Male Leiobunum vittatum, from here.

Belongs within: Palpatores.
Contains: Jussara, Zaleptus, Hypsibunus, Ceratobunus, Verpulus, Harmanda, Gagrella, Dentobunus, Eugagrella, Strandia, Aurivilliola, Leiobunum, Holcobunus, Geaya, Prionostemma, HomalenotusAstrobunus, 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).

The Gagrella problem cranked up to eleven
Published 6 January 2012
A Leiobunum eating a cricket, photographed in North Carolina by Jeffrey Pippen.

In previous posts on this site, I have referred to the problem of Gagrella: a large genus of Asian harvestmen diagnosed on the basis of characters long since recognised as unreliable that remains unrevised because no-one has yet been in a position to take on the amount of work required. Well, a paper has come out in the last few weeks that indicates that the Gagrella issue is just part of a larger problem, one of truly demonic proportions.

The British Nelima gothica, photographed by Gordon.

Gagrella belongs to a family of harvestmen called the Sclerosomatidae. Sclerosomatids are distinguished by having a more heavily sclerotised dorsum than most other long-legged harvestmen, and by the morphology of the male genitalia, with many (but not all) species having lateral extensions (like wings) on the shaft just behind the glans. Within the Sclerosomatidae, most authors have recognised four subfamilies: Sclerosomatinae, Gagrellinae, Leiobuninae and Gyinae. The boundaries between the subfamilies have long been realised to be a bit fuzzy, particularly between the Leiobuninae and Gagrellinae. A few molecular studies (for instance, Giribet et al. 2010) that have included more than one representative of more than one subfamily have failed to resolve them as separate. Nevertheless, the subfamilies have served as a convenient way to divide what is a quite large family of over 1200 described species, at least until a more extensive analysis can be conducted.

Enter Hedin et al. (2012) who take molecular data for six genes from seventy-odd sclerosomatids, mostly of the Holarctic genera Leiobunum and Nelima (both Leiobuninae). One suspects that the primary focus of the authors was originally just the phylogeny of these two genera, with the smattering of other sclerosomatids in the mix primarily intended as outgroups. But then they got a result that looks like this:

Yikes. That is a mess. The two main genera have not only failed to resolve as monophyletic, they have completely exploded. Gyas (the type species of the Gyinae) has apparently been so terrified by this show of opilionid pyrotechnics that it has scurried off to join the Phalangiidae. The Gagrellinae have also undergone something of a breakdown, appearing in four separate places on the tree (two of those including species supposedly of a single genus). But this is not simply a matter of poor resolution: many of the incongruent clades are reasonably supported. It is also worth noting that the results are not inconsistent with those recovered by Giribet et al. (2010) who included seven sclerosomatid species in their analysis.

As noted by the authors, the distribution of taxa in the results is not entirely random. There is a certain amount of biogeographic patterning, with the European and Japanese taxa included forming distinct clades, while the New World taxa form three clades. Within the Japanese taxa (which are some of the best studied sclerosomatids), there is some correlation with species groups recognised on morphological grounds, such as monophyly of the Leiobunum curvipalpe group. This latter example is interesting, as it represents a group of species that exhibit conservative features of external morphology despite showing broad variation in genitalic characters (Tsurusaki 1985b).

The sclerosomatine Astrobunus laevipes, photographed by Ch. Komposch.

Draining the morass of sclerosomatid taxonomy and phylogeny will doubtless be an arduous process, with all the large genera currently recognised likely to be polyphyletic. The support by Hedin et al. for previously recognised groups such as the Leiobunum curvipalpe complex suggest that morphological features will not be irrelevant in revising the family, but they will have to be analysed in their proper context. For instance, Hedin et al. suggest that the recovered polyphyly of Nelima may indicate that this genus, primarily defined by the absence of ornamentation found in other genera, may represent convergence through paedomorphosis (the acquisition of sexual maturity in a juvenile stage of development). In particular, this could explain how Giribet et al. (2009) found Nelima silvatica nested within two species of Sclerosomatinae, a primarily Mediterranean group of flattened, particularly heavily sclerotised species that might have been thought better supported than the other sclerosomatid subfamilies (unfortunately, Hedin et al.‘s analysis includes only a single sclerosomatine). The view of the road ahead might be daunting, but it promises to be a memorable journey.

