DOI: 10.5061/dryad.pnvx0k6wd
title: Ontogeny of superorganisms: Social control of queen specialization in ants

description: The functioning of biological systems relies on the cooperation of
 specialized components and understanding the processes that produce such
 specialization is a major challenge in biology. Here we study the ontogeny
 of biological systems at a new phenotypic level: the superorganisms (i.e.,
 insect societies with specialized individuals). We investigate how
 founding queens, the earliest developmental stage of ant colonies,
 transition from expressing behavioral pluripotency to becoming strictly
 specialized in egg production. We demonstrate that the presence of workers
 both initiates and maintains this queen specialization and propose that
 such a social control of queen behavior is common in ants and regulated by
 ancestral mechanisms. These findings contradict the traditional view of
 social insect queens as being intrinsically specialized in egg production
 and may reshape our understanding of division of labor in insect
 societies. 
description: All methods for the collection and analysis of the data are fully
 described in Majidifar et al (Functional Ecology, 2024).
description: # Ontogeny of superorganisms: Social control of queen specialization in
 ants The files contains
 all the data collected and analyzed in Majidifar et al. (Functional
 Ecology, 2024). All methods for the collection and analyses of the data
 are fully described in the article, and the R script for all statistical
 analyses of the datasets is available as Supporting Information. This
 README file was generated on 2024-02-09 by Romain LIBBRECHT. ## GENERAL
 INFORMATION ### Title of Dataset Ontogeny of superorganisms: Social
 control of queen specialization in ants ### Principal Investigator Contact
 Information Name: Romain Libbrecht Institution: Insect Biology Research
 Institute - UMR7261 - CNRS/University of Tours Address: Tours, France
 Email: romain.libbrecht@cnrs.fr ### Date of data collection 2017-2021 ###
 Geographic location of data collection Mainz, Germany ### Information
 about funding sources that supported the collection of the data Deutsche
 Forschungsgemeinschaft (DFG, German Research Foundation) - LI3051/2-1
 Deutsche Forschungsgemeinschaft (DFG, German Research Foundation) -
 GRK2526/1 - Project number 407023052 ## SHARING/ACCESS INFORMATION ###
 Licenses/restrictions placed on the data CC0 1.0 Universal (CC0 1.0)
 Public Domain ### Link to the publication using the data Majidifar, V.,
 Psalti, M. N., Coulm, M., Fetzer, E., Teggers, E-M., Rotering, F.,
 Grünewald, J., Mannella, L., Reuter, M., Unte, D., Libbrecht, R. (2024).
 Ontogeny of superorganisms: Social control of queen specialization in
 ants. Functional Ecology. ### Recommended citation for this dataset
 Majidifar, V., Psalti, M. N., Coulm, M., Fetzer, E., Teggers, E-M.,
 Rotering, F., Grünewald, J., Mannella, L., Reuter, M., Unte, D.,
 Libbrecht, R. (2024). Ontogeny of superorganisms: Social control of queen
 specialization in ants [Dataset]. Dryad.
 https://doi.org/10.5061/dryad.pnvx0k6wd ## DATA & FILE OVERVIEW ###
 File list 1. Brood_production.txt 2. Effect_of_worker_emergence.txt 3.
 Effect_of_worker_presence_and_food.txt 4.
 Reversibility_of_worker_effect_3_days.txt 5.
 Reversibility_of_worker_effect_2.5_years.txt 6.
 Reversibility_of_worker_effect_sequential_3.5_years.txt 7.
 Effect_of_workers_on_virgin_queens.txt 8.
 Effect_of_number_of_workers_5_larvae.txt 9.
 Effect_of_number_of_workers_15_larvae.txt 10.
 Effect_of_worker_cues_worker_CHCs.txt 11.
 Effect_of_worker_cues_former_worker_presence.txt 12.
 Effect_of_worker_cues_wire_mesh_BORIS_output.txt 13.
 Effect_of_worker_cues_wire_mesh_design_info.txt 14.
 Effect_of_worker_cues_dead_workers_BORIS_output.txt 15.
 Effect_of_worker_cues_dead_workers_design_info.txt 16.
