Cataglyphis bicolor

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Cataglyphis bicolor
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Hymenoptera
Family: Formicidae
Subfamily: Formicinae
Tribe: Formicini
Genus: Cataglyphis
Species group: bicolor
Species: C. bicolor
Binomial name
Cataglyphis bicolor
(Fabricius, 1793)

Cataglyphis bicolor casent0104612 profile 1.jpg

Cataglyphis bicolor casent0104612 dorsal 1.jpg

Specimen Label

Subspecies
Synonyms

This is one of two species of Cataglyphis (also Cataglyphis fortis) that have had their foraging behavior, and in particular their navigation abilities, intensively studied. The results of this body of work provide a remarkable accounting of how ants, and animals in general, can track and monitor their movement in ways that allow for sophisticated decision making. These studies have examined these two ants' sensory and nervous systems, their directional abilities, use of path integration, and view-based landmark guidance, and how all of this is brought together to determine where an individual decides to move. These ants are now a model organism for the study of animal navigation. Rüdiger Wehner has been at the center of much of this Cataglyphis research. Wehner and his colleagues have been conducting seasonal fieldwork studying C. bicolor and C. fortis in the area of Mahrès, Tunisia for 50 years (Wehner 2019). The book Desert Navigator:The Journey of an Ant (Wehner 2020) provides a fascinating summary of how this work was accomplished and its many important findings.

Identification

Keys including this Species

Distribution

Latitudinal Distribution Pattern

Latitudinal Range: 40° to 22.03642884°.

     
North
Temperate
North
Subtropical
Tropical South
Subtropical
South
Temperate

Distribution based on Regional Taxon Lists

Afrotropical Region: Eritrea, Nigeria.
Palaearctic Region: Algeria, Canary Islands.

Distribution based on AntMaps

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Distribution based on AntWeb specimens

Check data from AntWeb

Countries Occupied

Number of countries occupied by this species based on AntWiki Regional Taxon Lists. In general, fewer countries occupied indicates a narrower range, while more countries indicates a more widespread species.
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Estimated Abundance

Relative abundance based on number of AntMaps records per species (this species within the purple bar). Fewer records (to the left) indicates a less abundant/encountered species while more records (to the right) indicates more abundant/encountered species.
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Biology

Morphology

Penmetcha et al. (2019) - Desert ants of the genus Cataglyphis and Melophorus possess all three ocelli and are the only ants in which the function of the ocelli has been studied. Size of the ocellar lens and width of the ocellar rhabdoms in cross-section are typically larger in nocturnal insects (Warrant et al. 2006; Somanathan et al. 2009; Narendra et al. 2011; Narendra and Ribi 2017). This increases their optical sensitivity and allows individuals with larger ocelli to forage in slightly dim light conditions (Wellington 1974). The desert ants that we studied being strictly day-active had small lenses and narrow rhabdoms (in Cataglyphis spp) compared to the night-active Myrmecia.

Here we characterised the anatomical organisation of the ocelli in three species of desert ants (Cataglyphis bicolor, Cataglyphis fortis, and Melophorus bagoti). Cataglyphis bicolor, an ant that forages in both saltpans and around urban regions, had a fused rhabdom. Each rhabdom was composed of two retinula cells that contributed microvilli perpendicular to the long axis of the rhabdom. This organisation of the ocellar retina is likely to make them polarisation sensitive. Similarly organized rhabdoms are known from winged male ants (Narendra and Ribi 2017), bees, and wasps (Zeil et al. 2014; Ribi and Zeil 2018). Our anatomical data lend support to the behavioural (Fent and Wehner 1985; Fent 1986) and physiological evidence (Mote and Wehner 1980) of polarisation sensitivity in the ocelli of C. bicolor.

Association with Other Organisms

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Life History Traits

  • Mean colony size: 2,000 (Wehner et al., 1983; Beckers et al., 1989)
  • Foraging behaviour: solitary forager (Wehner et al., 1983; Beckers et al., 1989)

Castes

Queen

Images from AntWeb

Cataglyphis bicolor casent0104613 head 1.jpgCataglyphis bicolor casent0104613 head 2.jpgCataglyphis bicolor casent0104613 profile 1.jpgCataglyphis bicolor casent0104613 dorsal 1.jpgCataglyphis bicolor casent0104613 label 1.jpg
Paratype of unpublished name: Myrmecocystus bicolor gibbosusQueen (alate/dealate). Specimen code casent0104613. Photographer April Nobile, uploaded by California Academy of Sciences. Owned by ZMHB, Berlin, Germany.

