Cardiocondyla elegans

There are two queen castes in this species. Pashaei Rad et al. (2018) found this species in Iran on parkland soil in a Caspian moist littoral area.

Identification
Seifert (2003) - A member of the Cardiocondyla elegans group. Long scape, long PLG, moderately large eyes, and small postocular distance.

Seifert (2023) - Relatively large, CS 560 µm. Head moderately elongated, CL/CW 1.159. Postocular distance low, PoOc/CL 0.394. Scape rather long, SL/CS 0.842. Eye rather large, EYE/CS 0.252, with notable microsetae. Occipital margin suggestively concave to straight. Frons rather broad (FRS/CS 0.264), frontal carinae slightly converging immediately caudal of FRS level (FL/FR 1.051). Dorsal profile of promesonotum and of propodeum convex with a deep metanotal depression (Mgr/CS 4.82 %). Spines rather short and acute (SP/CS 0.119), their axis in profile deviating by about 35° from longitudinal axis of mesosoma, their bases rather distant (SPBA/CS 0.267). Petiole rather wide and slightly lower than wide (PeW/CS 0.348, PeH/CS 0.330); in profile with a moderately long peduncle and a steep anterior slope of the node (about 65° relative to ventral profile). Postpetiole wide and moderately high (PpW/CS 0.580, PpH/CS 0.300), in dorsal view suggestively heard-shaped, with a concave anterior margin and convex sides; postpetiolar sternite convex. Head in overall impression mildly shiny. Whole vertex with shallow, feebly bicoronate foveolae of 16–18 µm diameter, foveolar distance on paramedian vertex smaller than foveolar diameter, near to eyes larger; the interspaces between foveolae shiny and in places with fragments of a very delicate microreticulum (Fig. 9). Mesosoma shiny, with weakly developed microreticulum and microrugulae, a large number of foveolae present on dorsal promesonotum, their distance approximately equal to their diameter. Dorsal propodeum glabrous but with small foveolae and a very delicate microrugosity. Dorsum of waist glabrous, with scattered fragments of very fine microreticular structures. First gaster tergite glabrous. Pubescence on whole body long and dense, PLg/CS 7.92 %, sqPDg 3.95. Color of head, mesosoma, waist and gaster usually homogenously dark to medium brown; mandibles, scape, tibiae and tarsae yellowish brown.

The 85 available nest samples of the sister species Cardiocondyla elegans and Cardiocondyla dalmatica can be separated with a mean error of 1.2 % by the exploratory data analyses NC-Ward, NC-part.hclust, NC-part.kmeans and NC-NMDS.kmeans using the characters CS, CL/CW, PoOc/CL, EYE/CS, SL/CS, SP/CS, PeW/CS, PpW/CS, PeH/CS, PpH/CS, sqPDg, PLg/CS, Mgr/CS and dFOv (Fig. 133). A LDA using the same characters classified 97.8 % of 185 worker individuals of both species correctly. In the sample mean and when run in the LDA as wild-cards, the type samples of C. elegans, C. santschii, C. provincialis and C. gallica are allocated with posterior probabilities of 0.8443, 0.9166, 0.9972 and 0.9924 to the elegans cluster, whereas the lectotype of C. dalmatica is allocated with p=0.9961 to the dalmatica cluster.

Distribution
Cardiocondyla elegans is a western Mediterranean species, found from Portugal, southern France and Italy north to 47.5°N. In northern Italy east to 13.1°E and in southern Spain ascending to 930 m. Records of this species from the Balkans refer to Cardiocondyla dalmatica (Seifert, 2018, 2023; Lapeva-Gjonova & Antonova, 2022).

Distribution based on Regional Taxon Lists
Palaearctic Region: France, Iberian Peninsula, Italy, Portugal, Spain.