 Effect_of_methoprene_treatment.txt 17. Effect_of_precocene_treatment.txt
 18. Effect_of_larvae_on_repro.txt 19.
 Reversibility_in_other_species_Tnyl.txt #### Brood_production.txt Number
 of variables: 4 Number of rows: 3697 Variable list * Queen: Queen
 individual ID * Day: Number of days since start of the experiment *
 Brood_type: Type of brood item (Eggs, Larvae, Pupae, Workers) * Quantity:
 Quantity of brood item #### Effect_of_worker_emergence.txt Number of
 variables: 5 Number of rows: 25 Variable list * Treatment: Whether the
 observation was made before worker emergence (1_YQP) or after worker
 emergence (2_YQM) * Queen_Number: Queen individual ID * Touching: Number
 of times the queen touched the brood * Grooming: Number of times the queen
 groomed the brood * Carrying: Number of times the queen carried the brood
 #### Effect_of_worker_presence_and_food.txt Number of variables: 15 Number
 of rows: 841 Variable list * Colony: Colony ID * Tube: Tube label * Day:
 Number of days since start of the experiment * Time_of_day: Whether the
 observation was made in the morning or in the afternoon * Day_time:
 Concatenation of Day and Time_of_day * Time_increment: Number of half days
 since start of the experiment * Estimated_hours: Number of hours since
 start of the experiment * Feeding_state: Whether the queen was fed (food)
 or not (no_food) * Worker_presence: Whether the queen was experimentally
 provided with workers (workers) or not (no_workers) * Combination:
 Combination of Feeding_state and Worker_presence (NFNW = no_food and
 no_workers, FW = food and workers, NFW = no_food and workers, FNW = food
 and no_workers) * Observation_time: Duration of the observation in seconds
 * Nursing_events: Number of times the queen expressed brood care behavior
 * Nursing_events_per_min: Number of times per minute the queen expressed
 brood care behavior * Number_larvae_at_day0: Number of larvae at the start
 of the experiment * Number_pupae_removed_at_day0: Number of pupae removed
 at the start of the experiment ####
 Reversibility_of_worker_effect_3_days.txt Number of variables: 6 Number of
 rows: 145 Variable list * Box_label: Observation box ID * Treatment:
 Whether the queen was experimentally provided with workers (With_workers)
 or not (Without_workers) or with workers that were then removed
 (Workers_removed) * Tray: Tray ID. Boxes on the same tray were filmed
 together * Time: Number of hours since start of the experiment * Nursing:
 Number of observation scans where the queen expressed brood care behavior
 * Scans: Total number of observation scans ####
 Reversibility_of_worker_effect_2.5_years.txt Number of variables: 6 Number
 of rows: 43 Variable list * Box_label: Observation box ID * Tray: Tray ID.
 Boxes on the same tray were filmed together * Time: Number of hours since
 start of the experiment * Nursing: Number of observation scans where the
 queen expressed brood care behavior * Scans: Total number of observation
 scans #### Reversibility_of_worker_effect_sequential_3.5_years.txt Number
 of variables: 7 Number of rows: 145 Variable list * Box_label: Observation
 box ID * Number_of_workers: Number of workers in the box at the time of
 the observation * Batch: Batch ID. Boxes in the same batch were handled
 together * Tray: Tray ID. Boxes on the same tray were filmed together *
 Time: Number of hours since start of the experiment * Nursing: Number of
 observation scans where the queen expressed brood care behavior * Scans:
 Total number of observation scans ####
 Effect_of_workers_on_virgin_queens.txt Number of variables: 7 Number of
 rows: 57 Variable list * Colony_ID: Colony ID * Wings_removed: Whether the
 queens had their wings experimentally removed (yes) or not (no) * Workers:
 Whether the queen was experimentally provided with workers (yes) or not
 (no) * Time: Number of hours since start of the experiment * Treatment:
 Whether the queen was experimentally provided with workers (with worker)
 or not (without worker) * Batch: Batch ID. Queens in the same batch were
 handled together * Nursing_events: Number of times the queen expressed
 brood care behavior #### Effect_of_number_of_workers_5_larvae.txt Number
 of variables: 6 Number of rows: 229 Variable list * Box_label: Observation
 box ID * Number_workers: Number of workers experimentally provided to the
 queen * Tray: Tray ID. Boxes on the same tray were filmed together * Time:
 Number of hours since start of the experiment * Nursing: Number of
 observation scans where the queen expressed brood care behavior * Scans:
 Total number of observation scans ####
 Effect_of_number_of_workers_15_larvae.txt Number of variables: 7 Number of
 rows: 241 Variable list * Box_label: Observation box ID * Number_workers:
 Number of workers experimentally provided to the queen * Batch: Batch ID.
