Triaenonychidae

Soerensenella prehensor, photographed by Phil Bendle.

Belongs within: Triaenonychoidea.
Contains: HendeaEquitiusCeratomontia, AdaeinaeAcumontiaNunciellaNuncia, Triaenobuninae.

The Triaenonychidae are a family of harvestmen characterised by branched hind claws and found primarily on the southern continents (South America, Australasia and Africa). Kaolinonychus and the Nippononychinae from north-eastern Asian have also been assigned to this family, but their position here is open to doubt (Mendes & Kury 2008).

The Triaenonychini was established in its current form by Forster (1954) who suggested that the separate Southern Hemisphere subfamilies of Triaenonychidae previously recognised by Roewer (1923) were probably not natural groupings. Rather than eliminate them entirely, he reduced them to tribes of a single subfamily Triaenonychinae pending further analysis. The tribes remain untested, though Kury (2006) suggested that the previously triaenonychin genus Graemontia might be more closely related to the tribe Triaenobunini. Species of Triaenonychini are found in South Africa, Australia, New Zealand and South America. Fewer species have been described from the last locality than the others, though the South American taxa do include the type genus Triaenonyx.

Characters (from Suzuki 1975): Anterior margin of scute usually with three projections flanking chelicerae. Eyes situated on median tubercle. Dorsum with five tergal areas bounded by transverse grooves or rows of tubercles and three free tergites. Ninth tergite fused to anal plate. Palpi robust, without slender, elongated segments. Spiracles exposed or hidden in a groove between coxa and sternite. Metatarsi of legs usually divided by a constriction into two sections, a proximal astragalus and a distal calcaneus. Claws of third and fourth legs with uniform or stout central prong bearing two to six or less lateral branches in adults and three, four or five pairs of lateral spines or branches in juveniles. Pseudonychium, aroleum or scapulae not present on claws of adults or juveniles. Males with musculature in penis which apparently articulates distal process.

