Lacinius

Lacinius horridus, copyright AfroBrazilian.

Belongs within: Phalangiidae.

Lacinius is a western Palaearctic genus of generally spiny harvestmen.

Characters (from Martens 1978): Sexual dimorphism insignificant. Pedipalp primarily denticulate on femur and patella, rarely on other segments and then sparse, all segments lacking well-developed apophysis but sometimes with medial, brush-like hairy protrusions. Cheliceral lamellae smooth. Frontal margin with three median thorn-like denticles, either equally long or the middle longest. Tuber oculorum low, one or two times its own length from frontal margin. Truncus of penis slender to stocky, continuously narrowed distad from the base, sometimes slightly enlarged sub-distally; basal part of truncus rounded; mostly dorso-distally or ventro-distally flat or dorso-ventrally flattened; glans mostly sharply keeled below; musculature occupying at least basal half of trunk; truncus never with dorso-distal longitudinal groove.

<==Lacinius Thorell 1876 MDP15 [incl. Acantholophus Koch 1839 non Boisduval 1835 C92, Bidentolophus Roewer 1912 MDP15]
    |--*L. horridus (Panzer 1794) C92 (see below for synonymy)
    |--‘Opilio’ adungius Roewer 1956 [=Bidentolophus adungius; incl. O. birmanicus Roewer 1956] S03
    |--L. angulifer (Simon 1878) MDP15 [=Acantholophus angulifer S84]
    |--L. bidens (Simon 1880) MDP15 (see below for synonymy)
    |--L. bizleyi Mitov, Dunlop & Penney 2015 MDP15
    |--L. dentiger (Koch 1848) MDP15 (see below for synonymy)
    |--L. ephippiatus (Koch 1835) MDP15 (see below for synonymy)
    |--L. erinaceus Staręga 1966 MDP15
    |--L. insularis Roewer 1923 MDP15
    `--L. longisetus (Thorell 1876) [=Acantholophus longisetus] R23

Lacinius bidens (Simon 1880) MDP15 [=Acantholophus bidens MDP15, *Bidentolophus bidens C92, Egaenus bidens TTL00, Euphalangium bidens S03; incl. Opilio hexaspinulatus Saitô 1936 S03, O. koreanus Charitonov 1957 S03, O. peipingensis Wang 1941 S03, O. tchangi Wang 1941 S03, Phalangium nordenskioeldi var. transbaicalica Kulczyński 1901 ZSK99, Euphalangium nordenskioldi var. transbaicalicum R23, TTL00, Homolophus transbaicalicus ZSK99]

Lacinius dentiger (Koch 1848) MDP15 [=Acantholophus dentiger D04; incl. A. annulipes Koch 1867 D04, Bidentolophus bavaricus Roewer 1957 D04, Lacinius chelodentatus Hadži 1973 D04, L. labacensis Hadži 1931 D04]

Lacinius ephippiatus (Koch 1835) MDP15 [=Opilio ephippiatus D04, Acantholophus ephippiatus D04, Oligolophus ephippiatus D04; incl. Odiellus hungaricus Kolosvary 1941 D04, Lacinius oligodentatus Hadži 1931 D04, Odiellus rucneri Hadži 1973 D04, Opilio vittiger Meade 1855 D04, Ol. vittiger D04]

*Lacinius horridus (Panzer 1794) C92 [=Phalangium horridum D04, Acantholophus horridus D04, Opilio horridus D04; incl. A. bellicosus Sørensen 1894 D04, Lacinius horridus bulgaricus Šilhavý 1965 D04, A. coronatus Koch 1867 D04, Opilio coronatus Roewer 1911 D04, Odiellus coronatus M78, Lacinius dorsogranulatus Morin 1931 D04, L. gallipoliensis Roewer 1923 D04, Acantholophus granulatus Canestrini 1872 D04, Odiellus granulatus D04, A. hermanni Simon 1879 D04, Opilio hispidus Herbst 1798 D04, *A. hispidus C92, Lacinius hispidus D04, Phalangium hispidum D04, Acantholophus kochi Simon 1879 D04, A. lemniscatus Simon 1882 D04, Lacinius parisii Trossarelli 1934 D04, A. spinulosus Koch 1839 D04, Odiellus toscanus Roewer 1923 M78, Lacinius toscanus M78, L. zavalensis Hadži 1973 N05]

*Type species of generic name indicated

REFERENCES

[C92] Crawford, R. L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1–60.

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[M78] Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[MDP15] Mitov, P. G., J. A. Dunlop & D. Penney. 2015. A new species of Lacinius in amber (Arachnida: Opiliones). Fossil Record 18: 37–42.

[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.

[S84] Staręga, W. 1984. Revision der Phalangiidae (Opiliones), III. Die afrikanischen Gattungen der Phalangiinae, nebst Katalog aller afrikanischen Arten der Familie. Annales Zoologici 38 (1): 1–79.

[S03] Staręga, W. 2003. On the identity and synonymies of some Asiatic Opilioninae (Opiliones: Phalangiidae). Acta Arachnologica 52 (2): 91–102.

[TTL00] Tsurusaki, N., A. N. Tchemeris & D. V. Logunov. 2000. Two new species of Opiliones from southern Siberia and Mongolia, with an establishment of a new genus and redefinition of the genus Homolophus (Arachnida: Opiliones: Phalangiidae). Acta Arachnologica 49 (1): 73–86.

[ZSK99] Zhu, M., D. Song & J.-P. Kim. 1999. Notes on three species of the genus Homolophus (Opiliones: Phalangiidae) from China. Korean Arachnology 15 (1): 7–12.

Zachaeus

Zachaeus crista, copyright Ondřej Machač.

Belongs within: Phalangiidae.

Zacheus is a genus of large-bodied phalangiid harvestmen found in the eastern Mediterranean and western Asia (Kurt et al. 2015). Historically, there has been confusion regarding the correct spelling of the genus name; Crawford (1992) provides reasoning for use of the spelling 'Zachaeus'.

Characters (from Kurt et al. 2015): Body large, heavily denticulated dorsally; chelicerae usually strong, second segment enlarged; pedipalps normally structured, strong, robust; legs short, first pair much thicker than others; truncus of penis basally widened, parallel-sided on distal half, distally shallow spoon-shaped, glans usually banana-shaped (shorter in Z. anatolicus and Z. seyyari), stylus long.

<==Zachaeus Koch 1839 K14 [=Zacheus Koch 1839 C92]
    |--Z. crista (Brullé 1832) K14 (see below for synonymy)
    |--Z. anatolicus (Kulczyński 1903) K14 [=Egaenus crista var. anatolicus R23, Z. crista var. anatolicus R23]
    |--Z. birulai Redikorzev 1936 S99
    |--Z. canaliculatus Kollar in Roewer 1911 [=Phalangium canaliculatum ms] R23
    |--Z. caucasicus Roewer 1911 R23
    |--Z. cubanus Roewer 1923 R23
    |--‘Dasylobus’ egaenoides Simon 1885 (n. d.) R23
    |--Z. hebraicus (Simon 1884) K14 [=Phalangium hebraicum R23, Z. crista var. hebraicus R23]
    |--Z. kervillei (Sørensen 1912) (n. d.) KKY15 [=Egaenus kervillei R23]
    |--Z. leucomelas (Simon 1884) R23 [=Phalangium hebraicum leucomelas R11, Zachaeus crista var. leucomelas R23]
    |--Z. lupatus (Eichwald 1830) [incl. E. gulosus Simon 1878] S03
    |--Z. macrinus Roewer 1956 R56
    |--Z. mirabilis (Caporiacco 1949) KKY15
    |--Z. orchimonti (Giltay 1933) K14
    |--Z. palpipes Roewer 1915 R23
    |--‘Opilio’ punctipes Koch 1878 (n. d.) R23
    |--Z. redikorzevi (Staręga & Chevrizov 1978) K14
    |--Z. seyyari Kurt, Koç & Yağmur 2015 KKY15
    |--Z. shachdag (Snegovaya & Staręga 2008) KKY15
    `--Z. simferopolensis Chemeris & Kovblyuk 2005 [=Z. simpheropolensis] CK05

Zachaeus crista (Brullé 1832) K14 [=Phalangium crista C92, Egaenus crista C92; incl. E. hungaricus Lendl 1894 M78, Zachaeus crista var. hungaricus R23, Paropilio lineatus Roewer 1956 M78, *Zachaeus mordax Koch 1839 C92, Egaenus mordax R23, Z. trinotatus Koch 1839 R23, E. variegatus Lendl 1894 M78, Zachaeus crista var. variegatus R23]

*Type species of generic name indicated

REFERENCES

[CK05] Chemeris, A. N., & N. M. Kovblyuk. 2005. A contribution to the knowledge of the harvestman fauna of the Crimea (Arachnida: Opiliones). Arthropoda Selecta 14 (4): 305–328.

