Distribution and Ecology of Myrmecophilous Crickets in the Province of Granada (Spain)

Matvey Logachev; Nerea Megías-Valero; Mathias Welz-Sol; Paco Alarcón; Manuel Martín-Vivaldi

doi:10.13102/sociobiology.v73i3.12771

Authors

Matvey Logachev Universidad de Granada, Departamento de Zoología, Campus de Fuentenueva s/n, E-18003 Granada, Spain. https://orcid.org/0000-0001-9484-5108
Nerea Megías-Valero Universidad de Granada, Departamento de Zoología, Campus de Fuentenueva s/n, E-18003 Granada, Spain
Mathias Welz-Sol Universidad de Granada, Departamento de Zoología, Campus de Fuentenueva s/n, E-18003 Granada, Spain
Paco Alarcón Independent researcher, Sevilla, Spain https://orcid.org/0009-0003-9221-3950
Manuel Martín-Vivaldi Universidad de Granada, Departamento de Zoología - E-18003 & Unidad Asociada (CSIC): Coevolución - E-18071, Campus de Fuentenueva s/n, Granada, Spain https://orcid.org/0000-0002-5432-425X

DOI:

https://doi.org/10.13102/sociobiology.v73i3.12771

Keywords:

Ants, Bioclimatic zones, Granada, Myrmecophilus ochraceus, Spain, transition Europe-northern Africa

Abstract

Myrmecophily, broadly, refers to associations between certain organisms and ants. Among arthropods, one of the most specialized groups to live within ant nests is crickets of the genus Myrmecophilus Berthold, 1827. Worldwide distributed, the genus includes numerous species adapted to different hosts whose distribution and ecology have been scarcely studied in many regions. In the Iberian Peninsula, four species have been recorded, mainly from scattered locality records, while programmed samplings across ecosystems are lacking. In this context, this study documents, for the first time, the distribution of crickets of the genus Myrmecophilus in the province of Granada, Southeastern Spain, and describes their ecological preferences. Programmed sampling was carried out across different bioclimatic zones and habitat types, and a total of 1,156 ant nests of 19 genera were inspected, resulting in the recording of 78 cricket specimens, all assigned to the species Myrmecophilus ochraceus Fischer, 1853. The cricket was restricted to the thermo- and meso-Mediterranean zones and showed a marked preference for open grasslands. A high number of workers below stones and the presence of ant brood increased the likelihood of detecting the orthopteran. Considering all our records in Granada (more than 2000 ant nests inspected and 263 cricket specimens), only three out of 19 ant genera hosted crickets, including four new host ant species for M. ochraceus. Our results, together with previous studies, suggest that the only Myrmecophylus species inhabiting southern Iberia is M. ochraceus, which specializes on Messor, Monomorium, and Tetramorium species as main hosts. The possible implications of a geographical segregation of Myrmecophilus species between northern and southern Iberia are discussed.

Downloads

Download data is not yet available.

References

Aguirre-Segura, A., Barranco, P., & Pascual, F. (1995). La colección de ortópteros de la Estación Experimental de Zonas Áridas (C.S.I.C.) de Almería. (Insecta, Orthoptera). Boletín de la Asociación Española de Entomología, 19(1–2): 133–155.

Akino, T., Mochizuki, R., Morimoto, M., & Yamaoka, R. (1996). Chemical camouflage of Myrmecophilous cricket Myrmecophilus sp. To be integrated with several ant species. Japanese Journal of Applied Entomology and Zoology, 40(1): 39–46.

Alexiou, S., Gavalas, I., & Papapavlou, K. (2017). Orthoptera (Saltatoria) of Iraklia island, Cyclades, Greece: An annotated and illustrated catalogue. Israel Journal of Entomology, 47: 35–53.

Arcos, J., & García, F. (2023). Hormigas de la península Ibérica e islas Baleares. Independently published, 491 p.

Arcos, M. A., Edo-Botella, R., Balaguer-Beser, Á., & Ruiz, L. Á. (2023). Analyzing independent LFMC empirical models in the mid-Mediterranean region of Spain attending to vegetation types and bioclimatic zones. Forests, 14(7): Article 7.

