The extreme, workerless inquilines.

The extreme, workerless inquilines.

Teleutomyrmex Kutter, 1950

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

Not yet described species of extreme, workerless inquiline, from the genus Teleutomyrmex Kutter, 1950

05) The new, undescribed species from Teleutomyrmex Kutter, 1950 from Farab, Turkmenistan… See Dlussky, Soyunov, Zabelin, 1990 [“1989”].

Anergates Forel, 1874

06) Anergates atratulus (Schenck, 1852), by Forel, 1874
(= Myrmica atratula Schenck, 1852)
[Also described as new by Schenck, 1853b]
(= Tetramorium atratulum (Schenck, 1852), by Mayr, 1855)
[= Tomognathus atratulus (Schenck, 1852), by Mayr, 1863 following Mayr, 1861, obsolete combination.]
(= Tetramorium atratulum (Schenck, 1852), by Ward, Brady, Fisher & Schultz, 2015 ("2014"))
07) Anergates friedlandi Creighton, 1934
[= Tetramorium friedlandi (Creighton, 1934), by analogy]

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), by Wilson, 1984)
(= Anergatides kohli Wasmann, 1915)
(not Pheidole kohli Mayr, 1901)
11) Pheidole acutidens (Santschi, 1922), by Wilson, 1984
(= Bruchomyrma acutidens Santschi, 1922)
12) Pheidole argentina (Bruch, 1932), by Wilson, 1984
(= 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 Westwood, 1839.

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

14) Pheidole kusnezovi Wilson, 2003
(= Pheidole symbiotica (Kusnezov, 1952), by Wilson, 1984)
(= 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, T. staerckei Kratochvíl, in Kratochvíl, Novák, Šnoflák, 1944
and T. moravicum [Kratochvil, in] Novák & Sadil, 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. strobeli Emery, 1906
12) P. nitidula Emery, 1888
13) P. indica Mayr, 1879

14) P. obscurior 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)

- T. staerckei Kratochvíl, in Kratochvíl, Novák, Šnoflák, 1944 (= Tetramorium caespitum (Linnaeus, 1758) subsp. hungarica Röszler, 1935 ("1933-34") var. staerckei Röszler, 1936)
- 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
- The host species P. strobeli Emery, 1906 was known, during much of its history, as a subspecies of an other species, nl. P. nitidula Emery, 1888 subsp. richteri Forel, 1909 (a much used junior synonym of the older P. perversa Forel, 1908 subsp. richteri Forel, 1909) and known, in 1922, for a while as P. strobeli Emery, 1906 subsp. richteri Forel, 1909, this since Santschi, 1916. The species P. strobeli Emery, 1906 was revived from synonymy (with P. nitidula Emery, 1888 since Wilson, 2003) and the subspecies was synonymized with it in Casadei-Ferreira, Economo, Feitosa, 2020.
- P. obscurior Forel, 1886 was revived from synonymy with the 1 page older P. susannae Forel, 1886 and this in Casadei-Ferreira et al., 2020.

A new host and observations about copulation and adoption in Tetramorium-Strongylognathus nests.

Purkart, A., Wagner, H.CPurkart, A., Wagner, H.C., Goffová, K., Selnekovič, D., Holecová, M. 2021. Laboratory observations on Anergates atratulus (Schenck, 1852): mating behaviour, incorporation into host colonies, and competition with Strongylognathus testaceus (Schenck, 1852). Biologia.,

https://www.antwiki.org/wiki/i…s11756-021-00901-y%29.pdf

“Emery’s Rule” in its strict and its loose version.

In 1909, the entomologist/taxonomist Carlo Emery made an important generalization and noted that social parasites among insects and their hosts share common ancestry and hence tend to be parasites of species or genera to which they are closely related. Over time, this pattern has been recognized in many additional cases, and generalized to what is now known as “Emery's rule”. The pattern is best known for various taxa of Hymenoptera. The significance and general relevance of this pattern are still a matter of some debate, as a great many exceptions exist, though a common explanation for the phenomenon when it occurs is that the parasites may have started as facultative parasites within the host species itself (such forms of intraspecific parasitism are well-known.), but later became reproductively isolated and split off from the ancestral species, a form of sympatric speciation.

In the “strict version” of Emery’s rule, social parasitic species are their host’s closest relatives (a sister taxon to its host in a phylogenetic sense.), and likely evolved from their host’s lineage by sympatric speciation, or through a combination of allopatric and subsequent sympatric speciation. In the “loose version” of Emery’s rule, social parasites are close relatives of their host, but not sister species. Although few host-parasite pairs have been subjected to molecular phylogenetic analysis, studies to date support at least the loose version of Emery’s rule.