 Boxes in the same batch were handled together * Tray: Tray ID. Boxes on
 the same tray were filmed together * Time: Number of hours since start of
 the experiment * Nursing: Number of observation scans where the queen
 expressed brood care behavior * Scans: Total number of observation scans
 ## Effect_of_worker_cues_worker_CHCs.txt Number of variables: 11 Number of
 rows: 109 Variable list * Box_label: Observation box ID * Treatment:
 Whether the queen was provided with glass beads coated with CHC (CHC) or
 hexane (Hexane) * Tray: Tray ID. Boxes on the same tray were filmed
 together * Time: Number of hours since start of the experiment *
 Nursing_behavior: Number of observation scans where the queen expressed
 brood care behavior * Scans: Total number of observation scans *
 Number_of_eggs_in_queen_tube: Number of eggs in queen tube *
 Number_of_larvae_in_queen_tube: Number of larvae in queen tube *
 Number_of_pupae_in_queen_tube: Number of pupae in queen tube *
 Number_of_larvae_killed_or_disappeared_at_the_time_of_sample_collection:
 Number of larvae killed or disappeared at the time of sample collection *
 Number_of_eggs_in_petri_dish_at_the_time_of_sample_collection: Number of
 eggs in petri dish at the time of sample collection ####
 Effect_of_worker_cues_former_worker_presence.txt Number of variables: 13
 Number of rows: 109 Variable list * Box_label: Observation box ID *
 Treatment: Whether the queen was kept in a box that used to contain
 workers (Yes) or not (No) * Tray: Tray ID. Boxes on the same tray were
 filmed together * Time: Number of hours since start of the experiment *
 Nursing_behavior: Number of observation scans where the queen expressed
 brood care behavior * Scans: Total number of observation scans *
 Former_worker_presence: Whether the queen was kept in a box that used to
 contain workers (Yes) or not (No) * Tray: Tray ID. Boxes on the same tray
 were filmed together * Number_of_eggs_in_queen_tube: Number of eggs in
 queen tube * Number_of_larvae_in_queen_tube: Number of larvae in queen
 tube * Number_of_pupae_in_queen_tube: Number of pupae in queen tube *
 Number_of_larvae_killed_or_disappeared_at_the_time_of_sample_collection:
 Number of larvae killed or disappeared at the time of sample collection *
 Number_of_eggs_in_petri_dish_at_the_time_of_sample_collection: Number of
 eggs in petri dish at the time of sample collection ####
 Effect_of_worker_cues_wire_mesh_BORIS_output.txt Number of variables: 6
 Number of rows: 8571 Variable list * Observation: Observation ID *
 Behavior: Whether the queen expresses brood care (1) or not (0) in the
 scan * Start: Time of the start of the scan (in seconds) * Stop: Time of
 the start of the scan (in seconds) * Box_label: Observation box ID *
 Time_since_setup_h: Number of hours since start of the experiment ####
 Effect_of_worker_cues_wire_mesh_design_info.txt Number of variables: 13
 Number of rows: 61 Variable list * Box_label: Observation box ID * Tray:
 Tray ID. Boxes on the same tray were filmed together * Batch: Batch ID.
 Boxes in the same batch were handled together * Date: Date of experimental
 set-up * Treatment: Whether the queen was experimentally provided with
 workers (Callows_with_Q) or not (No_callows) or with workers separated
 from the queen and the brood with a wire mesh (Callows_separated) *
 Number_of_larvae: Number of larvae in the box * Number_of_callows: Number
 of workers in the box * Location_Queen: Location of queen collection *
 Queen_number: Queen individual ID * Number_of_eggs_in_queen_tube: Number
 of eggs in queen tube * Number_of_larvae_in_queen_tube: Number of larvae
 in queen tube * Number_of_pupae_in_queen_tube: Number of pupae in queen
 tube * Number_of_pupae_removed: Number of pupae that were removed from the
 queen tube prior to the experiment ####
 Effect_of_worker_cues_dead_workers_BORIS_output.txt Number of variables: 6
 Number of rows: 11017 Variable list * Observation: Observation ID *
 Behavior: Whether the queen expresses brood care (1) or not (0) in the
 scan * Start: Time of the start of the scan (in seconds) * Stop: Time of
 the start of the scan (in seconds) * Box_label: Observation box ID *
 Time_since_setup_h: Number of hours since start of the experiment ####
 Effect_of_worker_cues_dead_workers_design_info.txt Number of variables: 17
 Number of rows: 73 Variable list * Box_label: Observation box ID * Tray:
 Tray ID. Boxes on the same tray were filmed together * Batch: Batch ID.
