---
_id: '616'
abstract:
- lang: eng
text: Social insects protect their colonies from infectious disease through collective
defences that result in social immunity. In ants, workers first try to prevent
infection of colony members. Here, we show that if this fails and a pathogen establishes
an infection, ants employ an efficient multicomponent behaviour − "destructive
disinfection" − to prevent further spread of disease through the colony.
Ants specifically target infected pupae during the pathogen's non-contagious incubation
period, relying on chemical 'sickness cues' emitted by pupae. They then remove
the pupal cocoon, perforate its cuticle and administer antimicrobial poison, which
enters the body and prevents pathogen replication from the inside out. Like the
immune system of a body that specifically targets and eliminates infected cells,
this social immunity measure sacrifices infected brood to stop the pathogen completing
its lifecycle, thus protecting the rest of the colony. Hence, the same principles
of disease defence apply at different levels of biological organisation.
article_number: e32073
article_processing_charge: Yes
author:
- first_name: Christopher
full_name: Pull, Christopher
id: 3C7F4840-F248-11E8-B48F-1D18A9856A87
last_name: Pull
orcid: 0000-0003-1122-3982
- first_name: Line V
full_name: Ugelvig, Line V
id: 3DC97C8E-F248-11E8-B48F-1D18A9856A87
last_name: Ugelvig
orcid: 0000-0003-1832-8883
- first_name: Florian
full_name: Wiesenhofer, Florian
id: 39523C54-F248-11E8-B48F-1D18A9856A87
last_name: Wiesenhofer
- first_name: Anna V
full_name: Grasse, Anna V
id: 406F989C-F248-11E8-B48F-1D18A9856A87
last_name: Grasse
- first_name: Simon
full_name: Tragust, Simon
id: 35A7A418-F248-11E8-B48F-1D18A9856A87
last_name: Tragust
- first_name: Thomas
full_name: Schmitt, Thomas
last_name: Schmitt
- first_name: Mark
full_name: Brown, Mark
last_name: Brown
- first_name: Sylvia
full_name: Cremer, Sylvia
id: 2F64EC8C-F248-11E8-B48F-1D18A9856A87
last_name: Cremer
orcid: 0000-0002-2193-3868
citation:
ama: Pull C, Ugelvig LV, Wiesenhofer F, et al. Destructive disinfection of infected
brood prevents systemic disease spread in ant colonies. eLife. 2018;7.
doi:10.7554/eLife.32073
apa: Pull, C., Ugelvig, L. V., Wiesenhofer, F., Grasse, A. V., Tragust, S., Schmitt,
T., … Cremer, S. (2018). Destructive disinfection of infected brood prevents systemic
disease spread in ant colonies. ELife. eLife Sciences Publications. https://doi.org/10.7554/eLife.32073
chicago: Pull, Christopher, Line V Ugelvig, Florian Wiesenhofer, Anna V Grasse,
Simon Tragust, Thomas Schmitt, Mark Brown, and Sylvia Cremer. “Destructive Disinfection
of Infected Brood Prevents Systemic Disease Spread in Ant Colonies.” ELife.
eLife Sciences Publications, 2018. https://doi.org/10.7554/eLife.32073.
ieee: C. Pull et al., “Destructive disinfection of infected brood prevents
systemic disease spread in ant colonies,” eLife, vol. 7. eLife Sciences
Publications, 2018.
ista: Pull C, Ugelvig LV, Wiesenhofer F, Grasse AV, Tragust S, Schmitt T, Brown
M, Cremer S. 2018. Destructive disinfection of infected brood prevents systemic
disease spread in ant colonies. eLife. 7, e32073.
mla: Pull, Christopher, et al. “Destructive Disinfection of Infected Brood Prevents
Systemic Disease Spread in Ant Colonies.” ELife, vol. 7, e32073, eLife
Sciences Publications, 2018, doi:10.7554/eLife.32073.
