The Effects of Habitat Parameters on the Behavior, Ecology, and Conservation of the Udzungwa Red Colobus Monkey
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The Effects of Habitat Parameters on the Behavior, Ecology, and Conservation of the Udzungwa Red Colobus Monkey ( Procolobus gordonorum ) by Ruth Irene Steel Department of Biological Anthropology and Anatomy Duke University Date:_______________________ Approved: ___________________________ Kenneth Glander, Supervisor ___________________________ Joanna Lambert ___________________________ Daniel Schmitt ___________________________ Thomas Struhsaker ___________________________ Christine Wall Dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the Department of Biological Anthropology and Anatomy in the Graduate School of Duke University 2012 ABSTRACT The Effects of Habitat Parameters on the Behavior, Ecology, and Conservation of the Udzungwa Red Colobus Monkey ( Procolobus gordonorum ) by Ruth Irene Steel Department of Biological Anthropology and Anatomy Duke University Date:_______________________ Approved: ___________________________ Kenneth Glander, Supervisor ___________________________ Joanna Lambert ___________________________ Daniel Schmitt ___________________________ Thomas Struhsaker ___________________________ Christine Wall An abstract of a dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in the Department of Biological Anthropology and Anatomy in the Graduate School of Duke University 2012 Copyright by Ruth Irene Steel 2012 Abstract A central theme in animal ecology is the study of the relationship between ecology and behavior. This dissertation demonstrates how ecological parameters, particularly food and weather variables, correlate with ranging, activity budget, and diet in Udzungwa red colobus monkeys (URC, Procolobus gordonorum ), endemic to the Udzungwa Mountains, Tanzania. From April 2009 – March 2010, four URC groups were habituated and studied in Mwanihana and Magombera Forests. During all-day follows, ranging, activity budget, and diet data were collected using GPS units and ten-minute scan samples of sustained activities. Food characteristics were assessed using transect and phenology surveys. Preliminary nutritional analyses were conducted on species-specific parts fed upon during October. I designed and employed novel methods for estimating ranging statistics which were compared with conventional methods. Home range sizes were relatively small (8.8 – 20.8 ha). Home range size may relate to food species diversity and feeding selectivity for species-specific food parts. Variability in dietary composition was partly explained by plant phenology, forest composition, and selectivity differences among groups. URC primarily fed upon young leaves with higher fat content. Some non-nutritive items eaten may function as detoxifying agents. Data suggest that the URC dietary strategy involves nutrient iv balancing (i.e. maximizing energetic intake and nutrient acquisition while avoiding high concentrations of particular plant secondary compounds). Mean daily travel distance (DTD) was 970 m, longer than in other red colobus taxa. DTD was shorter in October and November when temperatures were warmer and a high abundance of young growth became available. Young growth comprised the majority of all groups’ diets in each sample period. When young growth abundances were highest, Magombera groups traveled shorter distances. One group’s DTD did not differ seasonally and was significantly longer compared to other groups. This group may have traveled longer distances and spent less time resting in order to consume young growth in a habitat with lower young growth abundances. A correlation between time spent resting and temperature suggests behavioral prevention of hyperthermia. URC energetic strategies may involve balancing thermoregulation with maximizing energy intake. Conservation recommendations are discussed and include stricter forest protection. v Contents Abstract .............................................................................................................................. iv List of Tables .................................................................................................................... xv List of Figures .................................................................................................................. xxi Acknowledgements ........................................................................................................ xxvi 1. Introduction ..................................................................................................................... 1 Species background: Biological importance of the URC and its habitat ...................... 3 Goals of ecological research ......................................................................................... 6 2. The influence of ranging behavior method choice on comparative studies, ecological interpretations, and conservation planning ....................................................................... 12 Introduction ................................................................................................................. 12 Materials and Methods ................................................................................................ 15 Study area .................................................................................................................. 15 Study groups ............................................................................................................. 18 Ranging data ............................................................................................................. 19 Methods for estimating daily travel distance ............................................................ 20 Methods for estimating home range.......................................................................... 22 Results ......................................................................................................................... 24 Daily Travel Distance ............................................................................................... 24 Home Range Area ..................................................................................................... 28 Discussion ................................................................................................................... 36 Daily travel distance methods comparison ............................................................... 36 Multiple Travel Paths Method .............................................................................. 36 vi Center of Mass method ......................................................................................... 38 Implications for DTD research ................................................................................. 39 Future considerations for applying the Multiple Travel Paths method ..................... 42 Home range area methods comparison and research application ............................. 44 Implications for cross-study comparisons with home range data ............................. 47 Measuring Ranging Statistics: Groups versus Individuals ....................................... 48 Conclusions ................................................................................................................. 49 3. Home Range Size and Characteristics .......................................................................... 52 Introduction ................................................................................................................. 52 Materials and Methods ................................................................................................ 57 Study area .................................................................................................................. 57 Study groups ............................................................................................................. 58 Methods for estimating home range size .................................................................. 59 Methods for estimating population size and density ................................................. 62 Vegetation transects .................................................................................................. 63 Habitat Variables ...................................................................................................... 65 Food tree species variables ....................................................................................... 67 Phenology assessments ............................................................................................. 68 Food availability ....................................................................................................... 69 Statistical analyses .................................................................................................... 70 Results ......................................................................................................................... 71 Home range size ........................................................................................................ 71 Home range overlap .................................................................................................. 78 vii URC population density and size estimates in Magombera Forest .......................... 85 Habitat Parameters .................................................................................................... 87 Comparing food tree variables across home ranges ................................................