Beiträge von Teleutotje

    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, in Turkmenistan 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, T. buschingeri and T. seiferti.


    The ants from the genus Teleutomyrmex are the most specialized ants on Earth. They are extreme, workerless inquilines that became 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 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!). Only Nylanderia belongs to a different subfamily, the Formicinae. Most of the known social parasitic ants belong to these two subfamilies…


    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 question the decision and wait for the ongoing genetic comparison of European and North American Anergates-samples. 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, for the moment, both names stay as species names on record. Now, with the description of two more species, things are getting clearer in this genus.


    Paratrechina and related genera were reviewed on genera-level in 2010 and one of the results was the division of Paratrechina into a few related genera, including Nylanderia. Some of these genera and part of the genus Nylanderia are recently revised and Nylanderia from North America was revised also but without the inquilines.


    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?

    Regulation of larval diapause by colony-founding queens of Crematogaster teranishii (Hymenoptera: Formicidae)


    Keiji NAKAMURA


    Department of Biosphere-Geosphere Science, Faculty of Biosphere-Geosphere Science, Okayama University of Science, Ridai-cho 1-1, Okayama 700-0005, Japan; e-mail: nakamura@big.ous.ac.jp


    Nuptial flights of Crematogaster teranishii Santschi occur in autumn. Queens and possibly larvae pass the first winter without workers in Okayama, Japan. This study examines and tests the hypothesis that C. teranishii queens regulate not only their own diapause, but also that of their larvae. Some queens collected immediately after their nuptial flight in early October were exposed to a low temperature of 10°C for 3 months before transfer to 25°C; these queens started oviposition approximately 8 days after the transfer. Larvae reared by these previously chilled queens pupated synchronously until day 50. Other field-collected queens reared at 25°C without prior chilling reproduced soon after collection. They started oviposition, but the development of their larvae was arrested and pupation occurred late or not at all within the120 day experimental period. The delay in pupation in colonies in which the queens were not previously exposed to low temperature strongly indicates that many larvae are in diapause. Because eggs and larvae were not exposed to low temperatures, they were unable to determine the developmental pathway providing the best response to environmental factors. Results indicate that queens regulate whether their larvae undergo diapause.


    Keywords: Hymenoptera, Formicidae, colony development, ant, larval diapause, reproductive diapause of queens, low temperature, social effect


    https://www.eje.cz/pdfs/eje/2018/01/68.pdf

    Solenopsis invicta Buren, 1972 and Solenopsis saevissima wagneri Santschi, 1916e.


    Bolton, 1995b did something very stirring to the usage of these species names. By recognizing the correct use of S. wagneri and giving it priority over S. invicta there was confusion about witch name to use. Some did use S. wagneri but it lasted till 1999/2000 till everybody accepted S. Invicta. So, those 5 to 6 years you can find also S. wagneri in some publications… but the correct name is S. invicta!


    Solenopsis metallica Pitts, Camacho, Gotzek, McHugh & Ross, 2018 and S. altipunctata by Pitts, 2002.


    In 2002 Pitts described S. altipunctata in his thesis, NOT in an official publication. Only in 2018 this species was described as S. metallica Pitts, Camacho, Gotzek, McHugh & Ross, 2018. In those 16 years the species was named in most publications S. n. sp. But… a few times S. altipunctata was used. This is not valid. Since 2018 the valid name is S. metallica Pitts, Camacho, Gotzek, McHugh & Ross, 2018.


    Solenopsis geminata var. micans Stitz, 1912, Solenopsis pylades var. itinerans Forel, 1911f and Solenopsis tridens var. lehmannnitschei Santschi, 1916e.


    These 3 subspecies escaped the revision of Trager and all the later ones… These should still be revised. Until then they stay as subspecies of S. geminate, S. saevissima and S. tridens respectively.

    Solenopsis pylades var. itinerans Forel, 1911f.


    - itinerans. Solenopsis pylades var. itinerans Forel, 1911f: 276 (w.) KENYA. Currently subspecies of saevissima: Wheeler, W.M. 1922a: 879.


    Solenopsis tridens var. substituta Santschi, 1925d.


    - substituta. Solenopsis tridens var. substituta Santschi, 1925d: 236 (w.q.) BRAZIL. Raised to species: Trager, 1991: 161.


    Solenopsis tridens Forel, 1911c.


    - tridens. Solenopsis tridens Forel, 1911c: 298 (w.) BRAZIL. Current subspecies: nominal plus lehmannnitschei. See also: Trager, 1991: 162.


    Solenopsis tridens var. lehmannnitschei Santschi, 1916e.


    - lehmannnitschei. Solenopsis tridens var. lehmannnitschei Santschi, 1916e: 378 (w.) ARGENTINA.


