Planned intervention: On Wednesday June 26th 05:30 UTC Zenodo will be unavailable for 10-20 minutes to perform a storage cluster upgrade.
Published April 8, 2021 | Version v1
Journal article Open

The head anatomy of Protanilla lini (Hymenoptera: Formicidae: Leptanillinae), with a hypothesis of their mandibular movement

  • 1. Institut für Zoologie und Evolutionsforschung, FSU Jena, 07743 Jena, Germany; Biodiversity and Biocomplexity Unit, Okinawa Institute of Science and Technology Graduate University, Onna-son, Okinawa, 904-0495, Japan.
  • 2. Biodiversity and Biocomplexity Unit, Okinawa Institute of Science and Technology Graduate University, Onna-son, Okinawa, 904-0495, Japan.
  • 3. Biodiversity and Biocomplexity Unit, Okinawa Institute of Science and Technology Graduate University, Onna-son, Okinawa, 904-0495, Japan; MUHNAC / cE3c – Centre for Ecology, Evolution and Environmental Changes, Faculdade de Ciências da Universidade de Lisboa, 1749-016 Lisbon, Portugal. Johan Billen, Zoological Institute, KU Leuven, 3000 Leuven, Belgium.
  • 4. Zoological Institute, KU Leuven, 3000 Leuven, Belgium.
  • 5. Institut für Zoologie und Evolutionsforschung, FSU Jena, 07743 Jena, Germany; Department of Entomology & Nematology, University of California, Davis, One Shields Ave, Davis, CA 95618.
  • 6. Institut für Zoologie und Evolutionsforschung, FSU Jena, 07743 Jena, Germany


Richter, Adrian, Garcia, Francisco Hita, Keller, Roberto A., Billen, Johan, Katzke, Julian, Boudinot, Brendon E., Economo, Evan P., Beutel, Rolf G. (2021): The head anatomy of Protanilla lini (Hymenoptera: Formicidae: Leptanillinae), with a hypothesis of their mandibular movement. Myrmecological News 31: 85-114, DOI: 10.25849/myrmecol.news_031:085



Files (12.0 MB)

Name Size Download all
12.0 MB Preview Download

Linked records

Additional details




  • AGOSTI, D., MAJER, J., ALONSO, E. & SCHULTZ, T.R. 2000: Ants: standard methods for measuring and monitoring biodiversity. - Smithsonian Institution Press, Washington, DC, XXI + 280 pp.
  • BAIDYA, P. & BAGCHI, S. 2020: Anew species of Protanilla TAYLOR 1990 (Hymenoptera: Formicidae: Leptanillinae) from India. - Halteres 11: 19-24.
  • BARDEN,P.2017:Fossilants (Hymenoptera:Formicidae):ancient diversity and the rise of modern lineages. - Myrmecological News 24: 1-30.
  • BARDEN, P. & GRIMALDI, D. 2013: Anew genus of highly specialized ants in Cretaceous Burmese amber (Hymenoptera: Formicidae). - Zootaxa 3681: 405-412.
  • BARDEN,P. & GRIMALDI, D. 2014: Adiverse ant fauna from the mid-Cretaceous of Myanmar (Hymenoptera: Formicidae). - Public Library of Science One 9: art. e93627.
  • BARONI URBANI, C. 1977: Materiali per una revision della sottofamiglia Leptanillinae EMERY (Hymenoptera: Formicidae). - Entomologica Brasiliensa 2: 427-488.
  • BARONIURBANI,C., BOLTON,B.& WARD,P.S. 1992: The internal phylogeny of ants (Hymenoptera: Formicidae). - Systematic Entomology 17: 301-329.
  • BARONI URBANI, C. & DE ANDRADE, M. 2007: Theant tribe Dacetini: limits and constituent genera, with descriptions of new species. - Annali del Museo Civico di Storia Naturale Giacomo Doria (Genova) 99: 1-191.
