http://www.collembola.org/taxa/tullidae.htm - Last updated on 2017.02.22 by Frans Janssens
Checklist of the Collembola: Tullbergiidae

Familia Tullbergiidae Bagnall RS, 1935:238 key  key  key
Genus Number
of species
Ameritulla3
Anaphorura1
Boudinotia1
Clavaphorura1
Delamarephorura5
Dinaphorura16
Doutnacia4
Fissuraphorura7
Jevania2
Karlstejnia5
Marcuzziella1
Mesaphorura56
Metaphorura9
Mexicaphorura1
Mixturatulla1
Multivesicula6
Najtiaphorura4
Neonaphorura14
Neotullbergia5
Paratullbergia8
Pongeiella3
Prabhergia3
Psammophorura3
Rotundiphorura1
Scaphaphorura5
Sensilatullbergia1
Stenaphorura8
Stenaphorura21
Tasphorura1
Tillieria3
Tullbergia43
Tullbergiella2
Wankeliella6
Weinera1
Tullbergiidae231
Genera with number of species
  Syn.: Onychiuridae Tullberginae(sic) Salmon, JT, 1943:377 ISS
  Syn.: Onychiuridae Tullberginae(sic) Palacios-Vargas, JG et Díaz, M, 1996:166 ISS
  Syn.: Onychiuridae Tullberginae(sic) Skidmore, RE, 1995:53 ISS
  Syn.: Onychiuridae Tullberginae(sic) Mari Mutt, JA et Bellinger, PF, 1996:169 ISS
  Syn.: Onychiuridae Tullberginae(sic) González, V, Díaz, M et Palacios-Vargas, JG, 1997:46 ISS

Tullbergiidae from the USA
2012.02.16 © Kouri, J.
Tullbergiidae from the UK
2012.10.02 © Murray, A.

  Syn.: Collembola Tuubergiinae(sic) Kaprus, IJ et Goblyk, KM, 2015:8 ISS

Subfamilia Clavaphorurinae
Remark by Janssens, F. 2012.11.04: For genera with 4 sensory clubs in the ant.3 sense organ we tentatively propose the subfamily Clavaphorurinae with type genus Clavaphorura Salmon, 1943.


Genus Clavaphorura Salmon, JT, 1943:377
Species septemseta Salmon, JT, 1943:377, t.t.
Clavaphorura septemseta from New Zealand
After Salmon JT, 1943 Pl.41 Fig.22
            Syn.: Tullbergia septemspina Schaller, 1949 nn


Subfamilia Tullbergiinae Deharveng L, 2004:427
  Syn.: Tullbergiinae Group I Bagnall, RS, 1935:240
  Syn.: Tullbergiinae Tullbergiini Bagnall, 1935

Genus Tullbergia Lubbock, 1876
  Syn.: ... Börneria(sic) Willem, 1902 IOS
  Syn.: Tullbergiidae Protullbergia Bagnall, 1947
  Syn.: Tullbergiinae Bachatullbergia Wray, 1958
  Syn.: Tullberginae Chaetophorura Rusek, 1976
  Syn.: Tullbergiinae Granuliphorura Rusek, 1976
Tullbergia sp. from Tasmania
Post-antennal organ and pseudocellus
After Ireson JE, 1990 Fig.70
Tullbergia sp. from Tasmania
Ant.3 sensory organ
After Ireson JE, 1990 Fig.71

  Syn.: Onychiuridae (Tullbergiinae) Tullberia(sic) Greenslade, P et Rusek, J, 1996:33 ISS
Tullbergia sp. from Antarctica
2003 © Stevens, M.
Remark by ICZN. 2007:841: Placed on the Official List of Names in Zoology. Opinion 291.

Tullbergia? sp. from Tasmania
2014.04.04 © Murray, A.
Tullbergia? sp. from Tasmania
Anal spines
2014.04.04 © Murray, A.

Species africana Martynova, 1979
Species alcirae Palacios-Vargas, JG et Salazar-Martínez, AE, 2014:24
Species anops Christiansen, K et Bellinger, P, 1980:482
Species antarctica Lubbock, 1876, t.t.
Tullbergia antarctica from Antarctica
After Lubbock J, 1879 Pl.XI Fig.1

Species arctica Wahlgren, 1900
Species australica Womersley, 1933
Species bella Fjellberg, A, 1988:37
Tullbergia bella from Norway
After Fjellberg, A, 1988 Fig.2G

Species bisetosa Börner, 1903
            Syn.: Tullbergia insularis Womersley, H, 1935:212
            Syn.: Tullbergia insularis Salmon, JT, 1964:146
            Syn.: Tullbergia insularis Mari Mutt, JA et Bellinger, PF, 1990:55
            Syn.: Mesaphorura sp. Watson, 1967
Species collis Bacon, GA, 1914:84
Tullbergia collis from the USA
After Bacon GA, 1914 Fig.1
Tullbergia collis from the USA
After Bacon GA, 1914 Fig.2

