Description of Variables for the dataset "Data from: Queenless honey bees build infrastructure for direct reproduction until their new queen proves her worth" Author: Michael L. Smith Author Contact: Department of Collective Behavior Max Planck Institute for Ornithology / University of Konstanz  msmith@orn.mpg.de michael.smith@uni-konstanz.de E: smithleemichael@gmail.com (preferred) P: +49 1772188269 Alternate Contact: Abstract: The terminal investment hypothesis predicts that individuals will alter their reproductive investments based on reproductive prospects. This hypothesis, however, has never been tested at the colony-level, where reproductive prospects for thousands of individuals can change instantly with the death of a single individual: the queen. A honey bee queen’s death also changes the reproductive mechanism; if the queen is not replaced, then workers reproduce directly, by producing males in reproductive comb – drone comb – before the colony dies. To test how workers respond to reproductive uncertainty, I made colonies queenless and measured comb building throughout the stages of rearing a replacement queen. Queenless workers built only drone comb throughout queen rearing, even when the colony possessed a virgin or mated queen. Only when the new queen started laying fertilized eggs did workers stop building drone comb. Despite queenless colonies being more likely to eventually rear a replacement queen (88%), than to fail (12%), workers still built drone comb for direct reproduction, even when the colony was provided with ample drone comb. Therefore, workers “pessimistically” invest in drone comb to prepare for direct reproduction, even while rearing a replacement queen to prevent that outcome. When faced with reproductive uncertainty, honey bees may “hope” for the best, but they prepare for the worst. This dataset supports the conclusions expressed in this paper. Keywords: Superorganism, colony state, reproductive uncertainty, queen failure, queen loss, Apis mellifera License Statement: These data are shared under a Creative Commons Attribution 4.0 International (CC-By 4.0) License. The data may be shared, used, and adapted freely, but attibution to the original authors must be given (see suggested citation below) and no more restrictive terms may be placed on it. Suggested Citation: Smith, Michael L. (2018) Data from: Queenless honey bees build infrastructure for direct reproduction until their new queen proves her worth. [Dataset] Cornell University Library's eCommons Repository. https://doi.org/10.7298/X4QZ285R Related Publications: Smith, Michael L. (2018) Queenless honey bees build infrastructure for direct reproduction until their new queen proves her worth. Evolution 72-12: 2810–2817. Published online: 17 Oct 2018. https://doi.org/10.1111/evo.13628 Smith, M.L., Ostwald, M.M., Loftus, J.C. et al. Naturwissenschaften (2014) 101: 783. https://doi.org/10.1007/s00114-014-1215-x Funding Details: This research was funded by an NSF-GRFP fellowship (DGE-1144153) and a NSF-DDIG grant (1600775). Geographic Location: This research was performed at the Liddell Field Station of Cornell University, in Ithaca, NY (42°27.6′ N, 76°26.7′ W), using colonies of honey bees headed by naturally mated queens (Apis mellifera lingustica). File Specific Details: 2013 EXPERIMENT: The data that were used in 2013 came from a larger experiment that was published in 2014, citation info: Smith ML, *Ostwald MM, *Loftus JC, Seeley TD. (2014). A critical number of workers in a honeybee colony triggers investment in reproduction. Naturwissenschaften. 101(10):783-790. DOI:10.1007/s00114-014-1215-x date: the date that the data were collected in format MM/DD/YY version: the experiment was done in two versions (see Smith et al 2014), version one refers to the first experiment, version two: refers to the repeated experiment, when all colonies had more workers day: experimental day that the data were collected sampling: sampling date that the data were collected (ie, sampling 1 = first time the colony was inspected to collect data, 2 = second time, etc) colony: unique ID for each colony letter refers to treatment group (see below) colonyid: unique ID for each colony where the letter does not refer to treatment group, instead just ""n"" as in the total sample size treatment: treatment groups for the experiment (see Smith et al 2014) ref = reference; honey = colonies with additional honey added; brood = colonies with additional brood added; comb = colonies with additional worker comb added (the comb devoid of honey or brood); pop = colonies with additional workers, so have a larger population of workers. WCA: Worker