short: C. Pull, L.V. Ugelvig, F. Wiesenhofer, A.V. Grasse, S. Tragust, T. Schmitt,
M. Brown, S. Cremer, ELife 7 (2018).
corr_author: '1'
date_created: 2018-12-11T11:47:31Z
date_published: 2018-01-09T00:00:00Z
date_updated: 2026-04-08T14:19:10Z
day: '09'
ddc:
- '570'
- '590'
department:
- _id: SyCr
doi: 10.7554/eLife.32073
ec_funded: 1
external_id:
isi:
- '000419601300001'
file:
- access_level: open_access
checksum: 540f941e8d3530a9441e4affd94f07d7
content_type: application/pdf
creator: system
date_created: 2018-12-12T10:10:43Z
date_updated: 2020-07-14T12:47:20Z
file_id: '4832'
file_name: IST-2018-978-v1+1_elife-32073-v1.pdf
file_size: 1435585
relation: main_file
file_date_updated: 2020-07-14T12:47:20Z
fulldoi: https://doi.org/10.7554/eLife.32073
has_accepted_license: '1'
intvolume: ' 7'
isi: 1
language:
- iso: eng
month: '01'
oa: 1
oa_version: Published Version
project:
- _id: 25DC711C-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '243071'
name: 'Social Vaccination in Ant Colonies: from Individual Mechanisms to Society
Effects'
- _id: 25DDF0F0-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '302004'
name: 'Collective disease defence and pathogen detection abilities in ant societies:
a chemo-neuro-immunological approach'
publication: eLife
publication_status: published
publisher: eLife Sciences Publications
publist_id: '7188'
pubrep_id: '978'
quality_controlled: '1'
related_material:
record:
- id: '819'
relation: dissertation_contains
status: public
scopus_import: '1'
status: public
title: Destructive disinfection of infected brood prevents systemic disease spread
in ant colonies
tmp:
image: /images/cc_by.png
legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
short: CC BY (4.0)
type: journal_article
user_id: c635000d-4b10-11ee-a964-aac5a93f6ac1
volume: 7
year: '2018'
...
---
_id: '1830'
abstract:
- lang: eng
text: To prevent epidemics, insect societies have evolved collective disease defences
that are highly effective at curing exposed individuals and limiting disease transmission
to healthy group members. Grooming is an important sanitary behaviour—either performed
towards oneself (self-grooming) or towards others (allogrooming)—to remove infectious
agents from the body surface of exposed individuals, but at the risk of disease
contraction by the groomer. We use garden ants (Lasius neglectus) and the fungal
pathogen Metarhizium as a model system to study how pathogen presence affects
self-grooming and allogrooming between exposed and healthy individuals. We develop
an epidemiological SIS model to explore how experimentally observed grooming patterns
affect disease spread within the colony, thereby providing a direct link between
the expression and direction of sanitary behaviours, and their effects on colony-level
epidemiology. We find that fungus-exposed ants increase self-grooming, while simultaneously
decreasing allogrooming. This behavioural modulation seems universally adaptive
and is predicted to contain disease spread in a great variety of host–pathogen
systems. In contrast, allogrooming directed towards pathogen-exposed individuals
might both increase and decrease disease risk. Our model reveals that the effect
of allogrooming depends on the balance between pathogen infectiousness and efficiency
of social host defences, which are likely to vary across host–pathogen systems.
acknowledgement: We thank Meghan L. Vyleta for the genetical fungal strain characterization
and Eva Sixt for ant drawings, Matthias Konrad for discussion and Christopher D.
Pull, Barbara Casillas-Peréz, Sebastian Novak, as well as three anonymous reviewers
and the theme issue editors Peter Kappeler and Charlie Nunn for valuable comments
on the manuscript.
article_processing_charge: No
article_type: original
author:
- first_name: Fabian
full_name: Theis, Fabian
last_name: Theis
- first_name: Line V
full_name: Ugelvig, Line V
id: 3DC97C8E-F248-11E8-B48F-1D18A9856A87
last_name: Ugelvig
orcid: 0000-0003-1832-8883
- first_name: Carsten
full_name: Marr, Carsten
last_name: Marr
- first_name: Sylvia
full_name: Cremer, Sylvia
id: 2F64EC8C-F248-11E8-B48F-1D18A9856A87
last_name: Cremer
orcid: 0000-0002-2193-3868
citation:
ama: Theis F, Ugelvig LV, Marr C, Cremer S. Opposing effects of allogrooming on
disease transmission in ant societies. Philosophical Transactions of the Royal
Society of London Series B, Biological Sciences. 2015;370(1669). doi:10.1098/rstb.2014.0108
apa: Theis, F., Ugelvig, L. V., Marr, C., & Cremer, S. (2015). Opposing effects
of allogrooming on disease transmission in ant societies. Philosophical Transactions
of the Royal Society of London. Series B, Biological Sciences. Royal Society.
