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Exploring Atlantic Puffin Ecological segregation between two closely related species: exploring Atlantic puffin and razorbill foraging hotspots. by Stephanie C. Symons B.Sc. Zoology, University of Guelph, 2014 A Thesis Submitted in Partial Fulfillment of the Requirements for the Degree of Master of Science in the Graduate Academic Unit of Biology Supervisors: Antony Diamond, Ph.D., Biology Department and Forestry and Environmental Management Graham Forbes, Ph.D., Biology Department and Forestry and Environmental Management Examining Board: Les Cwynar, Ph.D., Biology Department, Chair Joseph Nocera, Ph.D., Forestry and Environment Management Heather Major, Ph.D., Biology Department at UNBSJ This thesis is accepted by the Dean of Graduate Studies THE UNIVERSITY OF NEW BRUNSWICK January, 2018 ©Stephanie C. Symons, 2018 ABSTRACT Threats to the marine environment are on the rise in the Bay of Fundy and the Gulf of Maine. Significant changes in marine food webs are already affecting the most important seabird breeding site in New Brunswick, Machias Seal Island. During the breeding season, this island is home to several seabird species, including regionally important colonies of Atlantic Puffins (Fratercula arctica) and Razorbills (Alca torda). These two auks coexist sympatrically during the breeding season, occupying similar dietary niches. Additionally, these birds are central place foragers, restricting their foraging ranges by the need to make several return trips (~10 per day) to feed their young or relieve their mate. In recent years, their food supply has deteriorated and there is concern for the future of auks in this region. Current knowledge of their at-sea distribution in the Bay of Fundy is especially fragmented, depending mainly on casual observations by passersby or the occasional survey. Until recently, GPS loggers providing fine scale data, were too large to deploy on any but the largest seabirds. Using a mix of GPS technology and direct observation of chick diet, I determined that puffins and razorbills avoid niche overlap by foraging in different places. Slight differences in diet and dive depth are likely consequences of foraging location. Despite detecting no differences in nest attendance, provisioning, and chick growth at the burrow-level, individual behavioural observations revealed that mates of individuals equipped with loggers provisioned significantly more than their tagged mate. These findings show that important tag effects can be overlooked with conventional methods. ii DEDICATION To my parents. Thank you for always supporting and encouraging my adventurous spirit. In loving memory of Ernest Vautour (1930-2013), Jeannita Vautour (1942-2017), and Charlene Vautour (1971-2017). iii ACKNOWLEDGEMENTS Thank you to my supervisor, Dr. Tony Diamond and my supervisory committee, Dr. Graham Forbes and Dr. Laura McFarlane Tranquilla. Your guidance and constructive criticism were greatly appreciated. Thank you to my ALAR lab mates for helping me work through several struggles and providing feedback and insightful ideas. Thank you to my fellow field technicians and colleagues, Katherine Shlepr, Linda Takahashi, Jennifer Symons, Stefanie Collar, and Michelle Valiant for their patience and commitment to data collection. I would also like to thank captains, Andy Patterson and Peter Wilcox, along with Durlan Ingersoll and Renee Patterson for transport and ferrying supplies to their island. Thank you to Dr. Karel Allard for lending his expertise and insight during the early stages of my project. For help DNA sexing puffins and razorbill, I would like to thank Jin- Hong Kim and Dr. Jason Addison. Finally, a heartfelt thank you to my parents and many friends that I have made along the way; thank you for keeping me laughing, and for always listening to my banter. iv Table of Contents ABSTRACT ........................................................................................................................ ii DEDICATION ................................................................................................................... iii ACKNOWLEDGEMENTS ............................................................................................... iv Table of Contents ................................................................................................................ v List of Tables ..................................................................................................................... vi List of Figures ................................................................................................................... vii Chapter 1 : Introduction and Overview .............................................................................. 