[C92] Crawford, R. L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1–60.

[K14] Kurt, K. 2014. Updated checklist of harvestmen (Arachnida: Opiliones) in Turkey. Arch. Biol. Sci. 66 (4): 1617–1631.

[KKY15] Kurt, K., H. Koç & E. A. Yağmur. 2015. A new species of Zachaeus C. L. Koch from Turkey (Opiliones, Phalangiidae). ZooKeys 514: 15–23.

[M78] Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

[R11] Roewer, C.-F. 1911. Übersicht der Genera der Subfamilie der Phalangiini der Opiliones Palpatores nebst Beschreibung einiger neuer Gattungen und Arten. Archiv für Naturgesichte 77 (Suppl. 2): 1–106.

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

[R56] Roewer, C. F. 1956. Über Phalangiinae (Phalangiidae, Opiliones Palpatores). (Weitere Weberknechte XIX). Senckenbergiana Biologica 37 (3–4): 247–318.

[S99] Snegovaya, N. 1999. Contribution to the harvest spider (Arachnida, Opiliones) fauna of the Caucasus. Tr. J. of Zoology 23: 453–459.

[S03] Staręga, W. 2003. On the identity and synonymies of some Asiatic Opilioninae (Opiliones: Phalangiidae). Acta Arachnologica 52 (2): 91–102.

Ischyropsalis

Male Ischyropsalis hellwigii hellwigii, copyright Axel Schönhofer.

Belongs within: Ischyropsalidoidea.

Ischyropsalis is a genus of harvestmen found in Europe, members of which have conspicuously enlarged chelicerae in both sexes. Members of this genus are specialist predators of snails, using their large chelicerae to break open the snail's shell.

Characters (from Schönhofer 2013): Palps long and thin, elongated to enable interaction with the chelicerae. Chelicerae massive and enlarged, longer than the body; in most species glandular fields of males associated with knob-like apophyses exposing the field to dorsal and/or distal side. Truncus penis simple, filled up to two thirds with a single muscle; muscular tendon short; base of truncus divided into two short root-like structures; glans penis usually simple and conical, but inflated and distinct from the truncus in many species; the sclerotised ventral plate mostly short, from roughly rhombic to deeply bi-lobed at the base; sclerotised area of glans with several defined regions exhibiting dense fields of mostly very thick, long and backward-pointing hairs; glans gradually tapering into a long stylus, the latter simple and pointed, bent at the base but without contortion or additional structures. Tuber oculorum without spine. Dorsal armature mostly lacking, dorsum smooth, often glossy, in few species opisthosomal areas raised to low, pointed bumps. Two or many spine-like metapeltidial structures, arranged in a transversal row.

<==Ischyropsalis Koch 1839 (see below for synonymy) S13
    |--*I. kollari Koch 1839 (see below for synonymy) S13
    |--I. adamii Canestrini 1873 (see below for synonymy) S13
    |--I. cantabrica Luque & Labrada 2012 S13
    |--I. carli Lessert 1905 [incl. I. pentelica Roewer 1950] S13
    |--I. dentipalpis Canestrini 1872 [incl. I. helvetica Roewer 1916] S13
    |--I. dispar Simon 1872 [incl. I. archeri Roewer 1950, I. goodnighti Roewer 1950, I. noltei Dresco 1972] S13
    |--I. gigantea Dresco 1968 S13
    |--I. hadzii Roewer 1950 [=Phalangium cancroides Schmidt 1851 non Acarus cancroides Linnaeus 1758] S13
    |--I. hellwigii (Panzer 1794) S13 [=Phalangium hellwigii S13, Opilio helwigii D04]
    |    |--I. h. hellwigii S13 (see below for synonymy)
    |    `--I. h. lucantei Simon 1879 [incl. I. superbus Simon 1881, I. superba] S13
    |--I. hispanica Roewer 1953 [=I. hispanum] S13
    |--I. lithoclasica Schönhofer & Martens 2010 S13
    |--I. luteipes Simon 1872 S13 (see below for synonymy)
    |--I. magdalenae Simon 1881 S13 [=I. madalenae (l. c.) S13, I. madelenae D04]
    |--I. manicata Koch 1869 (see below for synonymy) S13
    |--I. muellneri Hamann 1898 [=I. muelleri (l. c.); incl. I. alfkeni Roewer 1950, I. corcyraea Roewer 1914] S13
    |--I. navarrensis Roewer 1950 [incl. I. espagnoli Dresco 1968, I. nicaea Roewer 1950, I. turki Roewer 1950] S13
    |--I. nodifera Simon 1879 (see below for synonymy) S13
    |--I. petiginosa Simon 1913 [=I. pectiginosa (l. c.)] S13
    |--I. pyrenaea Simon 1872 [=I. helwigi var. pyrenaea] S13
    |    |--I. p. pyrenaea [incl. I. moreana Roewer 1950, I. pyrenaea venasquensis Dresco 1967] S13
    |    `--I. p. alpinula Martens 1978 S13
    |--I. ravasinii Hadži 1942 S13
    |--I. redtenbacheri Doleschall 1852 R23
    |--I. robusta Simon 1872 [=Ischiropsalis robusta; incl. I. lusitanica Roewer 1923] S13
    `--I. strandi Kratochvíl 1936 [incl. I. knirschi Roewer 1950, I. ruffoi Caporiacco 1947] S13

Ischyropsalis Koch 1839 [=Ischiropsalis (l. c.); incl. Lhermia Lucas 1866, Lermia (l. c.), Odontopalpa Hadži 1931; Ischyropsalidinae] S13

Ischyropsalis adamii Canestrini 1873 [incl. I. amseli Roewer 1950, I. apuanus Caporiacco 1930, I. apuana, I. kratochvili Roewer 1950, I. apuanus nanus Dresco 1968] S13

Ischryosalis hellwigii hellwigii (Panzer 1794) S13 [incl. I. crassichelis Roewer 1950 S13, I. frankenbergi Bartoš 1938 S13, I. franzi Roewer 1950 S13, I. herbstii Koch 1847 S13, I. pectinifera Hadži 1928 S13, I. plicata Roewer 1923 S13, I. taunica Müller 1924 S13, I. tunica D04]

*Ischyropsalis kollari Koch 1839 [incl. I. bosnica Roewer 1914, I. danubia Roewer 1950, I. nivalis Roewer 1950, I. reimoseri Roewer 1950, I. scutata Roewer 1950, I. segregata Roewer 1950, I. spinichelis Roewer 1950, I. strasseri Roewer 1950, I. styriaca Roewer 1950, I. tirolensis Roewer 1950, I. (Odontopalpa) triglavensis Hadži 1931, I. troglodytes Roewer 1950] S13

Ischyropsalis luteipes Simon 1872 S13 [incl. I. caporiaccoi Roewer 1950 S13, I. caporioccoi (l. c.) S13, I. pestae Roewer 1950 S13, *Lhermia spinipes Lucas 1866 (n. o.) S13, Lermia (l. c.) spinipes R23]

Ischyropsalis manicata Koch 1869 [=I. manicatus (l. c.); incl. I. austriaca Roewer 1950, I. balcanica Roewer 1950, I. dacica Roewer 1916, I. ionatae Wajgel 1874 (n. n.), I. ionatai, I. kaestneri Roewer 1940, I. helvetica milleri Kratochvíl 1934] S13

Ischyropsalis nodifera Simon 1879 [incl. I. asturica Roewer 1950, I. corsica Roewer 1950, I. janetscheki Roewer 1950, I. sharpi Simon 1879] S13

*Type species of generic name indicated

REFERENCES

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[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.