Nomenclature

The following information is derived from Barry Bolton's Online Catalogue of the Ants of the World.

  • bicolor. Formica bicolor Fabricius, 1793: 351 (m.) (no state data, “Habitat in Barbaria”, = North African coast from Morocco to Libya).
    • [Misspelled as biceler by Menozzi, 1929e: 128; misspelled as bicolora by Borowiec, L. & Salata, 2012: 486.]
    • Athias-Henriot, 1947: 249 (l.); Wheeler, G.C. & Wheeler, J. 1980: 538 (l.).
    • Combination in Myrmecocystus: Emery & Forel, 1879: 449;
    • combination in Myrmecocystus (Cataglyphis): Santschi, 1910f: 234;
    • combination in Cataglyphis: Roger, 1863b: 12; Mayr, 1863: 402; Emery, 1912f: 98; Emery, 1915g: 22;
    • combination in Cataglyphis (Cataglyphis): Santschi, 1929b: 41.
    • Junior synonym of viatica: Nylander, 1856b: 59; Smith, F. 1858b: 9; Smith, F. 1861a: 32; Mayr, 1863: 402; Roger, 1863b: 12; Emery & Forel, 1879: 449; Nasonov, 1889: 14; Dalla Torre, 1893: 218.
    • Subspecies of viatica: Forel, 1907e: 15; Forel, 1908a: 14; Forel, 1909e: 386; Forel 1910c: 266; Forel, 1910e: 450; Wheeler, W.M. & Mann, 1916: 173; Stitz, 1917: 348; Wheeler, W.M. 1922a: 944.
    • Status as species: Latreille, 1802c: 123; Fabricius, 1804: 398; Lucas, H. 1849: 302; Smith, F. 1858b: 31; Emery & Forel, 1879: 449; Emery, 1906d: 58; Mayr, 1907a: 389; Forel, 1907e: 15; Santschi, 1908: 518; Santschi, 1910f: 234; Karavaiev, 1910b: 39; Karavaiev, 1911: 10; Karavaiev, 1912b: 590; Emery, 1915g: 22; Viehmeyer, 1923: 92; Müller, 1923b: 149; Karavaiev, 1924: 306; Emery, 1925b: 264; Menozzi, 1927g: 380; Santschi, 1929b: 41; Finzi, 1930b: 16; Finzi, 1930c: 24; Karavaiev, 1932: 250; Menozzi, 1933b: 87 (in key); Menozzi, 1934: 166Finzi, 1936: 192; Santschi, 1936c: 210; Finzi, 1939a: 160; Finzi, 1940: 165; Donisthorpe, 1942a: 33; Bernard, 1945: 137; Athias-Henriot, 1947: 249; Donisthorpe, 1947e: 111; Donisthorpe, 1950e: 1065; Bernard, 1953a: 198; Cagniant, 1964: 93; Cagniant, 1966b: 281; Cagniant, 1968a: 145; Cagniant, 1970c: 30; Bernard, 1971: 8; Hamann & Klemm, 1976: 674; Agosti, 1990b: 1486; Wehner, et al. 1994: 168; Bolton, 1995b: 134; Cagniant, 2006a: 194; Cagniant, 2009: 49; Legakis, 2011: 34; Borowiec, L. & Salata, 2012: 486; Kiran & Karaman, 2012: 9; Borowiec, L. 2014: 52 (see note in bibliography); Tohmé, G. & Tohmé, 2014: 139; Madl, 2019: 14.
    • Senior synonym of fairmairei: Santschi, 1929b: 41; Agosti, 1990b: 1487; Bolton, 1995b: 134.