Biology
Seifert (2003) - Cardiocondyla elegans shows a discrete gyne bimorphism in dimensions and shape of mesosoma and wing length while other measurements and characters are monomorphic. A cluster of gynes with MW 418 ± 16 [392, 442] (n = 10) can be separated from one with MW 485 ± 17 [465, 531] (n = 1 2) within 12 gyne samples investigated. The full similarity of the associated workers and the occurrence of both gyne types within the same nest (Barcelona: Bellaterra) indicate an intraspecific polymorphism. The microsomatic gynes with (ML*MW)/(CL*CW) 0.903 ± 0.024 [0.870, 0.940] are supposed to have weaker flight muscles and short wings. They should be mated intranidally by ergatoid males and should either stay within the mother colony, propagate by colony fission, or seek adoption in conspecific neighboured colonies. The macro somatic gynes with (ML*MW)/(CL*CW) 1.129 ± 0.046 [1.086, 1.254], strong flight muscles, and normal wing length are suspected to disperse in the air and to found a new colony independently after having been inseminated by an ergatoid male within the mother nest.

Colonies contain less then 500 workers and are monogynous.

Nomenclature

 * . Cardiocondyla elegans Emery, 1869b: 21, pl. 1, fig. 10 (w.q.) ITALY.
 * Type-material: lectotype worker (by designation of Seifert, 2003a: 225), 5 paralectotype workers.
 * Type-locality: lectotype Italy: Naples, Capodimonte, 18.vi.1866 (C. Emery); paralectotypes with same data but lacking date.
 * [Note: original description also mentions syntype workers: Italy: Naples, Portici, and a syntype putative queen: Italy: Lucca (Haliday).]
 * Type-depositories: MSNG (lectotype); MHNG (paralectotypes).
 * Menozzi, 1918: 83 (ergatoid m.); Kugler, J. 1984: 14 (ergatoid m.).
 * Status as species: Mayr, 1877: 18; Emery, 1878b: 50; Emery & Forel, 1879: 456; Mayr, 1880: 37; Forel, 1881: 7; André, 1881b: 69; André, 1883a: 328 (in key); Emery, 1889a: 441; Mayr, 1889: 280; Nasonov, 1889: 38; Dalla Torre, 1893: 70; Ruzsky, 1902d: 25; Ruzsky, 1905b: 624; Emery, 1909a: 21; Karavaiev, 1910b: 56; Emery, 1916b: 153; Menozzi, 1918: 83; Bondroit, 1918: 147; Emery, 1922e: 125; Karavaiev, 1934: 114; Grandi, 1935: 100; Finzi, 1936: 166; Finzi, 1939b: 88; Donisthorpe, 1950e: 1060; Consani & Zangheri, 1952: 40; Bernard, 1956c: 301 (in key); Ceballos, 1956: 308; Bernard, 1967: 157 (redescription); Pisarski, 1967: 388; Collingwood & Yarrow, 1969: 75; Baroni Urbani, 1971c: 68; Baroni Urbani, 1971c: 76; Tarbinsky, 1976: 73; Hamann & Klemm, 1976: 673; Aktaç, 1977: 121; Arnol'di & Dlussky, 1978: 538 (in key); Collingwood, 1978: 86 (in key); Kugler, J. 1984: 14; Agosti & Collingwood, 1987a: 56; Agosti & Collingwood, 1987b: 276 (in key); Kugler, J. 1988: 258; Dlussky, Soyunov & Zabelin, 1990: 194; Atanassov & Dlussky, 1992: 172; Collingwood, 1993b: 194; Arakelian, 1994: 43; Bolton, 1995b: 132; Poldi, et al. 1995: 