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MAMMAL ASSEMBLAGES OF THE CUYAHOGA VALLEY NATIONAL PARK: AN UPDATE AFTER 30 YEARS A thesis submitted To Kent State University in partial Fulfillment of the requirements for the Degree of Master of Sciences By Doug J. Marcum December, 2017 © Copyright All rights reserved Except for previously published materials Thesis written by Douglas J. Marcum B.S., Kent State University, 2011 M.S., Kent State University, 2017 Approved by ____________________________________, Advisor Oscar J. Rocha, Ph.D. ____________________________________, Chair, Department of Biological Sciences Laura G. Leff, Ph.D. ____________________________________, Dean, College of Arts and Sciences James L. Blank, Ph.D. TABLE OF CONTENTS……………………………………………………………………...... iii LIST OF FIGURES……………………………………………………………………………… v LIST OF TABLES…………………………………………………………………………...... viii ACKNOLEDGEMENTS………………………………………………………………………… x I. INTRODUCTION TO MAMMALIAN STUDIES IN THE CUYAHOGA VALLEY NATIONAL PARK, OHIO………………………………………...… 1 BACKGROUND………………………………………………………………… 1 STUDY AREA…………………………………………………………………... 2 MAMMALS OF CVNP………………………………………………………….. 5 STUDY DESIGN……………………………………………………………...…. 9 IMPLICATIONS OF RESEARCH…………………………………………...... 11 REFERENCES…………………………………………………………………. 12 II. DISTRIBUTION OF SMALL MAMMAL COMMUNITIES IN CUYAHOGA VALLEY NATIONAL PARK, OHIO AT TWO SPATIAL SCALES……………………………………………………...……... 14 INTRODUCTION…………………………………………………………….... 14 METHODS……………………………………………………………………... 17 STUDY SITES……………………………………………………….… 17 DATA COLLECTION…………………………………………………. 21 RESULTS………………………………………………………………………. 26 DISCUSSION………………………………………………………………..…. 40 REFERENCES…………………………………………………………………. 47 III. COMPARING METHODOLOGIES FOR STUDYING MAMMAL ASSEMBLAGES IN CUYAHOGA VALLEY NATIONAL PARK, OHIO….. 51 INTRODUCTION…………………………………………………………….... 51 METHODS……………………………………………………………………... 55 STUDY AREA…………………………………………………………. 55 DATA COLLECTION…………………………………………………. 56 RESULTS………………………………………………………………………. 66 iii DISCUSSION…………………………………………………………………... 85 REFERENCES…………………………………………………………………. 96 VI. CONCLUSIONS, SYNTHESIS, AND FUTURE STUDIES……………….... 100 CONCLUSIONS…………………………………………………………….... 100 FUTURE STUDIES AND MANAGEMENT IMPLICATIONS …………….. 108 REFERENCES………………………………………………………………... 111 APPENDICES A. HISTORICAL SPECIES LISTS FOR MAMMALS IN CUYAHOGA AND SUMMIT COUNTIES, OHIO…………………………………………………….. 114 B. MAMMALS OF CUYAHOGA VALLEY NATIONAL PARK RECORDED BY MAZZER ET AL. 1984………………………………………………………. 116 C. CHAPTER III CONTINGENCY TABLES………………………………………. 117 D. INVERTEBRATE DATA COLLECTED BY JAVIER OJEDA DURING PITFALL TRAPPING EFFORTS……………………………………………….... 118 E. EQUIPMENT SPECIFICATIONS………………………………………………... 120 iv LIST OF FIGURES Figure 1.1 Map of Cuyahoga Valley National Park……………………………………………... 3 Figure 2.1 Example of vegetation cover maps in CVNP from 1975 and 2013……………….... 18 Figure 2.2 Example of a topographic map indicating the sampling locations for small mammals used by Mazzer et al. (1984) in the CVNP……………………………… 19 Figure 2.3 Satellite image of northeast Ohio encompassing Cuyahoga Valley National Park… 20 Figure 2.4 Schematic representation of the standard trapping grid for small mammals……….. 23 Figure 2.5 Proportions of small mammals captured during Sherman trap efforts (2015) in CVNP………………………………………………………………………………. 27 Figure 2.6 Mean capture rate of small mammals by month……………………………………. 27 Figure 2.7 Average proportion of recaptures for Peromyscus leucopus and Microtus pennsylvanicus during subsequent nights of trapping in the summer season……….. 30 Figure 2.8 Mean canopy openness soil moisture and soil compaction for each successional stage………………………………………………………………………………… 33 Figure 2.9 Mean number of trees in each size class, standing dead trees, and woody debris in the vicinity of each trap for each successional stage…………………………….. 35 Figure 2.10 Mean percent cover for shrubs and herbaceous vegetation in the vicinity of each trap for each successional stage……………………………………………… 36 Figure 2.11 Correlation matrices among small mammal community data and between community data and microhabitat measures………………………………………. 37 Figure 2.12 Canonical Correspondence Analysis relating microhabitat variables to small mammal captures…………………………………………………………………. 38 Figure 3.1 Map of the Cuyahoga Valley watershed and surrounding region of northeast Ohio, United States………………………………………………………………….. 56 Figure 3.2 Photo of a camera trap placement used to inventory mammals in CVNP………….. 58 Figure 3.3 Map of Cuyahoga Valley National Park indicating road transects…………………. 60 Figure 3.4 Schematic representation of pitfall trapping design……………………………….... 62 v Figure 3.5 Photos of raptor pellets and an arrangement of identifiable mammal bones……….. 63 Figure 3.6 Abundance and frequency of mammal captures via camera traps………………….. 