1 Chapter 2 : Competitive exclusion in Atlantic puffins and razorbills in the face of declining food – feeding areas, depth, and diet. ............................................................... 19 Chapter 3 : Short-term GPS tag attachment disrupts feeding by Atlantic puffins, Fratercula arctica, and razorbills, Alca torda. ................................................................... 52 Chapter 4 : Summary ........................................................................................................ 87 Curriculum Vitae v List of Tables Table 3.1. Mean growth rates during the linear growth period (± standard deviation) and number of chicks fledged for Atlantic puffins and razorbills tagged at different breeding stages during preliminary field season in 2014. Adults of control burrows were not manipulated and chicks were measured throughout their development in the same way as adults manipulated while incubating, early-, mid-, and late-chick rearing. ...................... 78 Table 3.2 Burrow-level mean biomass per load, fish per load and feedings per day brought in to Atlantic puffin and razorbill chicks on Machias Seal Island, NB. Experimental burrows had one adult equipped with a GPS logger. Puffin feeding rates for procedural control burrows were collected by conducting dawn to dusk feeding watches. Days represent the total number of daylight hours recorded divided by the number of daylight hours in a day during the breeding season. Observations of biomass per load (g) and fish per load were made from video in 2015, whereas feedings per day were observed from video in 2014 and 2015 ............................................................................................ 79 Table 3.3 Individual-level mean biomass per load, fish per load and feedings per day brought in to Atlantic puffin and razorbill chicks in experimental burrows on Machias Seal Island, NB. Days represent the total number of daylight hours recorded divided by the number of daylight hours in a day during the breeding season. Observations of biomass per load (g) and fish per load were made from video in 2015, whereas feedings per day were observed from video in 2014 and 2015. ...................................................... 80 vi List of Figures Figure 2.1. Attachment method for GPS loggers on Atlantic puffins and razorbills, using 2 cable ties (1. and 4.) and 2 pieces of TESA® tape (2. and 3.). ...................................... 34 Figure 2.2. Ecotone loggers attached to the back feathers of an Atlantic puffin (a) and a razorbill (b). ...................................................................................................................... 35 Figure 2.3. Autocorrelated kernel density estimates (AKDE) for Atlantic puffins (orange) and razorbills (purple) breeding at Machias Seal Island, NB, in 2014 and 2015. Filled polygons represent the core foraging areas (50% AKDE contour), while the 90% AKDE contour shows the total foraging area. .............................................................................. 36 Figure 2.4. Bathymetry map overlaid with foraging points of puffins (orange) and razorbills (purple) from Machias Seal Island. Depths in the Gulf of Maine and Bay of Fundy range from -5 (light blue) to -300 m (dark blue). Puffins foraged in deeper waters and further from shore than razorbills. ............................................................................. 37 Figure 2.5. Frequency of dive durations (seconds). Approximately 56.8% of razorbill dives were 20-40 seconds, whereas puffins showed a more even range of dive duration.38 Figure 2.6. Subsample of available depths compared with depths at points used by puffins and razorbills diving around Machias Seal Island. Available depths are plotted in front of used depths. Depths at diving locations were significantly shallower than available depths in both species (puffins: df=1263, t=2.2, p=0.02; razorbills: df=909, t=33.4, p<0.001) .. 39 Figure 2.7. Dive duration (a) and percentage of dives based on time of day (hour ADT) for Atlantic puffins (ATPU) and razorbills (RAZO) studied on Machias Seal Island (MSI), Canada. Local time (Atlantic Daylight Savings) is UTC-3 hours. Red stars on (a) mark significant differences in length of dives when comparing hours of the day, holding vii species constant. Blue stars on (a) mark significant differences in dive length between species (boxplots) during that hour. .................................................................................. 40 Figure 2.8. Percent by mass of prey species brought in to puffin and razorbill chicks at Machias Seal Island in 2014 and 2015. Diet was variable for both species in 2014, whereas sand lance made up 92.5% of the puffin diet in 2015. In both years, razorbills brought in more herring than puffins. The Other group consists
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