4; Radchenko, 1995b: 449; Espadaler, 1997b: 29; Collingwood & Prince, 1998: 16 (in key); Collingwood & Heatwole, 2000: 7; Rigato, 2002: 173 (in key); Seifert, 2003a: 225 (redescription); Petrov, 2006: 99 (in key); Bračko, 2007: 17; Gratiashvili & Barjadze, 2008: 138; Paknia, et al. 2008: 153; Casevitz-Weulersse & Galkowski, 2009: 487; Vonshak, et al. 2009: 41; Lapeva-Gjonova, et al. 2010: 26; Csösz, et al. 2011: 57; Karaman, M.G. 2011b: 21; Legakis, 2011: 16; Borowiec, L. & Salata, 2012: 484; Guénard & Dunn, 2012: 40; Kiran & Karaman, 2012: 17; Borowiec, L. 2014: 46; Bračko, et al. 2014: 13; Tohmé, G. & Tohmé, 2014: 135; Lebas, et al. 2016: 266; Salata & Borowiec, 2018c: 44; Seifert, 2018: 186.
 * Senior synonym of gallica: Baroni Urbani, 1973a: 200; Kugler, J. 1984: 14; Bolton, 1995b: 132; Radchenko, 1995b: 449; Seifert, 2003a: 225; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * Senior synonym of provincialis: Radchenko, 1995b: 449; Seifert, 2003a: 225; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * Senior synonym of santschii: Bondroit, 1918: 147; Emery, 1922e: 125; Bolton, 1995b: 132; Radchenko, 1995b: 449; Seifert, 2003a: 225; Casevitz-Weulersse & Galkowsky, 2009: 487; Legakis, 2011: 16.
 * Distribution: France, Italy, Portugal, Spain.
 * [Note: distribution from Borowiec, L. 2014: 46, modified after Seifert, 2018:186-187.]
 * gallica. Xenometra gallica Bernard, 1957b: 101, fig. 1 (q.) FRANCE.
 * Type-material: 3 syntype males.
 * [Note: the three syntypes are ergatoid males, not queens (Baroni Urbani, 1973a: 200).]
 * Type-locality: France: edge of Dordogne, Lot, Pinsac, ix.1951 (G. Lepointe).
 * Type-depository: MNHN.
 * Status as species: Bernard, 1967: 158 (redescription); Kutter, 1968b: 203.
 * Junior synonym of elegans: Baroni Urbani, 1973a: 200; Kugler, J. 1984: 14; Bolton, 1995b: 132; Radchenko, 1995b: 449; Seifert, 2003a: 225; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * provincialis. Cardiocondyla provincialis Bernard, 1956c: 303, fig. 6 (w.) FRANCE.
 * Type-material: 2 syntype workers.
 * Type-locality: France: Var, Frejus, ix.1934 (F. Bernard).
 * Type-depository: MNHN.
 * Status as species: Bernard, 1967: 158 (redescription); Bolton, 1995b: 133.
 * Junior synonym of elegans: Radchenko, 1995b: 449; Seifert, 2003a: 225; Casevitz-Weulersse & Galkowsky, 2009: 487.
 * santschii. Cardiocondyla elegans r. santschii Forel, 1905b: 174 (w.) FRANCE.
 * Type-material: syntype workers (number not stated).
 * Type-locality: France: Marseille, viii.1904 (F. Santschi).
 * Type-depository: MHNG.
 * Forel, 1910a: 23 (q.).
 * Subspecies of elegans: Forel, 1910a: 23; Forel, 1911f: 276; Forel, 1911h: 457; Santschi, 1926f: 293.
 * Junior synonym of elegans: Bondroit, 1918: 147; Emery, 1922e: 125; Bolton, 1995b: 133; Radchenko, 1995b: 449; Seifert, 2003a: 225; Casevitz-Weulersse & Galkowsky, 2009: 487; Legakis, 2011: 16.

