Hadzinia Šilhavý, 1966a

SCHÖNHOFER, AXEL L., 2013, Axel L. Schönhofer, Zootaxa 3679 (1), pp. 1-68 : 29-48

Genus Hadzinia Šilhavý, 1966a (1 species)

Hadzinia Šilhavý, 1966a: 71 ; type species Nemastoma karamani Hadži, 1940 by monotypy and original designation.

Remarks. Novak (2005: 313) erroneously gave Roewer as describing author.

Etymology. Gender feminine (not specified, inferred from ending; ICZN 30.2.4). In honour of the Yugoslavian zoologist J. Hadži, who described the type species.

karamani ( Hadži, 1940) ; loc. typ.: SW Bosnia, Drvar, cave Resanovačka špilja Ledenica (clarification by Novak 2005: 313).

Nemastoma karamani Hadži, 1940: 2 .

Buresiolla karamani — Kratochvíl 1958: 532.

Hadzinia karamani — Šilhavý 1966a: 71; Novak 2005: 313.

Hadzinia caramani — Šilhavý 1966a: 71 (misspelling).

Genus Histricostoma Kratochvíl, 1958 (8 species)

Histricostoma (Histricostoma) Kratochvíl, 1958: 529 ; type species Histricostoma drenskii Kratochvíl, 1958 by original designation.

Centetostoma (Cretostoma) Kratochvíl, 1958: 530 ; type species Cretostoma creticum Roewer, 1927 by monotypy and original designation;—syn. Martens 1978: 129.

Histricostoma — Gruber 1976: 799.

Remarks. Gruber (1976) rejected Kratochvíl’s subgenera and restricted Histricostoma to include argenteolunulatum , dentipalpe and drenskii . By including creticum, Martens (1978) synonymised the monotypic Cretostoma (see creticum ).

Etymology. Neuter; probably derived from Latin “histrio” (actor) combined with „stoma“ in reference to Nemastoma . Maybe derived from “hystrix” (porcupine). The origin of this name remains uncertain.

anatolicum ( Roewer, 1962) ; loc. typ.: Turkey, Western Taurus, two caves 20 km NW Dag (the small one referred to as “piccola”, the large one as “ In Dag ”, Turkish for “ Dag Cave ”). Dag is on the road from Antalya to Isparta, at the border of both provinces. Remarks. Brignoli reported details about the type locality to Gruber, who made the information available for this catalogue .

Nemastoma anatolicum Roewer, 1962: 15 .

Histricostoma anatolicum comb. n.: The syntype series (SMF 13903) comprises females and juveniles only. Females are very characteristic in exhibiting transverse rows of anvil-shaped tubercles and pairs of very long spines which terminate into one or two smaller spines. According to Gruber (pers. comm.) the leg microsculpture is similar to that of H. creticum and the presence of similar Histricostoma from Anatolia is confirmed. Until males are available, a tentative placement in Histricostoma is advised. Staręga (1973: 132) mentioning anatolicum close to Mediostoma cypricum , which should be considered when revising these species.

argenteolunulatum ( Canestrini, 1875) ; loc. typ.: Italy, Calabria.

Nemastoma dentipalpe var. argenteo-lunulata Canestrini, 1875: 4.

Nemastoma argenteolunulatum — Roewer 1923: 668.

Carinostoma argenteolunulatum — Šilhavý 1969: 496.

Histricostoma argenteolunulatum — Gruber 1976: 799.

caucasicum ( Redikortsev, 1936) ; loc. typ.: Georgia, Adsharia, Suchumi Distr., Ažary and Lata.

Nemastoma caucasicum Redikortsev, 1936: 54 .

Histricostoma (Histricostoma) caucasicum — Staręga 1966a: 394.

Histricostoma caucasicum — Staręga 1978: 201; Martens 2006: 192 (redescr).

creticum ( Roewer, 1927) ; loc. typ.: Greece, Crete, Topolia.

Nemastoma creticum Roewer, 1927: 454 ;— Gruber 1963: 310.

Mitostoma creticum — Roewer 1951: 148; Martens 1966: 350 (redescr.).

Centetostoma (Cretostoma) creticum — Kratochvíl 1958: 530.

Histricostoma creticum — Martens 1978: 129; Gruber 1978: 569.

dentipalpe ( Ausserer, 1867) ; loc. typ.: Austria, Innsbruck, Stubai-Valley, locality Husslhof.

Nemastoma dentipalpe Ausserer, 1867: 168 .

Carinostoma dentipalpe — Šilhavý 1969: 496.

Nemastoma (Lugubrostoma) dentipalpe — Kratochvíl 1958: 539.

Histricostoma slovenicum Hadži, 1973a: 49 ;—syn. Martens 1978: 130.

Histricostoma dentipalpe — Gruber 1976: 799.

drenskii Kratochvíl, 1958: 570 ; loc. typ.: Bulgaria, Rhodope Mountains, Chvojna, locality “Peštera“.

gruberi Snegovaya & Marusik, 2012: 62 ; loc. typ.: Turkey, Aydın, KuŞadası, Dilek Peninsula, Büyük Menderes

Delta National Park.

mitovi Snegovaya & Marusik, 2012: 61; loc. typ.: Turkey, Bursa, İnegöl, Great Oylat Cave.

Genus Mediostoma Kratochvíl, 1958 (11 species)

Histricostoma (Basostoma) Kratochvíl, 1958: 529 ; type species Nemastoma topolium Roewer, 1951 by original designation;— syn. Gruber 1976: 799.

Histricostoma (Mediostoma) Kratochvíl, 1958: 529 ; type species Nemastoma graecum Roewer, 1917 by original designation (see humerale ).

Mediostoma — Gruber 1976: 799.

Etymology. Gender neuter; Latin “medius” (in the middle) combined with „stoma“, in reference to Nemastoma . The name probably refers to the position of pseudoarticulations in the middle of the leg femora. In the original composition of subgenera in Histricostoma ( Kratochvíl 1958) it was compared to the basal-positioned pseudoarticulations in Basostoma.

armatum Martens, 2006: 189 ; loc. typ.: Iran, Mazandaran, S Alamdeh.

ceratocephalum Gruber, 1976: 790. loc. typ.: Turkey, S Anatolia, Vilayet Içel, 10 km NE Silifke. Remarks. Gruber (1976: 794) discussed a placement of ceratocephalum in either Giljarovia or Mediostoma and decided for a tentative placement in the latter. Males of ceratocephalum are unavailable to confirm other generic characters.

cypricum ( Roewer, 1951) loc. typ.: Cyprus (not further specified).

Nemastoma cypricum Roewer, 1951: 136 .

Histricostoma (Mediostoma) cypricum — Kratochvíl 1958: 569.

Mediostoma cypricum — Gruber 1976: 799; Martens 2006: 184.

globuliferum (L. Koch, 1867) ; loc. typ.: Greece, Syros (=Syra).

Nemastoma globuliferum L. Koch, 1867: 893 — Roewer 1923: 656.

Mediostoma globuliferum comb. n.: According to existing descriptions (L. Koch 1867: 893, Roewer 1923: 656) and the geographic origin of the type, this is most likely a Mediostoma species. The type seems lost ( Roewer 1923: 656). Roewer’s re-description is based on material from Naxos. It is here tentatively transferred to Mediostoma to exclude it from Nemastoma where it certainly does not belong. A revision is necessary.

haasi ( Roewer, 1953a) ; loc. typ.: Israel, Jerusalem.

Mitostoma haasi Roewer, 1953a: 209 .

Nemastoma haasi — Staręga 1973: 132.

Mediostoma haasi — Gruber 1976: 799; Martens 2006: 184.

humerale (C.L. Koch, 1839b) ; loc. typ.: Greece, Nauplia.

Nemastoma humerale C.L. Koch, 1839b: 38 ;—C.L. Koch 1847: 66.

Nemastoma quadripunctatum humerale — Roewer 1914: 148.

Nemastoma graecum Roewer, 1917: 152 ;—syn. Gruber 1976: 799.

Histricostoma (Mediostoma) graecum — Kratochvíl 1958: 529

Mediostoma humerale — Gruber 1976: 799; Mitov 2002: 1640.

nigrum Martens, 2006: 190 ; loc. typ.: Iran, Gilan Prov., seaside of Talysch Mountains.

pamiricum Staręga, 1986 ; loc. typ.: Tajikistan, West-Pamir, Vanč River Valley.

Centetostoma pamiricum — Staręga 1978: 200 (as nomen nudum).

Mediostoma pamiricum Staręga, 1986: 301 .

Remarks. Staręga (1978: 200) mentioned the species name without giving a description) and tentatively placed it in Centetostoma . The intended description (cited as “Staręga 1979” in Staręga 1978) was postponed until 1986, objecting to the previous generic placement, as only a female was available. Snegovaya (2010: 353) had no females to compare pamiricum with Starengovia which both occur in geographic proximity in an area with no other known members of the family.

stussineri ( Simon, 1885a) ; loc. typ.: Greece, Ossa Mountains, Kokkino Vracho Cave.