Azevedo, L., Emberson, R., Esteca, F., & Moraes, G. (2015). Macrochelid mites (Mesostigmata: Macrochelidae) as biological control agents. Prospects for Biological Control of Plant Feeding Mites and Other Harmful Organisms, 19: 103–132.

Bergamaschi, D., Moore, W., & Giulio, A.D. (2025). Co-phylogeny and biogeography of the myrmecophilous beetle Paussus favieri (Carabidae, Paussinae) and its host ant Pheidole pallidula (Hymenoptera, Myrmicinae). Insect Science 0: 1–17.

Blender Development Team (2026) Blender (Version 5.2.0 LTS) [Computer software]. Blender Foundation. Available from: https://www.blender.org/download/releases/5-2/ (accessed 30 July 2026).

Bolton, B. (1987). A review of the Solenopsis genus-group and revision of Afrotropical Monomorium Mayr (Hymenoptera: Formicidae). Bulletin of the British Museum (Natural History). Entomology, 54: 263–452.

Cassar, L., Bartolo, A., Stalling, T., Schembri, S., & Massa, B. (2025). Myrmecophilus baronii Baccetti, 1966: Biogeography and genetics (Orthoptera: Myrmecophilidae). Fragmenta Entomologica, 57: 9–20.

Cassar, T., Lapeva-Gjonova, A., & Mifsud, D. (2023). The intranidal myrmecophiles of the Maltese Islands with notes on Messor nests as repositories of biodiversity. Insects, 14: 45.

Cigliano, M. M., Braun, H., Eades, D. C., & Otte, D. (2025). Orthoptera Species File. Version 5.0/5.0. (Accessed: 18 March 2026). http://Orthoptera.SpeciesFile.org (Accessed: 1 December 2025)

Collingwood, C. A. (1979). The Formicidae (Hymenoptera) of Fennoscandia and Denmark (Vol. 8). Scandinavian Science Press, 174 p.

Consejería de Medio Ambiente. (2015). 3ª Adecuación del Plan Forestal Andaluz, Horizonte 2015 [Acuerdo de 7 de septiembre de 2010, del Consejo de Gobierno]. Junta de Andalucía.

De Almeida, T., Blight, O., Mesléard, F., Bulot, A., Provost, E., & Thierry, D. (2020). Harvester ants as ecological engineers for Mediterranean grassland restoration: Impacts on soil and vegetation. Biological Conservation, 245: 1–8.

Dean, W. R. J., & Turner, J. S. (1991). Ants nesting under stones in the semi-arid Karoo, South Africa: Predator avoidance or temperature benefits? Journal of Arid Environments, 21(1): 59–69.

Dekoninck, W., Lock, K., & Janssens, F. (2007). Acceptance of two native myrmecophilous species, Platyarthrus hoffmannseggii (Isopoda: Oniscidea) and Cyphoderus albinus (Collembola: Cyphoderidae) by the introduced invasive garden ant Lasius neglectus (Hymenoptera: Formicidae) in Belgium. European Journal of Entomology, 104: 159–161.

Del Río Sánchez, J., Peñas, J., Lorite, J., & Muñoz, J. M. (2019). Itinerario botánico por Sierra Nevada. 9º Congreso de Biología de la Conservación de Plantas, 50 p.

Elmes, G. W. (1996). Biological diversity of ants and their role in ecosystem function. Biodiversity Research and Its Perspectives in East Asia, Chonbuk National University, Jeonju-Si, Korea, 33–48.

Espadaler, X., & Olmo-Vidal, J. (2011). The myrmecophilic cricket Myrmecophilus in Spain (Orthoptera, Myrmecophilidae). Sociobiology, 57: 321-328.

Fischer, L. H. (1853). Orthoptera Europaea: Accedunt tabulae lapidibus incisae XVIII quarum ultima coloribus partim illustrata. Wilhelm Engelmann, 500 p.

Franc, V., Majzlan, O., Krištín, A., & Wiezik, M. (2015). On the distribution and ecology of the ant cricket (Myrmecophilus acervorum) (Orthoptera: Myrmecophilidae) in Slovakia. Matthias Belivs University Proceedings, 5: 40–50.

Geiselhardt, S., Peschke, K., & Nagel, P. (2007). A review of myrmecophily in ant nest beetles (Coleoptera: Carabidae: Paussinae): Linking early observations with recent findings. Die Naturwissenschaften, 94: 871–894.