Type Material
Cardiocondyla elegans (examined by Seifert, 2023) Cardiocondyla elegans santschii (examined by Seifert, 2003, 2023) Cardiocondyla provincialis (examined by Seifert, 2023) Xenometra gallica (examined by Seifert, 2023)
 * The species has been described from Napoli / Italy. Investigated were one lectotype worker from MCSN genova and five paralectotype worker from MHN genève (a gift of Emery to Forel). The lectotype worker is labelled “Naples♀”, Bosco di Capodimonte 18. VI 1866”, “Cardiocondyla elegans Em. Napoli”, “TYPUS”, “Lectotype Cardiocondyla elegans Emery 1869, designated by B.Seifert 1998” and “ANTWEB CASENT 0904460”.
 * Lectotype worker designated by Seifert (2003). This taxon has been described from Marseille / France. Investigated were 5 worker syntypes labelled: “C. elegans Em. r. santschii. type For. Marseille vIII. 04 (Santschi)”, “TYPUS”, ANTWEB CASENT 0908353”; MHN genève. Three worker and two gynes from MHN genève labelled by Forel “C. elegans santschii Forel, type, Candia (Biso)” have no type status as the locality Candia is not mentioned in the original description.
 * The type locality of this taxon is Fréjus / France. Investigated were two worker syntypes labelled “Fréjus (var) plage IX. 34 F. Bernard”, “Cardiocondyla provincialis F. Bernard”, “Type”, “ANTWEB CASENT 0915398”; MNHN Paris.
 * This taxon has been described from Pinsac / France. Investigated were 3 ergatoid male syntypes plus 5 workers and 5 gynes labelled “Xenometra gallica F. Bernard”, “Pinsac Lepointe 1951”, “Type”, “ANTWEB CASENT 0915399”, MNHN Paris. This taxon was erected because F. Bernard erroneously considered the ergatoid males found within a nest sample of C. elegans as gynes of a different socially parasitic species.

Worker
Seifert (2003) - Rather large species, CS 560 ± 30. Head of medium length, CL/CW 1.153. Postocular distance smaller than in related species, PoOC/CL 0.397. Eye of medium size, EYE 0.248. Scape longer than in related species, SL/CS 0.847. Pubescence length on 1st tergite much longer than in other species, PLG 41.8. Clypeus more or less glabrous, with few weak longitudinal microrugae in posterior and lateral areas. Frontal lobes and frontocentral vertex posterior of frontal carinae finely longitudinal microrugulose. Frontal carinae usually slightly converging immediately caudal of the FRS level. Whole vertex with shallow but well-demarcated simple or bicoronate foveolae of 16 - 19 mm diameter; interspaces between foveolae on paramedian vertex smaller than their diameter. Dorsal area of promesonotum with shallow foveolae of 10 - 15 mm diameter around bases of pubescence hairs, interspaces glabrous and larger than foveolar diameter. Dorsal area of propodeum glabrous but with small foveolae and very delicate microrugosity. Lateral area of mesosoma in overall impression shining but finely microreticulate-rugulose. Dorsum of waist glabrous, with scattered fragments of very tine microreticular structures. First gaster tergite glabrous. Spines well-developed but rather blunt. Petiole node wider than long. Postpetiolar sternite with anteromedian portion significantly more bulging than anteroparamedian portion; in lateral view this anteromedian bulge forming small, obtusely-angled, rounded corner and changing into helcium with distinct angle. Head, mesosoma, waist, and gaster concolorous dark to blackish brown.

Queen
Seifert (2003) - Rather large, CS 635. Head of medium length (CL/CW 1.1 37 ± 0.015) and with slightly excavated occipital margin. Postocular distance smaller than in related species, PoOc/CL 0.390 ± 0.010. Scape much longer than in related species, SL/CS 0.803 ± 0.014. Whole body covered with very long and dense pubescence. Vertex with rather shallow but well-demarcated, closely-set, bicoronate foveolae; interspaces clearly smaller than foveolar diameter and on frontocentral and paramedian vertex with longitudinal rugulae. Frontal lobes densely rugulose. Lateral area of clypeus with few longitudinal carinulae. Promesonotum, prescutellum, and scutellum densely foveolate. Cuticular surface of dorsal waist and gaster glabrous, with scattered fragments of microreticular structures and numerous basal pits of pubescence hairs. Metapleurae microreticulate and with few longitudinal carinulae on the bulla glandulae metapleuralis. Propodeal spines well-developed but rather blunt. Petiole node in lateral view not produced caudad, in dorsal view nearly twice as wide than long. Postpetiole node twice as wide than long and with weakly concave anterodorsal margin; its sternite with a conspicuous anteroventromedial corner. Whole body concolourous medium to blackish brown. Gynes with mesosoma dimorphism.

