Travunioidea

Sclerobunus nondimorphicus, copyright Marshal Hedin.

Belongs within: Laniatores.
Contains: Cladonychiidae, Metanonychus.

The Travunioidea are a group of Holarctic laniatorean harvestmen characterised by male genitalia in which the expansive muscles tend to be restricted to the base of the truncus, often associated with a simplification of the glans (Derkarabetian et al. 2018).

Travunioidea [Travunoidea]
    |--Travuniidae [Travuniinae] DS18
    |    |--Dinaria Roewer 1935 KM07
    |    |    `--*D. vjetrenicae (Hadži 1932) [=Travunia vjetrenicae] KM07
    |    |--Trojanella Karaman 2005 DS18
    |    |    `--T. serbica Karaman 2005 KMS14
    |    `--Travunia Absolon 1920 [=Absolonia Roewer 1915 non Börner 1901, Abasola Strand 1928] KM07
    |         |--T. troglodytes (Roewer 1915) KMS14 (see below for synonymy)
    |         |--T. borisi (Hadži 1973) KMS14 [=Abasola borisi K07]
    |         |--T. hofferi (Šilhavý 1937) KMS14 [=Abasola hofferi KM07]
    |         `--T. jandai Kratochvíl 1937 KMS14
    `--+--Yuria Suzuki 1964 DS18, KM07
       |    `--Y. pulcra Suzuki 1964 KMS14
       |         |--Y. p. pulcra KMS14
       |         `--Y. p. briggsi Suzuki 1975 KMS14
       |--Cladonychiidae DS18
       `--+--Cryptomastridae DS18
          |    |--Cryptomaster Briggs 1969 UD05
          |    |    |--*C. leviathan Briggs 1969 K03
          |    |    `--C. behemoth Starrett & Derkarabetian 2016 DS18
          |    `--Speleomaster Briggs 1974 DS18
          |         |--*S. lexi Briggs 1974 K03
          |         `--S. pecki Briggs 1974 KMS14
          `--Paranonychidae (see below for synonymy) DS18
               |--Paranonychus Briggs 1971 DS18, K03
               |    |--*P. brunneus (Banks 1893) K03 (see below for synonymy)
               |    |--P. concolor Briggs 1971 K03
               |    `--P. fuscus (Suzuki 1976) [=Mutsunonychus fuscus] DS18
               `--+--+--Metanonychus DS18
                  |  `--Sclerobunus Banks 1893 DS18, K03 (see below for synonymy)
                  |       |  i. s.: S. jemez Derkarabetian & Hedin 2014 KMS14
                  |       |--+--S. idahoensis Briggs 1971 [=S. robustus idahoensis] DH14
                  |       |  `--S. nondimorphicus Briggs 1971 DH14
                  |       `--+--+--S. cavicolens (Banks 1905) (see below for synonymy) DH14
                  |          |  `--+--S. madhousensis (Briggs 1971) (see below for synonymy) DH14
                  |          |     `--S. ungulatus (Briggs 1971) [=Cyptobunus ungulatus] DH14
                  |          `--+--+--S. speoventus Derkarabetian & Hedin 2014 DH14
                  |             |  `--S. steinmanni Derkarabetian & Hedin 2014 DH14
                  |             `--+--*S. robustus (Packard 1877) (see below for synonymy) DH14
                  |                `--+--S. skywalkeri Derkarabetian & Hedin 2014 DH14
                  |                   `--+--S. glorietus Briggs 1971 [=S. robustus glorietus] DH14
                  |                      `--S. klomax Derkarabetian & Hedin 2014 DH14
                  `--+--+--Kaolinonychus Suzuki 1975 DS18, S75
                     |  |    `--*K. coreanus (Suzuki 1966) [=Peltonychia coreana] S75
                     |  |         |--K. c. coreanus S75
                     |  |         `--K. c. longipes (Suzuki 1966) [=Peltonychia coreana longipes] S75
                     |  `--Metanippononychus Suzuki 1975 DS18, S75
                     |       |  i. s.: M. daisenensis Suzuki 1975 S75
                     |       |         M. iyanus Suzuki 1975 S75
                     |       |--M. iriei Suzuki 1975 DS18, S75
                     |       |    |--M. i. iriei S75
                     |       |    `--M. i. yakuensis Suzuki 1975 S75
                     |       `--+--*M. tomishimai Suzuki 1975 S75, DS18, S75
                     |          |    |--M. t. tomishimai S75
                     |          |    `--M. t. awanus Suzuki 1975 S75
                     |          `--Nippononychus Suzuki 1975 DS18, S75
                     |               `--*N. japonicus (Miyoshi 1957) S75, KMS14 [=Peltonychia japonica S75]
                     `--+--Zuma Goodnight & Goodnight 1942 DS18, K03
                        |    |--*Z. acuta Goodnight & Goodnight 1942 [=Z. acuba (l. c.)] K03
                        |    `--Z. tioga Briggs 1971 K03
                        `--+--Kainonychus Suzuki 1975 DS18, S75
                           |    `--*K. akamai (Suzuki 1972) [=Peltonychia akamai] S75
                           |         |--K. a. akamai S75
                           |         `--K. a. esoensis Suzuki 1975 S75
                            `--Izunonychus Suzuki 1975 DS18, S75
                                 `--*I. ohruii Suzuki 1975 S75

Paranonychidae [Kaolinonychinae, Nippononychidae, Nippononychinae, Paranonychinae, Paranychinae, Sclerobunidae, Sclerobuninae] DS18

*Paranonychus brunneus (Banks 1893) K03 [=Sclerobunus brunneus K03; incl. S. parvus Roewer 1931 BH09]

Sclerobunus Banks 1893 DS18, K03 [incl. Cyptobunus Banks 1905 K03, Cryptobunus (l. c.) non Lawrence 1931 K03]

Sclerobunus cavicolens (Banks 1905) [=*Cyptobunus cavicolus, Cyp. cavicolens, Cytobunus (l. c.) cavicolens] DH14

Sclerobunus madhousensis (Briggs 1971) [=Cyptobunus ungulatus madhousensis, Cryptobunus (l. c.) ungulatus madhousensis, Cytobunus (l. c.) ungulatus madhousensis] DH14

*Sclerobunus robustus (Packard 1877) [=Scotolemon robustum, Phalangodes robusta, P. robustus] DH14

Travunia troglodytes (Roewer 1915) KMS14 [=*Absolonia troglodytes KM07, *Abasola troglodytes KM07, Scotolemon anophthalmum Absolon 1916 (n. n.) KM07, *Travunia anophthalma Absolon 1920 KM07]

*Type species of generic name indicated

REFERENCES

[BH09] Bragg, P. D., & R. G. Holmberg. 2009. The harvestmen (Arachnida, Opiliones) of British Columbia. J. Entomol. Soc. Brit. Columbia 106: 29–37.

[DH14] Derkarabetian, S., & M. Hedin. 2014. Integrative taxonomy and species delimitation in harvestmen: a revision of the western North American genus Sclerobunus (Opiliones: Laniatores: Travunioidea). PLoS One 9 (8): e104982.

[DS18] Derkarabetian, S., J. Starrett, N. Tsurusaki, D. Ubick, S. Castillo & M. Hedin. 2018. A stable phylogenomic classification of Travunioidea (Arachnida, Opiliones, Laniatores) based on sequence capture of ultraconserved elements. ZooKeys 760: 1–36.

[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.

[K07] Kury, A. B. 2007. Travuniidae Absolon and Kratochvíl, 1932. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 237–239. Harvard University Press: Cambridge (Massachusetts).

[KM07] Kury, A. B., & A. C. Mendes. 2007. Taxonomic status of the European genera of Travuniidae (Arachnida, Opiliones, Laniatores). Munis Entomology & Zoology 2 (1): 1–14.

[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.

[S75] Suzuki, S. 1975. The harvestmen of family Triaenonychidae in Japan and Korea (Travunoidea, Opiliones, Arachnida). Journal of Science of the Hiroshima University, Series B, Division 1 (Zoology) 26: 65–101.