Triaenonychidae [Triaenochini, Triaenonychinae, Triaenonychini, Triaenonychoidae]
    |--+--+--Pyenganella Hickman 1958DB21, BS20
    |  |  |    `--P. striata Hickman 1958KMS14
    |  |  `--HendeaDB21
    |  `--+--Hendeola Forster 1954DB21, F54
    |     |    |--*H. bullata Forster 1954F54
    |     |    |    |--H. b. bullataNS00
    |     |    |    `--H. b. pterna Forster 1954NS00
    |     |    `--H. woodwardi Forster 1954NS00
    |     `--EquitiusDB21
    `--+--+--+--Lizamontia Kury 2004DB21, K04
       |  |  |    |--*L. starengai Kury 2004K04
       |  |  |    |--L. draconensis (Lawrence 1939)KMS14 [=Acumontia draconensisK04]
       |  |  |    `--L. natalensis (Lawrence 1931) [=Acumontia natalensis]K04
       |  |  `--Paramontia Lawrence 1934DB21, S92
       |  |       |--*P. lisposoma (Lawrence 1931) [=Rostromontia lisposoma]S92
       |  |       `--P. infinita Lawrence 1934S92
       |  `--+--CeratomontiaDB21
       |     `--+--Amatola Lawrence 1931DB21, S92 [incl. Cryptobunus Lawrence 1931S92]
       |        |    |--*A. dentifrons Lawrence 1931S92
       |        |    |--A. armata (Lawrence 1938) [=Cryptobunus armatus]L02
       |        |    |--A. durbanica (Lawrence 1937) [=Cryptobunus durbanicus]S92
       |        |    |--A. maritima (Lawrence 1937) [=Cryptobunus maritimus]S92
       |        |    |--A. setifemur (Lawrence 1931)S92 [=*Cryptobunus setifemurL31; incl. C. silvicolus Lawrence 1937S92]
       |        |    `--A. unidentata (Lawrence 1937) [=Cryptobunus unidentatus]S92
       |        `--Biacumontia Lawrence 1931DB21
       |             |--*B. paucidens Lawrence 1931S92
       |             |--B. cornuta Lawrence 1931S92
       |             |--B. elata Kauri 1961S92
       |             |--B. fissidens Lawrence 1931S92
       |             |--B. maculata Lawrence 1938S92
       |             |--B. truncatidens Lawrence 1931S92
       |             `--B. variegata Lawrence 1934S92
       `--+--+--+--Picunchenops Maury 1988DB21, K03
          |  |  |    `--*P. spelaeus Maury 1988K03
          |  |  `--Hedwiga Roewer 1931DB21, F54
          |  |       `--*H. manubriata Roewer 1931F54
          |  `--+--+--Araucanobunus Muñoz-Cuevas 1973DB21, K03
          |     |  |    `--*A. juberthiei Muñoz-Cuevas 1973K03
          |     |  `--+--+--+--Valdivionyx Maury 1988DB21, K04
          |     |     |  |  |    `--*V. crassipes Maury 1988K03
          |     |     |  |  `--+--Gunvoria Kauri 1961MK08, S92
          |     |     |  |     |    `--*G. spatulata Kauri 1961S92, MK08
          |     |     |  |     `--Nahuelonyx Maury 1988DB21, K03
          |     |     |  |          `--*N. nasutus (Ringuelet 1959)K03, MK08 [=Diasia nasutaK03]
          |     |     |  `--Triaenonychoides Soares 1968DB21, K03 [=Trianonychoides (l. c.)K03]
          |     |     |       |--*T. cekalovici Soares 1968 [=Trianonychoides (l. c.) cekalovici]K03
          |     |     |       `--T. breviops Maury 1987K03
          |     |     `--Diasia Sørensen 1902DB21, K04 [=Diaria (l. c.)K03, Diasa (l. c.)K03]
          |     |          |--*D. michaelsenii Sørensen 1902 (see below for synonymy)K03
          |     |          |--D. araucana Maury 1987K03
          |     |          `--D. platnicki Maury 1987K03
          |     `--Triaenonyx Sørensen 1886DB21, K04 [incl. Pythora Koch in Sørensen 1886R23]
          |          |  i. s.: *T. rapax Sørensen 1886K03, R31 [=Pythora rapaxR23]
          |          |         T. corralensis Roewer 1915K03
          |          |         T. valdiviensis Sørensen 1902R31 [=Nuncia valdiviensisH20]
          |          |--T. dispersus Roewer 1915BS20, R31
          |          `--+--T. arrogans Soares 1968BS20, K03
          |             `--T. chilensis Sørensen 1902BS20, K03
          `--+--+--+--+--Speleomontia Lawrence 1931DB21, BS20
             |  |  |  |    `--*S. cavernicola Lawrence 1931S92
             |  |  |  `--Mensamontia Staręga 1992DB21, KMS14
             |  |  |       |--*M. morulifera Lawrence 1931S92
             |  |  |       `--M. melanophora Lawrence 1931S92
             |  |  `--+--+--‘Nuncia’ rostrata Maury 1990DB21
             |  |     |  |--‘Nuncia’ spinulosa Maury 1990BS20
             |  |     |  `--‘Nuncia’ verrucosa Maury 1990DB21, K03
             |  |     `--AdaeinaeDB21
             |  `--+--+--+--Micromontia Lawrence 1939DB21, K61
             |     |  |  |    `--*M. flava Lawrence 1939S92
             |     |  |  `--Roewerania Lawrence 1934DB21, S92
             |     |  |       |--*R. lignicola Lawrence 1934S92
             |     |  |       |--R. guduana Kauri 1961L02
             |     |  |       |--R. montana Kauri 1961L02
             |     |  |       `--R. spinosa Lawrence 1938L02
             |     |  `--Graemontia Staręga 1992DB21, KMS14
             |     |       |--*G. bifidens Lawrence 1931K06
             |     |       |--G. bicornigera Lawrence 1963K06
             |     |       |--G. decorata Lawrence 1938K06
             |     |       |--G. dentichelis Lawrence 1931K06