References based on Global Ant Biodiversity Informatics

 * Agosti, D. and C.A. Collingwood. 1987. A provisional list of the Balkan ants (Hym. Formicidae) and a key to the worker caste. I. Synonymic list. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 60: 51-62
 * Aktaç, N.. "Studies on the myrmecofauna of Turkey I. Ants of Siirt, Bodrum and Trabzon." Istanbul Universitesi Fen Fakultesi Mecmuasi. Seri B 41 (1977): 115-135.
 * Aldawood AS, Sharaf MR (2011) Monomorium dryhimi sp. n., a new ant species (Hymenoptera, Formicidae) of the M. monomorium group from Saudi Arabia, with a key to the Arabian Monomorium monomorium-group. ZooKeys 106: 4754. doi: 10.3897/zookeys.106.139
 * André, E.. "Species des Hyménoptères d'Europe et d'Algérie. Les fourmis." in: André, Edm. 1881-1886. Species des Hyménoptères d'Europe et d'Algérie. Tome Deuxième. Beaune: Edmond André, 919 + 48 pp. Tomo II (1881): 1016.
 * AntArea. Accessed on February 5th 2014 at http://antarea.fr/fourmi/
 * Antarea (Personal Communication - Rumsais Blatrix- 27 April 2018)
 * Antarea (at www.antarea.fr on June 11th 2017)
 * Barrett K. E. J. 1970. Ants in France, 1968-69. Entomologist 103: 270-274.
 * Bernard F. 1956. Révision des fourmis paléarctiques du genre Cardiocondyla Emery. Bull. Soc. Hist. Nat. Afr. Nord 47: 299-306.
 * Bernard F. 1957. Xenometra Emery, genre de fourmis parasite nouveau pour l'Ancien Monde (Hym. Formicidae). Bull. Soc. Entomol. Fr. 62: 100-103.
 * Bernard F. 1967. Faune de l'Europe et du Bassin Méditerranéen. 3. Les fourmis (Hymenoptera Formicidae) d'Europe occidentale et septentrionale. Paris: Masson, 411 pp.
 * Borowiec L. 2014. Catalogue of ants of Europe, the Mediterranean Basin and adjacent regions (Hymenoptera: Formicidae). Genus (Wroclaw) 25(1-2): 1-340.
 * Borowiec L., and S. Salata. 2012. Ants of Greece - Checklist, comments and new faunistic data (Hymenoptera: Formicidae). Genus 23(4): 461-563.
 * Borowiec L., and S. Salata. 2017. Ants of the Peloponnese, Greece (Hymenoptera: Formicidae). Polish Journal of Entomology 86: 193-236.
 * Bracko G. 2007. Checklist of the ants of Slovenia (Hymenoptera: Formicidae). Natura Sloveniae 9: 15-24
 * Bracko G., H. C. Wagner, A. Schulz, E. Gioahim, J. Maticic, and A. Tratnik. 2014. New investigation and a revised checklist of the ants (Hymenoptera: Formicidae) of the Republic of Macedonia. North-Western Journal of Zoology 10(1): 10-24.
 * Casevitz-Weulersse J., and C. Galkowski. 2009. Liste actualisee des Fourmis de France (Hymenoptera, Formicidae). Bull. Soc. Entomol. Fr. 114: 475-510.
 * Collingwood C., and H. Heatwole. 2000. Ants from Northwestern China (Hymenoptera, Formicidae). Psyche 103 (1-2): 1-24.
 * Collingwood, C. A. 1993. A Comparitive Study of the Ant Fauna of Five Greek Islands. Biologia Gallo-hellenica. 20,1:191-197
 * Collingwood, C. A., and I. H. H. Yarrow. "A survey of Iberian Formicidae." EOS (Revista española de entomología) 44 (1969): 53-101.
