Skip to main content Skip to main navigation menu Skip to site footer
Type: Article
Published: 2025-09-09
Page range: 257-276
Abstract views: 137
PDF downloaded: 4

Inflated taxonomy in the West Mediterranean Temnothorax algiricus complex (Hymenoptera, Formicidae) revealed by quantitative morphology

Institute of Evolutionary Biology (CSIC-Univ. Pompeu Fabra); Passeig Marítim de la Barceloneta; 37-49; 08003 Barcelona; Spain; Department of Chemistry; Life Sciences; and Environmental Sustainability; University of Parma; Parco Area delle Scienze 11/A; 43124 Parma; Italy
Institute of Evolutionary Biology (CSIC-Univ. Pompeu Fabra); Passeig Marítim de la Barceloneta; 37-49; 08003 Barcelona; Spain; Museum für Naturkunde; Leibniz Institute for Evolution and Biodiversity Science; Center for Integrative Biodiversity Discovery; Invalidenstraße 43; Berlin
HUN-REN-ELTE-MTM Integrative Ecology Research Group; Pázmány Péter ave 1/C; Budapest 1117; Hungary; Department of Systematic Zoology and Ecology; Institute of Biology; ELTE-Eötvös Loránd University; Pázmány Péter ave 1/C; Budapest 1117; Hungary
Institute of Evolutionary Biology (CSIC-Univ. Pompeu Fabra); Passeig Marítim de la Barceloneta; 37-49; 08003 Barcelona; Spain
Hymenoptera ant taxonomy Myrmicinae Crematogastrini morphometrics

Abstract

Temnothorax is one of the largest ant genera, with most taxa described from the Mediterranean. The western Mediterranean Temnothorax algiricus complex (Hymenoptera, Formicidae) is a well-defined lineage of dark-reddish arboreal species, whose taxonomy has never been comprehensively addressed. We conducted a comprehensive taxonomic revision using quantitative morphological analysis complemented by qualitative observations to clarify the systematic relationships within this group. Our analysis of 13 previously described taxa reveals only three valid species: T. algiricus (Forel, 1894), T. atlantis (Santschi, 1911), and T. trabutii (Forel, 1894). We establish several new synonymies: Leptothorax angustulus brunea Santschi, 1918, Leptothorax gazella Santschi, 1932, Leptothorax gazella monticola Santschi, 1932, Temnothorax mediterraneus Ward, Brady, Fisher, & Schultz, 2014, and Temnothorax atlantis veneris Galkowski & Cagniant, 2017 are considered junior subjective synonyms of T. algiricus. Leptothorax kiudiria Espadaler, 1997, Leptothorax angustulus silvanus (Forel, 1907), Temnothorax atlantis suturalis Galkowski & Cagniant, 2017, and Temnothorax continentalis Galkowski & Cagniant, 2017 are synonymized under T. atlantis. Leptothorax lindbergi Santschi, 1931 is considered a synonym of T. trabutii. Leptothorax convexus timida Santschi, 1912 is a junior synonym of Temnothorax convexus (Forel, 1894), and excluded from the complex. This revision resolves the oversplit nomenclature characteristic of North African myrmecology and establishes taxonomic consistency with European classifications. Our quantitative morphology-based framework provides the foundation for future integrative taxonomic studies investigating intraspecific variation and potential cryptic diversity within this ecologically important ant lineage.

 

