Extreme, workerless inquilinism.

My thread:

https://www.ameisenportal.eu/v…05e1dc6be68c6a65508ef1b39

The extreme, workerless inquilines.

Teleutomyrmex Kutter, 1950

01) Teleutomyrmex schneideri Kutter, 1950
(= Tetramorium schneideri (Kutter, 1950))
(not Tetramorium schneideri Emery, 1898)
(= Tetramorium inquilinum Ward, Brady, Fisher & Schultz, 2015 ("2014"))
02) Teleutomyrmex kutteri Tinaut, 1990
(= Tetramorium kutteri (Tinaut, 1990))
(not Tetramorium semilaeve kutteri Santschi, 1927)
03) Teleutomyrmex seiferti Kiran & Karaman, in Kiran, et al. 2017
(= Tetramorium seiferti (Kiran & Karaman, in Kiran, et al. 2017))
04) Teleutomyrmex buschingeri Lapeva-Gjonova, in Kiran, et al. 2017
(= Tetramorium buschingeri (Lapeva-Gjonova, in Kiran, et al. 2017))

Not yet described species of extreme, workerless inquiline

05) The new, undescribed species from Teleutomyrmex from Farab, Turkmenistan… See Dlussky, et al. 1990.

Anergates Forel, 1874

06) Anergates atratulus (Schenck, 1852)
(= Myrmica atratula Schenck, 1852)
(= Tetramorium atratulum (Schenck, 1852))
[= Tomognathus atratulus (Schenck, 1852), obsolete combination.]
07) Anergates friedlandi Creighton, 1934
[= Tetramorium friedlandi (Creighton, 1934)]

Tetramorium Mayr, 1855
(Only a few species in a big genus.)

08) Tetramorium microgyna Santschi, 1918
09) Tetramorium parasiticum Bolton, 1980

Pheidole Westwood, 1839
(Only a few species in a big genus.)

10) Pheidole neokohli Wilson, 1984
(= Pheidole kohli (Wasmann, 1915))
(= Anergatides kohli Wasmann, 1915)
(not Pheidole kohli Mayr, 1901)
11) Pheidole acutidens (Santschi, 1922)
(= Bruchomyrma acutidens Santschi, 1922)
12) Pheidole argentina (Bruch, 1932)
(= Gallardomyrma argentina Bruch, 1932)
13) Pheidole parasitica Wilson, 1984

Excluded from the extreme, workerless inquilines
Once this species was included in the extreme, workerless inquilines but now it is considered to be a workerless inquiline without extreme reductions, e.g. no pupoid males but normal ones. The decision to exclude it was made by Edward Osborne Wilson in 1984 in a study of the inquilines in the genus Pheidole.

Pheidole Westwood, 1839
(Only one species in a big genus.)

14) Pheidole kusnezovi Wilson, 2003
(= Pheidole symbiotica (Kusnezov, 1952))
(= Eriopheidole symbiotica Kusnezov, 1952)
(not Pheidole symbiotica Wasmann, 1909)

Distribution

01) Europe (Alps, Pyrenees and Northern Spain)
02) Europe (Southern Iberia)
03) Turkey (Anatolia)
04) Europe (Southern Balkans)

05) Turkmenistan

06) Europe
07) North America

08) Southern Africa
09) Southern Africa

10) Central Africa
11) South America
12) South America
13) India

14) South America

Host species

01), 02), 03), 04), 05), 06), 07), 08) and 09) Certain species of the genus Tetramorium Mayr, 1855

01) T. alpestre Steiner, Schlick-Steiner & Seifert, 2010 and T. impurum (Förster, 1850)
and maybe T. caespitum (Linnaeus, 1758)?
02) T. cf. caespitum (Linnaeus, 1758)
03) T. cf. chefketi Forel, 1911
04) T. cf. chefketi Forel, 1911

05) A species from the genus Tetramorium Mayr, 1855…

06) T. impurum (Förster, 1850), T. caespitum (Linnaeus, 1758), T. immigrans Santschi, 1927
and T. moravicum Kratochvil, 1941, T. diomedeum Emery, 1908, T. chefketi Forel, 1911
07) T. immigrans Santschi, 1927

08) T. sericeiventre Emery, 1877 and T. sepositum Santschi, 1918
09) T. avium Bolton, 1980

10), 11), 12), 13) and 14) Certain species of the genus Pheidole Westwood, 1839

10) P. megacephala (Fabricius, 1793) subsp. melancholica Santschi, 1912
11) P. nitidula Emery, 1888
12) P. nitidula Emery, 1888
13) P. indica Mayr, 1879

14) P. susannae Forel, 1886

A remark about synonymy

Anergates friedlandi Creighton, 1934 is now a synonym from Anergates atratulus (Schenck, 1852), more precisely an introduced form in North Americe. So, the name is Anergates atratulus (Schenck, 1852) or, if you follow Ward et al. 2015 ("2014"), Tetramorium atratulum (Schenck, 1852)...

Synonyms of host species

- T. caespitum (Linnaeus, 1758) (= Formica caespitum Linnaeus, 1758)
- T. impurum (Förster, 1850) (= Myrmica impura Förster, 1850)
- P. megacephala (Fabricius, 1793) (= Formica megecephala Fabricius, 1793)
- P. megacephala (Fabricius, 1793) subsp. melancholica Santschi, 1912 was originally described as P. punctulata Mayr, 1866 st. melancholica Santschi, 1912.
- P. susannae Forel, 1886 (= P. obscurior Forel, 1886)

An official literature list of Teleutomyrmex Kutter, 1950.

Baroni Urbani, C., 1967, “Le distribuzioni geografiche discontinue dei Formicidi mirmecobiotici.” Archivio Botanico e Biogeografico Italiano, vol. 43, [=(4)12], p. 355-365.

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Baur, A., Chalwatzis, N., Buschinger, A., Zimmermann, F. K., 1995, “Mitochondrial DNA sequences reveal close relationships between social parasitic ants and their host species.” Current Genetics, vol. 28, p. 242-247.

Baur, A., Sanetra, M., Chalwatzis, N., Buschinger, A., Zimmermann, F. K., 1996, “Sequence comparisons of the internal transcribed spacer region of ribosomal genes support close relationships between parasitic ants and their respective host species (Hymenoptera: Formicidae).” Insectes Sociaux, vol. 43, p. 53-67.

Bernard, F., 1967 ["1968"], “Faune de l'Europe et du Bassin Méditerranéen. 3. Les fourmis (Hymenoptera Formicidae) d'Europe occidentale et septentrionale.” Paris, Masson, 411 pp.

Blaimer, B. B., Ward, P. S., Schultz, T. R., Fisher, B. L., Brady, S. G., 2018, “Paleotropical Diversification Dominates the Evolution of the Hyperdiverse Ant Tribe Crematogastrini (Hymenoptera: Formicidae).” Insect Systematics and Diversity, vol. 2, no. 5, art. 3, p. 1-14.

Bolton, B., 1976, “The ant tribe Tetramoriini (Hymenoptera: Formicidae). Constituent genera, review of smaller genera and revision of Triglyphothrix Forel.” Bulletin of the British Museum (Natural History) (Entomology), vol. 34, p. 281-379.

Bolton, B., 1994, “Identification guide to the ant genera of the world.” Cambridge, Mass., Harvard University Press, 222 pp.

Bolton, B., 1995, “A taxonomic and zoogeographical census of the extant ant taxa (Hymenoptera: Formicidae).” Journal of Natural History, vol. 29, p. 1037-1056.

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-> Dobrzanska, J. & Dobrzanski, J., 1988, “1. Inkwilinizm.”, or “[Inquilinism.]” Kosmos, vol. 37, no. 4, p. 617-629.

-> Dobrzanska, J. & Dobrzanski, J., 1988, “2. Niewolnictwo.”, or “[Slavery.]” Kosmos, vol. 37, no. 4, p. 629-639.

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- His name is spelled “Göszwald” in this publication!

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Stumper, R., 1956, “Etudes myrmécologiques LXXVII. Les sécrétions attractives des reines de fourmis.” Mitteilungen der Schweizerischen Entomologischen Gessellschaft, vol. 29, no. 4, p.373-380.

Stumper, R., Kutter, H., 1950, “Sur le stade ultime du parasitisme social chez les fourmis, atteint par Teleutomyrmex Schneideri (subtrib. nov.; gen. nov.; spec. nov. Kutter).” Comptes Rendus (Hebdomadaires) des Séances de l'Academie des Sciences, vol. 231, p. 876-878.

Tinaut, A., 1981, “Estudio de los Formícidos de Sierra Nevada.” Tesis Doctoral, Universidad de Granada, 463 pp.