Species crozetensis Deharveng, 1981
Species duops Christiansen, K et Bellinger, P, 1980:489
             Syn.: Tullbergia doups(sic) Cutz-Pool, LQ et Vázquez-González, M, 2012:109,ca ISS
Species gambiensis Womersley, H, 1935:211
            Syn.: Tullbergia gambiense(sic) Womersley, H, 1935:211 IOS
            Syn.: Tullbergia scalpellata Salmon, JT, 1949:17
            Syn.: Tullbergia subantarctica Salmon, JT, 1949:18
            Syn.: Tullbergia tillyardi Womersley, H, 1939:317
            Syn.: Tullbergia trisetosa Womersley, H, 1939:130
             Syn.: Tullbergia gambiense(sic) Womersley, 1939 ISS
             Syn.: Tullbergia gambiense(sic) Salmon, JT, 1964:146 ISS
             Syn.: Tullbergia gambiense(sic) Lawrence, 1968 ISS
             Syn.: Tullbergia gambiense(sic) Greenslade, P in Houston, WWK, 1994:62 ISS
             Syn.: Tullbergia gambiense(sic) Greenslade, P et Rusek, J, 1996:33 ISS
             Syn.: Tullbergia gambiense(sic) Najt, J et Weiner, WM in Najt, J et Matile, L, 1997:12 ISS
Species ghilarovi (Khanislamova, 1987) Dunger, W & Schlitt, B, 2011:144
Species harti (Rusek, 1991) Waltz & Hart, 1997
Species huetheri da Gama, 1968
            Syn.: Tullbergia hütheri(sic) da Gama, 1968 IOS
Species inconspicua Izarra, 1965
Species insularis Wahlgren, E, 1906:8
Species kilimanjarica (Delamare Deboutteville, 1953) Paclt, 1959
Species latens Christiansen, K et Bellinger, P, 1980:497
Species mala Christiansen, K et Bellinger, P, 1980:500
Species massoudi Hermosilla & Rubio, 1976, n.n.
Species maxima Deharveng, 1981
Species mediantarctica Wise, 1967
Species meridionalis Cassagnau & Rapoport, 1962
            Syn.: Tullbergia meridionalis f. bisetosa Cassagnau & Rapoport, 1962
Tullbergia meridionalis from Chile
After Rapoport EH & Rubio IR, 1963 Fig.73
Tullbergia meridionalis from Argentina
After Arbea JI, 2016 Fig.20-21

Species meridionalis2 Barra, J-A, 1995:132
Species mexicana Handschin, 1928
Species minensis Arlé, R, 1960:165
Species mixta Wahlgren, E, 1906:8
Species nearctica Bernard, EC, 2016:472
Tullbergia nearctica from the USA
After Bernard EC, 2016 Fig.13AB

Species nulla Christiansen, K et Bellinger, P, 1980:502
Species obtusochaeta (Rusek, 1976) Christiansen, K & Bellinger, P, 1980:504
             Syn.: Tullbergia obtusichaeta(sic) Bernard, EC, 2016:477 ISS
Species ouatilou Najt, J et Weiner, WM in Najt, J et Matile, L, 1997:11
Species paranensis Izarra, 1969
Species pomorskii (Smolis, 2010)
Species quadrisetosa (Willem, 1902) Börner, 1902
Börneria(sic) quadrisetosa from Antarctica
After Willem V, 1902 Pl.I Fig.1
             Syn.: Tullbergia quadriseta(sic) Womersley, H, 1935:212 ISS
Species rapoporti Arbea, JI, 2016:65
Tullbergia rapoporti from Argentina
After Arbea JI, 2016 Fig.1-2
Tullbergia rapoporti from Argentina
After Arbea JI, 2016 Fig.18

Species schaefferi Salmon, 1974
Species templei Wise, KAJ, 1970:210
Species tolanara Thibaud, J-M, 2008:510
Species trisetosa Schäffer, 1897
            Syn.: Aphorura trisetosa ssp. quadrisetosa Salmon, JT, 1964:146
             Syn.: Aphorura trisetosa ssp. quadriseta(sic) Womersley, H, 1935:212 ISS
            Syn.: Tullbergia tullbergi Womersley, 1939
Species vancouverica (Rusek, 1976) Christiansen, K & Bellinger, P, 1980:508
             Syn.: Tullbergia vancourverica(sic) Poole, RW in Poole, RW et Gentili, P, 1997:54 ISS
Species ventanensis Rapoport, 1963
Species womersleyi (Bagnall, 1947) Greenslade P in Houston, 1994:63

Genus Anaphorura Izarra, 1972
Species lavadoi (Izarra, 1972) Palacios-Vargas & Díaz, 1995:171, t.t.
             Syn.: Anaphorura lavadori(sic) Palacios-Vargas & Díaz, 1995 ISS

Genus Fissuraphorura Rusek, 1991
Species sp.1 Barra, J-A, 1998:526
Species cubanica Rusek, 1991
             Syn.: Fissuraphorura Cubanica(sic) Palacios-Vargas & Díaz, 1996 ISS
Species deharvengi Rusek, 1991
Species duplex (da Gama, 1962) Lucianez & Simon, 1992
Species gisini (Selga, 1963) Lucianez & Simon, 1992
Species miscellanea Barra, J-A, 1995:135
Species selgae (Prabhoo, 1971) Lucianez & Simon, 1992

Genus Tillieria Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:123 key
  Syn.: .. Tilleria(sic) Greenslade P, [1996] ISS
Species araucariensis Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:124
Species insularis Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:124, t.t.
Species penai Weiner & Najt, 1994

Genus Dinaphorura Bagnall, RS, 1935:241
  Syn.: Onychiuroidea (Tullbergiidae) Austraphorura Bagnall, 1947
  Syn.: Onychiuridae (Tullbergiinae) Astraphorura(sic) Rodgers, D et Greenslade, P, 1996:1370 ISS
Species americana Rapoport, 1962
Species chazeaui Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:126
Species diversispina Womersley, H, 1935:213
            Syn.: Dinaphorura harrisoni Bagnall, 1947
            Syn.: Tullbergia septemspina Schaller, 1949 nn =?
            Syn.: Protullbergia willemi Bagnall, 1947
Species harrisoni Bagnall, 1947
Species jarai Najt & Rubio, 1978
Species kerguelensis Deharveng, 1981

Dinaphorura laterospina from New Zealand
After Salmon, J.T. 1941 Pl.48 Fig.132.
Species laterospina Salmon, JT, 1941:311
Species magellanica Rubio, 1974
Species matileorum Najt, J in Tillier, S, 1988:29
Species najtae Arbea, JI, 2016:78
Dinaphorura najtae from Argentina
After Arbea JI, 2016 Fig.36-37

Species nerudai Arbea, JI, 2016:75
Dinaphorura nerudai from Argentina
After Arbea JI, 2016 Fig.27-28

