BRIDGING AQUATIC AND TERRESTRIAL ECOSYSTEMS: ECOLOGY OF SEMI-AQUATIC MAMMALS IN SOUTHERN ILLINOIS

Author

Angela Holland, Southern Illinois University CarbondaleFollow

Date of Award

8-1-2016

Degree Name

Doctor of Philosophy

Department

Zoology

First Advisor

Hellgren, Eric

Abstract

Mammals in freshwater aquatic systems play important roles as ecosystem engineers, trophic transfer agents, and apex predators, thus acting as indicators of freshwater ecosystem function. Watersheds inhabited by semi-aquatic mammals have increased links between adjacent terrestrial and aquatic ecosystems compared to watersheds where they are not present. Semi-aquatic mammals not only exert top-down influences on streams, but are affected by bottom-up forces from the riparian system itself. The goal of this study was to identify variables that correlate with the presence of beaver (Castor canadensis), muskrat (Ondatra zibethicus), mink (Neovison vison), and river otter (Lontra canadensis), including their interactions, resulting in a better understanding of the areas where these semi-aquatic mammals occur and their effects on the riparian system. The objectives of this study were (1) to identify variables related to the probability of detection, initial occupancy, colonization, and extinction of the 4 semi-aquatic mammals in southern Illinois; and (2) to test if the reintroduction of river otter has changed stream food webs. To address my first objective, I sampled 120 bridge sites in 2 periods (winter: Jan-Feb; and spring: Mar-Apr) during 2012–2014 in 11 major watersheds in the southern third of Illinois (44,526 km2) to estimate multi-season occupancy. Each survey unit was a 400-m stream segment visited twice by 2 observers for a total of 4 observations per site per period. Observers recorded all mammal signs found, including sign species and type. Sites were Intensive Basin Survey Sites sampled by Illinois Department of Natural Resources (IDNR) and Illinois Environmental Protection Agency (IEPA), allowing data collected by the state to be available for explanatory variables for mammal occupancy. Data collected by the Illinois Natural History Survey (INHS) also were available for a subset of sites. I collected local- and landscape-scale habitat and weather variables for each site. I developed hypotheses regarding occupancy of sites based on land-cover, human disturbance, and stream attributes for each species. I developed additional hypotheses regarding prey availability and water quality for river otter and mink. Sites used in each analysis were dependent on data available to address the hypotheses of interest. Beaver and muskrat were present at ≥100 of 103 sites for ≥1 observation. Naïve occupancy was high (≥82%) every year for both species. Detection probabilities for beaver and muskrat were best predicted by survey period. Beaver detection remained fairly constant across survey periods except for a decrease in winter 2014, whereas muskrat detection was generally lower during winter and higher during spring. Beaver were more likely to occupy larger streams than smaller streams during the initial survey period. Sites that lacked a dominant land-cover had a lower probability of beaver colonization than sites dominated by agriculture or woody vegetation at the landscape scale. In addition, the probability that a site would be colonized by beaver during the study increased with availability of water within 1 km of the surveyed segment, increased stream density, sites at larger streams, and river otter presence during the previous period. The probability of site extinction decreased as stream size, stream density within 1 km, and water availability within 500 m of the survey segment increased. Occupancy of muskrat during the initial survey period was negatively predicted by % forest in the 1-km riparian area, channel incision, and amount mercury in the sediment. Colonization by muskrats was lower during the long spring to winter intervals than the short winter to spring intervals, and was positively related to the amount of organic carbon in the sediment. The probability of site extinction by muskrat increased with increasing % forest within the riparian area around the stream segment, decreasing 1-km stream density, and when land-cover within 1 km of sites was dominated by agriculture or woody vegetation. Multi-season occupancy of river otter and mink were assessed in 2 separate analyses that used either land-cover and management variables or prey availability and riparian community composition, respectively. In the first analysis, river otter and mink were present at ≥84 of 103 sites. Naïve occupancy was higher every year for mink (≥88.3%) than for river otter (≥55.3%). Detection of river otter and mink in the first analysis increased as substrate availability increased. Occupancy of river otter during the initial survey period was predicted by large stream size, low % developed area within 250 m of the surveyed segment, and proximity to nearest river otter reintroduction point. Probability of colonization by river otter varied by survey period and was higher at sites with larger streams, higher stream density, lower % developed area, and within a known river otter population area. Site extinction by river otter in the first analysis varied by survey period and was linked to increased organic carbon in the sediment, and decreased road density within 1 km of the surveyed segment. River otter harvest was not found to affect site colonization or extinction. Mink occupancy during the initial period was negatively associated with water availability within 100 m of the survey segment. Site colonization by mink varied by month and increased with increasing developed area within 100 m of the surveyed segment, increasing channel incision, and decreasing rainfall. Probability of site extinction for mink increased as stream size and developed area within 500 m of the surveyed segment increased, and when woody vegetation was the dominant land-cover type within 1 km of the surveyed segment. The second analysis of multi-season occupancy of river otter and mink used 77 sites, 81.8% of which had ≥1 river otter detection in the study and 98.7% of which had ≥1 mink detection. Naïve occupancy differed between years but gradually increased for river otter and remained high (≥93.5%) for mink. Increasing substrate availability increased the probability of river otter detection, whereas mink detection varied by survey period. Occupancy during the initial survey period was higher in sites closer to the reintroduction points for river otter. Probability of colonization of river otter was positively associated with macroinvertebrate IBI and fish species richness, sites with high fish species richness of fish families preferred by river otter also had reduced otter extinction probability. No tested variables predicted initial occupancy for mink, but mink were more likely to colonize sites with increased fish richness and when muskrat were present during the previous period. Mink had decreased probability of extinction in sites with increasing mussel community index. My results indicate that semi-aquatic mammals in Illinois were affected by a riparian habitat, water availability, and stream community variables at both the landscape and local scale. I found high occupancy of mink, beaver, and muskrat across the entire landscape of southern Illinois, and my results suggest that the geographic range of river otter continues to expand. Relationships of occupancy of these semi-aquatic mammals to measurements of urban areas and human disturbance were not consistent across all species. Mink and river otter occupancy were both predicted by aspects of prey availability, indicating the importance of predator-prey relationships in occupancy dynamics of riparian predators. Hypotheses regarding predator pressure and changes in environmental variables were used to test the effects of river otter reintroduction on stream communities. For this objective, I used structural equation models. I compared fish and macroinvertebrate communities from before (1982-1995) and after (2005-2013) reintroduction of river otter, which occurred in 1994–1996. Fish and macroinvertebrate community data for 35 sites located throughout 6 major watersheds in southern Illinois (25,550 km2) were obtained from state agencies. Changes in stream communities were evaluated using 4 metrics (species richness, species dominance, skewness in size distribution of prey, and proportion of individuals in the size class preferred by river otter). Neither the inclusion of river otter site use nor change in stream quality, measured by change in % forest, improved models over the simple model which only included fish and macroinvertebrate communities. Overall, I found no evidence that river otter presence or change in forest cover affected stream fish and macroinvertebrate communities.