References

  1. Arcos, J., Chaves, D., Alarcón, P. & Rosado, Á. (2022) First record of Temnothorax convexus (Forel, 1894) in Portugal (Hymenoptera: Formicidae) with an updated checklist of the ants from the country. Sociobiology, 692, e7623. https://doi.org/10.13102/sociobiology.v69i2.7623
  2. Arcos, J. & García, F. (2023) Hormigas de la península Ibérica e islas Baleares. Independently published, Spain, 491 pp.
  3. Arcos, J. & Alarcón, P. (2024) Confirmed presence of the enigmatic ant Aphaenogaster gemella (Hymenoptera: Formicidae) in Iberia. The Journal of Basic and Applied Zoology, 85 (1), 21. https://doi.org/10.1186/s41936-024-00375-9
  4. Baroni Urbani, C. (1971) Catalogo delle specie di Formicidae d’Italia (Studi sulla mirmecofauna d’Italia X). Memorie della Societa Entomologica Italiana, 50, 5–287.
  5. Bernard, F. (1967) Faune de l’Europe et du Bassin Méditerranéen. 3. Les fourmis (Hymenoptera Formicidae) d’Europe occidentale et septentrionale. Masson, Paris, 411 pp.
  6. Bishop, T.R., Robertson, M.P., Gibb, H., van Rensburg, B.J., Braschler, B., Chown, S.L., Ford, S.H., Munyai, T.C. & Parr, C.L. (2016) Ant assemblages have darker and larger members in cold environments. Global Ecology and Biogeography, 25 (12), 1489–1499. https://doi.org/10.1111/geb.12516
  7. Bolton, B. (1995) A new general catalogue of the ants of the world. Harvard University Press, Cambridge, Mass., 504 pp.
  8. Bolton, B. (2025) An online catalog of the ants of the world. Available from: https://antcat.org.
  9. Bondroit, J. (1918) Les fourmis de France et de Belgique. Annales de la Société Entomologique de France, 87, 1–174. https://doi.org/10.1080/21686351.1918.12279608
  10. Borowiec, L. (2014) Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus, 25, 1–340.
  11. Borowiec, L., Lebas, C. & Salata, S. (2022) Notes on ants (Hymenoptera: Formicidae) from three northern Aegean islands – Lemnos, Samothraki and Thasos. Annals of the Upper Silesian Museum in Bytom Entomology, 31, 1–14.
  12. Bračko, G. (2006) Review of the ant fauna (Hymenoptera: Formicidae) of Croatia. Acta Entomologica Slovenica, 14 (2), 131–156.
  13. Cagniant, H. (1968) Liste préliminaire de fourmis forestières d’Algérie. Résultats obtenus de 1963 à 1966. Bulletin de la Société d’Histoire Naturelle de Toulouse, 104, 138–147.
  14. Cagniant, H. (1970) Deuxième liste de fourmis d’Algérie récoltées principalement en forêt. (1re partie). Bulletin de la Société d’Histoire Naturelle de Toulouse, 105, 405–430.
  15. Cagniant, H. (2006) Liste actualisee des fourmis du Maroc (Hymenoptera: Formicidae). Myrmecologische Nachrichten, 8, 193–200.
  16. Cagniant, H. & Espadaler, X. (1997) Les Leptothorax, Epimyrma et Chalepoxenus du Maroc (Hymenoptera: Formicidae). Clé et catalogue des espèces. Annales de la Société Entomologique de France, New Series, 33 (3), 259–284. https://doi.org/10.1080/21686351.1997.12277888
  17. Casevitz-Weulersse, J. & Galkowski, C. (2009) Liste actualisée des Fourmis de France (Hymenoptera, Formicidae). Bulletin de la Société Entomologique de France, 114 (4), 475–510. https://doi.org/10.3406/bsef.2009.2724
  18. Csősz, S., Heinze, J. & Mikó, I. (2015) Taxonomic synopsis of the Ponto-Mediterranean ants of Temnothorax nylanderi species-group. PLoS One, 10 (11), e0140000. https://doi.org/10.1371/journal.pone.0140000
  19. Csősz, S., Báthori, F., Rádai, Z., Herczeg, G. & Fisher, B.L. (2023) Comparing ant morphology measurements from microscope and online AntWeb.org 2D z‐stacked images. Ecology and Evolution, 13 (3), e9897. https://doi.org/10.1002/ece3.9897
  20. Csősz, S., Alicata, A., Báthori, F., Galkowski, C., Schifani, E., Yusupov, Z., Herczeg, G. & Prebus, M.M. (2025a) Integrative taxonomy reveals inflated biodiversity in the European Temnothorax unifasciatus complex (Hymenoptera: Formicidae). Zoologica Scripta, 54 (1), 33–49. https://doi.org/10.1111/zsc.12690
