Redescription of Proformica nasuta (Nylander, 1856) (Hymenoptera, Formicidae) using an integrative approach

Christophe Galkowski, Claude Lebas, Philippe Wegnez, Alain Lenoir, Rumsaïs Blatrix


The taxonomy of the Palaearctic ant genus Proformica Ruzsky, 1902 is confused and in need of revision. The type specimen for P. nasuta (Nylander, 1856), the type species of the genus, was from Beaucaire, southern France, and is presumably lost. Based on extensive sampling of Proformica nests in southern France, including the type locality, we show that the concept of P. nasuta has been erroneous for more than a century. We integrate information from the morphology of workers and sexual castes, DNA markers, and cuticular hydrocarbons to re-define species in southern France. This allowed us to provide a new, accurate description of P. nasuta and designate a neotype, as well as reference individuals for all castes. In addition, we propose a name, P. longipilosa sp. nov., for a species that since the end of the 19th century has mistakenly been included in P. nasuta.


ants; Proformica longipilosa sp. nov.; Western Palaearctic; molecular markers; cuticular hydrocarbons

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Agosti D. 1994. The phylogeny of the ant tribe Formicini (Hymenoptera, Formicidae) with the description of a new genus. Systematic Entomology 19: 93–117.

Agosti D. & Collingwood C.A. 1987. A provisional list of the Balkan ants (Hym. Formicidae) with a key to the worker caste. II. Key to the worker caste, including the European species without the Iberian. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 60: 261–293.

Ballard J.W.O. 2000. When one is not enough: Introgression of mitochondrial DNA in Drosophila. Molecular Biology and Evolution 17: 1126–1130.

Belshaw R. & Quicke D.L.J. 1997. A molecular phylogeny of the Aphidiinae (Hymenoptera: Braconidae). Molecular Phylogenetics and Evolution 7: 281–293.

Bernard F. 1968. Faune de l’Europe et du Bassin Méditerranéen. 3. Les Fourmis (Hymenoptera Formicidae) d’Europe occidentale et septentrionale. Masson, Paris.

Bolton B. 2014. An Online Catalog of the Ants of the World. Available from [accessed 26 Jan. 2017].

Bondroit J. 1918. Les fourmis de France et de Belgique. Annales de la Société Entomologique de France 87: 1–174.

Chan K.M.A. & Levin S.A. 2005. Leaky prezygotic isolation and porous genomes: Rapid introgression of maternally inherited DNA. Evolution 59: 720–729.

Collingwood C.A. 1956. Ant hunting in France. L’entomologiste 89: 105–108.

Collingwood C.A. 1976. A provisional list of Iberian Formicidae with a key to the worker caste. EOS – Revista Española de Entomologia 52: 65–95.

Collingwood C.A. & Yarrow I.H.H. 1969. A survey of Iberian Formicidae. EOS – Revista Espanola de Entomologia 44: 53–101.

Dahbi A., Cerda X., Hefetz A. & Lenoir A. 1996. Social closure, aggressive behavior, and cuticular hydrocarbon profiles in the polydomous ant Cataglyphis iberica (Hymenoptera, Formicidae). Journal of Chemical Ecology 22: 2173–2186.

Dlussky G.M. 1969. Ants of the genus Proformica Ruzs. of the USSR and contiguous countries (Hymenoptera: Formicidae). Zoologicheskii Zhurnal 48: 217–232. [in Russian]

Edgar R.C. 2004. MUSCLE: multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32: 1792–1797.

Emery C. 1909. Beiträge zur Monographie der Formiciden des paläarktischen Faunengebietes (Hym.). Teil VII. Deutsche Entomologische Zeitschrift 1909: 179–204.

Emery C. 1925. Hymenoptera, Fam. Formicidae, subfam. Formicinae. Genera Insectorum 183: 1–302.

Errard C., Ruano F., Richard F.J., Lenoir A., Tinaut A. & Hefetz A. 2006. Co-evolution-driven cuticular hydrocarbon variation between the slave-making ant Rossomyrmex minuchae and its host Proformica longiseta (Hymenoptera: Formicidae). Chemoecology 16: 235–240.