Ischyropsalidoidea

Sabacon cf. cavicolens, copyright Marshal Hedin.

Belongs within: Palpatores.
Contains: Ischyropsalis.

The Ischyropsalidoidea are a clade of harvestmen found in the Holarctic region, members of which have an unsegmented ovipositor in females, and a penis with a single muscle in males (Shultz 1998).

<==Ischyropsalidoidea [Ischropsalidae, Ischyropsalidae, Ischyropsalidoidae]
    |  i. s.: Piankhi Dunlop, Bartel & Mitov 2012 DBM12
    |           `--*P. steineri Dunlop, Bartel & Mitov 2012 DBM12
    |--Ischyropsalididae S13
    |    |--Ischyropsalis S13
    |    `--Ceratolasmatinae [Ceratolasmatidae] S13
    |         |--Ceratolasma Goodnight & Goodnight 1942 S98
    |         |    `--*C. tricantha Goodnight & Goodnight 1942 GG42
    |         `--Acuclavella Shear 1986 G07b, G07a
    |              |--*A. cosmetoides Shear 1986 S13
    |              |--A. merickeli Shear 1986 BH09
    |              |--A. quattuor Shear 1986 E90
    |              `--A. shoshone Shear 1986 E90
    `--+--Taracidae S13
       |    |--Taracus Simon 1879 GV09, S98
       |    |    |--*T. packardi Simon 1879 S79
       |    |    |--T. birsteini Ljovuschkin 1971 DBM12
       |    |    |--T. gertschi Goodnight & Goodnight 1942 GG42
       |    |    |--T. malkini Goodnight & Goodnight 1945 GG45
       |    |    |--T. nigripes Goodnight & Goodnight 1943 GG43
       |    |    |--T. pallipes Banks 1894 R23
       |    |    |--T. silvestrii Roewer 1929 S13
       |    |    `--T. spinosus Banks 1894 [=T. spinosa] R23
       |    `--+--Crosbycus Roewer 1914 S10
       |       |    |--*C. dasycnemus (Crosby 1911) [=Nemastoma dasycnemum] S13
       |       |    |--C. pentelicus Roewer 1951 (n. d.) S13
       |       |    `--C. speluncarum Roewer 1951 (n. d.) S13
       |       `--Hesperonemastoma Gruber 1970 GV09, S10
       |            |--*H. pallidimaculosum (Goodnight & Goodnight 1945) (see below for synonymy) S10
       |            |--H. kepharti (Crosby & Bishop 1924) [=Nemastoma kepharti] S13
       |            |--H. modestum (Banks 1894) BH09 [=Nemastoma modesta BH09, N. modestum R23]
       |            |--H. packardi (Roewer 1914) (see below for synonymy) S10
       |            `--H. smilax Shear 2010 S10
       `--Sabacon Simon 1879 (see below for synonymy) S13
            |--*S. paradoxus Simon 1879 G02 [=Sa. paradoxum G02, Scotolemon paradoxus D04]
            |--S. akiyoshiensis Suzuki 1963 [=S. akiyoshiense] S13
            |--S. altomontanus Martens 1983 [=S. altomontanum] S13
            |--S. astoriensis Shear 1975 E90
            |--S. briggsi Shear 1975 E90
            |--S. bryantii (Banks 1898) (see below for synonymy) S13
            |--S. cavicolens (Packard 1884) (see below for synonymy) S13
            |--S. chomolongmae Martens 1972 S13
            |--S. claviger (Menge 1854) [=Nemastoma clavigerum; incl. S. bachofeni Roewer 1939] D06
            |--S. crassipalpis (Koch 1879) G07a, CL00 (see below for synonymy)
            |--S. crassipes S66
            |--S. dentipalpis Suzuki 1949 [=S. dentipalpe] S13
            |--S. dhaulagiri Martens 1972 S13
            |--S. distinctus Suzuki 1974 [=S. distinctum] S13
            |--S. franzi Roewer 1953 S13
            |--S. gonggashan Tsurusaki & Song 1993 TS93
            |--S. imamurai Suzuki 1964 T06
            |--S. iriei Suzuki 1974 S13
            |--S. ishizuchi Suzuki 1974 S13
            |--S. jiriensis Martens 1972 S13
            |--S. makinoi Suzuki 1949 T06
            |    |--S. m. makinoi [incl. S. habei Suzuki 1965] S13
            |    `--S. m. sugimotoi Suzuki & Tsurusaki 1983 S13
            |--S. martensi Tsurusaki & Song 1993 TS93
            |--S. mitchelli Crosby & Bishop 1924 E90
            |--S. occidentalis (Banks 1894) [=Phlegmacera occidentalis] BH09
            |--S. okadai Suzuki 1941 DBM12
            |--S. palpogranulatus Martens 1972 [=S. palpogranulatum] S13
            |--S. pasonianus Luque 1991 (see below for synonymy) S13
            |--S. picosantrum Martens 1983 S13
            |--S. pygmaeus Miyosi 1942 TT05 [=S. pygmaeum TM00]
            |--S. relictus Marten 1972 [=S. relictum] S13
            |--S. satoikioi Miyosi 1942 S13
            |--S. sergeidedicatus Martens 1989 [=S. sergeidedicatum] S13
            |--S. sheari Cokendolpher 1984 E90
            |--S. simoni Dresco 1952 S13
            |--S. siskiyou Shear 1975 E90
            |--S. unicornis Martens 1972 DBM12
            `--S. viscayanus Simon 1881 [=S. viscayanum, S. viscayasanum, S. vizcayanus] S13
                 |--S. v. viscayanus [incl. S. inflatus Dresco 1955 (n. n.)] S13
                 `--S. v. ramblaianus Martens 1983 [=S. viscayanum ramblaianum] S13

Hesperonemastoma packardi (Roewer 1914) [=Nemastoma packardi, N. troglodytes Packard 1877 non Leiobunum troglodytes Wankel 1861] S10

*Hesperonemastoma pallidimaculosum (Goodnight & Goodnight 1945) [=Nemastoma pallidimaculosa, N. pallidimaculosum] S10

Sabacon Simon 1879 [incl. Parasabacon Hansen & Sørensen 1904, Phlegmacera Packard 1884, Tomicomerus Pavesi 1899; Sabaconidae] S13

Sabacon bryantii (Banks 1898) [=Phlegamacera bryantii, S. bryanti (l. c.), *Tomicomerus bryantii, T. bryanti (l. c.); incl. T. bispinosus Pavesi 1899] S13

Sabacon cavicolens (Packard 1884) [=*Phlegmacera cavicolens, P. cavicoleus (l. c.); incl. Nemastoma inops Packard 1884, Hesperonemastoma inops, Sabacon jonesi Goodnight & Goodnight 1942, S. spinosus Weed 1893] S13

Sabacon crassipalpis (Koch 1879) G07a, CL00 [=Nemastoma crassipalpe G07a, *Parasabacon crassipalpis G07a, Sabacon crassipalpe CL00]

Sabacon pasonianus Luque 1991 [=S. picosantrum pasonianum, S. picosantrum asonianum (l. c.), S. picosantrum asoniarum (l. c.)] S13

*Type species of generic name indicated

REFERENCES

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

[CL00] Chemeris, A. N., & D. V. Logunov. 2000. Notes on two species of Sabacon Simon, 1879 from the mountains of South Siberia (Arachnida: Opiliones: Sabaconidae). Arthropoda Selecta 9 (3): 207–213.