https://doi.org/10.1098/rstb.2014.0108
chicago: Theis, Fabian, Line V Ugelvig, Carsten Marr, and Sylvia Cremer. “Opposing
Effects of Allogrooming on Disease Transmission in Ant Societies.” Philosophical
Transactions of the Royal Society of London. Series B, Biological Sciences.
Royal Society, 2015. https://doi.org/10.1098/rstb.2014.0108.
ieee: F. Theis, L. V. Ugelvig, C. Marr, and S. Cremer, “Opposing effects of allogrooming
on disease transmission in ant societies,” Philosophical Transactions of the
Royal Society of London. Series B, Biological Sciences, vol. 370, no. 1669.
Royal Society, 2015.
ista: Theis F, Ugelvig LV, Marr C, Cremer S. 2015. Opposing effects of allogrooming
on disease transmission in ant societies. Philosophical Transactions of the Royal
Society of London. Series B, Biological Sciences. 370(1669).
mla: Theis, Fabian, et al. “Opposing Effects of Allogrooming on Disease Transmission
in Ant Societies.” Philosophical Transactions of the Royal Society of London.
Series B, Biological Sciences, vol. 370, no. 1669, Royal Society, 2015, doi:10.1098/rstb.2014.0108.
short: F. Theis, L.V. Ugelvig, C. Marr, S. Cremer, Philosophical Transactions of
the Royal Society of London. Series B, Biological Sciences 370 (2015).
corr_author: '1'
date_created: 2018-12-11T11:54:15Z
date_published: 2015-05-26T00:00:00Z
date_updated: 2026-08-12T06:27:32Z
day: '26'
department:
- _id: SyCr
doi: 10.1098/rstb.2014.0108
ec_funded: 1
external_id:
isi:
- '000354143400010'
pmid:
- '25870394'
fulldoi: https://doi.org/10.1098/rstb.2014.0108
intvolume: ' 370'
isi: 1
issue: '1669'
language:
- iso: eng
main_file_link:
- open_access: '1'
url: http://www.ncbi.nlm.nih.gov/pmc/articles/PMC4410374/
month: '05'
oa: 1
oa_version: Submitted Version
pmid: 1
project:
- _id: 25DC711C-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '243071'
name: 'Social Vaccination in Ant Colonies: from Individual Mechanisms to Society
Effects'
- _id: 25DDF0F0-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '302004'
name: 'Collective disease defence and pathogen detection abilities in ant societies:
a chemo-neuro-immunological approach'
- _id: 25E0E184-B435-11E9-9278-68D0E5697425
name: Schnellboot Antnet Junge Akademie
- _id: 25E24DB2-B435-11E9-9278-68D0E5697425
name: Fellowship of Wissenschaftskolleg zu Berlin
publication: Philosophical Transactions of the Royal Society of London. Series B,
Biological Sciences
publication_identifier:
eissn:
- 1471-2970
issn:
- 0962-8436
publication_status: published
publisher: Royal Society
publist_id: '5273'
quality_controlled: '1'
related_material:
record:
- id: '9721'
relation: research_data
status: public
scopus_import: '1'
status: public
title: Opposing effects of allogrooming on disease transmission in ant societies
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 370
year: '2015'
...
---
_id: '2926'
abstract:
- lang: eng
text: To fight infectious diseases, host immune defenses are employed at multiple
levels. Sanitary behavior, such as pathogen avoidance and removal, acts as a first
line of defense to prevent infection [1] before activation of the physiological
immune system. Insect societies have evolved a wide range of collective hygiene
measures and intensive health care toward pathogen-exposed group members [2].
One of the most common behaviors is allogrooming, in which nestmates remove infectious
particles from the body surfaces of exposed individuals [3]. Here we show that,
in invasive garden ants, grooming of fungus-exposed brood is effective beyond
the sheer mechanical removal of fungal conidiospores; it also includes chemical
disinfection through the application of poison produced by the ants themselves.