Tinaut, A., 1990, “Teleutomyrmex kutteri, spec. nov. A new species from Sierra Nevada (Granada, Spain).” Spixiana, vol. 13, no. 2, p. 201-208.

Tinaut, A., 2013, “Las Hormigas. (Hymenoptera: Formicidae).” P. 392-408 and p. 517-518 in Vol. 2 in: Ruano, F., Tierno de Figueroa, M., Tinaut, A. (eds., Asociación Española de Entomología, Universidad de León), 2013, “Los Insectos de Sierra Nevada. 200 Años de Historia. Vol. 1 & 2.” Asociación Española de Entomología, 544 pp. and 524 pp.

Tinaut, A., Martínez-Ibáñez, M. D., Ruano, F., 2007, “Inventário de las especies de formícidos de Sierra Nevada, Granada (España) (Hymenoptera, Formicidae).” or “Inventory of the ant species of Sierra Nevada, Granada (Spain) (Hymenoptera, Formicidae).” Zoologica Baetica, vol. 18, p. 49-68.

Tinaut, A., Ruano, F., 2021, “Biogeography of Iberian Ants (Hymenoptera: Formicidae).” Diversity, vol. 13, no. 2, article no. 88, p. 1-25.

Tinaut, A., Ruano, F., Martínez, M. D., 2005, “Biology, Distribution and Taxonomic Status of the Parasitic Ants of the Iberian Peninsula (Hymenoptera: Formicidae, Myrmicinae).” Sociobiology, vol. 46, no. 3, p. 449-489.

Wagner, H. C., Arthofer, W., Seifert, B., Muster, C., Steiner, F. M., Schlick-Steiner, B. C., 2017, “Light at the end of the tunnel: Integrative taxonomy delimits cryptic species in the Tetramorium caespitum complex (Hymenoptera: Formicidae).” Myrmecological News, vol. 25, p. 95–129.

Ward, P. S., Brady, S. G., Fisher, B. L., Schultz, T. R., 2015 (“2014”), “The evolution of myrmicine ants: Phylogeny and biogeography of a hyperdiverse ant clade (Hymenoptera: Formicidae).” Systematic Entomology, vol. 40, no. 1, p. 61-81. (Article first published online: 23 July 2014).

Ward, P. S., Brady, S. G., Fisher, B. L., Schultz, T. R., 2016, “Phylogenetic classifications are informative, stable, and pragmatic: the case for monophyletic taxa.” Insectes Sociaux, vol. 63, no. 4, p. 489-492.

Wegnez, P., Ignace, D., Lommelen, E., Hardy, M., Bogaert, J., Nilsson, C., 2015, “Redécouverte de Teleutomyrmex schneideri Kutter, 1950 dans les Alpes françaises (Hymenoptera: Formicidae).” Bulletin de la Société royale belge d’Entomologie/Bulletin van de Koninklijke Belgische Vereniging voor Entomologie, vol. 151, p. 52-57.

Wheeler, G. C., Wheeler, J., 1985, “A simplified conspectus of the Formicidae.” Transactions of the American Entomological Society, vol. 111, p. 255-264.

Wilson, E. O., 1963, “The Social Biology of Ants.” Annual Review of Entomology, vol. 8, no. 1, p. 345-368.

Wilson, E. O., 1963, “Social modifications related to rareness in ant species.” Evolution, vol. 17, p. 249-253.

Wilson, E. O., 1971, “The insect societies.” Cambridge, Mass., Harvard University Press, x + 548 pp.

Wilson, E. O., 1975, “Sociobiology. The new synthesis.” Cambridge, Mass., Harvard University Press, 697 pp.

Wilson, E. O., 1984, “Tropical social parasites in the ant genus Pheidole, with an analysis of the anatomical parasitic syndrome (Hymenoptera: Formicidae).” Insectes Sociaux, vol. 31, no. 3, p. 316-334.

107 references.

T1 Teleutomyrmex.

Teleutomyrmex [junior synonym of Tetramorium]

- Teleutomyrmex Kutter, 1950: 82. Type-species: Teleutomyrmex schneideri, by original designation.

- Teleutomyrmex junior synonym of Tetramorium: Ward et al., 2015: 76.

T2 schneideri.

schneideri. Teleutomyrmex schneideri Kutter, 1950a: 82, figs. 1-23 (q.m.) Switzerland.

- [Junior secondary homonym of Tetramorium schneideri Emery, 1898c: 145.]

- Combination in Tetramorium: Ward, et al. 2015: 76.

- Status as species: Stumper, 1951: 129; Brun, 1952: 73; Gösswald, 1953: 81; Bernard, 1967: 240 (redescription); Bolton, 1976: 309 (redescription); Kutter, 1977c: 167; Dlussky, Soyunov & Zabelin, 1990: 210; Bolton, 1995b: 403; Casevitz-Weulersse & Galkowski, 2009: 494; Borowiec, L. 2014: 170; Kiran, et al. 2017: 146.

- Replacement name: Tetramorium inquilinum Ward, et al. 2015: 76.

T3 inquilinum.

schneideri. Teleutomyrmex schneideri Kutter, 1950a: 82, figs. 1-23 (q.m.) Switzerland.

- [Junior secondary homonym of Tetramorium schneideri Emery, 1898c: 145.]

- Combination in Tetramorium: Ward, et al. 2015: 76.

- Status as species: Stumper, 1951: 129; Brun, 1952: 73; Gösswald, 1953: 81; Bernard, 1967: 240 (redescription); Bolton, 1976: 309 (redescription); Kutter, 1977c: 167; Dlussky, Soyunov & Zabelin, 1990: 210; Bolton, 1995b: 403; Casevitz-Weulersse & Galkowski, 2009: 494; Borowiec, L. 2014: 170; Kiran, et al. 2017: 146.

- Replacement name: Tetramorium inquilinum Ward, et al. 2015: 76.

inquilinum. Tetramorium inquilinum Ward, Brady, Fisher & Schultz, 2015: 76.

- Replacement name for schneideri Kutter, 1950a: 82. [Junior secondary homonym of Tetramorium schneideri Emery, 1898c: 145.]

T4 kutteri.

kutteri. Teleutomyrmex kutteri Tinaut, 1990b: 202, figs. 1-3, photos. 1-2 (q.m.) Spain.

- Combination in Tetramorium: Ward et al., 2015: 76.

- [Not in AntWiki: No replacement name for Tetramorium kutteri (Tinaut, 1990b, 202), junior secondary homonym of Tetramorium semilaeve kutteri Santschi, 1927b: 57.]

T5 seiferti.

seiferti. Teleutomyrmex seiferti Kiran & Karaman, in Kiran, et al. 2017: 148, figs. 3a, 4a, 5a, 6-8 (q.m.) Turkey.

- [Note: Kiran, et al. 2017: 146, retain the paraphyletic genus Teleutomyrmex.]

T6 buschingeri.

buschingeri. Teleutomyrmex buschingeri Lapeva-Gjonova, in Kiran, et al. 2017: 151, figs. 3b, 4b, 5b (q.) Bulgaria.

- [Note: Kiran, et al. 2017: 146, retain the paraphyletic genus Teleutomyrmex.]

T1 Teleutomyrmex.

- Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 226, Teleutomyrmex in Myrmicinae, Tetramoriini).

- Kutter, H. 1950a. Über eine neue, extrem parasitische Ameise. 1. Mitteilung. Mitt. Schweiz. Entomol. Ges. 23: 81-94 (page 82, Teleutomyrmex in Myrmicinae, Tetramoriini [subtribe Teleutomyrmini]).

- Sanetra, M.; Buschinger, A. 2000. Phylogenetic relationships among social parasites and their hosts in the ant tribe Tetramoriini (Hymenoptera: Formicidae). Eur. J. Entomol. 97: 95-117 (page 95, phylogeny).

- 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). Syst. Entomol. 40: 61-81 (page 76).

T2 schneideri.

- Brun, R., 1952. Das zentralnervensystem von Teleutomyrmex schneideri Kutt. (Hym. Formicid.). III. Mitteilung. Mitt. Schweiz. Entomol. Ges. 25: 73-86 (Not in AntWiki!).

- Buschinger, A. 1987: Teleutomyrmex schneideri KUTTER 1950 and other parasitic ants found in the Pyrenees (Hymenoptera, Formicidae). Spixiana 10, 81-83.

- Buschinger, A. 1995: Nicht am Ende: Die “Endameise” Teleutomyrmex schneideri. Ameisenschutz aktuell 9, 1-7.

- Buschinger, A. 1999: Wiederfund der sozialparasitischen Ameise Teleutomyrmex schneideri in der Schweiz. Mitt. Schweiz. Entomol. Ges. 72, 227-279. (Rediscovery in Switzerland after 50 years, at Simplon Pass).