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[D06] Dunlop, J. A. 2006. Baltic amber harvestman types (Arachnida: Opiliones: Eupnoi and Dyspnoi). Fossil Record 9 (2): 167–182.

[DBM12] Dunlop, J. A., C. Bartel & P. G. Mitov. 2012. An enigmatic spiny harvestman from Baltic amber. Fossil Record 15 (2): 91–101.

[E90] Edgar, A. L. 1990. Opiliones (Phalangida). In: Dindal, D. L. (ed.) Soil Biology Guide pp. 529–581. John Wiley & Sones: New York.

[GV09] Giribet, G., L. Vogt, A. Pérez González, P. Sharma & A. B. Kury. 2009. A multilocus approach to harvestman (Arachnida: Opiliones) phylogeny with emphasis on biogeography and the systematics of Laniatores. Cladistics 25: 1–30.

[GG42] Goodnight, C. J., & M. L. Goodnight. 1942. New American Phalangida. American Museum Novitates 1164: 1–4.

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

[GG45] Goodnight, C. J., & M. L. Goodnight. 1945. Phalangida from the United States. Journal of the New York Entomological Society 53: 239–245.

[G02] Gruber, J. 2002. Origin and gender of the name Sabacon Simon, 1879 (Opiliones: Palpatores: Ischyropsalidoidea). Newsl. Brit. Arachnol. Soc. 96: 6.

[G07a] Gruber, J. 2007a. Dyspnoi: historical systematic synopsis. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 131–135. Harvard University Press: Cambridge (Massachusetts).

[G07b] Gruber, J. 2007b. Ceratolasmatidae Shear, 1986. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 136–142. Harvard University Press: Cambridge (Massachusetts).

[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. Hesperonemastoma smilax, n. sp., a remarkable new harvestman from a cave in West Virginia, with comments on other reported cave-dwelling Hesperonemastoma species (Opiliones, Ischyropsalidoidea, Sabaconidae). Journal of Cave and Karst Studies 72 (2): 105–110.

[S98] Shultz, J. W. 1998. Phylogeny of the Opiliones (Arachnida): an assessment of the “Cyphopalpatores” concept. Journal of Arachnology 26: 257–272.

[S79] Simon, E. 1879. Descriptions d’Opiliones nouveaux. Annales de la Société Entomologique de Belgique 22 (Comptes-Rendus): lxx–lxxv.

[S66] Suzuki, S. 1966. Four remarkable phalangids from Korea. Annotationes Zoologicae Japonenses 39 (2): 95–106.

[T06] Tsurusaki, N. 2006. Harvestmen of Shimane Prefecture, western Honshu, Japan: an annotated list of species with special reference to geographic variation (Arachnida: Opiliones). In: Board of the Shimane Meeting of the Japanese Society of Biological Education (ed.) Shin Shimane no Seibutsu pp. 153–170. Board of the Shimane Meeting of the Japanese Society of Biological Education: Izuno (Japan).

[TM00] Tsurusaki, N., & H. Minato. 2000. Opiliones of Wakayama Prefecture, Honshu, Japan. Nankiseibutu 42 (1): 3–8.

[TS93] Tsurusaki, N., & D. Song. 1993. Two new species of Sabacon from Sichuan Province, China (Arachnida: Opiliones: Sabaconidae). Zoological Science 10: 155–159.

[TT05] Tsurusaki, N., M. Takanashi, N. Nagase & T. Shimada. 2005. Fauna and biogeography of harvestmen (Arachnida: Opiliones) of the Oki Islands, Japan. Acta Arachnologica 54 (1): 51–63.

Troguloidea

Pair of Dicranolasma soerensenii, copyright Salvatore Canu.

Belongs within: Palpatores.
Contains: Nemastomatidae, Trogulidae.

The Troguloidea is a clade of harvestmen found in Holarctic region, primarily characterised by features of the genitalia. Members of this clade possess two muscles in the penis, and aciniform vaginal glands and outer circular muscles in the ovipositor (Giribet & Kury 2007).

<==Troguloidea [Eupagosterni, Troguloidae]
    |  i. s.: Eotrogulus [Eotrogulidae] K13
    |           `--E. fayoli Thevenin 1901 D07
    |         Nemastomoides Thevenin 1901 D07 [incl. Protopilio D07; Nemastomoididae K13]
    |           |--N. elaveris Thevenin 1901 D07
    |           `--N. longipes (Petrunkevitch 1913) [=Protopilio longipes] D07
    |--Nipponopsalis Martens & Suzuki 1966 M76 [Nipponopsalididae GE02]
    |    |--*N. abei (Sato & Suzuki 1939) S73 [=Ischyropsalis abei M76]
    |    |    |--N. a. abei S73
    |    |    `--N. a. longipes Suzuki 1973 S73
    |    |--N. coreana (Suzuki 1966) [=Ischyropsalis coreana] S13
    |    `--N. yezoensis (Suzuki 1958) [=Ischyropsalis yezoensis] S13
    `--+--+--Nemastomatidae GE02
       |  `--Ameticos Garwood, Dunlop et al. 2011 GS14, GD11
       |       `--*A. scolos Garwood, Dunlop et al. 2011 GD11
       `--+--Trogulidae GE02
          `--Dicranolasma Sørensen 1873 GE02, R23 (see below for synonymy)
               |--*D. scabrum (Herbst 1799) [=Opilio scaber; incl. D. schmidti Hadži 1928] S13
               |--D. apuanum Marcellino 1971 G98
               |--D. cretaeum Gruber 1998 G98
               |--D. cristatum Thorell 1876 G98
               |--D. giljarovi Šilhavý 1966 G98
               |--D. hirtum Loman 1894 (see below for synonymy) S13
               |--D. hoberlandti Šilhavý 1956 G98
               |--D. kurdistanum Staręga 1970 G98
               |--D. mladeni Karaman 1990 G98
               |--D. opilionoides (Koch 1867) (see below for synonymy) S13
               |--D. pauper Dahl 1903 G98
               |--D. ponticum Gruber 1998 G98
               |--D. ressli Gruber 1998 G98
               |--D. soerensenii Thorell 1876 S13 (see below for synonymy)
               |--D. thracium Staręga 1976 G98
               `--D. verhoeffi Dahl 1903 G98

Dicranolasma Sørensen 1873 GE02, R23 [incl. Amopaum Sørensen 1873 R23; Dicranolasmatidae, Dicranolasmatinae, Dicranolasmatini, Dicranolasmidae]

Dicranolasma hirtum Loman 1894 [incl. D. diomedeum Kulczyński 1907, D. diomedaeum (l. c.), D. napoli Goodnight & Goodnight 1944] S13

Dicranolasma opilionoides (Koch 1867) [=Trogulus opilionoides; incl. D. kettyi Marcellino 1968, D. kettyae, *Amopaum spinipalpe Sørensen 1873] S13

Dicranolasma soerensenii Thorell 1876 S13 [=Amopaum soerenseni D04, D. soerenseni S13; incl. D. latifrons Simon 1879 D04, D. wiehlei Kraus 1959 D04]

*Type species of generic name indicated

REFERENCES

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[D07] Dunlop, J. A. 2007. Paleontology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 247–265. Harvard University Press: Cambridge (Massachusetts).

[GD11] Garwood, R. J., J. A. Dunlop, G. Giribet & M. D. Sutton. 2011. Anatomically modern Carboniferous harvestmen demonstrate early cladogenesis and stasis in Opiliones. Nature Communications 2 (444): 1–7.

[GS14] Garwood, R. J., P. P. Sharma, J. A. Dunlop & G. Giribet. 2014. A Paleozoic stem group to mite harvestmen revealed through integration of phylogenetics and development. Current Biology 24 (9): 1017–1023.

[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.

Giribet, G., & A. B. Kury. 2007. Phylogeny and biogeography. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 62–87. Harvard University Press: Cambridge (Massachusetts).