Nemastoma stussineri Simon, 1885a: 217 .

Histricostoma (Mediostoma) stussineri — Kratochvíl 1958: 569.

Mediostoma stussineri — Gruber 1976: 799; Mitov 2002: 1639.

Remarks. M. stussineri is the largest species within Mediostoma , matching body size of the related genus Pyza and also showing some genital-morphological affinities ( Gruber 1979, Mitov 2002). Its relationships should be reinvestigated.

variabile Martens, 2006: 185; loc. typ.: Iran, Azerbaijan, Makidi near Kaleybar.

vitynae ( Roewer, 1927) ; loc. typ.: Greece, Crete, Topolia Canyon.

Nemastoma vitynae Roewer, 1927: 455 .

Mitostoma omalosum Roewer, 1951: 146 ;—syn. Martens 1966: 350.

Nemastoma topolium Roewer, 1951: 135 ;—syn. Gruber 1976: 799.

Mediostoma vitynae — Gruber 1976: 799.

Remarks. Gruber (1976) stated that two of the three species of Basostoma, topolium and vitynae (excluding funebre ), are synonymous, and referred to Martens (1966). Martens listed only vitynae and Gruber probably referred to his description, thus synonymy of topolium with vitynae has to be attributed to Gruber. The type localities of topolium (Crete) and vitynae (Peleponnes) are geographically isolated and warrant reinvestigation.

Genus Mitostoma Roewer, 1951 (17 species, 3 subspecies)

Mitostoma Roewer, 1951: 140 ; type species Phalangium chrysomelas Hermann, 1804 by original designation;— Tedeschi & Sciaky 1997: 504.

Mitostoma (Mitostoma) — Kratochvíl 1958: 530.

Remarks. Mitostoma stands out of the bulk of Nemastomatidae in exhibiting a very complex, three-dimensionally structured penial glans. Molecular data ( Schönhofer & Martens 2010a) corroborated a placement of Mitostoma in a basal and isolated position to the rest of the Palearctic Nemastomatinae . However, Kratochvíl’s (1958) Mitostomatinae have not been accepted ( Gruber 1976: 800), as his definition included unrelated genera.

Martens (1978) synonymised many of Hadži's (1973b) late nominal species. However, several species from the Central Balkan, rarely mentioned subsequently, appear to be valid (I. Karaman, pers. comm.) and are here treated as such. Types of Hadži are mainly lost or in bad shape (T. Novak, pers. comm.), so collecting at the type localities is necessary to redefine these species.

Etymology. Gender neuter; from Greek „mitos“ (thread) and „stoma“ (mouth) referring to the slender, glistening pedipalps, with regards to the etymology of Nemastoma .

alpinum ( Hadži, 1931) ; loc. typ.: Slovenia, Triglav Massiv.

Nemastoma chrysomelas alpinum Hadži, 1931: 109 .

Mitostoma chrysomelas multidenticulatum Hadži, 1973a: 54 ;—syn. Martens 1978: 147.

Mitostoma chrysomelas michielii Hadži, 1973a: 55 ;—syn. Martens 1978: 147.

Mitostoma alpinum — Martens 1978: 147.

anophthalmum ( Fage, 1946) ; loc. typ.: Italy, Bergamo, Grotta grande della Cava di Burligo.

Nemastoma anophthalmum Fage, 1946: 328 .

Buresiolla anophthalmum — Kratochvíl 1958: 533.

Mitostoma anophthalmum — Martens 1978: 149; Tedeschi & Sciaky 1997: 507.

Remarks. From the complex penis of this otherwise character-poor troglobiont, Fage (1946: 328) already inferred a closer relationship to chrysomelas , which later became the type species of Mitostoma . Considerable size difference within this species warrants further investigation ( Martens 1978: 149, Tedeschi & Sciaky 1997: 507).

atticum ( Roewer, 1927) ; loc. typ.: Greece, Pentelikon Mountain, way up to Penteli Monastery.

Nemastoma atticum Roewer, 1927: 453 .

Mitostoma atticum — Roewer 1951: 144.

Remarks. The type (SMF RII/239) is a female with only two remaining legs. Judging from this poor material, it seems morphologically very similar to specimens of Mitostoma chrysomelas . The same type locality was given by Roewer in the description of several invalid species (see Crosbycus pentelicus ; Helversen & Martens 1972). It should be neglected if collections at this site cannot confirm the presence of similar animals.

cancellatum ( Roewer, 1917) ; loc. typ.: Bosnia and Herzegovina, Sarajevo.

Nemastoma cancellatum Roewer, 1917: 154 .

Mitostoma cancellatum — Roewer 1951: 147; Mitov 2000: 161.

carneluttii Hadži, 1973a: 57 ; loc. typ.: Montenegro, Vasojevičkem Komu. Remarks. Martens (pers. comm.) investigated the type in the Hadži Collection and assumed that it is close to cancellatum . A revision of both species is recommended.

chrysomelas ( Hermann, 1804) ; loc. typ.: France, Alsace, Strasbourg.

Remarks. Many names have been assigned to this widespread Central European species, based on differences in the variable colouration and pattern of anvil-shaped tubercles. Most of them were synonymised by Martens (1978). He also mentioned that all Mitostoma described by Avram (1969, 1970) fall within the variation of chrysomelas , but did not list them as synonyms, having not seen the type material. They were officially synonymised by Weiss (1996: 263) based on good knowledge of the Romanian fauna. Subsequent publications (e.g. by Weiss and Babalean) did only cite chrysomelas for Romania. Tedeschi and Sciaky (1997: 510) considered some alpine populations of chrysomelas as distinct species, which remained unattended.

Phalangium chrysomelas Hermann, 1804: 108 .

Nemastoma chrysomelas —C.L. Koch 1839b: 38.

Nemastoma quadricorne L. Koch, 1861: 142 ;—syn. Roewer 1914: 159.

Leiobunum troglodytes Wankel, 1861: 257 ;—syn. Šilhavý 1956a: 79. Remarks. According to Wankel’s (1861) drawings the type is a juvenile Mitostoma , and according to the type locality ( Czech Republic, Moravia caves) only chrysomelas applies (for troglodytes being preoccupied see also Hesperonemastoma packardi ). Šilhavý (1956a) did not list troglodytes in the synonymy of chrysomelas .

Nemastoma troglodytes — Roewer 1923: 657.

Nemastoma saxonica Hnatewytsch in Kästner, 1928: 18;—syn. Spoek 1963: 17.

Nemastoma spinosum Hnatewytsch, 1929: 231 ;—syn. Spoek 1963: 18. Remarks. Spoek (1963: 17) first placed spinosum in the synonymy of saxonica and later (1963: 18) both in chrysomelas . See also extensive discussion in Spoek (1963).

Nemastoma janetscheki Schenkel, 1950: 766 ;—syn. Martens 1978: 142.

Mitostoma chrysomelas — Roewer 1951: 142.

Nemastoma silhavyi Roewer, 1951: 143 ;—syn. Staręga 1976: 335.

Mitostoma spinosum — Kratochvíl (1934: 8): 146; syn. Spoek 1963: 18.

Mitostoma hadzii Roewer, 1951: 146 ;—syn. Martens 1978: 142.

Nemastoma chrisomelas — Cîrdei 1956: 74 (misspelling).

Mitostoma vosagorum Rambla, 1956: 152 ;—syn. Meijer 1973: 125.

Mitostoma (Mitostoma) chrysomelas — Kratochvíl 1958: 530.

Crosbycus bükkensis Loksa, 1962: 265 ;—syn. Martens 1978: 142.

Crosbycus transdanubicus Loksa, 1962: 267 ;—syn. Martens 1978: 142.

Mitostoma chrysomelas confusum Spoek, 1963: 21 ;—syn. Martens 1978: 142.

Mitostoma chrysomelas romanicum Avram, 1965: 181 ;—syn. Staręga 1976: 335.

Mitostoma chrysomelas romanium — Šilhavý 1966a: 70 (misspelling).

Crosbycus buekkensis — Gruber & Martens 1968: 138.

Mitostoma dumitrescui Avram, 1969: 140 ;— Martens 1978: 144; syn. Weiss 1996: 263.

Mitostoma orghidani Avram, 1969: 144 ;— Martens 1978: 144; syn. Weiss 1996: 263.

Mitostoma moldavica Avram, 1970: 172 ;— Martens 1978: 144; syn. Weiss 1996: 263.

Mitostoma helenae Avram, 1970: 174 ;— Martens 1978: 144; syn. Weiss 1996: 263.

Mitostoma rodnae Avram, 1970: 178 ;— Martens 1978: 144; syn. Weiss 1996: 263.

Mitostoma (Mitostoma) sketi Hadži, 1973a: 50 ; syn. n.: According to Hadži’s (1973a: 51) drawings and the locality (Zlot, Serbia) this is very likely Mitostoma chrysomelas .

