Hoplopholcus forskali ( Thorell, 1871 )

Huber, Bernhard A., 2020, Revision of the spider genus Hoplopholcus Kulczyński (Araneae, Pholcidae), Zootaxa 4726 (1), pp. 1-94 : 44-54

Hoplopholcus forskali ( Thorell, 1871)

Figs 194–197 , 200–245

Pholcus Forskålii Thorell, 1871: 151 (♂ ♀).

Pholcus Forskalii— Chyzer & Kulczyński 1891: 149, pl. 6, figs 18a, b.

Holocnemus Forskalii— Chyzer & Kulczyński 1896: 11. Chyzer & Kulczyński 1897: 310. Damin 1900: 21.

Hoplopholcus Forskålii — Kulczyński 1908: 63.

Holocnemus forskali — Drensky 1936: 55.

Pholcus (Hoplopholcus) forskali— Drensky 1929: 28.

Haplopholcus [sic] Forskali — Dudich 1933: 126.

Haplopholcus forskalli [sic]— Fuhn & Niculescu-Burlacu 1970: 415, fig. 11.

Hoplopholcus Forskali — Loksa 1969: 70, figs 46, 47 c–d.

Hoplopholcus forskoeli — Brignoli 1976: 561 , figs 34–36, 38, 40–41. Brignoli 1978: 490, fig. 58.

Hoplopholcus forskalii — Nicolić & Polenec 1981: 20 . Senglet 2001: 62, figs 6–7.

Hoplopholcus forskali — Stojićević 1929: 16 . Drensky 1939: 248, fig. 22, map 3 (but see Distribution below). Szinetár et al. 1999: 162. Ćurčić et al. 1999: 8 P. Deltshev et al. 2003: 12. Kenyeres & Szinetár 2003: 57. Ćurčić et al. 2004: 104. Kovács et al. 2006: 13. Kovács & Szinatár 2016: 171, figs 1–2. Moscaliuc 2012: 11. Eberle et al. 2018 (molecular data). Huber et al. 2018: fig. 6.

Probable misidentifications (see Distribution below)

Hoplopholcus forskali: Drensky 1939 : map 3 (at least records from Crete, Iraq, and Turkmenistan shown in map), Caporiacco 1949: 123.

Holocnemus Forschkali : Kolosváry 1938: 65.

Pholcus Forskåli : Kolosváry 1939: 133.

Diagnosis. Easily distinguished from known congeners by shape of procursus ( Figs 212–213 ): very long distal part (i.e. part beyond attachment site of ventral spine), long and slender transparent process, small process between ventral spine and transparent process (similar in H. minotaurinus ), and protruding spiny membrane distally on prolateral side. Females are difficult to distinguish externally from congeners; median pouch of uterus externus folded toward anteriorly producing distinctive duplicature of cuticle ( Figs 204 , 207, 210 , 220 ) that is usually visible in uncleared specimens; pair of lateral pouches of uterus externus usually with sclerotized external margins clearly visible in uncleared specimens ( Figs 203 , 206, 209 ).

Type material. Unknown number of male and female syntypes, depository unknown (see Note below), from “ Hungary (the Banat)” (see Note below), date not given, leg. L. von Kempelen; not examined.

Notes. The type locality “Banat” was in the 1860ies part of the Habsburg Kingdom of Hungary but is currently divided among three countries: the eastern part lies in western Romania, the western part in northeastern Serbia, and only a small northern part lies within southeastern Hungary (B in Fig. 443 ). The exact type locality is thus unknown. Since no other Hoplopholcus species seems to occur in this region, the uncertainty about the type locality appears unproblematic.

The NHMW has a vial with 4♂ 4♀ 2 juvs that were collected by Kempelen in 1866 (examined). This might be topotypical material even though both labels in the vial give “ Wien ” as only locality information and both give a wrong identification (“ Holocnemus pluchei ”). I have not seen any Hoplopholcus forskali specimen that reliably originates from Austria and the locality information on the label may be erroneous. Since the identity of the species is beyond doubt, the uncertainty about the type specimens also appears unproblematic.

Other material examined. HUNGARY, Veszprém: 1♀ in pure ethanol, ZFMK (Ast 4), Balaton Lake, Tihany Peninsula (46.917°N, 17.883°E), Levendulas, under rocks at shore, 13.ix.2003 (J.J. Astrin) . Budapest: 1♂ 2♀, ZFMK (Ar 20936), Molnár János Cave [47.518°N, 19.036°E], 105 m a.s.l., 13.ix.2000 (F. Gasparo) . Csongrad: 1♂ 3♀ in pure ethanol, ZFMK (Kov 1), Bordány (46.322°N, 19.922°E), in building, 15.i.2014 (G. Kovacs) . 1♂ in pure ethanol, ZFMK (Kral 150), Vac (47.796°N, 19.140°E), 7.vi.2019 (J. Král) .