Henderson, G., & Akre, R. D. (1986). Biology of the myrmecophilous cricket, Myrmecophila manni (Orthoptera: Gryllidae). Journal of the Kansas Entomological Society, 59: 454–467.

Hsu, P.-W., Hugel, S., Wetterer, J. K., Tseng, S.-P., Ooi, C.-S. M., Lee, C.-Y., & Yang, C.-C. S. (2020). Ant crickets (Orthoptera: Myrmecophilidae) associated with the invasive yellow crazy ant Anoplolepis gracilipes (Hymenoptera: Formicidae): Evidence for cryptic species and potential co-introduction with hosts. Myrmecological News, 30: 103–129.

Hölldobler, B., & Kwapich, C. L. (2022). The Guests of Ants: How Myrmecophiles Interact with Their Hosts. Cambridge: Harvard University Press, 576 p.

Iorgu, I. Ștefan, Iorgu, E., Stalling, T., Puskás, G., Chobanov, D., Szövényi, G., Moscaliuc, L., Motoc, R., Tăușan, I., & Fusu, L. (2021). Ant crickets and their secrets: Myrmecophilus acervorum is not always parthenogenetic (Insecta: Orthoptera: Myrmecophilidae). Zoological Journal of the Linnean Society, 20: 1–18.

Jiménez, J., & Tinaut, A. (1992). Mirmecofauna de la Sierra de Loja (Granada) (Hymenoptera, Formicidae). Orsis: Organismes i Sistemes, 7: 97–111.

Junker, E. (1997). Untersuchungen zur Lebensweise und Entwicklung von Myrmecophilus acervorum (Panzer, 1799) (Saltatoria: Myrmecophilidae). Articulata, 12(2): 93–106.

Juvé, Y., Weyna, A., Lauroua, E., Nidelet, S., Khaldi, M., Barech, G., Lebas, C., Rasplus, J.Y., Cruaud, A., Condamine, F.L. & Romiguier, J. (2025). Phylogenomics of Messor harvester ants (Hymenoptera: Formicidae: Stenammini) unravels their biogeographical origin and diversification patterns. Systematic Entomology, 50(4): 1025-1040.

Kaczmarczyk-Ziemba, A., Wagner, G., Staniec, B., Zagaja, M., Pietrykowska-Tudruj, E., Iorgu, E., & Iorgu, I. Ștefan. (2024). Intraspecific diversity of Myrmecophilus acervorum (Orthoptera: Myrmecophilidae) indicating an ongoing cryptic speciation. Scientific Reports, 14: 1–18.

Komatsu, T., Maruyama, M., & Itino, T. (2010). Differences in host specificity and behavior of two ant cricket species (Orthoptera: Myrmecophilidae) in Honshu, Japan. Journal of Entomological Science, 45(3): 227–238.

Komatsu, T., Munetoshi, M., Hattori, M., & Itino, T. (2017). Morphological characteristics reflect food sources and degree of host ant specificity in four Myrmecophilus crickets. Insectes Sociaux, 65: 47–57.

Komatsu, T., Munetoshi, M., & Itino, T. (2013). Nonintegrated host association of Myrmecophilus tetramorii, a specialist myrmecophilous ant cricket (Orthoptera: Myrmecophilidae). Psyche: A Journal of Entomology, 2013: 1-5.

Kovář, P., Vojtı́šek, P., & Zentsova, I. (2013). Ants as ecosystem engineers in natural restoration of human made habitats. Journal of Landscape Ecology, 6: 18-31.

Lapeva-Gjonova, A., Csősz, S., & Báthori, F. (2022). Fungi associated with Messor ants on the Balkan Peninsula: first biogeographical data. Diversity, 14: 1–8.

Logachev, M. (2024). Ant-associated crickets of the genus Myrmecophilus Berthold, 1827 in the Iberian Peninsula and Balearic Islands. Iberomyrmex, 13: 30–42.

Loidi, J. (2017). The Vegetation of the Iberian Peninsula. Cham: Springer Nature, 676.

Maggenti, M. A. B., Maggenti, A. R., & Gardner, S. L. (2017). Dictionary of Invertebrate Zoology. Zea E-Books Collection, 61 p.

