Abstract
A new fossil species of the ant genus Cardiocondyla Emery, 1869, C. primitiva sp. nov., is described based on a winged gyne from the late Eocene Rovno amber (Ukraine, 37.8–33.9 Ma). It is the first Cardiocondyla found in the late Eocene European ambers, and the earliest known record of this genus. Most features of C. primitiva sp. nov. fit the modern diagnosis of Cardiocondyla. However, it is clearly distinguished from all known modern congeners by its much more developed forewing venation. In particular, the cells mcu, 1+2r and 3r are closed and the free branches 4+5RS and 2+3+4M are well developed, whereas forewing in modern species has only one closed cell 1+2r, and the free branches 4+5RS and 2+3+4M are strongly or completely reduced. The discovery of C. primitiva sp. nov. pushes the date of the earliest known fossil Cardiocondyla back at least 18 million years, and may force a reconsideration of its phylogenetic relationship with other Myrmicinae genera.
References
- Ashmead, W.H. (1905) A skeleton of a new arrangement of the families, subfamilies, tribes and genera of the ants, or the superfamily Formicoidea. Canadian Entomologist, 37, 381–384. https://doi.org/10.4039/Ent37381-11
- Blanchard, B.D. & Moreau, C.S. (2017) Defensive traits exhibit an evolutionary trade-off and drive diversification in ants. Evolution, 71 (2), 315–328. https://doi.org/10.1111/evo.13117
- Blaimer, B.B., Ward, P.W., Schultz, T.R., Fisher, B.L. & Brady, S.G. (2018) Paleotropical diversification dominates the evolution of the hyperdiverse ant tribe Crematogastrini (Hymenoptera: Formicidae). Insect Systematics and Diversity, 2 (5), 3, 1–14. https://doi.org/10.1093/isd/ixy013
- Bolton, B. (1981) A revision of six minor genera of Myrmicinae (Hymenoptera: Formicidae) in the Ethiopian zoogeographic region. Bulletin of the British Museum (Natural History) (Entomology), 43, 245–307.
- Bolton, B. (1982) Afrotropical species of the myrmicine ant genera Cardiocondyla, Leptothorax, Melissotarsus, Messor and Cataulacus. Bulletin of the British Museum (Natural History) (Entomology), 45, 307–370.
- Bolton, B. (1994) Identification guide to the ant genera of the world. Harvard University Press, Cambridge, Massachusetts & London, 222 pp.
- Bolton, B. (2003) Synopsis and classification of Formicidae. Memoirs of the American Entomological Institute, 71, 1–370.
- Bolton, B. (2025) An online catalog of the ants of the world. Available from: http://antcat.org (accessed May 2025).
- Bolton, B. & Marsh, A.C. (1989) The Afrotropical thermophilic genus Ocymyrmex (Hymenoptera: Formicidae). Journal of Natural History, 23, 1267–1308. https://doi.org/10.1080/00222938900770681
- Boudinot, B.E., Perrichot, V. & Chaul, J.C.M. (2020) †Camelosphecia gen. nov., lost ant-wasp intermediates from the mid-Cretaceous (Hymenoptera, Formicoidea). ZooKeys, 1005, 21–55. https://doi.org/10.3897/zookeys.1005.57629
- Boudinot, B.E., Bock, B.L., Weingardt, M., Tröger, D., Batelka, J., Li, D., Richter, A., Poh, H., Moosdorf, O.T.D., Jandausch, K., Hammel, J.U. & Beutel, R.G. (2024) Et latet et lucet: Discoveries from the Phyletisches Museum amber and copal collection in Jena, Germany. Deutsche Entomologische Zeitschrift, 71, 111–176. https://doi.org/10.3897/dez.71.112433
- Brady, S.G., Schultz, T.R., Fisher, B.L. & Ward, P.S. (2006) Evaluating alternative hypotheses for the early evolution and diversification of ants. Proceedings of the National Academy of Sciences U.S.A., 103, 18172–18177. https://doi.org/10.1073/pnas.0605858103
- Branstetter, M.G., Longino J.T., Ward, P.S. & Faircloth, B.C. (2017) Enriching the ant tree of life: enhanced UCE bait set for genome-scale phylogenetics of ants and other Hymenoptera. Methods in Ecology and Evolution, 8, 768–776. https://doi.org/10.1111/2041-210X.12742
- Brown, W.L. Jr. & Nutting, W.L. (1950) Wing venation and the phylogeny of the Formicidae. Transactions of the American Entomological Society, 75, 113–132.
- Dlussky, G.M. (1981) The ants of deserts. Nauka, Moscow, 230 pp. [In Russian]
- Dlussky, G.M. (1983) A new family of late Cretaceous Hymenoptera—“intermediate link” between ants and scolioids. Paleontologicheskii Zhurnal, 3, 65–78. [In Russian]
- Dlussky, G.M. (1987) New Formicoidea (Hymenoptera) from late Cretaceous. Paleontologicheskii Zhurnal, 1, 131–135. [In Russian]
- Dlussky, G.M. & Perfilieva, K.S. (2003) Paleogene ants of the genus Archimyrmex Cockerell, 1923. Paleontological Journal, 37, 39–47.