- Buschinger, A. 2000: Die “Endameise” Teleutomyrmex schneideri in der Schweiz: Erster Wiederfund nach 50 Jahren. Ameisenschutz aktuell 14, 43-45.

- Buschinger, A. (2009) Social parasitism among ants: a review. (Hymenoptera: Formicidae). Myrmecological News 12: 219-235.

- Emery, C. 1898c. Beiträge zur Kenntniss der palaearktischen Ameisen. Öfvers. Fin. Vetensk.-Soc. Förh. 20: 124-151 (Not in AntWiki!).

- Gösswald, K. 1953. Histologische Untersuchungen an der arbeiterlosen Ameise Teleutomyrmex schneideri Kutter (Hym. Formicidae). Mitt. Schweiz. Entomol. Ges. 26: 81-128 (page 81).

- Hölldobler, B.; Wilson, E. O. 1990. The ants. Cambridge, Mass.: Harvard University Press, xii + 732 pp.

- Kiran, K., Karaman, C., Lapeva-Gjonova, A. & Aksoy, V. 2017. Two new species of the “ultimate” parasitic ant genus Teleutomyrmex Kutter, 1950 from the Western Palaearctic. Myrmecological News 25: 145-155.

- Kutter, H. 1950a. Über eine neue, extrem parasitische Ameise. 1. Mitteilung. Mitt. Schweiz. Entomol. Ges. 23: 81-94 (page 82, figs. 1-23 queen, male described).

- Kutter, H. 1977c. Hymenoptera, Formicidae. Insecta Helv. Fauna 6: 1-298 (page 167).

- Stumper, R. 1951. Teleutomyrmex schneideri Kutter. 2 Mitteilung. Über die Lebensweise der neuen Schmarotzerameise. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 24: 129-152.

- 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). Syst. Entomol. 40: 61-81 (page 76).

T3 inquilinum.

- Brun, R., 1952. Das zentralnervensystem von Teleutomyrmex schneideri Kutt. (Hym. Formicid.). III. Mitteilung. Mitt. Schweiz. Entomol. Ges. 25: 73-86 (Not in AntWiki!).

- Buschinger, A. 1987: Teleutomyrmex schneideri KUTTER 1950 and other parasitic ants found in the Pyrenees (Hymenoptera, Formicidae). Spixiana 10, 81-83.

- Buschinger, A. 1995: Nicht am Ende: Die “Endameise” Teleutomyrmex schneideri. Ameisenschutz aktuell 9, 1-7.

- Buschinger, A. 1999: Wiederfund der sozialparasitischen Ameise Teleutomyrmex schneideri in der Schweiz. Mitt. Schweiz. Entomol. Ges. 72, 227-279. (Rediscovery in Switzerland after 50 years, at Simplon Pass).

- Buschinger, A. 2000: Die “Endameise” Teleutomyrmex schneideri in der Schweiz: Erster Wiederfund nach 50 Jahren. Ameisenschutz aktuell 14, 43-45.

- Buschinger, A. (2009) Social parasitism among ants: a review. (Hymenoptera: Formicidae). Myrmecological News 12: 219-235.

- Emery, C. 1898c. Beiträge zur Kenntniss der palaearktischen Ameisen. Öfvers. Fin. Vetensk.-Soc. Förh. 20: 124-151 (Not in AntWiki!).

- Espadaler, X. & Cuesta, D. 2006. Teleutomyrmex schneideri Kutter, 1950 en España (Hymenoptera, Formicidae). Graellsia, 62(2): 261-262.

- Gösswald, K. 1953. Histologische Untersuchungen an der arbeiterlosen Ameise Teleutomyrmex schneideri Kutter (Hym. Formicidae). Mitt. Schweiz. Entomol. Ges. 26: 81-128 (page 81).

- Hölldobler, B.; Wilson, E. O. 1990. The ants. Cambridge, Mass.: Harvard University Press, xii + 732 pp.

- Kiran, K., Karaman, C., Lapeva-Gjonova, A. & Aksoy, V. 2017. Two new species of the “ultimate” parasitic ant genus Teleutomyrmex Kutter, 1950 from the Western Palaearctic. Myrmecological News 25: 145-155.

- Kutter, H. 1950a. Über eine neue, extrem parasitische Ameise. 1. Mitteilung. Mitt. Schweiz. Entomol. Ges. 23: 81-94 (page 82, figs. 1-23 queen, male described).

- Kutter, H. 1977c. Hymenoptera, Formicidae. Insecta Helv. Fauna 6: 1-298 (page 167).

- Stumper, R. 1951. Teleutomyrmex schneideri Kutter. 2 Mitteilung. Über die Lebensweise der neuen Schmarotzerameise. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 24: 129-152.

- 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). Syst. Entomol. 40: 61-81 (page 76).

- Wegnez, P., Ignace, D., Lommelen, E., Hardy, M., Bogaert, J. & Nilsson, C. 2015. Redécouverte de Teleutomyrmex schneideri Kutter, 1950 dans les Alpes françaises (Hymenoptera: Formicidae). Bulletin de la Société royale belge d’Entomologie, 151: 52-57.

T4 kutteri.

- Buschinger, A. (2009). Social parasitism among ants: a review. (Hymenoptera: Formicidae). Myrmecological News 12: 219-235.

- Kiran, K., Karaman, C., Lapeva-Gjonova, A. & Aksoy, V. 2017. Two new species of the “ultimate” parasitic ant genus Teleutomyrmex Kutter, 1950 from the Western Palaearctic. Myrmecological News 25: 145-155.

- Santschi, F. 1927b. A propos du Tetramorium caespitum L. Folia Myrmecol. Termit. 1: 52-58 (Not in AntWiki!).

- Tinaut, A. 1990b. Teleutomyrmex kutteri, spec. nov. A new species from Sierra Nevada (Granada, Spain). Spixiana 13: 201-208 (page 202, figs. 1-3, photos. 1-2 queen, male described).

- 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). Syst. Entomol. 40: 61-81 (page 76).

T5 seiferti.

- Kiran, K., Karaman, C., Lapeva-Gjonova, A. & Aksoy, V. 2017. Two new species of the “ultimate” parasitic ant genus Teleutomyrmex Kutter, 1950 from the Western Palaearctic. Myrmecological News 25: 145-155.

T6 buschingeri.

- Kiran, K., Karaman, C., Lapeva-Gjonova, A. & Aksoy, V. 2017. Two new species of the “ultimate” parasitic ant genus Teleutomyrmex Kutter, 1950 from the Western Palaearctic. Myrmecological News 25: 145-155.

Key to parasitic Tetramorium species.

This queen key is based on: Kiran, K., Karaman, C., Lapeva-Gjonova, A. & Aksoy, V. 2017. Two new species of the “ultimate” parasitic ant genus Teleutomyrmex Kutter, 1950 from the Western Palaearctic. Myrmecological News 25: 145-155.

Originally assigned their own genus, Teleutomyrmex, these ants prey on other Tetramorium species. The species Tetramorium atratulum is not included in this key and bears no resemblance to the other parasites in the genus.

Males of T. buschingeri have yet to be collected.

1

Gynes . . . . . 2

Males . . . . . 5

2

Carinae or teeth on dorsal surface of propodeum absent, dorsal profile of propodeum much shorter than the declivitous one. All lateral surfaces of mesosoma and petiole covered by a well-developed reticulate or alveolate microsculpture. Head length index CL / CW < 0.945. Southern Balkans . . . . . Tetramorium buschingeri.

Carinae or teeth on dorsal surface of propodeum present, dorsal profile of propodeum not much shorter than the declivitous one. Surfaces of lateral mesosoma and petiole only in patches covered by a reticulate or alveolate microsculpture or completely smooth. Head length index CL / CW > 0.945 . . . . . 3

3

Scape long, SL / CS > 1.00. Distance of frontal carinae clearly larger than petiolar width, DFC / PW > 1.096. Size small, CW < 464 μm. Scapes and tibiae with weaker, largely decumbent pilosity. Southern Iberia . . . . . Tetramorium kutteri.

Scape shorter, SL / CS < 1.00. Distance of frontal carinae not much larger than petiolar width, DFC/PW < 1.096. Size larger, CW > 464 μm. Scapes and tibiae with profuse erect or suberect pilosity . . . . . 4

4

Ratio of distance between lateral ocelli and large diameter of complex eye larger: DLO / EL 0.93 - 1.11. Katepisternum with many long decumbent hairs, posterior corners of head posterior of the eyes smooth, absolute scape length larger: SL > 457 μm. Anatolia . . . . . Tetramorium seiferti.