  21. Csősz, S., Taheri, A., Schifani, E., Reyes-López, J-L., Alicata, A., Báthori, F. & Prebus, M.M. (2025b) Taxonomic revision of the Mediterranean Temnothorax rottenbergii species group via integrated morphological and molecular approaches. Insect Systematics and Diversity. [in press]
  22. Emery, C. (1916) Fauna entomologica italiana. I. Hymenoptera-Formicidae. Bollettino della Società Entomologica Italiana, 47, 79–275.
  23. Emery, C. (1924) Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [concl.]. Genera Insectorum, 174C, 207–397.
  24. Espadaler, X. & Collingwood, C.A. (1982) Notas sobre Leptothorax Mayr, 1855, con descripción de L. gredosi n. sp. (Hym. Formicidae). Boletín de la Asociación Española de Entomologia, 6, 41–48.
  25. Forel, A. (1894) Les formicides de la province d’Oran (Algerie). Bulletin de la Société Vaudoise des Sciences Naturelles, 30, 1–45.
  26. Forel, A. (1907) Fourmis nouvelles de Kairouan et d’Orient. Annales de la Société Entomologique de Belgique, 51, 201–208. https://doi.org/10.5962/bhl.part.1582
  27. Galkowski, C. & Lebas, C. (2016) Temnothorax conatensis nov. sp., décrite des Pyrénées-Orientales (France) (Hymenoptera, Formicidae). Revue de l’Association Roussillonnaise d’Entomologie, 25, 80–87.
  28. Galkowski, C. & Cagniant, H. (2017) Contribution à la connaissance des fourmis du groupe angustulus dans le genre Temnothorax (Hymenoptera, Formicidae). Revue de l’Association Roussillonnaise d’Entomologie, 26, 180–191.
  29. Giannetti, D., Schifani, E., Gugliuzzo, A., Zappalà, L., Biondi, A. & Grasso, D.A. (2022) Native European ants can discourage host colonization and reduce reproductive success of the invasive ambrosia beetle Xylosandrus compactus. Biological Control, 174, 105032. https://doi.org/10.1016/j.biocontrol.2022.105032
  30. Guénard, B., Weiser, M., Gomez, K., Narula, N. & Economo, E.P. (2017) The Global Ant Biodiversity Informatics (GABI) database: a synthesis of ant species geographic distributions. Myrmecological News, 24, 83–89.
  31. Guillem, R. (2009) A survey of the ants of Minorca (Hymenoptera: Formicidae) with two new species for the island: Hypoponera punctatissima (Roger, 1859) and Temnothorax algiricus (Forel, 1894). Boletin de la Asociacion Espanola de Entomologia, 33 (3–4), 447–460.
  32. Henin, J.M., Paiva, M.R. & Collingwood, C.A. (2001) The Iberian Leptothorax Mayr, 1855 (Hymenoptera, Formicidae): Discovery of a new species and first record of L. atlantis Santschi, 1911. Boletim da Sociedade Portuguesa de Entomologia, 7 (14), 161–166.
  33. Kahle, D. & Wickham, H. (2013) ggmap: Spatial Visualization with ggplot2. The R Journal, 5, 1. https://doi.org/10.32614/RJ-2013-014
  34. Lebas, C., Galkowski, C., Blatrix, R. & Wegnez, P. (2016) Guide Delachaux: Fourmis d’Europe occidentale. Delachaux et Niestlé, Luçon, 415 pp.
  35. Law, S.J., Bishop, T.R., Eggleton, P., Griffiths, H., Ashton, L. & Parr, C. (2020) Darker ants dominate the canopy: Testing macroecological hypotheses for patterns in colour along a microclimatic gradient. Journal of Animal Ecology, 89 (2), 347–359. https://doi.org/10.1111/1365-2656.13110
  36. Oberski, J.T., Griebenow, Z.H., Adams, R.M.M., Andersen, A., Andrade-Silva, J., Barden, P., Borowiec, M., Brady, S., Casadei-Ferreira, A., Csősz, S., Martins Dias, A., Dias, R.K.S., Feitosa, R.M., Fernandez, F., Fisher, B.L., General, D.E.M., Gomez, K., Hammel, J.U., Hawkes, P.G., Janda, M., Khalife, A., Ladino, N., Lieberman, Z.E., Lucky, A., Menchetti, M., Pires do Prado, L., Prebus, M.M., Probst, R.S., Punnath, A., Richter, A., Salata, S., Sánchez-Restrepo, A.F., Schifani, E., Schultz, T.R., Silva, R.R., Sosa-Calvo, J., Tocora, M.C., Ulysséa, M.A., van de Kamp, T., Wang, W.Y., Williams, J.L., Procopio Camacho, G. & Boudinot, B.E. (2025) Ant systematics: Past, present, and future. Insect Systematics and Diversity. [in press]
  37. Ogle, D.H., Doll, J.C., Wheeler, A.P. & Dinno, A. (2025) FSA: Simple Fisheries Stock Assessment Methods. Available from: https://cran.r-project.org/web/packages/FSA/index.html (accessed 3 August 2025) https://doi.org/10.32614/CRAN.package.FSA