    Myrmica virulens Smith, F. 1858b.


    - virulens. Myrmica virulens Smith, F. 1858b: 132 (w.) BRAZIL. Combination in Solenopsis: Dalla Torre, 1893: 76. Junior synonym of geminata: Mayr, 1886d: 460; Dalla Torre, 1893: 76; Creighton, 1930b: 59; Creighton, 1950a: 231; of bondari: Kempf & Brown, 1968: 99. Revived from synonymy and senior synonym of bondari: Trager, 1991: 155.
    - bondari. Solenopsis bondari Santschi, 1925d: 236 (w.) BRAZIL. Junior synonym of virulens: Trager, 1991: 155.


    Solenopsis bondari Santschi, 1925d.


    - bondari. Solenopsis bondari Santschi, 1925d: 236 (w.) BRAZIL. Junior synonym of virulens: Trager, 1991: 155.


    Solenopsis weyrauchi Trager, 1991.


    - weyrauchi. Solenopsis weyrauchi Trager, 1991: 190, figs. 76, 77, 80, 81 (w.) PERU.


    Solenopsis xyloni McCook, 1879.


    - xyloni. Solenopsis xyloni McCook, 1879: 188, figs. 37, 38 (s.w.q.) U.S.A. Wheeler, W.M. 1915b: 396 (m.); Wheeler, G.C. & Wheeler, J. 1955c: 133 (l.); Taber & Cokendolpher, 1988: 95 (k.). Junior synonym of geminata: Mayr, 1886d: 460; Dalla Torre, 1893: 76. Revived from synonymy as subspecies of geminata: Wheeler, W.M. 1910g: 563; Wheeler, W.M. 1915b: 395. Revived status as species: Creighton, 1930b: 98; Creighton, 1950a: 232; Snelling, R.R. 1963: 9. Senior synonym of maniosa, pylades: Trager, 1991: 166.
    - pylades. Solenopsis geminata r. pylades Forel, 1904d: 172 (q.) MEXICO. Forel, 1909a: 267 (w.); Forel, 1911c: 297 (m.); Bruch, 1916: 315 (m.). Raised to species: Forel, 1909a: 267; Bruch, 1914: 223; Forel, 1916: 459. Subspecies of saevissima: Santschi, 1923c: 267. Junior synonym of geminata: Emery, 1922e: 198; of saevissima: Wheeler, W.M. 1915b: 395; Wheeler, W.M. 1916g: 143; Wheeler, W.M. 1919c: 272; Creighton, 1930b: 70; of xyloni: Trager, 1991: 166.
    - maniosa. Solenopsis geminata subsp. maniosa Wheeler, W.M. 1915b: 396 (w.q.m.) U.S.A. Subspecies of xyloni: Creighton, 1930b: 102. Junior synonym of xyloni: Creighton, 1950a: 232. Revived from synonymy and raised to species: Wheeler, G.C. & Wheeler, J. 1986g: 13. Junior synonym of xyloni: Trager, 1991: 166.


    Solenopsis geminata r. pylades Forel, 1904d.


    - pylades. Solenopsis geminata r. pylades Forel, 1904d: 172 (q.) MEXICO. Forel, 1909a: 267 (w.); Forel, 1911c: 297 (m.); Bruch, 1916: 315 (m.). Raised to species: Forel, 1909a: 267; Bruch, 1914: 223; Forel, 1916: 459. Subspecies of saevissima: Santschi, 1923c: 267. Junior synonym of geminata: Emery, 1922e: 198; of saevissima: Wheeler, W.M. 1915b: 395; Wheeler, W.M. 1916g: 143; Wheeler, W.M. 1919c: 272; Creighton, 1930b: 70; of xyloni: Trager, 1991: 166.


    Solenopsis geminata subsp. maniosa Wheeler, W.M. 1915b.


    - maniosa. Solenopsis geminata subsp. maniosa Wheeler, W.M. 1915b: 396 (w.q.m.) U.S.A. Subspecies of xyloni: Creighton, 1930b: 102. Junior synonym of xyloni: Creighton, 1950a: 232. Revived from synonymy and raised to species: Wheeler, G.C. & Wheeler, J. 1986g: 13. Junior synonym of xyloni: Trager, 1991: 166.

    Solenopsis invicta Buren, 1972.