Ratio of distance between lateral ocelli and large diameter of complex eye smaller: DLO / EL 0.70 - 0.80. Katepisternum without or only with a few decumbent hairs, posterior corners of head posterior of the eyes densely microreticulate, absolute scape length smaller: SL < 457 μm. Alps and Pyrenees . . . . . Tetramorium inquilinum.

5

Anterior clypeal margin straight . . . . . 6

Anterior clypeal margin concave medially . . . . . Tetramorium seiferti.

6

Subgenital plate broadly convex, sagitta with sinusoidal shape . . . . . Tetramorium kutteri.

Subgenital plate slightly concave, sagitta broadly convex . . . . . Tetramorium inquilinum.

From: AntWiki - Where Ant Biologists Share Their Knowledge.

Hölldobler, B. K., Wilson, E. O., 1990, “The Ants.” Cambridge, Mass., Harvard University Press. Text used with permission of the authors.

From Chapter 12:

The “ultimate” social parasite.

There is no better way to begin a survey of the social symbioses than by considering the most extreme example known, that of the “ultimate” parasitic ant Teleutomyrmex schneideri. This remarkable species was discovered by Heinrich Kutter (1950a) at Saas-Fee, in an isolated valley of the Swiss Alps near Zermatt. Its behavior has been studied by Stumper (1950) and Kutter (1969), its neuroanatomy by Brun (1952), and its general anatomy and histology by Gösswald (1953). A second population has been reported from near Briançon in the French Alps by Collingwood (1956), a third in the French Pyrenees by Buschinger (1987c), and still others in the Spanish Sierra Nevada by Tinaut Ranera (1981). Appropriately, the name Teleutomyrmex means “final ant.”

The populations of Teleutomyrmex schneideri, like those of most workerless parasitic ant species (Wilson, 1963), are small and isolated. The Swiss population appears to be limited to the eastern slope of the Saas Valley, in juniper-Arctostaphylos woodland ranging from 1,800 to 2,300 m in elevation. The ground is covered by thick leaf litter and sprinkled with rocks of various sizes, providing, in short, an ideal environment for ants The ant fauna is of a typically boreal European complexion, comprising the following free-living species listed in the order of their abundance (Stumper, 1950): Formica fusca, Formica lugubris, Tetramorium caespitum, Leptothorax acervorum, Leptothorax tuberum, Camponotus ligniperda, Myrmica lobicornis, Myrmixa sulcinodis, Camponotus herculeanus, Formica sanguinea, Formica rufibarbis, Formica pressilabris, and Manica rubida. For some unexplained reason this little assemblage is extremely prone to social parasitism. Formica sanguinea is a facultative slavemaking species, preying on the other species of Formica. Doronomyrmex pacis, a workerless parasite living with Leptothorax acervorum, was discovered by Kutter as a genus new to science in the Saas-Fee forest in 1945. In addition, Kutter and Stumper found Epimyrma stumperi in nests of Leptothorax tuberum, as well as two parasitic Leptothorax, goesswaldi and kutteri, in nests of Leptothorax acervorum (Kutter, 1969).

Teleutomyrmex schneideri is a parasite of Tetramorium caespitum and Tetramorium impurum. Like so many other social parasites, it is phylogenetically closer to its host than to any of the other members of the ant fauna to which it belongs. In fact, it may have been derived directly from a temporarily free-living offshoot of this species, since Tetramorium caespitum and Tetramorium impurum (the host species at Briançon and in the Pyrenees) are the only nonparasitic tetramoriines known to exist at the present time through most of central Europe. It is difficult to conceive of a stage of social parasitism more advanced than that actually reached by Teleutomyrmex schneideri. The species occurs only in the nests of its hosts. It lacks a worker caste, and the queens contribute in no visibly productive way to the economy of the host colonies. The queens are tiny compared with most ants, especially other tetramoriines; they average only about 2.5 mm in total length. They are unique among all known social insects in being ectoparasitic. In other words, they spend much of their time riding on the backs of their hosts (Figure 12-1). The Teleutomyrmex queens display several striking morphological features that are correlated with this peculiar habit. The ventral surface of the gaster (the large terminal part of the body) is strongly concave, permitting the parasites to press their bodies close to those of their hosts. The tarsal claws and arolia are unusually large, permitting the parasites to secure a strong grip on the smooth chitinous body surface of the hosts. The queens have a marked tendency to grasp objects. Given a choice, they will position themselves on the top of the body of the host queen, either on the thorax or the abdomen. Deprived of the nest queen, they will then seize a virgin Tetramorium queen, or a worker, or a pupa, or even a dead queen or worker. Stumper observed a case in which six to eight Teleutomyrmex queens simultaneously grasped one Tetramorium queen, completely immobilizing her. The mode of feeding of the Teleutomyrmex is not known with certainty. The adults are evidently either fed by the host workers through direct regurgitation or else share in the liquid regurgitated to the host queen. In any case, they are almost completely inactive most of the time. The Teleutomyrmex adults, especially the older queens, are highly attractive to the host workers, who lick them frequently. According to Gösswald, large numbers of unicellular glands are located just under the cuticle of the thorax, pedicel, and abdomen of the queens; these are associated with glandular hairs and are believed to be the source of a special attractant for the host workers. The abdomens of older Teleutomyrmex queens become swollen with fat body and ovarioles, as is shown in Figure 12-1. This physogastry is made possible by the fact that the intersegmental membranes are thicker and more sclerotized than is usually the case in ant queens and can therefore be stretched more. Also, the abdominal sclerites themselves are widely overlapping in the virgin queen, so that the abdomen can be distended to an unusual degree before the sclerites are pulled apart. The ovarioles increase enormously in length, discard their initial orientation, and infiltrate the entire abdomen and even the postpetiolar cavity.

From one to several physogastric queens are found in each parasitized nest, usually riding on the back of the host queen. Each lays an average of one egg every thirty seconds. The infested Tetramorium colonies are typically smaller than uninfested ones, but they still contain up to several thousand workers. The Tetramorium queens also lay eggs, and these are capable of developing into either workers or sexual forms (Buschinger, personal communication). Consequently the brood of a parasitized colony consists typically of eggs, larvae, and pupae of Teleutomyrmex queens and males mixed with those of Tetramorium workers.

The bodies of the Teleutomyrmex queens bear the mark of extensive morphological degeneration correlated with their loss of social functions. The labial and postpharyngeal glands are reduced, and the maxillary and metapleural glands are completely absent. The mandibular glands, on the other hand, are apparently normal. In addition, the queens possess a tibial gland, the function of which is unknown. The integument is thin and less pigmented and sculptured in comparison with that of Tetramorium; as a result of these reductions the queens are shining brown, an appearance that contrasts with the opaque blackish brown of their hosts. The sting and poison apparatus are reduced; the mandibles are so degenerate that the parasites are probably unable to secure food on their own; the tibial-tarsal cleaning apparatus is underdeveloped; and, of even greater interest, the brain is reduced in size with visible degeneration in the associative centers. In the central nerve cord, ganglia 9-13 are fused into a single piece. The males are also degenerate. Their bodies, like those of the males of a few other extreme social parasites, are “pupoid,” meaning that the cuticle is thin and depigmented, actually greyish in color; the petiole and postpetiole are thick and provided with broad articulating surfaces; and the abdomen is soft and deflected downward at the tip.

In its essentials the life cycle of Teleutomyrmex schneideri resembles that of other known extreme ant parasites. Mating takes place within the host nest. The fecundated queens then either shed their wings and join the small force of egg layers within the home nest or else fly out in search of new Tetramorium nests to infest. Stumper found that the queens could be transferred readily from one Tetramorium colony to another, provided the recipient colony originated from the Saas-Fee. However, Tetramorium colonies from Luxembourg were hostile to the little parasites. Less surprisingly, ant species from the Saas-Fee other than Tetramorium caespitum always rejected the Teleutomyrmex. However, Buschinger (personal communication) has pointed out that the Saas-Fee population could be caespitum or impurum, or a mixture of both. In other words, the transfer might have been attempted across species.

From: AntWiki - Where Ant Biologists Share Their Knowledge.

Hölldobler, B. K., Wilson, E. O., 1990, “The Ants.” Cambridge, Mass., Harvard University Press. Text used with permission of the authors.

When do we know an ant is an extreme, workerless inquiline?

From Chapter 12:

1. The worker caste is lost.

2. The queen is either replaced by an ergatogyne, or ergatogynes appear together with a continuous series of intergrades connecting them morphologically to the queens.

3. There is a tendency for multiple egg-laying queens to coexist in the same host nest.

4. The queen and male are reduced in size, often dramatically so; in some cases (for example, Teleutomyrmex schneideri, Plagiolepis ampeloni, Plagiolepis xene) the queen is actually smaller than the host worker.