[UD05] Ubick, D., & J. A. Dunlop. 2005. On the placement of the Baltic amber harvestman Gonyleptes nemastomoides Koch & Berendt, 1854, with notes on the phylogeny of Cladonychiidae (Opiliones, Laniatores, Travunioidea). Mitteilungen Museum für Naturkunde, Geowissenschaftlichen Reihe 8: 75–82.

Samooidea

Features of Anaceros pauliani, from Lawrence (1959).

Belongs within: Grassatores.
Contains: Stenostygninae, Hinzuanius, Metabiantes, Biantes, Samoidae, Stygnomma.

The Samooidea are a group of laniatorean harvestmen found in Africa and the Neotropics that commonly possess a scopula and a sexually dimorphic metatarsus III (Sharma & Giribet 2011). Families of the Samooidea include the Biantidae, in which the eyes are not borne on a central eyemound but widely separated on the carapace.

See also: Biantidae: the importance of titillators.

<==Samooidea [Biantoidea]
    |--Biantidae [Biantoidae, Hinzuanidae] SG11
    |    |--Zairebiantes Kauri 1985 S92 [Zairebiantinae KPG07]
    |    |    `--*Z. microphthalmus Kauri 1985 S92
    |    |--Lacurbsinae KSP-G15
    |    |    |--Lacurbs Sørensen 1896 S92 [Lacurbsinae KPG07]
    |    |    |    |--L. nigrimana Roewer 1912 [=L. nigrimanus] S92
    |    |    |    `--L. spinosa Sørensen 1896 S92
    |    |    |--Eulacurbs Roewer 1949 KSP-G15, S92
    |    |    |    `--*E. paradoxa Roewer 1949 S92
    |    |    |--Metalacurbs Roewer 1914 KSP-G15, S92
    |    |    |    |--*M. simoni Roewer 1915 S92, KSP-G15
    |    |    |    |--M. cornipes (Roewer 1958) [=Prolacurbs cornines (l. c.)] S92
    |    |    |    |--M. oedipus (Roewer 1958) [=Prolacurbs oedipus] S92
    |    |    |    `--M. villiersi (Roewer 1953) [=Prolacurbs villiersi] S92
    |    |    `--Prolacurbs Roewer 1949 KSP-G15, S92
    |    |         `--*P. singularis Roewer 1949 S92
    |    |--Stenostygninae P-GAB09
    |    `--Biantinae KPG07
    |         |--Hinzuanius R23
    |         |--Metabiantes KPG07
    |         |--Eubiantes Roewer 1915 R23, S92
    |         |    `--*E. africanus Roewer 1915 S92
    |         |--Acudorsum Loman 1902 R23, S92
    |         |    `--*A. albimanum Loman 1902 S92
    |         |--Hoplobiantes Roewer 1915 R23
    |         |--Cryptobiantes Kauri 1961 K61
    |         |    `--*C. protector Kauri 1961 S92
    |         |--Fageibiantes Roewer 1949 KPG07, S92
    |         |    |--*F. bicornis (Fage 1946) [=Biantes bicornis, Fageibantes (l. c.) bicornis] S92
    |         |    `--F. bispina Lawrence 1959 [=Fageibantes (l. c.) bispina] S92
    |         |--Biantessus Roewer 1949 K61, S92
    |         |    |--*B. nigrotarsus (Lawrence 1933) [=Metabiantes nigrotarsus] S92
    |         |    `--B. vertebralis (Lawrence 1933) S92 [=Metabiantes vertebralis S92, *Biantanius vertebralis L63]
    |         |--Biantes S89
    |         |--Monobiantes Lawrence 1962 L62
    |         |    `--*M. benoiti Lawrence 1962 S92
    |         |--Biantomma Roewer 1942 L62, S92
    |         |    `--*B. nigrospinosum Roewer 1942 S92
    |         |--Clinobiantes Roewer 1927 L62, S92
    |         |    `--*C. paradoxus Roewer 1927 S92
    |         |--Biantella Roewer 1927 L62, S92
    |         |    `--*B. reticulata Roewer 1927 S92
    |         |--Anaceros Lawrence 1959 KSP-G15
    |         |    |--*A. humilis Lawrence 1959 S92
    |         |    |--A. anodonta Lawrence 1959 S92
    |         |    |--A. canidens Lawrence 1959 S92
    |         |    `--A. pauliani Lawrence 1959 S92
    |         |--Hirstienus Roewer 1949 KSP-G15, SPG11
    |         |    `--*H. nanus (Hirst 1913) [=Phalangodes nanus] SPG11
    |         |--Hovanoceros Lawrence 1959 KSP-G15, S92
    |         |    `--*H. bison Lawrence 1959 S92
    |         |--Ivobiantes Lawrence 1965 KSP-G15, S92
    |         |    `--*I. spinipalpis Lawrence 1965 S92
    |         |--Malgaceros Lawrence 1959 KSP-G15, KP-G15
    |         |    `--*M. boviceps Lawrence 1959 S92
    |         |--Probiantes croceus Roewer 1927 KSP-G15
    |         `--Tetebius Roewer 1949 KSP-G15, KP-G15
    |              `--*T. latibunus Roewer 1949 S92
    `--+--Samoidae SG11
       `--Stygnommatidae SG11
            |--Stygnomma SG11
            `--Stygnomimus Roewer 1927 SG11, KP-G15
                 |--*S. conopygus Roewer 1927 [=Stignomimus conopygus] KP-G15
                 `--S. malayensis Suzuki 1969 [=Stignomimus malayensis] KP-G15

Nomen nudum: Lacurbs fernandopoensis Prieto 1999 SP10

*Type species of generic name indicated

REFERENCES

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

[KPG07] Kury, A. B., & Pérez González, A. 2007. Biantidae Thorell, 1889. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 176–179. Harvard University Press: Cambridge (Massachusetts).

[KP-G15] Kury, A. B., & A. Pérez-González. 2015. A companion to part 2 of the world checklist of Opiliones species (Arachnida): Laniatores—Samooidea, Zalmoxoidea and Grassatores incertae sedis. Biodiversity Data Journal 3: e6663.

[KSP-G15] Kury, A. B., D. R. Souza & A. Pérez-González. 2015. World checklist of Opiliones species (Arachnida). Part 2: Laniatores—Samooidea, Zalmoxoidea and Grassatores incertae sedis. Biodiversity Data Journal 3: e6482.

[L62] Lawrence, R. F. 1962. Resultats scientifiques des missions zoologiques de l'I.R.S.A.C. en Afrique orientale (P. Basilewsky et N. Leleup, 1957). LXXIV. Opiliones. Annales du Musée Royal de l'Afrique Centrale: Sciences Zoologiques 110: 9–89.

[L63] Lawrence, R. F. 1963. The Opiliones of the Transvaal. Annals of the Transvaal Museum 24 (4): 275–304.

[P-GAB09] Pérez-González, A., & A. Alegre Barroso. 2009. On the enigmatic Heterolacurbs ovalis Roewer, 1912 (Opiliones, Laniatores, Biantidae). Zootaxa 2269: 65–67.

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

[SP10] Santos, R., & C. E. Prieto. 2010. Los Assamiidae (Opiliones: Assamiidae) de Río Muni (Guinea Ecuatorial), con la descripción de ocho nuevas especies. Revista de Biologia Tropical 58 (1): 203–243.

[SG11] Sharma, P. P. & G. Giribet. 2011. The evolutionary and biogeographic history of the armoured harvestmen—Laniatores phylogeny based on ten molecular markers, with the description of two new families of Opiliones (Arachnida). Invertebrate Systematics 25: 106–142.

[SPG11] Sharma, P. P., C. E. Prieto & G. Giribet. 2011. A new family of Laniatores (Arachnida: Opiliones) from the Afrotropics. Invertebrate Systematics 25: 143–154.

[S89] Staręga, W. 1989. Harvestmen (Opiliones) from the Mascarene Islands Indian Ocean and resurrection of the family Zalmoxidae. Annals of the Natal Museum 30: 1–8.

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

Grassatores

Microcina edgewoodensis, copyright Friends of Edgewood.

Belongs within: Laniatores.
Contains: Sandokanidae, Epedanoidea, Assamioidea, Zalmoxoidea, Samooidea, Gonyleptoidea, Calicina, Sitalcina, Banksula, Texella.

