Histricostoma dentipalpe, copyright Alberto Gozzi.

Belongs within: Troguloidea.
Contains: Mitostoma, Paranemastoma, Nemastomella, Mediostoma, Nemastoma, Giljarovia.

The Nemastomatidae are a diverse group of often complexly ornamented harvestmen found in the Holarctic region. Living nemastomatids may be divided between the western Palaearctic Nemastomatinae, and the eastern Asian and North American Ortholasmatinae. In most Ortholasmatinae (except the basal Martensolasma jocheni) the ocularium bears a complex anterior process that forms a hood above the chelicerae.

Characters (from Gruber 2007): Body length 1.2–5.6 mm. Dorsum flattened to strongly arched; with scutum magnum, or scutum parvum with tendency to fusion; ocularium near anterior border of scutum, sometimes with processes, elaborated into a hood, or divided into two columns; three laminae suprachelicerales generally well developed and delimited from carapace; ozopores not freely visible; scutum finely or coarsely granulated, variously sculptured or armed with macrosculptural elements. Venter with corona analis well developed, as are free sternites; anterior sternite narrowed to base of genital operculum; genital operculum with well-developed basal suture in females, without it in males; spiracles not visible; prosomal sternal area fused with coxae; leg coxae well delimited by immovable, with marginal rows of tubercles, often anvil-shaped. Chelicerae of normal proportions, often sexually dimorphic; dentition of chela with narrow, diaphanous teeth. Pedipalp length and girth variable; femora with ‘basal bend’; tibiae and tarsi with narrowed bases and somewhat inflated, less markedly so if pedipalp is long and thin; tarsal claw absent, but with nipple-like rudiment; clavate glandular setae present especially on distal segments, sometimes reduced in adults. Legs generally moderately short to long; femora often with pseudoarticulations; tarsi moderately pluriarticulated. Penis long, shaft with incrassate bulb-shaped base containing two intrinsic penis muscles, their tendons therefore long; shaft may bear lateral wings. Ovipositur unsegmented, truncus soft-skinned, short to very long, with non-glandular setae, apical portion (furca) with two weakly delimited segments.

