Sclerosomatidae

Male Leiobunum vittatum, from here.


Belongs within: Phalangioidea.
Contains: Jussara, Zaleptus, Hypsibunus, Ceratobunus, Harmanda, Gagrella, Dentobunus, Eugagrella, Strandia, Aurivilliola, Leiobunum, Holcobunus, Geaya, Prionostemma, Homalenotus, Nelima, Pseudogagrella, Psathyropus, Melanopa, Marthana, Gagrellula.

The Sclerosomatidae is a hyperdiverse group of long-legged harvestmen that is most diverse in Eurasia and the Americas, with relatively few species in Africa and Australasia. Though largely supported by molecular phylogenetic analyses, the clade is poorly defined morphologically; notable features found in many (though not all) species include a more or less straight penis, often with membranous winglets on either side of the shaft behind the glans, a ventral tooth-comb on the pedipalpal claw, and fusion of the dorsal segmentation to form a more or less hardened scutum. Most members of the Sclerosomatidae have been divided in the past between the subfamilies Sclerosomatinae, Leiobuninae and Gagrellinae, but recent phylogenetic analyses have indicated that the Leiobuninae and Gagrellinae, at least, are broadly polyphyletic (monophyly of the Sclerosomatinae remains largely uncertain).

<==Sclerosomatidae (see below for synonymy) VFG14
    |--Protolophus Banks 1893 HT12, C92 [Protolophidae, Protolophinae, Protolophini]
    |    |--*P. tuberculatus Banks 1893 C92
    |    |--P. cockerelli Goodnight & Goodnight 1942 GG42a
    |    |--P. differens Goodnight & Goodnight 1942 GG42a
    |    |--P. dixiensis Chamberlin 1925 GG42a
    |    |--P. longipes Schenkel 1951 E90
    |    |--P. niger Goodnight & Goodnight 1942 BH09
    |    |--P. rossi Goodnight & Goodnight 1943 GG43
    |    `--P. singularis Banks 1893 R23
    `--+--+--Leiobunum HT12
       |  |--+--+--Holcobunus HT12
       |  |  |  `--+--Geaya HT12
       |  |  |     |--Prionostemma HT12
       |  |  |     `--Carmenia Roewer 1915 HT12, T-D04 [=Carmenis (l. c.) C84b]
       |  |  |          `--*C. bunifrons Roewer 1915 C84b
       |  |  `--+--Trachyrhinus Weed 1892 HT12, T-D04
       |  |     |    |--*T. favosus (Wood 1868) [=Phalangium favosum, Astrobunus favosus] C81
       |  |     |    |--T. dicropalpus Cokendolpher 1981 C81
       |  |     |    |--T. horneri Cokendolpher 1981 C81
       |  |     |    |--T. marmoratus Banks 1894 [=T. maculatus (l. c.); incl. T. sonoranus Chamberlin 1925] C81
       |  |     |    |--T. mesillensis Cokendolpher 1981 C81
       |  |     |    `--T. rectipalpus Cokendolpher 1981 C81
       |  |     `--Krusa Goodnight & Goodnight 1947 HT12, C92 [incl. Krusella Roewer 1953 C92]
       |  |          |  i. s.: K. amazonica Roewer 1953 KP-R02
       |  |          |         *Krusella’ pilipes Roewer 1953 C92
       |  |          |         K. tuberculata Goodnight & Goodnight 1946 GG46
       |  |          |--*K. mexicana Goodnight & Goodnight 1947 C92, HT12, C92
       |  |          `--+--K. annulata Goodnight & Goodnight 1945 HT12, GG46
       |  |             `--K. stellata Goodnight & Goodnight 1946 HT12, GG46
       |  `--Sclerosomatinae [Sclerosomatini, Sclerosominae] C92
       |       |  i. s.: Mastobunus Simon 1879 C92
       |       |           `--*M. tuberculifer (Lucas 1847) C92, E12 (see below for synonymy)
       |       |         Granulosoma Martens 1973 K06
       |       |           `--*G. umidulum Martens 1973 C92
       |       |         Pseudastrobunus Martens 1973 K06
       |       |           `--*P. perpusillus Martens 1973 C92
       |       |         Pseudohomalenotus Caporiacco 1935 C92
       |       |           `--*P. bicornutus Caporiacco 1935 C92
       |       |         Pygobunus Roewer 1957 C92
       |       |           |--*P. formosanus Roewer 1957 C92
       |       |           `--P. okadai T07
       |       |         Metasclerosoma Roewer 1912 [=Neosclerosoma Roewer 1923; incl. Prosclerosoma Roewer 1915] C92
       |       |           |--*M. sardum (Thorell 1876) (see below for synonymy) C92
       |       |           |--M. depressum (Canestrini 1872) [=Homalenotus depressus, Sclerosoma depressum] R23
       |       |           |--M. remyi (Dresco 1950) [=Prosclerosoma remyi] D04
       |       |           `--‘Prosclerosoma’ simile Roewer 1916 [=P. similis (l. c.)] R23
       |       |--Homalenotus HT12
       |       `--+--+--Nelima HT12
       |          |  `--‘Leiobunum’ blackwalli Meade 1861 HT12, D04 (see below for synonymy)
       |          `--Astrobunus Thorell 1876 HT12, C92 (see below for synonymy)
       |               |--A. helleri (Ausserer 1867) D04 (see below for synonymy)
       |               |--A. bernardinus Simon 1879 [=A. bernardicus (l. c.); incl. A. simoni Hadzi 1928] D04
       |               |--A. dinaricus Roewer 1915 [incl. A. macedonicus Hadži 1973, A. vitynae Roewer 1957] M78