5. The male becomes “pupoid”: its body is thickened, the petiole and postpetiole become much more broadly attached, the genitalia are more externally exposed when not in use, the cuticle becomes thin and depigmented, and the wings are reduced or lost. The extreme examples of this trend are displayed by Anergates atratulus, Pheidole neokohli, and Pheidole acutidens (see Figures 12-19 and 12-20).

6. There is a tendency for the nuptial flights to be curtailed, and to be replaced by mating activity among nestmates (“adelphogamy”) within or near the host nest. Dispersal of the queen afterward is very limited.

7. Probably as a consequence of the curtailment of the nuptial flight just cited, the populations of inquiline species are usually very fragmented and limited in their geographic distribution.

8. The wing venation is reduced.

9. Mouthparts are reduced, with the mandibles becoming smaller and toothless and the palps losing segments. Concomitantly, the inquilines lose the ability to feed themselves and must be sustained by liquid food regurgitated to them by the host workers.

10. Antennal segments are fused and reduced in number.

11. The occiput, or rear portion of the head, of the queen is narrowed.

12. The central nervous system is reduced in size and complexity, usually through reduction of associative centers.

13. The petiole and postpetiole are thickened, especially the latter, and the postpetiole acquires a broader attachment to the gaster.

14. A spine is formed on the lower surface of the postpetiole (the Parasitendorn of Kutter).

15. The propodeal spines (if present in the ancestral species) “melt,” that is, they thicken and often grow shorter, and their tips are blunted.

16. The cuticular sculpturing is reduced or lost altogether over most of the body; in extreme cases the body surface becomes strongly shining.

17. The exoskeleton becomes thinner and less pigmented.

18. Many of the exocrine glands are reduced or lost, a trait already described in some detail in the earlier account of Teleutomyrmex schneideri.

19. The queens become highly attractive to the host workers, which lick them frequently. This is especially true of the older, physogastric individuals, and it appears to be due to the secretion of special attractant substances which are as yet chemically unidentified.

Sometimes, even general names change or are different than intended. Just like Teleutomyrmex.

First part means: das Ende, die Vollendung.

Last part means: die Ameise.

Since Brun, 1951-1952, everybody in the German hemisphere calls the ant: die Endameise.

Kutter wanted to call it, and did so in his 1950 article: die Grenzameise.

In English it always was the final ant...

Extreme, Workerless Inquiline ants.

Ants: Probably one of the most important animal-groups on Earth. Together with wasps and bees they play some of the most important roles in ecosystems around the world. Without these three groups of insects, most animal- and plant-species will become extinct on this little globe in space in the near future. So remember them well: Ants - Formicidae, Wasps - Vespidae and Bees (including digger-wasps) – Apidae. And if you want to know much about them, be prepared to read a lot! Almost every conceivably down-to-earth lifestyle you can think of, somewhere one or other ant lives like that, and for the airborne lifestyles, go to the wasps and bees! Have a lifetime of fun to discover all these animals! And for the most specialized animals on earth, you must go to the ants and discover the species in the genera Teleutomyrmex and Anergates and a few other, mostly related, species!

The extreme, workerless inquilines.

The genus Teleutomyrmex was described in 1950 by Heinrich Kutter, based on ants discovered in Saas-Fee (a small town in the canton Wallis, Switzerland.) in 1949 and 1950. The species was named T. schneideri. Later, the species was also found in the French Alps and Pyrenees, the Spanish Pyrenees and Cantabria and in a nearby place in the Swiss Alps. The type-meadow in Saas-Fee was destroyed between 1950 and 1971. A second species, T. kutteri, was described in 1990 by Alberto Tinaut based on animals from the Sierra Nevada, Spain and the third and forth species, from 2017, are from Bulgaria and Turkey, respectively T. buschingeri Lapeva-Gjonova, in Kiran, et al. 2017 and T. seiferti Kiran & Karaman, in Kiran, et al. 2017. The new species of Farab, Turkmenistan, is still not described.

The ants from the genera Teleutomyrmex and Anergates are the most specialized ants on Earth. Teleutomyrmex are extreme, workerless inquilines that became one more thing than Anergates, nl. ectoparasites. Let me explain.

- Parasite: An animal that is dependent on another species to survive. This dependence is temporary (during a certain period of its life.) or permanent.

- Social parasites: Social animals (like ants!) that are dependent on other social animals to survive. This can be temporary (during colony-foundation.) or permanent.

- Inquilines: Permanent social parasites among ants are also called inquilines.

- Workerless inquilines: The worker-caste, not needed by the inquilines, has disappeared. Only females/queens and males exist.

- Extreme, workerless inquilines: The females/queens and males have undergone some important morphological changes. Through these changes the ants are more adapted to their specialized way of life but, at the same time, they make sure that the ants can’t survive without their host. Some examples are: reduction of the mouthparts, development of appeasement-glands, over-development of the reproduction-organs, becoming weak and “soft”, males that become pupoid (show characteristic modifications that makes the male look “like a pupa”, e.g. yellowish color, downward curved gaster, big external genital plates,...),…

- Ectoparasites: Parasites that need to be carried around by their hosts. They are not capable to or have great difficulty with walking very short distances.

Edward Osborne Wilson, in 1971, wrote down a list with almost all the characters that determine social parasites and, in 1990, completed the list together with Berthold K. Hölldobler. In it 41 characters are listed that extreme, workerless inquilines can have. Not all those inquilines have all the characters but they have most of them. The ants of the genus Teleutomyrmex display 36 characters of the list but have also a few adaptations that are special for their ectoparasitic lifestyle.

Only around fourteen ant species are known that are extreme, workerless inquilines and four/five of them, the Teleutomyrmex species, have become ectoparasites. Teleutomyrmex females/queens have, for example, unique morphological adaptations like the dorsoventrally compressed gaster that easily fit around the gaster of the host-queen. Also, the queens and males of Teleutomyrmex-species have the terminal tarsal segments of their legs adapted/modified to be able to grip firmly the body of queens and workers of the host species and are so almost completely unable to walk alone.

One remarkable fact is that the males of Teleutomyrmex still have rudimentary, unusable wings while the males of the other extreme, workerless inquilines have lost the wings completely. In all its other characteristics it is further evolved compared with the other species. Strange but true!

A peculiar taxonomic fact: The genera Anergates and Teleutomyrmex and the host-genus Tetramorium are closely related ant-genera belonging to the tribe (group of closely related genera.) Tetramoriini (now the Crematogastrini). This tribe is part of the subfamily Myrmicinae which also includes the genus Pheidole (in the tribe Pheidolini, now the Attini!). Most, or all, of the known extreme, workerless inquilines belong to this subfamily…

Some taxonomic problems in this group of specialized ants.

In 1950, William Steel Creighton placed Anergates friedlandi as a synonym of A. atratulus. Although he recognized certain morphological differences between the two species in 1934 when he described A. friedlandi, he based his 1950 decision on the speculation of William L. Brown Jr. that the host-species of both inquilines was the same (yes) and that the North American population wasn’t native to that continent (maybe). So, no morphological data but host-species distribution was used to establish the synonymy. A few myrmecologists still questioned the decision… Now it is clear that the North American form is an introduced population of A. atratulus!

The last few years some myrmecologists (like Alfred Buschinger) think that the morphological differences between Teleutomyrmex schneideri and T. kutteri are very minimal and question if T. kutteri should be placed as a synonym of T. schneideri. Most still think both deserve species status (clear morphological differences between the queens and the males of both species!) and both names stay as species names on record. Now, with the description of two more species, things are getting clearer in this genus.

Tetramorium and related parasitic genera underwent a very thorough genetic phylogenetic study by Matthias Sanetra and Alfred Buschinger in 2000. When you read the paper only two possibilities exists. The first one: Tetramorium, Anergates and Teleutomyrmex should be considered to be synonyms of Strongylognathus and all the species together form one big genus (as Ward et al., 2015 ("2014"), say!). The other one: Tetramorium should be divided in at least eight different genera, all standing together with the parasitic ones in one compact tribe. The authors of the article still can’t follow either of the possibilities. Who will take a decision?

First descriptions of, and last revisions of the species complexes of the senior homonyms of Teleutomyrmex species when they are placed in the genus Tetramorium.

- Tetramorium schneideri Emery, 1898, valid.

Tetramorium schneideri Emery, 1898c: 145 (w.) KAZAKHSTAN. Palearctic.

Tarbinsky, 1976: 115 (q.); Radchenko & Scupola, 2015: 234 (m.).

Subspecies of Tetramorium striativentre: Ruzsky, 1905a: 518; Ruzsky, 1905b: 767; Emery, 1909f: 706; Karavaiev, 1911a: 55; Karavaiev, 1912b: 585; Kuznetsov-Ugamsky, 1923b: 253; Emery, 1924f: 278; Pisarski, 1967a: 403; Tarbinsky, 1976: 115 (redescription).