Species novaezealandiae Womersley, H, 1935:213
            Syn.: Dinaphorura novae-zealandeae(sic) Womersley, H, 1935:213 IOS
             Syn.: Dinaphorura novae-zealandiae(sic) Salmon, JT, 1941:310 ISS
             Syn.: Dinaphorura novae-hollandiae(sic) Womersley ISS
             Syn.: Dinaphorura novae-hollandae(sic) Womersley ISS
             Syn.: Dinaphorura novae-zealandeae(sic) Salmon, JT, 1964:147 ISS
             Syn.: Dinaphorura nova-zealandiae(sic) Najt, J in Tillier, S, 1988:32 ISS
             Syn.: Dinaphorura novaezealandeae(sic) Rodgers, D et Greenslade, P, 1996:1369 ISS
Species pefauri Rubio & Najt, 1979
Species spinosissima (Wahlgren, E, 1906:9) Bagnall, RS, 1935:242, t.t.
            Syn.: Tullbergia septemspina Schaller, 1949 nn =?
            Syn.: Dinaphorura diversispina Greenslade, P, 2015:9 laps
Species tooheyensis Rodgers, D et Greenslade, P, 1996:1374
Species wahlgreni (Bagnall, 1947) Najt & Rubio, 1978, s.d.

Genus Neonaphorura Bagnall, RS, 1935:240 key  key
  Syn.: ... Tullbergia Denis, 1932
  Syn.: Tullbergia Bachatullbergia Wray, 1958
  Syn.: Tullbergiinae Bachytullbergia(sic) Salmon, JT, 1964:149 ISS
  Syn.: Tullbergiidae Spinaphorura Arbea, 1987 =i

Neonaphorura sp. from France
2006 © Boldsystems.org
Neonaphorura sp. from France
2006 © Boldsystems.org

Species adulta (Gisin, 1944) Salmon, JT, 1964:149
            Syn.: Tullbergia (Neonaphorura) duboscqi f. adulta Gisin, 1944
             Syn.: Neonaphorura adula(sic) Traser, G et Thibaud, J-M, 1999:106 ISS
Species alicatai Giuga & Jordana, 2013
Species ariegica Arbea, JI, 1991:179
Species duboscqi (Denis, 1932) Salmon, JT, 1964:149, t.t.
            Syn.: Neonaphorura anglicana Bagnall, 1936
             Syn.: Neonaphorura anglicanus(sic) Salmon, JT, 1964:149 ISS
             Syn.: Neonaphorura duboscqui(sic) Bagnall, RS, 1935:241,tt ISS
             Syn.: Tullbergia dubosqi(sic) Haarlon, 1957 ISS
             Syn.: Neonaphorura duboscqui(sic) Salmon, JT, 1964:149 ISS
             Syn.: Neonaphorura dubosqui(sic) Kopeszki, H et Meyer, E, 1996:225 ISS
             Syn.: Neonaphorura dubosqi(sic) ssp. dubosqi(sic) Traser, G et Thibaud, J-M, 1999:106 ISS
Neonaphorura duboscqi from the UK
Two large and 4 small anal spines
2013.04.27 © Murray, A.

Species dungeri Schulz, 1994
Species graffi Kaprus, IJ et Goblyk, KM, 2015:2
Neonaphorura graffi from Ukraine
After Kaprus IJ & Goblyk KM, 2015 Fig.1

Species hexaspina Arbea, JI et Mateos, E, 1991:169
Species howdeni (Wray, 1958) Salmon, JT, 1964:149
Species loksai Traser, G et Thibaud, J-M, 1999:105
Remark by Kaprus, I.J. & Goblyk K.M. 2015:1-2: Among Neonaphorura species cited by Dunger and Schlitt (2011), only N. loksai Traser et Thibaud, 1999 does not fully fit with the diagnosis of the subfamily Tullbergiinae, because antennal III sense organ has only two sensory clubs [therefore matching subfamily Mesaphorurinae]. However, other characters of N. loksai are typical for the genus Neonaphorura.
Species moravica (Rusek, 1966) Khanislamova, 1988
Species navarrensis Arbea, JI, 1991:184
Species novemspina (Gisin, 1963) Christian, 1987
            Syn.: Tullbergia septemspina Schaller, 1949 nn =?
Species ortali Palissa, 2006
Species zakarpatica Kaprus, IJ et Goblyk, KM, 2015:4
Neonaphorura zakarpatica from Ukraine
After Kaprus IJ & Goblyk KM, 2015 Fig.12


Genus Tasphorura Greenslade, P et Rusek, J, 1996:28
Species vesiculata Greenslade, P et Rusek, J, 1996:28, t.t.

Genus Stenaphorura Absolon, K, 1900:411
  Syn.: Tullbergiidae Tullbergia Börner, 1902, nec Lubbock, 1876
  Syn.: Tullbergiini Stenaphorurella Luciañez & Simón, 1992
Species sp. Greenslade, P et Rusek, J, 1996:33
Species denisi Bagnall, 1935, B
            Syn.: Tullbergia parisi Denis, 1943
             Syn.: Stenaphorura parisii(sic) Stach, 1954 ISS
             Syn.: Stenaphorura parisii(sic) Salmon, JT, 1964:152 ISS
             Syn.: Stenaphorurella Denisi(sic) Khanislamova G M, Sterzynska M et Weiner W M, 1997:503 ISS
Stenaphorura denisi from the UK
2012.11.28 © Murray, A.
Stenaphorura denisi from the UK
2012.11.28 © Murray, A.
Stenaphorura denisi from the UK
Two large and two small anal spines; in front two bumps (*)
2012.11.28 © Murray, A.
Stenaphorura denisi from the UK
2012.11.28 © Murray, A.
Stenaphorura denisi from the UK
2013.01.11 © Murray, A.
Stenaphorura denisi from the UK
Two large and two small anal spines; in front two bumps
2013.01.11 © Murray, A.
Stenaphorura denisi from the UK
2013.01.11 © Murray, A.