SLOVAKIA, Nitra: 1♂, ZFMK (Ar 20937), Nitra City, Marianska Street [48.312°N, 18.084°E], in house, 6.vi.2017 (M. Fedoriak) . 1 juv., ZFMK (Ar 20938), Nitra City, Levická Street [48.310°N, 18.122°E], in house, 13.vi.2017 (M. Fedoriak) .

SERBIA, South Banat: 2♂ 3♀, MHNG, Alibunar [45.080°N, 20.967°E], 26.v.1972 (A. Senglet) .

ROMANIA, Tulcea: 1♂ 1♀ 5 juvs (2 vials), MGAB, Letea (45.281°N, 29.525°E), 2013 (L.A. Moscaliuc) .

Prahova: 1♂ 4♀ 3 juvs (2 vials), MGAB, Urleta (45.084°N, 25.795°E), 1954/1955, collector not given .

BULGARIA, Dobrich: 1♂, ZMMU, Krushari Municipality, Dunav (= Danube ) River valley , Kapitan Dimitrovo (43.953°N, 27.692°E), 120 m a.s.l., buildings, 20.viii.2005 (A.V. Gromov) . Lovech: 2♂ 1♀ 1 juv., ZMB (48585), Drenski [42.879°N, 24.779°E], “durch Prof. Deckert”, no further data . Vidin: 2♂ 1♀ 1 juv., NMNHS, “Belogradchik” [should be Chuprene Municipality], Varbovo village [~ 43.544°N, 22.650°E], Leponishki Pech, cave, 3.ix.1973 (“ P.B. ”) . Pazardzhik: 1♂ 1♀, NMNHS, Panagyurïşte, Fetenci (42.560°N, 24.208°E), 20.iii.2010 (S. Lazarov) . Blagoevgrad: 1♀, ZMMU, Sim- itli Municipality, N foothills of Krupnishna Planina Mt. Range , ~ 0.2 km NNE Krupnik (41.851°N, 23.126°E), 375 m a.s.l., hill, “under stones and building”, 12.viii.2005 (A.V. Gromov) . 3♀, ZMMU, Simitli Municipality, N foothills of Krupnishna Planina Mt. Range , ~ 1.1 km SSE Charnichna [Cherniche] (41.848°N, 23.135°E), 290 m a.s.l., steppe and meadow, un- der stones and in grass, 13–14.viii.2005 (A.V. Gromov) . 1♀, ZMMU, Razlog Municipality, S env. of Banya (41.877°N, 23.527°E), 850 m a.s.l., hill, Pinus forest, 8.viii.2005 (A.V. Gromov).

TURKEY, Konya: 1♂, MHNG, Lake Beyşehir, Haci Akif Island [37.62°N, 31.48°E], 24.iv.1973 (P. Brignoli) . 1♂ 1♀, ZFMK (Ar 20939), and 1♂ 1 juv. in pure ethanol, ZFMK (Tur 41), Beyşehir District, at Lake Beyşehir (37.621°N, 31.452°E), 1125 m a.s.l., among rocks near road, 21.vii.2016 (H. Öztürk) . 1♂ 1♀, ZFMK (Ar 20940), and 1♀ in pure ethanol, ZFMK (Tur 40), Aksu District, Zindan Mağarası (37.812°N, 31.085°E), 1300 m a.s.l., among rocks outside of cave, 21.vii.2016 (H. Öztürk) . 3♂ 2♀, ZFMK (Ar 20941), and 2♀ 1 juv. in pure ethanol, ZFMK (Tur 45), Seydişehir District, forest below Ferzene Mağarası (37.383°N, 31.837°E), 1230 m a.s.l., under rocks, 23.vii.2016 (H. Öztürk) . 3♂ 8♀, ZFMK (Ar 20942), and 1♂ 1♀ 1 juv. in pure ethanol, ZFMK (Tur 50), Taşkent District, Taşkent (36.922°N, 32.493°E), 1450 m a.s.l., under rocks in abandoned gardens, 25.vii.2016 (H. Öztürk) .

Isparta: 1♀, MHNG, 10 km N Isparta [~ 37.85°N, 30.53°E], no further data .