Status as species: Ruzsky, 1905b: 517; Dlussky, 1981b: 17; Dlussky & Zabelin, 1985: 232; Dlussky et al., 1990: 207; Radchenko, 1992a: 45 (in key); Radchenko, 1992b: 52; Bolton, 1995b: 414; Dietrich, 2004: 322; Schultz et al., 2006: 203; Paknia et al., 2010: 35; Guénard & Dunn, 2012: 57; Borowiec, 2014: 203; Radchenko & Scupola, 2015: 232 (redescription).

Material of the unavailable name Tetramorium striativentre schneideri longispina referred here by Dlussky et al., 1990: 207; Radchenko, 1992b: 52.

Tetramorium striativentre schneideri longispina Karavaiev, 1912, unavailable.

Tetramorium striativentre subsp. schneideri var. longispina Karavaiev, 1912b: 585 (w.) TRANSCASPIA. Palearctic.

Unavailable name; material referred to Tetramorium schneideri by Dlussky et al., 1990: 207; Radchenko, 1992b: 52.

- Dlussky, G. M., Soyunov, O. S., Zabelin, S. I., 1990 [“1989”], “Muravji Turkmenistana.”, or “[Ants of Turkmenistan.]” [In Russian.]. Ashkabad, Ylym Press, 275 pp.

- Emery, C., 1898c, “Beiträge zur Kenntniss der palaearktischen Ameisen.” Öfvers. Fin. Vetensk.-Soc. Förh. (= Öfversigt af Finska Vetenskaps-Societetens Förhandlingar (Helsinki)), vol. 20, p. 124-151.

- Karavaiev, V., 1912b, “Ameisen aus dem paläarktischen Faunengebiete.” Russkoe Entomologicheskoe Obozrenie, vol. 12, p. 581-596.

- Radchenko, A. G., 1992b, “[Ants of the genus Tetramorium (Hymenoptera, Formicidae) of the USSR fauna. Report 2.]” [In Russian.]. Zoologicheskii Zhurnal, vol. 71, no. 8, p. 50-58.

- Radchenko, A. G., Scupola, A., 2015, “Taxonomic revision of the striativentre species group of the genus Tetramorium (Hymenoptera, Formicidae).” Vestnik Zoologii, vol. 49, no. 3, p. 219-244.

- Tarbinsky, Y. S., 1976, “[The ants of Kirghizia.]” [In Russian.]. Frunze, Ilim, 217 pp.

- Tetramorium semilaeve kutteri Santschi, 1927, junior synonym of current valid taxon Tetramorium indocile Santschi, 1927.

Tetramorium semilaeve var. kutteri Santschi, 1927b: 57 (w.) SWITZERLAND. Palearctic.

Primary type information: Brig, Switzerland

Subspecies of Tetramorium semilaeve: Novák & Sadil, 1941: 85 (in key); Bolton, 1995b: 410.

Junior synonym of Tetramorium indocile: Wagner et al., 2017: 116.

- Santschi, F., 1927b, “A propos du Tetramorium caespitum L.” Folia Myrmecol. Termit., vol. 1, p. 52-58.

- Wagner, H. C., Arthofer, W., Seifert, B., Muster, C., Steiner, F. M., Schlick-Steiner, B. C., 2017, “Light at the end of the tunnel: Integrative taxonomy delimits cryptic species in the Tetramorium caespitum complex (Hymenoptera: Formicidae).” Myrmecological News, vol. 25, p. 95–129.

A1 Anergates.

Anergates [junior synonym of Tetramorium]

- Anergates Forel, 1874: 67 (see also p. 93). Type-species: Myrmica atratula, by monotypy.

- Anergates junior synonym of Tetramorium: Ward et al., 2015: 76.

A2 atratulum.

atratulum. Myrmica atratula Schenck, 1852: 91 (q.m.) Germany.

- [Also described as new by Schenck, 1853: 186.]

- Wheeler, W.M. 1909g: 182 (l.).

- Combination in Tetramorium: Mayr, 1855: 429.

- Combination in Tomognathus: Mayr, 1863. [obsolete combination.].

- Combination in Anergates: Forel, 1874: 68 (see also p. 93).

- Combination in Tetramorium: Ward et al., 2015: 76.

- Senior synonym of friedlandi: Creighton, 1950a: 243.

- See also: Donisthorpe, 1915d: 89; Boven, 1977: 81; Wheeler, G.C. & Wheeler, J. 1955c: 128; Atanassov & Dlussky, 1992: 161.

friedlandi. Anergates friedlandi Creighton, 1934: 193 (q.) U.S.A.

- Junior synonym of atratulum: Creighton, 1950a: 243.

A3 friedlandi.

friedlandi. Anergates friedlandi Creighton, 1934: 193 (q.) U.S.A.

- Junior synonym of atratulus: Creighton, 1950a: 243.

A1 Anergates.

- Bolton, B. 1976. The ant tribe Tetramoriini (Hymenoptera: Formicidae). Constituent genera, review of smaller genera and revision of Triglyphothrix Forel. Bull. Br. Mus. (Nat. Hist.) Entomol. 34: 281-379 (page 296, Anergates in Myrmicinae, Tetramoriini).

- Bolton, B. 1994. Identification guide to the ant genera of the world. Cambridge, Mass.: Harvard University Press, 222 pp. (page 106, Anergates in Myrmicinae, Tetramoriini).

- Bolton, B. 2003. Synopsis and Classification of Formicidae. Mem. Am. Entomol. Inst. 71: 370pp (page 224, Anergates in Myrmicinae, Tetramoriini).

- Dalla Torre, K. W. von. 1893. Catalogus Hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol. 7. Formicidae (Heterogyna). Leipzig: W. Engelmann, 289 pp. (page 64, Anergates in Myrmicinae).

- Dlussky, G. M.; Fedoseeva, E. B. 1988. Origin and early stages of evolution in ants. Pp. 70-144 in: Ponomarenko, A. G. (ed.) Cretaceous biocenotic crisis and insect evolution. Moskva: Nauka, 232 pp. (page 80, Anergates in Myrmicinae, Tetramoriini).

- Emery, C. 1877b. Saggio di un ordinamento naturale dei Mirmicidei, e considerazioni sulla filogenesi delle formiche. Bull. Soc. Entomol. Ital. 9: 67-83 (page 81, Anergates in Myrmicinae [Myrmicidae]).

- Emery, C. 1895l. Die Gattung Dorylus Fab. und die systematische Eintheilung der Formiciden. Zool. Jahrb. Abt. Syst. Geogr. Biol. Tiere 8: 685-778 (page 769, Anergates in Myrmicinae, Myrmicini).

- Emery, C. 1914e. Intorno alla classificazione dei Myrmicinae. Rend. Sess. R. Accad. Sci. Ist. Bologna Cl. Sci. Fis. (n.s.) 18: 29-42 (page 41, Anergates in Myrmicinae, Solenopsidini [subtribe Anergatini]).

- Emery, C. 1922c. Hymenoptera. Fam. Formicidae. Subfam. Myrmicinae. [part]. Genera Insectorum 174B: 95-206 (page 205, Anergates in Myrmicinae, Solenopsidini).

- Emery, C.; Forel, A. 1879. Catalogue des Formicides d'Europe. Mitt. Schweiz. Entomol. Ges. 5: 441-481 (page 457, Anergates in Myrmicinae [Myrmicidae]).

- Ettershank, G. 1966. A generic revision of the world Myrmicinae related to Solenopsis and Pheidologeton (Hymenoptera: Formicidae). Aust. J. Zool. 14: 73-171 (page 81, Anergates incertae sedis in Myrmicinae).

- Fisher and Cover. 2007. Ants of North America. A guide to the Genera. University of California Press.

- Forel, A. 1874. Les fourmis de la Suisse. Systématique, notices anatomiques et physiologiques, architecture, distribution géographique, nouvelles expériences et observations de moeurs. Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 26: 1-452 (page 67, Anergates as genus).

- Forel, A. 1893b. Sur la classification de la famille des Formicides, avec remarques synonymiques. Ann. Soc. Entomol. Belg. 37: 161-167 (page 165, Anergates in Myrmicinae, Formicoxenini).

- Forel, A. 1917. Cadre synoptique actuel de la faune universelle des fourmis. Bull. Soc. Vaudoise Sci. Nat. 51: 229-253 (page 243, Anergates in Myrmicinae, Solenopsidini).

- Heinze,J., B. Lautenschläger & A. Buschinger 2007. Female-biased sex ratios and unusually potent males in the social parasite Anergates atratulus (Hymenoptera: Formicidae). Myrmecological News 10, 1-5.

