Currie CR, Stuart AE (2001) Weeding and grooming of pathogens in agriculture by ants. Proc Biol
Sci 268(1471):1033–1039. https://doi.org/10.1098/rspb.2001.1605
De Fine Licht HH, Boomsma JJ, Tunlid A (2014) Symbiotic adaptations in the fungal cultivar of
leaf-cutting ants. Nat Commun 5:5675. https://doi.org/10.1038/ncomms6675
de Man TJ, Stajich JE, Kubicek CP, Teiling C, Chenthamara K, Atanasova L, Druzhinina IS,
Levenkova N, Birnbaum SS, Barribeau SM, Bozick BA, Suen G, Currie CR, Gerardo NM
(2016) Small genome of the fungus Escovopsis weberi, a specialized disease agent of ant
agriculture. Proc Natl Acad Sci U S A 113(13):3567–3572. https://doi.org/10.1073/pnas.
1518501113
Devine R, Hutchings MI, Holmes NA (2017) Future directions for the discovery of antibiotics from
actinomycete bacteria. Emerg Topics Life Sci 1(1):1–12. https://doi.org/10.1042/etls20160014
Dias DA, Urban S, Roessner U (2012) A historical overview of natural products in drug discovery.
Meta 2(2):303–336. https://doi.org/10.3390/metabo2020303
Engl T, Kroiss J, Kai M, Nechitaylo TY, Svatos A, Kaltenpoth M (2018) Evolutionary stability of
antibiotic protection in a defensive symbiosis. Proc Natl Acad Sci U S A 115(9):E2020–E2029.
https://doi.org/10.1073/pnas.1719797115
Fitzpatrick CR, Copeland J, Wang PW, Guttman DS, Kotanen PM, Johnson MTJ (2018) Assembly
and ecological function of the root microbiome across angiosperm plant species. Proc Natl Acad
Sci U S A 115(6):E1157–E1165. https://doi.org/10.1073/pnas.1717617115
Fleming A (1929) On the antibacterial action of cultures of a Penicillium, with special reference to
their use in the isolation of B. influenzæ. Br J Exp Pathol 10(3):226–236
Florez LV, Biedermann PH, Engl T, Kaltenpoth M (2015) Defensive symbioses of animals with
prokaryotic and eukaryotic microorganisms. Nat Prod Rep 32(7):904–936. https://doi.org/10.
1039/c5np00010f
Foster KR, Schluter J, Coyte KZ, Rakoff-Nahoum S (2017) The evolution of the host microbiome
as an ecosystem on a leash. Nature 548(7665):43–51. https://doi.org/10.1038/nature23292
Gil-Turnes MS, Fenical W (1992) Embryos of Homarus americanus are protected by epibiotic
bacteria. Biol Bull 182(1):105–108. https://doi.org/10.2307/1542184
Gil-Turnes MS, Hay ME, Fenical W (1989) Symbiotic marine bacteria chemically defend crustacean embryos from a pathogenic fungus. Science 246(4926):116–118
Gould K (2016) Antibiotics: from prehistory to the present day. J Antimicrob Chemother 71
(3):572–575. https://doi.org/10.1093/jac/dkv484
Haeder S, Wirth R, Herz H, Spiteller D (2009) Candicidin-producing Streptomyces support leafcutting ants to protect their fungus garden against the pathogenic fungus Escovopsis. Proc Natl
Acad Sci U S A 106(12):4742–4746
Hartmann A, Schmid M, van Tuinen D, Berg G (2009) Plant-driven selection of microbes. Plant
Soil 321(1–2):235–257. https://doi.org/10.1007/s11104-008-9814-y
Heine D, Holmes NA, Worsley SF, Santos ACA, Innocent TM, Scherlach K, Patrick EH, Yu DW,
Murrell JC, Vieria PC, Boomsma JJ, Hertweck C, Hutchings MI, Wilkinson B (2018) Chemical
warfare between leafcutter ant symbionts and a co-evolved pathogen. Nat Commun 9(1):2208.
https://doi.org/10.1038/s41467-018-04520-1
Hooper LV, Littman DR, Macpherson AJ (2012) Interactions between the microbiota and the
immune system. Science 336(6086):1268–1273. https://doi.org/10.1126/science.1223490
Hopwood DA (2007) Streptomyces in nature and medicine: the antibiotic makers. Oxford University Press, New York
Kaltenpoth M (2009) Actinobacteria as mutualists: general healthcare for insects? Trends Microbiol
17(12):529–535
Kaltenpoth M, Goettler W, Dale C, Stubblefield JW, Herzner G, Roeser-Mueller K, Strohm E
(2006) ‘Candidatus Streptomyces philanthi’, an endosymbiotic streptomycete in the antennae of
Philanthus digger wasps. Int J Syst Evol Microbiol 56(6):1403–1411
Kaltenpoth M, Goettler W, Koehler S, Strohm E (2010) Life cycle and population dynamics of a
protective insect symbiont reveal severe bottlenecks during vertical transmission. Evol Ecol 24
(2):463–477. https://doi.org/10.1007/s10682-009-9319-z
23 Defensive Microbiomes: A Widespread Phenomenon in Nature
509
Sci 268(1471):1033–1039. https://doi.org/10.1098/rspb.2001.1605
De Fine Licht HH, Boomsma JJ, Tunlid A (2014) Symbiotic adaptations in the fungal cultivar of
leaf-cutting ants. Nat Commun 5:5675. https://doi.org/10.1038/ncomms6675
de Man TJ, Stajich JE, Kubicek CP, Teiling C, Chenthamara K, Atanasova L, Druzhinina IS,
Levenkova N, Birnbaum SS, Barribeau SM, Bozick BA, Suen G, Currie CR, Gerardo NM
(2016) Small genome of the fungus Escovopsis weberi, a specialized disease agent of ant
agriculture. Proc Natl Acad Sci U S A 113(13):3567–3572. https://doi.org/10.1073/pnas.