  • BEUTEL, R.G., FRIEDRICH, F., HORNSCHEMEYER, T., POHL, H., HUNEFELD, F., BECKMANN,F., MEIER, R., MISOF, B., WHITING, M.F. & VILHELMSEN, L. 2011: Morphological and molecular evidenceconverge upona robust phylogenyof themegadiverse Holometabola. - Cladistics 27: 341-355.
  • BEUTEL, R.G., FRIEDRICH, F., YANG, X.-K. & GE, S.-Q. 2014: Insect morphology and phylogeny: a textbook for students of entomology. - Walter de Gruyter, Berlin, 532 pp.
  • BEUTEL, R.G. & VILHELMSEN,L. 2007:Headanatomy of Xyelidae (Hexapoda: Hymenoptera) and phylogenetic implications. - Organisms Diversity & Evolution 7: 207-230.
  • BHARTI, H. & AKBAR, S.A. 2015: First record of genus Protanilla (Hymenoptera: Formicidae: Leptanillinae) from India with description of a new species. - Journal of Asia-Pacific Entomology 18: 573-576.
  • BILLEN,J., BAUWELEERS,E., HASHIM, R.& ITO,F. 2013:Survey of the exocrine system in Protanilla wallacei (Hymenoptera, Formicidae). - Arthropod Structure & Development 42: 173-183.
  • BILLEN, J., ITO, F., MAILE, R. & MORGAN, E.D. 1998: The mandibular gland, probably the source of the alarm substance in Leptanilla sp. (Hymenoptera, Formicidae). - Naturwissenschaften 85: 596-597.
  • BLANKE,A. 2019: The earlyevolution of biting-chewing performance in Hexapoda. In: KRENN, H.W. (Ed.): Insect mouthparts. - Springer, Cham, pp. 175-202.
  • BOLTON,B. 1990:The higherclassification of theantsubfamily Leptanillinae (Hymenoptera: Formicidae). - Systematic Entomology 15: 267-282.
  • BOLTON,B. 1999: Ant genera of the tribe Dacetonini (Hymenoptera: Formicidae). - Journal of Natural History 33: 1639- 1689.
  • BOLTON, B. 2003: Synopsis and classification of Formicidae. - Memoirs of the American Entomological Institute 71: 1-370.
  • BOLTON, B. 2020: An online catalog of the ants of the world. - <>, retrieved on 14 September 2020.
  • BOLTON,B.& FISHER,B.L.2008:Afrotropicalantsof theponerine genera Centromyrmex MAYR, Promyopias SANTSCHI gen. rev. and Feroponera gen. n., with a revised key to genera of African Ponerinae (Hymenoptera: Formicidae). - Zootaxa 1929: 1-37.
  • BOOHER, D.B., GIBSON, J.C., LIU, C., LONGINO, J.T., FISHER, B.L., JANDA, M., NARULA, N., TOULKERIDOU, E., MIKHEYEV, A.S. & SUAREZ, A.V. 2021: Functional innovation promotes diversification of form in the evolution of an ultrafast trapjaw mechanism in ants. - Public Library of Science Biology 19: art. e3001031.
  • BOROWIEC, M.L., RABELING, C., BRADY, S.G., FISHER, B.L., SCHULTZ,T.R.& WARD,P.S.2019:Compositionalheterogeneity and outgroup choice influence the internal phylogeny of the ants. - Molecular phylogenetics and evolution 134: 111-121.
  • BOROWIEC, M.L., SCHULZ, A., ALPERT, G.D. & BANAR, P. 2011: Discovery of the worker caste and descriptions of two new speciesof Anomalomyrma (Hymenoptera: Formicidae: Leptanillinae) with unique abdominal morphology. - Zootaxa 2810: 1-14.
  • BORYSENKO, L.H. 2017: Description of a new genus of primitive ants from Canadian amber, with the study of relationships between crown ants and stem ants (Hymenoptera: Formicidae). -Insecta Mundi 570: 1-57.