- Sanetra, M.; Buschinger, A. 2000. Phylogenetic relationships among social parasites and their hosts in the ant tribe Tetramoriini (Hymenoptera: Formicidae). Eur. J. Entomol. 97: 95-117 (page 108, Anergates in Myrmicinae, Tetramoriini).

- Smith, D. R. 1979. Superfamily Formicoidea. Pp. 1323-1467 in: Krombein, K. V., Hurd, P. D., Smith, D. R., Burks, B. D. (eds.) Catalog of Hymenoptera in America north of Mexico. Volume 2. Apocrita (Aculeata). Washington, D.C.: Smithsonian Institution Pr (page 1401, Anergates in Myrmicinae, Tetramoriini).

- 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). Syst. Entomol. 40: 61-81 (page 76).

- Wheeler, G. C.; Wheeler, J. 1985b. A simplified conspectus of the Formicidae. Trans. Am. Entomol. Soc. 111: 255-264 (page 257, (anachronism)).

- Wheeler, W. M. 1910b. Ants: their structure, development and behavior. New York: Columbia University Press, xxv + 663 pp. (page 139, Anergates in Myrmicinae, Myrmicini).

- Wheeler, W. M. 1922i. Ants of the American Museum Congo expedition. A contribution to the myrmecology of Africa. VII. Keys to the genera and subgenera of ants. Bull. Am. Mus. Nat. Hist. 45: 631-710 (page 663, Anergates in Myrmicinae, Solenopsidini).

A2 atratulum.

- Atanassov, N.; Dlussky, G. M. 1992. Fauna of Bulgaria. Hymenoptera, Formicidae. Fauna Bûlg. 22: 1-310 (page 161).

- Adlerz, G. 1886. Myrmecologiska studier. II. Svenska myror och deras lefnadsförhållanden. Bihang till Kongliga Svenska Vetenskaps-Akademiens Handlingar. 11(18):1-329.

- Adlerz, G. 1908. Zwei Gynandromorphen von Anergates atratulus Schenck. Ark. Zool. 5(2 2: 1-6).

- Boven, J. K. A. van. 1977. De mierenfauna van België (Hymenoptera: Formicidae). Acta Zool. Pathol. Antverp. 67: 1-191 (page 81).

- Buschinger, A. (2009) Social parasitism among ants: a review. (Hymenoptera: Formicidae). Myrmecological News 12: 219-235.

- Buschinger, A., B. C. Schlick-Steiner, F. M. Steiner, and M. Sanetra. 2003. Anergates atratulus, eine ungewöhnlich seltene Parasiten-Ameise. Ameisenschutz Aktuell. 17:1-6.

- Collingwood, C. A. 1979. The Formicidae (Hymenoptera) of Fennoscandia and Denmark. Fauna Entomol. Scand. 8:1-174.

- Creighton, W. S. 1934. Descriptions of three new North American ants with certain ecological observations on previously described forms. Psyche (Camb.) 41: 185-200.

- Creighton, W. S. 1950a. The ants of North America. Bull. Mus. Comp. Zool. 104: 1-585 (page 243, Senior synonym of freidlandi).

- Crawley, W. C. 1912. Anergates atratulus, Schenk., a British ant, and the acceptance of a queen by Tetramorium caespitum, L. Entomologist's Record and Journal of Variation. 24:218-219.

- Dash, S.T. & Sanchez, L. 2009. New distribution record for the social parasitic ant Anergates atratulus (Schenck, 1852) (Hymenoptera: Formicidae): An IUCN red-listed species. Western North American Naturalist 69: 140-141.

- Donisthorpe, H. 1915f. British ants, their life-history and classification. Plymouth: Brendon & Son Ltd., xv + 379 pp. (page 89).

- Fisher and Cover. 2007. Ants of North America. A guide to the Genera. University of California Press.

- Forel, A. 1874. Les fourmis de la Suisse. Systématique, notices anatomiques et physiologiques, architecture, distribution géographique, nouvelles expériences et observations de moeurs. Neue Denkschr. Allg. Schweiz. Ges. Gesammten Naturwiss. 26: 1-452 (page 68, Combination in Anergates).

- Heinze, J., B. Lautenschläger, and A. Buschinger. 2007. Female-biased sex ratios and unusually potent males in the social parasite Anergates atratulus (Hymenoptera: Formicidae). Myrmecological News. 10:1-5.

- Lapeva-Gjonova, A.; Kiran, K.; Aksoy V. 2012. Unusual ant hosts of the socially parasitic ant Anergates atratulus (Schenck, 1852) (Hymenoptera, Formicidae). Psyche doi:10.1155/2012/391525.

- Mayr, G. 1855. Formicina austriaca. Beschreibung der bisher im österreichischen Kaiserstaate aufgefundenen Ameisen, nebst Hinzufügung jener in Deutschland, in der Schweiz und in Italien vorkommenden Arten. Verh. Zool.-Bot. Ver. Wien 5: 273-478 (page 429, Combination in Tetramorium).

- Mayr, G., 1861, “Die europäischen Formiciden. Nach der analytischen Methode bearbeitet.” Wien, C. Gerold’s Sohn, 80 pp.

- Mayr, G., 1863, “Formicidarum index synonymicus.” Verhandlungen der Kaiserlich-Königlichen Zoologisch-Botanischen Gesellschaft in Wien, vol. 13, p. 385-460.

- Schenck, C. F. 1852. Beschreibung nassauischer Ameisenarten. Jahrb. Ver. Naturkd. Herzogthum Nassau Wiesb. 8: 1-149 (page 91, queen, male described).

- Schenck, C. F. 1853b. Die nassauischen Ameisen-Species. (Fortsetzung.). Stett. Entomol. Ztg. 14: 185-198 (page 186, also described as new).

- 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). Syst. Entomol. 40: 61-81 (page 76).

- Wheeler, G. C.; Wheeler, J. 1955c. The ant larvae of the myrmicine tribe Solenopsidini. Am. Midl. Nat. 54: 119-141 (page 128).

- Wheeler, W. M. 1908. Comparative ethology of the European and North American ants. J. Psychol. Neurol. 13:404-435, pl. III-IV.

- Wheeler, W. M. 1909g. Observations on some European ants. J. N. Y. Entomol. Soc. 17: 172-187 (page 182, larva described).

A3 friedlandi.

- Bolton, B. 1995b. A new general catalogue of the ants of the world. Cambridge, Mass.: Harvard University Press, 504 pp. (page 63, catalogue).

- Creighton, W. S. 1934. Descriptions of three new North American ants with certain ecological observations on previously described forms. Psyche (Camb.) 41: 185-200.

- Creighton, W. S. 1950a. The ants of North America. Bull. Mus. Comp. Zool. 104: 1-585 (page 243, Junior synonym of atratulus).

A new article, not much about Teleutomyrmex but it is mentioned as an endemic...

Biogeography of Iberian Ants (Hymenoptera: Formicidae).

by Alberto Tinaut and Francisca Ruano.

Diversity 2021, 13(2), 88, p. 1-25.

Published: 19 February 2021.

https://www.mdpi.com/1424-2818/13/2/88/htm

or

https://www.mdpi.com/1424-2818/13/2/88/pdf with two supplements.

or the blog "Biogeography of Iberian ants":

https://blog.myrmecologicalnew…eography-of-iberian-ants/

100 references!!!

The other extreme, workerless inquilines…

Tetramorium Mayr, 1855.

Tetramorium [Myrmicinae: Tetramoriini].

- Tetramorium Mayr, 1855: 423. Type-species: Formica caespitum, by subsequent designation of Girard, 1879: 1016.

Tetramorium microgyna Santschi, 1918b.

microgyna. Tetramorium microgyna Santschi, 1918b: 132 (q., not w.) South Afrika.

- See also: Arnold, 1926: 253; Santschi, 1928f: 207; Bolton, 1980: 329.

Tetramorium parasiticum Bolton, 1980.

parasiticum. Tetramorium parasiticum Bolton, 1980: 281, figs. 65, 66 (q.) South Africa.

Pheidole Westwood, 1839.

Pheidole [Myrmicinae: Pheidolini].

- Pheidole Westwood, 1839: 219. Type-species: Atta providens, by monotypy.

Anergatides [junior synonym of Pheidole].

- Anergatides Wasmann, 1915b: 281. Type-species: Anergatides kohli (junior secondary homonym in Pheidole, replaced by Pheidole neokohli), by monotypy.

- Anergatides junior synonym of Pheidole: Smith, D.R. 1979: 1366; Wilson, 1984: 328.

Bruchomyrma [junior synonym of Pheidole].

- Bruchomyrma Santschi, 1922d: 248. Type-species: Bruchomyrma acutidens, by monotypy.

