Eur. J. Entomol. 122: 372-380, 2025 | DOI: 10.14411/eje.2025.042

Annual dynamics of subterranean ants (Hymenopetra: Formicidae) in a cork oak forest in Morocco (Northwest Africa)Original article

Bilal EL AJJOURI ORCID...1, Joaquin L. REYES-LÓPEZ ORCID...2, Ahmed TAHERI ORCID...1
1 LESCB URL-CNRST N◦ 18, Faculty of Sciences of Tetouan, Abdelmalek Essaadi University, Morocco; e-mails: elajjouri.bilal@gmail.com, a.taheri@uae.ac.ma
2 Area of Ecology, University of Cordoba, Building C-4 "Celestino Mutis", Rabanales Campus, Cordoba, Spain; e-mail: cc0reloj@uco.es

Subterranean ants play a critical role in soil ecosystems, yet their cryptic lifestyle and the challenges associated with their sampling hinder comprehensive understanding of their biodiversity. This study examines the seasonal dynamics of subterranean ant communities in a cork oak forest in northwestern Morocco. We employed an adapted soil-washing method. that enabled the collection of 1,318 individuals from 17 species, of which 41.18% were identified as hypogaeic. The analysis revealed significant differences in community composition across seasons, with summer exhibiting notably higher species richness and worker abundance. Non-hypogaeic ants showed strong positive correlations with temperature and negative correlations with rainfall, while hypogaeic ants exhibited weak and non-significant responses to climatic variables, likely due to their ecological specialization. These results suggest that climatic factors distinctly influence ant subgroups and highlight the ecological relevance of subterranean ants in soil biodiversity. The study underscores the importance of tailored sampling methodologies for capturing the diversity and dynamics of these often-overlooked communities.

Keywords: Biodiversity, hypogaeic fauna, seasonal dynamics, soil

Received: February 28, 2025; Revised: November 7, 2025; Accepted: November 7, 2025; Published online: December 11, 2025  Show citation

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EL AJJOURI, B., REYES-LÓPEZ, J.L., & TAHERI, A. (2025). Annual dynamics of subterranean ants (Hymenopetra: Formicidae) in a cork oak forest in Morocco (Northwest Africa). EJE, 122, Article 372-380. https://doi.org/10.14411/eje.2025.042
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    References

    1. Afi A., Achhal El Kadmiri A., Benabid A. & Rochdi M. 2005: Richesse et diversité floristique de la suberaie de la Mamora (Maroc). - Acta Bot. Malacit. 30: 127-138. Go to original source...
    2. Andersen A.N. & Brault A. 2010: Exploring a new biodiversity frontier: Subterranean ants in northern Australia. - Biodiv. Conserv. 19: 2741-2750. Go to original source...
    3. Andersen A.N. & Majer J.D. 2004: Ants show the way Down Under: Invertebrates as bioindicators in land management. - Front. Ecol. Environ. 2: 291-298. Go to original source...
    4. Andersen A.N., Hoffmann B.D., Müller W.J. & Griffiths A.D. 2002: Using ants as bioindicators in land management: Simplifying assessment of ant community responses. - J. Appl. Ecol. 39: 8-17. Go to original source...
    5. Anderson J.M. & Ingram J.S.I. 1989: Tropical Soil Biology and Fertility: A Handbook of Methods. CABI, Wallingford, 171 pp.
    6. Barroso Rodríguez Á. 2012: Habitat Use by the Mediterranean Ant Aphaenogaster senilis in Doñana: Competition and Resource Exploitation. PhD thesis, University of Sevilla, 174 pp. [in Spanish].
    7. Berghoff S.M., Maschwitz U. & Linsenmair K.E. 2003: Hypogaeic and epigaeic ant diversity on Borneo: Evaluation of baited sieve buckets as a study method. - Tropical Zool. 16: 153-163. Go to original source...