- Dlussky, G.M. & Perfilieva, K.S. (2014) Superfamily Formicoidea Latreille, 1802. In: Antropov, A.V., Belokobylskij, S.A., Compton, S.G. et al. The wasps, bees and ants from the Insect Limestone (Late Eocene) of the Isle of Wight, U.K. Earth and Environmental Science Transactions of the Royal Society of Edinburgh, 104 (3-4), 335–446. https://doi.org/10.1017/S1755691014000103
- Dlussky, G., Radchenko, A. & Dubovikoff, D. (2014) A new enigmatic ant genus from late Eocene Danish amber and its evolutionary and zoogeographic significance. Acta Palaeontologica Polonica, 59 (4), 931–939. https://doi.org/10.4202/app.2012.0028
- Economo, E.P., Narula, N., Friedman, N.R., Weiser, M.D. & Guénard, B. (2018) Macroecology and macroevolution of the latitudinal diversity gradient in ants. Nature Communications, 9 (1778), 1–44. https://doi.org/10.1038/s41467-018-04218-4
- Emery, C. (1895) Die Gattung Dorylus Fab. und die systematische Eintheilung der Formiciden. Zoologische Jahrbücher, Abtheilung für Systematik, Geographie und Biologie der Thiere, 8, 685–778.
- Emery, C. (1914) Intorno alla classificazione dei Myrmicinae. Rendiconto delle Sessioni della R. Accademia delle Scienze dell'Istituto di Bologna (N.S.), 18, 29–42.
- Heinze, J., Schrempl, A., Seifert, B. & Tinaut, A. (2002) Queen morphology and dispersal tactics in the ant, Cardiocondyla batesii. Insectes Sociaux, 49, 129–132. https://doi.org/10.1007/s00040-002-8291-1
- Heinze, J., Trindl, A., Seifert, B. & Yamauchi, K. (2005) Evolution of male morphology in the ant genus Cardiocondyla. Molecular Phylogenetics and Evolution, 37, 278– 288. https://doi.org/10.1016/j.ympev.2005.04.005
- Kück, P., Hita Garcia, F., Misof, B. & Meusemann, K. (2011) Improved phylogenetic analyses corroborate a plausible position of Martialis heureka in the ant tree of life. PLoS ONE, 6 (6), e21031. https://doi.org/10.1371/journal.pone.0021031
- Moreau, S. & Bell, C.D. (2013) Testing the museum versus cradle tropical biological diversity hypothesis: phylogeny, diversification, and ancestral biogeographic range evolution of the ants. Evolution, 67 (8), 2240–2257. https://doi.org/10.1111/evo.12105
- Perfilieva, K.S. (2010) Trends in evolution of ant wing venation (Hymenoptera, Formicidae). Entomological Review, 90 (7), 965–977. https://doi.org/10.1134/S0013873810070043
- Prebus, M. (2017) Insights into the evolution, biogeography and natural history of the acorn ants, genus Temnothorax Mayr (Hymenoptera: Formicidae). BMC Evolutionary Biology, 17, 1–22. https://doi.org/10.1186/s12862-017-1095-8
- Radchenko, A.G. (1995) Palaearctic ants of the genus Cardiocondyla Emery (Hymenoptera, Formicidae). Entomologicheskoe Obozrenie, 74 (2), 447–455. [English translation: Entomological Review, 1996, 75 (7), 99–109.]
- Radchenko, A. (2023a) A new extinct ant genus (Hymenoptera, Formicidae) from Baltic amber. Annales Zoologici, 73 (1), 41–49. https://doi.org/10.3161/00034541ANZ2023.73.1.004
- Radchenko, A.G. (2023b) New ant genus (Hymenoptera, Formicidae) from the Late Eocene Rovno amber. Zoodiversity, 57 (4), 323–336. https://doi.org/10.15407/zoo2023.04.323
- Radchenko, A. (2023c) Myrmica ants (Hymenoptera, Formicidae) in late Eocene European ambers. Annales Zoologici, 73 (4), 629–648. https://doi.org/10.3161/00034541ANZ2023.73.4.006
- Radchenko, A. (2024) Eotemnothorax gen. nov. (Hymenoptera, Formicidae)—a new ant genus from late Eocene European ambers. Annales Zoologici, 74 (4), 717–752. https://doi.org/10.3161/00034541ANZ2024.74.4.015
- Radchenko, A. (2025) Ants of the genus Aphaenogaster (Hymenoptera, Formicidae) in late Eocene European ambers. Annales Zoologici, 75 (2), 437–454. https://doi.org/10.3161/00034541ANZ2025.75.2.003
- Radchenko, A. & Dlussky, G.M. (2018) Two new fossil species of the ant genus Pristomyrmex Mayr (Hymenoptera, Formicidae) from the Rovno and Bitterfeld ambers. Annales Zoologici, 68 (2), 251–258. https://doi.org/10.3161/00034541ANZ2018.68.2.004
- Radchenko, A. & Dlussky, G.M. (2019) The first record of the ant genus Crematogaster (Hymenoptera, Formicidae) from the late Eocene European amber. Annales Zoologici, 69 (2), 417–421. https://doi.org/10.3161/00034541ANZ2019.69.2.008
- Radchenko, A., Dlussky, G.M. & Perfilieva, K.S. (2018) A new extinct ant genus (Hymenoptera, Formicidae, Myrmicinae) from the Late Eocene Rovno amber—a putative ancestor of the Leptothorax genus group. Myrmecological News, 27, 111–117.