  • BOUDINOT, B.E. 2015: Contributions to the knowledge of Formicidae (Hymenoptera, Aculeata): a new diagnosis of the family, the first global male-based key to subfamilies, and a treatment of early branching lineages. - European Journal of Taxonomy 120: 1-62.
  • BOUDINOT, B.E., KHOURI, Z., RICHTER, A., VAN DE KAMP, T., BARDEN, P., PERRICHOT, V. & CHAUL, J.C.M. 2020a: Chapter II. The evolution of the ants: Extinct ant sister-group illuminates eusociality origin and post-K/Pg persistence. Pp. 174- 393. In: BOUDINOT, B.E. 2020: Systematic and evolutionary morphology; Case studies on Formicidae, Mesozoic Aculeata and Hexapodan genitalia. - Universityof California, Davis, CA, 562 pp.
  • BOUDINOT,B.E., PERRICHOT,V.& CHAUL,J.C.M. 2020b: †Camelosphecia gen. nov., lost ant-wasp intermediates from the mid Cretaceous (Hymenoptera, Formicoidea). - ZooKeys 1005: 21-55.
  • BRADY, S.G., SCHULTZ, T.R., FISHER, B.L. & WARD, P.S. 2006: Evaluating alternativehypotheses for the early evolution and diversificationof ants.- Proceedings of the National Academy of Sciences of the United States of America 103: 18172-18177.
  • BRADY, S.G. & WARD, P.S. 2005: Morphological phylogeny of army ants and other dorylomorphs (Hymenoptera: Formicidae). - Systematic Entomology 30: 593-618.
  • BRANDAO, C.R.F., DINIZ, J.L.M. & FEITOSA, R.M. 2010: The venom apparatus and other morphological characters of the ant Martialis heureka (Hymenoptera, Formicidae, Martialinae). - Papeis Avulsos de Zoologia (Sao Paulo) 50: 413-423.
  • BRANSTETTER, M.G., LONGINO, J.T., WARD, P.S. & FAIRCLOTH, B.C. 2017: Enriching the ant tree of life: enhanced UCE bait set forgenome-scale phylogeneticsof ants and other Hymenoptera. - Methods in Ecology and Evolution 8: 768-776.
  • BROWN, W.L. 1953: Revisionary studies in the ant tribe Dacetini. - The American Midland Naturalist 50: 1-137.
  • BROWN,W.L. & KEMPF,W.W. 1967:Tatuidris, a remarkable new genus of Formicidae (Hymenoptera). - Psyche 74: 183-190.
  • BROWN, W.L., LEVIEUX, J. & GOTWALD Jr., W.H. 1970: Anew genus of ponerineantsfrom West Africa.- Psyche 77:259-275.
  • CAO, H., BOUDINOT, B.E., WANG, Z., MIAO, X., SHIH, C., REN, D. & GAO,T. 2020: Two new iron maiden ants from Burmese amber (Hymenoptera: Formicidae: †Zigrasimeciini). - Myrmecological News 30: 161-173.
  • CHAPMAN, R.F., SIMPSON, S.J. & DOUGLAS, A.E. (Eds.) 2012: The insects: structure and function, 5th edn. - Cambridge University Press, Cambridge, UK, 929 pp.
  • DUNCAN, C.D. 1969: Acontribution to the biology of North American vespine wasps.- Stanford University Publications, Biological Sciences 8: 1-272.
  • EHMER, B. & GRONENBERG, W. 1997: Antennal muscles and fast antennal movements in ants. - Journal of Comparative Physiology B 167: 287-296.
  • EMERY,C. 1904: Le affinita del genere Leptanilla e i limiti delle Dorylinae. - Archivio Zoologico 2: 107-116.
  • ENGELKES, K., FRIEDRICH, F., HAMMEL, J.U. & HAAS, A. 2018: A simple setup for episcopic microtomy and a digital image processing workflowto acquire high-quality volumedataand 3D surface models of small vertebrates. - Zoomorphology 137: 213-228.
  • FISHER, B.L. 1999: Improving inventory efficiency: a case study of leaf-litter ant diversity in Madagascar. - Ecological Applications 9: 714-731.