Mitostoma chrysomelas francohorioni Hadži, 1973a: 52 ;—syn. Martens 1978: 142.

Mitostoma chrysomelas poleneci Hadži, 1973a: 56 ;—syn. Martens 1978: 142.

daccordii Tedeschi & Sciaky, 1997: 511; loc. typ.: Italy, Venetia, Monti Lessini, Cima Posta.

fabianae Tedeschi & Sciaky, 1997: 513; loc. typ.: Italy, Tuscany, Elba, N slopes of Mt. Tambora.

gracile ( Redikortsev, 1936) ; loc. typ.: Russia, Gagari, Sochi (restricted by Staręga 1978: 202).

Nemastoma gracile Redikortsev, 1936: 39 .

Mitostoma gracile — Roewer 1951: 150; Staręga 1978: 202; Martens 2006: 207.

macedonicum Hadži, 1973a: 58 ; loc. typ.: Republic of Macedonia (not further specified).

olgae olgae ( Šilhavý, 1939) ; loc. typ.: Montenegro, Risan, Golodražnica.

Nemastoma olgae olgae Šilhavý, 1939: 108 (for species), 109 (for subspecies).

Mitostoma olgae olgae — Roewer 1951: 144.

Remarks. Novak (2004) considered the subspecies decorum and kratochvili as invalid, being based on characters showing high variability. While his argumentation is reasonable, a reinvestigation might be worthwhile because these populations occur within a zone of high endemism (e.g. in Trogulus ; Schönhofer & Martens 2009). For this reason, the subspecies are maintained for the time being.

olgae decorum ( Šilhavý, 1939): 109; loc. typ.: Montenegro, Nikšiće, Dragova Luka.

Nemastoma olgae decorum Šilhavý, 1939: 109 .

Mitostoma olgae decorum — Roewer 1951: 144.

olgae kratochvili ( Šilhavý, 1939) : 109; loc. typ.: Croatia, Brač, Pučišća.

Nemastoma olgae kratochvili Šilhavý, 1939: 109 .

Mitostoma olgae kratochvili — Roewer 1951: 144.

olgae zorae Hadži, 1973a: 60; loc. typ.: Republic of Macedonia, Šar Planina, Karaorman Mt.

orobicum ( Caporiacco, 1949) ; loc. typ.: Italy, Bergamo, Monte Arera.

Nemastoma orobicum Caporiacco, 1949: 121 .

Mitostoma orobicum — Chemini 1985: 72; Tedeschi & Sciaky 1997: 508.

Remarks. The species was transferred to Mitostoma and synonymised with chrysomelas by Martens (1978), stating that J. Gruber had seen the juvenile type. Chemini (1985) redescribed and revalidated orobicum based on adult males.

patrizii Roewer, 1953b ; loc. typ.: Italy, Sicily, Grotta di San Giovanni Domusnovas.

Mitostoma patrizii Roewer, 1953b: 39 ;— Tedeschi & Sciaky, 1997: 505.

pyrenaeum ( Simon, 1879a) ; loc. typ.: France, Ariége, Moulis, Château de la Brûlerie, Grotte d'Aubert. Nemastoma pyrenaeum Simon, 1879a: 287 .

Mitostoma asturicum Roewer, 1951: 147 ; syn. n.: The type locality ( Spain, Asturia; Cave near Oviedo) is doubtful as no new material from there is known ( Prieto 2008). The male type was investigated (SMF RII/ 6210) and is very similar to specimens of M. pyrenaeum .

Mitostoma pyrenaeum — Roewer 1951: 149.

sabbadinii Tedeschi & Sciaky, 1997: 508; loc. typ.: Italy, Sardinia, Sassari, N slope of Mt. Limbara.

valdemonense Marcellino, 1974 ; loc. typ.: Italy, Sicily, Nebrodi, Portella di Femmina morta.

Mitostoma valdemonense Marcellino, 1974: 187 ;— Tedeschi & Sciaky 1997: 512.

zmajevicae Hadži, 1973a: 59; loc. typ.: Republic of Macedonia, Zmajevici.

Genus Nemaspela Šilhavý, 1966a (7 species)

Nemaspela Šilhavý, 1966a: 72 ; type species Burešiolla sokolovi Ljovuschkin & Starobogatov, 1963 by original designation;— Martens 2006: 169; Chemeris 2009: 286.

Remarks. Šilhavý mentioned only Nemaspela sokolovi in the definition of the genus, but also illustrated abchasica and caeca as Nemaspela (1966a: 68) .

Etymology. Gender neuter; combination of “ Nemastoma ” and Latin “spelaeum” (cave), indicating a cave adaptation within Nemastomatidae .

abchasica (Ljovuschkin & Starobogatov, 1963) ; loc. typ.: Abchasia, Niznaja Sakuranskaja Cave.

Burešiolla abchasica Ljovuschkin & Starobogatov, 1963: 47 .

Nemaspela abchasica — Šilhavý 1966a: 68; Martens 2006: 173; Chemeris 2009: 287.

birsteini Ljovuschkin, 1972 ; loc. typ.: Abchasia, Gudauta, Tarkiladze Caves.

Nemaspela birsteini Ljovuschkin, 1972: 66 ;— Chemeris 2009: 289 (redescr.).

caeca ( Grese, 1911) ; loc. typ.: Ukraine, Crimea, unnamed cave near Skelja.

Nemastoma caecum Grese, 1911: 108 .

Burešiolla caecum —Ljovuschkin & Starobogatov 1963: 43.

Nemaspela coeca — Šilhavý 1966a: 68 (misspelling).

Nemaspela caeca —Ljovuschkin 1972: 66; Chemeris 2009: 289 (redescr.).

femorecurvata Martens, 2006: 171; Georgia, Ratscha Mt. Range, Ambrolauri Distr., Sakishore Cave near Upper

Tlugi.

kovali Chemeris, 2009: 293; loc. typ.: Russia, Central Caucasus, Kabardino-Balkaria, source of Belaya River,

Fontanka Cave.

sokolovi (Ljovuschkin & Starobogatov, 1963) ; loc. typ.: Russia, Krasnodar Prov., Sochi, 15 km N of Khosta, Vorontsovskaya Cave.

Burešiolla sokolovi Ljovuschkin & Starobogatov, 1963: 47 .

Nemaspela sokolovi — Šilhavý 1966a: 72; Martens 2006: 175; Chemeris 2009: 294.

taurica ( Lebedinski, 1914) ; loc. typ.: Ukraine, Crimea, Ajutišik Cave.

Phalangodes taurica Lebedinski, 1914: 115 ;—Ljovuschkin 1972: 65.

Nemaspela taurica — Staręga 1978: 202; Martens 2006: 169.

Remarks. Ljovuschkin (1972) mentioned Phalangodes taurica but excluded it from species he listed under Nemaspela .

Genus Nemastoma C.L. Koch, 1836b (7 species, 4 subspecies)

Nemastoma C.L. Koch, 1836b: 70 ; type species Phalangium bimaculatum Fabricius, 1775 by subsequent fixation ( Thorell 1876: 467) and subsequent delineation of the type species ( Gruber & Martens 1968: 139).

Nemastoma (Lugubrostoma) Kratochvíl, 1958: 529 ; type species Phalangium lugubre Müller, 1776 by original designation;— syn. Gruber & Martens 1968: 139.

Nemastoma (Stridulostoma) Hadži, 1973a: 48 ; type species Nemastoma (Stridulostoma) seliskari Hadži, 1973a by monotypy;—syn. Gruber 1976: 797.

Remarks. Nemastoma has long been the sole genus of the family, and accumulated large numbers of species. The subsequent decline of species assigned to Nemastoma is directly correlated with elaborating the systematics and taxonomy of the family. After Roewer (1951) assigned the first Nemastoma species to new genera, the original genus still comprised 107 species. Kratochvíl (1958) further elaborated the systematics, but counted 86 Nemastoma species , of which he included 23 in the subgenus Lugubrostoma , that came closest to today’s definition of Nemastoma . Gruber and Martens (1968) clarified the type species of Nemastoma (not Paranemastoma quadripunctatum as stated by Kratochvíl 1958) and restricted the generic name to a defined number of seven species.

Etymology. Gender neuter; from Greek „nema“ (thread) and „stoma“ (mouth), likely referring to the threadlike and glistening pedipalps.

bidentatum bidentatum Roewer, 1914 ; loc. typ.: The locality information ( Romania, Comana Vlasca) was considered erroneous, as the type series also contained triste and bidentatum sparsum, with only the latter occuring in Romania, but not bidentatum bidentatum ( Gruber & Martens 1968: 143) . The type locality is to be corrected, but should await a revision of the subspecies complex and use a locality within the known distribution.