The “Kutter-Wilson Paradox”.

There are only about 230 known parasitic species of ants among the 12,500 or so described ant species. Despite their rarity, they are common in a few subfamilies like the Myrmicinae and the Formicinae, and common in temperate ants but rare in tropical ants. Why is there such a strong bias in both the taxonomic and ecological distributions of social parasites and their hosts, the “Kutter-Wilson Paradox”? The selective forces and ecological conditions that favour social parasitism continue to be researched and discussed. Factors like cooler temperatures and polygyny (i.e., colonies with multiple queens.) are important considerations, but cannot explain the evolution of social parasitism in all cases.

The “Inquiline Syndrome”.

Wilson, E. O. [“The insect societies”, 1971, p. 374] and Hölldobler, B. K. and Wilson, E. O. [“The ants”, 1990, p. 467] defined the “Inquiline Syndrome” and listed 41 characters of morphological, behavioural and life history traits that evolved convergently in inquiline social parasites (a 19 points of special interesses-list.). 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.

Reference:

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

The new list:

Including this:

- T. impurum (Förster, 1850) (= Myrmica impura Förster, 1850)
- T. caespitum (Linnaeus, 1758) (= Formica caespitum Linnaeus, 1758)
- T. chefketi Forel, 1911 (= T. caespitum (Linnaeus, 1758) var. chefketi Forel, 1911)
- T. immigrans Santschi, 1927 (= T. caespitum (Linnaeus, 1758) var. immigrans Santschi, 1927)
- T. staerckei Kratochvíl, in Kratochvíl, Novák, Šnoflák, 1944 (= T. caespitum (Linnaeus, 1758) subsp. hungarica Röszler, 1935 ("1933-34") var. staerckei Röszler, 1936)
- T. diomedeum Emery, 1908 (= T. caespitum (Linnaeus, 1758) var. diomedea Emery, 1908)
- T. sepositum Santschi, 1918 (= T. gladstonei Forel, 1913 var. seposita Santschi, 1918)
- P. megacephala (Fabricius, 1793) (= Formica megecephala Fabricius, 1793) (= Formica edax Forskål, 1775, a nomen oblitum under Art. 23.9 of ICZN (1999))
- P. megacephala (Fabricius, 1793) subsp. melancholica Santschi, 1912 was originally described as P. punctulata Mayr, 1866 st. melancholica Santschi, 1912
- P. strobeli Emery, 1906 (= P. perversa Forel, 1908 subsp. richteri Forel, 1909, or, in 1922, at the moment the extreme, workerless inquiline species was described, = P. strobeli Emery, 1906 subsp. richteri Forel, 1909.)
- P. nitidula Emery, 1888 (= P. triconstricta Forel, 1886 var. nitidula Emery, 1888)
- P. obscurior Forel, 1886 (= P. susannae Forel, 1886 r. obscurior Forel, 1886)

The Extreme, Workerless Inquilines of the World.

The inquiline species.

The Tetramorium inquilinum species-group (“Degenerate workerless social parasites of several other Tetramorium species”, complete.) Before as Tetramorium Mayr, 1855 (Only a few species in a big genus.) (= Teleutomyrmex Kutter, 1950, by Ward, Brady, Fisher, Schultz, 2015 (“2014”), the old genus, complete.).

01) Tetramorium inquilinum Ward, Brady, Fisher, Schultz, 2015 (“2014”)

(= Teleutomyrmex schneideri Kutter, 1950)

(= Tetramorium schneideri (Kutter, 1950), by Ward, Brady, Fisher, Schultz, 2015 ("2014"))

(not Tetramorium schneideri Emery, 1898)

(= Tetramorium inquilinum Ward, Brady, Fisher, Schultz, 2015 ("2014"), replacement name)

02) Tetramorium kutteri (Tinaut, 1990)

(= Teleutomyrmex kutteri Tinaut, 1990)

(= Tetramorium kutteri (Tinaut, 1990), by Ward, Brady, Fisher, Schultz, 2015 ("2014"))

(not Tetramorium semilaeve André, 1883 var. kutteri Santschi, 1927)

03) Tetramorium seiferti (Kiran, Karaman, 2017, in Kiran, Karaman, Lapeva-Gjonova, Aksoy, 2017)