- Bruchomyrma junior synonym of Pheidole: Smith, D.R. 1979: 1366; Wilson, 1984: 327.

- [Bruchomyrma listed as genus: Dlussky & Fedoseeva, 1988: 80 (anachronism)].

- Bruchomyrma junior synonym of Pheidole: Bolton, 1994: 106; Wilson, 2003: 6.

Gallardomyrma [junior synonym of Pheidole].

- Gallardomyrma Bruch, 1932: 271. Type-species: Gallardomyrma argentina, by original designation.

- Gallardomyrma junior synonym of Pheidole: Smith, D.R. 1979: 1366; Wilson, 1984: 327.

- [Gallardomyrma listed as genus: Dlussky & Fedoseeva, 1988: 80 (anachronism)].

- Gallardomyrma junior synonym of Pheidole: Bolton, 1994: 106; Wilson, 2003: 6.

Pheidole neokohli Wilson, 1984 (= Pheidole kohli (Wasmann, 1915))

(= Anergatides kohli Wasmann, 1915)

(not Pheidole kohli Mayr, 1901).

kohli. Anergatides kohli Wasmann, 1915b: 283, pls. 7, 8 (q.m.) Democratic Republic Of Congo.

[Junior secondary homonym of kohli Mayr, 1901b; 11 (footnote).] Replacement name: Pheidole neokohli Wilson, 1984: 328. Wheeler, G.C. & Wheeler, J. 1955c: 130 (l.).

neokohli. Pheidole neokohli Wilson, 1984: 328. Replacement name for Anergatides kohli Wasmann, 1915b: 283. [Junior secondary homonym of Pheidole kohli Mayr, 1901b: 11 (footnote).]

Pheidole acutidens (Santschi, 1922) (= Bruchomyrma acutidens Santschi, 1922).

acutidens. Bruchomyrma acutidens Santschi, 1922d: 249, figs. A-D (q.) Argentina. Bruch, 1931: 31 (m.). Combination in Pheidole: Wilson, 1984: 327. See also: Wheeler, W.M. 1937c: 52; Wilson, 2003: 260.

Pheidole argentina (Bruch, 1932) (= Gallardomyrma argentina Bruch, 1932).

argentina. Gallardomyrma argentina Bruch, 1932: 273, figs. 1-8 (q.) Argentina. Combination in Pheidole: Wilson, 1984: 327. See also: Wilson, 2003: 267.

Pheidole parasitica Wilson, 1984.

parasitica. Pheidole parasitica Wilson, 1984: 321, fig. 3 (q.) India.

Excluded from the extreme, workerless inquilines

Eriopheidole [junior synonym of Pheidole].

- Eriopheidole Kusnezov, 1952a: 10. Type-species: Eriopheidole symbiotica (junior secondary homonym in Pheidole, replaced by Pheidole kusnezovi), by monotypy.

- Eriopheidole junior synonym of Pheidole: Smith, D.R. 1979: 1366; Bolton, 1994: 106.

Pheidole kusnezovi Wilson, 2003 (= Pheidole symbiotica (Kusnezov, 1952))

(= Eriopheidole symbiotica Kusnezov, 1952)

(not Pheidole symbiotica Wasmann, 1909).

symbiotica. Eriopheidole symbiotica Kusnezov, 1952a: 16, figs. 1-9 (q.m.) Argentina.

[Junior secondary homonym of Pheidole symbiotica Wasmann, 1909: 693 (footnoye).] Combination in Pheidole: Wilson, 1984: 328. Replacement name: Pheidole kusnezovi Wilson, 2003: 311.

kusnezovi. Pheidole kusnezovi Wilson, 2003: 311. Replacement name for Eriopheidole symbiotica Kusnezov, 1952a: 16. [Junior secondary homonym of Pheidole symbiotica Wasmann, 1909: 693 (footnote).]

Stumper, R., 1951, “Études myrmécologiques. X. La myrmécobiose.” (avec une figure). Bulletin de la Société des Naturalistes Luxembourgeois, vol. 55, p. 31-43.

Stumper, R., 1950, “Les Associations Complexes des Fourmis. Commensalisme, Symbiose et Parasitisme.” (Avec 6 figures). Bulletin biologique de la France et de la Belgique, vol. 84, p. 376-399.

102 references...

First: Yes, we keep Teleutomyrmex Kutter, 1950 and Anergates Forel, 1874.

All species in these little genera with the names Teleutomyrmex Kutter, 1950 and Anergates Forel, 1874 and, of course, the big genus Tetramorium Mayr, 1855.

Second: No, we don’t keep Teleutomyrmex Kutter, 1950 and Anergates Forel, 1874 but yes, we keep Tetramorium Mayr, 1855.

But: Only a few species in the genus Tetramorium Mayr, 1855 following Ward et al., 2015 (“2014”)…

Tetramorium inquilinum Ward, Brady, Fisher & Schultz, 2015
= Tetramorium schneideri (Kutter, 1950)
Tetramorium kutteri (Tinaut, 1990), still needs a new name
Tetramorium seiferti (Kiran & Karaman, in Kiran, et al. 2017)
Tetramorium buschingeri (Lapeva-Gjonova, in Kiran, et al. 2017)
Tetramorium atratulus (Schenck, 1852)
= Tetramorium friedlandi (Creighton, 1934)
Tetramorium microgyna Santschi, 1918
Tetramorium parasiticum Bolton, 1980

Third: No, we don’t keep Teleutomyrmex Kutter, 1950, Anergates Forel, 1874 and Tetramorium Mayr, 1855.

Or: …, if you follow the line further then Ward et al., 2015 (“2014”), a few species in the genus Strongylognathus Mayr, 1853.

Strongylognathus inquilinum (Ward, Brady, Fisher & Schultz, 2015)
= Strongylognathus schneideri (Kutter, 1950)
Strongylognathus kutteri (Tinaut, 1990), still needs a new name
Strongylognathus seiferti (Kiran & Karaman, in Kiran, et al. 2017)
Strongylognathus buschingeri (Lapeva-Gjonova, in Kiran, et al. 2017)
Strongylognathus atratulus (Schenck, 1852)
= Strongylognathus friedlandi (Creighton, 1934)
Strongylognathus microgyna (Santschi, 1918)
Strongylognathus parasiticum (Bolton, 1980)

The original descriptions.

Tetramorium Mayr, 1855.

Tetramorium [Myrmicinae: Tetramoriini].
- Tetramorium Mayr, 1855: 423. Type-species: Formica caespitum, by subsequent designation of Girard, 1879: 1016.

Strongylognathus Mayr, 1853e.

Strongylognathus [Myrmicinae: Tetramoriini].
- Strongylognathus Mayr, 1853e: 389. Type-species: Eciton testaceum, by monotypy. Replacement name for Myrmus Schenck, 1853b: 188. [Junior homonym of Myrmus Hahn, 1832: 81 (Hemiptera).].

Myrmus [junior homonym, see Strongylognathus].
- Myrmus Schenck, 1853b: 188. Type-species: Myrmus emarginatus (junior synonym of Eciton testaceum), by monotypy. [Junior homonym of Myrmus Hahn, 1832: 81 (Hemiptera).].
- Strongylognathus Mayr, 1853e: 389, replacement name for Myrmus Schenck.

Myrmus Schenck, 1853b.

Myrmus [junior homonym, see Strongylognathus].
- Myrmus Schenck, 1853b: 188. Type-species: Myrmus emarginatus (junior synonym of Eciton testaceum), by monotypy. [Junior homonym of Myrmus Hahn, 1832: 81 (Hemiptera).].
- Strongylognathus Mayr, 1853e: 389, replacement name for Myrmus Schenck.

The original references.

Girard, M., 1879, “Traité élémentaire d'entomologie. Volume 2.” Paris, Librairie J.-B. Baillière et Fils, 1028 pp.

Hahn, C. W., 1832, “Die Wanzenartigen Insecten. Getreu nach der Natur abgebildet und beschrieben. Band I, Heft 3.” Nürnberg, Zeh, p. 81-118.

Mayr, G., 1853e, “Ueber die Abtheilung der Myrmiciden, und eine neue Gattung derselben.” Verhandlungen der Zoologisch-Botanischen Vereins in Wien, vol. 3, p. 387-394.

Mayr, G., 1855, “Formicina austriaca. Beschreibung der bisher im österreichischen Kaiserstaate aufgefundenen Ameisen, nebst Hinzufügung jener in Deutschland, in der Schweiz und in Italien vorkommenden Arten.” Verhandlungen der Zoologisch-Botanischen Vereins in Wien, vol. 5, p. 273-478.

Schenck, C. F., 1853b, “Die nassauischen Ameisen-Species. (Fortsetzung.).” Stettiner Entomologische Zeitung, vol. 14, p. 185-198.