    - wagneri. Solenopsis saevissima var. wagneri Santschi, 1916e: 380 (w.) ARGENTINA. Junior synonym of saevissima: Wilson, 1952b: 55. Revived from synonymy, raised to species and senior synonym of invicta: Bolton, 1995b: 391. [Trager, 1991: 173 wrongly gave wagneri as an unavailable name, incorrectly citing its original notation as S. saevissima subsp. electra var. wagneri, and referring its material to invicta. But wagneri is available, with the original notation cited above, and has priority over invicta.] S. invicta conserved over wagneri because of usage, in accord with ICZN (1999): Shattuck, Porter & Wojcik, 1999: 27.
    - invicta. Solenopsis invicta Buren, 1972: 9, fig. 2 (w.q.m.) BRAZIL. Wheeler, G.C. & Wheeler, J. 1977: 588 (l.). Junior synonym of wagneri: Bolton, 1995b: 388. [Trager, 1991: 173 incorrectly gave wagneri as an unavailable name; the name is available and has priority over invicta, see note under wagneri.] S. invicta conserved over wagneri because of usage, in accord with ICZN (1999): Shattuck, Porter & Wojcik, 1999: 27. See also: Rhoades, 1977: 1; Smith, D.R. 1979: 1386.


    Solenopsis saevissima wagneri Santschi, 1916e.


    - wagneri. Solenopsis saevissima var. wagneri Santschi, 1916e: 380 (w.) ARGENTINA. Junior synonym of saevissima: Wilson, 1952b: 55. Revived from synonymy, raised to species and senior synonym of invicta: Bolton, 1995b: 391. [Trager, 1991: 173 wrongly gave wagneri as an unavailable name, incorrectly citing its original notation as S. saevissima subsp. electra var. wagneri, and referring its material to invicta. But wagneri is available, with the original notation cited above, and has priority over invicta.] S. invicta conserved over wagneri because of usage, in accord with ICZN (1999): Shattuck, Porter & Wojcik, 1999: 27.


    Solenopsis saevissima var. macdonaghi Santschi, 1916e.


    - macdonaghi. Solenopsis saevissima var. macdonaghi Santschi, 1916e: 379 (w.q.) ARGENTINA. Junior synonym of interrupta: Creighton, 1930b: 89. Revived from synonymy and raised to species: Trager, 1991: 179.


    Solenopsis megergates Trager, 1991.


    - megergates. Solenopsis megergates Trager, 1991: 181, figs. 68, 69, 72, 73 (w.q.) BRAZIL.


    Solenopsis metallica Pitts, Camacho, Gotzek, McHugh & Ross, 2018.


    - metallica. Solenopsis metallica Pitts, Camacho, Gotzek, McHugh & Ross, 2018: 331, figs. 16–20, 44, 89 (w.q.m.l.) BRAZIL.


    Solenopsis pusillignis Trager, 1991.


    - pusillignis. Solenopsis pusillignis Trager, 1991: 194, figs. 40, 41, 44, 45, 84 (w.) BRAZIL.


    Solenopsis (Solenopsis) pythia Santschi, 1934c.


    - pythia. Solenopsis (Solenopsis) pythia Santschi, 1934c: 30 (q.) ARGENTINA. Trager, 1991: 182 (w.). See also: Wilson, 1952b: 50.


    Solenopsis pylades var. quinquecuspis Forel, 1913l.


    - quinquecuspis. Solenopsis pylades var. quinquecuspis Forel, 1913l: 224 (w.) ARGENTINA. Subspecies of saevissima: Santschi, 1916e: 379; Creighton, 1930b: 86. Junior synonym of richteri: Wilson, 1952b: 57. Revived from synonymy and raised to species: Buren, 1972: 17. Senior synonym of blumi: Trager, 1991: 185.
    - blumi. Solenopsis blumi Buren, 1972: 20, fig. 5 (w.) URUGUAY. Junior synonym of quinquecuspis: Trager, 1991: 185.


    Solenopsis blumi Buren, 1972.


    - blumi. Solenopsis blumi Buren, 1972: 20, fig. 5 (w.) URUGUAY. Junior synonym of quinquecuspis: Trager, 1991: 185.


    Solenopsis pylades var. richteri Forel, 1909a.


    - richteri. Solenopsis pylades var. richteri Forel, 1909a: 267 (w.q.) ARGENTINA. Creighton, 1930b: 87 (m.); Wheeler, G.C. & Wheeler, J. 1977: 589 (l.). Subspecies of saevissima: Santschi, 1916e: 379; Creighton, 1950a: 232; Wilson, 1952b: 58. Raised to species: Buren, 1972: 4. Senior synonym of tricuspis: Creighton, 1930b: 87; of oblongiceps: Trager, 1991: 187. See also: Rhoades, 1977: 1.
    - tricuspis. Solenopsis pylades var. tricuspis Forel, 1912g: 4 (w.) ARGENTINA. Subspecies of saevissima: Santschi, 1916e: 379. Junior synonym of richteri: Creighton, 1930b: 87.
    - oblongiceps. Solenopsis saevissima st. oblongiceps Santschi, 1936d: 405, fig. 5 (w.) ARGENTINA. Junior synonym of saevissima: Wilson, 1952b: 55; of richteri: Trager, 1991: 187.