Systematics of Sclerosomatidae
<==Sclerosomatidae (see below for synonymy)VFG14
|--Protolophus Banks 1893HT12, C92 [Protolophidae, Protolophinae, Protolophini]
| |--*P. tuberculatus Banks 1893C92
| |--P. cockerelli Goodnight & Goodnight 1942GG42a
| |--P. differens Goodnight & Goodnight 1942GG42a
| |--P. dixiensis Chamberlin 1925GG42a
| |--P. longipes Schenkel 1951E90
| |--P. niger Goodnight & Goodnight 1942 [incl. Parodiellus projectus Goodnight & Goodnight 1942]CH18
| |--P. rossi Goodnight & Goodnight 1943GG43
| `--P. singularis Banks 1893GG42a
| |--+--+--HolcobunusHT12
| | | `--+--GeayaHT12
| | | |--PrionostemmaHT12
| | | `--Carmenia Roewer 1915HT12, T-D04 [=Carmenis (l. c.)C84b]
| | | `--*C. bunifrons Roewer 1915C84b
| | `--+--Trachyrhinus Weed 1892HT12, T-D04
| | | |--*T. favosus (Wood 1868) [=Phalangium favosum, Astrobunus favosus]C81
| | | |--T. dicropalpus Cokendolpher 1981C81
| | | |--T. horneri Cokendolpher 1981C81
| | | |--T. marmoratus Banks 1894 [=T. maculatus (l. c.); incl. T. sonoranus Chamberlin 1925]C81
| | | |--T. mesillensis Cokendolpher 1981C81
| | | `--T. rectipalpus Cokendolpher 1981C81
| | `--Krusa Goodnight & Goodnight 1947HT12, C92
| | | i. s.: K. amazonica Roewer 1953KP-R02
| | | K. flava Goodnight & Goodnight 1946GG46
| | | K. hidalguensis Gaona-Escamilla, Francke & Ponce-Saavedra 2016G-EFP-S16
| | | K. metallica Goodnight & Goodnight 1946GG46
| | | K. tuberculata Goodnight & Goodnight 1946GG46
| | |--*K. mexicana Goodnight & Goodnight 1947C92, HT12, C92
| | `--+--K. annulata Goodnight & Goodnight 1945HT12, GG46
| | `--K. stellata Goodnight & Goodnight 1946HT12, GG46
| `--Sclerosomatinae [Sclerosomatini, Sclerosominae]C92
| | i. s.: Mastobunus Simon 1879C92
| | `--*M. tuberculifer (Lucas 1847)C92, E12 (see below for synonymy)
| | Granulosoma Martens 1973K06
| | |--*G. umidulum Martens 1973C92
| | `--G. dissimile Martens 2018M18
| | Pseudastrobunus Martens 1973K06
| | `--*P. perpusillus Martens 1973C92
| | Pseudohomalenotus Caporiacco 1935C92
| | `--*P. bicornutus Caporiacco 1935C92
| | Pygobunus Roewer 1957C92
| | |--*P. formosanus Roewer 1957C92
| | `--P. okadaiT07
| | 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
| |--HomalenotusHT12
| `--+--AstrobunusHT12
| `--+--NelimaHT12
| `--‘Leiobunum’ blackwalli Meade 1861HT12, D04 (see below for synonymy)
`--+--+--+--Togwoteeus Roewer 1952BHS13, C92
| | | `--T. biceps (Thorell 1877)BHS12 (see below for synonymy)
| | `--+--+--‘Leiobunum’ patzquarum Goodnight & Goodnight 1942HT12, GG46
| | | `--‘Leiobunum’ townsendii Weed 1893BHS12 [=Liobunum townsendiiCZ04]
| | `--+--Leuronychus Banks 1900BHS13, C92 [incl. Microgyas Schenkel 1951C92]
| | | |--*L. pacificus (Banks 1894) [=Liobunum pacificum; incl. *Microgyas banksi Schenkel 1951]C92
| | | `--L. fulviventris (Cambridge 1904) [=Liobunum fulviventre]R23
| | `--+--‘Leiobunum’ bogerti Goodnight & Goodnight 1942HT12, GG45
| | |--‘Leiobunum’ colimae Goodnight & Goodnight 1945HT12, GG45
| | `--+--‘Leiobunum’ potosum Goodnight & Goodnight 1942BHS12
| | `--+--‘Leiobunum’ hoogstraali Goodnight & Goodnight 1942HT12, GG46
| | `--‘Leiobunum’ tascum Goodnight & Goodnight 1945HT12, GG45
| `--+--+--‘Leiobunum’ aldrichi Weed 1893BHS12 (see below for synonymy)
| | `--+--+--‘Leiobunum’ uxorium Crosby & Bishop 1924BHS12
| | | |--‘Leiobunum’ vittatum (Say 1821)BHS13 (see below for synonymy)
| | | `--+--‘Leiobunum’ crassipalpe Banks 1900BHS12 [=Liobunum crassipalpeE66]
| | | `--‘Leiobunum’ relictum Davis 1934BHS12
| | `--+--+--‘Leiobunum’ bracchiolum (McGhee 1977)BHS12
| | | `--‘Leiobunum’ politum (Wood 1868)BHS12 (see below for synonymy)
| | `--‘Leiobunum’ calcar groupIMS11
| | |--+--‘Leiobunum’ calcar (Wood 1868)BHS13, BHS12 (see below for synonymy)
| | | `--‘Leiobunum’ hoffmani Ingianni, McGhee & Shultz 2012BHS13
| | `--+--‘Leiobunum’ euserratipalpe Ingianni, McGhee & Schultz 2011BHS13, BHS12
| | `--‘Leiobunum’ nigropalpi (Wood 1868)BHS13, BHS12 (see below for synonymy)
| `--+--+--+--‘Leiobunum’ flavum Banks 1894BHS12 [=Liobunum flavumE66]
| | | `--‘Leiobunum’ verrucosum (Wood 1870)BHS13, IMS11 (see below for synonymy)
| | `--+--‘Leiobunum’ holtae McGhee 1977BHS12