Nemastoma bidentatum Roewer, 1914: 141 ;— Gruber & Martens 1968: 141 (redescr.).

bidentatum pluridentatum Hadži, 1973a; loc. typ.: Bosnia, Mt. Vlašić.

Nemastoma (Lugubrostoma) triste pluridentatum Hadži, 1973a: 43 .

Nemastoma (Stridulostoma) seliskari Hadži, 1973a: 48 ;—syn. Gruber 1976: 797.

Nemastoma seliskari — Gruber 1976: 797.

Nemastoma bidentatum pluridentatum — Novak 2005: 314.

Remarks. Hadži (1973a) mentioned Stridulostoma seliskari as distinct from N. (L.) triste pluridentatum by the presence or absence of assumed stridulatory structures only. Gruber (1976: 797) therefore rejected the subgenus Stridulostoma within Nemastoma and suggested a reinvestigation of the two “species”. Later, the sole type specimen of seliskari was assumed to be a hybrid between b. bidentatum and b. sparsum ( Novak & Gruber 2000: 286). Novak (2005) placed pluridentatum in the synonymy of N. bidentatum and regarded it as a valid subspecies, which is followed here. The subspecies in bidentatum are currently re-investigated.

bidentatum sparsum Gruber & Martens, 1968: 146; loc. typ.: Austria, Leitha Mountains, Sonnenberg.

bidentatum relictum Gruber & Martens, 1968: 147 ; loc. typ.: Austria, Salzburg, Kleinartal, Tappenkarsee.

bimaculatum ( Fabricius, 1775) ; loc. typ.: Great Britain (not further specified).

Phalangium bimaculatum Fabricius, 1775: 440 .

Nemastoma lugubre bimaculatum — Roewer 1923: 654.

Nemastoma bimaculatum —C.L. Koch 1836b: 71; Meade 1855: 412; Gruber & Martens 1968: 150 (redescr.).

dentigerum Canestrini, 1873: 51 ; loc. typ.: Italy, Padova, Altichiero.

Nemastoma dentigerum Canestrini, 1873: 51 ;— Gruber & Martens 1968: 153 (redescr.).

lugubre ( Müller, 1776) ; loc. typ.: Denmark (not further specified).

Phalangium lugubre Müller, 1776: 192 .

Nemastoma lugubre — Simon 1879a: 281; Gruber & Martens 1968: 156 (redescr.).

Nemastoma lugubre helvetica Lessert, 1917: 57 ;—syn. Gruber & Martens 1968: 156.

Nemastoma lugubre var. striatum Morin, 1934: 36 ;—syn. Gruber & Martens 1968: 156.

Nemastoma lugubre striatum — Roewer 1951: 127.

Nemastoma (Lugubrostoma) lugubre — Kratochvíl 1958: 529.

Lugubrostoma lugubre — Šilhavý 1966a: 68.

schuelleri Gruber & Martens, 1968: 160; loc. typ.: Austria, Muhr, Lungau.

transsylvanicum Gruber & Martens, 1968: 165; loc. typ.: Romania, Valea Galbenii.

triste (C.L. Koch, 1835) ; loc. typ.: Germany, Bavaria, Regensburg.

Phalangium triste C.L. Koch, 1835: 20 .

Nemastoma triste —C.L. Koch 1847: 63; Gruber & Martens 1968: 167 (redescr.).

Nemastoma lugubre unicolor Roewer, 1914: 134 ;—syn. Gruber 1966: 45.

Nemastoma (Lugubrostoma) triste — Gruber 1966: 45; Gruber & Martens 1968: 167.

Lugubrostoma triste — Šilhavý 1966a: 68; Gruber & Martens 1968: 167.

Genus Nemastomella Mello-Leitão, 1936 (14 species, 2 subspecies)

Nemastomella Mello-Leitão, 1936: 10 ; type species Nemastomella integripes Mello-Leitão, 1936 by monotypy and original designation;— Staręga 1986: 303; Prieto 2004; 2008.

Remarks. Staręga (1986: 303) used Nemastomella integripes as type species to make the genus name available, while he synonymised integripes with dubia in the same paper.

Prieto (2004, 2008) used a narrow and careful concept for Nemastomella , which is acknowledged but extended here, assuming the secondary loss of male cheliceral apophyses (as in cristinae , close to bacilliferum and dubium regarding genital morphology; in sexmucronata probably secondary loss because of cave adaptation) and allowing more variation in dorsal armament, to remove species from their definitely erroneous placement in Nemastoma . The species cristinae, maarebensis, monchiquensis and sexmucronata are hereby tentatively included in Nemastomella (see respective Remarks).

Etymology. Gender feminine; diminutive form of Nemastoma , using the suffix “-ella”.

armatissima ( Roewer, 1962) ; loc. typ.: Portugal, Coimbra. Mitostoma armatissimum Roewer, 1962: 15 ;— Prieto 2008: 62. Nemastomella armatissima comb. n.: The type (SMF RII/11156) seems to be a female of Nemastomella close to spinosissima , but differs in being uniformly dark in colour, the first two areae having a row of anvil-shaped tubercles and the cones terminating in two large, fork-like spines. A full redescription has to await the collection of males, and placement and locality data require confirmation.

bacillifera bacillifera ( Simon, 1879a) ; loc. typ.: France, Gers, Condom, La Plaigne du Mage.

Nemastoma bacilliferum Simon, 1879a: 287 .

Nemastomella bacillifera simoni Roewer, 1914: 155 ;—syn. Dresco 1967b: 368.

Nemastomella bacillifera simplex Simon, 1913: 385 ;—syn. Dresco 1967b: 368.

Nemastoma formosum Roewer, 1951: 134 ; syn. n.: After investigating the type (SMF RII/9525, France, Biarritz ), this is certainly bacillifera .

Histricostoma (Mediostoma) bacilliferum — Kratochvíl 1958: 569.

Histricostoma bacillifera — Šilhavý 1966a: 69.

Nemastomella bacillifera — Staręga 1986: 304; Prieto 2004: 110.

bacillifera carbonaria ( Simon, 1907) ; loc. typ.: Spain, Huesca, Vio, Llobrica Cave.

Nemastoma carbonarium Simon, 1907: 553 .

Histricostoma (Mediostoma) carbonarium — Kratochvíl 1958: 569.

Nemastomella carbonaria — Staręga 1986: 304.

Nemastomella bacillifera carbonaria — Prieto 2004: 111.

cristinae ( Rambla, 1969) ; loc. typ.: Spain, Léon, Puerto de Piedrafita.

Nemastoma cristinae Rambla, 1969: 91 ;— Prieto 2004: 119.

Nemastomella cristinae — Staręga 1986: 304.

Remarks. Oddly, Rambla (1969) did not mention the name of her new species throughout the written text but only in the legends of the figures. Staręga (1986) transferred the species to Nemastomella but Prieto (2004) placed it back in Nemastoma as it did not fit a number of external characters, e.g. having no cheliceral apophysis in males, and differences in granulation. External characters, as cheliceral apophyses, can get lost within a genus, as known from Giljarovia and Nemaspela ( Martens 2006) . The genital morphology of cristinae conforms with that of baciliferum and dubium , and cristinae is best placed in Nemastomella and Staręga’s combination is hereby revived.

dentipatellae ( Dresco, 1967b) ; loc. typ.: Spain, Santander, Arredondo, Cueva La Cañuela.

Nemastoma dentipatellae Dresco, 1967b: 382 .

Nemastomella dentipatellae — Staręga 1986: 304; Prieto 2004: 112.

dipentata ( Rambla, 1959) ; loc. typ.: Spain, Sierra Guadarrama, Lozoya Springs.

Nemastoma bacilliferum dipentatum Rambla, 1959: 79 .

Nemastoma hermanni Kraus, 1959: 297 ;—syn. Kraus 1961: 345.

Nemastomella dipentata — Staręga 1986: 304; Prieto 2004: 113.

dubia ( Mello-Leitao, 1936) ; loc. typ.: Catalogne (not further specified).

Nemastoma dubium Mello-Leitao, 1936: 8 .

Nemastomella integripes Mello-Leitao, 1936: 11 ;—syn. Staręga 1986: 304.

Nemastoma franzi Kraus, 1959: 299 ;—syn. Kraus 1961: 346.

Nemastomella dubia — Staręga 1986: 304.

gevia Prieto, 2004: 114; loc. typ.: Spain, Jaén, Peal de Becerro: Complejo del Arroyo de la Rambla.

hankiewiczii (Kulczyṅski, 1909); loc. typ.: Portugal, Torres Vedras, Barro.

Nemastoma hankiewiczii Kulczy ṅski, 1909: 465.

Nemastoma roeweri Kraus, 1961: 341 ;—syn. Rambla 1980: 199.