  • GOTWALD,W.H. 1969: Comparativemorphologicalstudies of the ants: with particular reference to the mouthparts (Hymenoptera: Formicidae). - New York State College of Agriculture, Ithaca, NY, 150 pp.
  • GRIEBENOW, Z.H. 2020a: Synonymization of the male-based ant genus Phaulomyrma (Hymenoptera, Formicidae) with Leptanilla based upon Bayesian total-evidence phylogenetic inference. - bioRxiv; doi: 10.1101/2020.08.28.272799.
  • GRIEBENOW, Z.H. 2020b: Delimitation of tribesin the subfamily Leptanillinae (Hymenoptera:Formicidae), with a description of the male of Protanilla lini TERAYAMA, 2009. - Myrmecological News 30: 229-250.
  • GRONENBERG, W. 1995:Thefastmandiblestrikeinthetrap-jaw ant Odontomachus. - Journal of Comparative Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology 176: 391-398.
  • GRONENBERG,W.1996:Thetrap-jaw mechanismin thedacetine ants Daceton armigerum and Strumigenys sp. - Journal of Experimental Biology 199: 2021-2033.
  • GRONENBERG, W., BRANDAO, C.R.F., DIETZ, B.H. & JUST, S. 1998: Trap-jaws revisited: themandible mechanismof theant Acanthognathus. - Physiological Entomology 23: 227-240.
  • GRONENBERG, W., PAUL, J., JUST, S. & HOLLDOBLER, B. 1997: Mandible muscle fibers in ants: fast or powerful? - Cell and Tissue Research 289: 347-361.
  • HITA-GARCIA, F., LIEBERMAN, Z., AUDISIO, T.L., LIU, C. & ECONOMO, E.P. 2019: Revision of the highly specialized ant genus Discothyrea (Hymenoptera: Formicidae) in the Afrotropics with X-ray microtomographyand 3D cybertaxonomy. - Insect Systematics and Diversity 3: art. 5.
  • HOLLDOBLER, B. & WILSON, E.O. 1990: The ants. - Harvard University Press, Cambridge, MA, 732 pp.
  • HSU, P.-W., HSU, F.-C., HSIAO, Y. & LIN, C.-C. 2017: Taxonomic notes on the genus Protanilla (Hymenoptera: Formicidae: Leptanillinae) from Taiwan. - Zootaxa 4268: 117-130.
  • ITO,F. & YAMANE,S. 2020: Behavior of the queen of Leptanilla clypeata YAMANE et Ito collected in the Bogor Botanical Gardens, West Java, Indonesia (Hymenoptera; Formicidae), with a note on colony composition and a description of the ergatoid queen. - Asian Myrmecology 12: art. e012004.
  • JACQUEMIN, J., DELSINNE, T., MARAUN, M. & LEPONCE, M. 2014: Trophic ecology of the armadillo ant, Tatuidristatusia, assessed by stable isotopes and behavioral observations. - Journal of Insect Science 14: art. 108.
  • JANET, C. 1905: Anatomie de la tete du Lasius niger. Reine. - Limoges Imprimerie-Librairie Ducourtieux et Gout, Li- moges, 40 pp.
  • KELLER, R.A. 2011: Aphylogenetic analysis of ant morphology (Hymenoptera: Formicidae) with special reference to the poneromorph subfamilies. - Bulletin of the American Museum of Natural History 355: 1-90.
  • KELLER,R.A.& PEETERS, C. 2020: Poneroid ants. In: STARR,C. (Ed.): Encyclopediaof social insects. - Springer International Publishing, Cham, pp. 1-6.
  • KRONAUER, D.J.C. 2009: Recent advances in army ant biology (Hymenoptera: Formicidae). - Myrmecological News 12: 51-65.
  • KUGLER, C. 1992: Stings of ants of the Leptanillinae (Hymenoptera: Formicidae). - Psyche 99: 103-115.