 * Collingwood, C. A.. "A comparative study of the ant fauna of five Greek islands." Biologia Gallo-Hellenica 20 (1993): 191-197.
 * Consani M., and P. Zangheri. 1952. Fauna di Romagna. Imenotteri - Formicidi. Memorie della Societa Entomologica Italiana 31: 38-48.
 * Csősz S., B. Markó, and L. Gallé. 2011. The myrmecofauna (Hymenoptera: Formicidae) of Hungary: an updated checklist. North-Western Journal of Zoology 7: 55-62.
 * Czechowski W., A. Radchenko, W. Czechowska and K. Vepsäläinen. 2012. The ants of Poland with reference to the myrmecofauna of Europe. Fauna Poloniae 4. Warsaw: Natura Optima Dux Foundation, 1-496 pp
 * Donisthorpe, H.. "A first instalment of the ants of Turkey." Annals and Magazine of Natural History (12)3 (1950): 1057-1067.
 * Emery C. 1916. Fauna entomologica italiana. I. Hymenoptera.-Formicidae. Bullettino della Società Entomologica Italiana 47: 79-275.
 * Emery, C.. "Beiträge zur Monographie der Formiciden des paläarktischen Faunengebietes. (Hym.) Teil VI. Cardiocondyla, Solenopsis, Oligomyrmex." Deutsche Entomologische Zeitschrift 1909 (1909): 19-37.
 * Emery, C.. "Catalogo delle formiche esistenti nelle collezioni del Museo Civico di Genova. Parte seconda. Formiche dell'Europa e delle regioni limitrofe in Africa e in Asia." Annali del Museo Civico di Storia Naturale 12 (1878): 43-59.
 * Emery, C.. "Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [part]." en Wytsman, P. "Genera Insectorum" 174B (1922): 95-206.
 * Espadaler X., F. Garcia, X. Roig, and R. Vila. 2013. Ants (Hymenoptera, Formicidae) from the Castell de Montesquiu park (Osona, north-east of the Iberian Peninsula). Boletín de la Sociedad Entomológica Aragonesa (S.E.A.) 53: 223-227.
 * Espadaler, X.. "Catalogos entomofauna aragonesa: Familia: Formicidae." Catalogos entomofauna aragonesa 13 (1997): 13-21.
 * Espadaler, X.. "Citas nuevas o interesantes de hormigas para España." Boletín de la Asociación Española de Entomología 3 (1979): 95-101.
 * Finzi B. 1939. Quinto contributo alla conoscenza della fauna mirmecologica della Venezia Giulia. Bollettino della Società Entomologica Italiana 71: 86-90.
 * Forel A. 1905. Miscellanea myrmécologiques II (1905). Ann. Soc. Entomol. Belg. 49: 155-185.
 * Forel A. 1911. Fourmis d'Afrique et d'Asie. I. Fourmis d'Afrique surtout du Musée du Congo Belge. Rev. Zool. Afr. (Bruss.) 1: 274-283.
 * Forel A. 1911. Sur le genre Metapone n. g. nouveau groupe des Formicides et sur quelques autres formes nouvelles. Rev. Suisse Zool. 19: 445-459.
 * Forel, A.. "Miscellanea myrmécologiques II (1905)." Annales de la Société Entomologique de Belgique 49 (1905): 155-185.
 * GRETIA. 2017. Bilan annuel de l'enquete sur la repartition des fourmis armoricaines. 23 pages.
 * Grandi G. 1935. Contributi alla conoscenza degli Imenotteri Aculeati. XV. Boll. R. Ist. Entomol. Univ. Studi Bologna 8: 27-121.
 * Gratiashvili N., Barjadze S. 2008. Checklist of the ants (Formicidae Latreille, 1809) of Georgia. Proceedings of the Institute of Zoology (Tbilisi) 23: 130-146.
