Belongs within: Phalangioidea.
Contains: Jussara, Zaleptus, Hypsibunus, Ceratobunus, Verpulus, 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). Instead, molecular phylogenies are more consistent with distinct European, South American, Asian and North American clades (Hedin et al. 2012).
<==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 [incl. Parodiellus projectus Goodnight & Goodnight 1942] CH18
| |--P. rossi Goodnight & Goodnight 1943 GG43
| `--P. singularis Banks 1893 GG42a
`--+--+--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
| | | i. s.: K. amazonica Roewer 1953 KP-R02
| | | K. flava Goodnight & Goodnight 1946 GG46
| | | K. hidalguensis Gaona-Escamilla, Francke & Ponce-Saavedra 2016 G-EFP-S16
| | | K. metallica Goodnight & Goodnight 1946 GG46
| | | 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
| | `--G. dissimile Martens 2018 M18
| | 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, B95 (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, C92 (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’ curvipalpe group T91
| | |--+--‘Leiobunum’ hikocola subgroup T91
| | | | |--‘Leiobunum’ hikocola Suzuki 1966 HT12, T06
| | | | `--‘Leiobunum’ oharai Tsurusaki 1991 T91
| | | `--+--‘Leiobunum’ platypenis subgroup T85b
| | | | |--‘Leiobunum’ globosum Suzuki 1953 HT12, T03
| | | | `--‘Leiobunum’ platypenis Suzuki 1953 (see below for synonymy) T85b
| | | `--‘Leiobunum’ manubriatum Karsch 1881 HT12, T06 (see below for synonymy)
| | `--+--‘Leiobunum’ hiraiwai (Sato & Suzuki 1939) HT12, T06 (see below for synonymy)
| | |--‘Leiobunum’ curvipalpe subgroup T90
| | | |--‘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 subgroup T85b
| | | |--‘Leiobunum’ hiasai Suzuki 1976 HT12, T85b
| | | |--‘Leiobunum’ sadoense Tsurusaki 1982 T85b
| | | `--‘Leiobunum’ simplum Suzuki 1976 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 BHS13, C92
| | `--T. biceps (Thorell 1877) BHS12 (see below for synonymy)
| `--+--+--‘Leiobunum’ patzquarum Goodnight & Goodnight 1942 HT12, GG46
| | `--‘Leiobunum’ townsendii Weed 1893 BHS12 [=Liobunum townsendii CZ04]
| `--+--+--‘Leiobunum’ bogerti Goodnight & Goodnight 1942 HT12, GG45
| | |--‘Leiobunum’ colimae Goodnight & Goodnight 1945 HT12, GG45
| | `--+--‘Leiobunum’ potosum Goodnight & Goodnight 1942 BHS12
| | `--+--‘Leiobunum’ hoogstraali Goodnight & Goodnight 1942 HT12, GG46
| | `--‘Leiobunum’ tascum Goodnight & Goodnight 1945 HT12, GG45
| `--Leuronychus Banks 1900 BHS13, C92 [incl. Microgyas Schenkel 1951 C92]
| |--*L. pacificus (Banks 1894) (see below for synonymy) C92
| `--L. fulviventris (Cambridge 1904) [=Liobunum fulviventre] R23
`--+--+--‘Leiobunum’ aldrichi Weed 1893 BHS12 (see below for synonymy)
| `--+--+--+--‘Leiobunum’ bracchiolum (McGhee 1977) BHS12
| | | `--‘Leiobunum’ politum (Wood 1868) BHS12 (see below for synonymy)
| | `--‘Leiobunum’ calcar group IMS11
| | |--+--‘Leiobunum’ calcar (Wood 1868) BHS13, BHS12 (see below for synonymy)
| | | `--‘Leiobunum’ hoffmani Ingianni, McGhee & Shultz 2012 BHS13
| | `--+--‘Leiobunum’ euserratipalpe Ingianni, McGhee & Schultz 2011 BHS13, BHS12
| | `--‘Leiobunum’ nigropalpi (Wood 1868) BHS13, BHS12 (see below for synonymy)
| `--+--‘Leiobunum’ uxorium Crosby & Bishop 1924 BHS12
| |--‘Leiobunum’ vittatum (Say 1821) BHS13 (see below for synonymy)
| `--+--‘Leiobunum’ crassipalpe Banks 1900 BHS12 [=Liobunum crassipalpe E66]
| `--‘Leiobunum’ relictum Davis 1934 BHS12
`--+--+--+--‘Leiobunum’ flavum Banks 1894 BHS12 [=Liobunum flavum E66]
| | `--‘Leiobunum’ verrucosum (Wood 1870) BHS13, IMS11 (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. hiemale (Weed 1890) CH18 (see below for synonymy)
`--+--‘Eumesosoma’ ocalensis Cokendolpher 1980 HT12, E90