    • Senior synonym of megalocola: Mayr, 1855: 382; Santschi, 1929b: 41; Agosti, 1990b: 1486; Bolton, 1995b: 134.
    • Senior synonym of rotundinodis: Santschi, 1929b: 41; Agosti, 1990b: 1487; Bolton, 1995b: 134.
    • Current subspecies: nominal plus adusta, basalis, congolensis, pubens, seticornis, sudanica.
  • fairmairei. Cataglyphis fairmairei Foerster, 1850b: 494 (m.) ALGERIA.
    • Junior synonym of viatica: Nylander, 1856b: 59; Smith, F. 1858b: 9; Mayr, 1861: 45 (in key); Smith, F. 1861a: 32; Mayr, 1863: 402; Roger, 1863b: 12; Dours, 1873: 166; André, 1874: 202 (in list).
    • Junior synonym of megalocola: Emery & Forel, 1879: 449; Dalla Torre, 1893: 217; Emery, 1925b: 266.
    • Junior synonym of bicolor: Santschi, 1929b: 41; Agosti, 1990b: 1487; Bolton, 1995b: 134.
  • megalocola. Formica megalocola Foerster, 1850b: 490 (w.) ALGERIA.
    • [Misspelled as megalochola by Smith, F. 1861a: 32.]
    • Emery, 1891b: 16 (q.m.).
    • Combination in Myrmecocystus: Emery & Forel, 1879: 449; Forel, 1890a: lxvii;
    • combination in Cataglyphis: Roger, 1863b: 12; Mayr, 1863: 402; Emery, 1912f: 98; Emery, 1925b: 265.
    • As unavailable (infrasubspecific) name: Stitz, 1917: 348.
    • Subspecies of viatica: Emery & Forel, 1879: 449; Emery, 1880: 389; André, 1882b: 167 (in key); Emery, 1884a: 376; Forel, 1890a: lxvii; Emery, 1891b: 16; Forel, 1892i: 306; Emery, in Dalla Torre, 1893: 218 (footnote); Forel, 1894d: 9; Ruzsky, 1905b: 435; Forel, 1907e: 15; Forel, 1909e: 377.
    • Status as species: Dalla Torre, 1893: 217.
    • Subspecies of bicolor: Emery, 1906d: 58; Emery, 1908g: 217; Karavaiev, 1912a: 17; Emery, 1925b: 265; Donisthorpe, 1950e: 1066.
    • Junior synonym of viatica: Mayr, 1855: 382; Nylander, 1856b: 59; Smith, F. 1858b: 9; Smith, F. 1861a: 32; Mayr, 1863: 402; Roger, 1863b: 12; André, 1874: 202 (in list).
    • Junior synonym of nodus: Agosti & Collingwood, 1987a: 59.
    • Junior synonym of bicolor: Mayr, 1855: 382; Santschi, 1929b: 41; Agosti, 1990b: 1486; Bolton, 1995b: 136.
  • rotundinodis. Myrmecocystus albicans subsp. rotundinodis Karavaiev, 1912a: 16 (w.) ALGERIA.
    • Combination in Cataglyphis: Emery, 1925b: 262.
    • Subspecies of albicans: Karavaiev, 1924: 305; Emery, 1925b: 262.
    • Junior synonym of bicolor: Santschi, 1929b: 41; Agosti, 1990b: 1487; Bolton, 1995b: 137.