| | `--‘Leiobunum’ ventricosum (Wood 1868)BHS12 [=Phalangium ventricosumD34, Liobunum ventricosumE66]
| | |--L. v. ventricosumD34
| | |--L. v. floridanum Davis 1934D34
| | `--L. v. hiemale (Weed 1890)CH18 (see below for synonymy)
| `--+--‘Eumesosoma’ ocalensis Cokendolpher 1980HT12, E90
| `--+--‘Leiobunum’ royali Goodnight & Goodnight 1946BHS12
| `--+--+--‘Leiobunum’ bimaculatum Banks 1893BHS12 [=Liobunum bimaculatumR23]
| | |--‘Leiobunum’ peninsulare Davis 1934BHS12
| | `--‘Leiobunum’ speciosum Banks 1900BHS12, D34 [=Liobunum speciosumE66]
| `--+--Eumesosoma Cokendolpher 1980BHS13, T-D03 [=Mesosoma Weed 1892 non Otto 1821C92; Mesosomatinae]
| | |--E. roeweri (Goodnight & Goodnight 1943)BHS12 (see below for synonymy)
| | |--E. arnetti Cokendolpher 1980E90
| | |--E. ephippiatum (Roewer 1923)E90 [=Mesosoma ephippiatumR23]
| | |--E. nigrum (Say 1821)C92 [=Phalangium nigrumC92, Astrobunus nigerR23, Mesosoma nigrumCZ04]
| | `--E. sayi Cokendolpher 1980E90
| `--+--‘Hadrobunus’ maculosus (Wood 1868)BHS13 (see below for synonymy)
| `--+--Forbesium Weed 1890CZ04
| | `--*F. formosum (Wood 1868)CZ04 (see below for synonymy)
| `--Hadrobunus Banks 1900BHS13, C92
| |--*H. grandis (Say 1821)C92 (see below for synonymy)
| | |--H. g. grandisR23
| | `--H. g. similis (Wood 1892)R23 (see below for synonymy)
| |--H. davisi Goodnight & Goodnight 1942GG42b
| |--H. fusiformis Shultz 2010S10
| |--H. knighti Goodnight & Goodnight 1942GG42b
| `--H. nonsacculatus Shultz 2012S12
`--+--+--‘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 paessleriBH09, Liobunum paessleriR23]
`--+--+--Gagrellopsis Sato & Suzuki 1939HT12, C92
| | `--*G. nodulifera Sato & Suzuki 1939C92
| `--‘Leiobunum’ curvipalpe groupT91
| |--+--‘Leiobunum’ hikocola subgroupT91
| | | |--‘Leiobunum’ hikocola Suzuki 1966HT12, T06
| | | `--‘Leiobunum’ oharai Tsurusaki 1991T91
| | `--+--‘Leiobunum’ manubriatum Karsch 1881HT12, T06 [=Liobunum manubriatumR10, Zaleptus manubriatusR10]
| | `--‘Leiobunum’ platypenis subgroupT85b
| | |--‘Leiobunum’ globosum Suzuki 1953HT12, T03
| | `--‘Leiobunum’ platypenis Suzuki 1953 (see below for synonymy)T85b
| `--+--‘Leiobunum’ hiraiwai (Sato & Suzuki 1939)HT12, T06 (see below for synonymy)
| |--‘Leiobunum’ curvipalpe subgroupT90
| | |--‘Leiobunum’ curvipalpe Roewer 1910HT12, T06 [=Liobunum curvipalpi T90]
| | `--‘Leiobunum’ tohokuense Suzuki 1976HT12, T06
| `--+--‘Leiobunum’ kohyai Suzuki 1953HT12, T85b [=Liobunum kohyaiT85b]
| `--+--‘Leiobunum’ montanum Suzuki 1953HT12, T06 (see below for synonymy)
| `--‘Leiobunum’ hiasai subgroupT85b
| |--‘Leiobunum’ hiasai Suzuki 1976HT12, T85b
| |--‘Leiobunum’ sadoense Tsurusaki 1982T85b
| `--‘Leiobunum’ simplum Suzuki 1976T85b
| | `--‘Leiobunum’ japonicum Müller 1914HT12, T03 (see below for synonymy)
| | |--L. j. japonicumT91
| | |--L. j. taiwanum Suzuki 1977T91
| | `--L. j. uenoi Suzuki 1964T85b, S73
| `--+--PsathyropusHT12
| |--MelanopaHT12
| |--+--MarthanaHT12
| | `--GagrellulaHT12
| `--Systenocentrus Simon 1886HT12, C92 [incl. Oncobunus Thorell 1889C92]
| |--*S. quinquedentatus Simon 1886C92
| |--S. confucianus Hirst 1911R55a
| |--S. galeatus (Thorell 1889)R55a, C92 [=*Oncobunus galeatusC92]
| |--S. japonicus Hirst 1911R55a
| |--S. luteobiseriatus Suzuki 1982S85
| `--S. rufus Roewer 1955R55a
`--+--+--Pseudoliobunum Müller 1914 [incl. Metaliobunum Suzuki 1940]C92
| | `--*Pseudoliobunum japanense Müller 1914C92 [=Leiobunum japanenseHT12]
| | |--P. j. japanenseC92
| | `--‘Leiobunum’ j. suzukii Roewer 1957 (see below for synonymy)C92
| `--‘Leiobunum’ maximum Roewer 1910HT12, T91 [=Liobunum maximumR23]
| |--L. m. maximumS77b
| |--L. m. distinctum Suzuki 1973S73
| |--L. m. ‘formosum’ Suzuki 1976 non Phalangium formosum Wood 1868S77b
| `--L. m. yushan Suzuki 1976S77b
`--+--‘Leiobunum’ rubrum Suzuki 1966HT12, TT05
`--+--‘Nelima’ satoi Suzuki 1944HT12, TM00
`--+--‘Nelima’ aokiiHT12
|--‘Nelima’ genufusca (Karsch 1881)HT12, T03 (see below for synonymy)
| |--N. g. genufuscaS83
| |--N. g. pumilio Suzuki 1950S73
| `--N. g. tuberculata Suzuki 1983S83
|--‘Nelima’ nigricoxa Sato & Suzuki 1939HT12, T03
|--‘Nelima’ suzukii Tsurusaki 2003HT12, T03
`--+--‘Nelima’ parvaHT12
`--‘Nelima’ similisHT12