 Boxes in the same batch were handled together * Date: Date of experimental
 set-up * Treatment: Whether the queen was experimentally provided with
 live workers (Callows_with_Q) or not (No_Callows) or with freshly killed
 workers (Dead_Workers) or with freshly killed callow workers
 (Dead_Callows) * Number_of_larvae: Number of larvae in the box *
 Number_of_Workers: Number of dead workers in the box *
 Number_of_dead_callows: Number of dead callows in the box *
 Number_of_callows: Number of live callow workers in the box *
 Location_Queen: Location of queen collection *
 Number_of_eggs_in_queen_tube: Number of eggs in queen tube *
 Number_of_larvae_in_queen_tube: Number of larvae in queen tube *
 Number_of_pupae_in_queen_tube: Number of pupae in queen tube *
 Number_of_pupae_removed: Number of pupae that were removed from the queen
 tube prior to the experiment * Number_of_larvae_at_the_end: Number of
 larvae at the end of the experiment * Number_of_eggs_at_the_end: Number of
 eggs at the end of the experiment * Number_of_callows_at_the_end: Number
 of callow workers at the end of the experiment ####
 Effect_of_methoprene_treatment.txt Number of variables: 15 Number of rows:
 44065 Variable list * Observation_id: Observation ID * Nursing: Whether
 the queen expresses brood care (1) or not (0) in the scan * Start: Time of
 the start of the scan (in seconds) * day_setup: Day of the experimental
 set-up * day_experiment: Day of the experiment * Hours_last_treatment:
 Number of hours since the last treatment * Hours_first_treatment: Number
 of hours since the first treatment * Box_Label: Observation box ID *
 Treatment: Treatment received by the queen * callows: Whether the queen
 was experimentally provided with workers (yes) or not (no) * queen_label:
 Queen individual ID * number_of_eggs_before_treatment: Number of eggs in
 the queen tube before treatment * number_of_larvae_before_treatment:
 Number of larvae in the queen tube before treatment * queen_location:
 Location of queen collection * collection_date: Date of queen collection
 #### Effect_of_precocene_treatment.txt Number of variables: 18 Number of
 rows: 56305 Variable list * Observation_id: Observation ID * Nursing:
 Whether the queen expresses brood care (1) or not (0) in the scan * Start:
 Time of the start of the scan (in seconds) * day_setup: Day of the
 experimental set-up * day_experiment: Day of the experiment *
 Hours_last_treatment: Number of hours since the last treatment *
 Hours_first_treatment: Number of hours since the first treatment *
 Box_Label: Observation box ID * Treatment: Treatment received by the queen
 * callows: Whether the queen was experimentally provided with workers
 (yes) or not (no) * queen_label: Queen individual ID *
 number_of_eggs_before_treatment: Number of eggs in the queen tube before
 treatment * number_of_larvae_before_treatment: Number of larvae in the
 queen tube before treatment * queen_location: Location of queen collection
 * collection_date: Date of queen collection * larvae_origin: Colony used
 as source of larvae * eggs_after_treatment_: Number of eggs after
 treatment * larvae_after_treatment: Number of larvae after treatment ####
 Effect_of_larvae_on_repro.txt Number of variables: 12 Number of rows: 331
 Variable list * Queen_ID: Queen individual ID * Larvae: Whether the queen
 was provided with larvae (Yes) or not (No) * Workers: Queens from
 established colonies were tested without workers (No), while this variable
 did not apply to founding queens (NA) * Founding_stage: Whether the queen
 tested was a founding queen (With_eggs) or an established queen
 (With_workers) * Initial_number_of_eggs: Number of eggs in the queen tube
 before the experimental set-up * Initial_number_of_larvae: Number of
 larvae in the queen tube before the experimental set-up *
 Initial_number_of_pupae: Number of pupae in the queen tube before the
 experimental set-up * Initial_number_of_workers: Number of workers in the
 queen tube before the experimental set-up * Number_of_eggs: Number of eggs
 in the experimental box * Number_of_larvae: Number of larvae in the
 experimental box * Number_of_workers: Number of workers in the
 experimental box * Day: Number of days since start of experiment ####
 Reversibility_in_other_species_Tnyl.txt Number of variables: 11 Number of
 rows: 106 Variable list * Code: Source colony ID * Date: Date of
 experimental set-up * Original_number_workers: Number of workers in source
 colony * Queens: Number of queens in source colony * Colony: Queen
 individual ID * Workers: Whether the queen was experimentally provided
 with workers (yes) or not (no) * broodcare: Number of observation scans
 where the queen expressed brood care behavior * antennating: Number of
 observation scans where the queen antennated the brood * no_broodcare:
 Number of observation scans where the queen did not express brood care
 behavior * scans: Total number of observation scans * Day: Number of days
 since start of experiment