       |               |--A. grallator Simon 1879 [incl. Prosclerosoma hispanicum Mello-Leitão 1936] D04
       |               |--A. kochi Thorell 1876 D04
       |               |--A. laevipes (Canestrini 1872) B95 (see below for synonymy)
       |               `--A. spinosus (Herbst 1799) (n. d.) (see below for synonymy) R23
       `--+--+--+--‘Leiobunum’ exilipes (Wood 1870) HT12, D34 (see below for synonymy)
          |  |  `--+--‘Nelima’ elegans (Weed 1889) HT12, C84c (see below for synonymy)
          |  |     `--‘Nelima’ paessleri (Roewer 1910) HT12, D34 [=Leiobunum paessleri BH09, Liobunum paessleri R23]
          |  `--+--+--Gagrellopsis Sato & Suzuki 1939 HT12, C92
          |     |  |    `--*G. nodulifera Sato & Suzuki 1939 C92
          |     |  `--+--+--‘Leiobunum’ hikocola Suzuki 1966 HT12, T06
          |     |     |  `--+--‘Leiobunum’ globosum Suzuki 1953 HT12, T03
          |     |     |    `--‘Leiobunum’ manubriatum Karsch 1881 HT12, T06 (see below for synonymy)
          |     |     `--+--‘Leiobunum’ hiraiwai (Sato & Suzuki 1939) HT12, T06 (see below for synonymy)
          |     |        |--+--‘Leiobunum’ curvipalpe Roewer 1910 HT12, T06 [=Liobunum curvipalpi T90]
          |     |        |  `--‘Leiobunum’ tohokuense Suzuki 1976 HT12, T06
          |     |        `--+--‘Leiobunum’ kohyai Suzuki 1953 HT12, T85b [=Liobunum kohyai T85b]
          |     |           `--+--‘Leiobunum’ hiasai Suzuki 1976 HT12, T85b
          |     |              `--‘Leiobunum’ montanum Suzuki 1953 HT12, T06 (see below for synonymy)
          |     `--+--+--+--Pseudogagrella HT12
          |        |  |  `--‘Leiobunum’ japonicum Müller 1914 HT12, T03 (see below for synonymy)
          |        |  |       |--L. j. japonicum T91
          |        |  |       |--L. j. taiwanum Suzuki 1977 T91
          |        |  |       `--L. j. uenoi Suzuki 1964 T85b, S73
          |        |  `--+--Psathyropus HT12
          |        |     |--Melanopa HT12
          |        |     |--+--Marthana HT12
          |        |     |  `--Gagrellula HT12
          |        |     `--Systenocentrus Simon 1886 HT12, C92 [incl. Oncobunus Thorell 1889 C92]
          |        |          |--*S. quinquedentatus Simon 1886 C92
          |        |          |--S. confucianus Hirst 1911 R55a
          |        |          |--S. galeatus (Thorell 1889) R55a, C92 [=*Oncobunus galeatus C92]
          |        |          |--S. japonicus Hirst 1911 R55a
          |        |          |--S. luteobiseriatus Suzuki 1982 S85
          |        |          `--S. rufus Roewer 1955 R55a
          |        `--+--+--Pseudoliobunum Müller 1914 [incl. Metaliobunum Suzuki 1940] C92
          |           |  |    `--*Pseudoliobunum japanense Müller 1914 C92 [=Leiobunum japanense HT12]
          |           |  |         |--P. j. japanense C92
          |           |  |         `--‘Leiobunum’ j. suzukii Roewer 1957 (see below for synonymy) C92
          |           |  `--‘Leiobunum’ maximum Roewer 1910 HT12, T91 [=Liobunum maximum R23]
          |           |       |--L. m. maximum S77
          |           |       |--L. m. distinctum Suzuki 1973 S73
          |           |       |--L. m. ‘formosum’ Suzuki 1976 non Phalangium formosum Wood 1868 S77
          |           |       `--L. m. yushan Suzuki 1976 S77
          |           `--+--‘Leiobunum’ rubrum Suzuki 1966 HT12, TT05
          |              `--+--‘Nelima’ satoi Suzuki 1944 HT12, TM00
          |                 `--+--‘Nelima’ aokii HT12
          |                    |--‘Nelima’ genufusca (Karsch 1881) HT12, T03 (see below for synonymy)
          |                    |    |--N. g. genufusca S83
          |                    |    |--N. g. pumilio Suzuki 1950 S73
          |                    |    `--N. g. tuberculata Suzuki 1983 S83
          |                    |--‘Nelima’ nigricoxa Sato & Suzuki 1939 HT12, T03
          |                    |--‘Nelima’ suzukii Tsurusaki 2003 HT12, T03
          |                    `--+--‘Nelima’ parva HT12
          |                       `--‘Nelima’ similis HT12
          `--+--Togwoteeus Roewer 1952 HT12, C92
             |    `--T. biceps (Thorell 1877) C92 (see below for synonymy)
             `--+--+--‘Leiobunum’ veracruzensis HT12
                |  |--‘Krusa’ flava Goodnight & Goodnight 1946 HT12, GG46
                |  `--‘Krusa’ metallica Goodnight & Goodnight 1946 HT12, GG46
                `--+--+--+--‘Leiobunum’ patzquarum Goodnight & Goodnight 1942 HT12, GG46
                   |  |  `--‘Leiobunum’ townsendii Weed 1893 HT12, D34 [=Liobunum townsendii CZ04]
                   |  |--+--‘Leiobunum’ bogerti Goodnight & Goodnight 1942 HT12, GG45
                   |  |  |--‘Leiobunum’ colimae Goodnight & Goodnight 1945 HT12, GG45
                   |  |  `--+--‘Leiobunum’ potosum Goodnight & Goodnight 1942 HT12, GG44