- Radchenko, Gröhn, C. & Ribbecke H.-W. (2024) Two new Aphaenogaster species (Hymenoptera, Formicidae) from Baltic Amber. Zoodiversity, 58 (3), 195–202. https://doi.org/10.15407/zoo2024.03.195
- Radchenko, A.G., Khomych, M.R. & Kalmer, M.M. (2025) First record of Bradoponera wunderlichi (Hymenoptera, Formicidae) from Rovno amber, with description of the worker caste. Zoodiversity, 59 (2), 113–120. https://doi.org/10.15407/zoo2025.02.113
- Santschi, F. (1930) Formicides de l’Angola. Revue Suisse de Zoologie, 37 (3), 53–82. https://doi.org/10.5962/bhl.part.117942
- Seifert, B. (2003a) The ant genus Cardiocondyla (Insecta: Hymenoptera: Formicidae)—a taxonomic revision of the C. elegans, C. bulgarica, C. batesii, C. nuda, C. shuckardi, C. stambuloffii, C. wroughtonii, C. emeryi, and C. minutior species groups. Annalen des Naturhistorischen Museums, Wien, Serie B, 104, 203–338.
- Seifert, B. (2023b) The ant genus Cardiocondyla (Hymenoptera: Formicidae): the species groups with Oriental and Australasian origin. Diversity, 15, 25, 1–60. https://doi.org/10.3390/d15010025
- Seifert, B. (2023c) A revision of the Palaearctic species of the ant genus Cardiocondyla Emery 1869 (Hymenoptera: Formicidae). Zootaxa, 5274 (1), 1–64. https://doi.org/10.11646/zootaxa.5274.1.1
- Seifert, B., Okita, I. & Heinze, J. (2017) A taxonomic revision of the Cardiocondyla nuda group (Hymenoptera: Formicidae). Zootaxa, 4290 (2), 324–356. https://doi.org/10.11646/zootaxa.4290.2.4
- Sharaf, M.R., Al Dhafer, H.M., Abdel-Dayem, M.S. & Aldawood, A.S. (2024) Cardiocondyla hashemi sp. n., a new species of the C. batesii species-group (Hymenoptera: Formicidae) from Saudi Arabia, with a key to the Saudi species. Zoology in the Middle East, February 2024, 1–11. https://doi.org/10.1080/09397140.2024.2321640
- Skalski, A.W. & Veggiani, A. (1990) Fossil resin in Sicily and the Northern Apennines: geology and organic content. Prace Muzeum Ziemi, 41, 37–49.
- Wang, B., Shi, G.L., Xu, C.P., Spicer, R.A., Perrichot, V., Schmidt, A.R., Feldberg, K., Heinrichs, J., Chény, C., Pang, H., Liu, X.Y., Gao, T.P., Wang Z.X., Ślipiński, A., Solórzano-Kraemer, M.M., Heads, S.W., Thomas, M.J., Sadowski, E.-M., Szwedo, J., Azar, D., Nel, A., Liu, Y., Chen, J., Zhang, Q., Zhang, Q.Q., Luo, C.H., Yu, T.T., Zheng D.R., Zhang, H.C. & Engel, M.S. (2021) The mid-Miocene Zhangpu biota reveals an outstandingly rich rainforest biome in East Asia. Science Advances, 7 (18), eabg0625. https://doi.org/10.1126/sciadv.abg0625
- Ward, P.S., Brady, S.G., Fisher, B.L. & Schultz, T.R. (2015) The evolution of myrmicine ants: phylogeny and biogeography of a hyperdiverse ant clade. Systematic Entomology, 40, 61–81. https://doi.org/10.1111/syen.12090
- Yoshimura, M. & Fisher, B.L. (2009) A revision of the male ants of the Malagasy region: key to genera of the subfamily Proceratiinae. Zootaxa, 2216 (1), 1–21. https://doi.org/10.11646/zootaxa.2216.1.1