Nemastomella hankiewiczii comb. n.: Rambla (1980: 199) regarded the differences between N. roeweri and N. hankiewiczii as intraspecific variation, while Kraus (1961: 343) depicted differences in the structure of the male cheliceral apophyses that, based on current systematics, would warrant placement in different genera. The issue cannot be resolved without checking more material and meanwhile Rambla's synonymy is retained. It is not clear what Kraus (1961) or Rambla (1980) perceived when discussing these species, but both forms show characters present in the Iberian radiation of Nemastomatidae , and a tentative placement in Nemastomella seems reasonable.

iberica (Rambla in Dresco, 1967b); loc. typ.: Spain, Sierra de Guadarrama (see Remarks).

Nemastoma ibericus Rambla in Dresco, 1967b: 382.

Nemastomella iberica — Staręga 1986: 304; Prieto 2004: 117. Remarks. The species was established based on material used for a re-description of N. bacilliferum bacilliferum based on non-type material ( Rambla 1959: 73). Dresco (1967b) recognised differences to the original bacilliferum , suggested the name ibericus and assigned Rambla as official author. Dresco specified the year of description as 1965, recited by Rambla (1967: 7), but it was not published in that year. Prieto (2004: 117) interprets the authorship of the species as “Rambla in Dresco, 1967” which is followed here. Dresco (1967) did not specify types or a type locality, but pointed solely to Rambla’s description, of which the illustrated male of the two series used for the description is to be preferred as lectotype (Collection Rambla, Barcelona: specimens n°335, 336). Depending on the formal lectotype selection, a more specific type locality is either Regajo del puerto de Navacerrada or Sanatorio Forest. Prieto (2004: 117) mentioned the species to be poorly characterised and advised revision.

maarebensis ( Simon, 1913) ; loc. typ.: Algier, Ifri Maareb, near Djebel Azerou Tidjer.

Nemastoma maarebense Simon, 1913: 385 .

Nemastomella maarebensis comb. n.: Known from one female only, any generic association remains preliminary, while placement within Nemastoma can be rejected. The type shows superficial similarity with Nemastomella sexmucronatum and is therefore tentatively placed in Nemastomella . Delfosse (2004: 48) erroneously reported the species for France.

manicata ( Simon, 1913) ; loc. typ.: Spain, Lérida, Llastarri, Minas dei Canal-B.

Nemastoma manicata Simon, 1913: 386 .

Nemastomella manicata — Prieto 2004: 112.

monchiquensis ( Kraus, 1961) ; loc. typ. Portugal: N slope Sierra de Monchique.

Nemastoma monchiquense Kraus, 1961: 342 ;— Rambla 1967: 6.

Nemastomella monchiquensis comb. n.: According to the form of the cheliceral apophysis and the penial stylus ( Rambla 1967) monchiquense is clearly a member of the Iberian radiation including Acromitostoma , Centetostoma and Nemastomella . However, it differs from the first two genera by the lack of characteristic penial wings, further from Acromitostoma in lacking anvil-shaped tubercles, and from Centetostoma in lacking large, knob-like tubercles on the dorsum and on the supracheliceral sclerites. N. monchiquense is transferred to Nemastomella for lacking these characters, but also for the penial glans armed with long, slender spines, for posessing rows of elongated papillae on the dorsal opisthosoma and for having a similar body size ( Kraus 1961, Rambla 1967).

sexmucronata ( Simon, 1911) ; loc. typ.: Spain, Santander, Santillana del Mar, Cueva de Altamira.

Nemastoma sexmucronatum Simon, 1911: 205 ;— Prieto 2004: 119

Nemastomella sexmucronata — Staręga 1986: 304. Remarks. Placing sexmucronatum in Nemastoma has to be rejected and it is tentatively reinstated in Nemastomella following Staręga 1986. Still, several features of sexmucronatum deviate from this genus. The penial truncus is narrowed below the glans, with deep lateral furrows, a stylus is almost absent, and large spines are missing from the glans ( Prieto 2004). Dorsal spines and male cheliceral apophyses may be reduced in this cave-dwelling species, but its position in Nemastomella is isolated, requiring investigation, as acknowledged by Prieto (2004).

spinosissima ( Kraus, 1961) ; typ. loc.: Spain, Navarra, Picos de Europa, foot of Peña Vieja Mt.

Nemastoma spinosissima Kraus, 1961: 345 .

Nemastomella spinosissima — Staręga 1986: 304; Prieto 2004: 117.

Genus Paranemastoma Redikorzev, 1936 (21 species, 5 subspecies)

Paranemastoma Redikorzev, 1936: 40 ; type species Paranemastoma superbum Redikorzev, 1936 by monotypy and original designation;— Mcheidze 1959: 111; Staręga 1966a: 392; Gruber & Martens 1968: 139; Martens 1978: 116.

Nemastoma (Dromedostoma) Kratochvíl, 1958: 529 , 538; type species Nemastoma radewi Roewer, 1926 by monotypy;—syn. Staręga 1966a: 392.

Buresiolla Kratochvíl, 1958: 529 ; type species Nemastoma bureschi Roewer, 1926 by original designation.

Paranemastoma (Buresiolla) — Staręga 1976: 332. Remarks. Staręga (1976: 332) transferred the type of Buresiolla , bureschi , to Paranemastoma , thus effecting synonymy with Paranemastoma (ICZN 61.3.1). Preserving the subgenus was not subsequently approved ( Mitov 2011).

Remarks. While the genus Paranemastoma is nowadays accepted and well-defined upon genital morphology, its revalidation and redefinition are difficult to pinpoint. Roewer (1951: 95) and Kratochvíl (1958: 526) rejected Paranemastoma as defined by Redikorzew (1936), who delineated his monotypic genus upon the presence or absence of pseudosegmentation of the legs. Based on genital morphology, and also on large body size, Šilhavý (1966a: 70) assumed close relationship between seven Nemastoma species , including quadripunctatum , all later accepted as Paranemastoma . He further delineated this assembly from species which are today in the sister genera Carinostoma ( elegans ) and Histricostoma ( dentipalpe ). Mcheidze (1959: 111) re-established Paranemastoma as a subgenus for superbum , and Staręga (1966a: 392) placed Dromedostoma in its synonymy. Staręga did so for superbum only, while quadripunctatum and others remained in the subgenus Nemastoma . In 1968, Gruber and Martens redefined Nemastoma for a narrow ensemble of species. To reinforce this, they associated the earlier concepts of Nemastoma by Kratochvíl (1958: 529; excluding Lugubrostoma ), Roewer and Šilhavý (for the latter two no explicit references were given) to conform better to Paranemastoma and its type superbum . While this could be regarded as redefining Paranemastoma , the concepts of Kratochvíl, Roewer and Šilhavý can hardly be seen in agreement, and Gruber and Martens (1968) gave no further definition or listed associated species. Yet, later authors have followed their proposal (see statement in Avram 1973a: 31), using mainly the conclusive concept of Šilhavý (1966a), and accepted species close to quadripunctatum as Paranemastoma ( Helversen & Martens 1971, Dumitrescu 1972, Avram 1973a). Martens (1978: 116) redefined the genus, stating the species-count to be twenty, but did not present a list, and his bibliography for Paranemastoma and the four discussed species is unusually incomplete (e.g. not listing his own contributions: Gruber & Martens 1968, Helversen & Martens 1971). In summary, assignment of species to Paranemastoma has always been a stepwise process, adding single species or small groups of species, and a complete list of species was never published. Taxonomy represents a nightmare, having to deal with about 100 names, often based on variable external characters as spination and colouration. Informative characters for species delineation, as male genital morphology and cheliceral apophysis structures, are complex and need careful re-evaluation.

Etymology. Gender neuter; Greek; combination of “para” (“close to”) and “ Nemastoma ” indicating the close relationship perceived at the time of separation.

ancae Avram, 1973b ; loc. typ.: Romania, Apuseni Mountains , RăcaŞ region (from different mines; no explicit type or type locality was specified).

Paranemastoma ancae Avram, 1973b: 135 ;— Babalean & Ilie 2003: 64; Weiss 1996: 265. Remarks. Weiss (1996: 265) treated P. ancae as valid and related to P. quadripunctatum but suggested reinvestigation.

armatum (Kulczyṅski, 1909); loc. typ.: Hercegovina, Domanović.

Nemastoma armatum Kulczy ṅski, 1909: 468.

Nemastoma quadripunctatum armatum — Roewer 1923: 661.

Paranemastoma armatum — Novak 2005: 316 (stated revision needed).

aurigerum aurigerum ( Roewer, 1951) ; loc. typ.: Bulgaria, Burgas.

Nemastoma aurigerum Roewer, 1951: 119 .

Paranemastoma aurigerum aurigerum — Staręga 1976: 322.

aurigerum ryla ( Roewer, 1951) ; loc. typ.: Bulgaria, Tscham Kuria, Borovec (restricted by Staręga 1976: 324). Nemastoma ryla Roewer, 1951: 120 .

Nemastoma (Dromedostom) ryla — Kratochvíl 1958: 562.