             |     |       |--G. erecta Kauri 1961K06
             |     |       |--G. natalensis Lawrence 1937K06
             |     |       `--G. viridiceps Kury 2006K06
             |     `--+--+--Paulianyx Lawrence 1959DB21, K04
             |        |  |    |--*P. brevipes Lawrence 1959S92
             |        |  |    `--P. ferruginea Lawrence 1959S92
             |        |  `--+--Hovanuncia Lawrence 1959DB21, S92
             |        |     |    |--*H. monticola Lawrence 1959S92
             |        |     |    `--H. bidentata Lawrence 1959S92
             |        |     `--+--Antongila Roewer 1931DB21, S92
             |        |        |    `--*A. spinigera Roewer 1931S92
             |        |        `--+--Decarynella Fage 1945DB21, S92
             |        |           |  `--*D. gracilipes Fage 1945S92
             |        |           `--AcumontiaDB21
             |        `--+--+--Ankaratrix Lawrence 1959DB21, S92
             |           |  |    |--*A. illota Lawrence 1959S92
             |           |  |    |--A. cancrops Lawrence 1959S92
             |           |  |    |--A. fisheri Porto & Pérez-González 2020DB21
             |           |  |    |--A. lawrencei Porto & Pérez-González 2020DB21
             |           |  |    |--A. makamba Porto & Pérez-González 2020DB21
             |           |  |    `--A. maloto Porto & Pérez-González 2020DB21
             |           |  `--+--Yatala Roewer 1942DB21
             |           |     |    |--Y. aspera (Forster 1949)KMS14
             |           |     |    `--Y. hirsti Roewer 1942KMS14
             |           |     `--+--Breviacantha Kauri 1954DB21, K54
             |           |        |    `--*B. gisleni Kauri 1954K54
             |           |        `--Dingupa Forster 1952DB21, BS20
             |           |             `--*D. glauerti Forster 1952F50, KMS14
             |           `--+--+--Notonuncia Hickman 1958DB21
             |              |  |    |--N. arvensis Hickman 1958KMS14
             |              |  |    |--N. diversa Hickman 1958KMS14
             |              |  |    `--N. obscura Hickman 1958KMS14
             |              |  `--+--Bryonuncia Hickman 1958DB21
             |              |     |    `--B. distincta Hickman 1958KMS14
             |              |     `--Ankylonuncia Hickman 1958DB21
             |              |          |--A. barrowensis Hickman 1958KMS14
             |              |          |--A. fallax Hickman 1958KMS14
             |              |          `--A. mestoni Hickman 1958KMS14
             |              `--NunciellaDB21
             `--+--+--‘Nuncia’ americana Roewer 1961DB21, K03
                |  `--+--‘Nuncia’ chilensis (Soares 1968)BS20, K03 (see below for synonymy)
                |     `--+--Calliuncus Roewer 1931BS20, H72
                |        |    |--*C. ferrugineus Roewer 1931H72, R31
                |        |    |--C. ephippiatus Roewer 1931R31
                |        |    |--C. glaber Kauri 1954 [=C. glabra]H59
                |        |    |--C. labyrinthus Hunt 1972H72
                |        |    |--C. odoratus Hickman 1958KMS14
                |        |    `--C. vulsus Hickman 1959 [=C. glaber Hickman 1958 non Kauri 1954]H59
                |        `--+--Callihamina Roewer 1942DB21
                |           |    `--C. adelaidia Roewer 1942KMS14
                |           `--Callihamus Roewer 1931DB21
                |                |--*C. badius Roewer 1931KM20
                |                `--C. inermis Roewer 1931R31
                `--+--NunciaDB21
                   `--+--Americobunus Muñoz-Cuevas 1972DB21, K03
                      |    `--*A. ringueleti Muñoz-Cuevas 1972K03
                      `--TriaenobuninaeDB21
Triaenonychidae incertae sedis:
  Leionuncia levis Hickman 1958KMS14
  Parattahia Roewer 1915KM20
    `--*P. u-signatum Roewer 1915 [=P. u-signata]KM20
  Promecostethus unifalculatus Enderlein 1909KMS14 [=Nuncia unifalculataF54]
  Psalenoba Roewer 1931F54
    `--*P. nunciaeformis Roewer 1931F54
  Perthacantha Roewer 1931R31
    `--*P. jugata Roewer 1931R31
  Brasiloctis Mello-Leitão 1938K03
    `--*B. bucki Mello-Leitão 1938K03
  Neonuncia Roewer 1915DB21
    |--*N. enderbyi (Hogg 1909)F54, KMS14 (see below for synonymy)
    |--N. blacki Forster 1954NS00
    |--N. campbelli Foster 1954NS00
    |--N. eastoni Forster 1954NS00
    `--N. opaca (Roewer 1931) [=Nuncia opaca]F54
  Austronuncia Lawrence 1931S92
    |--*A. spinipalpis Lawrence 1931S92, DB21
    `--A. leleupi Lawrence 1963S92
  Lispomontia Lawrence 1937K61
    `--L. coxidens Lawrence 1937K61
  Conoculus Forster 1949F49
    `--*C. asperus Forster 1949F49
  Yulella Lawrence 1939K04
    `--*Y. natalensis (Lawrence 1937) [=Roewerania natalensis]S92
  Millomontia Lawrence 1959K04
    |--*M. brevispina Lawrence 1959S92
    `--M. vadoni Lawrence 1959S92
  Millotonyx Lawrence 1959K04
    `--*M. tenuipes Lawrence 1959S92
  Ivohibea Lawrence 1959S92
    `--*I. cavernicola Lawrence 1959S92