The Grassatores are a clade of laniatorean harvestmen in which the penis lacks intrinsic muscles, with the glans expanding through hydraulic pressure alone.

Grassatores [Camptonoti] SG14
    |--+--Sandokanidae SG11
    |  `--+--Epedanoidea SG14
    |     `--+--+--Assamioidea SG14
    |        |  `--+--Zalmoxoidea SG11
    |        |     `--Samooidea SG11
    |        `--Gonyleptoidea SG14
    `--Phalangodidae [Lolinae, Paralolidae, Phalangodinae, Phalangodoidea] GV09
         |  i. s.: Ausobskya Martens 1972 UB08
         |           |--A. athos Martens 1972 T96
         |           |--A. brevipes Thaler 1996 T96
         |           |--A. hauseri Silhavy 1976 T96
         |           `--A. mahnerti Silhavy 1976 T96
         |         Idzubius akiyamae TI87
         |         Spalicus oeditarsus Roewer 1949 S01, KSP-G15
         |         Bindoona glauerti Roewer 1929 K54, KSP-G15
         |         Paralola Kratochvil 1951 K18
         |           `--P. buresi Kratochvíl 1951 UD05
         |         Haasus Roewer 1942 U07, M76
         |           `--H. judaeus Roewer 1949 S02
         |         Babrius Thorell 1890 R23
         |           `--B. murcidus Thorell 1891 KSP-G15
         |         Heterobabrius Roewer 1915 R23
         |           |--H. atroluteus Roewer 1915 R23
         |           `--H. granulatus Roewer 1927 KSP-G15
         |         Biconibunus Rower 1915 R23
         |           `--B. fuscus Roewer 1915 R23
         |         Euzalmoxis Roewer 1915 R23
         |           `--E. neoguinensis Roewer 1915 R23
         |         Hoplozalmoxis Roewer 1915 R23
         |           `--H. pallida Roewer 1915 R23
         |         Parazalmoxida Roewer 1916 R23
         |           `--P. solitaria Roewer 1916 R23
         |         Proscotolemon Roewer 1916 U07, S73
         |           `--*P. sauteri Roewer 1916 S73
         |                |--P. s. sauteri S73
         |                `--P. s. latens Suzuki 1973 S73
         |         Metaconomma Pickard-Cambridge 1905 R23, K03
         |           `--*M. femorale Pickard-Cambridge 1905 K03
         |         Belonisculus Roewer 1923 R23
         |           `--*B. jacobsoni Roewer 1923 R23
         |         Isaeolus Roewer 1954 R54, K03
         |           `--*I. zilchi Roewer 1954 K03
         |         Glennhuntia Shear 2001 S01
         |           `--*G. glennhunti Shear 2001 S01
         |         Phalangodinus Roewer 1912 R23, K03
         |           `--*P. surinamensis Roewer 1912 K03
         |         Metapellobunus Roewer 1923 R23, K03
         |           |--*M. unicolor (Roewer 1912) [=Pellobunus unicolor] K03
         |           `--M. lapidosus Roewer 1928 KSP-G15
         |         Neoscotolemon Roewer 1912 R23, K03
         |           |--*N. pictipes (Banks 1908) [=Scotolemon pictipes] K03
         |           `--N. lutzi Goodnight & Goodnight 1942 K03
         |         Pseudomitraceras Roewer 1912 R23, KA-Z11
         |           |--*P. brasiliense Roewer 1912 K03, KA-Z11 (see below for synonymy)
         |           |--P. curvatum Goodnight & Goodnight 1942 KA-Z11 [=P. curvatus K03]
         |           `--P. minutum Goodnight & Goodnight 1942 KA-Z11 [=P. minutus K03]
         |         Paraconomma Roewer 1915 R23, K03
         |           |--*P. argentinum Roewer 1915 [=P. argentina] K03
         |           |--P. ovale Goodnight & Goodnight 1942 [=P. ovala] K03
         |           `--P. spinooculorum Goodnight & Goodnight 1942 K03
         |         Guerrobunus Goodnight & Goodnight 1945 U07, K03 [incl. Caecoa Šilhavý 1974 K03]
         |           |--*G. minutus Goodnight & Goodnight 1945 [=Cynortina minutus, Dapessus minutus] K03
         |           |--G. arganoi (Šilhavý 1974) [=*Caecoa arganoi] K03
         |           `--G. vallensis Vázquez & Cokendolpher 1997 K03
         |         Savoa Forster 1949 F49
         |           `--*S. bonka Forster 1949 F49
         |         Ptychosoma Sørensen 1873 UO05, R23
         |           |--P. catalonicum T96
         |           `--P. vitellinum Sørensen 1873 T96, R23 [=Scotolemon vitellinus R23]
         |         Metagjellerujia Goodnight & Goodnight 1947 GG47
         |           `--*M. jewetti Goodnight & Goodnight 1947 GG47
         |         Foella Goodnight & Goodnight 1948 GG48
         |           `--*F. remingtoni Goodnight & Goodnight 1948 GG48
         |         Spinolatum Goodnight & Goodnight 1942 GG42a
         |           `--*S. mediale Goodnight & Goodnight 1942 GG42a, KA-Z11 [=S. medialis GG42a]
         |         Lola Kratochvíl 1937 non Hamel & Hamel 1929 (ICBN) UO05
         |           `--*L. insularis Kratochvíl 1937 UO05
         |         Nimbadus Roewer 1953 S92
         |           `--*N. femoralis Roewer 1953 SPG11
         |         Epedanestus Roewer 1938 W53
         |           `--E. geniculatus (Pocock 1903) W53 (see below for synonymy)
         |         Tweedielus S72
         |           |--T. brevipes Roewer 1949 S72
         |           `--T. longipes Roewer 1933 KSP-G15
         |--Remyus Roewer 1949 SG11, S92
         |    `--R. gracillimus Roewer 1949 S92
         `--+--Scotolemon Lucas 1860 SG11, M78
            |    |--*S. lespesii Lucas 1860 M78 [=Phalangodes lespesi R23, Ptychosoma lespesi R23]
            |    |--S. balearicus Rambla 1977 T96
            |    |--S. doriae Pavesi 1878 M78
            |    |--S. jaqueti Roewer 1912 R23
            |    |--S. krausi T96
            |    |--S. lucasi Simon 1872 [=Phalangodes lucasi] R23
            |    |--S. roeweri T96
            |    `--S. terricola Simon 1872 [=Phalangodes terricola] R23
            `--+--Calicina UB08
               `--+--Microcinella Ubick & Briggs 2008 UB08
                  |    |--*M. homi (Briggs & Ubick 1989) [=Microcina homi] UB08
                  |    `--M. coensis Ubick & Briggs 2008 UB08
                  `--+--+--+--Sitalcina UB08
                     |  |  `--Enigmina Ubick & Briggs 2008 UB08
                     |  |       |--*E. granita (Briggs 1968) [=Sitalcina granita] UB08
                     |  |       `--E. warrenorum Ubick & Briggs 2008 UB08
                     |  `--Microcina Briggs & Ubick 1989 UB08
                     |       |--*M. tiburona (Briggs & Hom 1966) [=Sitalcina tiburona] UB08
                     |       |--M. edgewoodensis Briggs & Ubick 1989 UB08
                     |       |--M. jungi Briggs & Ubick 1989 UB08
                     |       |--M. leei Briggs & Ubick 1989 UB08
                     |       |--M. lumi Briggs & Ubick 1989 UB08
                     |       |--M. potrero Ubick & Briggs 2008 UB08
                     |       |--M. sanbruno Ubick & Briggs 2008 UB08
                     |       |--M. stanford Ubick & Briggs 2008 UB08
                     |       `--M. tamalpais Ubick & Briggs 2008 UB08
                     `--+--Tularina Ubick & Briggs 2008 UB08
                        |    |--*T. tularensis Ubick & Briggs 2008 UB08
                        |    |--T. plumosa Ubick & Briggs 2008 UB08
                        |    `--T. scopula (Briggs 1968) [=Sitalcina scopula] UB08
                        `--+--Megacina Ubick & Briggs 2008 UB08
                           |    |--*M. cockerelli (Goodnight & Goodnight 1942) [=Sitalcina cockerelli] UB08
                           |    `--+--M. madera (Briggs 1968) [=Sitalcina madera] UB08
                           |       |--M. mayacma Ubick & Briggs 2008 UB08
                           |       `--M. schusteri Ubick & Briggs 2008 UB08
                           `--+--Banksula UB08
                              |--Texella UB08
                              |--Tolus Goodnight & Goodnight 1942 UB08, GG42b
                              |    `--*T. appalachius Goodnight & Goodnight 1942 GG42b [=Phalangodes appalachius K03]
                              |--Undulus Goodnight & Goodnight 1942 UB08, GG42b
                              |    `--*U. formosus Goodnight & Goodnight 1942 GG42b [=Phalangodes formosus K03]
                              |--Wespus Goodnight & Goodnight 1942 UB08, GG42b
                              |    `--*W. arkansasensis Goodnight & Goodnight 1942 GG42b (see below for synonymy)
                              |--Bishopella Roewer 1927 UB08, K03
                              |    |--*B. laciniosa (Crosby & Bishop 1924) K03, GG42b (see below for synonymy)
                              |    `--B. jonesi Goodnight & Goodnight 1942 GG42b
                              |--Phalangodes Tellkampf 1844 UB08, K03 (see below for synonymy)
                              |    |--*P. armata Tellkampf 1844 (see below for synonymy) K03
                              |    `--P. flavipes (Banks 1908) [=Scotolemon flavipes] K03
                              `--Crosbyella Roewer 1927 UB08, K03 (see below for synonymy)
                                   |--*C. spinturnix (Crosby & Bishop 1924) K03, GG42b [=Phalangodes spinturnix GG42b]
                                   |--C. distincta Goodnight & Goodnight 1942 GG42b (see below for synonymy)
                                   |--C. montana Goodnight & Goodnight 1942 GG42b [=Phalangodes montana K03]
                                   |--C. roeweri Goodnight & Goodnight 1942 GG42b (see below for synonymy)
                                   `--C. tuberculata Goodnight & Goodnight 1942 GG42b [=Phalangodes tuberculata K03]