    Solenopsis pylades var. tricuspis Forel, 1912g.


    - tricuspis. Solenopsis pylades var. tricuspis Forel, 1912g: 4 (w.) ARGENTINA. Subspecies of saevissima: Santschi, 1916e: 379. Junior synonym of richteri: Creighton, 1930b: 87.


    Solenopsis saevissima st. oblongiceps Santschi, 1936d.


    - oblongiceps. Solenopsis saevissima st. oblongiceps Santschi, 1936d: 405, fig. 5 (w.) ARGENTINA. Junior synonym of saevissima: Wilson, 1952b: 55; of richteri: Trager, 1991: 187.


    Myrmica saevissima Smith, F. 1855c


    - saevissima. Myrmica saevissima Smith, F. 1855c: 166, pl. 13, fig. 18 (w.) BRAZIL. Combination in Solenopsis: Mayr, 1862: 751. Goni, Zolessi & Imai, 1983: 365 (k.). Junior synonym of geminata: Mayr, 1865: 109; Mayr, 1886d: 460; Dalla Torre, 1893: 76. Revived from synonymy as subspecies of geminata: Wheeler, W.M. 1915b: 395. Revived status as species: Mann, 1916: 447; Santschi, 1916e: 378; Santschi, 1923c: 265; Creighton, 1930b: 80. Senior synonym of gracilior: Creighton, 1930b: 80; of moelleri (and its junior synonyms incrassata, morosa), perfida: Wilson, 1952b: 55. Material of the unavailable name picea referred here by Trager, 1991: 189. Current subspecies: nominal plus itinerans. See also: Buren, 1972: 15; Trager, 1991: 188.
    - gracilior. Solenopsis moelleri var. gracilior Forel, 1904d: 174 (w.) BRAZIL. Junior synonym of saevissima: Creighton, 1930b: 80.
    - moelleri. Solenopsis moelleri Forel, 1904d: 173 (w.q.m.) BRAZIL. Subspecies of saevissima and senior synonym of incrassata, morosa: Creighton, 1930b: 83. Junior synonym of saevissima: Wilson, 1952b: 55.
    -incrassata. Solenopsis geminata var. incrassata Forel, 1908c: 362 (w.) BRAZIL. Subspecies of pylades: Forel, 1909a: 268; of saevissima: Gallardo, 1919b: 247. Junior synonym of moelleri: Creighton, 1930b: 83.
    - morosa. Solenopsis saevissima var. morosa Santschi, 1916e: 380 (w.) BRAZIL. Santschi, 1923c: 265 (q.). Junior synonym of moelleri: Creighton, 1930b: 83.
    - perfida. Solenopsis saevissima var. perfida Santschi, 1923c: 266 (w.) BRAZIL. Junior synonym of saevissima: Wilson, 1952b: 55.


    Solenopsis moelleri var. gracilior Forel, 1904d.


    - gracilior. Solenopsis moelleri var. gracilior Forel, 1904d: 174 (w.) BRAZIL. Junior synonym of saevissima: Creighton, 1930b: 80.


    Solenopsis moelleri Forel, 1904d.


    - moelleri. Solenopsis moelleri Forel, 1904d: 173 (w.q.m.) BRAZIL. Subspecies of saevissima and senior synonym of incrassata, morosa: Creighton, 1930b: 83. Junior synonym of saevissima: Wilson, 1952b: 55.


    Solenopsis geminata var. incrassata Forel, 1908c.


    - incrassata. Solenopsis geminata var. incrassata Forel, 1908c: 362 (w.) BRAZIL. Subspecies of pylades: Forel, 1909a: 268; of saevissima: Gallardo, 1919b: 247. Junior synonym of moelleri: Creighton, 1930b: 83.


    Solenopsis saevissima var. morosa Santschi, 1916e.


    - morosa. Solenopsis saevissima var. morosa Santschi, 1916e: 380 (w.) BRAZIL. Santschi, 1923c: 265 (q.). Junior synonym of moelleri: Creighton, 1930b: 83.


    Solenopsis saevissima var. perfida Santschi, 1923c.


    - perfida. Solenopsis saevissima var. perfida Santschi, 1923c: 266 (w.) BRAZIL. Junior synonym of saevissima: Wilson, 1952b: 55.

    Atta geminata Fabricius, 1804.