Formic acid is the main active component of the poison. It inhibits fungal growth
of conidiospores remaining on the brood surface after grooming and also those
collected in the mouth of the grooming ant. This dual function is achieved by
uptake of the poison droplet into the mouth through acidopore self-grooming and
subsequent application onto the infectious brood via brood grooming. This extraordinary
behavior extends the current understanding of grooming and the establishment of
social immunity in insect societies.
acknowledgement: "Funding for this project was obtained by the German Research Foundation
(DFG, to S.C.) and the European Research Council (ERC, through an ERC-Starting Grant
to S.C. and an Individual Marie Curie IEF fellowship to L.V.U.).\r\nWe thank Jørgen
Eilenberg, Bernhardt Steinwender, Miriam Stock, and Meghan L. Vyleta for the fungal
strain and its characterization; Volker Witte for chemical information; Eva Sixt
for ant drawings; and Robert Hauschild for help with image analysis. We further
thank Martin Kaltenpoth, Michael Sixt, Jürgen Heinze, and Joachim Ruther for discussion
and Daria Siekhaus, Sophie A.O. Armitage, and Leila Masri for comments on the manuscript.
\r\n"
article_processing_charge: No
author:
- first_name: Simon
full_name: Tragust, Simon
id: 35A7A418-F248-11E8-B48F-1D18A9856A87
last_name: Tragust
- first_name: Barbara
full_name: Mitteregger, Barbara
id: 479DDAAC-E9CD-11E9-9B5F-82450873F7A1
last_name: Mitteregger
- first_name: Vanessa
full_name: Barone, Vanessa
id: 419EECCC-F248-11E8-B48F-1D18A9856A87
last_name: Barone
orcid: 0000-0003-2676-3367
- first_name: Matthias
full_name: Konrad, Matthias
id: 46528076-F248-11E8-B48F-1D18A9856A87
last_name: Konrad
- first_name: Line V
full_name: Ugelvig, Line V
id: 3DC97C8E-F248-11E8-B48F-1D18A9856A87
last_name: Ugelvig
orcid: 0000-0003-1832-8883
- first_name: Sylvia
full_name: Cremer, Sylvia
id: 2F64EC8C-F248-11E8-B48F-1D18A9856A87
last_name: Cremer
orcid: 0000-0002-2193-3868
citation:
ama: Tragust S, Mitteregger B, Barone V, Konrad M, Ugelvig LV, Cremer S. Ants disinfect
fungus-exposed brood by oral uptake and spread of their poison. Current Biology.
2013;23(1):76-82. doi:10.1016/j.cub.2012.11.034
apa: Tragust, S., Mitteregger, B., Barone, V., Konrad, M., Ugelvig, L. V., &
Cremer, S. (2013). Ants disinfect fungus-exposed brood by oral uptake and spread
of their poison. Current Biology. Cell Press. https://doi.org/10.1016/j.cub.2012.11.034
chicago: Tragust, Simon, Barbara Mitteregger, Vanessa Barone, Matthias Konrad, Line
V Ugelvig, and Sylvia Cremer. “Ants Disinfect Fungus-Exposed Brood by Oral Uptake
and Spread of Their Poison.” Current Biology. Cell Press, 2013. https://doi.org/10.1016/j.cub.2012.11.034.
ieee: S. Tragust, B. Mitteregger, V. Barone, M. Konrad, L. V. Ugelvig, and S. Cremer,
“Ants disinfect fungus-exposed brood by oral uptake and spread of their poison,”
Current Biology, vol. 23, no. 1. Cell Press, pp. 76–82, 2013.
ista: Tragust S, Mitteregger B, Barone V, Konrad M, Ugelvig LV, Cremer S. 2013.
Ants disinfect fungus-exposed brood by oral uptake and spread of their poison.