67 Figure 3.7 Loess regression showing positive relationship between species richness detected and number of camera trap nights…………………………………………. 68 Figure 3.8 Species accumulation curves for 28 camera trap locations in CVNP………………. 70 Figure 3.9 Proportion of three mammal groups represented in three successional stages of vegetation during camera trap surveys…………………………………………...… 71 Figure 3.10 Proportion of three mammal groups (individuals) found in three major forest types……………………………………………………………………………….. 72 Figure 3.11 Total detections for the eight most abundant mammal species during 13 road survey dates in the Cuyahoga Valley……………………………………………… 74 Figure 3.12 Number of individuals captured for each of six small mammals trapped during pitfall trapping across five sites in CVNP…………………………………………. 75 Figure 3.13 Small mammal richness by leaf litter depth across 15 pitfall trap arrays in CVNP……………………………………………………………………………… 77 Figure 3.14 Small mammal abundance by leaf litter depth across 15 pitfall trap arrays in CVNP……………………………………………………………………………… 77 Figure 3.15 Number of individuals identified for the five most common mammal species found in 71 raptor pellets………………………………………………………….. 79 Figure 3.16 Sample-based species rarefaction curve for five mammal inventory methods used in CVNP……………………………………………………………………... 81 Figure 3.17 Raw and corrected richness for Sherman trap, pitfall, and raptor pellet survey methods……………………………………………………………………………. 82 Figure 3.18 Detection biases for three small mammal survey methods by functional mammal groups…………………………………………………………………..... 82 Figure 3.19 Proportion of functional species groups detected by camera traps and road survey methods in CVNP corrected for sampling rate differences………………... 83 vi Figure 3.20 Principal components analysis comparing small mammal assemblages detected via Sherman trapping, pitfall trapping, and raptor pellet analysis ..………………....... 89 vii LIST OF TABLES Table 2.1 Names and descriptions of sites surveyed for small mammals in CVNP………….… 19 Table 2.2 Descriptor and methodology used for the assessment of microhabitat characteristics……………………………………………………………………….... 23 Table 2.3 Descriptor and methodologies used for the assessment of macrohabitat characteristics……………………………………………………………………….... 24 Table 2.4 Analysis of variance for capture rates observed in spring, summer, and fall………... 28 Table 2.5 Comparison of capture rates observed in the first and second trapping session, by sampling site…………………………………………………………………….... 29 Table 2.6 Lincoln-Peterson population density estimates per hectare for Peromyscus leucopus and Microtus pennsylvanicus………………………………………………. 29 Table 2.7 Summary of macrohabitat characteristics and small mammal community data by successional stage……………………………………………………………………. 31 Table 3.1 Functional classification of small mammals detected via Sherman trapping, pitfall trapping, and raptor pellet analysis………………………………………………….. 65 Table 3.2 Functional classification of mammals detected via camera trapping and road surveys……………………………………………………………………………….. 65 Table 3.3 Comparison of effort and success for five methods used to survey mammals in CVNP…………………………………………………………………………….... 66 Table 3.4 Comparison of mammal diversity metrics for three forest types in CVNP………….. 70 Table 3.5 Average detections per sample in forest versus non-forest habitats and wetland versus upland habitats for three functional groups of mammals in CVNP………….. 74 Table 3.6 Summary of pitfall captures and environmental variables measured in the CVNP by array and successional stage……………………………………………… 76 Table 3.7 Detectability rates for all mammal species documented by five survey methodologies in CVNP……………………………………………………………... 80 Table 3.8 Average proportion of species detected by camera traps and road survey methods in the Cuyahoga Valley…………………………………………………….. 84 viii Table 4.1 Small mammal species detected among three plant community successional stage categories during two inventories in CVNP………………………………... 101 Table 4.2 Small mammal species detected among three re-sampled sites from Mazzer et al. (1984) in CVNP…………………………………………………………….. 101 ix Acknowledgements The completion of this thesis is a monumental achievement for me. Throughout my entire career in school, I’ve been a procrastinator by nature, and I’ve often spent more time and energy in experiencing the world uninhibited by formal projects. I was known as the guy who “is always out in the woods”. My grandfather and mother would say that I always have “too many irons in the fire”. Knowing this about myself, I have strived in recent years to correct these tendencies by following through with ideas and projects that I’ve started before starting more projects and moving to new ideas. The challenge of

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