1518501113
Devine R, Hutchings MI, Holmes NA (2017) Future directions for the discovery of antibiotics from
actinomycete bacteria. Emerg Topics Life Sci 1(1):1–12. https://doi.org/10.1042/etls20160014
Dias DA, Urban S, Roessner U (2012) A historical overview of natural products in drug discovery.
Meta 2(2):303–336. https://doi.org/10.3390/metabo2020303
Engl T, Kroiss J, Kai M, Nechitaylo TY, Svatos A, Kaltenpoth M (2018) Evolutionary stability of
antibiotic protection in a defensive symbiosis. Proc Natl Acad Sci U S A 115(9):E2020–E2029.
https://doi.org/10.1073/pnas.1719797115
Fitzpatrick CR, Copeland J, Wang PW, Guttman DS, Kotanen PM, Johnson MTJ (2018) Assembly
and ecological function of the root microbiome across angiosperm plant species. Proc Natl Acad
Sci U S A 115(6):E1157–E1165. https://doi.org/10.1073/pnas.1717617115
Fleming A (1929) On the antibacterial action of cultures of a Penicillium, with special reference to
their use in the isolation of B. influenzæ. Br J Exp Pathol 10(3):226–236
Florez LV, Biedermann PH, Engl T, Kaltenpoth M (2015) Defensive symbioses of animals with
prokaryotic and eukaryotic microorganisms. Nat Prod Rep 32(7):904–936. https://doi.org/10.
1039/c5np00010f
Foster KR, Schluter J, Coyte KZ, Rakoff-Nahoum S (2017) The evolution of the host microbiome
as an ecosystem on a leash. Nature 548(7665):43–51. https://doi.org/10.1038/nature23292
Gil-Turnes MS, Fenical W (1992) Embryos of Homarus americanus are protected by epibiotic
bacteria. Biol Bull 182(1):105–108. https://doi.org/10.2307/1542184
Gil-Turnes MS, Hay ME, Fenical W (1989) Symbiotic marine bacteria chemically defend crustacean embryos from a pathogenic fungus. Science 246(4926):116–118
Gould K (2016) Antibiotics: from prehistory to the present day. J Antimicrob Chemother 71
(3):572–575. https://doi.org/10.1093/jac/dkv484
Haeder S, Wirth R, Herz H, Spiteller D (2009) Candicidin-producing Streptomyces support leafcutting ants to protect their fungus garden against the pathogenic fungus Escovopsis. Proc Natl
Acad Sci U S A 106(12):4742–4746
Hartmann A, Schmid M, van Tuinen D, Berg G (2009) Plant-driven selection of microbes. Plant
Soil 321(1–2):235–257. https://doi.org/10.1007/s11104-008-9814-y
Heine D, Holmes NA, Worsley SF, Santos ACA, Innocent TM, Scherlach K, Patrick EH, Yu DW,
Murrell JC, Vieria PC, Boomsma JJ, Hertweck C, Hutchings MI, Wilkinson B (2018) Chemical
warfare between leafcutter ant symbionts and a co-evolved pathogen. Nat Commun 9(1):2208.
https://doi.org/10.1038/s41467-018-04520-1
Hooper LV, Littman DR, Macpherson AJ (2012) Interactions between the microbiota and the
immune system. Science 336(6086):1268–1273. https://doi.org/10.1126/science.1223490
Hopwood DA (2007) Streptomyces in nature and medicine: the antibiotic makers. Oxford University Press, New York
Kaltenpoth M (2009) Actinobacteria as mutualists: general healthcare for insects? Trends Microbiol
17(12):529–535
Kaltenpoth M, Goettler W, Dale C, Stubblefield JW, Herzner G, Roeser-Mueller K, Strohm E
(2006) ‘Candidatus Streptomyces philanthi’, an endosymbiotic streptomycete in the antennae of
Philanthus digger wasps. Int J Syst Evol Microbiol 56(6):1403–1411
Kaltenpoth M, Goettler W, Koehler S, Strohm E (2010) Life cycle and population dynamics of a
protective insect symbiont reveal severe bottlenecks during vertical transmission. Evol Ecol 24
(2):463–477. https://doi.org/10.1007/s10682-009-9319-z
23 Defensive Microbiomes: A Widespread Phenomenon in Nature
509