Hygienic Behavior, Liquid-Foraging, and Trophallaxis in the Leaf-Cutting Ants, Acromyrmex subterraneus and Acromyrmex octospinosus Author(s): Freddie-Jeanne Richard and Christine Errard Source: Journal of Insect Science, 9(63):1-9. 2009. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1673/031.009.6301 URL: http://www.bioone.org/doi/full/10.1673/031.009.6301

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

Hygienic behavior, liquid-foraging, and trophallaxis in the leaf- cutting ants, Acromyrmex subterraneus and Acromyrmex octospinosus Freddie-Jeanne Richard1,2 and Christine Errard2 1 University of Poitiers, Laboratoire Ecologie, Evolution, Symbiose, UMR CNRS 6556, 40 avenue du Recteur Pineau, Poitiers, France 2 Institut de Recherche sur la Biologie de l’Insecte, IRBI - UMR CNRS 6035, Faculté des Sciences et Techniques, Université François Rabelais, F-37200 Tours, France

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Keywords: fungus-growing ants, grooming, fungus privation, foragers Abbreviations: LW: large workers, SW: small workers Correspondence: [email protected], Received: 31 October 2007 | Accepted: 15 October 2008 | Published: 3 December 2009 Associate Editor: Michael Breed was editor of this paper. Copyright: This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. ISSN: 1536-2442 | Vol. 9, Number 63 Cite this paper as: Richard F, Errard C. 2009. Hygienic behavior, liquid-foraging, and trophallaxis in the leaf-cutting ants, Acromyrmex subterraneus and Acromyrmex octospinosus. 9pp. Journal of Insect Science 9:63, available online: insectscience.org/9.63

Journal of Insect Science | www.insectscience.org 1 Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

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Journal of Insect Science | www.insectscience.org 2 Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

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Journal of Insect Science | www.insectscience.org 3 Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

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Figure 1. Experimental set-up for the fungus-less worker observations.N : nest area with the fungus, workers and queen and E: foraging area of workers belonging to the nest N; N’: nest area with only workers and E’: the foraging area of workers belonging to the nest N’.

Journal of Insect Science | www.insectscience.org 4 Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

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Table 1. Behaviors displayed by A. subterraneus ants with and without fungus. Data are presented as Mean ± SEM of behavior duration in seconds.

Nest workers With Nest workers Without Foragers (LW only) With fungus fungus fungus Replete Empty All LW (n=40) SW (n=20) LW (n=22) SW (n=10) (n=35) (n=25) (n=60) Self-grooming 12.59±1.22 13.33±3.06 12.9±1.43 8.86±0.66 5.99±0.62 6.53±0.8 7.32±2 Social behaviour 26.78±3.07 13.35±1.53 21.18±2.07 4.44±0.79 4.36±1.77 4.86±0.8 2.98±0.67 Immobility 2.7±0.77 6.15±2.83 4.14±1.25 13.74±2.41 12.69±3.37 32.89±6.37 38.91±8.59 Other activities (including fungus 57.92±3.45 67.16±4.37 61.77±2.73 72.5±2.34 76.99±3.48 58.06±6.10 51.21±7.29 contact)

Table 2. Behaviors displayed by A. octospinosus ants with fungus. Data are presented as Mean ± SEM of behavior duration in seconds.

Foragers (LW only) Nest workers Replete (n=35) Empty (n=25) All (n=60) LW (n=20) SW (n=10) Self-grooming 10.96±0.96 9.68±1.74 10.42±0.90 5.74±0.78 2.78±0.63 Social behaviour 22.92±2.98 14.3±3.91 19.33±2.4 4.47±1.22 1.27±0.69 Immobile 4.78±1.62 6.05±3.22 5.31±1.61 23.52±4.02 24.98±7.97 Other activities (including fungus contact) 61.33±3.93 69.97±5.38 64.93±3.2 66.27±3.65 70.98±7.96

Journal of Insect Science | www.insectscience.org 5 Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

Figure 2. Relative proportion of each behavior (in %) of Acromyrmex subterraneus ants and A. octospinosus returning foragers in the nest. Returning foragers are separated in two groups, “replete” or “empty”. Data are presented as Mean ± SEM of behavior in percentage. Bars above columns represent groups being compared; astericks indicate level of significance, * = p<0.05; *** = p<0.01; *** = p<0.001.