    - geminata. Atta geminata Fabricius, 1804: 423 (q.) CENTRAL AMERICA. Roger, 1862c: 289 (w.m.); Mayr, 1867a: 110 (w.q.m.); Wheeler, W.M. 1900b: 21 (l.); Wheeler, G.C. & Wheeler, J. 1955c: 132 (l.); Crozier, 1970: 116 (k.). Combination in Solenopsis: Mayr, 1863: 453. Senior synonym of clypeata, mandibularis: Roger, 1862c: 289; of cephalotes, paleata: Roger, 1863b: 32; Trager, 1991: 163; of glaber, laboriosus, polita: Mayr, 1863: 453; Mayr, 1886c: 362; Trager, 1991: 163; of drewseni: Mayr, 1870b: 996 (footnote); Mayr, 1886d: 460; of coloradensis: Mayr, 1886c: 365; of lincecumii, saxicola: Emery, 1895c: 276; of innota: Wheeler, W.M. 1922a: 877; of nigra: Creighton, 1930b: 59; of laevissima, mellea: Donisthorpe, 1932c: 463, 455 respectively; of rufa (and its junior synonym diabola): Ettershank, 1966: 141; of eduardi (and its junior synonym perversa), medusa (and its junior synonym bahiaensis), galapageia: Trager, 1991: 163. See also: Smith, D.R. 1979: 1386. Current subspecies: nominal plus micans.
    - paleata. Myrmica paleata Lund, 1831a: 116 (footnote) (w.) BRAZIL. Junior synonym of geminata: Roger, 1863b: 32.
    - mandibularis. Solenopsis mandibularis Westwood, 1840b: 87, pl. 2, fig. 5 (w.) "America aequinoctiali". Junior synonym of geminata: Roger, 1862c: 289.
    - rufa. Atta rufa Jerdon, 1851: 106 (w.q.) INDIA. Bingham, 1903: 158 (m.). Combination in Solenopsis: Emery, 1892b: 166. Junior synonym of geminata: Dalla Torre, 1893: 76. Revived from synonymy as subspecies of geminata: Forel, 1899a: 119; Wheeler, W.M. 1915b: 397; Creighton, 1930b: 66. Senior synonym of diabola: Creighton, 1930b: 66. Junior synonym of geminata: Ettershank, 1966: 141.
    - clypeata. Atta clypeata Smith, F. 1858b: 169 (q.m.) MEXICO. Junior synonym of geminata: Roger, 1862c: 289.
    - cephalotes. Solenopsis cephalotes Smith, F. 1859a: 149 (s.w.) INDONESIA (Aru I.). Junior synonym of geminata: Roger, 1863b: 32; of rufa: Creighton, 1950a: 231; of geminata: Trager, 1991: 163.
    - mellea. Myrmica mellea Smith, F. 1859a: 148 (w.) INDONESIA (Aru I.). Junior synonym of geminata: Donisthorpe, 1932c: 455.
    - laboriosus. Crematogaster laboriosus Smith, F. 1860b: 109 (q.) INDONESIA (Batjan I.). Combination in Solenopsis: Mayr, 1863: 453. Junior synonym of rufa: Creighton, 1950a: 231; of geminata: Trager, 1991: 163.
    - laevissima. Myrmica laevissima Smith, F. 1860b: 108 (w.) INDONESIA (Batjan I.). Junior synonym of geminata: Donisthorpe, 1932c: 463.
    - drewseni. Diplorhoptrum drewseni Mayr, 1861: 73 (diagnosis in key) (w.) ITALY. Junior synonym of geminata: Mayr, 1870b: 996; Mayr, 1886d: 460.
    - glaber. Myrmica glaber Smith, F. 1862b: 34 (w.) PANAMA. Junior synonym of geminata: Mayr, 1886c: 362; Trager, 1991: 163.
    - polita. Myrmica polita Smith, F. 1862b: 34 (w.) PANAMA. [Unresolved junior primary homonym of Myrmica polita Smith, F. 1860b: 108 (now in Pheidologeton).] Junior synonym of geminata: Mayr, 1863: 453; Mayr, 1886c: 362.
    - coloradensis. Atta coloradensis Buckley, 1867: 346 (s.w.) U.S.A. Junior synonym of geminata: Mayr, 1886c: 365.
    - lincecumii. Atta lincecumii Buckley, 1867: 344 (s.w.) U.S.A. Junior synonym of geminata: Emery, 1895c: 276.
    - saxicola. Myrmica (Monomorium) saxicola Buckley, 1867: 341 (w.) U.S.A. Junior synonym of geminata: Emery, 1895c: 276.
    - diabola. Solenopsis geminata var. diabola Wheeler, W.M. 1908e: 424 (w.q.m.) U.S.A. Junior synonym of rufa: Creighton, 1930b: 66.
    - nigra. Solenopsis geminata var. nigra Forel, 1908b: 45 (w.) COSTA RICA. Junior synonym of geminata: Creighton, 1930b: 59.
    - eduardi. Solenopsis eduardi Forel, 1912g: 12 (w.) COLOMBIA. Subspecies of geminata and senior synonym of perversa: Creighton, 1930b: 67. Junior synonym of geminata: Trager, 1991: 163.
    - innota. Solenopsis geminata var. innota Santschi, 1915c: 257, fig. 6 (s.w.q.m.) GABON. Junior synonym of geminata: Wheeler, W.M. 1922a: 877.
    - medusa. Solenopsis geminata subsp. medusa Mann, 1916: 447, pl. 4, fig. 31 (w.) BRAZIL. Creighton, 1930b: 69 (q.). Senior synonym of bahiaensis: Creighton, 1930b: 68. Junior synonym of geminata: Trager, 1991: 163.
    - galapageia. Solenopsis geminata var. galapageia Wheeler, W.M. 1919c: 272 (w.q.) ECUADOR (Galapagos Is). Subspecies of geminata: Creighton, 1930b: 65. Junior synonym of geminata: Trager, 1991: 163.
    - bahiaensis. Solenopsis edouardi var. bahiaensis Santschi, 1925d: 237 (w.) BRAZIL. Junior synonym of medusa: Creighton, 1930b: 68.
    - perversa. Solenopsis edouardi var. perversa Santschi, 1925b: 13 (s.w.q.) BRAZIL. Junior synonym of eduardi: Creighton, 1930b: 67.