<==Nemastomatidae [Nemastomidae]
    |    |  i. s.: Asiolasma Martens 2019M19
    |    |           |--*A. angka (Schwendinger & Gruber 1992)M19 [=Dendrolasma angkaM19, Cladolasma angkaS10]
    |    |           |--A. ailaoshan (Zhang, Zhao & Zhang 2018) [=Cladolasma ailaoshan]M19
    |    |           |--A. billsheari Martens 2019M19
    |    |           |--A. damingshan (Zhang & Zhang 2013) [=Cladolasma damingshan]M19
    |    |           |--A. juergengruberi Martens 2019M19
    |    |           `--A. schwendingeri Martens 2019M19
    |    |         Cryptolasma Cruz-López, Cruz-Bonilla & Francke 2018M19
    |    |--Martensolasma Shear 2006S13
    |    |    `--*M. jocheni Shear 2006S10
    |    |--+--Dendrolasma Banks 1894S13
    |    |  |    |--*D. mirabile Banks 1894 [=D. mirabilis]S13
    |    |  |    `--D. dentipalpe Shear & Gruber 1983S10
    |    |  `--Cladolasma Suzuki 1963S13
    |    |       `--*C. parvulum Suzuki 1963 [=C. parvula, Dendrolasma parvula, D. parvulum]S13
    |    `--+--Ortholasma Banks 1894S13
    |       |    |--*O. rugosum Banks 1894 [=O. rugosa]S10
    |       |    |--O. colossus Shear 2010S10
    |       |    |--O. coronadense Cockerell 1916 [=O. coronadensis; incl. O. setulipes Shear & Gruber 1983]S13
    |       |    |--O. levipes Shear & Gruber 1983S10
    |       |    `--O. pictipes Banks 1911R23
    |       `--Trilasma Goodnight & Goodnight 1942 [incl. Ruaxphilos Goodnight & Goodnight 1945]S13
    |            |--*T. bolivari Goodnight & Goodnight 1942S13 (see below for synonymy)
    |            |--T. chipinquensis Shear 2010S10
    |            |--T. hidalgo Shear 2010S10
    |            |--T. petersprousei Shear 2010S10
    |            |--T. ranchonuevo Shear 2010S10
    |            |--T. sbordonii Šilhavý 1973S13 [=Ortholasma sbordoniiS10]
    |            |--T. tempestado Shear 2010S10
    |            |--T. trispinosum Shear 2010S10
    |            `--T. tropicum Shear 2010S10
    `--Nemastomatinae [Giljaroviinae, Mitostomatinae]G07
         |  i. s.: Sinostoma Martens 2016M17
         |           `--S. yunnanicum Martens 2016M17
            |  `--+--Carinostoma Kratochvíl 1958S13
            |     |    |--*C. carinatum (Roewer 1914) [=Nemastoma carinatum, Mitostoma (Carinostoma) carinatum]S13
            |     |    |--C. elegans (Sørensen in Lendl 1894)KM20 (see below for synonymy)
            |     |    `--C. ornatum (Hadži 1940) [=Nemastoma ornatum, Mitostoma (Mitostoma) ornatum]S13
            |     `--Histricostoma Kratochvíl 1958 [incl. Cretostoma Kratochvíl 1958]S13
            |          |--*H. drenskii Kratochvíl 1958M06
            |          |--H. anatolicum (Roewer 1962) [=Nemastoma anatolicum]S13
            |          |--H. argenteolunulatum (Canestrini 1875) (see below for synonymy)S13
            |          |--H. caucasicum (Redikortsev 1936) [=Nemastoma caucasicum]S13
            |          |--H. creticum (Roewer 1927) (see below for synonymy)S13
            |          |--H. dentipalpe (Ausserer 1867) (see below for synonymy)S13
            |          |--H. gruberi Snegovaya & Marusik 2012S13
            |          |--H. mitovi Snegovaya & Marusik 2012S13
            |          `--H. tuberculatum (Koch & Berendt 1854) [=Nemastoma tuberculatum]S02
            `--+--+--Saccarella Schönhofer & Martens 2012S13
               |  |    `--*S. schilleri Schönhofer & Martens 2012S13
               |  |--NemastomellaS13
               |  |--Starengovia Snegovaya 2010S13
               |  |    |--*S. kirgizica Snegovaya 2010 [=S. kirgisica]S13
               |  |    |--S. ivanloebli Martens 2017M17
               |  |    |--‘Mediostoma’ pamiricum Staręga 1986M17 [=Centetostoma pamiricum Staręga 1978 (n. n.)S13]
               |  |    `--S. quadrituberculata Zhang & Martens 2018ZM18
               |  `--+--Acromitostoma Roewer 1951 [incl. Hispanostoma Kratochvíl 1958]S13
               |     |    |--*A. rhinoceros (Roewer 1917) [=Nemastoma rhinoceros, A. rhinocerus (l. c.)]S13
               |     |    `--A. hispanum (Roewer 1917) [=Nemastoma hispanum, Carinostoma (*Hispanostoma) hispanum]S13
               |     `--Centetostoma Kratochvíl 1958S13
               |          |--*C. centetes (Simon 1881) [=Nemastoma centetes, Mitostoma centetes, Nemastomella centetes]S13
               |          |--C. dubium (Mello-Leitao 1936)GE02
               |          |--C. juberthiei Martens 2011S13
               |          |--C. scabriculum (Simon 1879) [=Nemastoma scabriculum]S13
               |          `--C. ventalloi (Mello-Leitao 1936) [=Nemastoma ventalloi]S13
               `--+--Hadzinia Roewer 1966S13, N05
                  |    `--*H. karamani (Hadži 1940) [=Nemastoma karamani, Buresiolla karamani, Hadzinia caramani (l. c.)]S13
                  |--Pyza Staręga 1976S13
                  |    |--*P. bosnica (Roewer 1917) (see below for synonymy)S13
                  |    |--P. anatolica (Roewer 1959) [=Crosbycus anatolicus]S13
                  |    |--P. navarrense (Roewer 1951) [=Nemastoma navarrense; incl. N. lindbergi Roewr 1959]S13
                  |    `--P. taurica Gruber 1979S13
                  |  `--Vestiferum Martens 2006S13
                  |       |--*V. funebre (Redikorzev 1936) (see below for synonymy)M06
                  |       `--V. alatum Martens 2006M06
                     |--Caucnemastoma Martens 2006S13
                     |    |--*C. golovatchi Martens 2006S13
                     |    `--C. martensi Snegovaya in Snegovaya & Chumachenko 2011S13
                     `--Nemaspela Šilhavý 1966S13
                          |--*N. sokolovi (Ljovushkin & Starobogatov 1963) [=Buresiolla sokolovi]M06
                          |--N. abchasica (Ljovushkin & Starobogatov 1963) [=Buresiolla abchasica]M06
                          |--N. birsteini Ljovuschkin 1972S13
                          |--N. caeca (Grese 1911) [=Nemastoma caecum, Buresiolla caecum, Nemaspela coeca (l. c.)]S13
                          |--N. femorecurvata Martens 2006M06
                          |--N. kovali Chemeris 2009S13
                          `--N. taurica (Lebedinski 1914) [=Phalangodes taurica]S13

