An introduction to Teleutomyrmex.....

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    • Strongylognathus Mayr, 1853d.

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

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

      Myrmus Schenck, 1853.

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

      Some know why I putted this one here..............

      The post was edited 1 time, last by Teleutotje ().

    • Now, let me explain. It was shown by Ward et al. 2015 ("2014") that Teleutomyrmex and Anergates should be synonymized under Tetramorium but, surprisingly, they kept Strongylognathus out of it. It should be a synonym of Tetramorium but the name should not be Strongylognathus but Tetramorium ("better name'). But if you do it, do it good and synonymize it like it should... so: Strongylognathus.

      Tetramorium Mayr, 1855.

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

      The post was edited 1 time, last by Teleutotje ().

    • In 1951 Walter Linsenmaier made a crude black-and-white drawing of three Teleutomyrmex schneideri females riding on the back of a Tetramorium-queen. In front of this group, a Tetramorium-worker stands alongside her queen. This drawing was published in the 1951-article that Robert Stumper wrote about the life-style of the Teleutomyrmex schneideri-queen.

      In 1951 a color-version of this drawing was also made by Walter Linsenmaier for an article in "Du: kulturelle Monatsschrift" by Heinrich Kutter, “Von dufttäuschenden Mörderinnen und berittener Königin.”. Heinrich Kutter used it again in 1968 as the front-plate of his little book about social parasitic ants of Switzerland. Later Walter Linsenmaier used it in his general review-book of insects of the world, published in 1973/4.

      In 1977 a better version of the black-and-white drawing appeared in Heinrich Kutter’s determination-book for Swiss ants. This version is the most-copied drawing when Teleutomyrmex is depicted in an article or a book.

      And on June 12, 2011 a colored version of the black-and-white/color-version drawing appeared as a tattoo on the back of my right shoulder… 60 years after its first publication!

      The post was edited 1 time, last by Teleutotje ().

      Boletín de la Sociedad Entomológica Aragonesa (S.E.A.), nº 63 (31/12/2018): 235–239.…ca_Hymenoptera_Formicidae

      (Thank you Merkur!)
    • Philippe WEGNEZ, David IGNACE, Els LOMMELEN, Maximilien HARDY, Johan BOGAERT & Carin NILSSON (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, 151: 52-57.…Alpes_fran%C3%A7aises.pdf
    • From: AntWiki - Where Ant Biologists Share Their Knowledge.

      Hölldobler, B. and 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. and 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.