    Myrmica paleata Lund, 1831a.


    - paleata. Myrmica paleata Lund, 1831a: 116 (footnote) (w.) BRAZIL. Junior synonym of geminata: Roger, 1863b: 32.


    Solenopsis mandibularis Westwood, 1840b.


    - mandibularis. Solenopsis mandibularis Westwood, 1840b: 87, pl. 2, fig. 5 (w.) "America aequinoctiali". Junior synonym of geminata: Roger, 1862c: 289.


    Atta rufa Jerdon, 1851.


    - rufa. Atta rufa Jerdon, 1851: 106 (w.q.) INDIA. Bingham, 1903: 158 (m.). Combination in Solenopsis: Emery, 1892b: 166. Junior synonym of geminata: Dalla Torre, 1893: 76. Revived from synonymy as subspecies of geminata: Forel, 1899a: 119; Wheeler, W.M. 1915b: 397; Creighton, 1930b: 66. Senior synonym of diabola: Creighton, 1930b: 66. Junior synonym of geminata: Ettershank, 1966: 141.


    Atta clypeata Smith, 1858b.


    - clypeata. clypeata. Atta clypeata Smith, F. 1858b: 169 (q.m.) MEXICO. Junior synonym of geminata: Roger, 1862c: 289.: 169 (q.m.) MEXICO. Junior synonym of geminata: Roger, 1862c: 289.


    Solenopsis cephalotes Smith, F. 1859a.


    - cephalotes. Solenopsis cephalotes Smith, F. 1859a: 149 (s.w.) INDONESIA (Aru I.). Junior synonym of geminata: Roger, 1863b: 32; of rufa: Creighton, 1950a: 231; of geminata: Trager, 1991: 163.


    Myrmica mellea Smith, F. 1859a.


    - mellea. Myrmica mellea Smith, F. 1859a: 148 (w.) INDONESIA (Aru I.). Junior synonym of geminata: Donisthorpe, 1932c: 455.


    Crematogaster laboriosus Smith, F. 1860b.


    - laboriosus. Crematogaster laboriosus Smith, F. 1860b: 109 (q.) INDONESIA (Batjan I.). Combination in Solenopsis: Mayr, 1863: 453. Junior synonym of rufa: Creighton, 1950a: 231; of geminata: Trager, 1991: 163.


    Myrmica laevissima Smith, F. 1860b


    - laevissima. Myrmica laevissima Smith, F. 1860b: 108 (w.) INDONESIA (Batjan I.). Junior synonym of geminata: Donisthorpe, 1932c: 463.


    Diplorhoptrum drewseni Mayr, 1861.


    - drewseni. Diplorhoptrum drewseni Mayr, 1861: 73 (diagnosis in key) (w.) ITALY. Junior synonym of geminata: Mayr, 1870b: 996; Mayr, 1886d: 460.


    Myrmica glaber Smith, F. 1862b.


    - glaber. Myrmica glaber Smith, F. 1862b: 34 (w.) PANAMA. Junior synonym of geminata: Mayr, 1886c: 362; Trager, 1991: 163.


    Myrmica polita Smith, F. 1862b.