Taxonomic Notes

Borowiec & Salata (2020): Cataglyphis bicolor group is a complex of similar species difficult to identify. Eyer et al. (2017) support the occurrence of at least four distinct species in the C. bicolor group in Israel, one of which may be a complex of three more species.

Description

Karyotype

  • n = 26 (Israel; Tunisia) (Hauschteck-Jungen & Jungen, 1983).

References

References based on Global Ant Biodiversity Informatics

  • Agosti D. 1990. Review and reclassification of Cataglyphis (Hymenoptera, Formicidae). Journal of Natural History 24: 1457-1505.
  • Agosti D., C. Austin, O. A. Gökcen, W. A. König, E. D. Morgan, E. D. Scott, R. Wehner. 1996. (S)-2-methyl-1-hexanol, characteristic mandibular gland substance of ants of the Cataglyphis bicolor group. Chemoecology 7: 57-60.
  • Agosti, D. 1990. Review and reclassification of Cataglyphis (Hymentopera: Formicidae) . Journal of Natural History 24:1457-1505.
  • Aissat L., and R. Moulai. 2016. Analysis of insect diversity of three islets in the North East Algeria. J. Entomol. Res. Soc. 18(2): 1-21.
  • Alfieri A. 1931. Contribution à l'étude de la faune myrmécologique de l'Egypte. Bulletin. Société Entomologique d'Egypte 15: 42-48.
  • Alfieri, A.. "Contribution à l'étude de la faune myrmécologique de l'Egypte." Bulletin de la Société Entomologique d'Egypte 15 (1931): 42-48.
  • Andoni V. 1977. Kontribut mbi Himenopteret e familjes Formicidae te vendit tone. Buletini I Shkencave te Natyres 31(2): 93-101.
  • Barech G., M. Khaldi, F. Zahra Boujelal, and X. Espadaler. 2018. Diversity and structure of the ant fauna of the El Ksob dam in Algeria: new record for Aphaenogaster rupestris Forel, 1909 (Hymenoptera: Formicidae). Boletin de la Sociedad Entomologica Aragonesa (S. E. A.) 62: 253-258.
  • Barech G., M. Khaldi, S. Ziane, A. Zedam, S. Doumadji, M. Sharaf, and X. Espadaler. 2016. A first checklist and diversity of ants (Hymenoptera: Formicidae) of the saline dry lake Chott El Hodna in Algeria, a Ramsar Conservation Wetland. African Entomology 24(1): 143–152.
  • Bernard F. 1970. Les fourmis de la forêt de Mâmora (Maroc). Revue d'Écologie et de Biologie du Sol 6:n483-513.
  • 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.
  • Bouzekri M. A. 2011. Bioecologie des Formicidae dans la region de Djelfa: nidification et relation avec les plantes. Mphil Ecole National Superieure agronomique. 153pp.
  • Bouzekri M., S. Daoudi-Hacini, H. Cagniant, and S. E. Doumandji. 2015. Etude comparative des associations (plantes-fourmis) dans une region steppique (cas de la region de Djelfa, Algerie). Lebanese Science Journal 16(1): 69-77.
  • Cagniant H. 2009. Le genre Cataglyphis Foerster, 1850 au Maroc (Hyménoptères Formicidae). Orsis 24: 41-71.
  • Cagniant, H. 1968. Liste preliminaire de fourmis forestieres d'Algerie. Resultats obtenus de 1963 a 1964. Bulletin de la Société d'Histoire Naturelle de Toulouse 104: 138-147
  • Cagniant, H.. "Deuxième liste de fourmis d'Algérie, récoltées principalement en forêt (Deuxième partie)." Bulletin de la Société d'Histoire Naturelle de Toulouse 106 (1970): 28-40.
  • Cagniant, H.. "Liste actualisee des fourmis du Maroc (Hymenoptera: Formicidae)." Myrmecologische Nachrichten 8 (2006): 193-200.
  • Cagniant, H.. "Liste préliminaire de fourmis forestières d'Algérie. Résultats obtenus de 1963 à 1966." Bulletin de la Societe d'Histoire Naturelle de Toulouse 104 (1968): 138-147.
  • Cagniant, H.. "Note sur le peuplement en fourmis d'une montagne de la région d'Alger, l'Atlas de Blida." Bulletin de la Société d'Histoire Naturelle de Toulouse 102 (1966): 278-284.