Carinostoma elegans (Sørensen in Lendl 1894)KM20 [=Nemastoma elegansKM20, Mitostoma (Carinostoma) elegansS13; incl. N. elegans var. batorligetiense Szalay 1952S13, Carinostoma elegans batorligetienseS13]

Histricostoma argenteolunulatum (Canestrini 1875) [=Nemastoma dentipalpe var. argenteolunulata, Carinostoma argenteolunulatum, N. argenteolunulatum]S13

Histricostoma creticum (Roewer 1927) [=Nemastoma creticum, Centetostoma (*Cretostoma) creticum, Mitostoma creticum]S13

Histricostoma dentipalpe (Ausserer 1867) [=Nemastoma dentipalpe, Carinostoma dentipalpe, N. (Lugubrostoma) dentipalpe; incl. Histricostoma slovenicum Hadži 1973]S13

*Pyza bosnica (Roewer 1917) [=Nemastoma bosnicum, N. (Lugubrostoma) bosnicum; incl. N. gallwitzi Roewer 1923, N. (L.) goliae Hadži 1973, N. (L.) bosnicum orientale Kratochvíl 1958, N. (L.) sarae Hadži 1973]S13

*Trilasma bolivari Goodnight & Goodnight 1942S13 [=Ortholasma (*Trilasma) bolivariS10; incl. *Ruaxphilos petrunkevitchou Goodnight & Goodnight 1945S13]

*Vestiferum funebre (Redikorzev 1936) [=Nemastoma funebre, N. (Lugubrostoma) funebre; incl. N. brunneum Redikorzev 1936]M06

*Type species of generic name indicated


[GE02] Giribet, G., G. D. Edgecombe, W. C. Wheeler & C. Babbitt. 2002. Phylogeny and systematic position of Opiliones: a combined analysis of chelicerate relationships using morphological and molecular data. Cladistics 18: 5–70.

[G07] Gruber, J. 2007. Nemastomatidae Simon, 1872. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 148–151. Harvard University Press: Cambridge (Massachusetts).

[KM20] Kury, A. B., A. C. Mendes, L. Cardoso, M. S. Kury & A. A. Granado. 2020. WCO-Lite: Online world catalogue of harvestmen (Arachnida, Opiliones). Version 1.0—Checklist of all valid nomina in Opiliones with authors and dates of publication up to 2018. Published by the author: Rio de Janeiro.

[M06] Martens, J. 2006. Weberknechte aus dem Kaukasus (Arachnida, Opiliones, Nemastomatidae). Senckenbergiana Biologica 86 (2): 145–210.

[M17] Martens, J. 2017. On Starengovia Snegovaya, a genus of Asian nemastomatines (Arachnida: Opiliones: Nemastomatidae). Revue Suisse de Zoologie 124 (2): 187–201.

[M19] Martens, J. 2019. An ancient radiation: ortholasmatine harvestmen in Asia—a new genus, three new species and a revision of the known species (Arachnida, Opiliones, Nemastomatidae). Revue Suisse de Zoologie 126 (1): 79–110.

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

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

[S13] Schönhofer, A. L. 2013. A taxonomic catalogue of the Dyspnoi Hansen and Sørensen, 1904 (Arachnida: Opiliones). Zootaxa 3679 (1): 1–68.

[S10] Shear, W. A. 2010. New species and records of ortholasmatine harvestmen from México, Honduras, and the western United States (Opiliones, Nemastomatidae, Ortholasmatinae). ZooKeys 52: 9–45.

[S02] Staręga, W. 2002. Baltic amber harvestmen (Opiliones) from Polish collections. Annales Zoologici 52 (4): 601–604.

[ZM18] Zhang, C., & J. Martens. 2018. Ancient home or in exile? The easternmost species of genus Starengovia Snegovaya, 2010 found in China (Opiliones, Nemastomatidae, Nemastomatinae). ZooKeys 770: 105–115.

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