Species gibsoni Murphy, 1965
Species japygiformis Absolon, K, 1900:412, t.t.
             Syn.: Stenaphorura iapygiformis(sic) Bagnall, RS, 1935:241,tt ISS
             Syn.: Tullbergia iapygiformis(sic) Bagnall, RS, 1935:237 ISS
             Syn.: Stenaphorura iapygiformis(sic) Bagnall, 1935 ISS
             Syn.: Tullbergia japygidiformis(sic) Salmon, 1964 ISS
Species lubbocki Bagnall, 1935
Species metaparisi (Traser & Weiner, 1999) Dunger, W & Schlitt, B, 2011:138
Species quadrispina Börner, 1901, B
            Syn.: Aphorura 4-spina(sic) ssp. taurica Lebedinsky, 1904
            Syn.: Stenaphorura axelsoni Bagnall, 1935
            Syn.: Stenaphorura absoloni Bagnall, 1936
Stenaphorura quadrispina (bar=2µm)
Hexagonal epicuticular ultrastructure with secondary granules
After Nickerl J & al, 2012 Fig.2C'
Stenaphorura quadrispina (bar=500nm)
Epicuticular secondary granule
After Nickerl J & al, 2012 Fig.2C''

Species shaconage Bernard, EC, 2016:467
Stenaphorura shaconage from the USA
After Bernard EC, 2016 Fig.10AB


Subfamilia Mesaphorurinae Dunger, W & Schlitt B, 2011:19
  Syn.: Tullbergiinae Stenaphorurini Luciáñez & Simón, 1992
  Syn.: Tullbergiidae Stenaphorurinae Deharveng, L, 2004:427
Mesaphorurinae juv. from Canada
2014.02.02 © Hochhalter, R.


Genus Mesaphorura Börner, 1901 key
Mesaphorura sp. juv. from the UK
2013.01.12 © Murray, A.
Mesaphorura sp. juv. from the UK
2013.01.12 © Murray, A.

Species amazonica Oliveira & Thibaud, 1992
Species arbeai Simón Ruiz Martinez & Lucianez, 1994
Species atlantica (Rusek, 1979) Thibaud, J-M & Palacios-Vargas, JG, 1999:29
Species baconae Bagnall, 1947
Species bassolsae Palacios-Vargas, JG et Díaz, M, 1996:12
Species betschi Rusek, 1979, B
            Syn.: Mesaphorura massoudi Rusek, 1979
             Syn.: Tullbergia betchi(sic) Fjellberg, A, 1988:39 ISS
Species choudhurii Yosii, 1966
Mesaphorura choudhurii from India
After Mandal GP & Hazra AK, 2009 Fig.15a+b

             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:73 ISS
             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:77 ISS
             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:115 ISS
             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:126 ISS
             Syn.: Mesaphorura choudhuri(sic) Mandal, GP et Hazra, AK, 2009:128 ISS
Species critica Ellis, WN, 1976:230
            Syn.: Mesaphorura krausbaueri Ardanaz, 1986, partim
Mesaphorura critica from Australia
After Greenslade P, 1992 Fig.2

Species delamarei Weiner, 1991
Species dubia Börner, 1902
Species florae Simón, Ruiz, Martinez & Lucianez, 1994
             Syn.: Mesaphorura phlorae(sic) García-Gómez, A, Castaño-Meneses, G et Palacios-Vargas, JG, 2009:911 ISS
Species foveata Bonet, 1944
Species guyana Thibaud, 2004
Species himalayensis Yosii, 1971
Remark by Simón Benito, J.C. 2006.10.16:in litt.: This species should be revised, it does not belong in Mesaphorura (pseudocelli are not star shaped), but in Chaetophorura or Weinera.

Species hygrophila (Rusek, J, 1971:199) Ellis, 1976:232
             Syn.: Mesaphorura hygrophyla(sic) Ellis, 1976 ISS
Species hylophila Rusek, J, 1982:14, B
            Syn.: Mesaphorura krausbaueri Jordana & Beruete, 1983, partim
Species intermedia Prabhoo, 1971
Species iowensis (Mills, 1932) Bagnall, 1937
Species isochaeta Arbea, JI et Jordana, R, 1989:8
  Subspecies leitzaensis Jordana, 1993
Species italica (Rusek, J, 1971:203) Ellis, 1974:109, B
Species jarmilae Rusek, J, 1982:20
Species javieri (Palacios-Vargas, JG & Díaz, M, 1996:18) Palacios-Vargas, JG in Sánchez-Ramos, G, Reyes-Castillo, P & Dirzo, R, 2005:364
Species jeremiei Thibaud, J-M et Weiner, WM in Najt, J et Matile, L, 1997:74
Species jevanica Rusek, J, 1996:208, B
Species jirii Rusek, J, 1982:18
Species krausbaueri (Börner, 1901) Absolon, K, 1901:389, t.t., B
            Syn.: Tullbergia iowensis Mills, 1932
            Syn.: Paratullbergia macdougalli Bagnall, 1936
            Syn.: Mesaphorura baconae Bagnall, 1947
            Syn.: Mesaphorura krausbaueri var. bonariensis Rapoport, 1959
             Syn.: Mesaphorura Krausbaueri(sic) Absolon, K, 1901:389 ISS
             Syn.: Mesaphorura Krausbaueri(sic) Absolon, K, 1901:580 ISS
             Syn.: Tullbergia Krausbaueri(sic) Loksa, I, 1961:224 ISS
             Syn.: Tullbergia krausboueri(sic) Salmon, JT, 1964:150 ISS
             Syn.: Tullbergia kraubaueri(sic) Wibo, C, 1971:107 ISS
             Syn.: Mesaphorura krausberi(sic) Hopkin, 1997 ISS
             Syn.: Folsomia krausberi(sic) Moore, JC, Saunders, P, Selby, G, Horton, H, Chelius, MK, Chapman, A et Horrocks, RD, 2005:117 ISS
             Syn.: Mesaphorura krausbauerii(sic) Thibaud, J-M, 2008:510,gr ISS
  Subspecies atypica Denis, 1924
  Subspecies inermis Axelson, WM, 1905:790
Species macrochaeta Rusek, 1976, B
            Syn.: Metaphorura affinis f. B Ellis, WN, 1976:225
            Syn.: Mesaphorura krausbaueri Jordana & Beruete, 1983, partim
            Syn.: Mesaphorura krausbaueri Arbea & Jordana, 1985, partim
            Syn.: Mesaphorura krausbaueri Jordana & al., 1987, partim
            Syn.: Tullbergia krausbaueri Petersen, H, 1971:133
            Syn.: Tullbergia krausbaueri Greenslade, 1992 gr
            Syn.: Mesaphorura (krausbaueri-group) sp. Greenslade, 1990
Mesaphorura macrochaeta from Australia
After Greenslade P, 1992 Fig.1
             Syn.: Mesaphorura macrocheta(sic) D'Haese, CA, 2003:570 ISS
             Syn.: Mesaphorura macrocheta(sic) D'Haese, CA, 2003:397 ISS
             Syn.: Mesaphorura macrocheata(sic) Moore, JC, Saunders, P, Selby, G, Horton, H, Chelius, MK, Chapman, A et Horrocks, RD, 2005:117 ISS
             Syn.: Mesaphorura macrocheata(sic) Rochefort, S, Therrien, F, Shetlar, DJ et Brodeur, J, 2006:63 ISS
Mesaphorura macrochaeta (bar=2µm)
Hexagonal epicuticular ultrastructure with secondary granules
After Nickerl J & al, 2012 Fig.5D

