, Volume 74, Issue 1, pp 65–89 | Cite as

Parasitic cockroaches indicate complex states of earliest proved ants

  • Peter VršanskýEmail author
  • Lucia Šmídová
  • Hemen Sendi
  • Peter Barna
  • Patrick Müller
  • Sieghard Ellenberger
  • Hao Wu
  • Xiaoyin Ren
  • Xiaojie Lei
  • Dany Azar
  • Juraj Šurka
  • Tao Su
  • Weiyudong Deng
  • Xianhui Shen
  • Jun Lv
  • Tong Bao
  • Günter Bechly
Original Article


Myrmecophilous and termitophilous interactions likely contributed to the competitive advantage and evolutionary success of eusocial insects, but how these commensal and parasitic relationships originated is unclear due to absence of fossil records. New extinct cockroaches of the still living family Blattidae are reported here from the Cretaceous Myanmar amber (99 Ma) and are the earliest known inhabitants of complex ant nests, demonstrating that this specialised myrmecophily originated shortly after ant eusociality and appeared in the fossil record. Cretaceous stem aposematic Blattidae are known from the amber of Myanmar and Lebanon and we report them here also from the Syrian amber. Concurrent evolution suggests that the collective internal defence of early ants was weak and allowed infiltrations by numerous unrelated organisms, At the same time, the contemporary presence of ant mimicking myrmecomorphs suggests a need for strong external protection against visually hunting predators. Myrmecophily is supported by morphological adaptations (lack of wide fat body and feeding of adult male; short, fossorial legs; shortened cerci; oligomerised antenna; hairy surface structures) and camouflage behaviour, documented by sediment and own feces covering. Moreover the same piece of amber contains ants, ant mimics and other undescribed ant nest-visiting insects as syninclusions. Another species preserved along with two termites is a putative termitophile. Abundant comparatively large parasitic cockroaches influenced Mesozoic tropical forest ecosystems by affecting the early evolution of complex nests of eusocial insects. Rainforest rudiments in South Yunnan yielded observation of analogical still living, formally undescribed species.


Fossil insect Mesozoic Cretaceous amber Myanmar Syria New genera New species 



We thank Ing. Robert Oružinský, Dr. Mária Kazimírová, Dr. Ľubomír Vidlička, Martin Styan (Bratislava), Prof. Bo Wang (Nanjing) and Dr. Karin Wolfschwenninger (Stuttgart) for technical help and linguistic revision. This work was supported by the Slovak Research and Development Agency under the contract no. APVV-0436-12, and by UNESCO-Amba/ MVTS supporting grant of Presidium of the Slovak Academy of Sciences; VEGA 0012-14, 2/0042/18; Literary Fund. This research was supported by the Strategic Priority Research Program (B) of the Chinese Academy of Sciences (XDB26000000) and the National Natural Science Foundation of China (41572010, 41622201, 41688103).

Author’s contribution

We collected the material (S.E., P.M., D.A., L.J., W.H., B.T.); took the photographs (S.E., P.M., P.B., L.Š., P.V., X.R., X.L.), produced drawings (L.Š., P.B., P.V.), CT (P.B., L.Š., H.S.), descriptions and comparison (P.V., L.Š., P.B., H.S.); observations in the living ecosystem (P.V., T.S., W.D., T.B., X.S.); designed research (P.V.), wrote and edited the paper (P. V., G.B., D.A.) with contributions from all authors.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.


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© Institute of Zoology, Slovak Academy of Sciences 2018

Authors and Affiliations

  • Peter Vršanský
    • 1
    • 2
    • 3
    • 4
    • 5
    Email author
  • Lucia Šmídová
    • 6
  • Hemen Sendi
    • 7
  • Peter Barna
    • 3
  • Patrick Müller
    • 8
  • Sieghard Ellenberger
    • 9
  • Hao Wu
    • 10
  • Xiaoyin Ren
    • 5
  • Xiaojie Lei
    • 5
  • Dany Azar
    • 5
    • 11
  • Juraj Šurka
    • 12
  • Tao Su
    • 13
  • Weiyudong Deng
    • 13
  • Xianhui Shen
    • 13
  • Jun Lv
    • 14
  • Tong Bao
    • 5
    • 15
  • Günter Bechly
    • 16
  1. 1.Institute of ZoologySlovak Academy of SciencesBratislavaSlovakia
  2. 2.Institute of Physics, Research Centre of Quantum InformaticsSlovak Academy of SciencesBratislavaSlovakia
  3. 3.Earth Science InstituteSlovak Academy of SciencesBratislavaSlovakia
  4. 4.Palaeontological InstituteRussian Academy of SciencesMoscowRussia
  5. 5.State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and PalaeontologyChinese Academy of SciencesNanjingChina
  6. 6.Institute of Geology and Palaeontology, Faculty of ScienceCharles University in PraguePrague 2Czech Republic
  7. 7.Faculty of Natural SciencesComenius UniversityBratislavaSlovakia
  8. 8.KäshofenGermany
  9. 9.KasselGermany
  10. 10.Zhejiang Museum of Natural HistoryHangzhouChina
  11. 11.Natural Sciences Department, Faculty of Science II, FanarLebanese UniversityFanar – MatnLebanon
  12. 12.Earth Science InstituteSlovak Academy of SciencesBanská BystricaSlovakia
  13. 13.Paleoecology Research Group, CAS MenglunXishuangbanna Tropical Botanical GardenMenglaChina
  14. 14.Dian Jiang Collection, Zhao Jia Yue TaiBaoshanChina
  15. 15.Steinmann-InstitutAbteilung PaläontologieBonnGermany
  16. 16.BöblingenGermany

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