[G98] Gruber, J. 1998. Beiträge zur Systematik der Gattung Dicranolasma (Arachnida: Opiliones, Dicranolasmatidae). I. Dicranolasma thracium Staręga und verwandte Formen aus Südosteuropa und Südwestasien. Annalen des Naturhistorischen Museum in Wien. Serie B. Botanik und Zoologie 100: 489–537.

[K13] Kury, A. B. 2013. Order Opiliones Sundevall, 1833. Zootaxa 3703: 27–33.

[M76] Martens, J. 1976. Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida: Opiliones). Entomologica Germanica 3 (1–2): 51–68.

[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.

[S73] Suzuki, S. 1973. Opiliones from the South-west Islands, Japan. Journal of Science of the Hiroshima University, Series B, Division 1, 24: 205–279.

Palpatores

Acropsopilio neozelandiae, copyright Stephen Thorpe.

Belongs within: Opiliones.
Contains: Ischyropsalidoidea, Troguloidea, Phalangioidea.

The Palpatores, the long-legged harvestmen, are a diverse group of harvestmen including many of the most familiar members of the order. Members of this clade have ozopores flush with the body surface (not raised on ozophores) and simple, single claws on the legs; most (but not all) also have particularly elongate legs. Basal members of the Palpatores with small bodies and greatly enlarged eye mounds were classified until recently in a family Caddidae, but phylogenetic analyses have recently indicated that this group is para- or polyphyletic, and may represent the plesiomorphic appearance of the clade as a whole (Groh & Giribet 2015).

Palpatores [Apagosterni, Caddidae, Caddoidea, Plagiostethi]
    |--Dyspnoi [Nemastomini, Nemastomoidae, Trogulini] GG15
    |    |  i. s.: Echinopustulus samuelnelsoni Dunlop 2004 D07
    |    |--Acropsopilionidae [Acropsopilioninae, Acropsopilionoidea] GFB14
    |    |    |--Caddella Hirst 1925 SC06 [=Cadella (l. c.) K61; incl. Oonopsopilio Lawrence 1931 SC03]
    |    |    |    |--*C. capensis Hirst 1925 S75 [=C. (Oonopsopilio) capensis K61]
    |    |    |    |--C. africana (Lawrence 1931) [=*Oonopsopilio africanus] S75
    |    |    |    |--C. croeseri Staręga 1988 S92
    |    |    |    |--C. haddadi Lotz 2011 L11
    |    |    |    |--C. jocquei Staręga 2008 L11
    |    |    |    `--C. spatulipilis Lawrence 1934 S92 (see below for synonymy)
    |    |    |--Austropsopilio Forster 1955 [incl. Tasmanopilio Hickman 1957] GG15
    |    |    |    |--*A. novaehollandiae Forster 1955 SC03
    |    |    |    |--A. altus Cantrell 1980 C80
    |    |    |    |--A. cygneus Hickman 1957 H57
    |    |    |    |--A. fuscus (Hickman 1957) GG15 [=*Tasmanopilio fuscus H57]
    |    |    |    |--A. inermis Cantrell 1980 C80
    |    |    |    |--A. megalops (Hickman 1957) [=Tasmanopilio megalops] GG15
    |    |    |    `--A. sudamericanus Shultz & Cekalovic 2003 SC03
    |    |    `--Acropsopilio Silvestri 1904 SC06 [incl. Zeopsopilio Forster 1948 SC03]
    |    |         |--*A. chilensis Silvestri 1904 S75 (see below for synonymy)
    |    |         |--A. australicus Cantrell 1980 C80
    |    |         |--A. boopis (Crosby 1904) [=Caddo boopis] S75
    |    |         |--A. chomulae (Goodnight & Goodnight 1948) [=Caddo chomulae, A. chommulae (l. c.)] S75
    |    |         |--A. neozelandiae (Forster 1948) GFB14 [=*Zeopsopilio neozelandiae F48]
    |    |         |--A. talishensis Morin 1937 S04
    |    |         `--A. venezuelensis González-Sponga 1992 GG15
    |    `--+--Ischyropsalidoidea GG15
    |       `--Troguloidea GG15
    `--Eupnoi [Phalangioidae] GG15
         |  i. s.: Brigantibunum Dunlop & Anderson 2005 DA05
         |           `--*B. listoni Dunlop & Anderson 2005 DA05
         |         Daohugopilio Huang, Selden & Dunlop 2009 HSD09
         |           `--*D. sheari Huang, Selden & Dunlop 2009 HSD09
         |--Mesobunus Huang, Selden & Dunlop 2009 GS14, HSD09
         |    |--*M. martensi Huang, Selden & Dunlop 2009 HSD09
         |    `--M. dunlopi Giribet, Tourinho et al. 2012 GT12
         `--+--Phalangioidea GS14
            `--+--Macrogyion Garwood, Dunlop et al. 2011 GS14, GD11
               |    `--*M. cronus Garwood, Dunlop et al. 2011 GD11
               `--Caddo Banks 1892 GS14, S75 [Caddinae, Caddini, Caddoinae]
                    |--*C. agilis Banks 1892 [incl. C. glaucopis Crosby 1904] S75
                    |--C. dentipalpus (Koch & Berendt 1854) [=Platybunus dentipalpus] S02
                    `--C. pepperella Shear 1975 S75

Nomen nudum: Caddo medama Suzuki 1958 GG15

*Acropsopilio chilensis Silvestri 1904 S75 [incl. A. normae Cekalovic 1974 GG15, A. chilensis var. ogloblini Canals 1932 S75]

Caddella spatulipilis Lawrence 1934 S92 [=Cadella spatulipilis L34, Caddella spatulipalpis (l. c.) S75; incl. Cadella spatulipilis var. caledonica Lawrence 1934 L34]

*Type species of generic name indicated

REFERENCES

[C80] Cantrell, B. K. 1980. Additional Australian harvestmen (Arachnida: Opiliones). Journal of the Australian Entomological Society 19: 241–253.

[D07] Dunlop, J. A. 2007. Paleontology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 247–265. Harvard University Press: Cambridge (Massachusetts).

[DA05] Dunlop, J. A., & L. I. Anderson. 2005. A fossil harvestman (Arachnida, Opiliones) from the Mississippian of East Kirkton, Scotland. Journal of Arachnology 33: 482–489.

[F48] Forster, R. R. 1948. A new genus and species of the family Acropsopilionidae (Opiliones) from New Zealand. Transactions of the Royal Society of New Zealand 77 (1): 139–141.

[GD11] Garwood, R. J., J. A. Dunlop, G. Giribet & M. D. Sutton. 2011. Anatomically modern Carboniferous harvestmen demonstrate early cladogenesis and stasis in Opiliones. Nature Communications 2 (444): 1–7.

[GS14] Garwood, R. J., P. P. Sharma, J. A. Dunlop & G. Giribet. 2014. A Paleozoic stem group to mite harvestmen revealed through integration of phylogenetics and development. Current Biology 24 (9): 1017–1023.

[GFB14] Giribet, G., R. Fernández & S. L. Boyer. 2014. On four poorly known harvestmen from New Zealand (Arachnida: Opiliones: Cyphophthalmi: Eupnoi: Dyspnoi: Laniatores). New Zealand Journal of Zoology 41 (4): 223–233.

[GT12] Giribet, G., A. L. Tourinho, C. Shih & D. Ren. 2012. An exquisitely preserved harvestman (Arthropoda, Arachnida, Opiliones) from the Middle Jurassic of China. Organisms, Diversity & Evolution 12: 51–66.

[GG15] Groh, S., & G. Giribet. 2015. Polyphyly of Caddoidea, reinstatement of the family Acropsopilionidae in Dyspnoi, and a revised classification system of Palpatores (Arachnida, Opiliones). Cladistics 31 (3): 277–290.