                   |  |     `--+--‘Leiobunum’ hoogstraali Goodnight & Goodnight 1942 HT12, GG46
                   |  |        `--‘Leiobunum’ tascum Goodnight & Goodnight 1945 HT12, GG45
                   |  `--Paranelima Caporiacco 1938 HT12, C92 [incl. Glabrurus Goodnight & Goodnight 1942 C92]
                   |       |--*P. taibeli Caporiacco 1938 C92
                   |       |--P. albalineata Goodnight & Goodnight 1942 GG45
                   |       |--P. cerrana Goodnight & Goodnight 1942 GG42c
                   |       |--P. correa Goodnight & Goodnight 1945 GG45
                   |       |--P. lutzi (Goodnight & Goodnight 1942) GG45 [=*Glabrurus lutzi C92]
                   |       `--P. mexicana (Goodnight & Goodnight 1942) [=Leptobunus mexicanus] C84a
                   `--+--+--‘Leiobunum’ aldrichi Weed 1893 HT12, C84c (see below for synonymy)
                      |  `--+--+--+--‘Leiobunum’ bracchiolum (McGhee 1977) BHS12
                      |     |  |  `--‘Leiobunum’ politum (Wood 1868) HT12, BHS12 (see below for synonymy)
                      |     |  `--+--+--‘Leiobunum’ calcar (Wood 1868) HT12, BHS12 (see below for synonymy)
                      |     |     |  `--‘Leiobunum’ hoffmani Ingianni, McGhee & Shultz 2012 BHS13
                      |     |     `--+--‘Leiobunum’ euserratipalpe Ingianni et al. 2011 BHS13, BHS12
                      |     |        `--‘Leiobunum’ nigropalpi (Wood 1868) BHS13, BHS12 (see below for synonymy)
                      |     `--+--‘Leiobunum’ uxorium Crosby & Bishop 1924 HT12, D34
                      |        |--‘Leiobunum’ vittatum (Say 1821) HT12, D34 (see below for synonymy)
                      |        `--+--‘Leiobunum’ crassipalpe Banks 1900 BHS12 [=Liobunum crassipalpe E66]
                      |           `--‘Leiobunum’ relictum Davis 1934 BHS12
                      `--+--Leuronychus Banks 1900 HT12, C92 [incl. Microgyas Schenkel 1951 C92]
                         |    |--*L. pacificus (Banks 1894) (see below for synonymy) C92
                         |    `--L. fulviventris (Cambridge 1904) [=Liobunum fulviventre] R23
                         `--+--+--+--‘Leiobunum’ flavum Banks 1894 HT12, D34 [=Liobunum flavum E66]
                            |  |  `--‘Leiobunum’ verrucosum (Wood 1868) HT12, S08 (see below for synonymy)
                            |  `--+--‘Leiobunum’ holtae McGhee 1977 BHS12
                            |     `--‘Leiobunum’ ventricosum (Wood 1868) BHS12 (see below for synonymy)
                            |          |--L. v. ventricosum D34
                            |          |--L. v. floridanum Davis 1934 D34
                            |          `--L. v. hyemale (Weed 1890) CZ04 (see below for synonymy)
                            `--+--‘Eumesosoma’ ocalensis Cokendolpher 1980 HT12, E90
                               `--+--‘Leiobunum’ royali Goodnight & Goodnight 1946 BHS13, GG46
                                  `--+--+--‘Leiobunum’ bimaculatum Banks 1893 BHS13, BHS12 [=Liobunum bimaculatum R23]
                                     |  |--‘Leiobunum’ peninsulare Davis 1934 BHS12
                                     |  `--‘Leiobunum’ speciosum Banks 1900 BHS12, D34 [=Liobunum speciosum E66]
                                     `--+--Eumesosoma Cokendolpher 1980 BHS13, T-D03 (see below for synonymy)
                                        |    |--E. roeweri (Goodnight & Goodnight 1943) (see below for synonymy) C92
                                        |    |--E. arnetti Cokendolpher 1980 E90
                                        |    |--E. ephippiatum (Roewer 1923) E90 [=Mesosoma ephippiatum R23]
                                        |    |--E. nigrum (Say 1821) C92 (see below for synonymy)
                                        |    `--E. sayi Cokendolpher 1980 E90
                                        `--+--+--‘Leiobunum’ aurugineum Crosby & Bishop 1924 HT12, D34
                                           |  `--Forbesium Weed 1890 CZ04
                                           |       `--*F. formosum (Wood 1868) CZ04, S08 (see below for synonymy)
                                           `--Hadrobunus Banks 1900 BHS13, C92
                                                |--*H. grandis (Say 1821) C92 (see below for synonymy)
                                                |    |--H. g. grandis R23
                                                |    `--H. g. similis (Wood 1892) R23 (see below for synonymy)
                                                |--H. davisi Goodnight & Goodnight 1942 GG42b
                                                |--H. fusiformis BHS12
                                                |--H. knighti Goodnight & Goodnight 1942 GG42b
                                                `--H. maculosus (Wood 1868) GE02 (see below for synonymy)