Paranemastoma aurigerum ryla — Staręga 1976: 323.

aurigerum joannae Staręga, 1976: 324; loc. typ.: Bulgaria, Belasica Mt., 30 km WSW Petrič, Ključ.

aurosum (L. Koch, 1869) ; loc. typ.: Greece.

Nemastoma aurosum L. Koch, 1869: 165 .

Nemastoma quadripunctatum aurosum — Roewer 1951: 661.

Paranemastoma aurosum — Novak 2005: 318 (stated revision needed).

beroni Mitov, 2011: 304; loc. typ.: Bulgaria, Sandanski Distr., Slavyanka Mountains, cave Stoykova Dupka 1. bicuspidatum (C.L. Koch, 1835) ; loc. typ.: Austria, Salzburg, Gastein.

Phalangium bicuspidatum C.L. Koch, 1835: 16 .

Nemastoma bicuspidatum —C.L. Koch 1836b: 70; Šilhavý 1956a: 124, 141 (erroneously listed as type species of Nemastoma ).

Nemastoma quadripunctatum bicuspidatum — Roewer 1923: 663.

Nemastoma reimoseri Roewer, 1951: 108 ; syn. n.: According to size, colouration and distribution stated for reimoseri , species other than biscupidatum can be excluded.

Paranemastoma bicuspidatum — Avram 1973a: 31; Martens 1978: 121 (redescr.).

bureschi ( Roewer, 1926) ; loc. typ.: Bulgaria, Isker Defilé, cave above Tzerowo.

Nemastoma bureschi Roewer, 1926: 300 .

Nemastoma tunetanum Roewer, 1951: 130 ; syn. n.: Investigation of the syntypes of N. tunetanum (SMF RII/ 241) show that these are without doubt Paranemastoma that perfectly fit the redescription of P. bureschi by Mitov (2011). N. tunetanum from “ Tunisia ” is another misplaced Roewer species.

Crosbycus bureschi — Roewer 1951: 152.

Buresiolla bureschi — Kratochvíl 1958: 529.

Buresiolla tunetorum — Kratochvíl 1958: 527 (misspelling).

Buresiolla tunetanum — Kratochvíl 1958: 532.

Paranemastoma (Buresiolla) bureschi — Staręga 1976: 332.

Paranemastoma bureschi — Karaman 1995b: 334; Mitov 2011: 303 (redescr.).

corcyraeum ( Roewer, 1917) ; loc. typ.: Greece, Corfu.

Nemastoma quadripunctatum corcyraeum Roewer, 1917: 143 .

Paranemastoma corcyraeum — Novak 2004: 248.

filipes ( Roewer, 1917) ; loc. typ.: Azerbaijan, Lenkoran.

Nemastoma quadripunctatum var. filipes Roewer, 1917: 144 .

Nemastoma filipes — Roewer 1923: 665.

Nemastoma (Dromedostoma) filipes — Kratochvíl 1958: 538.

Paranemastoma filipes — Martens 2006: 203 (redescr.)

iranicum Martens, 2006: 204; loc. typ.: Iran, Mazandaran, 11 km E Alasht.

kalischevskyi ( Roewer, 1951) ; loc. typ.: Georgia, Suchumi Distr., Azhazy.

Nemastoma kalischevskyi Roewer, 1951: 122 .

Nemastoma suchumium Roewer, 1951: 122 ;—syn. Martens 2006: 196.

Nemastoma charitonovi Mcheidze, 1952: 545 ;—syn. Martens 2006: 196.

Paranemastoma kalischevskyi — Staręga 1966a: 390.

Paranemastoma charitonovi — Staręga 1978: 204.

Paranemastoma suchumium — Staręga 1978: 205.

kochii ( Nowicki, 1870) ; loc. typ.: High Tatras Mountains (mostly in Slovakia).

Nemastoma kochii Nowicki, 1870: 57 .

Nemastoma quadripunctatum kochii — Roewer 1923: 662.

Nemastoma nicaeum Roewer, 1951: 110 ;—syn. Martens 1978: 124.

Nemastoma hungaricum Roewer, 1951: 114 ;—syn. Martens 1978: 124.

Paranemastoma kochii — Avram 1973a: 31; Martens 1978: 124 (redescr.).

longipes ( Schenkel, 1947) ; loc. typ.: Albania, Tirana.

Nemastoma gigas longipes Schenkel, 1947: 10 .

Nemastoma longipes — Roewer 1951: 116.

Nemastoma tenue Hadži, 1973a: 31 ;—syn. Novak 2004: 248. Paranemastoma longipes — Novak 2004: 248; Mitov, 2000: 160. Remarks. Novak (2004) considers longipes as valid and related it to corcyraeum .

quadripunctatum ( Perty, 1833) ; loc. typ.: Germany, vicinity of Munich.

Phalangium quadripunctatum Perty, 1833: 204 .

Phalangium flavimanum C.L. Koch, 1835: 22 ; syn. n.: Judging from the description (C.L. Koch 1835) and the area it was collected from (between Bad Gastein and Triest), a Paranemastoma , already associated with quadripunctatum by Roewer (1914: 144).

Nemastoma flavimanum —C.L. Koch 1847: 64; Roewer 1914: 144.

Nemastoma quadripunctatum — Simon 1879a: 279.

Nemastoma quadripunctatum var. moesiacum Roewer, 1917: 143 ; syn. Novak & Gruber 2000: 286. Nemastoma quadripunctatum quadripunctatum — Roewer 1923: 660.

Nemastoma nervosum Roewer, 1923: 668 ;—syn. Martens 1978: 117.

Nemastoma quadripunctatum var. lessinensis Caporiacco, 1940: 34 ;—syn. Martens 1978: 117.

Nemastoma chaucicum Roewer, 1951: 105 ;—syn. Martens 1978: 117.

Nemastoma alfkeni Roewer, 1951: 106 ;—syn. Martens 1978: 117.

Nemastoma gertschi Roewer, 1951: 107 ;—syn. Martens 1978: 117.

Nemastoma austriacum Roewer, 1951: 108 ;—syn. Martens 1978: 117.

Nemastoma lessinensis — Roewer 1951: 110.

Nemastoma strasseri Roewer, 1951: 110 ;—syn. Novak 2005: 318.

Nemastoma armeniacum Roewer, 1951: 122 ;—syn. Martens 2006: 206. Remarks. Martens (2006) did not propose the synonymy but pointed to the similarity with quadripunctatum and the high possibility of the type locality of armeniacum being wrong. Martens is followed here, and the synonymy attributed to him. Crosbycus lengersdorfi Roewer, 1951: 152 ;—syn. Martens 1978: 117.

Paranemastoma quadripunctatum — Helversen & Martens 1971: 380; redescr. Martens 1978: 117. Nemastoma mediosignatum Hadži, 1973a: 24 ;—syn. Martens 1978: 117.

Nemastoma slovenicum Hadži, 1973a: 26 ;—syn. Martens 1978: 117.

Nemastoma triglavense Hadži, 1973a: 27 ;—syn. Martens 1978: 117.

Nemastoma quadripunctatum carniolicum Hadži, 1973a: 28 ;—syn. Martens 1978: 117.

Nemastoma emonense Hadži, 1973a: 29 ;—syn. Martens 1978: 117.

radewi ( Roewer, 1926) ; loc. typ.: Bulgaria, Trnowo Distr., near Monastir Dranowski, Wodnata Pestera Cave. Nemastoma radewi Roewer, 1926: 299 .

Nemastoma radevi — Wolf 1938: 630 (misspelling); Šilhavý 1966a: 68.

Nemastoma riparium Roewer, 1951: 108 ;—syn. Komposch & Gruber 2004: 525.

Nemastoma schenkeli Roewer, 1951: 108 ;—syn. Komposch & Gruber 2004: 525.

Nemastoma polonicum Roewer, 1951: 112 ;—syn. Novak & Gruber 2000: 287.

Nemastoma insulare Roewer, 1951: 116 ;—syn. Novak 2004: 247. Remarks. Roewer used specimens of radewi for several false or questionable descriptions of new species. Gruber (in Novak 2004: 247) mentioned the type locality Hvar (Lesina) as “dubious”.

Nemastoma (Dromedostoma) radewi — Kratochvíl, 1958: 555.

Nemastoma (Dromedostoma) atanasovi Kratochvíl, 1958: 562 ;—syn. Staręga 1976: 326.

Nemastoma (Dromedostoma) markovi Kratochvíl, 1958: 548 ;—syn. Staręga 1976: 326.

Nemastoma markovi — Šilhavý 1966a: 68.

Nemastoma (Dromedostoma) paspalevi Kratochvíl, 1958: 558 ;—syn. Staręga 1976: 326.

Nemastoma (Dromedostoma) paspalevi var. nigrum Kratochvíl, 1958: 560 ;—syn. Staręga 1976: 326. Nemastoma (Dromedostoma) atanasovi balcanica Kratochvíl, 1958: 562 ;—syn. Staręga 1976: 326. Paranemastoma radevi — Avram 1973a: 34.