  • Cagniant, H.. Les peuplements de fourmis des forêts algériennes: écologie, biocénotique, essai biologique. Universite de Toulouse, 1973.
  • Chemala A., M. Benhamacha, D. M. Ould el Hadj, F. Marniche, and S. Daoudi. 2017. A preliminary list of the ant fauna in Northeastern Sahara of Algeria (Hymenoptera: Formicidae). Sociobiology 64(2): 146-154.
  • Delye, G., and J. L. Bonaric. "Fourmis du sud Marocain." Etude de certains milieux du Maroc et de leux evolution recente RCP 249 (1973).
  • Dietrich, B. and R. Wehner. 2003. Sympatry and Allopatry in Two Desert Ant Sister Species: How Do Cataglyphis bicolor and C. savignyi Coexist? Oecologia136 (1) :63-72
  • Dillier, F.-X. and R. Wehner. 2004. Spatio-temporal patterns of colony distribution in monodomous and polydomous species of North African desert ants, genus Cataglyphis. Insectes Sociaux 51(2):186-196.
  • Donisthorpe H. 1942. The Formicidae (Hym.) of the Armstrong College Expedition to the Siwa Oasis. Annals and Magazine of Natural History (11)9: 26-33.
  • Donisthorpe H. 1947. Results of the Armstrong College Expedition to Siwa Oasis (Libyan Desert), 1935: A second list of the Formicidae (Hymenoptera). Bulletin de la Société Fouad Ier d'Entomologie 31: 109-111.
  • Emery C. 1906. Rassegna critica delle specie paleartiche del genere Myrmecocystus. Memorie della Reale Accademia delle Scienze dell'Istituto di Bologna (6)3: 47-61.
  • Emery C. 1915. Formiche raccolte nell'Eritrea dal Prof. F. Silvestri. Bollettino del Laboratorio di Zoologia Generale e Agraria della Reale Scuola Superiore d'Agricoltura. Portici 10: 3-26.
  • Emery, C.. "Crociera del Violante, comandato dal capitano armatore Enrico d'Albertis, durante l'anno 1877. Formiche." Annali del Museo Civico di Storia Naturale 15 (1880): 389-398.
  • Emery, C.. "Escursioni zoologiche del Dr. Enrico Festa nell'Isola di Rodi. XII. Formiche." Bollettino del Museo di Zoologia ed Anatomia Comparatadella Reale Università di Torino 30 (1915): 1-7.
  • Emery, C.. "Exploration scientifique de la Tunisie. Zoologie. - Hyménoptères. Révision critique des fourmis de la Tunisie." Explor. Scient. De la Tunisie Zoll. Hym. (Folleto) Paris. Imp (1891): iii + 21 pp.
  • Forel A. 1890. Fourmis de Tunisie et de l'Algérie orientale. Annales de la Société Entomologique de Belgique 34: lxi-lxxvi.
  • Forel A. 1907. Formiciden aus dem Naturhistorischen Museum in Hamburg. II. Teil. Neueingänge seit 1900. Mitt. Naturhist. Mus. Hambg. 24: 1-20.
  • Forel A. 1909. Études myrmécologiques en 1909. Fourmis de Barbarie et de Ceylan. Nidification des Polyrhachis. Bull. Soc. Vaudoise Sci. Nat. 45: 369-407.
  • Forel A. 1910. Note sur quelques fourmis d'Afrique. Annales de la Société Entomologique de Belgique 54: 421-458.
  • Forel, A. 1894. Les formicides de la province d'Oran (Algerie). Bulletin de la Société Vaudoise des Sciences Naturelles 30: 1-45.
  • Forel, A. 1908. Fourmis de Ceylan et d'Égypte récoltées par le Prof. E. Bugnion. Lasius carniolicus. Fourmis de Kerguelen. Pseudandrie? Strongylognathus testaceus. Bull. Soc. Vaudoise Sci. Nat. 44: 1-22
  • Forel, A.. "Fourmis de Tunisie et de l'Algerie orientale recoltees et decrites par Auguste Forel." Annales de la Société Entomologique de Belgique 34 (1890): lxi-lxxvi.
  • Forel, A.. "Les Formicides de la Province d'Oran (Algérie)." Bulletin de la Société Vaudoise des Sciences Naturelles 30 (1894): 1-45.
  • Guehef Z. H., Y.Kherbouche, L. Ababsa, L. Eddoud, O. Guezoul, and M. Sekour. 2018. Comparison of myrmicofaune in two agricultural environments in the region of Oued Souf. Revue des BioRessources 8(1): 53-60.