Mesaphorura macrochaeta from the UK
2012.11.15 © Murray, A.
Mesaphorura macrochaeta from the UK
2012.11.15 © Murray, A.
Mesaphorura macrochaeta from the UK
2012.11.15 © Murray, A.
Mesaphorura macrochaeta from the UK
2012.11.15 © Murray, A.
Mesaphorura macrochaeta from Holland
Prominent anal spines and macrosetae on abd.6
2013.11.27 © van Bezouw, R.

Species mani Thibaud, 2002
Species maricaensis Fernandes, LH et de Mendonça, MC, 2004:17
Species matilei Thibaud & Palacios-Vargas, 2000
Species minutissima Salmon, JT, 1944:141
Species occidafricana Murphy, 1965
Species orousseti Najt, Thibaud & Weiner, 1990
            Syn.: Mesaphorura longispina Arbea, JI et Jordana, R, 1991:118
            Syn.: Mesaphorura longispina ssp. stellae Arbea, JI et Selga, D in Jordana, R et Ariño, AH, 1994:76
Species pacifica Rusek, 1976
Species petterdassi (Fjellberg, A, 1988:38) Sterzynska & Ehrnsberger, 1998
Tullbergia petterdassi from Norway
After Fjellberg, A, 1988 Fig.3A

Species pongei Rusek, 1982, B
Species redondoi Jordana, R et Arbea, JI in Jordana, R et Ariño, AH, 1994:4
Mesaphorura redondoi from Spain
After Jordana R & Arbea JI, 1994 Fig.2a
Remark by Dányi, L. & Traser, G. 2008:28: Jordana et al. (1997) considered this taxon as a subspecies of Mesaphorura isochaeta Arbea & Jordana, 1990, endemic of the Iberian Peninsula. However, this status is questionable in the light of the Hungarian occurences.


Species rudolfi Rusek, 1986
Species ruseki (Christiansen & Bellinger, 1980) Miranda Rangel & Palacios-Vargas, 1992
Species schembrii Thibaud, J-M et Christian, E, 1989:76
Species sensibilis Rusek, J, 1973:79
Species silvicola (Folsom, JW, 1932:57) Stach, 1954
            Syn.: Tullbergia granulata Mills, HB, 1934:37
Species simoni Jordana, R et Arbea, JI in Jordana, R et Ariño, AH, 1994:2
Mesaphorura simoni from Spain
After Jordana R & Arbea JI, 1994 Fig.1a

Species simplex (Gisin, 1958)
Species sp. Maes & Palacios-Vargas, 1988:3
Species sp. Thibaud, 1993
Species sp. Palacios-Vargas & Díaz, 1995:171
Species sp. Palacios-Vargas & Díaz, 1995:172
Species sp. Mejía-Ortiz, LM, Palacios-Vargas J, Cardona, L et Viccon-Pale, JA, 1997:26
Species spelaea (Nosek & Neuherz, 1976)
Species subitalica Thibaud, 1996
Species sylvatica (Rusek, J, 1971:195) Dunger, 1977:180
             Syn.: Mesaphorura silvatica(sic) Kaprus, IJ, Pomorski, RJ, Skarzynski, D et Potapov, MB, 2005:893 ISS
Species tenuisensillata Rusek, 1974, B
             Syn.: Mesaphorura tenuisensillana(sic) Kaprus, IJ, Pomorski, RJ, Skarzynski, D et Potapov, MB, 2005:897 ISS
Species thalassophila Bagnall, 1937
Species tuberosa Bonet, 1944
Species yosiii (Rusek, J, 1967:191) Rusek, 1973, B
            Syn.: Mesaphorura krausbaueri Arbea & Jordana, 1985, partim
            Syn.: Mesaphorura krausbaueri Jordana & al., 1987, partim
             Syn.: Tullbergia (Mesaphorura) yosii(sic) Rusek, J, 1971:192 ISS
             Syn.: Mesaphorura yosii(sic) Rusek, 1973 ISS
             Syn.: Mesaphorura yosii(sic) Rusek, 1979 ISS
             Syn.: Mesaphorura yosii(sic) Ponge in Pesson, 1980 ISS
             Syn.: Mesaphorura yossii(sic) Palacios-Vargas et al., 1982 ISS
             Syn.: Mesaphorura yosii(sic) Hågvar, 1983 ISS
             Syn.: Mesaphorura yosii(sic) Ponge, J-F, 1983:363 ISS
             Syn.: Mesaphorura yosii(sic) Hågvar, 1984 ISS
             Syn.: Mesaphorura yosii(sic) Hågvar & Abrahamsen, 1984 ISS
             Syn.: Mesaphorura yosii(sic) Pozo, 1986 ISS
             Syn.: Tullbergia yosii(sic) Takeda, 1987 ISS
             Syn.: Tullbergia vosii(sic) Leinaas, HP, 1988:278 ISS
             Syn.: Mesaphorura yosii(sic) Szeptycki & Weiner in Razowski, 1990 ISS
             Syn.: Mesaphorura yosii(sic) Arbea, JI et Jordana, R, 1991:116 ISS
             Syn.: Mesaphorura yosii(sic) Najt, J, Thibaud, J-M et Jacquemart, S (+), 1991:165 ISS
Mesaphorura yosiii from Australia
After Greenslade P, 1992 Fig.3