Sclerosomatidae incertae sedis:
  Diguetinus Roewer 1912 C92
    `--*D. raptator Roewer 1912 C92
  Lanthanopilio Cokendolpher & Cokendolpher 1984 C92
    `--*L. chickeringi (Roewer 1956) [=Opilio chickeringi] C92
  ‘Mitopus’ mobilis Karsch 1881 (n. d.) R23
  ‘Phalangium’ ortoni Wood 1869 (n. d.) R23
  ‘Phalangium spiniferum’ Cantor 1842 non Lucas 1840 R23
  Munequita Mello-Leitão 1941 T-D04
    |--*M. pulchra Mello-Leitão 1941 C92 [incl. Holcobunus albianus Mello-Leitão 1944 T-D03]
    |--‘Holcobunus’ dentatus Roewer 1910 T-D04
    |--‘Prionostemma’ luteoscutum Roewer 1910 T-D04
    `--‘Prionostemma’ u-sigillatum Mello-Leitão 1938 T-D04
  Pectenobunus Roewer 1910 [incl. Acropiliops Mello-Leitão 1933, Caiza Roewer 1925] T-D04
    |--*P. paraguayensis (Canestrini 1888) [=Opilio paraguayensis] C92
    |--‘Caiza’ argentina Ringuelet 1959 T-D04
    |--*Caiza’ colliculosa Roewer 1925 T-D04
    `--P. ruricola (Mello-Leitão 1933) T-D04 [=*Acropiliops ruricola C92]
  Garleppa Roewer 1912 T-D04
    |--*G. granulata Roewer 1912 C92
    `--G. insperata Soares 1972 T-D04
  Bastioides Mello-Leitão 1931 (n. d.) T-D04
    `--*B. coxopunctatus Mello-Leitão 1931 (n. d.) [=B. coxopunctata (l.c.)] T-D03
  Jussara T07
  Tamboicus Roewer 1912 C92
    `--*T. fuhrmanni Roewer 1912 C92
  Amazonesia Soares 1970 C84b
    |--*A. quadriprocessigera Soares 1970 C92
    `--A. pulchra Soares 1970 KP-R02
  Azucarella Roewer 1959 C84b
    `--*A. weyrauchi Roewer 1959 C92
  Caluga Roewer 1959 C84b
    `--*C. rupicola Roewer 1959 C92
  Parageaya Mello-Leitão 1933 T-D04 [incl. Corderobunus Roewer 1938 C92]
    |--*P. ciliata Mello-Leitão 1933 C92
    |--P. albifrons Goodnight & Goodnight 1942 GG42c
    |--P. corderoi (Mello-Leitão 1936) T-D04 [=Hypsibunus corderoi C92, *Corderobunus corderoi C92]
    `--P. vittata Mello-Leitão 1939 T-D04 [=Corderobunus vittatus KP-R02]
  Fesa Roewer 1953 C92
    `--*F. tricolor Roewer 1953 C92
  Globulosoma Martens 1987 M87
    |--*G. gandakense Martens 1987 M87
    `--G. montivaga Martens 1987 K06
  Harmandina Schenkel 1954 C92
    `--*H. sinensis Schenkel 1954 C92
  Hexazaleptus Suzuki 1966 C92
    `--*H. junbesi Suzuki 1966 C92
  Himaldroma Martens 1987 M87
    |--*H. altus Martens 1987 M87, K06 [=H. alta C92]
    `--H. pineti Martens 1987 K06
  Himalzaleptus Martens 1987 M87
    `--*H. quinqueconicus Martens 1987 M87
  Holmbergiana Mello-Leitão 1931 T-D04 (see below for synonymy)
    |--*H. weyenberghii (Holmberg 1876) (see below for synonymy) C92
    |--*Guaranobunus’ guaraniticus Ringuelet 1959 T-D04
    |--H. orientalis Ringuelet 1963 T-D04
    `--H. tibialis Ringuelet 1959 T-D04
  Nepalkanchia Martens 1990 [=Kanchia Martens 1987 non Moore 1883] C92
    |--*N. pluviosilvestris (Martens 1987) [=*Kanchia pluviosilvestris] C92
    `--‘Kanchia’ silvicola Martens 1987 M87
  Koyamaia Suzuki 1972 C92
    `--*K. curvipes Suzuki 1972 C92
  Liopagus Chamberlin 1916 C92
    `--*L. simplex Chamberlin 1916 C92
  Nepalgrella Martens 1987 M87
    |--*N. kortaliensis Martens 1987 M87
    `--N. yamputhini Martens 1987 M87
  Octozaleptus Suzuki 1966 C92
    `--*O. harai Suzuki 1966 C92
  Onostemma Mello-Leitão 1938 C92
    `--*O. imitans Mello-Leitão 1938 C92
  Oobunus Kishida 1930 C92
    `--*O. schizops Kishida 1930 C92
  Paragagrellina Schenkel 1963 C92
    `--*P. legendrei Schenkel 1963 C92
  Paratamboicus Mello-Leitão 1940 C92
    `--*P. bicornutus Mello-Leitão 1940 C92
  Paraumbogrella Suzuki 1963 C92
    `--P. pumilio (Karsch 1881) T03 (see below for synonymy)
  Paruleptes Soares 1970 C84b
    `--*P. coronatus Soares 1970 C92
  Pokhara Suzuki 1970 M87
    |--*P. lineata Suzuki 1970 M87
    |--P. kathmandica Martens 1987 M87
    |--P. minuta Martens 1987 M87
    |--P. occidentalis Martens 1987 M87
    |--P. quadriconica Martens 1987 M87
    |--P. trisulensis Martens 1987 M87
    |--P. uenoi Martens 1987 M87
    `--P. yodai (Suzuki 1966) [=Hypsibunus yodai; incl. H. gibber Suzuki 1966] M87
  Psammogeaya Mello-Leitão 1946 T-D04
    `--*P. arenata Mello-Leitão 1946 C92
  Pseudarthromerus Karsch 1892 C92
    `--*P. spurius Karsch 1892 C92
  Romerella Goodnight & Goodnight 1943 C92
    |--*R. punctata Goodnight & Goodnight 1943 C92
    |--R. bicolor Goodnight & Goodnight 1944 GG53
    |--R. brasiliensis Goodnight & Goodnight 1944 GG44a
    `--R. catharina Goodnight & Goodnight 1944 GG44b
  Sericicorpus Martens 1987 M87
    `--*S. nigrum Martens 1987 M87
  Taperina Roewer 1953 C92
    `--*T. nigripes Roewer 1953 C92
  Varinodulia Canals 1935 TK01
    `--*V. insularis Canals 1935 C92
  Xerogrella Martens 1987 M87
    `--*X. dolpensis Martens 1987 M87
  Abaetetuba Tourinho-Davis 2004 T-D04
    |--*A. plaumanni (Roewer 1953) [=Prionostemma plaumanni] T-D04