[H57] Hickman, V. V. 1957. Some Tasmanian harvestmen of the sub-order Palpatores. Papers and Proceedings of the Royal Society of Tasmania 91: 65–79.

[HSD09] Huang, D., P. A. Selden & J. A. Dunlop. 2009. Harvestmen (Arachnida: Opiliones) from the Middle Jurassic of China. Naturwissenschaften 96 (8): 955–962.

[K61] Kauri, H. 1961. Opiliones. In: Hanström, B., P. Brinck & G. Rudebeck. South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 9–197. Almqvist & Wiksell: Uppsala.

[L34] Lawrence, R. F. 1934. New South African Opiliones. Annals of the South African Museum 30 (4): 549–586.

[L11] Lotz, L. N. 2011. Three new harvestmen species from southern Africa (Arachnida: Opiliones: Caddidae, Neopilionidae, Assamiidae). Journal of Afrotropical Zoology 7: 3–8.

[S75] Shear, W. A. 1975. The opilionid family Caddidae in North America, with notes on species from other regions (Opiliones, Palpatores, Caddoidea). Journal of Arachnology 2: 65–88.

[SC03] Shultz, J. W., & T. Cekalovic. 2003. First species of Austropsopilio (Opiliones, Caddoidea, Caddidae) from South America. Journal of Arachnology 31: 20–27.

[SC06] Shultz, J. W., & T. Cekalovic. 2006. First species of Hesperopilio (Opiliones, Caddoidea) from South America. Journal of Arachnology 34 (1): 46–50.

[S04] Snegovaya, N. Y. 2004. Preliminary notes on the harvestman fauna (Opiliones) of Azerbaijan. Arthropoda Selecta Special Issue 1: 307–318.

[S92] Staręga, W. 1992. An annotated check-list of Afrotropical harvestmen, excluding the Phalangiidae (Opiliones). Annals of the Natal Museum 33 (2): 271–336.

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

Nemastoma

Nemastoma sp., from Axel L. Schönhofer.

Belongs within: Nemastomatidae.

Nemastoma is a widespread genus of small harvestmen in Europe.

Characters (from Martens 1978): Male genital morphology with truncus penis long and narrow, usually almost at right angles to the dilated base; Glans in a straight extension of trunk, scarcely wider than latter; dorsoventrally flattened. Glans symmetrical or slightly asymmetrical, laterally with one to eight strong, basad-directed hooks, exceptionally missing. Scutum primarily a scutum magnum, old and strongly sclerotized individuals with scutum compositum (abdominal tergites VI, VII and rarely VIII fused with scutum magnum). Smooth-shiny to roughly granulated, without dorsal spines; deep black, carapace often with gold or silver markings.

<==Nemastoma Koch 1836 [incl. Lugubrostoma Kratochvíl 1958, Stridulostoma Hadži 1973] S13
    |--*N. bimaculatum (Fabricius 1775) M06 (see below for synonymy)
    |--N. bidentatum Roewer 1914 M06 [=N. (Lugubrostoma) bidentatum N05]
    |    |--N. b. bidentatum M06
    |    |--N. b. pluridentatum Hadži 1973 (see below for synonymy) S13
    |    |--N. b. relictum Gruber & Martens 1968 M06
    |    `--N. b. sparsum Gruber & Martens 1968 M06
    |--N. daciscum Koch 1869 (n. d.) S13
    |--N. dentigerum Canestrini 1873 M06
    |--‘Crosbycus’ goodnighti Roewer 1951 (n. d.) S10, S13 [=C. goodnightorum (l. c.) S13]
    |--N. incertum Koch & Berendt 1854 D07
    |--N. lilliputanum (Lucas 1849) [=Goniosoma lilliputanum] S13
    |--N. lugubre (Müller 1776) M06 (see below for synonymy)
    |--N. rude Simon 1881 D04
    |--N. schuelleri Gruber & Martens 1968 M06
    |--N. superbum Koch 1869 M06
    |--N. transsylvanicum Gruber & Martens 1968 M06
    `--N. triste (Koch 1835) M06 (see below for synonymy)

Nemastoma bidentatum pluridentatum Hadži 1973 [=N. (Lugubrostoma) triste pluridentatum; incl. N. (*Stridulostoma) seliskari Hadži 1973] S13

*Nemastoma bimaculatum (Fabricius 1775) M06 [=Phalangium bimaculatum M06, N. lugubre bimaculatum D04, Opilio bimaculatus D04]

Nemastoma lugubre (Müller 1776) M06 [=Phalangium lugubre D04, Histricostoma lugubre D04, Nemastoma (*Lugubrostoma) lugubre S13; incl. N. lugubre helvetica Lessert 1917 S13, N. lugubre var. striatum Morin 1934 S13]

Nemastoma triste (Koch 1835) M06 [=Phalangium triste R23, N. (Lugubrostoma) triste S13; incl. N. kollari Doleschall 1852 R23, N. lugubre unicolor Roewer 1914 S13]

*Type species of generic name indicated

REFERENCES

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[D07] Dunlop, J. A. 2007. Paleontology. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 247–265. Harvard University Press: Cambridge (Massachusetts).

Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

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

[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. Hesperonemastoma smilax, n. sp., a remarkable new harvestman from a cave in West Virginia, with comments on other reported cave-dwelling Hesperonemastoma species (Opiliones, Ischyropsalidoidea, Sabaconidae). Journal of Cave and Karst Studies 72 (2): 105–110.

Paranemastoma

Paranemastoma sp., from Axel L. Schönhofer.

Belongs within: Nemastomatidae.

Paranemastoma is a genus including some of the largest nemastomatid harvestmen, found in central and eastern Europe.

Characters (from Martens 1978): Male genital morphology with truncus penis long and extremely narrow, base widened, markedly separated from the rest of the trunk, but not angled; deeply divided, the two muscular strands concentrated in the two halves, glans in a straight extension of truncus, scarcely wider than latter, differentiated only by stronger sclerotization and tiny sensory bristles. Tip of glans divided into two branches, the right branch (in ventral view) narrower and slightly overarching the shorter and wider left branch. Scutum with para-median spine pairs, most developed on abdominal area II, weaker in female. Body always deep black, usually with gold or silver spotting.