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Journal of Insect Science | www.insectscience.org 6 Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

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C2C6 Acknowledgements  2?5  @7 C6A=6E6 7@C286CD 6?82865 :? EC@A92==2I:D 06 E92?< ,@A9:6 &2C>:8?@? 2?5 ':4@=2D @F:I 7@C C6DA64E:G6=J :? A. subterraneus 2?5 A. octospinosus  4@?4FC E96:C 4@?EC:3FE:@?D E@ E96 @3D6CG2E:@?D &:D492 :;6C@? FCC:6 2?5 #2>6D ! !F?E 7@C E96:C 4@> =2I:D  :? A. subterraneus &@C6:C2 6E 2=  6DA6 >6?ED C?2F5 F36CE 7@C DE2E:DE:42= 25G:D6D 2?5 #24@3FD 4:2==J 4@>A2C65 E@ @E96C DA64:6D DF49 2D  :? # @@>D>2 2?5 E96 ,>:E9D@?:2? -C@A:42= +6D62C49 "? Acromyrmex subterraneus subterraneus $F2 2?5 E96 FE@C:525 '24:@?2= 56= >3: &@C6@G6C :? E96 AC6D6?E 6IA6C:>6?ED E96 >62? E:>6 @7 6?E6 J 6= &2C '& 7@C A6C>:DD:@? E@ D2>A=6 2?E E96 EC@A92==2I:D H2D 23@FE  D64@?5D 7@C 3@E9 DA64:6D 4@=@?:6D @7 Acromyrmex octospinosus :? )2?2>2 2?5 6IA@CE 4@>A2C65 E@  D64@?5D C6A@CE65 :? &@C:6C2 6E 2= E96> E@ FC@A6 06 E92?< E96 FE@C:525 '24:@?2= 56=   >3:6?E6 '& 2?5 -6C6K:?92 6==2 %F4:2 ), ( .   4@?G6?E:@? 7@C A6C>:DD:@? E@ 4@==64E -96 =@H 7C6BF6?4J @7 EC@A92==2I:D 36EH66? ?@?7@C286CD 2?E 4@=@?:6 Acromyrmex subterraneus :? C2K:= 06 E92?< 92D 2? :?E6C6DE:?8 C6=2E:@?D9:A E@ E96 AC6D6?46 @7 E96 :==6D @DBF6E 7@C E96 56D:8? @7 :8FC6 06 2C6 8C2E67F= 7F?8FD 82C56? -96 7F?8FD H9:49 :D 2? 6DD6?E:2= D@FC46 E@
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 8 = 2?5

Journal of Insect Science | www.insectscience.org 7 Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

