The extreme, workerless inquilines.

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I noticed two days ago that I did miss a part about T. schneideri in the general review. It was written by Stumper in 1951. I'll put it later online as a small supplement.....

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Here is the page. It is the beginning of page 22, before "Tinaut, A., 1990.".

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Included is an update. Stumper is in it and my name article. The name is not valid any more, published in an unofficial way but it could not be in a valid way....

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Two updates for additional lit.:

Gray, K. W., Rabeling, C., 2023 (“2022”), “Global biogeography of ant social parasites: Exploring patterns and mechanisms of an inverse latitudinal diversity gradient.” Journal of Biogeography, vol. 50, no. 2, p. 316-329. (+ 4 supporting files.). (Article first published online: 08 December 2022).

Finally all info!

Gösswald, K., 1938, “Über den Sozialparasitismus der Ameisen.” P. 1149-1155 in Vol. 2 in: Jordan, K., Hering, E. M. (editors.), 1939 (vol. 1-4)-1940 (vol. 5), “VII. Internationaler Kongress für Entomologie. Herausgegeben im Auftrage des Kongresses von K. Jordan und E. M. Hering.“ Selbstverläge der Internationalen Kongresse für Entomologie. Weimar, G. Uschmann, vol. 1-5, 3130 + ccxliv pp.

For the old: Gösswald, K., 1938, “Über den Sozialparasitismus der Ameisen.” VII. Internationaler Entomo¬ logenkongress, p. 1149—1155.

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Not jet included in the above manuscript but I am looking for an easy alternative....

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A few more inclusions:

“Creighton, W. S., 1950.”, for Anergates.

“This extraordinary workerless parasite was first described by Schenck in 1852. This observer was also the first to publish on the habits of Anergates. It would be hard to imagine a more remarkable set of structural modifications than those which occur in both sexes of A. atratulus. The virgin female possesses a deep, median sulcus on the dorsum of the gaster. After fertilization the excessive development of the ovaries separates the gastric sclerites and stretches the intersegmental membranes between them. The lateral expansion is greater than the dorsoventral stretching, with the result that the gaster of the fertile female finally assumes a shape rather like a biscuit. The separated sclerites appear like islands on the top and bottom surfaces. The structure of the male is even more aberrant. This sex is apterous and pupoidal. The body apparently never becomes fully chitinized. The gaster is curved under at the tip and provided with a disproportionally large set of genitalia. In both sexes the mandibles are poorly developed and apparently quite useless for purposes of feeding. It is no wonder that this insect has attracted much interest or that, despite its rarity, its habits have been repeatedly studied. While the significance of some of the observations is not altogether clear, there is good general agreement as to the behavior of this remarkable species. Anergates is a parasite of Tetramorium caespitum. At maturity a parasitized nest consists of a single fertile female of Anergates, a considerable number of T. caespitum workers and a large number of pupoidal males and virgin females of Anergates. The T. caespitum workers feed and care for the parasites but are particularly attentive to the males and much less interested in the females, (Adlerz 1913, Wasmann 1908). It was shown by Janet (1897) that the male of Anergates possesses unusually large mandibular glands and Forel (1922) supposes that the secretion from these glands is relished by the T. caespitum workers. Because the Anergates male is apterous, the nuptial flight has undergone some peculiar modifications. Copulation takes place between sisters and brothers (adelphogamy) within the confines of the nest. Mating is difficult for the clumsy, pupoidal male and is facilitated, as Forel has pointed out, (1922) by the docility of the female. After fertilization the female emerges from the nest for the nuptial flight. It is obvious that at the completion of the nuptial flight the Anergates female must find a nest of T. caespitum and gain access to it. Repeated experiments have been performed to ascertain how this is accomplished but the results have been uncertain in most cases. It is easy to show that the fertilized Anergates female will try to gain access to the T. caespitum nest. This she does by making an unobserved entry or, in many cases, by seizing the antenna of a T. caespitum worker who thereupon drags her into the nest. What follows next is not so clear. In the great majority of cases the Anergates female is killed by the T. caespitum workers if we may judge from experimental data. In one case, however, there has been a different result. In 1912 Crawley succeeded in getting a colony of Tetramorium to accept an Anergates female. This colony contained sexual forms of the host. At the end of a week following the introduction of the Anergates female to the nest, the T. caespitum workers had killed and cut to pieces the T. caespitum males and females. The introduced female of Anergates later showed the characteristic gastric enlargement which marks the mature queen but died without having laid any eggs. In summing up the evidence concerning this and other experiments Donisthorpe (1915) expressed the opinion that the destruction of the Tetramorium female by her own workers is probably the normal sequence of events following the entrance of the Anergates female into a T. caespitum colony. It is difficult to see how else the T. caespitum female could be eliminated, since it is unlikely that the Anergates female could kill her.”