  • 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.
  • Hauschteck-Jungen E., and H. Jungen. 1983. Ant chromosomes. II. Karyotypes of western palearctic species. Insectes Soc. 30: 149-164.
  • Karavaiev V. 1912. Ameisen aus Tunesien und Algerien, nebst einigen unterwegs in Italien gesammelten Arten. Rus. Entomol. Obozr. 12: 1-22.
  • Karavaiev V. 1924. Zur Systematik der paläarktischen Myrmecocystus (Formicidae), nebst einigen biologischen Notizen. Konowia 3: 301-308.
  • Kiran K., and C. Karaman. 2012. First annotated checklist of the ant fauna of Turkey (Hymenoptera: Formicidae). Zootaxa 3548: 1-38.
  • Knaden M., and R. Wehner. 2005. The coexistence of two large-sized thermophilic desert ants: the question of niche differentiation in Cataglyphis bicolor and Cataglyphis mauritanica (Hymenoptera: Formicidae). Myrmecologische Nachrichten 7: 31-42.
  • Kuznetsov G. T. 1990. Comparative analysis of Hymenoptera (Formicidae) population on altitudinal zones of central Kopetdag. Izvestiya Akademii Nauk Turkmenskoi SSR. Seriya Biologicheskikh Nauk 1990(3): 64-67.
  • Madl M. 2019. Notes on the ant fauna of Eritrea (Insecta: Hymenoptera: Formicidae): type specimens deposited in the Natural History Museum Vienna (Austria) and a preliminary checklist. Ann. Naturhist. Mus. Wien, B 121: 9-18.
  • Mayr, G.. "Ameisen aus Tripolis und Barka." Zoologische Jahrbücher, Abteilung für Systematik, Geographie und Biologie der Tiere 26 (1908): 415-418.
  • Medler J. T. 1980: Insects of Nigeria - Check list and bibliography. Mem. Amer. Ent. Inst. 30: i-vii, 1-919.
  • Menozzi C. 1927. Risultati zoologici della Missione inviata dalla R. Società Geografica Italiana per l'esplorazione dell'Oasi di Giarabub (1926-1927). Formicidae (Hymenoptera).. Annali del Museo Civico di Storia Naturale Giacomo Doria. 52: 379-382.
  • Menozzi C. 1929. Ricerche faunistiche nelle isole italiane dell'Egeo. Imenotteri (formiche). Archivio Zoologico Italiano. 13: 145-146.
  • Menozzi C. 1934. Reperti mirmecofaunistici raccolti dal Prof. L di Caporiacco nelle oasi di Cufra e in altre localita del deserto Libico. Atti della Societa dei Naturalisti e Matematici di Modena (6)13(65): 153-166
  • Santschi, F. 1938. Mission Robert Ph. Dollfus en Égypte. XV. Hymenoptera: Formicidae. Mémoires de l'Institut d'Égypte 37: 253
  • Santschi, F.. "Étude sur les Cataglyphis." Revue Suisse de Zoologie 36 (1929): 25-70.
  • Santschi, F. 1908. Nouvelles fourmis de l'Afrique du Nord (Égypte, Canaries, Tunisie). Annales de la Société Entomologique de France 77: 517-534.
  • Tak N. 2009. Ants Formicidae of Rajasthan. Records of the Zoological Survey of India, Occasional Paper No. 288, iv, 46 p
  • Tohme G., and H. Tohme. 2014. Nouvelles liste des especes de fourmis du Liban (Hymenoptera, Formicoidea). Lebanese Science Journal 15(1): 133-141.
  • Viehmeyer H. 1923. Wissenschaftliche Ergebnisse der mit Unterstützung der Akademie der Wissenschaften in Wien aus der Erbschaft Treitl von F. Werner unternommenen zoologischen Expedition nach dem anglo-ägyptischen Sudan (Kordofan) 1914. VII. Hymenoptera A. Formicidae. Denkschriften der Akademie der Wissenschaften in Wien. Mathematisch-Naturwissenschaftliche Klasse 98: 83-94.
  • Wehner R., S. Wehner, and D. Agosti. 1994. Patterns of biogeographic distribution within the bicolor species group of the North African desert ant, Cataglyphis Foerster 1850 (Insecta: Hymenoptera: Formicidae). Senckenb. Biol. 74: 163-191.
  • Wheeler W. M. 1922. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VIII. A synonymic list of the ants of the Ethiopian region. Bulletin of the American Museum of Natural History 45: 711-1004