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.