    |--A. bahiensis (Mello-Leitão 1931) [=Geaya bahiensis] T-D04
    |--A. citrina (Pocock 1903) [=Prionostemma citrinum, Holcobunus citrinus] T-D04
    `--A. lisei Tourinho-Davis 2004 T-D04
  Zaleptus M87
  Hypsibunus R55b
  Ceratobunus R55b
  Echinobunus Roewer 1912 C92
    `--*E. elegans Roewer 1912 C92
  Diangathia Roewer 1955 M87
    `--*D. bovifrons Roewer 1955 M87
  Euceratobunus Roewer 1923 C92
    `--*E. pulcher (With 1903) [=Ceratobunus pulcher] C92
  Orissula Roewer 1955 C92
    `--*O. heterospinulata Roewer 1955 C92
  Obigrella Roewer 1955 C92
    `--*O. nigra Roewer 1955 C92
  Bakerinulus Roewer 1955 C92
    `--*B. luzonicus Roewer 1955 C92
  Biceropsis Roewer 1935 C92
    `--*B. maculata Roewer 1935 C92
  Sinadroma Roewer 1955 C92
    `--*S. sherriffsi Roewer 1955 C92
  Carmichaelus Roewer 1929 C92
    `--*C. maculatus Roewer 1929 C92
  Carinobius Roewer 1955 C92
    `--*C. bicornutus Roewer 1955 C92
  Microzaleptus Roewer 1955 C92
    |--*M. quadratus Roewer 1955 C92
    `--M. montanus (Banks 1930) [=Metazaleptus montanus] R55b
  Metaverpulus Roewer 1912 M87
    |--*M. hirsutus Roewer 1912 (see below for synonymy) M87
    |--M. albipunctatus (Roewer 1915) [=Harmandella albipunctata, Harmanda albipunctata] M87
    |--M. bhutanicus Martens 1987 M87
    |--M. kanchensis Martens 1987 K06
    |--M. laevis Roewer 1955 M87
    |--M. multidentatus Martens 1987 M87
    `--M. persimilis Martens 1987 M87
  Pentazaleptus Roewer 1955 C92
    |--*P. adspersus Roewer 1955 C92
    `--P. guttatus Roewer 1955 R55b
  Adungrella Roewer 1955 C92
    `--*A. aenea Roewer 1955 C92
  Bastia Roewer 1910 C92
    |--*B. lineata Roewer 1910 C92
    `--B. guttata Banks 1930 R55b
  Zaleptiolus Roewer 1955 M87
    |--*Z. laevipes Roewer 1955 M87
    |--Z. ater Martens 1987 M87
    `--Z. implicatus Suzuki 1970 M87
  Ceratobunoides Roewer 1923 C92
    |--*C. sumatranus Roewer 1923 C92
    |--C. bandarensis Roewer 1955 R55b
    `--C. mentaweiensis Roewer 1955 R55b
  Ceratobunellus Roewer 1911 C92
    |--*C. calcuttensis (With 1903) [=Ceratobunus calcuttensis] C92
    |--C. brevipes (With 1903) R55b [=Ceratobunus brevipes R23]
    `--C. philippinus Roewer 1955 R55b
  Tetraceratobunus Roewer 1915 C92
    |--*T. lineatus Roewer 1915 C92
    |--T. lithobius Roewer 1955 R55b
    `--T. marmoratus Roewer 1955 R55b
  Zaleptanus Roewer 1929 C92
    |--*Z. gravelyi Roewer 1929 C92
    `--Z. promeus Roewer 1955 R55b
  Cervibunus Roewer 1912 C92
    |--*C. maculatus Roewer 1912 C92
    |--C. ater Roewer 1955 R55b
    `--C. ornatus Roewer 1929 R55b
  Metazaleptus Roewer 1912 M87
    |--*M. hirsutus (With 1903) [=Zaleptus hirsutus] M87
    |--M. borneensis Banks 1930 R55b
    |--M. rufescens Banks 1930 R55b
    `--M. striatus Suzuki 1985 S85
  Euzaleptus Roewer 1911 M87
    |--*E. minutus (With 1903) (see below for synonymy) M87
    |--E. muticus Roewer 1929 M87
    `--E. sarawakensis Roewer 1911 M87
  Bullobunus Roewer 1910 C92
    |--*B. luteovittatus Roewer 1910 C92
    |--B. ater Roewer 1910 R55b
    |--B. bakeri Roewer 1926 R55b
    |--B. rufus Roewer 1955 R55b
    `--B. similis Roewer 1910 R55b
  Zaleptulus Roewer 1955 C92
    |--*Z. unicolor Roewer 1955 C92
    |--Z. annulipes (Banks 1930) [=Zaleptus annulipes] R55b
    |--Z. banksi Roewer 1955 R55b
    |--Z. lineatus Roewer 1955 R55b
    `--Z. pulchellus (Banks 1930) [=Zaleptus pulchellus] R55b
  Verpulus Simon 1901 C92
    |--*V. spumatus Simon 1901 C92
    |--V. brunneus Roewer 1955 R55b
    |--V. fulvus Suzuki 1985 M87
    |--V. gracilis Roewer 1955 R55b
    |--V. laevipes Roewer 1955 R55b
    |--V. magnus Roewer 1955 R55b
    |--V. marginatus Roewer 1912 R55b
    `--V. peguensis Roewer 1955 R55b
  Harmanda M87
  Gagrella M87
  Dentobunus R55a
  Eugagrella R54b
  Strandia R54b
  Aurivilliola R55a
  Metahehoa Suzuki 1985 C92
    `--*M. granulata Suzuki 1985 S85
  Pseudosystenocentrus Suzuki 1985 C92
    `--*P. foveolatus Suzuki 1985 S85
  Antigrella Roewer 1954 C92
    `--*A. orissana Roewer 1954 C92
  Palniella Roewer 1929 C92
    `--*P. virididorsata Roewer 1929 C92
  Verrucobunus Roewer 1931 C92
    `--*V. trispinosus Roewer 1931 C92
  Scotomenia Thorell 1889 C92
    `--*S. cetrata Thorell 1889 C92
  Metadentobunus Roewer 1915 C92
    `--*M. formosae Roewer 1915 C92
         |--M. f. formosae S77
         `--M. f. garampiensis Suzuki 1944 S77
  Hamitergum Crawford 1992 [=Lophacanthus Redikorzev 1936 non Stock 1880] C92
    `--*H. eobium (Redikorzev 1936) C92 (see below for synonymy)
  Hehoa Roewer 1929 C92
    `--*H. bunigera Roewer 1929 C92
  Baturitia Roewer 1931 C92
    `--*B. trispinosa Roewer 1931 C92
  Umbogrella Roewer 1955 C92
    `--*U. minuta Roewer 1954 C92
  Chebabius Roewer 1935 K06
    `--*C. angulatus Roewer 1935 C92