Grassatores incertae sedis:
  Anamota Šilhavý 1979 K03
    `--*A. custodiens Šilhavý 1979 K03
  Belemarua Roewer 1949 K03
    `--*B. nitens (Soares & Soares 1946) [=Paramitraceras nitens, Timoleon nitens] K03
  Bissopius Roewer 1949 K03
    `--*B. rugosus Roewer 1949 K03
  Caecobunus Roewer 1927 K03
    `--*C. termitarum Roewer 1927 K03
  Caribula Šilhavý 1979 K03
    `--*C. longimana Šilhavý 1979 K03
  Cleombrotus Sørensen 1932 K03
    `--*C. minutus Sørensen 1932 K03
  Contuor Roewer 1963 K03
    `--*C. novum Roewer 1963 K03
  Detlefilus Roewer 1949 K03
    `--*D. trispinosus Roewer 1949 K03
  Ignacianulus Roewer 1957 K03
    `--*I. pumilio Roewer 1957 K03
  Jimeneziella Kury & Alonso-Zarazaga 2011 KA-Z11
    |--*J. decui Avram 1970 K03
    `--J. negreai Avram 1970 K03
  Alpazia Özdikmen & Kury 2006 [=Lapazia Roewer 1949 non Ferris 1937] OK06
    `--*A. minima (Roewer 1949) [=*Lapazia minima] OK06
  Manuelangelia Kury & Alonso-Zarazaga 2011 KA-Z11 (see below for synonymy)
    `--*M. tuberosa (González-Sponga 1998) KA-Z11 [=*Limonia tuberosa K03]
  Liomma Roewer 1959 K03
    `--*L. laeve Roewer 1959 K03
  Micrisaeus Roewer 1957 K03
    `--*M. gracillimus Roewer 1957 K03
  Mirda Šilhavý 1973 K03
    `--*M. insulanus (Banks 1901) [=Stygnus insulanus] K03
  Munis Roewer 1963 K03
    `--*M. multivirgatus Roewer 1963 K03
  Octophthalmus Wood 1869 K03
    `--*O. marginatus Wood 1869 (n. d.) K03
  Ortizia Roewer 1952 K03
    `--*O. gracilipes Roewer 1952 K03
  Neoparalus Özdikmen 2006 [=Paralus Roewer 1949 non Rafinesque 1815] O06
    `--*N. granitus (Roewer 1949) [=*Paralus granitus] O06
  Pentos Roewer 1952 K03
    `--*P. pygoplus Roewer 1952 K03
  Pucallpana Avram & Soares 1983 K03
    `--*P. pullex Avram & Soares 1983 K03
  Siryseus Roewer 1949 K03
    `--*S. tibialis Roewer 1949 K03
  Tarmaops Roewer 1956 K03
    `--*T. koepckei Roewer 1956 K03
  Valifema Šilhavý 1979 K03
    `--*V. blanda Šilhavý 1979 K03

*Bishopella laciniosa (Crosby & Bishop 1924) K03, GG42b [=Phalangodes laciniosa GG42b; incl. B. marianna Goodnight & Goodnight 1942 K03]

Crosbyella Roewer 1927 UB08, K03 [incl. Goodnightiella Roewer 1949 non Soares & Soares 1945 K03]

Crosbyella distincta Goodnight & Goodnight 1942 GG42b [=*Goodnightiella distincta K03, Phalangodes distincta K03]

Crosbyella roeweri Goodnight & Goodnight 1942 GG42b [=Crosbiella (l. c.) roeweri K03, Phalangodes roeweri K03]

Epedanestus geniculatus (Pocock 1903) W53 [=Epedanus geniculatus R23, Heterobiantes geniculatus W53, Pseudobiantes geniculatus W53]

Manuelangelia Kury & Alonso-Zarazaga 2011 KA-Z11 [=Limonia González-Sponga 1998 nec Meigen 1803 nec Agassiz 1846 nec Carvalho 1985 K03]

Phalangodes Tellkampf 1844 UB08, K03 [=Acanthocheir Lucas 1861 K03, KA-Z11; incl. Phrixis Cope 1872 K03]

*Phalangodes armata Tellkampf 1844 [=*Acanthocheir armata, A. armatum; incl. *Phrixis longipes Cope 1872, Phryxus (l. c.) longipes] K03

*Pseudomitraceras brasiliense Roewer 1912 K03, KA-Z11 [=Preudomitraceras (l. c.) brasiliensis K03, Ps. brasiliensis K03]

*Wespus arkansasensis Goodnight & Goodnight 1942 GG42b [=Phalangodes arkansasensis K03]

*Type species of generic name indicated

REFERENCES

[F49] Forster, R. R. 1949. Opiliones from the Solomon Islands. Records of the Australian Museum 22 (2): 141–147.

[GV09] Giribet, G., L. Vogt, A. Pérez González, P. Sharma & A. B. Kury. 2009. A multilocus approach to harvestman (Arachnida: Opiliones) phylogeny with emphasis on biogeography and the systematics of Laniatores. Cladistics 25: 1–30.

[GG42a] Goodnight, C. J., & M. L. Goodnight. 1942a. Phalangids from British Guiana. American Museum Novitates 1167: 1–13.

[GG42b] Goodnight, C. J., & M. L. Goodnight. 1942b. New Phalangodidae (Phalangida) from the United States. American Museum Novitates 1188: 1–18.

[GG47] Goodnight, C. J., & M. L. Goodnight. 1947. Report on a collection of phalangids from New Guinea. Transactions of the American Microscopical Society 66 (4): 328–338.

[GG48] Goodnight, C. J., & M. L. Goodnight. 1948. New phalangids from the Southwest Pacific. American Museum Novitates 1371: 1–14.

[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.

[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.

[K18] Kury, A. B. 2018. Familial nomina in harvestmen (Arachnida, Opiliones). Bionomina 13: 1–27.

[KA-Z11] Kury, A. B. & M. A. Alonso-Zarazaga. 2011. Addenda and corrigenda to the "Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones)". Zootaxa 3034: 47–68.

[KSP-G15] Kury, A. B., D. R. Souza & A. Pérez-González. 2015. World checklist of Opiliones species (Arachnida). Part 2: Laniatores—Samooidea, Zalmoxoidea and Grassatores incertae sedis. Biodiversity Data Journal 3: e6482.