  38. 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
  39. R Core Team (2025) R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna. Available from: https://www.R-project.org/ (accessed 3 August 2025)
  40. Rigato, S. & Toni, I. (2011) Short notes 21. Hymenoptera, Formicidae. In: Nardi, G., Whitmore, D., Bardiani, M., Birtele, D., Mason, F., Spada, L. & Cerretti, P. (Eds.), Biodiversity of Marganai and Montimannu (Sardinia). Cierre Edizioni, Sommacampagna, Verona, pp. 873–882.
  41. RStudio Team (2025) RStudio: Integrated Development for R. RStudio. PBC, Boston, Massachusetts. Available from: http://www.rstudio.com/ (accessed 3 August 2025)
  42. Salata, S. & Borowiec, L. (2018) Taxonomic and faunistic notes on Greek ants (Hymenoptera: Formicidae). Annals of the Upper Silesian Museum in Bytom Entomology, 27, 1–51.
  43. Santschi, F. (1911) Formicides nouveaux de l’Afrique Mineure (4e note). Bulletin de la Société d’Histoire Naturelle de l’Afrique du Nord, 2, 11–14.
  44. Santschi, F. (1912) Quelques nouvelles variétés de fourmis africaines. Bulletin de la Société d’Histoire Naturelle de l’Afrique du Nord, 3, 147–149.
  45. Santschi, F. (1918) Leptothorax nouveaux de l’Afrique Mineure. Bulletin de la Société d’Histoire Naturelle de l’Afrique du Nord, 9, 31–38.
  46. Santschi, F. (1923) Messor et autres fourmis paléarctiques. Revue Suisse de Zoologie, 30, 317–336.
  47. Schembri, S.P. & Collingwood, C.A. (1995) The myrmecofauna of the Maltese Islands. Remarks and additions (Hymenoptera Formicidae). Bollettino della Società Entomologica Italiana, 127, 153–158.
  48. Schifani, E., Giannetti, D., Csősz, S., Castellucci, F., Luchetti, A., Castracani, C., Spotti, F.A., Mori, A. & Grasso, D.A. (2022) Is mimicry a diversification-driver in ants? Biogeography, ecology, ethology, genetics and morphology define a second West-Palaearctic Colobopsis species (Hymenoptera: Formicidae). Zoological Journal of the Linnean Society, 194, 1424–1450. https://doi.org/10.1093/zoolinnean/zlab035
  49. Schifani, E., Prebus, M.M. & Alicata, A. (2022) Integrating morphology with phylogenomics to describe four island endemic species of Temnothorax from Sicily and Malta (Hymenoptera, Formicidae). European Journal of Taxonomy, 833, 143–179. https://doi.org/10.5852/ejt.2022.833.1891
  50. Schifani, E. & Alicata, A. (2023) Nomenclatural changes on some Mediterranean Aphaenogaster Mayr, 1853 taxa (Hymenoptera, Formicidae). Zootaxa, 5277 (1), 59–70. https://doi.org/10.11646/zootaxa.5277.1.2
  51. Schifani, E., Alicata, A., Prebus, M.M. & Csősz, S. (2025) Integrative description of Temnothorax siculus sp. n.: a new ant species endemic to Sicily, Italy (Hymenoptera, Formicidae). Diversity, 17 (4), 194. https://doi.org/10.3390/d17040294
  52. Schneider, C.A., Rasband, W.S. & Eliceiri, K.W. (2012) NIH Image to ImageJ: 25 years of image analysis. Nature Methods, 9, 671–675. https://doi.org/10.1038/nmeth.2089
  53. Seifert, B. (2020) The gene and gene expression (GAGE) species concept: An universal approach for all eukaryotic organisms. Systematic Biology, 69, 1033–1038. https://doi.org/10.1093/sysbio/syaa032
  54. Venables, W.N. & Ripley, B.D. (2002) Modern Applied Statistics with S. 4th Edition. Springer, New York, New York, 510 pp. https://doi.org/10.1007/978-0-387-21706-2
  55. Wang, R., Kass, J.M., Galkowski, C., Garcia, F., Hamer, M.T., Radchenko, A., Salata, S., Schifani, E., Yusupov, Z.M., Economo, E.P. & Guénard, B. (2023) Geographic and climatic constraints on bioregionalization of European ants. Journal of Biogeography, 50, 503–514. https://doi.org/10.1111/jbi.14546
  56. 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 (Hymenoptera: Formicidae). Systematic Entomology, 40, 61–81. https://doi.org/10.1111/syen.12090

How to Cite

Schifani, E., Menchetti, M., Csősz, S. & Vila, R. (2025) Inflated taxonomy in the West Mediterranean Temnothorax algiricus complex (Hymenoptera, Formicidae) revealed by quantitative morphology. Zootaxa, 5691 (2), 257–276. https://doi.org/10.11646/zootaxa.5691.2.4