(= Teleutomyrmex seiferti Kiran, Karaman, 2017, in Kiran, Karaman, Lapeva-Gjonova, Aksoy, 2017)

(= Tetramorium seiferti (Kiran, Karaman, 2017, in Kiran, Karaman, Lapeva-Gjonova, Aksoy, 2017), by analogy)

04) Tetramorium buschingeri (Lapeva-Gjonova, 2017, in Kiran, Karaman, Lapeva-Gjonova, Aksoy, 2017)

(= Teleutomyrmex buschingeri Lapeva-Gjonova, 2017, in Kiran, Karaman, Lapeva-Gjonova, Aksoy, 2017)

(= Tetramorium buschingeri (Lapeva-Gjonova, 2017, in Kiran, Karaman, Lapeva-Gjonova, Aksoy, 2017), by analogy)

05) Tetramorium albenae Salata, van Delft, Borowiec, 2023, in Salata, van Delft, van Delft, Georgiadis, Borowiec, 2023

Not yet described species of extreme, workerless inquiline, from the Tetramorium inquilinum species-group (= Teleutomyrmex Kutter, 1950.).

06) The new, undescribed species from Tetramorium Mayr, 1855 (= Teleutomyrmex Kutter, 1950) from Farab, Turkmenistan… See

Dlussky, Soyunov, Zabelin, 1990 [“1989”].

Tetramorium Mayr, 1855 (Only a few species in a big genus.) (= Anergates Forel, 1874, by Ward, Brady, Fisher, Schultz, 2015 (“2014”), the old genus, complete.).

07) Tetramorium atratulum (Schenck, 1852)

(= Myrmica atratula Schenck, 1852)

[Also described as new by Schenck, 1853]

(= Tetramorium atratulum (Schenck, 1852), by Mayr, 1855)

[= Tomognathus atratulus (Schenck, 1852), by Mayr, 1863 following Mayr, 1861, obsolete combination.]

(= Anergates atratulus (Schenck, 1852), by Forel, 1874)

(= Tetramorium atratulum (Schenck, 1852), by Ward, Brady, Fisher, Schultz, 2015 ("2014"))

08) Tetramorium friedlandi (Creighton, 1934)

(= Anergates friedlandi Creighton, 1934)

[= Tetramorium friedlandi (Creighton, 1934), by analogy]

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

09) Tetramorium microgyna Santschi, 1918

10) Tetramorium parasiticum Bolton, 1980

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

11) Pheidole neokohli Wilson, 1984

(= Anergatides kohli Wasmann, 1915)

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

(not Pheidole kohli Mayr, 1901)

(= Pheidole neokohli Wilson, 1984, replacement name)

12) Pheidole acutidens (Santschi, 1922)

(= Bruchomyrma acutidens Santschi, 1922)

(= Pheidole acutidens (Santschi, 1922), by Wilson, 1984)

13) Pheidole argentina (Bruch, 1932)

(= Gallardomyrma argentina Bruch, 1932)

(= Pheidole argentina (Bruch, 1932), by Wilson, 1984)

14) 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 Westwood, 1839.

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

15) Pheidole kusnezovi Wilson, 2003

(= Eriopheidole symbiotica Kusnezov, 1952)

(= Pheidole symbiotica (Kusnezov, 1952), by Wilson, 1984)

(not Pheidole symbiotica Wasmann, 1909)

(= Pheidole kusnezovi Wilson, 2003, replacement name)

Distribution.

01) Europe (Alps, Pyrenees and Northern Spain)

02) Europe (Southern Iberia)

03) Turkey (Anatolia)

04) Europe (Southern Balkans or, more precisely, Bulgaria)

05) Europe (Balkan Peninsula or, more precisely, Greece)

06) Turkmenistan

07) Palaearctic region (most important: Europe)

08) North America

09) Southern Africa

10) Southern Africa

11) Central Africa

12) South America

13) South America

14) India

15) South America

The host species.

01), 02), 03), 04), 05), 06), 07), 08), 09) and 10) 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) T. kephalosi Salata, Borowiec, 2017

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

07) T. impurum (Förster, 1850), T. caespitum (Linnaeus, 1758), T. immigrans Santschi, 1927, T. staerckei Kratochvíl, 1944, in

Kratochvíl, Novák, Šnoflák, 1944, T. moravicum [Kratochvil, 1941, in] Novák, Sadil, 1941, T. diomedeum Emery, 1908, T. chefketi

Forel, 1911

08) T. immigrans Santschi, 1927

09) T. sericeiventre Emery, 1877 and T. sepositum Santschi, 1918

10) T. avium Bolton, 1980

11), 12), 13), 14) and 15) Certain species of the genus Pheidole Westwood, 1839.