 * Guénard B., and R. R. Dunn. 2012. A checklist of the ants of China. Zootaxa 3558: 1-77.
 * Hamann H. H. F., and W. Klemm. 1976. Ergebnisse der von Dr. O. Paget und Dr. E. Kritscher auf Rhodos durchgeführten zoologischen Exkursionen. XVI. Formicidae. Annalen des Naturhistorischen Museums in Wien 80: 669-679.
 * Karaman M. G. 2009. An introduction to the ant fauna of Macedonia (Balkan Peninsula), a check list (Hymenoptera, Formicidae). Natura Montenegrina 8(3): 151-162.
 * Karaman M. G. 2011. A catalogue of the ants (Hymenoptera, Formicidae) of Montenegro. Podgorica: Catalogues 3, Volume 2, Montenegrin Academy of Sciences and Arts, 140 pp.
 * Kiran K., and C. Karaman. 2012. First annotated checklist of the ant fauna of Turkey (Hymenoptera: Formicidae). Zootaxa 3548: 1-38.
 * Kugler J. 1984. The males of Cardiocondyla Emery (Hymenoptera: Formicidae) with the description of the winged male of Cardiocondyla wroughtoni (Forel). Israel Journal of Entomology 17: 1-21.
 * Kugler J. 1988. The zoogeography of Israel. 9. The zoogeography of social insects of Israel and Sinai. Monographiae biologicae 62: 251-275.
 * Lapeva-Gjonova, L., V. Antonova, A. G. Radchenko, and M. Atanasova. "Catalogue of the ants (Hymenoptera, Formicidae) of Bulgaria." ZooKeys 62 (2010): 1-124.
 * Legakis Collection Database
 * Luque-García, G., J. L. Reyes-López, and J. Fernández-Haeger. "Estudio faunístico de las hormigas (Hymenoptera: Formicidae) de la cuenca del río Guadiamar: Primeras aportaciones." Boletín de la Sociedad Entomológica Aragonesa 30 (2002): 153-159.
 * Magboul R., A. Khider, E. Idris, and T. H. Alam. 2013. Ant diversity at Sunut forest, Khartoum, Sudan. Egypt. Acad. J. Biolog. Sci. 6(1): 43-46.
 * Marikovsky P. I. 1979. Ants of the Semireche Desert. [In Russian.]. Alma Ata: Nauka, 263 pp.
 * Medina, M.. "Catálogo provisional de las hormigas de Andalucía." Actas de la Sociedad Española de Historia Natural 20 (1891): 95-104.
 * Menozzi C. 1918. Primo contributo alla conoscenza della fauna mirmecologica del Modenese. Atti della Società dei Naturalisti e Matematici di Modena. (5)4: 81-88.
 * Mercier, J.-L.,J.-C. Lenoir, A. Eberhardt, S. Frohschammer, C. Williams and J. Heinze. 2007. Hammering, mauling, and kissing: stereotyped courtship behavior in Cardiocondyla ants. Insectes Sociaux 54(4):403-411.
 * Ordóñez-Urbano C., J. Reyes-López, and S. Carpintero-Ortega. 2007. Estudio faunísidos de los formícidos (Hymenoptera: Formicidae) asociados a los bosques de ribera en la Provincia de Córdoba (España). Primeras Aportaciones. Boletin Sociedad Entomológica Aragonesa 40: 367-375.
 * Paknia O., A. Radchenko, H. Alipanah, and M. Pfeiffer. 2008. A preliminary checklist of the ants (Hymenoptera: Formicidae) of Iran. Myrmecological News 11: 151-159.
 * Paraschivescu D. 1978. Elemente balcanice in mirmecofauna R. S. Romania. Nymphaea 6: 463- 474.