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A remark about the literature lists of Anergates and Teleutomyrmex.

A small remark about the “* ” for some of the references in those literature lists. This sign is placed there when the reference is about the hosts of the genus named in the title or when the reference is about a part of the systematic places of the species around the species but not about it directly.

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Additional references.
(A few remarks: Adlerz, G., 1913 is untraceable. Crawley, W. C., 1912, Gösswald, K., 1938, Gösswald, K., 1951, Janet, C., 1897 and Wasmann, E., 1908 are not sure so a few alternatives are also given.).

Adlerz, G., 1886, “Myrmecologiska studier. II. Svenska myror och deras lefnadsförhållanden.” Bihang till Kongliga Svenska Vetenskaps-Akademiens Handlingar, vol. 11, no. 18, p. 1-329.

Arnold, G., 1926, “A monograph of the Formicidae of South Africa. Appendix.” Annals of the South African Museum, vol. 23, p. 191-295.

Buschinger, A., 1999, “Bemerkenswerte ameisenfunde aus Sudtirol (Hymenoptera: Formicidae).“ Myrmecologische Nachrichten, vol. 3, p. 1-8.

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

Crawley, W. C., 1912, “Anergates atratulus, Schenk, a british ant, and the acceptance of a queen by Tetramorium caespitum, L.“ The Entomologist´s Record and Journal of Variation, London, vol. 24, nr. 9, p. 218-219.

Crawley, W. C., Donisthorpe, H., 1913, “The Founding of Colonies by Queen Ants.” (Anergates atratulus, p. 68-73.) P. 11-77, in: Jordan, K., Eltringham, H. (editors.), 1913, “2nd International Congress of Entomology, Oxford August 1912. Volume 2. Transactions.“ Oxford. London and Aylesbury, Hazell, Watson & Viney, Ld., 489 pp. (+ 34 plates!).

Dlussky, G. M., 1981, “Muravi pustyn.”, or “[Desert Ants.]” [In Russian.]. Moskva, Nauka, 230 pp.

Emery, C., 1909, “Über den Ursprung der dulotischen, parasitischen und myrmekophilen Ameisen.” Biologisches Centralblatt, vol. 29, p. 352-362.

Emery, C., 1913, “Über die Abstammung der europäischen arbeiterinnenlosen Ameise “Anergates“.“ Biologisches Centralblatt, vol. 33, p. 258-260.

Forel, A., 1893, “Sur la classification de la famille des Formicides, avec remarques synonymiques.” Annales de la Société Entomologique de Belgique, vol. 37, p. 161-167.

Forel, A., 1922, “Le monde social des fourmis comparé à celui de l'homme. Tome 3. Appareils d'observation. Fondation des foumilières. Moeurs à l'intérieur des nids. Bétail, jardins, fourmis parasites. (Avec appendice du Dr. E. Bugnion.).” Genève, Librairie Kundig, vii + 227 pp.

Gösswald, K., 1938, “Grundsätzliches über parasitische Ameisen unter besonderer Berücksichtigung der abhängigen Koloniegründung von Lasius umbratus mixtus Nyl.” Zeitschrift für Wissenschaftliche Zoologie, vol. 151, p. 101-148.

Gösswald, K., 1938, “Über den Sozialparasitismus der Ameisen.” P. 1149-1155 in Vol. 2 in: Jordan, K., Hering, E. M. (editors.), 1939 (vol. 1-4)-1940 (vol. 5), “VII. Internationaler Kongress für Entomologie. Herausgegeben im Auftrage des Kongresses von K. Jordan und E. M. Hering.“ Selbstverläge der Internationalen Kongresse für Entomologie. Weimar, G. Uschmann, vol. 1-5, 3130 + ccxliv pp.

Gösswald, K., 1951, “Zur Biologie, Ökologie und Morphologie einer neuen Varietät der Kleinen Roten Waldameise: Formica minor pratensoides.“ Zeitschrift für Angewandte Entomologie, vol. 32, p. 433457.

Gösswald, K., 1951, “Versuche zum Sozialparasitismus der Ameisen bei der Gattung Formica L.“ Zoologische Jahrbücher. Abteilung für Systematik, Ökologie und Geographie der Tiere, vol. 80, p. 533582.