Mérel, V., Vogel, S., Lavanchy, G., Dumas, Z., Labédan, M., Stalling, T., & Schwander, T. (2025). Recent neo-X and Y sex chromosomes in an ant cricket. Journal of Evolutionary Biology, 38: 1000–1007.

Moyano Ayala, L. (2014). Estudio y seguimiento de la fauna de Orthoptera de un entorno natural sometido a un programa de restauración ecológica en el sur de la Península Ibérica (Doctoral dissertation). University of Cordoba, 284 p.

Offenberg, J. (2001). Balancing between mutualism and exploitation: The symbiotic interaction between Lasius ants and aphids. Behavioral Ecology and Sociobiology, 49: 304–310.

Ortiz, F. J., & Tinaut, J. A. (1988). Formícidos del litoral granadino. Orsis: Organismes i Sistemes, 3: 145–163.

Pardo, G., Del Prado, A., Martínez-Mena, M., Bustamante, M., Rodriguez Martin, J. A., Álvaro-Fuentes, J., & Moral, R. (2016). Orchard and horticulture systems in Spanish Mediterranean coastal areas: Is there a real possibility to contribute to C sequestration? Agriculture, Ecosystems & Environment, 238(1): 153–167.

Pierce, N., Braby, M., Heath, A., Lohman, D., Mathew, J., Rand, D., & Travassos, M. (2002). The ecology and evolution of ant association in the Lycaenidae (Lepidoptera). Annual Review of Entomology, 47: 733–771.

Posit team. (2026). RStudio: Integrated Development Environment for R (Version 4.5.2) [Computer software]. Posit Software, PBC. http://www.posit.co/

QGIS Development Team. (2026). QGIS Geographic Information System (Version 4.2) [Computer software]. QGIS Association. http://qgis.org

Reznikova, Z. (2001). Interaction of the ant Myrmica rubra L. as a predator with springtails (Collembola) as a mass prey. Doklady Biological Sciences: Proceedings of the Academy of Sciences of the USSR, Biological Sciences Sections / Translated from Russian, 380: 475–477.

Rodríguez-Castilla, G. & Stalling, T. (2025). The Mediterranean ant cricket Myrmecophilus myrmecophilus (Savi, 1819), new species for Portugal (Orthoptera: Myrmecophilidae). Heteropterus Revista de Entomología, 25(2): 303–306.

Salata, S., Kalarus, K., Borowiec, L., Trichas, A., Kujawa, K. (2020). How estimated ant diversity is biased by the sampling method? A case study of Crete: a Mediterranean biodiversity hotspot. Biodiversity and Conservation, 29: 3031–3050.

Sandoval Cortés, P. J., Aguayo Becerra, D., Tinaut, A., & Ruano Díaz, F. (2024). Dataset of the arthropodofauna of Sierra Nevada from the zoological collections of the University of Granada. Version 1.8. Dept. Of Zoology, Faculty of Science, University of Granada. Occurrence dataset. [Dataset].

Sanno, R., Kataoka, K., Hayakawa, S., Ide, K., Chuong, N., Nguyen, T., Le, B., Kim, O., Mineta, K., Takeyama, H., Takeda, M., Sato, T., Suzuki, T., Yura, K., & Asahi, T. (2021). Comparative analysis of mitochondrial genomes in Gryllidea (Insecta: Orthoptera): Implications for adaptive evolution in ant-loving crickets. Genome Biology and Evolution, 13(10): 1–8.

Schembri, S. P. (1984). Notes on the Orthoptera of the Maltese Islands: The genus Myrmecophilus (Orthoptera : Gryllidae). The Central Mediterranean Naturalist, 1(3): 57–59.

Schifani, E., Nalini, E., Gentile, V., Alamanni, F. M., Ancona, C., Caria, M., Cillo, D., & Bazzato, E. (2021). Ants of Sardinia: An updated checklist based on new faunistic, morphological and biogeographical notes. Journal of Zoology, 104: 21–35.

Schöning, C., Espadaler, X., Hensen, I., & Roces, F. (2004). Seed predation of the tussock-grass Stipa tenacissima L. by ants (Messor spp.) in southeastern Spain: The adaptive value of trypanocarpy. Journal of Arid Environments, 56: 43–61.