[M76] Martens, J. 1976. Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida: Opiliones). Entomologica Germanica 3 (1–2): 51–68.

[M78] Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[O06] Özdikmen, H. 2006. Nomenclatural changes for some Laniatores (Opiliones) genera: New substitute names and new combinations. Mun. Ent. Zool. 1 (1): 63–68.

[OK06] Özdikmen, H., & A. B. Kury. 2006. Three homonymous generic names in Araneae and Opiliones. Journal of Arachnology 34 (1): 279–280.

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

[R54] Roewer, C. F. 1954. Spinnentiere aus El Salvador, I. (Arachnoidea: Pedipalpi, Solifuga, Opiliones-Laniatores). Senckenbergiana Biologica 35: 57–73.

[S02] Schwendinger, P. J. 2002. Penis morphology in Oncopodidae (Opiliones, Laniatores): Evolutionary trends and relationships. Journal of Arachnology 30 (2): 425–434.

[SG11] Sharma, P. P., & G. Giribet. 2011. The evolutionary and biogeographic history of the armoured harvestmen—Laniatores phylogeny based on ten molecular markers, with the description of two new families of Opiliones (Arachnida). Invertebrate Systematics 25: 106–142.

[SG14] Sharma, P. P., & G. Giribet. 2014. A revised dated phylogeny of the arachnid order Opiliones. Frontiers in Genetics 5 (255): 1–13.

[SPG11] Sharma, P. P., C. E. Prieto & G. Giribet. 2011. A new family of Laniatores (Arachnida: Opiliones) from the Afrotropics. Invertebrate Systematics 25: 143–154.

[S01] Shear, W. A. 2001. Two new cave-dwelling harvestmen from Western Australia (Arachnida: Opiliones: Assamiidae and “Phalangodidae”). Records of the Western Australian Museum Supplement 64: 153–158.

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

[S72] Suzuki, S. 1972. Opiliones of Semangkok Forest Reserve, Malaysia. Journal of Science of the Hiroshima University, Series B, Division 1 (Zoology) 24 (1): 1–37.

[S73] Suzuki, S. 1973. Opiliones from the South-west Islands, Japan. Journal of Science of the Hiroshima University, Series B, Division 1 (Zoology) 24: 205–279.

[T96] Thaler, K. 1996. Neue Funde europäischer Krallenweberknechte (Arachnida, Opiliones: Phalangodidae, Travuniidae). Berichte des Naturwissenschaftlich-Medizinischen Vereins in Innsbruck 83: 135–148.

[TI87] Tsurusaki, N., & H. Ikeda. 1987. Opiliones of the Hakone district, Japan. Report of the Owakidani Natural History Museum 7: 1–16.

[U07] Ubick, D. 2007. Phalangodidae Simon, 1879. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 217–221. Harvard University Press: Cambridge (Massachusetts).

[UB08] Ubick, D., & T. S. Briggs. 2008. The harvestman family Phalangodidae. 6. Revision of the Sitalcina complex (Opiliones: Laniatores). Proceedings of the California Academy of Sciences, series 4, 59 (1): 1–108.

[UD05] Ubick, D., & J. A. Dunlop. 2005. On the placement of the Baltic amber harvestman Gonyleptes nemastomoides Koch & Berendt, 1854, with notes on the phylogeny of Cladonychiidae (Opiliones, Laniatores, Travunioidea). Mitteilungen Museum für Naturkunde, Geowissenschaftlichen Reihe 8: 75–82.

[UO05] Ubick, D., & R. Ozimec. 2005. On the harvestman genus Lola Kratochvíl (Opiliones: Laniatores). Natura Croatica 14 (3): 161–174.

[W53] Wang F.-C. 1953. A list of Chinese spiders of the order Opiliones. Kunchong Xuebao [Acta Entomologica Sinica] 3: 503–512.

Lacinius

Lacinius horridus, copyright AfroBrazilian.

Belongs within: Phalangiidae.

Lacinius is a western Palaearctic genus of generally spiny harvestmen.

Characters (from Martens 1978): Sexual dimorphism insignificant. Pedipalp primarily denticulate on femur and patella, rarely on other segments and then sparse, all segments lacking well-developed apophysis but sometimes with medial, brush-like hairy protrusions. Cheliceral lamellae smooth. Frontal margin with three median thorn-like denticles, either equally long or the middle longest. Tuber oculorum low, one or two times its own length from frontal margin. Truncus of penis slender to stocky, continuously narrowed distad from the base, sometimes slightly enlarged sub-distally; basal part of truncus rounded; mostly dorso-distally or ventro-distally flat or dorso-ventrally flattened; glans mostly sharply keeled below; musculature occupying at least basal half of trunk; truncus never with dorso-distal longitudinal groove.

<==Lacinius Thorell 1876 MDP15 [incl. Acantholophus Koch 1839 non Boisduval 1835 C92, Bidentolophus Roewer 1912 MDP15]
    |--*L. horridus (Panzer 1794) C92 (see below for synonymy)
    |--‘Opilio’ adungius Roewer 1956 [=Bidentolophus adungius; incl. O. birmanicus Roewer 1956] S03
    |--L. angulifer (Simon 1878) MDP15 [=Acantholophus angulifer S84]
    |--L. bidens (Simon 1880) MDP15 (see below for synonymy)
    |--L. bizleyi Mitov, Dunlop & Penney 2015 MDP15
    |--L. dentiger (Koch 1848) MDP15 (see below for synonymy)
    |--L. ephippiatus (Koch 1835) MDP15 (see below for synonymy)
    |--L. erinaceus Staręga 1966 MDP15
    |--L. insularis Roewer 1923 MDP15
    `--L. longisetus (Thorell 1876) [=Acantholophus longisetus] R23

Lacinius bidens (Simon 1880) MDP15 [=Acantholophus bidens MDP15, *Bidentolophus bidens C92, Egaenus bidens TTL00, Euphalangium bidens S03; incl. Opilio hexaspinulatus Saitô 1936 S03, O. koreanus Charitonov 1957 S03, O. peipingensis Wang 1941 S03, O. tchangi Wang 1941 S03, Phalangium nordenskioeldi var. transbaicalica Kulczyński 1901 ZSK99, Euphalangium nordenskioldi var. transbaicalicum R23, TTL00, Homolophus transbaicalicus ZSK99]

Lacinius dentiger (Koch 1848) MDP15 [=Acantholophus dentiger D04; incl. A. annulipes Koch 1867 D04, Bidentolophus bavaricus Roewer 1957 D04, Lacinius chelodentatus Hadži 1973 D04, L. labacensis Hadži 1931 D04]

Lacinius ephippiatus (Koch 1835) MDP15 [=Opilio ephippiatus D04, Acantholophus ephippiatus D04, Oligolophus ephippiatus D04; incl. Odiellus hungaricus Kolosvary 1941 D04, Lacinius oligodentatus Hadži 1931 D04, Odiellus rucneri Hadži 1973 D04, Opilio vittiger Meade 1855 D04, Ol. vittiger D04]

*Lacinius horridus (Panzer 1794) C92 [=Phalangium horridum D04, Acantholophus horridus D04, Opilio horridus D04; incl. A. bellicosus Sørensen 1894 D04, Lacinius horridus bulgaricus Šilhavý 1965 D04, A. coronatus Koch 1867 D04, Opilio coronatus Roewer 1911 D04, Odiellus coronatus M78, Lacinius dorsogranulatus Morin 1931 D04, L. gallipoliensis Roewer 1923 D04, Acantholophus granulatus Canestrini 1872 D04, Odiellus granulatus D04, A. hermanni Simon 1879 D04, Opilio hispidus Herbst 1798 D04, *A. hispidus C92, Lacinius hispidus D04, Phalangium hispidum D04, Acantholophus kochi Simon 1879 D04, A. lemniscatus Simon 1882 D04, Lacinius parisii Trossarelli 1934 D04, A. spinulosus Koch 1839 D04, Odiellus toscanus Roewer 1923 M78, Lacinius toscanus M78, L. zavalensis Hadži 1973 N05]

*Type species of generic name indicated

REFERENCES

[C92] Crawford, R. L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1–60.

