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Alloparental Care in a Solitary Bee

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Alloparental Care in a Solitary Bee Alloparental care in a solitary bee Vern Lewis, B.Sc. Submitted in partial fulfillment of the requirements for the degree of Master of Science Department of Biological Sciences, Brock University St. Catharines, Ontario © 2014 i Abstract Research into the evolutionary origins of sociality in insect colonies is changing emphasis from understanding how eusociality is maintained to how insects transition from solitary to social lifestyles. The pygmy carpenter bees (Ceratina spp.) offer an excellent model for investigating such factors as they have been historically thought of as solitary but have recently been shown to be socially polymorphic, which may indicate that they are currently in a transitive phase. By utilizing behavioural observation and experimental removal protocols, I show that extended parental care, as well as sibling care in Ceratina calcarata plays an important role in offspring development. I found, upon removal of the mother, that specifically produced ‘dwarf’ female offspring take over parental care roles in the nest. The existence of alloparental care and generational overlap suggests that although they are classified as solitary bees, C. calcarata possess the prerequisite behavioural repertoire for sociality. ii Acknowledgments First and foremost I would like to thank my supervisor, Dr. Miriam Richards. Without her guidance and infinite patience my dream of being a scientist would have dissolved into dull regret. I came here excited for the opportunity to use the knowledge I had already acquired, and leave in awe of that which I have yet to learn. Not only has Professor Richards made me a better scientist, but also a better student. Thank-you. I would also like to thank my committee members, Dr. Joffre Mercier and Dr. Gaynor Spencer. Their influence and feedback have helped make my project something I can be truly proud of. To my lab mates and friends, you have made my time here something much more than simply a stage in my career. To Tommy Onuferko, who delights in the small things, thank you for making me look forward to spending my whole day in the lab (and stop eating my candy). To Jess Vickruck, sometimes you have the mind of a sailor and that’s why we love you. To Rola Kutby, you were the most generous person I’ve met here, and the first to get me a birthday cake in almost 20 years. Thank you! Lastly to David Awde, who showed me that even ‘molecular’ people can be interesting. I will remember you all! Finally, to Tanya Kosanovic. My love, with whom I share all things, I dedicate this to you. Smooches! iii Contents Abstract .............................................................................................................................. ii Acknowledgments .............................................................................................................. iii List of tables ....................................................................................................................... vi List of figures ..................................................................................................................... vi Introduction ....................................................................................................................... 1 Parental care in insects .................................................................................................... 1 Social categories in insects ............................................................................................. 5 Solitary ........................................................................................................................ 5 Parasociality ................................................................................................................ 6 Eusociality ................................................................................................................... 6 Presocial ...................................................................................................................... 7 Routes to eusociality ....................................................................................................... 7 General description of nesting biology in the genus Ceratina ...................................... 10 Parental care in Ceratina ............................................................................................... 12 Social variation in the genus Ceratina .......................................................................... 13 Current state of sociobiological understanding in Ceratina calcarata ........................... 15 Hypotheses .................................................................................................................... 19 Objectives and predictions ............................................................................................ 20 Section 1: Baseline foraging behaviour and overwintering survival ........................ 20 Section 2: Offspring response to orphaning .............................................................. 20 Methods ............................................................................................................................ 22 Site location .................................................................................................................. 22 Field season timeline ..................................................................................................... 22 Preparation of nesting substrates .................................................................................. 25 Preparation of observation nests ................................................................................... 30 Spring phase .............................................................................................................. 30 Summer phase ........................................................................................................... 32 Marking of foragers ...................................................................................................... 34 Behavioural observations .............................................................................................. 35 Daily activity schedule .............................................................................................. 35 Quantifying foraging effort ....................................................................................... 36 Types of foraging trips .............................................................................................. 38 Brood survival ............................................................................................................... 38 Assessing gut contents .................................................................................................. 39 Overwintering survival experiment .............................................................................. 41 Data analysis ................................................................................................................. 42 Results .............................................................................................................................. 43 Section 1: Foraging patterns of Ceratina calcarata mothers ......................................... 43 Rates of nest founding .............................................................................................. 43 Daily patterns of foraging activity by C. calcarata mothers ..................................... 43 Spring foraging patterns of C. calcarata mothers ...................................................... 47 Summer foraging patterns of C. calcarata mothers ................................................... 53 Activity rates of C. calcarata mothers ....................................................................... 57 Section 2: Offspring response to brood orphaning ....................................................... 61 Brood survival ........................................................................................................... 61 iv Identity of foraging daughter .................................................................................... 62 Comparison of foraging mothers vs. foraging daughters .............................................. 64 Seasonal foraging patterns of foraging daughters ..................................................... 64 Daily foraging patterns of foraging daughters .......................................................... 64 Brood feeding by the forager ........................................................................................ 69 Necessity of summer phase brood provisioning ........................................................... 71 Discussion ......................................................................................................................... 74 Section 1: Baseline reproductive behaviour in Ceratina calcarata females .................. 74 Seasonality in foraging activity .................................................................................... 74 Parental care and foraging activity ........................................................................... 75 Resource abundance and foraging activity ............................................................... 76 Necessity for summer phase brood provisioning .......................................................... 78 Section 2: Ceratina calcarata’s response to brood orphaning ....................................... 79 Foraging rates in mothers vs. alloparents ................................................................
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