             Syn.: Mesaphorura yosii(sic) Ponge, J-F, 1993:242 ISS
             Syn.: Mesaphorura yosii(sic) Barra, 1994 ISS
             Syn.: Mesaphorura yosii(sic) Greenslade, P in Houston, WWK, 1994:59 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M, Najt, J et Jaquemart(sic), S (+), 1994:200 ISS
             Syn.: Mesaphorura yosii(sic) Barra, 1995 ISS
             Syn.: Mesaphorura yosii(sic) Kopeszki, H et Meyer, E, 1996:225 ISS
             Syn.: Mesaphorura yosii(sic) Rusek, J, 1996:214 ISS
             Syn.: Mesaphorura yosii(sic) Hopkin, 1997 ISS
             Syn.: Mesaphorura yosii(sic) Najt, J et Weiner, WM in Najt, J et Matile, L, 1997:10 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M et Weiner, WM in Najt, J et Matile, L, 1997:76 ISS
             Syn.: Mesaphorura yosii(sic) Ponge, J-F, 1999:58 ISS
             Syn.: Mesaphorura yosii(sic) Palacios-Vargas, JG, 2000:29 ISS
             Syn.: Mesaphorura yosii(sic) Ponge, J-F, 2000:510,B ISS
             Syn.: Mesaphorura yosii(sic) Park, K-H, Kim, K-G, Park, S-J, Kim, B-J et Lee, B-H, 2000:415 ISS
             Syn.: Mesaphorura yosii(sic) Sabatini & Innocenti, 2000 ISS
             Syn.: Mesaphorura yosii(sic) Peck, SB, 2001:103 ISS
             Syn.: Mesaphorura yosii(sic) Castaño-Meneses, G, Palacios-Vargas, JG et Cutz-Pool, L, 2004:138 ISS
             Syn.: Mesaphorura yosii(sic) Fernandes, LH et de Mendonça, MC, 2004:16 ISS
             Syn.: Mesaphorura yosii(sic) Tamura & al., 2004 ISS
             Syn.: Mesaphorura yosii(sic) Janssens, F et Dethier, M, 2005:154,B ISS
             Syn.: Mesaphorura yosii(sic) Boitaud, L, Salmon, S Bourlette, C et Ponge J-F, 2006:451 ISS
             Syn.: Mesaphorura yosii(sic) Fiera, C, 2007:18 ISS
             Syn.: Mesaphorura yosii(sic) Dányi, L et Traser, G, 2008:29 ISS
             Syn.: Mesaphorura yosii(sic) Thibaud, J-M, 2008:510 ISS
             Syn.: Mesaphorura yosii(sic) Xiong, Y, Gao, Y, Yin, W-Y et Luan, Y-X, 2008:730 ISS
             Syn.: Mesaphorura yosii(sic) Fernandes, LH, Nessimian, JL et de Mendonça, MC, 2009:1036 ISS
             Syn.: Mesaphorura yosii(sic) Nakamori, T, Fujiwara, N, Matsumoto, N et Okada, H, 2009:7 ISS
             Syn.: Mesaphorura yosii(sic) Busmachiu, G, 2010:153 ISS
             Syn.: Tullbergia yosii(sic) Suetsugu, N, Satoh, F et Koike, T, 2010:60 ISS
             Syn.: Mesaphorura yosii(sic) Chauvat, M, Trap, J, Perez, G, Delporte, P et Aubert, M, 2011:411 ISS
             Syn.: Mesaphorura yosii(sic) Dunger, W et Schlitt, B, 2011:86 ISS
             Syn.: Mesaphorura yosii(sic) Palacios-Vargas, JG, Castaño-Meneses, G et Estrada, DA, 2011:120 ISS
             Syn.: Mesaphorura yosii(sic) Peck, SB et Jacquemart, J, 2011:5 ISS
             Syn.: Mesaphorura yossii(sic) Cutz-Pool, LQ et Vázquez-González, M, 2012:109 ISS
             Syn.: Mesaphorura yosii(sic) Busmachiu, G, Bedos, A et Deharveng, L, 2015:101 ISS
             Syn.: Mesaphorura yosii(sic) Janion-Scheepers, C, Deharveng, L, Bedos, A et Chown, L, 2015:64 ISS
             Syn.: Mesaphorura yosii(sic) Bernard, EC, 2016:451 ISS
             Syn.: Mesaphorura yosii(sic) Bernard, EC, 2016:457 ISS
             Syn.: Mesaphorura yosii(sic) Bernard, EC, 2016:460 ISS
             Syn.: Mesaphorura yosii(sic) Bernard, EC, 2016:476 ISS
             Syn.: Mesaphorura yosii(sic) Yu, D-Y, Zhang, F, Stevens, MI, Yan, Q-B, Liu, M-Q et Hu, F, 2016:288 ISS

Genus Neotullbergia Bagnall, RS, 1935:241
Remark by Dunger, W & Schlitt B. 2011:111: The genus Neotullbergia Bagnall needs a revision.
Remark by Janssens, F. 2012.11.04: The genus Neotullbergia Bagnall possibly is a genus complex given some species such as ramicuspis and crassicuspis have 2 sensory clubs while others such as staudacheri and tricuspis have 3 sensory clubs in the ant.3 sense organ.
Species americana Bonet, 1944
Species crassicuspis (Gisin, 1943) Salmon, JT, 1964:148, B
Species ramicuspis (Gisin, 1953) Salmon, 1964:148
            Syn.: Tullbergia tricuspis Strebel, 1957
            Syn.: Tullbergia tricuspis Paclt, 1961
Species staudacheri (Kos, 1940) Dunger, W & Schlitt, B, 2011:112
            Syn.: Tullbergia Neotullbergia tricuspis Gisin, 1944
            Syn.: Neotullbergia laingi Bagnall, 1936
Species tricuspis (Börner, 1902) Bagnall, RS, 1935:241, t.t.