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 1840R23
Munequita Mello-Leitão 1941T-D04
|--*M. pulchra Mello-Leitão 1941C92 [incl. Holcobunus albianus Mello-Leitão 1944T-D03]
|--‘Holcobunus’ dentatus Roewer 1910T-D04
|--‘Prionostemma’ luteoscutum Roewer 1910T-D04
`--‘Prionostemma’ u-sigillatum Mello-Leitão 1938T-D04
Pectenobunus Roewer 1910 [incl. Acropiliops Mello-Leitão 1933, Caiza Roewer 1925]T-D04
|--*P. paraguayensis (Canestrini 1888) [=Opilio paraguayensis]C92
|--P. argentinus (Ringuelet 1959) [=Caiza argentina]KM20
|--*Caiza’ colliculosa Roewer 1925T-D04
`--P. ruricola (Mello-Leitão 1933)T-D04 [=*Acropiliops ruricolaC92]
Garleppa Roewer 1912T-D04
|--*G. granulata Roewer 1912C92
`--G. insperata Soares 1972T-D04
Bastioides Mello-Leitão 1931 (n. d.)T-D04
`--*B. coxopunctatus Mello-Leitão 1931 (n. d.) [=B. coxopunctata (l.c.)]T-D03
Tamboicus Roewer 1912C92
`--*T. fuhrmanni Roewer 1912C92
Amazonesia Soares 1970C84b
|--*A. quadriprocessigera Soares 1970C92
`--A. pulchra Soares 1970KP-R02
Azucarella Roewer 1959C84b
`--*A. weyrauchi Roewer 1959C92
Caluga Roewer 1959C84b
`--*C. rupicola Roewer 1959C92
Parageaya Mello-Leitão 1933T-D04 [incl. Corderobunus Roewer 1938C92]
|--*P. ciliata Mello-Leitão 1933C92
|--P. albifrons Goodnight & Goodnight 1942GG42c
|--P. corderoi (Mello-Leitão 1936)T-D04 [=Hypsibunus corderoiC92, *Corderobunus corderoiC92]
`--P. vittata Mello-Leitão 1939T-D04 [=Corderobunus vittatusKP-R02]
Fesa Roewer 1953C92
`--*F. tricolor Roewer 1953C92
Globulosoma Martens 1987M87
|--*G. gandakense Martens 1987M87
`--G. montivaga Martens 1987K06
Harmandina Schenkel 1954C92
`--*H. sinensis Schenkel 1954C92
Hexazaleptus Suzuki 1966C92
`--*H. junbesi Suzuki 1966C92
Himaldroma Martens 1987M87
|--*H. altus Martens 1987M87, K06 [=H. altaC92]
`--H. pineti Martens 1987K06
Himalzaleptus Martens 1987M87
`--*H. quinqueconicus Martens 1987M87
Holmbergiana Mello-Leitão 1931T-D04 [incl. Guaranobunus Ringuelet 1959T-D04, Simplicibunus Ringuelet 1954C84b]
|--*H. weyenberghii (Holmberg 1876) (see below for synonymy)C92
|--*Guaranobunus’ guaraniticus Ringuelet 1959T-D04
|--H. orientalis Ringuelet 1963T-D04
`--H. tibialis Ringuelet 1959T-D04
Nepalkanchia Martens 1990 [=Kanchia Martens 1987 non Moore 1883]C92
|--*N. pluviosilvestris (Martens 1987) [=*Kanchia pluviosilvestris]C92
`--‘Kanchia’ silvicola Martens 1987M87
Koyamaia Suzuki 1972C92
`--*K. curvipes Suzuki 1972C92
Liopagus Chamberlin 1916C92
`--*L. simplex Chamberlin 1916C92
Nepalgrella Martens 1987M87
|--*N. kortaliensis Martens 1987M87
`--N. yamputhini Martens 1987M87
Octozaleptus Suzuki 1966C92
`--*O. harai Suzuki 1966C92
Onostemma Mello-Leitão 1938C92
`--*O. imitans Mello-Leitão 1938C92
Oobunus Kishida 1930C92
`--*O. schizops Kishida 1930C92
Paragagrellina Schenkel 1963C92
`--*P. legendrei Schenkel 1963C92
Paratamboicus Mello-Leitão 1940C92
`--*P. bicornutus Mello-Leitão 1940C92
Paraumbogrella Suzuki 1963C92
`--P. pumilio (Karsch 1881)T03 (see below for synonymy)
Paruleptes Soares 1970C84b
`--*P. coronatus Soares 1970C92
Pokhara Suzuki 1970M87
|--*P. lineata Suzuki 1970M87
|--P. kathmandica Martens 1987M87
|--P. minuta Martens 1987M87
|--P. occidentalis Martens 1987M87
|--P. quadriconica Martens 1987M87
|--P. trisulensis Martens 1987M87
|--P. uenoi Martens 1987M87
`--P. yodai (Suzuki 1966) [=Hypsibunus yodai; incl. H. gibber Suzuki 1966]M87