Paranemastoma atanasovi — Avram 1973a: 35.

Paranemastoma paspalevi — Avram 1973a: 35.

Nemastoma (Dromedostoma) megarae Hadži, 1973a: 42 ;—syn. Novak 2005: 317.

sillii sillii ( Herman, 1871) ; loc. typ.: Romania, Sibiu.

Nemastoma sillii Herman, 1871: 28 .

Nemastoma lineatum Sørensen in Lendl 1894: 20 —syn. Martens 1978: 126.

Nemastoma gigas Sørensen in Lendl 1894: 28 —syn. Staręga 1978: 205.

Nemastoma quadripunctatum sillii — Roewer 1923: 662.

Nemastoma bishopi Roewer, 1951: 106 ;—syn. Martens 1978: 126. Remarks. The type locality ( Germany, Black Forest, Titi Lake) is situated within the geographical range of P. quadripunctatum , not of sillii . The issue is only of academic interest as it would place bishopi in the synonymy of another Paranemastoma species. Nemastoma carpathicum Roewer, 1951: 112 ;—syn. Avram 1973a: 60.

Nemastoma politum Roewer, 1951: 112 ;—syn. Staręga 1978: 205.

Nemastoma wiehlei Roewer, 1951: 114 ;—syn. Martens 1978: 126.

Nemastoma zilchi Roewer, 1951: 120 ;—syn. Martens 1978: 126.

Nemastoma bulgaricum Roewer, 1951: 120 ;—syn. Staręga 1976: 331.

Nemastoma rumelium Roewer, 1951: 120 ;—syn. Staręga 1976: 331.

Nemastoma mohilevum Roewer, 1951: 125 ;—syn. Staręga 1978: 205.

Nemastoma romanium Roewer, 1951: 125 ;—syn. Martens 1978: 126.

Nemastoma rumanicum — Martens 1978: 126 (misspelling).

Crosbycus cavernae Roewer, 1951: 152 ;—syn. Staręga 1978: 205.

Paranemastoma sillii — Avram 1973a: 31; Martens 1978: 126; (both: redescr.).

Paranemastoma gigas — Avram 1973a: 30.

Paranemastoma silli silli Babalean, 2011: 42 .

silli monticola Babalean, 2011: 42; loc. typ.: Romania, Parâng Mts., Mount Vârful Mohorul, at lake Câlcescu.

simplex ( Giltay, 1932) ; loc. typ.: Greece, Taygetos Mts., Ladas (orig. Lada).

Nemastoma simplex Giltay, 1932: 1 ; comb. n.: Roewer (1951) related simplex to Nemastoma filipes , which is unlikely for geographical reasons. He seems the only one to have discussed this species, which, according to the original description, is a Paranemastoma and distinct from others from the Greece mainland. Paranemastoma is confirmed for the Peloponnese (Martens pers. comm.).

Crosbycus graecus Giltay, 1932: 2 ; syn. n.: Judging from Giltay’s (1932) drawings (single opisthosomal sclerite) and from the body size (3 mm), the type of Crosbycus graecus is a juvenile Paranemastoma . Rambla (1968: 69) and Gruber and Martens (1968: 138) came to the same conclusion, but did not assign C. graecus to a valid species, although it is likely conspecific with P. simplex specimens collected from the very same spot.

superbum Redikorzev, 1936 ; loc. typ.: Georgia, Batumi (Batum).

Paranemastoma superbum Redikorzev, 1936: 40 ;— Gruber & Martens 1968: 139 (see remarks on Paranemastoma ); Martens 2006: 200 (redescr.).

Nemastoma supersum Roewer, 1951: 126 (invalid nomen novum for superbum );— Staręga 1978: 206. Nemastoma superbum bacuriana Mcheidze, 1959: 111 ;—syn. Staręga 1978: 206.

Nemastoma superbum bacurianum — Mcheidze 1964: 117 (misspelling).

Nemastoma (Paranemastoma) superbum — Staręga 1966a: 392.

thessalum ( Simon, 1885a) ; loc. typ.: Greece, Ossa Mountains, summit.

Nemastoma thessalum Simon, 1885a: 216 .

Nemastoma quadripunctatum thessalum — Roewer 1923: 662.

Paranemastoma teshalum — Avram 1973a: 31 (misspelling).

Paranemastoma thessalum — Novak 2005: 318.

titaniacum ( Roewer, 1914) ; loc. typ.: Bosnia (not further specified).

Nemastoma titaniacum Roewer, 1914: 162 .

Nemastoma bimaculosum Roewer, 1951: 118 ; syn. n.: Novak 2005: 316 related bimaculosum to P. titaniacum , which is followed here.

Nemastoma acrospinosum Roewer, 1951: 125 . Remarks. Novak (2004: 248) referred to the thesis of Karaman (1995a: 33) who cited Gruber (in litt.) to be responsible for this synonymy, though he never officially published it.

Nemastoma (Dromedostoma) titaniacum — Kratochvíl 1958: 538.

Nemastoma acrospinosum pretneri Hadži, 1973a: 34 ;—syn. Karaman 1995a: 35.

Nemastoma grabovicae Hadži, 1973a: 37 ;—syn. Karaman 1995a: 35.

Nemastoma bipunctatum Hadži, 1973a: 36 ;—syn. Karaman 1995a: 35.

Paranemastoma titaniacum — Karaman 1995b: 334; Mitov 2000: 161.

werneri (Kulczyṅski, 1903); loc. typ.: Turkey, Uludağ Mt. (orig. Olympus Bithynicus).

Nemastoma werneri Kulczy ṅski, 1903: 673;— Cîrdei 1960: 80.

Nemastoma quadripunctatum werneri — Roewer 1923: 663.

Paranemastoma quadripunctatum werneri — Cîrdei 1960: 80.

Paranemastoma werneri ; new rank: Cîrdei (1960: 80) recorded werneri from former Yugoslavia and Šilhavý (1956a: 142) cited the name for Poland. Staręga (1978: 205) interpreted these specimens as silli , and regarded werneri , described from the Anatolian Peninsula, as separate. Unlikely to be a subspecies of quadripunctatum for geographical reasons, werneri is treated here as species untill it is revised. The presence of Paranemastoma at the type locality has been confirmed (NHMW).

Paranemastoma considered as nomina dubia: Most of the following names have never been mentioned following the original publication and are not identifiable as distinct species. On the basis of their descriptions, e.g. body size and spination, they can be transferred to Paranemastoma . Many have been described by Roewer (1951), in detail difficult to recognise for likely using false localities and already described species. Others were described by Hadži (1973a) upon variants and many of his types are lost and, thus recognition of his species is also difficult. If these names cannot be assigned to distinct species, they should be treated as invalid.

aeginum ( Roewer, 1951) .

Nemastoma aeginum Roewer, 1951: 121 .

Paranemastoma aeginum comb. n.

amseli ( Roewer, 1951) .

Nemastoma amseli Roewer, 1951: 123 .

Paranemastoma amseli — Marcellino 1980: 333. Remarks. Never collected again, Paranemastoma probably does not occur in Sardinia and amseli is likely to be another “misplaced” Roewer species.

amuelleri ( Roewer, 1951) .

Nemastoma amülleri Roewer, 1951: 118 .

Paranemastoma amuelleri comb. n.

bolei ( Hadži, 1973a) .

Nemastoma (Dromedostoma) bolei Hadži, 1973a: 38 .

Paranemastoma bolei comb. n.

brevipalpatum ( Roewer, 1951) .

Nemastoma brevipalpatum Roewer, 1951: 122 .

Paranemastoma brevipalpatum — Prieto 2008: 56 (assumed false loc. typ.; not Spain).

caporiaccoi ( Roewer, 1951) .

Nemastoma caporiaccoi Roewer, 1951: 123 .

Paranemastoma caporiaccoi comb. n.: According to the original illustrations and size, it is definitely a Paranemastoma , a genus not present in Sicily.

carneluttii ( Hadži, 1973a) .

Nemastoma carneluttii Hadži, 1973a: 39 .

Paranemastoma carneluttii comb. n.

emigratum ( Roewer, 1959) .

Nemastoma emigratum Roewer, 1959: 37 .

Paranemastoma emigratum comb. n.: According to Roewer’s drawings (1959: 38) this is most likely a Paranemastoma . While the collector K. Lindberg insisted that his specimen is from India, this is unlikely given the known distribution of the genus and family. Confusion with Lindberg’s rich collections from Greece and Turkey, where Paranemastoma is present, seems likely.

ferkeri ( Roewer, 1951) .

Nemastoma ferkeri Roewer, 1951: 121 .

Paranemastoma ferkeri comb. n.

gigas montenegrinum ( Nosek, 1904)

Nemastoma gigas montenegrinum Nosek, 1904: 4 .