Espadaler X. & Cagniant H. 1987. Contribution à la connaissance des fourmis marocaines. Description du mâle de Proformica theryi Santschi, 1936 (Hymenoptera, Formicidae). Nouvelle Revue d’Entomologie 4: 133–138.

Ferchaud A.L., Lyet A., Cheylan M., Arnal V., Baron J.P., Montgelard C. & Ursenbacher S. 2011. High genetic differentiation among French populations of the Orsini’s Viper (Vipera ursinii ursinii) based on mitochondrial and microsatellite data: implications for conservation management. Journal of Heredity 102: 67–78.

Forel A. 1886. Études myrmécologiques en 1886. Annales de la Société Entomologique de Belgique 30: 131–215.

Goropashnaya A.V., Fedorov V.B., Seifert B. & Pamilo P. 2004. Limited phylogeographical structure across Eurasia in two red wood ant species Formica pratensis and F. lugubris (Hymenoptera, Formicidae). Molecular Ecology 13: 1849–1858.

Goropashnaya A.V., Fedorov V.B., Seifert B. & Pamilo P. 2007. Phylogeography and population structure in the ant Formica exsecta (Hymenoptera, Formicidae) across Eurasia as reflected by mitochondrial DNA variation and microsatellites. Annales Zoologici Fennici 44: 462–474.

Goropashnaya A.V., Fedorov V.B., Seifert B. & Pamilo P. 2012. Phylogenetic relationships of Palaearctic Formica species (Hymenoptera, Formicidae) based on mitochondrial cytochrome b sequences. PLoS ONE 7: e41697.

Hasegawa E., Tinaut A. & Ruano F. 2002. Molecular phylogeny of two slave-making ants: Rossomyrmex and Polyergus (Hymenoptera: Formicidae). Annales Zoologici Fennici 39: 267–271.

Hebert P.D.N., Penton E.H., Burns J.M., Janzen D.H. & Hallwachs W. 2004. Ten species in one: DNA barcoding reveals cryptic species in the neotropical skipper butterfly Astraptes fulgerator. Proceedings of the National Academy of Sciences 101: 14812–14817.

Lanfear R., Calcott B., Ho S.Y.W. & Guindon S. 2012. PartitionFinder: combined selection of partitioning schemes and substitution models for phylogenetic analyses. Molecular Biology and Evolution 29: 1695–1701.

Lenoir A., Depickere S., Devers S., Christides J.P. & Detrain C. 2009. Hydrocarbons in the ant Lasius niger: from the cuticle to the nest and home range marking. Journal of Chemical Ecology 35: 913–921.

Liautard C. & Keller L. 2001. Restricted effective queen dispersal at a microgeographic scale in polygynous populations of the ant Formica exsecta. Evolution 55: 2484–2492.

Linnen C.R. & Farrell B.D. 2007. Mitonuclear discordance is caused by rampant mitochondrial introgression in Neodiprion (Hymenoptera: Diprionidae) sawflies. Evolution 61: 1417–1438.

Martin A.P. & Palumbi S.R. 1993. Body size, metabolic rate, generation time, and the molecular clock. Proceedings of the National Academy of Sciences 90: 4087–4091.

Martin S.J., Helanterä H. & Drijfhout F.P. 2008. Evolution of species-specific cuticular hydrocarbon patterns in Formica ants. Biological Journal of the Linnean Society 95: 131–140.

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.

Nowbahari E., Lenoir A., Clément J.L., Lange C., Bagnères A.G. & Joulie C. 1990. Individual, geographical and experimental variation of cuticular hydrocarbons of the ant Cataglyphis cursor (Hymenoptera: Formicidae): their use in nest and subspecies recognition. Biochemical Systematics and Ecology 18: 63–73.

Nylander W. 1856. Synopsis des formicides de France et d’Algérie. Annales des Sciences Naturelles: Zoologie et Biologie Animale, 4th Series 5: 51–109. Available from [accessed 12 Feb. 2017].

Oppelt A., Spitzenpfeil N., Kroiss J. & Heinze J. 2008. The significance of intercolonial variation of cuticular hydrocarbons for inbreeding avoidance in ant sexuals. Animal Behaviour 76: 1029–1034.