    8. Bolton B. 2000: The ant tribe Dacetini. - Mem. Am. Entomol. Inst. 65: 1-1028.
    9. Bolton B. & Fisher B.L. 2011: Taxonomy of Afrotropical and West Palaearctic ants of the ponerine genus Hypoponera Santschi (Hymenoptera: Formicidae). - Zootaxa 2843: 1-118. Go to original source...
    10. Borowiec L. & Salata S. 2022: A monographic review of ants of Greece (Hymenoptera: Formicidae). - Nat. Hist. Monogr. Upper Silesian Mus. 1: 1-300.
    11. Cagniant H. 1996: Les Aphaenogaster du Maroc (Hymeno­ptera: Formicidae): Clé et catalogue des espèces. - Ann. Soc. Entomol. Fr. (N.S.) 32: 67-85. Go to original source...
    12. Cagniant H. 1997: Le genre Tetramorium au Maroc (Hymeno­ptera: Formicidae): Clé et catalogue des espèces. - Ann. Soc. Entomol. Fr. (N.S.) 33: 89-100. Go to original source...
    13. Cagniant H. 2005: Les Crematogaster du Maroc (Hym., Formicidae). Clé de détermination et commentaires. - Orsis 20: 7-12.
    14. Cagniant H. 2006: Liste actualisée des fourmis du Maroc (Hymenoptera: Formicidae). - Myrmecol. Nachr. 8: 193-200.
    15. Cagniant H. & Espadaler X. 1997: Les Leptothorax, Epimyrma et Chalepoxenus du Maroc (Hymenoptera: Formicidae). Clé et catalogue des espèces. - Ann. Soc. Entomol. Fr. (N.S.) 33: 259-284. Go to original source...
    16. Castro D., Fernández F., Meneses A., Tocora M., Sanchez S. & Peña-Venegas C. 2018: A preliminary checklist of soil ants (Hymenoptera: Formicidae) of Colombian Amazon. - Biodiv. Data J. 6: e29278, 23 pp. Go to original source...
    17. Cerda X., Retana J. & Cros S. 1997: Thermal disruption of transitive hierarchies in Mediterranean ant communities. - J. Anim. Ecol. 66: 363-374. Go to original source...
    18. Climate Data 2021: Climat Larache: Température de l'eau à, Température moyenne Larache, Pluviométrie, diagramme ombrothermique pour Larache. URL: https://fr.climate-data.org/afrique/maroc/larache/larache-4963/#climate-table
    19. Cros S., Cerdá X. & Retana J. 1997: Spatial and temporal variations in the activity patterns of Mediterranean ant communities. - Écoscience 4: 269-278. Go to original source...
    20. Culliney T. 2013: Role of arthropods in maintaining soil fertility. - Agriculture 3: 629-659. Go to original source...
    21. Culver D.C. & Pipan T. 2014: Shallow Subterranean Sabitats: Ecology, Evolution, and Conservation. 1st ed. Oxford University Press, Oxford, UK, 258 pp. Go to original source...
    22. Dunn R.R., Agosti D., Andersen A.N., Arnan X., Bruhl C.A., Cerdá X., Ellison A.M., Fisher B.L., Fitzpatrick M.C., Gibb H. et al. 2009: Climatic drivers of hemispheric asymmetry in global patterns of ant species richness. - Ecol. Lett. 12: 324-333. Go to original source...
    23. Eguchi K., Hashimoto Y. & Malsch A.K.F. 2006: Pheidole schoedli sp.n. (Hymenoptera: Formicidae), a subterranean species found from North Borneo. - Myrmecol. Nachr. 8: 31-34.
    24. Eguchi K., Bui T.V., General D.M. & Alpert G.D. 2010: Revision of the ant genus Anillomyrma Emery, 1913 (Hymeno­ptera: Formicidae: Myrmicinae: Solenopsidini). - Myrmecol. News 13: 31-36.
    25. Eisenbeis G. & Wichard W. 1987: Atlas on the Biology of Soil Arthropods. Springer, Berlin, Heidelberg, 437 pp. Go to original source...