    - polita. Myrmica polita Smith, F. 1862b: 34 (w.) PANAMA. [Unresolved junior primary homonym of Myrmica polita Smith, F. 1860b: 108 (now in Pheidologeton).] Junior synonym of geminata: Mayr, 1863: 453; Mayr, 1886c: 362.


    Atta coloradensis Buckley, 1867.


    - coloradensis. Atta coloradensis Buckley, 1867: 346 (s.w.) U.S.A. Junior synonym of geminata: Mayr, 1886c: 365.


    Atta lincecumii Buckley, 1867.


    - lincecumii. Atta lincecumii Buckley, 1867: 344 (s.w.) U.S.A. Junior synonym of geminata: Emery, 1895c: 276.


    Myrmica (Monomorium) saxicola Buckley, 1867.


    - saxicola. Myrmica (Monomorium) saxicola Buckley, 1867: 341 (w.) U.S.A. Junior synonym of geminata: Emery, 1895c: 276.


    Solenopsis geminata var. diabola Wheeler, W.M. 1908e.


    - diabola. Solenopsis geminata var. diabola Wheeler, W.M. 1908e: 424 (w.q.m.) U.S.A. Junior synonym of rufa: Creighton, 1930b: 66.


    Solenopsis geminata var. nigra Forel, 1908b.


    - nigra. Solenopsis geminata var. nigra Forel, 1908b: 45 (w.) COSTA RICA. Junior synonym of geminata: Creighton, 1930b: 59.


    Solenopsis eduardi Forel, 1912g.


    - eduardi. Solenopsis eduardi Forel, 1912g: 12 (w.) COLOMBIA. Subspecies of geminata and senior synonym of perversa: Creighton, 1930b: 67. Junior synonym of geminata: Trager, 1991: 163.


    Solenopsis geminata var. innota Santschi, 1915c.


    - innota. Solenopsis geminata var. innota Santschi, 1915c: 257, fig. 6 (s.w.q.m.) GABON. Junior synonym of geminata: Wheeler, W.M. 1922a: 877.


    Solenopsis geminata subsp. medusa Mann, 1916.


    - medusa. Solenopsis geminata subsp. medusa Mann, 1916: 447, pl. 4, fig. 31 (w.) BRAZIL. Creighton, 1930b: 69 (q.). Senior synonym of bahiaensis: Creighton, 1930b: 68. Junior synonym of geminata: Trager, 1991: 163.


    Solenopsis geminata var. galapageia Wheeler, W.M. 1919c.


    - galapageia. Solenopsis geminata var. galapageia Wheeler, W.M. 1919c: 272 (w.q.) ECUADOR (Galapagos Is). Subspecies of geminata: Creighton, 1930b: 65. Junior synonym of geminata: Trager, 1991: 163.


    Solenopsis edouardi var. bahiaensis Santschi, 1925d.


    - bahiaensis. Solenopsis edouardi var. bahiaensis Santschi, 1925d: 237 (w.) BRAZIL. Junior synonym of medusa: Creighton, 1930b: 68.


    Solenopsis edouardi var. perversa Santschi, 1925b.


    - perversa. Solenopsis edouardi var. perversa Santschi, 1925b: 13 (s.w.q.) BRAZIL. Junior synonym of eduardi: Creighton, 1930b: 67.


    Solenopsis geminata var. micans Stitz, 1912.


    - micans. Solenopsis geminata var. micans Stitz, 1912: 506, fig. 8 (w.) INDONESIA (Seram I.).


    Labauchena hostilis Borgmeier, 1959a.


    - hostilis. Labauchena hostilis Borgmeier, 1959a: 318, figs. 10, 11 (q.) BRAZIL. Combination in Solenopsis: Ettershank, 1966: 141.


    Solenopsis saevissima var. interrupta Santschi, 1916e.


    - interrupta. Solenopsis saevissima var. interrupta Santschi, 1916e: 379 (w.) ARGENTINA. Subspecies of saevissima: Creighton, 1930b: 89. Raised to species: Wilson, 1952b: 51. See also: Buren, 1972: 22; Trager, 1991: 178.

    Trager Pitts Pitts et al.
    amblychila xxx
    aurea xxx
    bruesi xxx
    daguerrei xxx xxx
    electra xxx xxx xxx
    gayi xxx
    geminata xxx
    geminata var. micans
    hostilis xxx xxx
    interrupta xxx xxx xxx
    invicta xxx xxx xxx
    macdonaghi xxx xxx xxx
    megergates xxx xxx xxx
    altipunctata / metallica xxx xxx
    pusillignis xxx xxx xxx
    pythia xxx xxx xxx
    quinquecuspis xxx xxx xxx
    richteri xxx xxx xxx
    saevissima xxx xxx xxx
    pylades var. itinerans
    substituta xxx
    tridens xxx
    tridens var. lehmannnitschei
    virulens xxx
    weyrauchi xxx xxx xxx
    xyloni xxx

    Solenopsis aurea subsp. amblychila Wheeler, W.M. 1915b.


