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Washboarding: Are Man-Managed Honeybees Performing a Vestigial Activity Fostered by Cryptogams?


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Fig. 1

Observations and experiments on hives: a) visual inspection of hive surfaces; b–c) lichen transplants composed of bark stripes glued on satin ribbons. b) D-type sample, collected in Portugal in 2018 and c) A-type sample, collected in Italy in 2019; d) placement of transplants. Photo record: Davide Romani.
Observations and experiments on hives: a) visual inspection of hive surfaces; b–c) lichen transplants composed of bark stripes glued on satin ribbons. b) D-type sample, collected in Portugal in 2018 and c) A-type sample, collected in Italy in 2019; d) placement of transplants. Photo record: Davide Romani.

Fig. 2

Beekeepers responding to a questionnaire indicate the frequency of their observations of washboarding. On the right, details on the number of times that each respondent observed the washboarding behaviour.
Beekeepers responding to a questionnaire indicate the frequency of their observations of washboarding. On the right, details on the number of times that each respondent observed the washboarding behaviour.

Fig. 3

Relative importance of possible explanations of washboarding, expressed as posts related to YouTube videos. They are grouped based on main themes.
Relative importance of possible explanations of washboarding, expressed as posts related to YouTube videos. They are grouped based on main themes.

Fig. 4

Bee responses to lichen stripes transplant: a) washboarding behaviour; b) lichen stripe sample conditions before placing; c) same sample as in b), after two months of exposure in front of the hive: evidence of lichen removal; d–f) decoloration of those lichens present on the bark slice after two months of exposure.
Bee responses to lichen stripes transplant: a) washboarding behaviour; b) lichen stripe sample conditions before placing; c) same sample as in b), after two months of exposure in front of the hive: evidence of lichen removal; d–f) decoloration of those lichens present on the bark slice after two months of exposure.

Fig. S1

Study area, where apiaries under experimental manipulation were placed, 45°13′59.96″N; 11°27′56.8″E, north-east of Italy.Maps credits: northern Italy at https://d-maps.com/carte.php?num_car=5880&lang=it;detail of the study area at https://www.google.it/maps (Immagini ©2022, TerraMetrics, Dati cartografici ©2022 Google)
Study area, where apiaries under experimental manipulation were placed, 45°13′59.96″N; 11°27′56.8″E, north-east of Italy.Maps credits: northern Italy at https://d-maps.com/carte.php?num_car=5880&lang=it;detail of the study area at https://www.google.it/maps (Immagini ©2022, TerraMetrics, Dati cartografici ©2022 Google)