Current Biology. 23(1), 76–82.
mla: Tragust, Simon, et al. “Ants Disinfect Fungus-Exposed Brood by Oral Uptake
and Spread of Their Poison.” Current Biology, vol. 23, no. 1, Cell Press,
2013, pp. 76–82, doi:10.1016/j.cub.2012.11.034.
short: S. Tragust, B. Mitteregger, V. Barone, M. Konrad, L.V. Ugelvig, S. Cremer,
Current Biology 23 (2013) 76–82.
corr_author: '1'
date_created: 2018-12-11T12:00:23Z
date_published: 2013-01-07T00:00:00Z
date_updated: 2026-04-08T14:22:39Z
day: '07'
department:
- _id: SyCr
- _id: CaHe
doi: 10.1016/j.cub.2012.11.034
ec_funded: 1
external_id:
isi:
- '000313383700026'
fulldoi: https://doi.org/10.1016/j.cub.2012.11.034
intvolume: ' 23'
isi: 1
issue: '1'
language:
- iso: eng
month: '01'
oa_version: None
page: 76 - 82
project:
- _id: 25DAF0B2-B435-11E9-9278-68D0E5697425
grant_number: CR-118/3-1
name: Host-Parasite Coevolution
- _id: 25DC711C-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '243071'
name: 'Social Vaccination in Ant Colonies: from Individual Mechanisms to Society
Effects'
- _id: 25DDF0F0-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '302004'
name: 'Collective disease defence and pathogen detection abilities in ant societies:
a chemo-neuro-immunological approach'
publication: Current Biology
publication_status: published
publisher: Cell Press
publist_id: '3811'
quality_controlled: '1'
related_material:
record:
- id: '9757'
relation: research_data
status: public
- id: '961'
relation: dissertation_contains
status: public
scopus_import: '1'
status: public
title: Ants disinfect fungus-exposed brood by oral uptake and spread of their poison
type: journal_article
user_id: 317138e5-6ab7-11ef-aa6d-ffef3953e345
volume: 23
year: '2013'
...
---
_id: '2284'
abstract:
- lang: eng
text: 'Background: The brood of ants and other social insects is highly susceptible
to pathogens, particularly those that penetrate the soft larval and pupal cuticle.
We here test whether the presence of a pupal cocoon, which occurs in some ant
species but not in others, affects the sanitary brood care and fungal infection
patterns after exposure to the entomopathogenic fungus Metarhizium brunneum. We
use a) a comparative approach analysing four species with either naked or cocooned
pupae and b) a within-species analysis of a single ant species, in which both
pupal types co-exist in the same colony. Results: We found that the presence of
a cocoon did not compromise fungal pathogen detection by the ants and that species
with cocooned pupae increased brood grooming after pathogen exposure. All tested
ant species further removed brood from their nests, which was predominantly expressed
towards larvae and naked pupae treated with the live fungal pathogen. In contrast,
cocooned pupae exposed to live fungus were not removed at higher rates than cocooned
pupae exposed to dead fungus or a sham control. Consistent with this, exposure
to the live fungus caused high numbers of infections and fungal outgrowth in larvae
and naked pupae, but not in cocooned pupae. Moreover, the ants consistently removed
the brood prior to fungal outgrowth, ensuring a clean brood chamber. Conclusion:
Our study suggests that the pupal cocoon has a protective effect against fungal
infection, causing an adaptive change in sanitary behaviours by the ants. It further
demonstrates that brood removal-originally described for honeybees as "hygienic
behaviour"-is a widespread sanitary behaviour in ants, which likely has important
implications on disease dynamics in social insect colonies.'
acknowledgement: "The study was funded by the European Research Council (Marie Curie
ERG 036569) and Marie Curie IEF 302204 to LVU\r\nCC BY 2.0\r\n"
article_number: '225'
article_processing_charge: No
author:
- first_name: Simon
full_name: Tragust, Simon
id: 35A7A418-F248-11E8-B48F-1D18A9856A87
last_name: Tragust
- first_name: Line V
full_name: Ugelvig, Line V
id: 3DC97C8E-F248-11E8-B48F-1D18A9856A87
last_name: Ugelvig
orcid: 0000-0003-1832-8883
- first_name: Michel
full_name: Chapuisat, Michel
last_name: Chapuisat
- first_name: Jürgen
full_name: Heinze, Jürgen
last_name: Heinze
- first_name: Sylvia
full_name: Cremer, Sylvia
id: 2F64EC8C-F248-11E8-B48F-1D18A9856A87
last_name: Cremer
orcid: 0000-0002-2193-3868
citation:
ama: Tragust S, Ugelvig LV, Chapuisat M, Heinze J, Cremer S. Pupal cocoons affect
sanitary brood care and limit fungal infections in ant colonies. BMC Evolutionary
Biology. 2013;13(1). doi:10.1186/1471-2148-13-225
apa: Tragust, S., Ugelvig, L. V., Chapuisat, M., Heinze, J., & Cremer, S. (2013).