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[M78] Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[MDP15] Mitov, P. G., J. A. Dunlop & D. Penney. 2015. A new species of Lacinius in amber (Arachnida: Opiliones). Fossil Record 18: 37–42.

[N05] Novak, T. 2005. An overview of harvestmen (Arachnida: Opiliones) in Bosnia and Herzegovina. Natura Croatica 14 (4): 301–350.

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

[S84] Staręga, W. 1984. Revision der Phalangiidae (Opiliones), III. Die afrikanischen Gattungen der Phalangiinae, nebst Katalog aller afrikanischen Arten der Familie. Annales Zoologici 38 (1): 1–79.

[S03] Staręga, W. 2003. On the identity and synonymies of some Asiatic Opilioninae (Opiliones: Phalangiidae). Acta Arachnologica 52 (2): 91–102.

[TTL00] Tsurusaki, N., A. N. Tchemeris & D. V. Logunov. 2000. Two new species of Opiliones from southern Siberia and Mongolia, with an establishment of a new genus and redefinition of the genus Homolophus (Arachnida: Opiliones: Phalangiidae). Acta Arachnologica 49 (1): 73–86.

[ZSK99] Zhu, M., D. Song & J.-P. Kim. 1999. Notes on three species of the genus Homolophus (Opiliones: Phalangiidae) from China. Korean Arachnology 15 (1): 7–12.

Zachaeus

Zachaeus crista, copyright Ondřej Machač.

Belongs within: Phalangiidae.

Zacheus is a genus of large-bodied phalangiid harvestmen found in the eastern Mediterranean and western Asia (Kurt et al. 2015). Historically, there has been confusion regarding the correct spelling of the genus name; Crawford (1992) provides reasoning for use of the spelling 'Zachaeus'.

Characters (from Kurt et al. 2015): Body large, heavily denticulated dorsally; chelicerae usually strong, second segment enlarged; pedipalps normally structured, strong, robust; legs short, first pair much thicker than others; truncus of penis basally widened, parallel-sided on distal half, distally shallow spoon-shaped, glans usually banana-shaped (shorter in Z. anatolicus and Z. seyyari), stylus long.

<==Zachaeus Koch 1839 K14 [=Zacheus Koch 1839 C92]
    |--Z. crista (Brullé 1832) K14 (see below for synonymy)
    |--Z. anatolicus (Kulczyński 1903) K14 [=Egaenus crista var. anatolicus R23, Z. crista var. anatolicus R23]
    |--Z. birulai Redikorzev 1936 S99
    |--Z. canaliculatus Kollar in Roewer 1911 [=Phalangium canaliculatum ms] R23
    |--Z. caucasicus Roewer 1911 R23
    |--Z. cubanus Roewer 1923 R23
    |--‘Dasylobus’ egaenoides Simon 1885 (n. d.) R23
    |--Z. hebraicus (Simon 1884) K14 [=Phalangium hebraicum R23, Z. crista var. hebraicus R23]
    |--Z. kervillei (Sørensen 1912) (n. d.) KKY15 [=Egaenus kervillei R23]
    |--Z. leucomelas (Simon 1884) R23 [=Phalangium hebraicum leucomelas R11, Zachaeus crista var. leucomelas R23]
    |--Z. lupatus (Eichwald 1830) [incl. E. gulosus Simon 1878] S03
    |--Z. macrinus Roewer 1956 R56
    |--Z. mirabilis (Caporiacco 1949) KKY15
    |--Z. orchimonti (Giltay 1933) K14
    |--Z. palpipes Roewer 1915 R23
    |--‘Opilio’ punctipes Koch 1878 (n. d.) R23
    |--Z. redikorzevi (Staręga & Chevrizov 1978) K14
    |--Z. seyyari Kurt, Koç & Yağmur 2015 KKY15
    |--Z. shachdag (Snegovaya & Staręga 2008) KKY15
    `--Z. simferopolensis Chemeris & Kovblyuk 2005 [=Z. simpheropolensis] CK05

Zachaeus crista (Brullé 1832) K14 [=Phalangium crista C92, Egaenus crista C92; incl. E. hungaricus Lendl 1894 M78, Zachaeus crista var. hungaricus R23, Paropilio lineatus Roewer 1956 M78, *Zachaeus mordax Koch 1839 C92, Egaenus mordax R23, Z. trinotatus Koch 1839 R23, E. variegatus Lendl 1894 M78, Zachaeus crista var. variegatus R23]

*Type species of generic name indicated

REFERENCES

[CK05] Chemeris, A. N., & N. M. Kovblyuk. 2005. A contribution to the knowledge of the harvestman fauna of the Crimea (Arachnida: Opiliones). Arthropoda Selecta 14 (4): 305–328.

[C92] Crawford, R. L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1–60.

[K14] Kurt, K. 2014. Updated checklist of harvestmen (Arachnida: Opiliones) in Turkey. Arch. Biol. Sci. 66 (4): 1617–1631.

[KKY15] Kurt, K., H. Koç & E. A. Yağmur. 2015. A new species of Zachaeus C. L. Koch from Turkey (Opiliones, Phalangiidae). ZooKeys 514: 15–23.

[M78] Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[R11] Roewer, C.-F. 1911. Übersicht der Genera der Subfamilie der Phalangiini der Opiliones Palpatores nebst Beschreibung einiger neuer Gattungen und Arten. Archiv für Naturgesichte 77 (Suppl. 2): 1–106.

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

[R56] Roewer, C. F. 1956. Über Phalangiinae (Phalangiidae, Opiliones Palpatores). (Weitere Weberknechte XIX). Senckenbergiana Biologica 37 (3–4): 247–318.

[S99] Snegovaya, N. 1999. Contribution to the harvest spider (Arachnida, Opiliones) fauna of the Caucasus. Tr. J. of Zoology 23: 453–459.

[S03] Staręga, W. 2003. On the identity and synonymies of some Asiatic Opilioninae (Opiliones: Phalangiidae). Acta Arachnologica 52 (2): 91–102.

Assamiidae

Dorsum of Erecella flava, from Roewer (1935).

Belongs within: Grassatores.
Contains: Irumuinae, Sidaminae, Assamiinae, Polycoryphinae, Dampetrinae, Selencinae, Trionyxellinae, Hypoxestinae, Metereca, Ereca, Chilon.

The Assamiidae are a family of short-legged harvestmen found in tropical and subtropical regions of the Old World.

Characters (from Santos & Prieto 2010): Prosomal shield with five processes on frontal border (central, two internal lateral and two lateral external or marginal); pedipalps not very long, tibia and tarsus dorsoventrally flattened and crossed one on another; penis with distally enlarged trunk (ventral lamina), sometimes bilobed, with variable envelope of membranes; glans cylindrical, subdistal, housing spiny or warty evaginable membrane in the shape of a funnel.