Radchenko A. 2007. The ants (Hymenoptera, Formicidae) in the collection of William Nylander. Fragmenta Faunistica (Warsaw) 50: 27–41.

Reyment R.A. 1989. Compositional data analysis. Terra Review 1: 29–34.

Ronquist F., Teslenko M., van der Mark P., Ayres D.L., Darling A., Hohna S., Larget B., Liu L., Suchard M.A. & Huelsenbeck J.P. 2012. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61: 539–542.

Ruano F., Devers S., Sanllorente O., Errard C., Tinaut A. & Lenoir A. 2011. A geographical mosaic of coevolution in a slave-making host–parasite system. Journal of Evolutionary Biology 24: 1071–1079.

Ruzsky M. 1905. The ants of Russia. Trudy Obshchestva Estestvoispytatelei pri Imperatorskom Kazanskom Universitete 38: 1–800. [in Russian]

Sanllorente O., Lorite P., Devers S., Ruano F., Lenoir A. & Tinaut A. 2012. The spatial distribution does not affect host–parasite coevolution in Rossomyrmex ants. Insectes Sociaux 59: 361–368.

Sanllorente O., Ruano F. & Tinaut A. 2015. Large-scale population genetics of the mountain ant Proformica longiseta (Hymenoptera: Formicidae). Population Ecology 57: 637–648.

Santschi F. 1925. Fourmis d’Espagne et autres espèces paléarctiques (Hymenopt.). EOS – Revista Española de Entomologia 1: 339–360.

Saux C., Fisher B.L. & Spicer G.S. 2004. Dracula ant phylogeny as inferred by nuclear 28S rDNA sequences and implications for ant systematics (Hymenoptera: Formicidae: Amblyoponinae). Molecular Phylogenetics and Evolution 33: 457–468.

Schlick-Steiner B.C., Steiner F.M., Seifert B., Stauffer C., Christian E. & Crozier R.H. 2010. Integrative taxonomy: a multisource approach to exploring biodiversity. Annual Review of Entomology 55: 421–438.

Seifert B. 2007. Die Ameisen Mittel- und Nordeuropas. Lutra, Görlitz, Germany.

Smith A.A., Millar J.G., Hanks L.M. & Suarez A.V. 2013. A conserved fertility signal despite population variation in the cuticular chemical profile of the trap-jaw ant Odontomachus brunneus. Journal of Experimental Biology 216: 3917–3924.

Stamatakis A., Hoover P. & Rougemont J. 2008. A rapid bootstrap algorithm for the RAxML web servers. Systematic Biology 57: 758–771.

Stumper R. 1957. Sur l’éthologie de la fourmi a miel Proformica nasuta Nyl. (Étude myrmécologique LXXVIII). Bulletin de la Société des Naturalistes Luxembourgeois 60: 87–97.

Tamura K., Peterson D., Peterson N., Stecher G., Nei M. & Kumar S. 2011. MEGA5: Molecular Evolutionary Genetics Analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods. Molecular Biology and Evolution 28: 2731–2739.

Ward P.S. & Downie D.A. 2005. The ant subfamily Pseudomyrmecinae (Hymenoptera: Formicidae): phylogeny and evolution of big-eyed arboreal ants. Systematic Entomology 30: 310–335.

Wheeler W.M. 1913. A revision of the ants of the genus Formica (Linné) Mayr. Bulletin of the Museum of Comparative Zoology 53: 379–565.

Zinenko O., Stümpel N., Mazanaeva L., Bakiev A., Shiryaev K., Pavlov A., Kotenko T., Kukushkin O., Chikin Y., Duisebajeva T., Nilson G., Orlov N.L., Tuniyev S., Ananjeva N.B., Murphy R.W. & Joger U. 2015. Mitochondrial phylogeny shows multiple independent ecological transitions and northern dispersion despite of Pleistocene glaciations in meadow and steppe vipers (Vipera ursinii and Vipera renardi). Molecular Phylogenetics and Evolution 84: 85–100.


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