    26. El Keroumi A., Naamani K., Soummane H. & Dahbi A. 2012: Seasonal dynamics of ant community structure in the Moroccan Argan Forest. - J. Insect Sci. 12: 94, 19 pp. Go to original source...
    27. Ellison A.M. 2012: Out of Oz: Opportunities and challenges for using ants (Hymenoptera: Formicidae) as biological indicators in north-temperate cold biomes. - Myrmecol. News 17: 105-119.
    28. Fisher J., Beames L., Rangers B.J., Rangers N.N., Majer J. & Heterick B. 2014: Using ants to monitor changes within and surrounding the endangered Monsoon Vine Thickets of the tropical Dampier Peninsula, northwestern Australia. - Forest Ecol. Manag. 318: 78-90. Go to original source...
    29. Folgarait P.J., Monjeau J.A. & Kittlein M. 1998: Solving the enigma of granivory rates in Patagonia and throughout other deserts of the world: Is thermal range the explanation? - Ecología Austral 8: 251-263.
    30. Fowler H.G. & Delabie J.H.C. 1995: Resource partitioning among epigaeic and hypogaeic ants (Hymenoptera: Formicidae) of a Brazilian cocoa plantation. - Ecología Austral 5: 117-124.
    31. Houadria M. & Menzel F. 2021: Digging deeper into the ecology of subterranean ants: Diversity and niche partitioning across two continents. - Diversity 13: 53, 15 pp. Go to original source...
    32. Jacquemin J., Roisin Y. & Leponce M. 2016: Spatio-temporal variation in ant (Hymenoptera: Formicidae) communities in leaf-litter and soil layers in a premontane tropical forest. - Myrmecol. News 22: 129-139.
    33. Jiménez-Valverde A., Gilgado J.D., Sendra A., Pérez-Suárez G., Herrero-Borgoñón J.J. & Ortuño V.M. 2015: Exceptional invertebrate diversity in a scree slope in Eastern Spain. - J. Insect Conserv. 19: 713-728. Go to original source...
    34. Kalule-Sabiti J.M.N. 1980: The ecological role of ants (Formicidae) in the ecosystem. - Afr. J. Ecol. 18: 113-121. Go to original source...
    35. Kaspari M., O'Donnell S. & Kercher J.R. 2000: Energy, density, and constraints to species richness: Ant assemblages along a productivity gradient. - Am. Nat. 155: 280-293. Go to original source...
    36. Kaspari M., Ward P.S. & Yuan M. 2004: Energy gradients and the geographic distribution of local ant diversity. - Oecologia 140: 407-413. Go to original source...
    37. Kottek M., Grieser J., Beck C., Rudolf B. & Rubel F. 2006: World Map of the Köppen-Geiger climate classification updated. - Meteorol. Zeitschr. 15: 259-263. Go to original source...
    38. Kugler J. & Ionescu-Hirsch A. 2007: Anochetus bytinskii, a new ant species from Israel (Hymenoptera: Formicidae). - Israel J. Entomol. 37: 287-298.
    39. Lavelle P. 1997: Faunal activities and soil processes: Adaptive strategies that determine ecosystem function. - Adv. Ecol. Res. 27: 93-132. Go to original source...
    40. Lavelle P. & Spain A.V. 2001: Soil organisms. In: Soil Ecology. Springer, Dordrecht, pp. 201-356. Go to original source...
    41. Lemagnen R. 2009: Techniques: Tamisages, lavages et autres berlèses. - Invertébrés Armoricains 3: 19-22.
    42. Longino J.T. & Colwell R.K. 1997: Biodiversity assessment using structured inventory: Capturing the ant fauna of a tropical rain forest. - Ecol. Appl. 7: 1263-1277. Go to original source...
    43. Majer J.D., Kitching R.L., Heterick B.E., Hurley K. & Brennan K.E.C. 2001: North-south patterns within arboreal ant assemblages from rain forests in Eastern Australia. - Biotropica 33: 643-661. Go to original source...