    - amblychila. Solenopsis aurea subsp. amblychila Wheeler, W.M. 1915b: 394 (w.q.m.) U.S.A. Subspecies of xyloni: Creighton, 1930b: 104; of aurea: Creighton, 1950a: 230. Junior synonym of aurea: Snelling, R.R. 1963: 7; Wheeler, G.C. & Wheeler, J. 1986g: 13. Revived from synonymy and raised to species: Trager, 1991: 168.


    Solenopsis geminata var. aurea Wheeler, W.M. 1906d.


    - aurea. Solenopsis geminata var. aurea Wheeler, W.M. 1906d: 336 (w.q.m.) U.S.A. Crozier, 1970: 116 (k.). Raised to species: Forel, 1909a: 269. Subspecies of xyloni: Creighton, 1930b: 103. Revived status as species: Creighton, 1950a: 230; Snelling, R.R. 1963: 6. Senior synonym of huachucana: Trager, 1991: 170.
    - huachucana. Solenopsis huachucana Wheeler, W.M. 1915b: 393 (w.q.) U.S.A. Junior synonym of aurea: Trager, 1991: 170.


    Solenopsis huachucana Wheeler, W.M. 1915b.


    - huachucana. Solenopsis huachucana Wheeler, W.M. 1915b: 393 (w.q.) U.S.A. Junior synonym of aurea: Trager, 1991: 170.


    Solenopsis (Solenopsis) gayi subsp. bruesi Creighton, 1930b.


    - bruesi. Solenopsis (Solenopsis) gayi subsp. bruesi Creighton, 1930b: 52, pl. 4, figs. 4, 5 (w.) PERU. Raised to species: Trager, 1991: 172.


    Labauchena daguerrei Santschi, 1930d.


    - daguerrei. Labauchena daguerrei Santschi, 1930d: 81, fig. 12 (q.m.) ARGENTINA. Combination in Solenopsis: Ettershank, 1966: 140. Senior synonym of acuminata: Borgmeier, 1959a: 318.
    - acuminata. Labauchena acuminata Borgmeier, 1949: 208, figs. 10-14 (w.) ARGENTINA. Junior synonym of daguerrei: Borgmeier, 1959a: 318.


    Labauchena acuminata Borgmeier, 1949.


    - acuminata. Labauchena acuminata Borgmeier, 1949: 208, figs. 10-14 (w.) ARGENTINA. Junior synonym of daguerrei: Borgmeier, 1959a: 318.


    Solenopsis pylades r. electra Forel, 1914d.


    - electra. Solenopsis pylades r. electra Forel, 1914d: 275 (w.) ARGENTINA. Creighton, 1930b: 93 (q.). Subspecies of geminata: Wheeler, W.M. 1915b: 397; of saevissima: Santschi, 1923c: 266; Santschi, 1929d: 299; Creighton, 1930b: 92. Junior synonym of saevissima: Wilson, 1952b: 55. Revived from synonymy and raised to species: Trager, 1991: 192.


    Myrmica gayi Spinola, 1851a.


    - gayi. Myrmica gayi Spinola, 1851a: 242 (w.) CHILE. Combination in Solenopsis: Mayr, 1886c: 365. Junior synonym of geminata: Mayr, 1886d: 460; Dalla Torre, 1893: 76. Revived from synonymy as subspecies of geminata: Emery, 1895i: 11; Emery, 1906c: 121. Revived status as species: Forel, 1909a: 269; Santschi, 1923c: 261; Creighton, 1930b: 48. Senior synonym of fazi: Brown, 1950e: 248. See also: Trager, 1991: 170.
    - fazi. Solenopsis gayi var. fazi Santschi, 1923c: 261 (w.) CHILE. Creighton, 1930b: 51 (q.). Junior synonym of gayi: Brown, 1950e: 248.


    Solenopsis gayi var. fazi Santschi, 1923c


    - fazi. Solenopsis gayi var. fazi Santschi, 1923c: 261 (w.) CHILE. Creighton, 1930b: 51 (q.). Junior synonym of gayi: Brown, 1950e: 248.

    Maybe, just like Teleutomyrmex, we should consider A. atratulus Schenck, 1852 not a member of Tetramorium, so T. atratulum (Schenck, 1852), but a member of Strongylognathus, so S. atratulus (Schenck, 1852).


    But then you also have S. microgyna (Santschi, 1918) and S. parasiticum (Bolton, 1980)!