11) P. megacephala (Fabricius, 1793) subsp. melancholica Santschi, 1912

12) P. strobeli Emery, 1906

13) P. nitidula Emery, 1888

14) P. indica Mayr, 1879

15) P. obscurior Forel, 1886

A remark about synonymy.

Tetramorium friedlandi (Creighton, 1934) is now a synonym from Tetramorium atratulum (Schenck, 1852), more precisely an introduced form in North America (see also Schär, Talavera, Espadaler, Rana, Andersen, Cover, Vila, 2018.). This synonymy was given by Creighton, 1950. So, the name is Tetramorium atratulum (Schenck, 1852)...

Synonyms of the host species.

- T. impurum (Förster, 1850) (= Myrmica impura Förster, 1850)

- T. caespitum (Linnaeus, 1758) (= Formica caespitum Linnaeus, 1758)

- T. chefketi Forel, 1911 (= T. caespitum (Linnaeus, 1758) var. chefketi Forel, 1911)

- T. immigrans Santschi, 1927 (= T. caespitum (Linnaeus, 1758) var. immigrans Santschi, 1927)

- T. staerckei Kratochvíl, 1944, in Kratochvíl, Novák, Šnoflák, 1944 (= T. caespitum (Linnaeus, 1758) subsp. hungarica Röszler, 1935

("1933-34") var. staerckei Röszler, 1936)

- T. diomedeum Emery, 1908 (= T. caespitum (Linnaeus, 1758) var. diomedea Emery, 1908)

- T. sepositum Santschi, 1918 (= T. gladstonei Forel, 1913 var. seposita Santschi, 1918)

- P. megacephala (Fabricius, 1793) (= Formica megecephala Fabricius, 1793) (= Formica edax Forskål, 1775, a nomen oblitum under

Art. 23.9 of ICZN (1999))

- P. megacephala (Fabricius, 1793) subsp. melancholica Santschi, 1912 was originally described as P. punctulata Mayr, 1866 st.

melancholica Santschi, 1912

- P. strobeli Emery, 1906 (= P. perversa Forel, 1908 subsp. richteri Forel, 1909, or, in 1922, at the moment the extreme, workerless

inquiline species was described, = P. strobeli Emery, 1906 subsp. richteri Forel, 1909.)

- P. nitidula Emery, 1888 (= P. triconstricta Forel, 1886 var. nitidula Emery, 1888)

- P. obscurior Forel, 1886 (= P. susannae Forel, 1886 r. obscurior Forel, 1886)

And then...

…, if you follow the line further that Ward, Brady, Fisher, Schultz, 2015 (“2014”) outlined, the first 10 extreme, workerless inquilines become a few species in the genus Strongylognathus Mayr, 1853.

01) Strongylognathus inquilinum (Ward, Brady, Fisher, Schultz, 2015 (“2014”))

(= Strongylognathus schneideri (Kutter, 1950))

02) Strongylognathus kutteri (Tinaut, 1990)

03) Strongylognathus seiferti (Kiran, Karaman, 2017, in Kiran, Karaman, Lapeva-Gjonova, Aksoy, 2017)

04) Strongylognathus buschingeri (Lapeva-Gjonova, 2017, in Kiran, Karaman, Lapeva-Gjonova, Aksoy, 2017)

05) Strongylognathus albenae (Salata, van Delft, Borowiec, 2023, in Salata, van Delft, van Delft, Georgiadis, Borowiec, 2023)

07) Strongylognathus atratulus (Schenck, 1852)

08) Strongylognathus friedlandi (Creighton, 1934)

09) Strongylognathus microgyna (Santschi, 1918)

10) Strongylognathus parasiticum (Bolton, 1980)

If you followed the systematics of ants in 2014-2015, you noticed that all the species of Teleutomyrmex Kutter, 1950 and Anergates Forel, 1874 became Tetramorium Mayr, 1855. But normally, they should have the name Strongylognathus Mayr, 1853. No, they, Ward, Brady, Fisher, Schultz, 2015 (“2014”), didn't like that! So, until the ICZN would say different, they kept Tetramorium Mayr, 1855.

And now you find under a few species, described in the "normal" genus “Teleutomyrmex Kutter, 1950”, this:

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

But they keep themselves a paraphyletic genus, nl. Tetramorium Mayr, 1855! So it should be Strongylognathus Mayr, 1853...