<==Paranemastoma Redikorzev 1936 [incl. Buresiolla Kratochvíl 1958, Dromedostoma Kratochvíl 1958] S13
    |--*P. superbum Redikorzev 1936 M11 (see below for synonymy)
    |--P. aeginum (Roewer 1951) [=Nemastoma aeginum] S13
    |--P. amseli (Roewer 1951) [=Nemastoma amseli] S13
    |--P. amuelleri (Roewer 1951) [=Nemastoma amuelleri] S13
    |--P. ancae Avram 1973 S13
    |--P. armatum (Kulczyński 1909) [=Nemastoma quadripunctatum armatum] N05
    |--P. aurigerum (Roewer 1951) [=Nemastoma aurigerum] S13
    |    |--P. a. aurigerum S13
    |    |--P. a. joannae Staręga 1976 S13
    |    `--P. a. ryla (Roewer 1951) [=Nemastoma ryla, N. (Dromedostoma) ryla] S13
    |--P. aurosum (Koch 1869) N05 [=Nemastoma quadripunctatum aurosum N05, N. quadripunctatum var. aurosa D04]
    |--P. beroni Mitov 2011 M11
    |--P. bicuspidatum (Koch 1835) (see below for synonymy) S13
    |--P. bolei (Hadži 1973) [=Nemastoma (Dromedostoma) bolei] S13
    |--P. brevipalpatum (Roewer 1951) [=Nemastoma brevipalpatum] S13
    |--P. bureschi (Roewer 1926) (see below for synonymy) S13
    |--P. caporiaccoi (Roewer 1951) [=Nemastoma caporiaccoi] S13
    |--P. carneluttii (Hadži 1973) [=Nemastoma carneluttii] S13
    |--P. corcyraeum (Roewer 1917) [=Nemastoma quadripunctatum corcyraeum] S13
    |--P. emigratum (Roewer 1959) [=Nemastoma emigratum] S13
    |--P. ferkeri (Roewer 1951) [=Nemastoma ferkeri] S13
    |--P. filipes (Roewer 1917) [=Nemastoma quadripunctatum var. filipes, N. (Dromedostoma) filipes] S13
    |--P. gostivarense (Hadži 1973) [=Nemastoma gostivarense] S13
    |--‘Crosbycus’ graecus Giltay 1932 (see below for synonymy) S13
    |--P. ikarium (Roewer 1951) [=Nemastoma ikarium] S13
    |--P. ios (Roewer 1917) [=Nemastoma quadripunctatum ios] S13
    |--P. iranicum Martens 2006 M06
    |--P. kaestneri (Roewer 1951) [=Nemastoma kaestneri] S13
    |--P. kalischevskyi (Roewer 1951) (see below for synonymy) S13
    |--P. kochii (Nowicki 1870) (see below for synonymy) S13
    |--P. longipalpatum (Roewer 1951) [=Nemastoma longipalpatum] S13
    |--P. longipes (Schenkel 1947) N05 (see below for synonymy)
    |--P. macedonicum (Hadži 1973) [=Nemastoma macedonicum] S13
    |--P. machadoi (Roewer 1951) [=Nemastoma machadoi] S13
    |--P. mackenseni (Roewer 1923) [=Nemastoma mackenseni] S13
    |--P. multisignatum (Hadži 1973) [=Nemastoma (Dromedostoma) multisignatum] S13
    |--P. ‘nigrum’ (Hadži 1973) [=Nemastoma nigrum non N. paspalevi var. nigrum Kratochvíl 1958] S13
    |--P. perfugium (Roewer 1951) [=Nemastoma perfugium] S13
    |--P. quadripunctatum (Perty 1833) M06 (see below for synonymy)
    |--P. radewi (Roewer 1926) (see below for synonymy) S13
    |--P. roeweri Starega 1978 [=Nemastoma redikorzevi Roewer 1951 non Charitonov 1946, P. redikorzevi] S13
    |--P. santorinum (Roewer 1951) [=Nemastoma santorinum] S13
    |--P. senussium (Roewer 1951) [=Nemastoma senussium] S13
    |--P. sillii (Herman 1871) S13
    |    |--P. s. sillii S13 (see below for synonymy)
    |    |--P. ‘gigas’ montenegrinum Nosek 1904 S13 [=Nemastoma gigas montenegrinum S13, N. g. montenegrina (l. c.) R23]
    |    `--P. s. monticola Babalean 2011 S13
    |--P. sketi (Hadži 1973) [=Nemastoma (Dromedostoma) sketi] S13
    |--P. spinosulum (Koch 1869) [=Nemastoma spinosulum, N. spinulosum (l. c.)] S13
    |--P. thessalum (Simon 1885) [=Nemastoma quadripunctatum thessalum, P. teshalum (l. c.)] S13
    |--P. titaniacum (Roewer 1914) S13 (see below for synonymy)
    |--P. umbo (Roewer 1951) [=Nemastoma umbo] M06
    `--P. werneri (Kulczyński 1903) [=Nemastoma quadripunctatum werneri, P. quadripunctatum werneri] S13

‘Crosbycus’ graecus Giltay 1932 [incl. Nemastoma simplex Giltay 1932 non N. bacilliferum simplex Simon 1913, Paranemastoma simplex] S13

Paranemastoma bicuspidatum (Koch 1835) [=Phalangium bicuspidatum, Nemastoma quadripunctatum bicuspidatum; incl. N. reimoseri Roewer 1951] S13

Paranemastoma bureschi (Roewer 1926) [=Nemastoma bureschi, Crosbycus bureschi, Paranemastoma (*Buresiolla) bureschi; incl. N. tunetanum Roewer 1951, B. tunetanum, B. tunetorum (l. c.)] S13

Paranemastoma kalischevskyi (Roewer 1951) [=Nemastoma kalischevskyi; incl. N. charitonovi Mcheidze 1952, Paranemastoma charitonovi, N. suchumium Roewer 1951, P. suchumium] S13

Paranemastoma kochii (Nowicki 1870) [=Nemastoma quadripunctatum kochii; incl. N. hungaricum Roewer 1951, N. nicaeum Roewer 1951] S13

Paranemastoma longipes (Schenkel 1947) N05 [=Nemastoma gigas longipes M00, N. (Dromedostoma) longipes N05; incl. N. tenue Hadži 1973 S13]

Paranemastoma quadripunctatum (Perty 1833) M06 [=Phalangium quadripunctatum D04, Nemastoma quadripunctatum N05; incl. N. alfkeni Roewer 1951 D04, Paranemastoma alfeki D04, N. armeniacum Roewer 1951 S13, P. armeniacum M06, N. austriacum Roewer 1951 D04, N. quadripunctatum carniolicum Hadži 1973 D04, N. chaucicum Roewer 1951 D04, Paranemastoma chaucicum D04, N. emonense Hadži 1973 D04, Phalangium flavimanum Koch 1835 K35, Nemastoma flavimanum D04, Opilio flavimanum D04, N. gertschi Roewer 1951 D04, Paranemastoma gertschi D04, Crosbycus lengersdorfi Roewer 1951 D04, Nemastoma quadripunctatum var. lessinensis Caporiacco 1940 S13, N. mediosignatum Hadži 1973 D04, N. quadripunctatum var. moesiacum Roewer 1917 S13, Paranemastoma moesiacum N05, N. nervosum Roewer 1923 S13, N. slovenicum Hadži 1973 D04, N. (Dromedostoma) strasseri Roewer 1951 N05, N. triglavense Hadži 1973 D04]

Paranemastoma radewi (Roewer 1926) [=Nemastoma radewi S13, N. (*Dromedostoma) radewi S13, N. radevi (l. c.), Paranemastoma radevi; incl. N. (D.) atanasovi Kratochvíl 1958, P. atanasovi, N. (D.) atanasovi balcanica Kratochvíl 1958, N. insulare Roewer 1951, N. (D.) markovi Kratochvíl 1958, N. (D.) megarae Hadži 1973, N. (D.) paspalevi var. nigrum Kratochvíl 1958, N. (D.) paspalevi Kratochvíl 1958, Paranemastoma paspalevi, N. polonicum Roewer 1951, N. riparium Roewer 1951, n. schenkeli Roewer 1951] S13

Paranemastoma sillii sillii (Herman 1871) S13 [=Nemastoma quadripunctatum sillii S13; incl. N. bishopi Roewer 1951 S13, N. bulgaricum Roewer 1951 S13, N. carpathicum Roewer 1951 S13, Crosbycus cavernae Roewer 1951 S13, N. gigas Sørensen in Lendl 1894 S13, N. sillii gigas, Paranemastoma gigas S13, N. lineatum Sørensen in Lendl 1894 S13, N. quadripunctatum lineatum, N. mohilevum Roewer 1951 S13, N. politum Roewer 1951 S13, N. romanium Roewer 1951 S13, N. rumanicum (l. c.) S13, N. rumelium Roewer 1951 S13, N. wiehlei Roewer 1951 S13, N. zilchi Roewer 1951 S13]

*Paranemastoma superbum Redikorzev 1936 non Nemastoma superbum Koch 1869 M11 [=P. supersum Roewer 1951 M06, Nemastoma supersum M06; incl. N. superbum bacuriana Mcheidze 1959 M06, N. superbum bacurianum M06]

Paranemastoma titaniacum (Roewer 1914) S13 [=Nemastoma quadripunctatum titaniacum N05, N. (Dromedostoma) titaniacum S13; incl. N. acrospinosum Roewer 1951 S13, N. bimaculosum Roewer 1951 S13, Paranemastoma bimaculosum N05, N. bipunctatum Hadži 1973 S13, N. (D.) grabrovicae Hadži 1973 N05, N. gabbrovicae (l. c.) N05, N. acrospinosum pretneri Hadži 1973 S13]

*Type species of generic name indicated

REFERENCES

[D04] Delfosse, E. 2004. Catalogue préliminaire des Opilions de France métropolitaine (Arachnida Opiliones). Bulletin de Phyllie 20: 34–58.