  • KUGLER, J. 1987: The Leptanillinae (Hymenoptera: Formicidae). - Israel Journal of Entomology 20: 45-57.
  • KUCK, P., GARCIA, F.H., MISOF, B. & MEUSEMANN, K. 2011: Improved phylogenetic analyses corroborate a plausible position of Martialis heureka in the ant tree of life. - Public Library of Science One 6: art. e21031.
  • LARABEE, F.J., GRONENBERG, W. & SUAREZ, A.V. 2017: Performance, morphologyand controlof power-amplifiedmandibles in the trap-jaw ant Myrmoteras (Hymenoptera:Formicidae). - Journal of Experimental Biology 220: 3062-3071.
  • LARABEE,F.J.& SUAREZ,A.V. 2014: Theevolution andfunctional morphology of trap-jaw ants (Hymenoptera: Formicidae). - Myrmecological News 20: 25-36.
  • LEONG, C.-M., YAMANE, S. & GUENARD, B. 2018: Lost in the city: discovery of the rare ant genus Leptanilla (Hymenoptera: Formicidae) in Macau with description of Leptanilla macauensis sp. nov. - Asian Myrmecology 10: art. e010001.
  • LOPEZ, F., MARTINEZ, M. & BARANDICA, J. 1994: Four new species of the genus Leptanilla (Hymenoptera: Formicidae) from Spain - relationships to other species and ecological issues. - Sociobiology 24: 179-212.
  • LOSEL,P.&HEUVELINE,V.2016:Enhancingadiffusionalgorithm for 4D image segmentation using local information. - SPIE Medical Imaging 9784: art. 97842L.
  • LOSEL, P.D., VAN DE KAMP, T., JAYME, A., ERSHOV, A., FARAGO, T., PICHLER, O., JEROME, N.T., AADEPU, N., BREMER, S., CHILINGARYAN, S.A., HEETHOFF, M., KOPMANN, A., ODAR, J., SCHMELZLE, S., ZUBER, M., WITTBRODT, J., BAUMBACH, T. & HAUVELINE, V. 2020: Introducing Biomedisa as an opensource online platform for biomedical image segmentation. - Nature Communications 11: art. 5577.
  • LUCKY, A., TRAUTWEIN, M.D., GUENARD, B.S., WEISER, M.D. & DUNN, R.R. 2013: Tracing the rise of ants-out of the ground. - Public Library of Science One 8: art. e84012.
  • MAN, P., RAN, H., CHEN, Z. & XU, Z. 2017: The northern-most record of Leptanillinaein Chinawith descriptionof Protanilla beijingensis sp. nov. (Hymenoptera: Formicidae). - Asian Myrmecology 9: 1-12.
  • MASUKO,K. 1989: Larval hemolymph feeding in the ant Leptanilla japonica by use of a specialized duct organ, the "larval hemolymph tap" (Hymenoptera: Formicidae). - Behavioral Ecology and Sociobiology 24: 127-132.
  • MASUKO, K. 1990: Behavior and ecology of the enigmatic ant Leptanilla japonica BARONI URBANI (Hymenoptera: Formicidae: Leptanillinae). - Insectes Sociaux 37: 31-57.
  • MICHENER, C.D. & FRASER, A. 1978: A comparative anatomical study of mandibular structure in bees (Hymenoptera: Apoidea).- Universityof Kansas Science Bulletin 51: 463-482.
  • MOREAU, C.S. & BELL, C.D. 2013: Testing the museum versus cradle tropical biological diversity hypothesis: phylogeny, diversification, and ancestralbiogeographic range evolution of the ants. - Evolution 67: 2240-2257.
  • MOREAU, C.S., BELL, C.D., VILA, R., ARCHIBALD, S.B. & PIERCE, N.E. 2006: Phylogeny of the ants: diversification in the age of angiosperms. - Science 312: 101-104.
  • NELSEN, M.P., REE, R.H. & MOREAU, C.S. 2018: Ant-plant interactions evolved through increasing interdependence. - Proceedings of the National Academy of Sciences of the United States of America 115: 12253-12258.