Stalling, T. (2010). First record of the ant-loving cricket Myrmecophilus ochraceus for Zakynthos (Greece) with notes on the synonymy of Myrmecophilus salomonis (Orthoptera: Myrmecophilidae). Entomologia Hellenica, 19(1): Article 1.

Stalling, T. (2013a). A new species of ant-loving cricket from Mallorca, Balearic Islands, Spain (Orthoptera, Myrmecophilidae). Graellsia, 69(2): 153–156.

Stalling, T. (2013b). Myrmecophilus balcanicus, a new species of ant-loving cricket from the Former Yugoslav Republic of Macedonia, with notes on the synonymy of Myrmecophilus zorae. Entomologia Hellenica, 22: 29–34.

Stalling, T. (2015). New records of the ant-loving cricket Myrmecophilus fuscus Stalling, 2013 from the Mediterranean Basin (Orthoptera: Myrmecophilidae). Articulata, 30: 105–108.

Stalling, T. (2016). The presence of the ant-loving cricket Myrmecophilus ochraceus in Sardinia, Italy (Orthoptera: Myrmecophilidae). Fragmenta Entomologica, 48: 17.

Stalling, T. (2017). Myrmecophilus gallicus, a new species of ant cricket from France (Orthoptera: Myrmecophilidae). Articulata, 32: 1–7.

Stalling, T. (2024). Myrmecophilus jordanicus, a new species of ant cricket from Jordan and Israel, with notes on the synonymy of Myrmecophilus nigricornis (Orthoptera: Myrmecophilidae). Israel Journal of Entomology, 53: 25–32.

Stalling, T., & Birrer, S. (2013). Identification of the ant-loving crickets, Myrmecophilus Berthold, 1827 (Orthoptera: Myrmecophilidae), in Central Europe and the northern Mediterranean Basin. Articulata, 28: 1–11.

Stalling, T., Espadaler, X., & Carron, N. (2015). First record of the ant-loving cricket Myrmecophilus fuscus Stalling, 2013 (Orthoptera: Myrmecophilidae) in mainland Spain. Sociobiology 62(4): 481-483.

Stalling, T., Iorgu, I. Ștefan, & Chobanov, D. (2020). The ant cricket Myrmecophilus orientalis on the Dodecanese Islands, Greece (Orthoptera: Myrmecophilidae). Travaux Du Muséum National d’Histoire Naturelle “Grigore Antipa"Antipa”, 63: 63–67.

Stalling, T., Sjödahl, M., & Ulrik, P. (2017). Records of the ant cricket Myrmecophilus acervorum from the northern border of the distribution range (Orthoptera: Myrmecophilidae). Entomologisk Tidskrift, 138: 97–101.

Steiner, F., Csősz, S., Markó, B., Gamisch, A., Rinnhofer, L., Folterbauer, C., Hammerle, S., Stauffer, C., Arthofer, W., & Schlick-Steiner, B. (2018). Turning one into five: Integrative taxonomy uncovers complex evolution of cryptic species in the harvester ant Messor “structor”. Molecular Phylogenetics and Evolution, 127: 387–404.

Temreshev, I. (2020). First record of the ant-loving cricket Myrmecophilus crenatus Gorochov, 1986 (Orthoptera, Myrmecophilidae) in Kazakhstan. Acta Biologica Sibirica, 6: 407–412.

Tinaut, A., Jiménez Rojas, J., & Pascual, R. (1994). Estudio de la mirmecofauna de los bosques de Quercus Linneo, 1753 de la provincia de Granada (Hymenoptera; Formicidae). Ecología, 8: 429–438.

Tinaut, A., Martínez-Ibáñez M. D. & Ruano, F. (2007). Inventario de las especies de formícidos de Sierra Nevada, Granada (España) (Hymenoptera, Formicidae). Zool. Baetica, 18: 49-68.

Tinaut, A., Ruano, F. (2021). Biogeography of Iberian ants (Hymenoptera: Formicidae). Diversity, 13(2): 88.

Valle, F. (1985). Mapa de las series de vegetación de Sierra Nevada (España).

Wetterer, J., & Hugel, S. (2008). Worldwide spread of the ant cricket Myrmecophilus americanus, a symbiont of the longhorn crazy ant, Paratrechina longicornis. Sociobiology, 52: 157–165.