostly forests dominated by white oak (Quercus alba) with abundant hickory (Carya spp.), pine (Pinus spp.), and black oak (Quercus velutina), while pine dominated what were mostly pine-oak forests in the mountains. Within the valleys, pine was strongly associated with hilltops, eastern hemlock (Tsuga canadensis) was abundant on north slopes, hickory was associated with south slopes, and riparian zones had high frequencies of ash (Fraxinus spp.) and hickory. In the mountains, white oak was infrequent on south slopes, chestnut (Castanea dentata) was more abundant on south slopes and ridgetops than north slopes and mountain coves, and white oak and maple (Acer spp.) were common in riparian zones. Marker density analysis suggests that trees were still common over most of the landscape by 1855. The findings suggest there were large differences in vegetation between valleys and mountains due in part to differences in elevation, and vegetation differed more by topographic classes in the valleys than in the mountains. Possible areas of tree-cutting were evenly distributed by topographic classes, suggesting Europeans settlers were clearing land and harvesting timber in most areas of Union County.