Gray, K. W., Rabeling, C., 2023 (“2022”), “Global biogeography of ant social parasites: Exploring patterns and mechanisms of an inverse latitudinal diversity gradient.” Journal of Biogeography, vol. 50, no. 2, p. 316-329. (+ 4 supporting files.). (Article first published online: 08 December 2022).

International Commission on Zoological Nomenclature (I.C.Z.N.) [Ride, W. D. L., Cogger, H. G., Dupuis, C., Kraus, O., Minelli, A., Thompson, F. C., Tubbs, P. K. (editors.)], 1999, “International Code of Zoological Nomenclature [= Code International de Nomenclature Zoologique]. Fourth Edition, adopted by the International Union of Biological Sciences.” London, The International Trust for Zoological Nomenclature, c/o The Natural History Museum, 1999. Padova, Tipografia La Garangola, xxix + 306 pp.

Janet, C., 1896, “Les fourmis.” Bulletin de la Société Zoologique de France, vol. 21, p. 60-93.

Janet, C., 1897, ”Études sur les fourmis, les guêpes et les abeilles. Note 13. Sur le Lasius mixtus, l'Antennophorus uhlmanni, etc.” Limoges, H. Ducourtieux, 62 pp.

Janet, C., 1897, “Études sur les fourmis, les guêpes et les abeilles. Note 16. Limites morphologiques des anneaux post-céphaliques et musculature des anneaux post-thoraciques chez la Myrmica rubra.” Lille, Le Bigot Frères, 36 pp.

Janet, C., 1898, ”Études sur les fourmis, les guêpes et les abeilles. Note 17. Système glandulaire tégumentaire de la Myrmica rubra. Observations diverses sur les fourmis.” Paris, G. Carré et C. Naud, 30 pp.

Latreille, P. A., 1804, “Tableau méthodique des insectes.” Pp. 129-200 in: Société de Naturalistes et d'Agriculteurs, “Nouveau dictionnaire d'histoire naturelle. Tome 24.” Paris, Déterville, 84 + 85 + 238 + 18 + 34 pp.

Latreille, P. A., 1810, “Considérations générales sur l'ordre natural des animaux composant les classes des Crustacés, des Arachnides et des Insectes; avec un tableau méthodique de leurs genres, disposés en familles.” Paris, F. Schoell, 444 pp.

Linsenmaier, W., 1972, “Knaurs Grosses Insektenbuch.” München, Th. Knaur Nachf., 396 Seiten, 1888 Insektendarstellungen (together with an English edition: Linsenmaier, W., 1972, “Insects of the World.” New York, Mc Graw-Hill. I have the French edition: Linsenmaier, W., 1973. “Insectes du Monde.” Collection Eugène Clarence Braun-Munk. Suisse, Stock, 380 pp.).

Lopez, F., 1988, “Aproximación al estudio del género Tetramorium Mayr, 1855 en la Peninsula Ibérica (Hymenoptera, Formicidae).” Mem. Licenciatura. Universidad Complutense, Madrid, 295 pp.

Lopez-Osorio, F., 2020, “Emery’s Rule.” P. 365-368 in: Starr, C. K. (editor.), 2021, “Encyclopedia of Social Insects.” Springer International Publishing, Cham., xxvi + 1049 pp.

Santschi, F., 1928, “Descriptions de nouvelles fourmis éthiopiennes (suite).” Revue de Zoologie et de Botanique Africaines, vol. 16, p. 191-213.

Schär, S., Talavera, G., Espadaler, X., Rana, J. D., Andersen, A. A., Cover, S. P., Vila, R., 2018, “Do Holarctic ant species exist? Trans-Beringian dispersal and homoplasy in the Formicidae.” Journal of Biogeography, vol. 45, no. 8, p. 1917-1928. (+ 6 supporting files.).

Smith, D. R., 1979, “Superfamily Formicoidea.” Pp. 1323-1467 in: Krombein, K. V., Hurd, P. D., Smith, D. R., Burks, B. D. (editors.), 1979, “Catalog of Hymenoptera in America north of Mexico. Volume 2. Apocrita (Aculeata).” Washington, D.C., Smithsonian Institution Press, pp. i-xvi, 1199-2209.

von Hagens, J., 1867, “Ueber Ameisen mit gemischten Colonien.” Berliner Entomologische Zeitschrift, vol. 11, nr. 1-2, p. 101-108.