Assamiidae [Assamioidae]
    |--+--Selencinae SG11
    |  `--+--+--Arulla SG11
    |     |  `--Tarnus pulcher SG11
    |     `--+--Trionyxellinae SG11
    |        `--+--Hypoxestinae SG11
    |           `--+--Paktongius SG11
    |              `--Mysoreinae R35
    |                   |--Typestus Roewer 1935 R35
    |                   |    `--*T. sulcatus Roewer 1935 R35
    |                   |--Attakattius Roewer 1929 R35
    |                   |    `--A. spinifrons Roewer 1929 R35
    |                   `--Mysorea Roewer 1935 SG11, R35
    |                        |--*M. brevipus Roewer 1935 R35
    |                        `--M. thaiensis Suzuki 1985 SG11
    `--Erecinae [Tsadseinae] SP10
         |--Metereca SP10
         |--Ereca SP10
         |--Chilon SG11
         |--Valpara Roewer 1929 R35
         |    `--V. albitarsis Roewer 1929 R35
         |--Kukkala Roewer 1929 R35
         |    `--K. trispinifrons Roewer 1929 R35
         |--Mandaria Roewer 1935 SP10
         |    `--*M. caeca Roewer 1935 R35
         |--Merucola Roewer 1935 R35
         |    `--*M. granulatus Roewer 1935 [=M. granulata] R35
         |--Dodabetta Roewer 1929 R35
         |    `--D. conigera Roewer 1929 R35
         |--Djemia Roewer 1935 R35
         |    `--*D. cooperi Roewer 1935 R35
         |--Wintonia Roewer 1923 SP10
         |    `--*W. scabra Roewer 1923 R35
         |--Sacesphorus Thorell 1889 R35
         |    `--*S. maculatus Thorell 1889 T89
         |--Parapygoplus Roewer 1912 R35
         |    `--P. variatus (Thorell 1889) [=Pygoplus variatus] R35
         |--Irnia Roewer 1935 R35
         |    `--*I. scabra Roewer 1935 R35
         |--Bibundina Roewer 1935 SP10
         |    `--*B. pectinata Roewer 1935 R35
         |--Baeorix Thorell 1889 R35 [=Boeorix R23]
         |    `--*B. manducus Thorell 1889 T89 [=Boeorix manducus R23]
         |--Lepchana Roewer 1927 R35
         |    `--L. spinipalpis Roewer 1927 M77
         |--Eubaeorix Roewer 1912 R35 [=Euboeorix (l. c.) R23]
         |    `--E. gravelyi Roewer 1912 R35 [=Euboeorix (l. c.) gravelyi R23]
         |--Sijucavernicus Roewer 1923 R35
         |    `--S. kempi Roewer 1923 R35
         |--Fakoa Roewer 1923 SP10
         |    `--*F. spinulata (Roewer 1915) [=Acanthophrysella spinulata] S92
         |--Buemba Roewer 1935 R35
         |    `--*B. filipes Roewer 1935 R35
         |--Tetecus Roewer 1935 R35
         |    `--*T. tenuis Roewer 1935 R35
         |--Metapygoplus Roewer 1923 R35
         |    `--*M. intermedius (Loman 1892) [=Pygoplus intermedius] R35
         |--Gapotus Roewer 1935 R35
         |    `--*G. frontalis Roewer 1935 R35
         |--Vandarawella Roewer 1935 R35
         |    `--*V. bicolor Roewer 1935 R35
         |--Cereoides Roewer 1935 SP10
         |    `--*C. nebulosa (Sørensen 1896) [=Cerea nebulosa] R35
         |--Cereipes Roewer 1935 SP10
         |    `--*C. angusta (Roewer 1912) S92 [=Cerea angusta R35]
         |--Acanthophrysella Strand 1911 R35 [=Acanthophrys Loman 1902 non Milne Edwards 1865 R23]
         |    `--*A. pectinata (Loman 1902) S92 [=Acanthophrys pectinata R35]
         |--Anjolus Goodnight & Goodnight 1948 GG48
         |    `--*A. malkini Goodnight & Goodnight 1948 GG48
         |--Mbinia Santos & Prieto 2010 SP10
         |    `--*M. xenophora Santos & Prieto 2010 SP10
         |--Cardwella Roewer 1935 SP10
         |    `--*C. atar (Sørensen in Henriksen 1932) [=Wintonia atar, Dampetrus atar ms] R35
         |--Ivocoryphus Lawrence 1965 SP10
         |    `--*I. jezequeli Lawrence 1965 S92
         |--Neocoryphus Lawrence 1965 SP10
         |    `--*N. niger Lawrence 1965 S92
         |--Cereodiscus Roewer 1935 SP10
         |    `--*C. lesserti Roewer 1940 S92
         |--Ereala Roewer 1950 SP10 [incl. Faradjea Roewer 1950 S92]
         |    `--*E. armata Roewer 1950 [incl. Faradjea schoutedeni Roewer 1950] S92
         |--Eregonda Roewer 1950 SP10
         |    `--*E. tenuis Roewer 1950 S92
         |--Termitereca Roewer 1940 SP10
         |    `--*T. singularis Roewer 1940 S92
         |--Angolyppa Lawrence 1957 SP10
         |    `--*A. scabra Lawrence 1957 S92
         |--Angopygoplus Lawrence 1951 SP10
         |    `--*A. dentichelis Lawrence 1951 S92
         |--Montereca Lawrence 1962 L62
         |    `--*M. paucidens Lawrence 1962 S92
         |--Bundukia Lawrence 1962 L62
         |    `--*B. nigra Lawrence 1962 S92
         |--Roewereca Lawrence 1962 L62
         |    `--*R. tenebrosa Lawrence 1962 S92
         |--Neobaeorix Lawrence 1962 L62
         |    `--*N. cornuta Lawrence 1962 S92
         |--Aberdereca Goodnight & Goodnight 1959 L62, S92
         |    `--*A. parva Goodnight & Goodnight 1959 S92
         |--Erebalda Roewer 1940 L62, S92
         |    `--*E. cryptostigma Roewer 1940 S92
         |--Erecops Roewer 1940 L62, S92
         |    `--*E. multispina Roewer 1940 S92
         |--Kakontwea Roewer 1951 L62, S92
         |    `--*K. leleupi Roewer 1951 S92
         |--Lubudia Roewer 1951 L62, S92
         |    `--*L. leleupi Roewer 1951 S92
         |--Kasaina Lawrence 1957 L62, S92
         |    `--*K. scabra Lawrence 1957 S92
         |--Allereca Roewer 1961 L62, S92
         |    `--*A. ruandana Roewer 1961 S92
         |--Comereca Roewer 1961 L62, S92
         |    `--*C. rectipes Roewer 1961 S92
         |--Buniabia Roewer 1961 L62, S92
         |    `--*B. filipes Roewer 1961 S92
         |--Bambereca Kauri 1985 K85
         |    `--*B. spinifrons Kauri 1985 S92
         |--Tubereca Kauri 1985 K85
         |    `--*T. biharaguana Kauri 1985 S92
         |--Neopygoplus Roewer 1923 R35
         |    |--N. jacobsoni Roewer 1923 R35
         |    `--N. siamensis Suzuki 1985 SG11
         |--Tundabia Roewer 1935 R35
         |    |--*T. semicaeca Roewer 1935 R35
         |    `--T. ugandensis Goodnight & Goodnight 1959 S92
         |--Bulobana Roewer 1935 R35
         |    |--*B. octopunctata Roewer 1935 R35
         |    `--B. infuscata Roewer 1940 S92
         |--Eupygoplus Roewer 1915 R35
         |    |--E. armatus Roewer 1915 R35
         |    `--E. gracilis Roewer 1927 R35
         |--Cerea Sørensen 1896 SP10
         |    |--*C. lugubris Sørensen 1896 S92
         |    `--C. feai Roewer 1927 S92 [=C. feae R35]
         |--Izea Roewer 1927 R35
         |    |--*I. pectinata Roewer 1927 S92
         |    `--I. armata Roewer 1935 R35
         |--Erecongoa Roewer 1950 SP10
         |    |--*E. gracilis Roewer 1950 S92
         |    `--E. granulata Roewer 1952 S92
         |--Lygippus Roewer 1940 SP10
         |    |--*L. abdominalis Roewer 1940 S92
         |    `--L. machadoi Lawrence 1951 S92
         |--Erecabia Roewer 1940 L62, S92
         |    |--*E. hartmanni Roewer 1940 S92
         |    `--E. pluridens Lawrence 1962 S92
         |--Erecomma Roewer 1940 L62, S92
         |    |--*E. montana Roewer 1940 S92
         |    `--E. laurenti Roewer 1961 S92
         |--Cereatta Roewer 1935 R35
         |    |--*C. celeripes (Loman 1910) S92 [=Cerea celeripes R35]
         |    |--C. elegans Roewer 1935 R35
         |    `--C. kivuensis Roewer 1961 S92
         |--Pygoplus Thorell 1889 R35
         |    |--*P. obscurus Thorell 1889 T89
         |    |--P. ferrugineus Thorell 1889 R35
         |    `--P. trifasciatus Thorell 1889 R35
         |--‘Macrobunus’ Roewer 1912 non Tullgren 1901 R35
         |    |--M. aborensis Roewer 1913 R35
         |    |--M. longipes Roewer 1913 R35
         |    `--M. singularis Roewer 1912 R35
         |--Callereca Roewer 1940 SP10
         |    |--*C. gracilis Roewer 1940 S92
         |    |--C. angolensis Lawrence 1949 S92
         |    `--C. teteana Roewer 1954 S92
         |--Cryptopygoplus Lawrence 1931 R35 [incl. Lawrenciola Roewer 1935 L11]
         |    |--*C. africanus Lawrence 1931 S92
         |    |--C. capriviensis Lotz 2011 L11
         |    |--C. coronatus Staręga & Snegovaya 2009 L11
         |    |--C. damaranus Lawrence 1931 [=*Lawrenciola damarana] S92
         |    `--C. rhodesianus Lawrence 1931 [=Lawrenciola rhodesiana] R35
         |--Propygoplus Roewer 1923 R35, M77 (see below for synonymy)
         |    |--*P. maculatus (Roewer 1912) [=Parapygoplus maculatus] M77
         |    |--‘*Karsianga’ rugosa Roewer 1940 M77
         |    |--‘*Sikkimula’ scabrisoma Roewer 1940 M77
         |    |--P. siwalik Martens 1977 M77
         |    `--‘*Sikkimella’ tenuipes Roewer 1927 M77
         |--Lygippulus Roewer 1954 L62, S92
         |    |--*L. major Roewer 1954 S92
         |    |--L. nigrescens Roewer 1961 S92
         |    |--L. parvulus Roewer 1954 S92
         |    |--L. scaber Roewer 1954 S92
         |    `--L. setipes Roewer 1961 S92
         |--Erecula Roewer 1935 R35
         |    |--*E. pachypes Roewer 1935 R35
         |    |--E. cincta Roewer 1961 S92
         |    |--E. crassipes Kauri 1985 S92
         |    |--E. leleupi Lawrence 1962 S92
         |    |--E. marmorata Roewer 1940 S92
         |    |--E. novemdentata Roewer 1961 S92
         |    `--E. septemdentata Lawrence 1957 S92
         `--Erecella Roewer 1935 R35
              |--*E. lutea Roewer 1935 S92
              |--E. basilewskyi Lawrence 1962 S92
              |--E. biseriata Roewer 1961 S92
              |--E. brunnea Roewer 1940 S92
              |--E. flava Roewer 1935 R35
              |--E. nigropicta Roewer 1961 S92
              |--E. signata Roewer 1950 S92
              |--E. transversalis Roewer 1961 S92
              `--E. walikaleana Kauri 1985 S92