`--+--‘Leiobunum’ royali Goodnight & Goodnight 1946 BHS12
`--+--+--‘Leiobunum’ bimaculatum Banks 1893 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) BHS12 (see below for synonymy)
| |--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
`--+--‘Hadrobunus’ maculosus (Wood 1868) BHS13 (see below for synonymy)
`--+--+--‘Leiobunum’ aurugineum Crosby & Bishop 1924 BHS12
| `--Forbesium Weed 1890 CZ04
| `--*F. formosum (Wood 1868) CZ04 (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
Sclerosomatidae incertae sedis:
‘Mitopus’ mobilis Karsch 1881 (n. d.) R23
‘Phalangium’ ortoni Wood 1869 (n. d.) R23
‘Phalangium’ rudipalpe Gervais 1849 (n. d.) CH18
‘Phalangium spiniferum’ Cantor 1842 non Lucas 1840 R23
Diguetinus Roewer 1912 C92
`--*D. raptator Roewer 1912 C92
Lanthanopilio Cokendolpher & Cokendolpher 1984 C92
`--*L. chickeringi (Roewer 1956) [=Opilio chickeringi] C92
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 C92
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. brevispinus Chen & Shih 2018 CS18
`--M. garampiensis Suzuki 1944 [=M. formosae garampiensis] CS18
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 ZZ09
|--*P. taiwana Suzuki 1974 ZZ09
`--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. sagani Palencia, Peñalver et al. 2019 PP19
`--C. unicolor Roewer 1910 [incl. C. unifasciatus Roewer 1923] PP19
Krusella Roewer 1953 G-EFP-S16
`--*K. pilipes Roewer 1953 C92
Goasheer Snegovaya, Cokendolpher & Mozaffarian 2018 SCM18
`--*G. iranus (Roewer 1952) [=Homolophus iranus, Microliobunum iranum] SCM18
Mesobunus Huang, Selden & Dunlop 2009 PP19, GT12
|--*M. martensi Huang, Selden & Dunlop 2009 HSD09
`--M. dunlopi Giribet, Tourinho et al. 2012 GT12
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
Nomen nudum: Systenocentrus uyemurai Kishida 1934 TM00
Astrobunus Thorell 1876 HT12, B95 [=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) BHS12 [=Mesosoma roeweri C92; incl. *M. nigrum Weed 1892 non Phalangium nigrum Say 1821 C92, 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 [=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 grande CH18, 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) BHS13 [=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 BHS12 [=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) BHS13, BHS12 [=Phalangium calcar D34, Liobunum calcar E66; incl. Li. brunnea Walker 1928 D34, Leiobunum cumberlandense McGhee in Ingianni, McGhee & Shultz 2011 (n. n.) BHS12, Le. serratipalpe Roewer 1910 IMS11, Liobunum serratipalpe E66]
‘Leiobunum’ exilipes (Wood 1870) HT12, D34 [=Phalangium exilipes D34, Lei. exillipes (l. c.) CH18, 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 nigripalpi IMS11, Li. nigripalpis non Leiobunum nigripalpe Simon 1889 IMS11]
‘Leiobunum’ platypenis Suzuki 1953 [=Liobunum platypenis; incl. Leiobunum esoense Suzuki 1976] T85b
‘Leiobunum’ politum (Wood 1868) 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 hiemale (Weed 1890) CH18 [=Phalangium ventricosum hiemale CH18, Forbesium hyemale CZ04, Leiobunum ventricosum hyemale CZ04, Liobunum hyemale R10]
‘Leiobunum’ verrucosum (Wood 1870) BHS13, IMS11 [=Phalangium verrucosum S08, Liobunum verrucosum S08; incl. Li. nigripes Weed 1892 S08, Leiobunum nigripes S08]
‘Leiobunum’ vittatum (Say 1821) BHS13 [=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, C92 [=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. Liobunum giganteum Loman 1902 ST83, T03, Leiobunum giganteum ST83, Nelima gigantea T03, N. tenuis Roewer 1957 ST83]
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) BHS12 [=Mitopus biceps BH09, Homolophus biceps BH09; incl. *Togwoteeus granipalpus Roewer 1952 C92]
*Type species of generic name indicated
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