  Paradentobunus Roewer 1915 C92
    `--*P. aureomaculatus Roewer 1915 C92
  Padangrella Roewer 1954 C92
    `--*P. jacobsoni Roewer 1954 C92
  Pergagrella Roewer 1954 C92
    `--*P. monticola Roewer 1954 C92
  Gagrellenna Roewer 1929 C92
    `--*G. bipunctata Roewer 1929 C92
  Coonoora Roewer 1929 C92
    `--*C. biceratops Roewer 1929 C92
  Hypogrella Roewer 1955 C92
    `--*H. poecilis Roewer 1955 C92
  Sarasinia Roewer 1913 C92
    `--*S. punctata Roewer 1913 C92
  Gagrellina Roewer 1913 C92
    `--*G. vestita Roewer 1913 C92
  Syngagrella Roewer 1913 C92
    `--*S. bistriata Roewer 1913 C92
  Heterogagrella Roewer 1954 C92
    |--*H. indica Roewer 1954 C92
    `--H. biseriata Suzuki 1981 S85
  Gagrellissa Roewer 1931 C92
    |--*G. jacobsoni Roewer 1931 C92
    `--G. luteobiseriata Suzuki 1985 S85
  Akalpia Roewer 1915 C92
    |--*A. oblonga Roewer 1915 C92
    `--A. nipponica Sato & Suzuki 1938 R55a
  Melanopella Roewer 1931 C92
    |--*M. feuerborni Roewer 1931 C92
    `--M. marginata Roewer 1955 R55a
  Chasenella Roewer 1955 C92
    |--*C. luma Roewer 1933 C92
    `--C. pakka Roewer 1933 R55a
  Altobunus Roewer 1910 C92
    |--*A. formosus Roewer 1910 C92
    |--A. inermis (Simon 1877) R54a [=Gagrella inermis R23]
    `--A. maculatus Roewer 1910 R54a
  Prodentobunus Roewer 1923 C92
    |--*P. unispinosus (Roewer 1912) [=Dentobunus unispinosus] C92
    |--P. nitidus Roewer 1955 R55a
    `--P. tao Roewer 1955 R55a
  Metasyleus Roewer 1955 C92
    |--*M. ephippiatus Roewer 1955 C92
    |--M. orissus Roewer 1955 R55a
    `--M. tenuis Roewer 1955 R55a
  Sataria Roewer 1915 C92
    |--*S. maculata Roewer 1915 C92
    |--S. coronata Roewer 1929 R55a
    `--S. unicolor Roewer 1914 R55a
  Syleus Thorell 1876 [=Acanthonotus Koch 1839 non Goldfuss 1809] C92
    |--*S. niger (Koch 1839) [=*Acanthonotus niger] C92
    |--S. mysoreus Roewer 1955 R55a
    `--S. rufus Roewer 1955 R55a
  Neogagrella Roewer 1913 C92
    |--*N. eximia Roewer 1913 C92
    |--N. balica Roewer 1931 R54b
    `--N. barnesi Roewer 1929 R54b
  Hologagrella Roewer 1910 C92
    |--*H. reticulata Roewer 1910 C92
    |--H. curvicornis Roewer 1913 R55a
    |--H. curvispina Banks 1930 B30
    |--H. luzonica Roewer 1910 R55a
    `--H. timorana Roewer 1955 R55a
  Bonthainia Roewer 1913 [incl. Apogrella Roewer 1955, Narmadella Roewer 1931, Nilgirisia Roewer 1929] C92
    |--B. aenescens Roewer 1913 C92
    |--*Apogrella’ celebensis Roewer 1955 C92
    |--*Narmadella’ elegans Roewer 1931 C92
    `--*Nilgirisia’ gravelyi Roewer 1929 C92
  Melanopula Roewer 1929 C92
    |--*M. biceps Roewer 1929 C92
    |--M. crassitarsis Roewer 1955 R55a
    |--M. insularis Roewer 1931 R55a
    |--M. shanensis Roewer 1955 R55a
    `--M. thaiensis Suzuki 1985 S85
  Hexomma Thorell 1876 [incl. Arthrocentrus Thorell 1889, Gagrella Roewer 1910 non Stoliczka 1869] C92
    |--*H. vulcanicum (Doleschall 1859) C92 [=Phalangium vulcanicum C92, Gagrella vulcanica R54a]
    |--‘Gagrella’ arthrocentra Roewer 1910 (see below for synonymy) C92
    `--*Gagrella’ feae Thorell 1889 C92
         |--G. f. feae R54a
         |--G. f. bispinosa Thorell 1889 R54a
         |--G. f. conjugata Roewer 1954 R54a
         `--G. f. humeralis Thorell 1889 R54a
  Paragagrella Roewer 1912 C92
    |--*P. typus Roewer 1912 C92
    |--P. bakeri Roewer 1954 R54b
    |--P. basalis Roewer 1929 R54b
    |--P. bicolor Roewer 1954 R54b
    |--P. brevispina Banks 1930 B30
    |--P. mysorea Roewer 1939 R54b
    |--P. palpalis Roewer 1954 R54b
    |--P. roeweri Giltay 1930 R54b
    `--P. siberutiana Roewer 1931 R54b
  Umbopilio Roewer 1956 K06
    |--*U. paradoxus Roewer 1956 K06
    `--U. martensi Klimeš 2006 K06
  Eusclera Roewer 1910 [=Eusclerella Roewer 1910] C92
    `--*E. aureomaculata Roewer 1910 [=*Eusclerella aureomaculata] C92
  Microliobunum Roewer 1912 C92
    `--*M. brevipes Roewer 1912 C92
  Dilophiocara Redikorzev 1931 C92
    `--*D. bactrianum Redikorzev 1931 [=D. bactriana] C92
  Micronelima Schenkel 1938 C92
    `--*M. brevipes Schenkel 1938 C92
  Schenkeliobunum Staręga 1964 [=Thrasychiroides Schenkel 1963 non Soares 1947] C92
    `--*S. tuberculatum (Schenkel 1963) [=*Thrasychiroides tuberculatus] C92
  Pseudomelanopa Suzuki 1974 C92
    |--*P. taiwana Suzuki 1974 C92
    `--P. liupan Zhang & Zhu 2009 ZZ09
  Amauropilio Mello-Leitão 1937 C92
    |--*A. lacoei (Petrunkevitch 1922) C92 [=Phalangium lacoei C92, A. lawei D04, P. lawei (l. c.) C92]
    `--A. atavus (Cockerell 1907) [=Leptobunus atavus] D07
  Cosmobunus Simon 1879 C92
    |--*C. granarius (Lucas 1847) C92, E12 (see below for synonymy)
    |--C. auratus Goodnight & Goodnight 1946 GG46
    |--C. unicolor Roewer 1910 R23
    `--C. unifasciatus Roewer 1923 R23