Psammogeaya Mello-Leitão 1946T-D04
`--*P. arenata Mello-Leitão 1946C92
Pseudarthromerus Karsch 1892C92
`--*P. spurius Karsch 1892C92
Romerella Goodnight & Goodnight 1943C92
|--*R. punctata Goodnight & Goodnight 1943C92
|--R. bicolor Goodnight & Goodnight 1944GG53
|--R. brasiliensis Goodnight & Goodnight 1944GG44a
`--R. catharina Goodnight & Goodnight 1944GG44b
Sericicorpus Martens 1987M87
`--*S. nigrum Martens 1987M87
Taperina Roewer 1953C92
`--*T. nigripes Roewer 1953C92
Varinodulia Canals 1935TK01
`--*V. insularis Canals 1935C92
Xerogrella Martens 1987M87
`--*X. dolpensis Martens 1987M87
Abaetetuba Tourinho-Davis 2004T-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 2004T-D04
Echinobunus Roewer 1912C92
`--*E. elegans Roewer 1912C92
Diangathia Roewer 1955M87
`--*D. bovifrons Roewer 1955M87
Euceratobunus Roewer 1923C92
`--*E. pulcher (With 1903) [=Ceratobunus pulcher]C92
Orissula Roewer 1955C92
`--*O. heterospinulata Roewer 1955C92
Obigrella Roewer 1955C92
`--*O. nigra Roewer 1955C92
Bakerinulus Roewer 1955C92
`--*B. luzonicus Roewer 1955C92
Biceropsis Roewer 1935C92
`--*B. maculata Roewer 1935C92
Sinadroma Roewer 1955C92
`--*S. sherriffsi Roewer 1955C92
Carmichaelus Roewer 1929C92
`--*C. maculatus Roewer 1929C92
Carinobius Roewer 1955C92
`--*C. bicornutus Roewer 1955C92
Microzaleptus Roewer 1955C92
|--*M. quadratus Roewer 1955C92
`--M. montanus (Banks 1930) [=Metazaleptus montanus]R55b
Metaverpulus Roewer 1912M87
|--*M. hirsutus Roewer 1912 [incl. Euzaleptus pilosus Roewer 1929, Adungrella punctulata Roewer 1955]M87
|--M. albipunctatus (Roewer 1915) [=Harmandella albipunctata, Harmanda albipunctata]M87
|--M. bhutanicus Martens 1987M87
|--M. kanchensis Martens 1987K06
|--M. laevis Roewer 1955M87
|--M. multidentatus Martens 1987M87
`--M. persimilis Martens 1987M87
Pentazaleptus Roewer 1955C92
|--*P. adspersus Roewer 1955C92
|--P. guttatus Roewer 1955R55b
|--P. luteomaculatus Suzuki 1977S77a
`--P. rufus Suzuki 1977S77a
Adungrella Roewer 1955C92
`--*A. aenea Roewer 1955C92
Bastia Roewer 1910C92
|--*B. lineata Roewer 1910C92
|--B. elegans Suzuki 1977S77a
`--B. guttata Banks 1930R55b
Zaleptiolus Roewer 1955M87
|--*Z. laevipes Roewer 1955M87
|--Z. ater Martens 1987M87
`--Z. implicatus Suzuki 1970M87
Ceratobunoides Roewer 1923C92
|--*C. sumatranus Roewer 1923C92
|--C. bandarensis Roewer 1955R55b
`--C. mentaweiensis Roewer 1955R55b
Ceratobunellus Roewer 1911C92
|--*C. calcuttensis (With 1903) [=Ceratobunus calcuttensis]C92
|--C. brevipes (With 1903)R55b [=Ceratobunus brevipesR23]
`--C. philippinus Roewer 1955R55b
Tetraceratobunus Roewer 1915C92
|--*T. lineatus Roewer 1915C92
|--T. lithobius Roewer 1955R55b
`--T. marmoratus Roewer 1955R55b
Zaleptanus Roewer 1929C92
|--*Z. gravelyi Roewer 1929C92
|--Z. curvitarsus Suzuki 1977S77a
|--Z. promeus Roewer 1955R55b
`--Z. ramosus Suzuki 1977S77a
Cervibunus Roewer 1912C92
|--*C. maculatus Roewer 1912C92
|--C. ater Roewer 1955R55b
`--C. ornatus Roewer 1929R55b
Metazaleptus Roewer 1912M87
|--*M. hirsutus (With 1903) [=Zaleptus hirsutus]M87
|--M. borneensis Banks 1930R55b
|--M. rufescens Banks 1930R55b
`--M. striatus Suzuki 1985S85
Euzaleptus Roewer 1911M87
|--*E. minutus (With 1903) (see below for synonymy)M87
|--E. muticus Roewer 1929M87
`--E. sarawakensis Roewer 1911M87
Bullobunus Roewer 1910C92