 * Pashaei Rad S., B. Taylor, R. Torabi, E. Aram, G. Abolfathi, R. Afshari, F. Borjali, M. Ghatei, F. Hediary, F. Jazini, V. Heidary Kiah, Z. Mahmoudi, F. Safariyan, and M. Seiri. 2018. Further records of ants (Hymenoptera: Formicidae) from Iran. Zoology in the Middle East 64(2): 145-159.
 * Peru L. 1999. Quelques ajouts et corrections a un catalogue des fourmis de la region Centre (Hymenoptera, Formicidae). Bulletin de l'Entomologie Tourangelles 20(1): 9-12.
 * Petrov I. Z. 2002. Contribution to the myrmecofauna (Formicidae, Hymenoptera) of the Banat Province (Serbia). Archives of Biological Sciences, Belgrade, 54(12): 57-64.
 * Petrov I. Z., and A. Legakis. 1996. Contribution to the Myrmecofauna (Formicidae, Hymenoptera) of Greece. Archives of biological sciences, Belgrade, 48 (3-4): 31-32.
 * Petrov I. Z., and C. A. Collingwood. 1992. Survey of the myrmecofauna (Formicidae, Hymenoptera) of Yugoslavia. Archives of Biological Sciences (Belgrade) 44: 79-91.
 * Radchenko A. G. 1996. Palaearctic ants of the genus Cardiocondyla Emery (Hymenoptera, Formicidae). Entomological Review (Washington) 75(7): 99-109.
 * Radchenko A. G. 1996. Palaearctic ants of the genus Cardiocondyla Emery (Hymenoptera, Formicidae). Entomological Review (Washington). 75(7): 99-109.
 * Reyes, J. L., and G. Luque. "Nuevos datos sobre el género Cardiocondyla (Hym.: Formicidae) en andalucía (Sur de la Península Ibérica)." Boletín de la Asociación Española de Entomología 25 (3-4) (2001): 125-126.
 * Salata S., and L Borowiec. 2017. Species of Tetramorium semilaeve complex from Balkans and western Turkey, with description of two new species of (Hymenoptera: Formicidae: Myrmicinae). Annales Zoologici (Warsaw) 62:279–313.
 * Salata S., and L. Borowiec. 2018. Taxonomic and faunistic notes on Greek ants (Hymenoptera: Formicidae). Annals of the Upper Silesian Museum in Bytom Entomology 27: 1-51.
 * Scupola A. 2009. Le specie italiane de genre Cardiocondyla (Hymenoptera Formicidae). Bollettino della Società Entomologica Italiana 141: 187-190.
 * Seifert B. 2003. 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 in Wien. B, Botanik, Zoologie 104: 203-338.
 * Seifert B., I. Okita,and J. Heinze. 2017. A taxonomic revision of the Cardiocondyla nuda group (Hymenoptera: Formicidae). Zootaxa 4290: 324-356.
 * Seifert, B.. "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." Ann. Naturhist. Mus. Wien 104B (2003): 203-338.
 * Suñer i Escriche, David. "Contribució al coneixement mirmecologic de Gavarres, Montgrí, Guilleríes i la Serralada Transversal." Tesis Doctoral Universida (1991): 577 pp.
 * Tohme G. 1996. Formicidae. Etude de la diversité biologique n° 4 . Ministère de lAgriculture à Beyrouth (Eds.). P85-87.
 * Vonshak M., and A. Ionescu-Hirsch. 2009. A checklist of the ants of Israel (Hymenoptera: Formicidae). Israel Journal of Entomology 39: 33-55.
 * Wagner H. C., B. Seifert, R. Borovsky, and W. Paill. 2018. First insight into the ant diversity of the Vjosa valley, Albania (Hymenoptera: Formicidae). Acta ZooBot Austria 155: 315–321.