Paranemastoma gigas montenegrinum comb. n.: The name seemingly was not used after the original description and the type locality, Montenegro, is not in the distribution area of silli , with which gigas was synonymised. Avram (1973a), discussing the synonymy of gigas , did not mention subspecies of gigas , some of which were later accepted as species (e.g. longipes ). According to the description, montenegrinum could conform to titaniacum , rendering the latter synonymous. For the lack of an explicit type locality and the likely loss of the montenegrinum type, titaniacum is preserved for now.

gostivarense ( Hadži, 1973a) .

Nemastoma gostivarense Hadži, 1973a: 32 .

Paranemastoma gostivarense comb. n.

ikarium ( Roewer, 1951) .

Nemastoma ikarium Roewer, 1951: 125 .

Paranemastoma ikarium comb. n.

ios ( Roewer, 1917) .

Nemastoma quadripunctatum ios Roewer, 1917: 143 .

Nemastoma ios — Roewer 1951: 121.

Paranemastoma ios comb. n.

kaestneri ( Roewer, 1951) .

Nemastoma kästneri Roewer, 1951: 121 .

Paranemastoma kaestneri comb. n.

longipalpatum ( Roewer, 1951) .

Nemastoma longipalpatum Roewer, 1951: 116 ;

Paranemastoma longipalpatum comb. n.

macedonicum ( Hadži, 1973a) .

Nemastoma macedonicum Hadži, 1973a: 25 .

Paranemastoma macedonicum comb. n.

machadoi ( Roewer, 1951) .

Nemastoma machadoi Roewer, 1951: 123 .

Paranemastoma machadoi — Prieto 2008: 55 (excluded from Spain).

mackenseni ( Roewer, 1923) .

Nemastoma mackenseni Roewer, 1923: 668 .

Paranemastoma mackenseni comb. n.

multisignatum ( Hadži, 1973a) .

Nemastoma (Dromedostoma) multisignatum Hadži, 1973a: 35

Paranemastoma multisignatum comb. n.

nigrum ( Hadži, 1973a) .

Nemastoma (Dromedostoma) nigrum Hadži, 1973a: 32 .

Paranemastoma nigrum comb. n.

perfugium ( Roewer, 1951) .

Nemastoma perfugium Roewer, 1951: 110 .

Paranemastoma perfugium comb. n.: Roewer (1951) placed perfugium in the group of Paranemastoma -like Nemastomatidae . The type locality (Elba, Italy) is otherwise not known to be inhabited by Paranemastoma and hence doubtful.

redikorzevi ( Roewer, 1951) .

Nemastoma redikorzevi Roewer, 1951: 125 .

Paranemastoma redikorzevi — Staręga 1978: 205.

Paranemastoma roeweri Staręga, 1978: 205 ;— Chevrizov 1979: 14; syn. n.: Staręga (1978: 205) found Nemastoma redikorzevi Roewer (loc. typ. Ukraine “ Krim ”) to be preoccupied by Nemastoma redikorzevi Charitonov , renamed it roeweri and transferred it to Paranemastoma . In this case of primary homonymy, the younger name is not to be automatically replaced, as Staręga transferred the species to different genera in the same paper, not notifying the commission (ICZN 23.9.5).

santorinum ( Roewer, 1951) .

Nemastoma santorinum Roewer, 1951: 124 .

Paranemastoma santorinum comb. n.

senussium ( Roewer, 1951) .

Nemastoma senussium Roewer, 1951: 123 .

Paranemastoma senussium comb. n.: According to drawing and size, it is definitely a Paranemastoma unlikely to occur in the North African Kufra Oasis.

sketi ( Hadži, 1973a) .

Nemastoma (Dromedostoma) sketi Hadži, 1973a: 41 .

Paranemastoma sketi comb. n.

spinosulum (L. Koch, 1869) .

Nemastoma spinosulum L. Koch, 1869: 165 .

Nemastoma spinulosum — Roewer 1951: 140 (misspelling).

Paranemastoma spinosulum comb. n. Remarks. L. Koch (1869) provided descriptions as a determination key, allowing association with Paranemastoma from the mentioned pair of prominent spines only. Roewer (1951: 140) mentioned it as N. spinulosum , while the species seems not recognisable any more.

umbo ( Roewer, 1951) .

Nemastoma umbo Roewer, 1951: 122 .

Paranemastoma umbo — Staręga 1978: 206; Martens 2006: 206. Remarks. Described from the Caucasus, Martens (2006: 207) questioned that umbo originates from this geographic area, but was unable to assign it to another species.

Genus Pyza Star ę ga, 1976 (4 species)

Pyza Staręga, 1976: 315 ; type species Nemastoma bosnicum Roewer, 1917 by monotypy and original designation;— Gruber 1979: 559.

Remarks. Gruber (1979) revised the genus and mentioned that the type species, P. bosnica , is distantly related to the remaining species and only tentatively placed them in Pyza . A phylogenetic comparison with Mediostoma and Vestiferum is advised.

Etymology. Gender feminine; although sounding similar to modern Neapolitan fast food, the true titular saints are globular Polish potato dumplings, in resemblance to the body form of these opilionids.

anatolica ( Roewer, 1959) ; loc. typ.: Turkey, Cheykhan, caves 30 km N Mardine.

Crosbycus anatolicus Roewer, 1959: 38 .

Pyza anatolica — Gruber 1979: 572.

bosnica ( Roewer, 1917) ; loc. typ.: Bosnia, Miljačka River Valley near Sarajevo.

Nemastoma bosnicum Roewer, 1917: 150 .

Nemastoma gallwitzi Roewer, 1923: 667 ;—syn. Staręga 1976: 315.

Nemastoma (Lugubrostoma) bosnicum orientale Kratochvíl, 1958: 565 ;—syn. Staręga 1976: 316.

Nemastoma (Lugubrostoma) goliae Hadži, 1973a: 45 ;—syn. Gruber 1979: 560.

Nemastoma (Lugubrostoma) sarae Hadži, 1973a: 46 ;—syn. Gruber 1979: 560.

Pyza bosnica — Staręga 1976: 315.

navarrense ( Roewer, 1951) ; loc. typ.: Spain, Navarra. Remarks. According to Gruber (1979: 561) the type locality is incorrect. He showed that the species is widely distributed in Greece and subsequent records from the Pyrenees are absent. Nemastoma navarrense Roewer, 1951: 130 . Nemastoma lindbergi Roewer, 1959: 36 ;—syn. Gruber 1979: 561. Pyza navarrense — Gruber 1979: 561.

taurica Gruber, 1979: 566 ; Turkey; loc. typ.: Turkey, Vilayet Antalya, 18 km W Topraktepe, E Manavgat.

Genus Saccarella Schönhofer & Martens, 2012 (1 species)

Saccarella Schönhofer & Martens, 2012: 412 ; type species Saccarella schilleri Schönhofer & Martens, 2012 by monotypy and original designation.

Etymology. Gender feminine; derived from the name of the type locality Monte Saccarello.

schilleri Schönhofer & Martens, 2012: 412 ; typ. loc.: Italy, Liguria, Imperia Prov., Monte Saccarello, Monesi di

Triora.

Genus Starengovia Snegovaya, 2010 (1 species)

Starengovia Snegovaya, 2010: 351 ; type species Starengovia kirgizica Snegovaya, 2010 by monotypy and original designation.

Etymology. Gender feminine; in honor of the Polish arachnologist Wojciech Staręga.

kirgizica Snegovaya, 2010 ; typ. loc.: Kyrgyzstan, Alash Mountain Ridge and River valley, near Alash. Starengovia kirgizica Snegovaya, 2010: 352 .

Starengovia kirgisica — Snegovaya, 2010: 353. Remarks. The spelling is hereby fixed to kirgizica (ICZN 24.2.3).

Genus Vestiferum Martens, 2006 (2 species)

Vestiferum Martens, 2006: 175 ; type species Nemastoma funebre Redikorzev, 1936 by original designation.

Remarks. Martens (2006) united two species in Vestiferum , which he considered as not very closely related. He mentioned affinities of alatum to the Turkish species of the heterogenous Pyza .

Etymology. Gender neuter; Latin “vestis” (dress, blanket) and the ending „ferum“ (bearing) referring to the soil incrusted body, which is unusual within Nemastomatidae .

alatum Martens, 2006: 176; loc. typ.: Georgia, Ajaria, Kintrish State Reserve, Zeraboseli.

funebre (Redikorzev, 1936) ; loc. typ.: Abchasia, Gagra, Kvemo-Ažara near Tkvarčeli.

Nemastoma funebre Redikorzev, 1936: 36 ;— Šilhavý 1966a: 69; Staręga 1966a: 391; 1978: 203.

Nemastoma brunneum Redikorzev, 1936: 35 ;—syn. Staręga 1978: 203.

Vestiferum funebre — Martens 2006: 175.

Remarks. As both names were published in the same paper, Staręga (1978; as first revising author) gave funebre precedence over brunneum (ICZN 24.2.2).