|--*B. luteovittatus Roewer 1910C92
|--B. ater Roewer 1910R55b
|--B. bakeri Roewer 1926R55b
|--B. culionicus Suzuki 1977S77a
|--B. punctatus Suzuki 1977S77a
|--B. rufus Roewer 1955R55b
|--B. similis Roewer 1910R55b
`--B. unicolor Suzuki 1977S77a
Zaleptulus Roewer 1955C92
|--*Z. unicolor Roewer 1955C92
|--Z. annulipes (Banks 1930) [=Zaleptus annulipes]R55b
|--Z. banksi Roewer 1955R55b
|--Z. lineatus Roewer 1955R55b
`--Z. pulchellus (Banks 1930) [=Zaleptus pulchellus]R55b
Metahehoa Suzuki 1985C92
`--*M. granulata Suzuki 1985S85
Pseudosystenocentrus Suzuki 1985C92
`--*P. foveolatus Suzuki 1985S85
Antigrella Roewer 1954C92
`--*A. orissana Roewer 1954C92
Palniella Roewer 1929C92
`--*P. virididorsata Roewer 1929C92
Verrucobunus Roewer 1931C92
`--*V. trispinosus Roewer 1931C92
Scotomenia Thorell 1889C92
`--*S. cetrata Thorell 1889C92
Metadentobunus Roewer 1915C92
|--*M. formosae Roewer 1915C92
|--M. brevispinus Chen & Shih 2018CS18
`--M. garampiensis Suzuki 1944 [=M. formosae garampiensis]CS18
Hamitergum Crawford 1992 [=Lophacanthus Redikorzev 1936 non Stock 1880]C92
`--*H. eobium (Redikorzev 1936)C92 [=*Lophacanthus eobiusC92; incl. L. borchseniusi Charitonov 1957S65]
Hehoa Roewer 1929C92
`--*H. bunigera Roewer 1929C92
Baturitia Roewer 1931C92
`--*B. trispinosa Roewer 1931C92
Umbogrella Roewer 1955C92
`--*U. minuta Roewer 1954C92
Chebabius Roewer 1935K06
`--*C. angulatus Roewer 1935C92
Paradentobunus Roewer 1915C92
`--*P. aureomaculatus Roewer 1915C92
Padangrella Roewer 1954C92
`--*P. jacobsoni Roewer 1954C92
Pergagrella Roewer 1954C92
`--*P. monticola Roewer 1954C92
Gagrellenna Roewer 1929C92
`--*G. bipunctata Roewer 1929C92
Coonoora Roewer 1929C92
`--*C. biceratops Roewer 1929C92
Hypogrella Roewer 1955C92
`--*H. poecilis Roewer 1955C92
Sarasinia Roewer 1913C92
`--*S. punctata Roewer 1913C92
Gagrellina Roewer 1913C92
`--*G. vestita Roewer 1913C92
Syngagrella Roewer 1913C92
`--*S. bistriata Roewer 1913C92
Heterogagrella Roewer 1954C92
|--*H. indica Roewer 1954C92
`--H. biseriata Suzuki 1981S85
Gagrellissa Roewer 1931C92
|--*G. jacobsoni Roewer 1931C92
`--G. luteobiseriata Suzuki 1985S85
Akalpia Roewer 1915C92
|--*A. oblonga Roewer 1915C92
`--A. nipponica Sato & Suzuki 1938R55a
Melanopella Roewer 1931C92
|--*M. feuerborni Roewer 1931C92
`--M. marginata Roewer 1955R55a
Chasenella Roewer 1955C92
|--*C. luma Roewer 1933C92
`--C. pakka Roewer 1933R55a
Altobunus Roewer 1910C92
|--*A. formosus Roewer 1910C92
|--A. inermis (Simon 1877)R54a [=Gagrella inermisR23]
`--A. maculatus Roewer 1910R54a
Prodentobunus Roewer 1923C92
|--*P. unispinosus (Roewer 1912) [=Dentobunus unispinosus]C92
|--P. luteus Suzuki 1977S77a
|--P. nitidus Roewer 1955R55a
`--P. tao Roewer 1955R55a
Metasyleus Roewer 1955C92
|--*M. ephippiatus Roewer 1955C92
|--M. orissus Roewer 1955R55a
`--M. tenuis Roewer 1955R55a
Sataria Roewer 1915C92
|--*S. maculata Roewer 1915C92
|--S. coronata Roewer 1929R55a
`--S. unicolor Roewer 1914R55a
Syleus Thorell 1876 [=Acanthonotus Koch 1839 nec Goldfuss 1809 nec Ross 1835]C92
|--*S. niger (Koch 1839) [=*Acanthonotus niger]C92
|--S. mysoreus Roewer 1955R55a
`--S. rufus Roewer 1955R55a
Neogagrella Roewer 1913C92
|--*N. eximia Roewer 1913C92
|--N. balica Roewer 1931R54b
`--N. barnesi Roewer 1929R54b
Hologagrella Roewer 1910C92
|--*H. reticulata Roewer 1910C92