Nomen nudum: Dicranopalpus nipponensis Kishida 1934 TM00
             Systenocentrus uyemurai Kishida 1934 TM00

Astrobunus Thorell 1876 HT12, C92 [=Hoplites Koch 1869 (nom. rej.) non Neumayr 1875 C92; incl. Roeweriolus Kolosváry 1933 C92]

Astrobunus helleri (Ausserer 1867) D04 [=Acantholophus helleri D04, Hoplites helleri R23; incl. *H. argentatus Koch 1869 C92, *Astrobunus argentatus C92, A. croaticus Sörensen 1894 D04, Hoplites pavesii Canestrini 1872 D04, A. pavesii D04, A. slovenicus Hadži 1927 D04]

Astrobunus laevipes (Canestrini 1872) B95 [=Hoplites laevipes B95; incl. Astrobunus bosnicus Roewer 1912 M78, Roeweriolus carpathicus Bartos 1938 D04, R. dudichi Szalay 1951 B95, Astrobunus dudichi B95, *R. hungaricus Kolosváry 1933 C92, Sclerosoma meadi Thorell 1876 D04, Astrobunus meadi B95, A. bernardicus simoni Hadži 1927 D04, Roeweriolus slavicus Kratochvil 1934 D04]

Astrobunus spinosus (Herbst 1799) (n. d.) [=Opilio spinosus, Phalangium spinosum nec Bosc 1792 nec Latreille 1798] R23

*Cosmobunus granarius (Lucas 1847) C92, E12 [=Phalangium granarium D04; incl. P. flavounilineatum Lucas 1847 D04, P. levipes Lucas 1847 D04]

Eumesosoma Cokendolpher 1980 BHS13, T-D03 [=Mesosoma Weed 1892 non Otto 1821 C92; Mesosomatinae]

Eumesosoma nigrum (Say 1821) C92 [=Phalangium nigrum C92, Astrobunus niger R23, Mesosoma nigrum CZ04]

Eumesosoma roeweri (Goodnight & Goodnight 1943) [=Mesosoma roeweri; incl. *M. nigrum Weed 1892 non Phalangium nigrum Say 1821, M. texanum Goodnight & Goodnight 1943] C92

*Euzaleptus minutus (With 1903) [=Zaleptus minutus; incl. Metaverpulus distinctus Suzuki 1970, Eugagrella minima Roewer 1954, Zaleptanus monticola Roewer 1955, Metagagrella tenuis Roewer 1954] M87

*Forbesium formosum (Wood 1868) CZ04, S08 [=Phalangium formosum D34, Leiobunum formosum HT12, Liobunum formosum E66]

‘Gagrella’ arthrocentra Roewer 1910 [=*Arthrocentrus atratus Thorell 1889 non G. atrata Stoliczka 1869] C92

Hadrobunus grandis (Say 1821) C92 [=Phalangium grandis BH09, P. grande C92, Astrobunus grandis R23, Leptobunus grandis R23, Liobunum grande R23]

Hadrobunus grandis similis (Wood 1892) R23 [=Liobunum grande var. simile CZ04, L. similis Wood 1890 (n. n.) CZ04]

Hadrobunus maculosus (Wood 1868) GE02 [=Phalangium maculosum R23, Leptobunus maculosus R23, Liobunum maculosum R23]

*Hamitergum eobium (Redikorzev 1936) C92 [=*Lophacanthus eobius C92; incl. L. borchseniusi Charitonov 1957 S65]

Holmbergiana Mello-Leitão 1931 T-D04 [incl. Guaranobunus Ringuelet 1959 T-D04, Simplicibunus Ringuelet 1954 C84b]

*Holmbergiana weyenberghii (Holmberg 1876) [=Opilio weyenberghii, O. weyemberghi (l. c.), O. weyenberghi (l. c.), O. weyenbergii (l. c.); incl. *Simplicibunus delicatus Ringuelet 1953] C92

‘Leiobunum’ aldrichi Weed 1893 HT12, C84c [=Liobunum longipes aldrichi C84c; incl. Le. gordoni Goodnight & Goodnight 1945 CZ04, Li. longipes Weed 1890 non Le. longipes Menge in Koch & Berendt 1854 C84c]

‘Leiobunum’ blackwalli Meade 1861 HT12, D04 [=Leiobunus blackwallii D04, Liobunum blackwalli D04, Liobunum blackwilli (l. c.) D04, Liobunus blackwallii D04; incl. Leiobunum granadinum Marcellino 1967 M78, Leiobunum hemisphaericum Koch 1872 D04]

‘Leiobunum’ calcar (Wood 1868) HT12, BHS12 [=Phalangium calcar D34, Liobunum calcar E66; incl. Li. brunnea Walker 1928 D34, Leiobunum cumberlandense McGhee in Ingianni et al. 2011 (n. n.) BHS12]