Pupal cocoons affect sanitary brood care and limit fungal infections in ant colonies.
BMC Evolutionary Biology. BioMed Central. https://doi.org/10.1186/1471-2148-13-225
chicago: Tragust, Simon, Line V Ugelvig, Michel Chapuisat, Jürgen Heinze, and Sylvia
Cremer. “Pupal Cocoons Affect Sanitary Brood Care and Limit Fungal Infections
in Ant Colonies.” BMC Evolutionary Biology. BioMed Central, 2013. https://doi.org/10.1186/1471-2148-13-225.
ieee: S. Tragust, L. V. Ugelvig, M. Chapuisat, J. Heinze, and S. Cremer, “Pupal
cocoons affect sanitary brood care and limit fungal infections in ant colonies,”
BMC Evolutionary Biology, vol. 13, no. 1. BioMed Central, 2013.
ista: Tragust S, Ugelvig LV, Chapuisat M, Heinze J, Cremer S. 2013. Pupal cocoons
affect sanitary brood care and limit fungal infections in ant colonies. BMC Evolutionary
Biology. 13(1), 225.
mla: Tragust, Simon, et al. “Pupal Cocoons Affect Sanitary Brood Care and Limit
Fungal Infections in Ant Colonies.” BMC Evolutionary Biology, vol. 13,
no. 1, 225, BioMed Central, 2013, doi:10.1186/1471-2148-13-225.
short: S. Tragust, L.V. Ugelvig, M. Chapuisat, J. Heinze, S. Cremer, BMC Evolutionary
Biology 13 (2013).
corr_author: '1'
das_tickbox: '1'
date_created: 2018-12-11T11:56:46Z
date_published: 2013-10-14T00:00:00Z
date_updated: 2026-07-28T09:35:11Z
day: '14'
ddc:
- '570'
department:
- _id: SyCr
doi: 10.1186/1471-2148-13-225
ec_funded: 1
external_id:
isi:
- '000326620500001'
file:
- access_level: open_access
checksum: c16ef36f2a10786a7885e19c4528d707
content_type: application/pdf
creator: system
date_created: 2018-12-12T10:13:41Z
date_updated: 2020-07-14T12:45:37Z
file_id: '5026'
file_name: IST-2016-402-v1+1_1471-2148-13-225.pdf
file_size: 281736
relation: main_file
file_date_updated: 2020-07-14T12:45:37Z
fulldoi: https://doi.org/10.1186/1471-2148-13-225
has_accepted_license: '1'
intvolume: ' 13'
isi: 1
issue: '1'
language:
- iso: eng
month: '10'
oa: 1
oa_version: Published Version
project:
- _id: 25DC711C-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '243071'
name: 'Social Vaccination in Ant Colonies: from Individual Mechanisms to Society
Effects'
- _id: 25DAF0B2-B435-11E9-9278-68D0E5697425
grant_number: CR-118/3-1
name: Host-Parasite Coevolution
- _id: 25DDF0F0-B435-11E9-9278-68D0E5697425
call_identifier: FP7
grant_number: '302004'
name: 'Collective disease defence and pathogen detection abilities in ant societies:
a chemo-neuro-immunological approach'
publication: BMC Evolutionary Biology
publication_status: published
publisher: BioMed Central
publist_id: '4647'
pubrep_id: '402'
quality_controlled: '1'
related_material:
record:
- id: '9753'
relation: research_data
status: public
scopus_import: '1'
status: public
title: Pupal cocoons affect sanitary brood care and limit fungal infections in ant
colonies
tmp:
image: /images/cc_by.png
legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
short: CC BY (4.0)
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 13
year: '2013'
...