|--H. curvicornis Roewer 1913R55a
|--H. curvispina Banks 1930B30
|--H. luzonica Roewer 1910R55a
`--H. timorana Roewer 1955R55a
Bonthainia Roewer 1913 [incl. Apogrella Roewer 1955, Narmadella Roewer 1931, Nilgirisia Roewer 1929]C92
|--B. aenescens Roewer 1913C92
|--B. annulata Suzuki 1977S77a
|--*Apogrella’ celebensis Roewer 1955C92
|--*Narmadella’ elegans Roewer 1931C92
`--*Nilgirisia’ gravelyi Roewer 1929C92
Melanopula Roewer 1929C92
|--*M. biceps Roewer 1929C92
|--M. crassipes Suzuki 1977S77a
|--M. crassitarsis Roewer 1955R55a
|--M. insularis Roewer 1931R55a
|--M. shanensis Roewer 1955R55a
`--M. thaiensis Suzuki 1985S85
Hexomma Thorell 1876 [incl. Arthrocentrus Thorell 1889, Gagrella Roewer 1910 non Stoliczka 1869]C92
|--*H. vulcanicum (Doleschall 1859)C92 [=Phalangium vulcanicumC92, Gagrella vulcanicaR54a]
|--‘Gagrella’ arthrocentra Roewer 1910 [=*Arthrocentrus atratus Thorell 1889 non G. atrata Stoliczka 1869]C92
`--*Gagrella’ feae Thorell 1889C92
|--G. f. feaeR54a
|--G. f. bispinosa Thorell 1889R54a
|--G. f. conjugata Roewer 1954R54a
`--G. f. humeralis Thorell 1889R54a
Paragagrella Roewer 1912C92
|--*P. typus Roewer 1912C92
|--P. bakeri Roewer 1954R54b
|--P. basalis Roewer 1929R54b
|--P. bicolor Roewer 1954R54b
|--P. brevispina Banks 1930B30
|--P. mysorea Roewer 1939R54b
|--P. palpalis Roewer 1954R54b
|--P. roeweri Giltay 1930R54b
`--P. siberutiana Roewer 1931R54b
Umbopilio Roewer 1956K06
|--*U. paradoxus Roewer 1956K06
`--U. martensi Klimeš 2006K06
Eusclera Roewer 1910 [=Eusclerella Roewer 1910]C92
`--*E. aureomaculata Roewer 1910 [=*Eusclerella aureomaculata]C92
Microliobunum Roewer 1912C92
`--*M. brevipes Roewer 1912C92
Dilophiocara Redikorzev 1931C92
`--*D. bactrianum Redikorzev 1931 [=D. bactriana]C92
Micronelima Schenkel 1938C92
`--*M. brevipes Schenkel 1938C92
Schenkeliobunum Staręga 1964 [=Thrasychiroides Schenkel 1963 non Soares 1947]C92
`--*S. tuberculatum (Schenkel 1963) [=*Thrasychiroides tuberculatus]C92
Pseudomelanopa Suzuki 1974ZZ09
|--*P. taiwana Suzuki 1974ZZ09
`--P. liupan Zhang & Zhu 2009ZZ09
Amauropilio Mello-Leitão 1937C92
|--*A. lacoei (Petrunkevitch 1922)C92 [=Phalangium lacoeiC92, A. laweiD04, P. lawei (l. c.)C92]
`--A. atavus (Cockerell 1907) [=Leptobunus atavus]D07
Cosmobunus Simon 1879C92
|--*C. granarius (Lucas 1847)C92, E12 (see below for synonymy)
|--C. americanus Roewer 1957R57
|--C. auratus Goodnight & Goodnight 1946GG46
|--C. sagani Palencia, Peñalver et al. 2019PP19
`--C. unicolor Roewer 1910 [incl. C. unifasciatus Roewer 1923]PP19
Krusella Roewer 1953G-EFP-S16
`--*K. pilipes Roewer 1953C92
Goasheer Snegovaya, Cokendolpher & Mozaffarian 2018SCM18
`--*G. iranus (Roewer 1952) [=Homolophus iranus, Microliobunum iranum]SCM18
Mesobunus Huang, Selden & Dunlop 2009PP19, GT12
|--*M. martensi Huang, Selden & Dunlop 2009HSD09
`--M. dunlopi Giribet, Tourinho et al. 2012GT12
Paranelima Caporiacco 1938HT12, C92 [incl. Glabrurus Goodnight & Goodnight 1942C92]
|--*P. taibeli Caporiacco 1938C92
|--P. albalineata Goodnight & Goodnight 1942GG45
|--P. cerrana Goodnight & Goodnight 1942GG42c
|--P. correa Goodnight & Goodnight 1945GG45
|--P. lutzi (Goodnight & Goodnight 1942)GG45 [=*Glabrurus lutziC92]
`--P. mexicana (Goodnight & Goodnight 1942) [=Leptobunus mexicanus]C84a
Leiolima Prieto & Wijnhoven 2020PW20
`--*L. iberica Prieto & Wijnhoven 2020PW20