Comb Area - the area of worker comb built by the workers, measured as the number of 2x2cm squares. *To convert to actual area of cm2, one must multiply WCA by 4 DCA: Drone Comb Area - the area of drone comb built by the workers, measured as the number of 2x2cm squares. *To convert to actual area of cm2, one must multiply DCA by 4 TCA: Total Comb Area - the area of total comb built by the workers, measured as the number of 2x2cm squares. *To convert to actual area of cm2, one must multiply TCA by 4 WCP: Worker Comb Proportion - the proportion of newly built comb by the workers that was worker comb (i.e., WCA/TCA) DCP: Drone Comb Proportion - the proportion of newly built comb by the workers that was drone comb (i.e., DCA/TCA) QR: QueenRight; whether a colony possessed a queen on the date of inspection Y = yes, N = no ReliablyQR: Reliably QueenRight - whether a colony was queenright (had a queen) or queenless (did not have a queen) for all the sampling dates. Y = yes, N = no AddedWC: Added Worker Comb - the amount of worker comb that the workers had added (built) to the colony since the last date of inspection (e.g., if a colony had 10 units of worker comb on sampling date 1, and 15 units of worker comb on sampling date 2, then AddedWC would be 5, because the colony has added 5 units of worker comb between the two dates. AddedDC: Added Drone Comb - the amount of drone comb that was added (built) to the colony since the last date of inspection (e.g., if a colony had 10 units of drone comb on sampling date 1, and 15 units of drone comb on sampling date 2, then AddedDC would be 5, because the colony has added 5 units of drone comb between the two dates. TotalAdded: Total Added - the total amount of comb that the workers built since the last date of inspection (i.e., sum of AddedWC and AddedDC) PropDCAdd: Proportion Drone Comb Added - the proportion of comb that the workers had added (built) since the last date of inspection (i.e., AddedDC/TotalAdded) 2015 EXPERIMENT: ColonyID: unique number ID for each colony ExpDay: experimental day that the data were collected. Colonies were installed on 3 May 2015, Day 0 of the experiment, and so day 4 would be 7 May 2015, day 8 would be 11 May 2015, etc. WC_squares: WC_squares = Worker Comb Squares; meausring the comb built by the bees, WC_Squares is the number of 2x2cm squares that were worker comb. DC_squares: DC_squares = Drone Comb Squares; meausring the comb built by the bees, DC_Squares is the number of 2x2cm squares that were drone comb. TC_squares: TC_squares = Total Comb Squares; meausring the comb built by the bees, TC_Squares is the total number of 2x2cm squares built by the bees (i.e. WC_squares + DC_squares). WCA: Worker Comb Area - the cm2 area of worker comb built by the bees (WCA = WC_squares * 4) DCA: Drone Comb Area - the cm2 area of drone comb built by the bees (DCA = DC_squares * 4) TCA: Total Comb Area - the cm2 area of total comb built by the bees (TCA = TC_squares * 4) DeltaWCA: Delta Worker Comb Area - the amount of worker comb that the workers had added (built) to the colony since the last date of inspection (e.g., if a colony had 10 cm2 of worker comb on sampling date 1, and 15 cm2 of worker comb on sampling date 2, then DeltaWCA would be 5 cm2, because the colony has added 5 cm2 of worker comb between the two dates. DeltaDCA: Delta Drone Comb Area - the amount of drone comb that the workers had added (built) to the colony since the last date of inspection (e.g., if a colony had 10 cm2 of drone comb on sampling date 1, and 15 cm2 of drone comb on sampling date 2, then DeltaDCA would be 5 cm2, because the colony has added 5 cm2 of drone comb between the two dates. DeltaTCA: Delta Total Comb Area - the amount of total comb that the workers had added (built) to the colony since the last date of inspection (e.g., if a colony had 10 cm2 of total comb on sampling date 1, and 15 cm2 of total comb on sampling date 2, then DeltaTCA would be 5 cm2, because the colony has added 5 cm2 of total comb between the two dates. DeltaWCAdaily: Delta Worker Comb Area Daily - the amount of comb that was added, divided by the number of days between the two inspection dates. E.g., if a colony added 5cm2 of worker comb over 5 days, then the daily amount of worker comb added would be 1cm2. DeltaDCAdaily: Delta Drone Comb Area Daily - the amount of comb that was added, divided by the number of days between the two inspection dates. E.g., if a colony added 5cm2 of drone comb over 5 days, then the daily amount of drone comb added would be 1cm2. DeltaTCAdaily: Delta Total Comb Area Daily - the amount of comb that was added, divided by the number of days between