Wagner, H. C., Karaman, C., Aksoy, V., Kiran, K., 2018, “A mixed colony of Tetramorium immigrans Santschi, 1927 and the putative social parasite Tetramorium aspina sp.n. (Hymenoptera: Formicidae).” Myrmecological News, vol. 28, p. 25-33.

Wasmann, E., 1891, “Die zusammengesetzten Nester und gemischten Kolonien der Ameisen. Ein Beitrag zur Biologie, Psychologie und Entwicklungsgeschichte der Ameisengesellschaften.“ Münster in Westfalen, Aschendorffschen Buchdruckerei, vii + 262 pp. (+ 2 plates!).

Wasmann, E., 1908, “Nachtrag zu: Weitere Beiträge zum sozialen Parasitismus und der Sklaverei bei den Ameisen.“ Biologisches Zentralblatt, vol. 28, p. 726-731.

Wasmann, E., 1908, “Weitere Beiträge zum sozialen Parasitismus und der Sklaverei bei den Ameisen.“ Biologisches Zentralblatt, vol. 28, p. 257-271, 290-306, 321-333, 353-382, 417-441.

Wasmann, E., 1909, “Über den Ursprung des sozialen Parasitismus, der Sklaverei und der Myrmekophilie bei den Ameisen.“ Biologisches Centralblatt (Berlin), vol. 29, p. 587-604.

Wasmann, E., 1909, “Über den Ursprung des sozialen Parasitismus, der Sklaverei und der Myrmekophilie bei den Ameisen. (Fortsetzung.).“ [part a] Biologisches Centralblatt (Berlin), vol. 29, p. 619-637.

Wasmann, E., 1909, “Über den Ursprung des sozialen Parasitismus, der Sklaverei und der Myrmekophilie bei den Ameisen. (Fortsetzung.).“ [part b] Biologisches Centralblatt (Berlin) vol. 29, p. 651-663.

Wasmann, E., 1909, “Über den Ursprung des sozialen Parasitismus, der Sklaverei und der Myrmekophilie bei den Ameisen. (Schluss.).“ Biologisches Centralblatt (Berlin), vol. 29, p. 683-703.

Wheeler, W. M., 1908, “Comparative ethology of the European and North American ants.” Journal für Psychologie und Neurologie, vol. 13, p. 404-435, pl. III-IV.

Wheeler, W. M., 1910, “Ants: Their Structure, Development and Behavior.” Columbia Biological Series, no. 9. New York, Columbia University Press, xxv + 663 pp.

Wheeler, W. M., 1937, “Mosaics and other anomalies among ants.” Cambridge, Mass., Harvard University Press, 95 pp.

Yarrow, I. H. H., 1955, “The type species of the ant genus Myrmica Latreille.” Proceedings of the Royal Entomological Society of London. Series B, vol. 24, p. 113-115.

42 references.

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And I worked also a little at the lay-out of it....

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Tryed it for the last time, this time by "Myrmecological News."....

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Now, it was put for inclusion in "Myrmecological News" on friday, today I got this answer:

"Dear Marc Van der Stappen,
thank you again for submitting your manuscript to Myrmecol. News. We have discussed among us editors how to proceed with this rather unusual manuscript and have arrived at the conclusion that it is valuable but not suitable for our journal. We have also tried to think of another outlet which might be better suited but not succesfully so.

We believe that you have been in touch with Professor Buschinger for a long while already, right? Maybe he could have in mind a potential oulet for this piece of work!?

Sorry that we do not have better news and hoping for your understanding for our decision,
on behalf of the editorial team,
warmest,
f"

So, no hard feelings but... I give up!

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I know Prof. Buschinger a long time, since 12/13 June 2002, so more then 20 years, and when I did ask him something, whenever he was in the possebility to help, he did that. But here he can't help so it will be the end for me, sad but so it is. I will put the final version of the document online when I have made pdf's of them and then.....

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The new version:

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And the second half:

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Now, this should not be here, but where do I put it then....

"Insecta Helvetica Fauna" is now online, viewable and for download, in parts or in total...

And who made vol. 6 and vol. 6a...

Yep, enjoy it!!!

https://www.e-periodica.ch/digbib/volumes?UID=ihf-001

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The complete Anergates pages from Kutter, 1977:

The complete and good Teleutomyrmex pages from Kutter, 1977:

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His thougths about the great variances in the genus Tetramorium:

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And further:

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And the differences in determination keys:

Females:

Males:

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Now, an update but not for the pictures (parts 2, 4, 6, 8 and 10.), For them, see viewtopic.php?f=23&t=1533&start=240#p23613 .