    44. Mammola S., Giachino P.M., Piano E., Jones A., Barberis M., Badino G. & Isaia M. 2016: Ecology and sampling techniques of an understudied subterranean habitat: The Milieu Souterrain Superficiel (MSS). - Sci Nat. 103: 88, 24 pp. Go to original source...
    45. Martins M.F. de O., Thomazini M.J., Baretta D., Brown G.G., Rosa M.G. da, Zagatto M.R.G., Santos A., Nadolny H.S., Cardoso G.B.X., Niva C.C., Bartz M.L.C. & Feitosa R.M. 2020: Accessing the subterranean ant fauna (Hymenoptera: Formicidae) in native and modified subtropical landscapes in the Neotropics. - Biota Neotrop. 20: e20190782, 16 pp. Go to original source...
    46. Nitzu E., Nae A., Giurginca A. & Popa I. 2010: Invertebrate communities from the mesovoid shallow substratum of the carpatho-euxinic area: Eco-faunistic and zoogeographic analysis. - Trav. Inst. Spéol. 49: 41-79.
    47. Nitzu E., Nae A., Bãncilã R., Popa I., Giurginca A. & Plãiaºu R. 2014: Scree habitats: Ecological function, species conservation and spatial-temporal variation in the arthropod community. - Syst. Biodiv. 12: 65-75. Go to original source...
    48. Nitzu E., Dorobãþ M.L., Popa I., Giurginca A. & Baba ª. 2018: The influence of geological substrate on the faunal structure of the superficial subterranean habitats. - Carpath. J. Earth Environ. Sci. 13: 383-393. Go to original source...
    49. Normand H. 1911: Description d'un nouveau procedé de capture des Coléopteres hypogés. - L'Echange 315: 114-116, 124-126.
    50. Oksanen J., Simpson G., Blanchet F., Kindt R., Legendre P., Minchin P., O'Hara R., Solymos P., Stevens M., Szoecs E. et al. 2024: _vegan: Community Ecology Package_. R Package Version 2.6-6.1. URL: http://www.CRAN.R-project.org/package=vegan
    51. Ortuño V.M., Gilgado J.D., Jiménez-Valverde A., Sendra A., Pérez-Suárez G. & Herrero-Borgoñón J.J. 2013: The "Alluvial Mesovoid Shallow Substratum", a new subterranean habitat. - PLoS ONE 8: e76311, 16 pp. Go to original source...
    52. Pipan T., López H., Oromí P., Polak S. & Culver D.C. 2010: Temperature variation and the presence of troglobionts in terrestrial shallow subterranean habitats. - J. Nat. Hist. 45: 253-273. Go to original source...
    53. R Core Team 2024: R: A Language and Environment for Statistical Computing (Version 4.4.0) [R]. R Foundation for Statistical Computing, URL: http://www.R-project.org
    54. Roeder K.A., Roeder D.V. & Bujan J. 2021: Ant thermal tolerance: A review of methods, hypotheses, and sources of variation. - Ann. Entomol. Soc. Am. 114: 459-469. Go to original source...
    55. Rohde K. 1992: Latitudinal gradients in species diversity: The search for the primary cause. - Oikos 65: 514-527. Go to original source...
    56. Ryder Wilkie K.T., Mertl A.L. & Traniello J.F.A. 2007: Biodiversity below ground: Probing the subterranean ant fauna of Amazonia. - Naturwissenschaften 94: 725-731. Go to original source...
    57. Samih A., Petit D., Maatouf N., Trócoli S., Habbaz H. & Rohi L. 2024: Diversity of beetle communities in cork oak forest of Larache frome Morocco. - Indian J. Entomol. 87: e2590, 12 pp. Go to original source...