[K35] Koch, C. L. 1835. Arachniden. In: Herrich-Schäffer, G. A. W. Deutschlands Insecten. Band 34, Hefte 128–133. F. Pustet: Regensburg.

Martens, J. 1978. Spinnentiere, Arachnida: Weberknechte, Opiliones. Gustav Fischer Verlag: Jena.

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

[M00] Mitov, P. 2000. Contribution to the knowledge of the harvestmen (Arachnida: Opiliones) of Albania. Ekológia 19 (Suppl. 3): 159–169.

[M11] Mitov, P. G. 2011. A new anophthalmous species of Paranemastoma from Bulgaria (Opiliones: Nemastomatidae). Journal of Arachnology 39 (2): 303–319.

[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.

Nemastomatidae

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]
    |--+--Halitherses Giribet & Dunlop 2005 GS14, GD05 [Halithersidae]
    |  |    `--*H. grimaldii Giribet & Dunlop 2005 GD05
    |  `--Ortholasmatinae S10
    |       |--Martensolasma Shear 2006 S13
    |       |    `--*M. jocheni Shear 2006 S10
    |       |--+--Dendrolasma Banks 1894 S13
    |       |  |    |--*D. mirabile Banks 1894 [=D. mirabilis] S13
    |       |  |    `--D. dentipalpe Shear & Gruber 1983 S10
    |       |  `--Cladolasma Suzuki 1963 S13
    |       |       |--*C. parvulum Suzuki 1963 [=C. parvula, Dendrolasma parvula, D. parvulum] S13
    |       |       `--C. angka (Schwendinger & Gruber 1992) [=Dendrolasma angka] S10
    |       `--+--Trilasma Goodnight & Goodnight 1942 [incl. Ruaxphilos Goodnight & Goodnight 1945] S13
    |          |    |--*T. bolivari Goodnight & Goodnight 1942 S13 (see below for synonymy)
    |          |    |--T. chipinquensis Shear 2010 S10
    |          |    |--T. hidalgo Shear 2010 S10
    |          |    |--T. petersprousei Shear 2010 S10
    |          |    |--T. ranchonuevo Shear 2010 S10
    |          |    |--T. sbordonii Šilhavý 1973 S13 [=Ortholasma sbordonii S10]
    |          |    |--T. tempestado Shear 2010 S10
    |          |    |--T. trispinosum Shear 2010 S10
    |          |    `--T. tropicum Shear 2010 S10
    |          `--Ortholasma Banks 1894 S13
    |               |--*O. rugosum Banks 1894 [=O. rugosa] S10
    |               |--O. colossus Shear 2010 S10
    |               |--O. coronadense Cockerell 1916 [=O. coronadensis; incl. O. setulipes Shear & Gruber 1983] S13
    |               |--O. levipes Shear & Gruber 1983 S10
    |               `--O. pictipes Banks 1911 R23
    `--Nemastomatinae [Giljaroviinae, Mitostominae] G07
         |--Mitostoma S13
         `--+--+--Paranemastoma S13
            |  `--+--Carinostoma Kratochvíl 1958 S13
            |     |    |--*C. carinatum (Roewer 1914) [=Nemastoma carinatum, Mitostoma (Carinostoma) carinatum] S13
            |     |    |--C. elegans (Sørensen 1894) (see below for synonymy) S13
            |     |    `--C. ornatum (Hadži 1940) [=Nemastoma ornatum, Mitostoma (Mitostoma) ornatum] S13
            |     `--Histricostoma Kratochvíl 1958 [incl. Cretostoma Kratochvíl 1958] S13
            |          |--*H. drenskii Kratochvíl 1958 M06
            |          |--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 2012 S13
            |          |--H. mitovi Snegovaya & Marusik 2012 S13
            |          `--H. tuberculatum (Koch & Berendt 1854) [=Nemastoma tuberculatum] S02
            `--+--+--Saccarella Schönhofer & Martens 2012 S13
               |  |    `--*S. schilleri Schönhofer & Martens 2012 S13
               |  |--Nemastomella S13
               |  |--Starengovia Snegovaya 2010 S13
               |  |    `--*S. kirgizica Snegovaya 2010 [=S. kirgisica] S13
               |  `--+--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 1958 S13
               |          |--*C. centetes (Simon 1881) (see below for synonymy) S13
               |          |--C. dubium (Mello-Leitao 1936) GE02
               |          |--C. juberthiei Martens 2011 S13
               |          |--C. scabriculum (Simon 1879) [=Nemastoma scabriculum] S13
               |          `--C. ventalloi (Mello-Leitao 1936) [=Nemastoma ventalloi] S13
               `--+--Pyza Staręga 1976 S13
                  |    |--*P. bosnica (Roewer 1917) (see below for synonymy) S13
                  |    |--P. anatolica (Roewer 1959) [=Crosbycus anatolicus] S13
                  |    |--P. navarrense (Roewer 1951) [=N. navarrense; incl. N. lindbergi Roewr 1959] S13
                  |    `--P. taurica Gruber 1979 S13
                  |--Hadzinia Roewer 1966 S13, N05
                  |    `--*H. karamani (Hadži 1940) (see below for synonymy) S13
                  |--+--Vestiferum Martens 2006 S13
                  |  |    |--*V. funebre (Redikorzev 1936) (see below for synonymy) M06
                  |  |    `--V. alatum Martens 2006 M06
                  |  `--Mediostoma S13
                  `--+--Nemastoma S13
                     |--Caucnemastoma Martens 2006 S13
                     |    |--*C. golovatchi Martens 2006 S13
                     |    `--C. martensi Snegovaya in Snegovaya & Chumachenko 2011 S13
                     |--Nemaspela Šilhavý 1966 S13
                     |    |--*N. sokolovi (Ljovushkin & Starobogatov 1963) [=Buresiolla sokolovi] M06
                     |    |--N. abchasica (Ljovushkin & Starobogatov 1963) [=Buresiolla abchasica] M06
                     |    |--N. birsteini Ljovuschkin 1972 S13
                     |    |--N. caeca (Grese 1911) [=Nemastoma caecum, Buresiolla caecum, Nemaspela coeca (l. c.)] S13
                     |    |--N. femorecurvata Martens 2006 M06
                     |    |--N. kovali Chemeris 2009 S13
                     |    `--N. taurica (Lebedinski 1914) [=Phalangodes taurica] S13
                     `--Giljarovia S13

Carinostoma elegans (Sørensen 1894) [=Nemastoma elegans, Mitostoma (Carinostoma) elegans; incl. N. elegans var. batorligetiense Szalay 1952, Carinostoma elegans batorligetiense] S13

*Centetostoma centetes (Simon 1881) [=Nemastoma centetes, Mitostoma centetes, Nemastomella centetes] S13

*Hadzinia karamani (Hadži 1940) [=Nemastoma karamani, Buresiolla karamani, Hadzinia caramani (l. c.)] S13

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 1942 S13 [=Ortholasma (*Trilasma) bolivari S10; incl. *Ruaxphilos petrunkevitchou Goodnight & Goodnight 1945 S13]

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

*Type species of generic name indicated

REFERENCES

[GS14] Garwood, R. J., P. P. Sharma, J. A. Dunlop & G. Giribet. 2014. A Paleozoic stem group to mite harvestmen revealed through integration of phylogenetics and development. Current Biology 24 (9): 1017–1023.

[GD05] Giribet, G., & J. A. Dunlop. 2005. First identifiable Mesozoic harvestman (Opiliones: Dyspnoi) from Cretaceous Burmese amber. Proceedings of the Royal Society of London Series B—Biological Sciences 272: 1007–1013.

[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).

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

[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.