  • OGATA, K., TERAYAMA, M. & MASUKO, K. 1995: The ant genus Leptanilla: discovery of the worker-associated male of L. japonica, and a description of a new species from Taiwan (Hymenoptera: Formicidae: Leptanillinae). - Systematic Entomology 20: 27-34.
  • PATEK, S., BAIO, J., FISHER, B. & SUAREZ, A. 2006: Multifunctionality and mechanical origins: ballistic jaw propulsion in trap-jaw ants. - Proceedings of the National Academy of Sciences of the United States of America 103: 12787-12792.
  • PAUL,J. & GRONENBERG,W. 1999: Optimizing force and velocity: mandible muscle fibre attachments in ants. - Journal of Experimental Biology 202: 797-808.
  • PAUL, J., ROCES, F. & HOLLDOBLER, B. 2002: Howdo ants stick out their tongues? - Journal of Morphology 254: 39-52.
  • PERRAULT,G.H.1999:Etudemorphoanatomiqueetbiometrique du mEtasoma antErieur des ouvrieres. Contribution a la systEmatique et a la phylogEnie des fourmis (Hymenoptera: Formicidae). - Annales de la Societe entomologique de France 40: 291-371.
  • PERRICHOT,V. 2014: Anew species of the Cretaceous ant Zigrasimecia based on the worker caste reveals placement of the genus in the Sphecomyrminae (Hymenoptera: Formicidae). - Myrmecological News 19: 165-169.
  • PETERSEN,B. 1968: Some novelties in presumed males of Leptanillinae (Hym., Formicidae). - Entomologiske Meddelelser 36: 577-598.
  • POHL,H.2010:Ascanningelectron microscopyspecimen holder forviewing differentanglesof a single specimen.- Microscopy Research and Technique 73: 1073-1076.
  • POLILOV, A.A. 2016:At thesizelimit-effectsof miniaturization in insects. - Springer International Publishing, Cham, 335pp.
  • POLILOV, A.A., RIBERA, I., YAVORSKAYA, M.I., CARDOSO, A., GREBENNIKOV, V.V. & BEUTEL, R.G. 2019: The phylogeny of Ptiliidae (Coleoptera: Staphylinoidea) - the smallest beetles and their evolutionary transformations. - Arthropod Systematics & Phylogeny 77: 433-455.
  • RABELING, C., BROWN, J.M. & VERHAAGH, M. 2008: Newly discovered sister lineage sheds light on early ant evolution. - Proceedings of the National Academy of Sciences of the United States of America 105: 14913-14917.
  • RICHTER, A., GARCIA, F.H., KELLER,R.A., BILLEN, J., ECONOMO, E.P. & BEUTEL, R.G. 2020: Comparative analysis of worker head anatomy of Formica and Brachyponera (Hymenoptera: Formicidae). - Arthropod Systematics & Phylogeny 78: 133-170.
  • RICHTER, A., KELLER, R.A., ROSUMEK, F.B., ECONOMO, E.P., HITA GARCIA, F. & BEUTEL, R.G. 2019: The cephalic anatomy of workers of the antspecies Wasmanniaaffinis (Formicidae, Hymenoptera, Insecta) and its evolutionary implications. - Arthropod Structure & Development 49: 26-49.
  • SCHINDELIN,J., ARGANDA-CARRERAS,I.,FRISE, E.,KAYNIG,V., LONGAIR, M., PIETZSCH, T., PREIBISCH, S., RUEDEN, C., SAAL- FELD, S., SCHMID, B., TINEVEZ, J.-Y., WHITE, D.J., HARTEN- STEIN, V., ELICEIRI, K., TOMANCAK, P. & CARDONA, A. 2012: Fiji: an open-source platform for biological-image analysis. - Nature Methods 9: 676-682.
  • SCHMIDT,F.& SOLAR,R. 2010: Hypogaeic pitfall traps: methodological advances and remarks to improve the sampling of a hidden ant fauna. - Insectes Sociaux 57: 261-266.