    58. Sanders N.J., Lessard J.-P., Fitzpatrick M.C. & Dunn R.R. 2007: Temperature, but not productivity or geometry, predicts elevational diversity gradients in ants across spatial grains. - Global Ecol. Biogeogr. 16: 640-649. Go to original source...
    59. Sauvage C. 1961: Flore des suberaies marocaines: Catalogue des cryptogames vasculaires et des phanerogames. Travaux de l'Institut scientifique cherifien, Série Botanique No. 22, Rabat, 252 pp.
    60. Schmidt F.A. & Solar R.R.C. 2010: Hypogaeic pitfall traps: Methodological advances and remarks to improve the sampling of a hidden ant fauna. - Insectes Soc. 57: 261-266. Go to original source...
    61. Sharaf M.R., Aldawood A.S. & Taylor B. 2011: The formicine ant genus Plagiolepis Mayr (Hymenoptera: Formicidae) in the Arabian peninsula, with description of two new species. -Trans. Am. Entomol. Soc. 137: 203-215. Go to original source...
    62. Silva R.R.D. & Silvestre R. 2004: Riqueza da fauna de formigas (Hymenoptera: Formicidae) que habita as camadas superficiais do solo em Seara, Santa Catarina. - Papéis Avulsos Zool. 44: 1-11. Go to original source...
    63. Szewczyk T. & McCain C.M. 2016: A systematic review of global drivers of ant elevational diversity. - PLoS ONE 11: e0155404, 18 pp. Go to original source...
    64. Taheri A., Reyes-Lopez J.L. & Bennas N. 2014: Contribution à l'étude de la faune myrmécologique du Parc National de Talassemtane (Nord du Maroc): Biodiversité, biogéographie et espèces indicatrices. - Bol. Soc. Entomol. Aragon. (S.E.A.) 54: 225-236.
    65. Taylor B. 2015: The Ants of (sub-Saharan) Africa (Hymeno­ptera: Formicidae) Profusely Illustrated with Original Drawings and Photographs and Catalogue Notes on all Ant Species Described from sub-Saharan Africa. 12th ed. URL: https://www.antsofafrica.org/
    66. Underwood E.C. & Fisher B.L. 2006: The role of ants in conservation monitoring: If, when, and how. - Biol. Conserv. 132: 166-182. Go to original source...
    67. Verdinelli M., Yakhlef S., Cossu C., Pilia O. & Mannu R. 2017: Variability of ant community composition in cork oak woodlands across the Mediterranean region: Implications for forest management. - iForest - Biogeosci. Forestry 10: 707-714. Go to original source...
    68. Villemant C. & Fraval A. 1993: La faune entomologique du chêne-liège en forêt de la Mamora (Maroc). - Ecol. Mediterr. 19: 89-98. Go to original source...
    69. Weather Spark 2022: Larache Climate, Weather by Month, Average Temperature (Morocco). URL: https://weatherspark.com/y/33173/Average-Weather-in-Larache-Morocco-Year-Round
    70. Weiser M.D. & Kaspari M. 2006: Ecological morphospace of New World ants. - Ecol. Entomol. 31: 131-142. Go to original source...
    71. Wickham H. 2016: ggplot2: Elegant Graphics for Data Analysis. Springer, New York, URL: http://www.ggplot2.tidyverse.org Go to original source...
    72. Wong M.K.L. & Guénard B. 2017: Subterranean ants: Summary and perspectives on field sampling methods, with notes on diversity and ecology (Hymenoptera: Formicidae). - Myrmecol. News 25: 1-16.
    73. Zouaki N., Ouattar H., Slim M. & Fadli M. 2018: First evaluation of the structure and the entomological biodiversity of the Maâmora forest and the Merja of Fouarat (Kenitra, Morocco). - J. Entomol. Zool. Stud. 6: 32-37.
    74. Zryanin V.A. 2015: The first subterranean ant species of the genus Meranoplus F. Smith, 1853 (Hymenoptera: Formicidae) from Vietnam. - Caucas. Entomol. Bull. 11: 153-160. Go to original source...

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