Assamiidae incertae sedis:
  Sokodea Roewer 1935 R35
    `--*S. caeca Roewer 1935 R35
  Maccabeesa Roewer 1936 S92
    `--*M. lawrencei Roewer 1936 S92
  Aburistella Lawrence 1947 S92
    `--A. flava Lawrence 1947 S92
  Afroassamia Caporiacco 1940 S92
    `--*A. laevipes Caporiacco 1940 S92
  Bancoella Lawrence 1947 S92
    `--*B. bimaculata Lawrence 1947 S92
  Banconyx Lawrence 1947 S92
    `--*B. dentichelis Lawrence 1947 S92
  Cleoxestus Roewer 1954 S92
    `--*C. luteipictus Roewer 1954 S92
  Irumuinae SP10
  Kungwea Roewer 1961 S92
    `--K. scabra Roewer 1961 S92
  Lukandamila Roewer 1961 S92
    `--L. cookei Roewer 1961 S92
  Simienatus Roewer 1956 S92
    `--*S. scotti Roewer 1956 S92
  Typhloburista Lawrence 1947 S92
    `--*T. pusilla Lawrence 1947 S92
  Acacinae SP10
    |--Acaca Roewer 1935 R35 [incl. Pseudoacaca Caporiacco 1949 S92]
    |    |--*A. albatra Roewer 1935 R35
    |    `--A. liobuniformis (Caporiacco 1949) [=Pseudoacaca liobuniformis] S92
    `--Acanthacaca Roewer 1952 L62, S92
         |--*A. upembensis Roewer 1952 S92
         `--A. katumbea Roewer 1961 S92
  Pungoica Roewer 1914 R35 [incl. Pungoiella Roewer 1914 S92, Pygoselenca Roewer 1953 S92; Pungoicinae]
    |--*P. simoni Roewer 1914 S92
    `--P. bifurcata (Roewer 1914) (see below for synonymy) S92
  Aburistinae SP10
    |--Aburista Roewer 1935 R35
    |    `--*A. termitarum Roewer 1935 R35
    `--Typhlobunellus Roewer 1927 R35
         |--*T. formicarum Roewer 1927 S92
         `--T. platypalpis Lawrence 1947 S92
  Opcochinae R35
    |--Drugius Roewer 1929 R35
    |    `--D. parvus Roewer 1929 R35
    |--Trionyxana Roewer 1927 R35
    |    `--T. gracilipes Roewer 1927 R35
    `--Opcochina Roewer 1927 R35
         |--O. carli Roewer 1929 R35
         `--O. gravelyi Roewer 1927 R35
  Sidaminae SP10
  Assamiinae SP10
  Polycoryphinae SP10
  Dampetrinae SP10
  Parazalmoxis Roewer 1913 S92
    `--*P. africana Roewer 1913 (see below for synonymy) S92
  Nsorkia dorsicana SG11

Dicoryphus Loman 1902 R35 [incl. Coelobunus Loman 1902 S92, Caelobunus R23, Metarhabdopygus Roewer 1913 R35]

Dicoryphus melanacanthus (Loman 1902) [=Coelobunus melanacanthus, Caelobunus melanacanthus; incl. Co. fuscus Roewer 1912, Ca. fuscus] S92

*Parazalmoxis africana Roewer 1913 [incl. Hypoxestus holmi Goodnight & Goodnight 1959, P. roeweri Caporiacco 1949] S92

Propygoplus Roewer 1923 R35, M77 [incl. Karsianga Roewer 1940 M77, Sikkimella Roewer 1927 M77, Sikkimula Roewer 1940 M77]

Pungoica bifurcata (Roewer 1914) [=Pungoiella bifurcata; incl. Pygoselenca albisignata Roewer 1953, Pungoica decorata Lawrence 1947] S92

*Type species of generic name indicated

REFERENCES

[GG48] Goodnight, C. J., & M. L. Goodnight. 1948. New phalangids from the Southwest Pacific. American Museum Novitates 1371: 1–14.

[K85] Kauri, H. 1985. Opiliones from central Africa. Koninklijk Museum voor Midden-Afrika, Turveren, België, Zoologische Wetenschappen, Annalen 245: 1–167.

[L62] Lawrence, R. F. 1962. Resultats scientifiques des missions zoologiques de l'I.R.S.A.C. en Afrique orientale (P. Basilewsky et N. Leleup, 1957). LXXIV. Opiliones. Annales du Musée Royal de l'Afrique Centrale: Sciences Zoologiques 110: 9–89.

[L11] Lotz, L. N. 2011. Three new harvestmen species from southern Africa (Arachnida: Opiliones: Caddidae, Neopilionidae, Assamiidae). Journal of Afrotropical Zoology 7: 3–8.

[M77] Martens, J. 1977. Opiliones aus dem Nepal-Himalaya. III. Oncopodidae, Phalangodidae, Assamiidae (Arachnida). Senckenbergiana Biologica 57 (4–6): 295–340.

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

[R35] Roewer, C.-F. 1935. Alte und neue Assamiidae. Weitere Weberknechte VIII. (8. Ergänzung der “Weberknechte der Erde” 1923). Veröffentlichungen aus dem Deutschen Kolonial- und Uebersee-Museum in Bremen 1: 1–168.

[SP10] Santos, R., & C. E. Prieto. 2010. Los Assamiidae (Opiliones: Assamiidae) de Río Muni (Guinea Ecuatorial), con la descripción de ocho nuevas especies. Revista de Biologia Tropical 58 (1): 203–243.

[SG11] Sharma, P. P., & G. Giribet. 2011. The evolutionary and biogeographic history of the armoured harvestmen—Laniatores phylogeny based on ten molecular markers, with the description of two new families of Opiliones (Arachnida). Invertebrate Systematics 25: 106–142.

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

[T89] Thorell, T. 1889. Viaggio di Leonardo Fea in Birmania e regioni vicine. XXI.—Aracnidi Artrogastri Birmani raccolti da L. Fea nel 1885–1887. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 521–729.