Genus Metaphorura Stach, J, 1954:204 key
Species affinis (Börner, 1902) Bagnall, 1936 sensu Stach, 1954
            Syn.: Metaphorura börneri(sic) Bagnall, 1936 sensu Stach, 1954 IOS
             Syn.: Tullbergia bipartita ssp. borneri(sic) Bagnall, 1939 ISS
            Syn.: Metaphorura affinis type B Ellis, WN, 1976:227
            Syn.: Metaphorura bipartita? Skarzynski, D, Pomorski, RJ, Smolis, A, Weiner, WM, Szeptycki, A, Slawska, M et Sterzynska, M, 2002:32
Metaphorura affinis
2006 © Boldsystems.org
Metaphorura affinis from the UK
Two anal spines with medial conical projection
2013.04.27 © Murray, A.

Species bipartita (Handschin, 1920) Bagnall, 1936, t.t.
            Syn.: Metaphorura börneri(sic) Bagnall, 1936 IOS
             Syn.: Metaphorura börneri(sic) Bagnall, 1937 ISS
             Syn.: Metaphorura börneri(sic) Kloet, GS et Hincks, WD, 1945:5 ISS
             Syn.: Metaphorura börneri(sic) Denis, 1948 ISS
            Syn.: Tullbergia bagnalli Denis, 1943
             Syn.: Tullbergia Bagnalli(sic) Delamare Deboutteville, C et Jacquemart, S, 1961:8 ISS
            Syn.: Metaphorura affinis Stach, 1955
Species denisi Simón Benito, 1985
            Syn.: Metaphorura affinis f. C Ellis, WN, 1976:225
Species ellisi Simón Benito, 1985
            Syn.: Metaphorura affinis f. A Ellis, WN, 1976:225
Species incisa (Bonet, 1944) Castaño-Meneses, G, Palacios-Vargas, JG & Traser, G, 2000:1266
Species knowltoni (Wray, DL, 1950:97) Palacios-Vargas, JG & Díaz, M, 1996:173
Tullbergia knowltoni from the USA
After Wray DL, 1950 Pl.5 Fig.1

Species orestia Pomorski, RJ, Skarzynski, D et Kaprus, I, 1998:265
Species riozoi Castaño-Meneses, G, Palacios-Vargas, JG et Traser, G, 2000:1263
Species triacantha (Börner, 1901) Dallai, R, Malatesta, E & Ramellini, P in Minelli, A, Ruffo, S & La Posta, S, 1995:8

Genus Delamarephorura Weiner & Najt, 1999
  Syn.: Tullbergiidae Spicatella Thibaud, 2002
Species bedosae (Thibaud, 2002) Janion, Deharveng & Weiner, 2013
Species capensis Janion, Deharveng & Weiner, 2013
Species salti (Delamare Deboutteville, 1953) Weiner & Najt, 1999, t.t.
Species szeptickii Barra & Weiner, 2009
Species tami Janion, Deharveng & Weiner, 2013

Genus Stenaphorura2 Thibaud, 2002 =i
Species marionae Thibaud, 2002

Genus Tullbergiella Izarra, 1965
Species allendei (Izarra, 1975) Mari Mutt & Bellinger, 1990:57
Species humilis (Izarra, 1965) Mari Mutt & Bellinger, 1990:57, t.t.

Genus Karlstejnia Rusek, 1974
Species annae Rusek, 1974
Species montana Rusek, 1978
Species norvegica (Fjellberg, 1974) Potapov, M & Stebaeva, S, 1997:124
Species rusekiana Weiner, 1983
Species sibirica Rusek, 1978

Genus Prabhergia Salmon, 1965
Species imadatei Tamura H in Tamura, H et Zhao, L-J, 1996:791
Species indonesiae Yoshii, R et Suhardjono, YR, 1989:29
Species nayarii Salmon, 1965, t.t.

Genus Boudinotia Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:121
Species prima Weiner, WM et Najt, J in Chazeau, J et Tillier, S, 1991:121, t.t.

Genus Marcuzziella Rusek, 1975
  Syn.: Tullergiinae Marcuziella(sic) Petersen, H, 2011:486 ISS
Species tripartita Rusek, 1975

Genus Wankeliella Rusek, 1975 key
Species bescidica Smolis, A et Skarzynski, D, 2003:451
Species intermedia Potapov, M et Stebaeva, S, 1997:123
Species medialis Simón-Benito, JC et Jordana, R in Jordana, R et Ariño, AH, 1994:16
Wankeliella medialis from Spain
After Simón JC & Jordana R, 1994 Fig.1c

Species mediochaeta Rusek, 1975
Species peterseni Rusek, 1975 tt
Species pongei Rusek, 1978

Genus Paratullbergia Womersley, 1930
Species brevispina Skarzynski & Pomorski, 1999
Species callipygos (Börner, 1902) Salmon, JT, 1964:147, B
            Syn.: Paratullbergia concolor Womersley, 1930 =?
            Syn.: Paratullbergia carpenteri Bagnall, 1935 =?
            Syn.: Paratullbergia womersleyi Bagnall, 1935
            Syn.: Tullbergia callipygos f. hirsuta Cassagnau, P, 1952:305
             Syn.: Tullbergia callipygas(sic) Heungens, A et Van Daele, E, 1973:136,B ISS
             Syn.: Paratullbergia calliphygos(sic) Moore, JC, Saunders, P, Selby, G, Horton, H, Chelius, MK, Chapman, A et Horrocks, RD, 2005:117 ISS
Paratullbergia callipygos
2006 © Boldsystems.org
Paratullbergia callipygos from the UK
2012.11.27 © Murray, A.
Paratullbergia callipygos from the UK
Elongate post-antennal organ
2012.11.27 © Murray, A.
Paratullbergia callipygos from the UK
Distinct large anal spines
2012.11.27 © Murray, A.