Nomen nudum: Systenocentrus uyemurai Kishida 1934TM00

*Cosmobunus granarius (Lucas 1847)C92, E12 [=Phalangium granariumD04; incl. P. flavounilineatum Lucas 1847D04, P. levipes Lucas 1847D04]

Eumesosoma roeweri (Goodnight & Goodnight 1943)BHS12 [=Mesosoma roeweriC92; incl. *M. nigrum Weed 1892 non Phalangium nigrum Say 1821C92, M. texanum Goodnight & Goodnight 1943C92]

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

*Forbesium formosum (Wood 1868)CZ04 [=Phalangium formosumD34, Leiobunum formosumHT12, Liobunum formosumE66]

Hadrobunus grandis (Say 1821)C92 [=Phalangium grandisS12, Astrobunus grandisR23, Hadrobunus grandeS12, Leptobunus grandisR23, Liobunum grandeR23, Phalangium grandeS12; incl. Leiobunum aurugineum Crosby & Bishop 1924S12]

Hadrobunus grandis similis (Wood 1892)R23 [=Liobunum grande var. similisS12, L. similis Wood 1890 (n. n.)CZ04]

‘Hadrobunus’ maculosus (Wood 1868)BHS13 [=Phalangium maculosumR23, Leptobunus maculosusR23, Liobunum maculosumR23]

*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 1893BHS12 [=Liobunum longipes aldrichiC84c; incl. Le. gordoni Goodnight & Goodnight 1945CZ04, Li. longipes Weed 1890 non Le. longipes Menge in Koch & Berendt 1854C84c]

‘Leiobunum’ blackwalli Meade 1861HT12, D04 [=Leiobunus blackwalliiD04, Liobunum blackwalliD04, Liobunum blackwilli (l. c.)D04, Liobunus blackwalliiD04; incl. Leiobunum granadinum Marcellino 1967M78, Leiobunum hemisphaericum Koch 1872D04]

‘Leiobunum’ calcar (Wood 1868)BHS13, BHS12 [=Phalangium calcarD34, Liobunum calcarE66; incl. Li. brunnea Walker 1928D34, Leiobunum cumberlandense McGhee in Ingianni, McGhee & Shultz 2011 (n. n.)BHS12, Le. serratipalpe Roewer 1910IMS11, Liobunum serratipalpeE66]

‘Leiobunum’ exilipes (Wood 1870)HT12, D34 [=Phalangium exilipesD34, Lei. exillipes (l. c.)CH18, Liobunum exilipesR23; incl. Leuronychus gertschiC84a]

‘Leiobunum’ hiraiwai (Sato & Suzuki 1939)HT12, T06 [=Nelima hiraiwaiT06; incl. L. hiraiwai fuji Suzuki 1976T06, L. hiraiwai izuense Suzuki 1976T06, L. hiraiwai longum Suzuki 1976T06, L. hiraiwai shiranense Suzuki 1976T06, L. tamanum Suzuki 1957T06, L. tsushimense Suzuki 1976T06]

‘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 1914HT12, T03 [=Liobunum japonicumTT05; incl. Le. wegneri Šilhavý 1976TT05]

‘Leiobunum’ montanum Suzuki 1953HT12, T06 [=Liobunum montanumT85a; incl. L. virgeum ontakense Suzuki 1976T85a, L. montanum sobosanum Suzuki 1976T85a, L. virgeum Suzuki 1976T85a]

‘Leiobunum’ nigropalpi (Wood 1868)BHS13, BHS12 [=Phalangium nigropalpiD34, Liobunum nigropalpiE66, Leiobunum nigripalpiIMS11, Li. nigripalpis non Leiobunum nigripalpe Simon 1889IMS11]

‘Leiobunum’ platypenis Suzuki 1953 [=Liobunum platypenis; incl. Leiobunum esoense Suzuki 1976]T85b

‘Leiobunum’ politum (Wood 1868)BHS12 [=L. politusD34; incl. L. cavernarum Roewer 1952CZ04, L. politum magnum Weed 1893D34]

‘Leiobunum’ ventricosum hiemale (Weed 1890)CH18 [=Phalangium ventricosum hiemaleCH18, Forbesium hyemaleCZ04, Leiobunum ventricosum hyemaleCZ04, Liobunum hyemaleR10]

‘Leiobunum’ verrucosum (Wood 1870)BHS13, IMS11 [=Phalangium verrucosumS08, Liobunum verrucosumS08; incl. Li. nigripes Weed 1892S08, Leiobunum nigripesS08]

‘Leiobunum’ vittatum (Say 1821)BHS13 [=Phalangium vittatumD34, Liobunum vittatumE66; incl. Phalangium dorsatum Say 1821D34, Leiobunum vittatum dorsatumD34, Liobunum vittatum minor Weed 1893CZ04]

*Mastobunus tuberculifer (Lucas 1847)C92, E12 [=Phalangium tuberculiferumD04, Sclerosoma tuberculiferaD04; incl. Homalenotus sicanus Pavesi 1876D04, S. sicanumD04]

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

‘Nelima’ elegans (Weed 1889)HT12, C92 [=Liobunum elegansCZ04, Leiobunum elegansD34; incl. Li. annulatum Walker 1928D34, Phalangium bicolor Wood 1868 non Fabricius 1793C84c, Astrobunus bicolorD34, Leiobunum bicolor non Koch 1848D34, Liobunum bicolorC84c]

‘Nelima’ genufusca (Karsch 1881)HT12, T03 [=Mitopus genufuscusR23; incl. Liobunum giganteum Loman 1902ST83, T03, Leiobunum giganteumST83, Nelima giganteaT03, N. tenuis Roewer 1957ST83]

Paraumbogrella pumilio (Karsch 1881)T03 [=Gagrella pumilioC92, Melanopa pumilioR23; incl. *Paraumbogrella huzitai Suzuki 1963C92]

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

Togwoteeus biceps (Thorell 1877)BHS12 [=Mitopus bicepsBH09, Homolophus bicepsBH09; incl. *Togwoteeus granipalpus Roewer 1952C92]

*Type species of generic name indicated


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