  • STOEV, P., AKKARI, N., KOMERICKI, A., EDGECOMBE, G.D. & BONATO,L. 2015: At the end of the rope: Geophilus hadesi sp. n. - the world's deepest cave-dwelling centipede (Chilopoda, Geophilomorpha, Geophilidae). - ZooKeys 510: 95-114.
  • WANG, Z., ZHANG, W., WU, J. & YANG, Y. 2020: An approach to observing and quantifying real-time mandibular muscle topologyin thetrap-jawant Odontomachusmonticola bysynchrotron imaging. - Journal of Insect Behavior 33: 174-183.
  • WARD,P.S. 2014: Thephylogeny andevolution of ants. - Annual Review of Ecology, Evolution, and Systematics 45: 23-43.
  • WARD, P.S. & FISHER, B.L. 2016: Tales of dracula ants: the evolutionary history of the ant subfamily Amblyoponinae (Hymenoptera: Formicidae). - Systematic Entomology 41: 683-693.
  • WHEELER, G.C. & WHEELER, E.W. 1930: Two new ants from Java. - Psyche 37: 193-201.
  • WILKIE, K.T.R., MERTL, A.L. & TRANIELLO, J.F.A. 2007:Biodiversity below ground: probing the subterranean ant fauna of Amazonia. - Naturwissenschaften 94: 725-731.
  • WILSON, E.O. 1955: A monographic revision of the ant genus Lasius. - Bulletin of the Museum of Comparative Zoology 113: 1-199.
  • WIPFLER, B., MACHIDA, R., MULLER, B. & BEUTEL, R.G. 2011: OntheheadmorphologyofGrylloblattodea (Insecta)andthe systematic position of the order, with a new nomenclature for the head muscles of Dicondylia. - Systematic Entomology 36: 241-266.
  • WONG, M.K.L. & GUENARD, B. 2016: Leptanilla hypodracos sp. n., a new species of the cryptic ant genus Leptanilla (Hymenoptera, Formicidae) from Singapore, with new distribution data and an updated key to Oriental Leptanilla species. - ZooKeys 551: 129-144.
  • WONG, M. & GUENARD, B. 2017: Subterranean ants: summary and perspectives on field sampling methods, with notes on diversity and ecology (Hymenoptera: Formicidae). - Myrmecological News 25: 1-16.
  • WONG, M.K.L. & GUENARD, B. 2020: Subterranean ants. In: STARR, C. (Ed.): Encyclopediaof social insects. - Springer International Publishing, Cham, pp. 1-6.
  • XU, Z.-H. 2012: Furcotanilla, a new genus of the ant subfamily Leptanillinae from China with descriptions of two new speciesof Protanilla and P. rafflesi Taylor (Hymenoptera: Formicidae). - Sociobiology 59: 477-491.
  • YAMADA, A., NGUYEN, D.D. & EGUCHI, K. 2020: Unveiling the morphology of the oriental rare monotypic ant genus Opamyrma Yamane, Bui & Eguchi, 2008 (Hymenoptera: Formicidae: Leptanillinae) and its evolutionary implications, with first descriptions of the male, larva, tentorium, and sting apparatus. - Myrmecological News 30: 27-52.
  • YAMANE,S., BUI,T.V.& EGUCHI,K.2008:Opamyrmahungvuong, a newgenusandspeciesof antrelatedto Apomyrma (Hymenoptera: Formicidae: Amblyoponinae). - Zootaxa 1767: 55-63.
  • ZHANG, W., LI, M., ZHENG, G., GUAN, Z., WU, J. & WU, Z. 2020: Multifunctional mandibles of ants: variation in gripping behavior facilitated byspecific microstructures andkinematics. - Journal of Insect Physiology 120: art. 103993.
  • ZIMMERMANN, D. & VILHELMSEN, L. 2016: The sister group of Aculeata (Hymenoptera) - evidence from internal head anatomy, with emphasis on the tentorium. - Arthropod Systematics & Phylogeny 74: 195-218.