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762EFC6D Ecology     ?5C256 )) @CE: % &@C6:C2  @2C6EE@ & +2>@D /& &2E@D ( 692G:@C @7 Atta sexdens rubropilosa !@H2C5 ## 0:6>6C   96>:42= 64@=@8J @7 9@DE A=2?E D6=64E:@? !J>6?@AE6C2 @C>:4:526 H@C<6CD 5FC:?8 E96 AC6A2C2E:@? @7 E96 3J E96 =6274FEE:?8 2?E Atta cephalotes "? %@78C6? , /2?56C =627
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2?E Acromyrmex subterraneus brunneus Sociobiology     E96 A@?6C:?6 2?E Ponera coarctata :>A=:42E:@?D 7@C E96 6G@=FE:@? @7 =:BF:5
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FCC:6 + 3 )C6G2=6?46 2?5 :>A24E @7 2 G:CF=6?E A2C2D:E6 @? 2 &6D<2=: & @?2G:E2@F8@FC52?  )C@G@DE  28?6C6D   FDE:4: EC:A2CE:E6
>FEF2=:D> Oecologia    6C  =6>6?E #%  &6492?:D> F?56C=J:?8 4FE:4F=2C 9J5C@ 42C3@? 9@>@86?6:EJ :? E96 2?E Camponotus vagus ,4@A  !J>6?@AE6C2 @C>:4:526 +@=6 @7 A@DEA92CJ?862= 8=2?5D Journ- FCC:6 + &F6==6C .  &2==@49  2 -96 28C:4F=EFC6 A2E9@=@8J al of Chemical Ecology    @7 2?E 7F?8FD 82C56? Proceedings of the National Academy of Sciences   &@C6:C2 ( CE92= #& 2CC6C2 &) ,:=G2 ) ,2>F6=D +"  (C2= EC@A92==2I:D :? 25F=E =6274FEE:?8 2?ED Acromyrmex subterraneus FCC:6 + ,4@EE # ,F>>6C36== + &2==@49  3 F?8FD subterraneus Insectes sociaux 8C@H:?8 2?ED FD6 2?E:3:@E:4 324E6C:2 E@ 4@?EC@= 82C56? A2C2D:E6D !J>6?@AE6C2
@C>:4:526   Nature     &FC2<2>: - !:82D9: ,  ,@4:2= @C82?:K2E:@? :? EH@ AC:>:E:G6 2E Cyphomyrmex rimosus Myrmicocrypta ednaella FCC:6 + ,EF2CE  0665:?8 2?5 8C@@>:?8 @7 A2E9@86?D :? E:?6 2?ED 2?5  H:E9 C676C Journal of Ethology 28C:4F=EFC6 3J 2?ED The Royal Society of London Series B  6?46 E@ E96:C 7F?8FD DF3DEC2E6D 2?5 7@@5 D@FC46D     

293:  !676EK  6C52 1 %6?@:C   -C@A92==2I:D >65:2E6D ':6=D6? # @@>D>2 ## (=592> '# )6E6CD6? ! &@C82?   F?:7@C>:EJ @7 4@=@?J @5@C :? Cataglyphis iberica 2?ED !J>6?@AE6C2 @=@?J=6G6= 2?5 D62D@?DA64:7:4 G2C:2E:@? :? 4FE:4F=2C 9J5C@42C Formica truncorum In- @C>:4:526 Journal of Insect Behavior   3@? AC@7:=6D @7 :?5:G:5F2= H@C<6CD :? E96 2?E sectes Sociaux   =6E496C # &:496?6C   Kin recognition in animals #@9? 0:=6J '@CE9 + #24A2CE>6?E2=:D2E:@? 4@==64E:G6=J :D@=2E6 92K2C5@FD H2DE6 :? E96 =6274FEE:?8 2?E Atta cephalotes Behavioral Ecology and Sociobiology )@F=D6? & @@>D>2 ##  &FEF2=:DE:4 7F?8: 4@?EC@= 4C@A 5:G6CD:EJ Science   :?
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Journal of Insect Science | www.insectscience.org 8 Journal of Insect Science: Vol. 9 | Article 63 Richard et al.

+:492C5 # )@F=D6? & C:;79@FE  #@?6D + @@>D>2 ## 2 -C2?:6==@ #  +@D6?82FD + ,2G@:6 $ -96 56G6=@A>6?E @7 ,A64:7:4:EJ :? 496>:42= AC@7:=6D @7 H@C<6CD 3C@@5 2?5 >FEF2=:DE:4 :>>F?:EJ :? 2 D@4:2= :?D64E 6G:56?46 7@C E96 8C@FA 724:=:E2E:@? @7 7F?8: :? Atta Acromyrmex 2?5 Sericomyrmex 7F?8FD8C@H:?8 2?ED 5:D62D6 C6D:DE2?46 Proceedings of the National Academy of Sciences of the Journal of Chemical Ecology     United States of America  

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Journal of Insect Science | www.insectscience.org 9