Website list

Website Brief summary of content Opinion/info
1 https://beekeep.info/a-treatise-on-modern-honey-bee-management/the-basis-for-management/mysterious-behaviors-of-honey-bees/washboard-behavior/Last accessed: 18 Jan. 2021 Site dedicated to informing beekeepersmanaged by The Apis Information Resource Center, developed by Dr. Malcolm T. Sanford, Professor Emeritus, University of Florida intriguing behaviour;presentation of results of Bohrer, K., & Pettis, J.S.;hosting 2 videos and a link too a google page listing results for washboarding behaviour
2 https://honeybeesuite.com/washboarding-bees-arockin-and-alickin/Last accessed: 18 Jan. 2021 Site dedicated to informing beekeepers and peoplemanaged by Sam Droege, USGS Patuxent Wildlife Research Center Brief description in the blog, reporting results ofBohrer, K., & PettisJ.S.;hosting 1 video22 replying posts
3 https://ucanr.edu/blogs/blogcore/postdetail.cfm?postnum=7586Last accessed: 18 Jan. 2021 Informative site of theAgriculture and Natural Resources, University of CaliforniaBlog-Article by Kathy Keatley Garvey. It would be interesting to find out;14 replying posts
4 https://beekeepingforum.co.uk/showthread.php?t=33125Lastaccessed: 18 Jan. 2021 UK Beekeeping Forum.Various posts. reporting info of Bohrer, K.,& Pettis, J.S.;10 replying posts
5 https://www.beesource.com/forums/showthread.php?230659-WashboardingLastaccessed: 18 Jan. 2021 Forum community dedicated to beekeeping, bee owners and enthusiasts.6 discussions following a starter post. reporting info of Bohrer, K., & Pettis, J.S.;70 replying posts
6 http://iussi.confex.com/iussi/2006/techprogram/P2650.HTMLastaccessed: 18 Jan. 2021 From the program of an International Conference (IUSSI 2006).Poster presentation related to washboarding behaviour by Bohrer, K., & Pettis, J.S. The function of this behaviour remains to be elucidated, after experimental observations of worker age and surface texture.
7 http://www.myrmecos.net/2010/08/19/washboarding-bees/Lastaccessed: 18 Jan. 2021 Informative website by Alex Wild.Posted video. Description, no hypotheses;13 replying posts
8 https://forum.honeyflow.com/t/theories-on-washboarding/9718Lastaccessed: 18 Jan. 2021 Informative website by a private company.Posts following introduction of topic: Theories on Washboarding Description, + hypotheses;20 replying posts1 video
9 http://www.keltronixinc.com/washboarding-bees/Lastaccessed: 18 Jan. 2021 Informative website by a private company.Posts following introduction of topic: Washboarding Bees Description, + videos, + hypotheses;
10 http://www.beeculture.com/ask-phil-november-2014/Lastaccessed: 18 Jan. 2021 Bee Culture The Magazine of American Beekeeping.Article/post by Phil Craft, November 1, 2014 Responding to post
11 https://forum.canberrabees.com/t/bees-washboarding/200/2Lastaccessed: 18 Jan. 2021 Site dedicated to informing beekeepersPosts following introduction of topic: Bees washboarding Description, + videos;2 replying posts

Hypotheses on reasons behind washboarding behaviour, based on scientific reports in the literature

Available hypotheses explaining washboarding, found in the literature at the end of the 19th-beginning of the 20th century, as cited in Alfonsus (1932)

Mentioned Author(s) Year of report Explanation for washboarding

Dönhoff 1860 playing instinct
Schmid & Kleine 1861 playing instinct
Mehring 1866 wax lining
Kleine 1866 scraping to remove fungi and lichens
Buttel-Reepen 1915 playing instinct
Haidak 1929 scraping
Vahan 1930 petting something
Betts 1931 propelling the content of the honey-sac and exuding other substances
Alfonsus 1932 mechanical cleaning

Available hypotheses explaining washboarding, found in the literature 21st - 22nd century, as cited in Taulmann (2017)

Author(s) Year of report Explanation for washboarding

Gary 1992 cleaning the surface around the hive entrance, removing any loose particles, parasites, fungi, or other debris
Bohrer and Pettis 2006
Garvey 2012 keeping active during scarcity of food sources
Taulmann 2017 applying an attractive scent to the surface around the entrance, to help foragers to orientate

Fv/Fm values (±SD; n=3) in X. parietina after exposure

Sample Fv/Fm Observations
D1 0.22±0.20 depigmentation, presence of propolis on the surface
D2 NA absent
D3 0.22±0.39 depigmentation, presence of propolis on the surface
D4 0.16±0.17 depigmentation
D5 0.15±0.15 depigmentation
D6 0 depigmentation
D7 0.21±0.19 presence of propolis on the surface
D8 0.42±0.13 depigmentation
D9 0.42±0.37 depigmentation, presence of propolis on the surface
D10 0.15±0.13 depigmentation
A1 0.27±0.25 presence of propolis on the surface
A2 0.28±0.08 presence of propolis on the surface
A3 0.06±0.03 depigmentation
A4 0.16±0.13 presence of propolis on the surface
A5 0.39±0.06 presence of propolis on the surface
A6 0.41±0.05 no changes observed
A7 0.15±0.12 depigmentation
A8 0.32±0.29 depigmentation
A9 0.41±0.08 no changes observed
A10 NA absent