‘Leiobunum’ exilipes (Wood 1870) HT12, D34 [=Phalangium exilipes D34, Liobunum exilipes R23; incl. Leuronychus gertschi C84a]

‘Leiobunum’ hiraiwai (Sato & Suzuki 1939) HT12, T06 [=Nelima hiraiwai T06; incl. L. hiraiwai fuji Suzuki 1976 T06, L. hiraiwai izuense Suzuki 1976 T06, L. hiraiwai longum Suzuki 1976 T06, L. hiraiwai shiranense Suzuki 1976 T06, L. tamanum Suzuki 1957 T06, L. tsushimense Suzuki 1976 T06]

‘Leiobunum’ japanense suzukii Roewer 1957 [incl. L. bifrons Roewer 1957, *Metaliobunum japonicum Suzuki 1940 non Leiobunum japonicum Müller 1914, L. japanense japonicum] C92

‘Leiobunum’ japonicum Müller 1914 HT12, T03 [=Liobunum japonicum TT05; incl. Le. wegneri Šilhavý 1976 TT05]

‘Leiobunum’ manubriatum Karsch 1881 HT12, T06 [=Liobunum manubriatum R10, Zaleptus manubriatus R10]

‘Leiobunum’ montanum Suzuki 1953 HT12, T06 [=Liobunum montanum T85a; incl. L. virgeum ontakense Suzuki 1976 T85a, L. montanum sobosanum Suzuki 1976 T85a, L. virgeum Suzuki 1976 T85a]

‘Leiobunum’ nigropalpi (Wood 1868) BHS13, BHS12 [=Phalangium nigropalpi D34, Liobunum nigropalpi E66]

‘Leiobunum’ politum (Wood 1868) HT12, BHS12 [=L. politus D34; incl. L. cavernarum Roewer 1952 CZ04, L. politum magnum Weed 1893 D34]

‘Leiobunum’ ventricosum (Wood 1868) BHS12 [=Phalangium ventricosum D34, Liobunum ventricosum E66]

‘Leiobunum’ ventricosum hyemale (Weed 1890) CZ04 [=Forbesium hyemale CZ04, Liobunum hyemale R10, Leiobunum ventricosum hiemale R23]

‘Leiobunum’ verrucosum (Wood 1868) HT12, S08 [=Phalangium verrucosum S08, Liobunum verrucosum S08; incl. Li. nigripes Weed 1892 S08, Leiobunum nigripes S08]

‘Leiobunum’ vittatum (Say 1821) HT12, D34 [=Phalangium vittatum D34, Liobunum vittatum E66; incl. Phalangium dorsatum Say 1821 D34, Leiobunum vittatum dorsatum D34, Liobunum vittatum minor Weed 1893 CZ04]

*Leuronychus pacificus (Banks 1894) [=Liobunum pacificum; incl. *Microgyas banksi Schenkel 1951] C92

*Mastobunus tuberculifer (Lucas 1847) C92, E12 [=Phalangium tuberculiferum D04, Sclerosoma tuberculifera D04; incl. Homalenotus sicanus Pavesi 1876 D04, S. sicanum D04]

*Metasclerosoma sardum (Thorell 1876) [=Sclerosoma sardum, *Neosclerosoma sardum; incl. *Prosclerosoma insignum Roewer 1915] C92

*Metaverpulus hirsutus Roewer 1912 [incl. Euzaleptus pilosus Roewer 1929, Adungrella punctulata Roewer 1955] M87

‘Nelima’ elegans (Weed 1889) HT12, C84c [=Liobunum elegans CZ04, Leiobunum elegans D34; incl. Li. annulatum Walker 1928 D34, Phalangium bicolor Wood 1868 non Fabricius 1793 C84c, Astrobunus bicolor D34, Leiobunum bicolor non Koch 1848 D34, Liobunum bicolor C84c]

‘Nelima’ genufusca (Karsch 1881) HT12, T03 [=Mitopus genufuscus R23; incl. N. gigantea (Loman 1902) T03]

Paraumbogrella pumilio (Karsch 1881) T03 [=Gagrella pumilio C92, Melanopa pumilio R23; incl. *Paraumbogrella huzitai Suzuki 1963 C92]

Sclerosomatidae [Gagrellinae, Gagrelleae, Gagrellidae, Gagrellinas, Gagrellini, Leiobunidae, Leiobuninae, Liobuninae, Liobunini, Sclerosomidae, Zalepteae, Zaleptinae, Zaleptini] VFG14

Togwoteeus biceps (Thorell 1877) C92 [=Mitopus biceps BH09, Homolophus biceps BH09; incl. *Togwoteeus granipalpus Roewer 1952 C92]

*Type species of generic name indicated

REFERENCES

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[B95] Bokor, Z. 1995. New ocurrence of Astrobunus laevipes (Canestrini, 1872) (Arachnoidea, Opiliones, Phalangiidae) in Hungary (in the valley of river Rakaca-Cserehát). Acta Universitatis Szegediensis Acta Biologica 41: 89–91.

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

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[C84a] Cokendolpher, J. C. 1984a. Revision of the harvestman genus Leptobunus and dismantlement of the Leptobunidae (Arachnida: Opiliones: Palpatores). Journal of the New York Entomological Society 92 (4): 371–402.

[C84b] Cokendolpher, J. C. 1984b. Clarification of the Colombian harvestman genus Carmenia, with a review of the New World Gagrellinae (Opiliones: Gagrellidae). Florida Entomologist 67 (3): 471–478.

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[GG43] Goodnight, C. J., & M. L. Goodnight. 1943. New and little known phalangids from the United States. American Midland Naturalist 29 (3): 643–656.

[GG44a] Goodnight, C. J., & M. L. Goodnight. 1944a. Duas espécies novas de opiliões Sul-Americanos. Revista de Entomologia 15 (3): 332–334.

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