             Syn.: Tullbergia callipygosa(sic) Falahati HosseinAbad, A, Mehr, M S et Khyroodin, A, 2013:260 ISS
Paratullbergia callipygos from the UK
2013.04.05 © Murray, A.
Paratullbergia callipygos juv. from the UK
Transverse dorsal suture in 4th abdominal segment
2013.04.15 © Murray, A.
Paratullbergia callipygos from the UK
6th abdominal segment with ridges bumps and large anal spines
2013.05.07 © Murray, A.
Paratullbergia callipygos juv. from Holland
Abd.4 with transverse suture
Abd.6 with ridges bumps and 2 anal spines
2014.10.06 © van Duinen, J.
Paratullbergia callipygos juv. from the UK
Transverse dorsal suture in 4th abdominal segment
2014.12.28 © Nurcombe, E.

Species caroli Lucianez Ortega & Simón, 1991
Species concolor Womersley, 1930, t.t.
Species indica Salmon, 1965
Species macdougalli Bagnall, 1936
Species salmoni Prabhoo, 1971
Species trivandrana Prabhoo, 1971

Genus Multivesicula Rusek, 1982
Species aliciae Palacios-Vargas, JG et Díaz, M, 1996:15
Species ampla (Christiansen & Bellinger, 1980)
Species columbica Rusek, 1982
Species dolomitica Rusek, 1982
Species giljarovi Rusek, 1982
Species punctata Rusek, 1982
             Syn.: Multivesicula puctata(sic) Skidmore, RE, 1995:52 ISS

Genus Pongeiella Rusek, 1991
Species falca (Christiansen & Bellinger, 1980) Rusek, 1991
  Subspecies europaea Rusek, 1991
Species luciaelvirae Thibaud & Peja, 1996
Species stojanovorum Pomorski & Skarzynski, 1997

Genus Psammophorura Thibaud & Weiner, 1994
Species gedanica Thibaud & Weiner, 1994
             Syn.: Psammophorura gedanicus(sic) Bernard, EC, 2016:467 ISS
Species miniclavata Bernard, EC, 2016:464
Psammophorura miniclavata from the USA
After Bernard EC, 2016 Fig.8AB

Species neocaledonica Thibaud, J-M et Weiner, WM in Najt, J et Matile, L, 1997:77

Genus Rotundiphorura Rusek, 1991
Species habanica Rusek, 1991

Genus Weinera Thibaud, 1993
Species ghislaineae Thibaud, 1993

Genus Sensilatullbergia Thibaud & Ndiaye, 2006
Species senegalensis Thibaud & Ndiaye, 2006

Genus Mexicaphorura Palacios-Vargas, JG et Catalán, E, 2013:92
Species guerrerensis Palacios-Vargas, JG et Catalán, E, 2013:93, t.t.
             Syn.: Mexicaphorura guerrensis(sic) Palacios-Vargas, JG et Catalán, E, 2013:94 ISS

Genus Ameritulla Bernard, EC, 2016:452
Species clavata (Mills, HB, 1934:38) Bernard, EC, 2016:454, t.t.
Ameritulla clavata from the USA
After Bernard EC, 2016 Fig.1AB

Species hades (Christiansen & Bellinger, 1980) Bernard, EC, 2016:453
Species obscura Bernard, EC, 2016:458
Ameritulla obscura from the USA
After Bernard EC, 2016 Fig.4AB


Genus Mixturatulla Bernard, EC, 2016:461
Species ozwini Bernard, EC, 2016:461, t.t.
Mixturatulla ozwini from the USA
After Bernard EC, 2016 Fig.6ABC


Subfamilia Austraphorurinae Deharveng L, 2004:427
  Syn.: Tullbergiinae Austraphorurini Luciáñez & Simón, 1992

Genus Doutnacia Rusek, 1974
Species sp. Fjellberg, A in Pomorski, RJ & Skarzynski, D, 1998:252
Species coineaui Massoud & Thibaud, 1985
Remark by Dunger, W. & Schlitt,B. 2011:35: This species probably belongs to another genus. Further characters separate the species from the genus Doutnacia. We consider the species a mere species inquirenda.

Species mols Fjellberg, A, 1998:146
            Syn.: Doutnacia ammophila Pomorski, RJ et Skarzynski, D, 1998:248
Species xerophila Rusek, 1974, t.t.
             Syn.: Doutnacia xerotherma(sic) Russell, DJ, 2008:124 lapsus ISS

Genus Jevania Rusek, 1978
Species fageticola Rusek, 1978, t.t.
Species weinerae Rusek, 1978
             Syn.: Jevania weineriae(sic) Busmachiu, G, 2010:153 ISS

Genus Scaphaphorura Petersen, 1965
Species arenaria (Petersen, 1965) Ellis, WN & Bellinger, PF, 1973:49, t.t.
Species corpesiensis Simón & Lucianez, 1990
Species cubana Thibaud, 1994
Species pakieti Pomorski, RJ et Skarzynski, D, 1997:3
Species sp. Dunger, 1977:181, s.i.

Genus Najtiaphorura Weiner & Thibaud, 1991
Species dobrolubovae Shveenkova, YB et Potapov, MB, 2003:?
Species marocana Thibaud & Boumezzough, 2006
Species olivieri (Weiner & Thibaud, 1991) Dunger, W & Schlitt, B, 2011:101
Species sandrinae Weiner & Thibaud, 1991



References