*Diasia michaelsenii Sørensen 1902 [=Diaria (l. c.) michaelsenii, Diasa michaelsenii (l. c.), Diasia michaelseni (l. c.)]K03

*Neonuncia enderbyi (Hogg 1909)F54, KMS14 [=Triaenonyx enderbeiF54, Neonuncia enderbeiF54, Nuncia enderbaeiH20; incl. Nu. insulana Roewer 1942F54]

‘Nuncia’ chilensis (Soares 1968)BS20, K03 [=Parattahia chilensisK03; incl. *Chilenuncia donosoi Muñoz-Cuevas 1971K03]

*Type species of generic name indicated

References

[BS20] Baker, C. M., K. Sheridan, S. Derkarabetian, A. Pérez-González, S. Vélez & G. Giribet. 2020. Molecular phylogeny and biogeography of the temperate Gondwanan family Triaenonychidae (Opiliones: Laniatores) reveals pre-Gondwanan regionalisation, common vicariance, and rare dispersal. Invertebrate Systematics 34: 637–660.

[DB21] Derkarabetian, S., C. M. Baker, M. Hedin, C. E. Prieto & G. Giribet. 2021. Phylogenomic re-evaluation of Triaenonychoidea (Opiliones: Laniatores), and systematics of Triaenonychidae, including new families, genera and species. Invertebrate Systematics 35: 133–157.

[F49] Forster, R. R. 1949. Australian Opiliones. Memoirs of the National Museum of Victoria 16: 59–89.

[F50] Forster, R. R. 1950. Western Australian Opiliones. Journal of the Royal Society of Western Australia 36: 23–29.

[F54] Forster, R. R. 1954. The New Zealand harvestmen (sub-order Laniatores). Canterbury Museum Bulletin 2: 1–329.

[H59] Hickman, V. V. 1959. Calliuncus vulsus nom. nov. (Opiliones). Papers and Proceedings of the Royal Society of Tasmania 93: vi.

[H20] Hogg, H. R. 1920. Some Australian Opiliones. Proceedings of the Zoological Society of London 1920: 31–48.

[H72] Hunt, G. S. 1972. A new cavernicolous harvestman from Western Australia (Arachnida: Opiliones: Triaenonychidae). Journal of the Australian Entomological Society 11: 232–236.

[K54] Kauri, H. 1954. Report from Professor T. Gislén’s expedition to Australia in 1951-1952. 9. Harvest-spiders from S. W. Australia. Lunds Universitets Ärsskrift. N. F. Avd. 2, 50 (11): 1–10.

[K61] Kauri, H. 1961. Opiliones. In: Hanström, B., P. Brinck & G. Rudebeck (eds) South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 9–197. Almqvist & Wiksell: Uppsala.

[K03] Kury, A. B. 2003. Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones). Revista Ibérica de Aracnología, volumen especial monográfico 1: 1–337.

[K04] Kury, A. B. 2004. A new genus of Triaenonychidae from South Africa (Opiliones, Laniatores). Revista Ibérica de Aracnología 9: 205–210.

[K06] Kury, A. B. 2006. A new species of Graemontia Lawrence, 1931, from the Western Cape, South Africa, with notes on the relationships of the genus (Opiliones: Laniatores: Triaenonychidae). African Zoology 41 (1): 45–50.

[KM20] Kury, A. B., A. C. Mendes, L. Cardoso, M. S. Kury & A. A. Granado. 2020. WCO-Lite: Online world catalogue of harvestmen (Arachnida, Opiliones). Version 1.0—Checklist of all valid nomina in Opiliones with authors and dates of publication up to 2018. Published by the author: Rio de Janeiro.

[KMS14] Kury, A. B., A. C. Mendes & D. R. Souza. 2014. World checklist of Opiliones species (Arachnida). Part 1: Laniatores—Travunioidea and Triaenonychoidea. Biodiversity Data Journal 2: e4094.

[L31] Lawrence, R. F. 1931. The harvest-spiders (Opiliones) of South Africa. Annals of the South African Museum 29 (2): 341–508.

[L02] Lotz, L. N. 2002. The Opiliones (Arachnida) of the Free State Province, South Africa. Navorsinge van die Nasionale Museum Bloemfontein 18 (8): 161–187.

[MK08] Mendes, A. C., & A. B. Kury. 2008. Intercontinental Triaenonychidae—the case of Ceratomontia (Opiliones, Insidiatores). Journal of Arachnology 36 (2): 273–279.

[NS00] Nicholls, D. C., P. J. Sirvid, S. D. Pollard & M. Walker. 2000. A list of arachnid primary types held in Canterbury Museum. Records of the Canterbury Museum 14: 37–48.

[R23] Roewer, C.-F. 1923. Die Weberknechte der Erde: Systematische Bearbeitung der bisher bekannnten Opiliones. Gustav Fischer: Jena.

[R31] Roewer, C. F. 1931. Über Triaenonychiden. Zeitschrift für Wissenschaftliche Zoologie 138: 137–185.

[S92] Staręga, W. 1992. An annotated check-list of Afrotropical harvestmen, excluding the Phalangiidae (Opiliones). Annals of the Natal Museum 33 (2): 271–336.

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