YouTube video listNeoSansIntel-Medium

Link to Videos Title /Author /date of post Visualizations /comments
1 https://www.youtube.com/watch?v=lbwumXVTOz8 Bees WashboardingAlexWild7320 Aug. 2010 45.313 / 24
2 https://www.youtube.com/watch?v=sxjc4tSKJFs&feature=emb_logo Washboard BehaviorDr. James E. Tew30 Aug. 2012 28.338 / 69
3 https://www.youtube.com/watch?v=tcJtJdke0AQ Washboarding/BeesDon The Fat Bee Man31 Aug. 2012 19.419 / 10
4 https://www.youtube.com/watch?v=0qncNbsh4_0&t=118s Washboard Movement of Honey Bees (Example and Explanation)Honey Bee Honey13 Nov. 2015 12.023 / 8
5 https://www.youtube.com/watch?reload=9&v=h6oaASApNk4&feature=emb_logo HONEY BEE Apis mellifera washboarding mystery solved?Dr. Tom Seeley is right.insidethehive.tv18 Aug. 2019 9.871 / 37
6 https://www.youtube.com/watch?v=yUDBk9VTypc&t=149s When Honey Bees Do The Washboarding ThingDavid Burns4 Aug. 2016 8756 / 20
7 https://www.youtube.com/watch?v=DBZpa-6rs8A Nectar Dearth, Wash Boarding Bees And Never Getting PoisonIvy AgainJason Chrisman16 Aug. 2020 7.644 / 98
Link to Videos correlating washboarding to propolis distribution Title / Author /date of post Visualizations /comments
8 https://www.youtube.com/watch?v=tXWefJhw2rA&feature=youtu.be Honeybee Propolis and Washboarding is there a connection?18 bees24 Aug. 2020 3.872 / 46
9 https://www.youtube.com/watch?v=y3gUqC694D4 Does bee washboarding on boxed hives mimic propolising in nature?18 bees10 Sept. 2020 2.774 / 27
10 https://www.youtube.com/watch?v=6NRjYNJEb6g Bees WashboardingJossamer Jedi22 Nov. 2015 126 / 0
Link to other interesting Videos Title / Author /date of post Visualizations /comments
11 https://www.youtube.com/watch?v=EavTZt8ybDc Wash boarding beesJustified Bee7 May 2017 0/0
12 https://www.youtube.com/watch?v=1t8JyQTUmSs Bees shimmering or washboardingJohn Haverson17 Aug. 2016 282/0
13 https://www.youtube.com/watch?v=2WIeAd_ATko Bee tree bees wash boarding todayHooterville Honey bees22 Aug. 2020 22/1
14 https://www.youtube.com/watch?v=K6LJyVnqFYU&fbclid=IwAR2FsbjtCHHzZi16cOaXWijeQUnuWdhKhCuJOKRCQDqIsIZqZvc8tKs6JOY Honeybees foraging on lichen? Very interesting honeybeebehaviorSwarmstead Bees & Gardening22 Oct. 2020 362/13

List of identified lichen species present on each bark stripe used during the field experiment

Lichens present (x) on bark stripes collected in Italy in May 2019

A1 A2 A3 A4 A5 A6 A7 A8 A9 A10

Amandinea punctata x x x x
Caloplaca cerina x
Candelaria concolor x x x x x x x x x x
Hyperphyscia adglutinata x x x x x x x x x x
Lecania cyrtella x x x
Lecidella elaeochroma x x x
Phaeophyscia orbicularis x x x x x x
Physcia adscendens x x x x x x x x
Physcia aipolia x
Physcia dubia x
Scoliciosporum sp. x
Xanthoria parietina x x x x x x x x x x

Lichens present (x) on bark stripes collected in Portugal in April 2018

D1 D2 D3 D4 D5 D6 D7 D8 D9 D10

Arthonia radiata x
Hyperphyscia adglutinata x x x x x
Lecanora chlarotera x x
Lecidella elaeochroma x
Pertusaria sp. x x
Physcia erumpens x x x
Xanthoria parietina x x x x x x x x
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