the two inspection dates. E.g., if a colony added 5cm2 of total comb over 5 days, then the daily amount of total comb added would be 1cm2. WCP_Total: Worker Comb Proportion Total - the total proportion of built comb that is worker comb DCP_Total: Drone Comb Proportion Total - the total proportion of built comb that is drone comb WCP_Delta: Worker Comb Proportion Delta - the proportion of comb that was built between two experimental dates that was worker comb (i.e., DeltaWCA/DeltaTCA)" DCP_Delta: Drone Comb Proportion Delta - the proportion of comb that was built between two experimental dates that was drone comb (i.e., DeltaDCA/DeltaTCA) Eggs?: Whether or not queen-laid eggs were present in the colony, Y = yes, N = no. QR_QX: QueenRight or QueenLess - whether a colony had a queen or not. QR includes virgin, mated, and laying queens" ColonyStatus: The colony's status with respect to the process of rearing a replacement queen. QX - Uncapped Q cell = colony is queenless and has an uncapped queen cell; QX - Capped Q cell = colony is queenless and has a capped queen cell; QR - Virgin = colony has a virgin queen; QR - mated = colony has a mated queen; QR - laying = colony has a laying queen 2016 EXPERIMENT: ColonyID: unique ID for each colony, each beginning with c (for colony) followed by a number Treatment: treatment group that the colony was in. QXwoDC = QueenLess without Drone Comb added to the colony; QXwDC = QueenLess with Drone Comb added to the colony ExpDay: experimental day that the data were collected. Colonies were installed on 25 April 2016, Day 0 of the experiment, and so day 5 would be 30 April 2016, day 8 would be 4 May 2016, etc. WC_squares: Worker Comb Squares - meausring the comb built by the bees, WC_Squares is the number of 2x2cm squares that were worker comb. DC_squarea: Drone Comb Squares - meausuring the comb built by the bees, DC_Squares is the number of 2x2cm squares that were drone comb. TC_squares: Total Comb Squares - meausring the comb built by the bees, TC_Squares is the total number of 2x2cm squares built by the bees (i.e. WC_squares + DC_squares). WCA: Worker Comb Area - the cm2 area of worker comb built by the bees (WCA = WC_squares * 4) DCA: Drone Comb Area - the cm2 area of drone comb built by the bees (DCA = DC_squares * 4) TCA: Total Comb Area - the cm2 area of total comb built by the bees (TCA = TC_squares * 4) DeltaWCA: Delta Worker Comb Area - the amount of worker comb that the workers had added (built) to the colony since the last date of inspection (e.g., if a colony had 10 cm2 of worker comb on sampling date 1, and 15 cm2 of worker comb on sampling date 2, then DeltaWCA would be 5 cm2, because the colony has added 5 cm2 of worker comb between the two dates. DeltaDCA: Delta Drone Comb Area - the amount of drone comb that the workers had added (built) to the colony since the last date of inspection (e.g., if a colony had 10 cm2 of drone comb on sampling date 1, and 15 cm2 of drone comb on sampling date 2, then DeltaDCA would be 5 cm2, because the colony has added 5 cm2 of drone comb between the two dates. DeltaTCA: Delta Total Comb Area - the amount of total comb that the workers had added (built) to the colony since the last date of inspection (e.g., if a colony had 10 cm2 of total comb on sampling date 1, and 15 cm2 of total comb on sampling date 2, then DeltaTCA would be 5 cm2, because the colony has added 5 cm2 of total comb between the two dates. WCP_Total: Worker Comb Proportion Total - the total proportion of built comb that is worker comb DCP_Total: Drone Comb Proportion Total - the total proportion of built comb that is drone comb WCP_Delta: Worker Comb Proportion Delta - the proportion of comb that was built between two experimental dates that was worker comb (i.e., DeltaWCA/DeltaTCA) DCP_Delta: Drone Comb Proportion Delta - the proportion of comb that was built between two experimental dates that was drone comb (i.e., DeltaDCA/DeltaTCA) Eggs?: Whether or not queen-laid eggs were present in the colony, Y = yes, N = no. QR_QX: QueenRight or QueenLess - whether a colony had a queen or not. QR includes virgin, mated, and laying queens ColonyStatus: The colony's status with respect to the process of rearing a replacement queen. QX - Uncapped Q cell = colony is queenless and has an uncapped queen cell; QX - Capped Q cell = colony is queenless and has a capped queen cell; QR - Virgin = colony has a virgin queen; QR - mated = colony has a mated queen; QR - laying = colony has a laying queen ColonyStatus_v2: The colony's status with respect to the process of rearing a replacement queen, combined with the treatment group the colony is in (QXwoDC or QXwDC).