Metacommunity Dynamics Of Benthic Macroinvertebrates In A Large River

Metacommunity ecology focuses on the interaction between local communities and is inherently linked to dispersal as a result. Within this framework, communities are structured by a combination of in-site responses to the immediate environment (species sorting), stochasticity (patch dynamics), and connections to other communities via distance between communities and dispersal (neutrality), and source-sink dynamics (mass effects; see Chapter 1 for a detailed description of metacommunity theory, the study site, and macroinvertebrate communities found). In Chapter 2 I describe spatial scale of study and dispersal ability as both have the ability to influence the degree to which communities interact. However, little is known about how these factors influence the importance of all metacommunity dynamics. I compared dispersal mode of immature aquatic insects and dispersal ability of winged adults across multiple spatial scales in a large river. The strongest drivers of river communities were patch dynamics, followed by species sorting, then neutrality. Active dispersers during aquatic lifestages on average exhibited lower patch dynamics, higher species sorting, and significant mass effects compared to passive dispersers. Active and strong dispersers also had a scale-independent influence of neutrality, while neutrality was stronger at broader spatial scale for passive and weak dispersers. These results indicate as dispersal ability increases patch dynamics decreases, species sorting increases, and neutrality should decrease. The perceived influence of neutrality may also be dependent on spatial scale and dispersal ability. In Chapter 3 I describe how river benthic macroinvertebrate communities may influence tributary invertebrate communities via adult flight and tributaries may influence mainstem communities via immature drift. This relationship may also depend on relative mainstem and tributary size, as well as abiotic tributary influence on mainstem habitat. To investigate the interaction between a larger river and tributary I sampled mainstem benthic invertebrate communities and quantified habitat of a 7th order river (West Branch Susquehanna River) above and below a 5th order tributary confluence, as well as 0.95-3.2 km upstream in the tributary. Non-metric multidimensional scaling showed similar patterns of clustering between sampling locations for both habitat characteristics and invertebrate communities. In addition, mainstem river communities and habitat directly downstream of the tributary confluence cluster tightly together, intermediate between tributary and mid-channel river samples. In Bray-Curtis dissimilarity comparisons between tributary and mainstem river communities the furthest upstream tributary communities were least similar to river communities. Middle tributary samples were also closest by Euclidean distance to the upstream mainstem riffle and exhibited higher similarity to mid-channel samples than the furthest downstream tributary communities. My results indicate river and tributary benthic invertebrate communities may interact and likely result in direct and indirect mass effects of a tributary on the downstream mainstem community by invertebrate drift and habitat restructuring via material delivery from the tributary. I also showed likely direct effects of adult dispersal from the river and oviposition in proximal tributary locations where Euclidian, rather than river, distance may be more important in determining river-tributary interactions.

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.