Bilecik: 1♂, NOHUAM, İnhisar, İnpazarcık Mağarası (40.04°N, 30.38°E, 240 m a.s.l.), 15.ix.2011 (A. Top- çu) .

Kütahya: 1♀, NOHUAM, İnli, İnli Mağarası (39.48°N, 30.21°E, 1085 m a.s.l.), 1 1.ix.2011 (A. Topçu) .

UNCLEAR/UNIDENTIFIED LOCALITIES: 1♂ 1♀, NHMW, “Palanka”, “Inv. No. 374”, no further data. 1♂, NHMW, Hungary, “A.D. 1884, A.N.I. 258”, “Inv. No. 375”, no further data. 1♂ 2♀, MNHN ( AR 10324), E. Simon collection number 2090, “Hungaria (Herm.)”, no further data.

Redescription. Male (Budapest, Molnár János Cave, ZFMK). MEASUREMENTS. Total length 5.3, carapace width 2.0. Distance PME-PME 120 µm; diameter PME 140 µm; distance PME-ALE 40 µm; diameter AME 85 µm; distance AME-AME 30 µm. Leg 1: 30.4 (8.3 + 0.8 + 8.4 + 10.9 + 2.0), tibia 2: 5.9, tibia 3: 4.4, tibia 4: 5.3; tibia 1 L/d: 42.

COLOR (in ethanol). Carapace mostly pale ochre-yellow, medially with light brown band including ocular area; clypeus light brown; sternum medially dark brown, lighter brown toward margins; legs ochre-yellow, with slightly darker brown rings subdistally on femora and tibiae and in patella area; abdomen gray, with regular pattern of oblique dark marks in posterior third above spinnerets, ventrally with diffuse dark pattern.

BODY. Habitus as in Figs 194–195 . Ocular area slightly elevated. Deep thoracic pit and indistinct pair of furrows diverging from pit toward posterior margin. Clypeus unmodified. Sternum wider than long (1.0/0.9), unmodified. Abdomen oval, dorso-posteriorly rounded. Gonopore in scanned male with six epiandrous spigots ( Fig. 232 ). ALS with only two spigots each ( Fig. 224 ).

CHELICERAE. As in Figs 214–215 , with pair of latero-distal apophyses provided with three modified coneshaped hairs each ( Figs 228–231 ); with fine stridulatory ridges.

PALPS. As in Figs 200–202 ; coxa with low retrolateral bulge, trochanter barely protruding ventrally, femur with retrolateral dark line and prolateral stridulatory pick; palpal tarsal organ exposed ( Fig. 226 ); procursus ( Figs 212–213 ) with small ventral ‘knee’, with distinctively long distal element, long transparent process and long and straight ventral spine ( Fig. 234 ), with small pointed process between ventral spine and transparent process, with hair-like distal processes ( Fig. 235 ), on prolateral side with protruding spiny membrane ( Figs 237–238 ), possibly with glandular pores (arrows in Fig. 239 ); genital bulb ( Figs 216–219 ) with distally widened ventral sclerite, embolar sclerite with small cone-shaped processes ( Fig. 240 ); with distinct dorsal membranous process ( Fig. 217 ).

LEGS. Femur 1 with single row of ~28 ventral spines ( Figs 242–243 ); without curved hairs (many hairs miss- ing), few vertical hairs; retrolateral trichobothrium of tibia 1 at 4%; prolateral trichobothrium present on all leg tibiae; tarsi without distinct pseudosegments but with many small platelets ( Fig. 244 ).

Male (variation). Tibia 1 in 25 other males: 7.1–11.3 (mean 9.1). Most males with two or three modified hairs on each cheliceral apophysis, sometimes asymmetric; rarely with four modified hairs, in one male with 3+5 modified hairs. Dorsal abdominal marks often also present in anterior part. Some males with spines also on femur 2 (up to 20; thinner than on femur 1). Some males with very few curved hairs on metatarsus 1 only (weakly curved). Small pointed process between ventral spine and transparent process slightly larger in Turkish specimens.

Female. In general similar to male ( Figs 196–197 ) but without spines on legs and with capsulate palpal tarsal organ ( Fig. 227 ). ALS as in male ( Fig. 225 ). Tibia 1 in 31 females: 6.8–10.7 (mean 8.1).

FEMALE GENITALIA. Epigynum as in Figs 203 , 206, 209 , posterior rim medially whitish, internal sclerotized arcs and pair of lateral pouches of uterus externus usually visible in uncleared specimens; with pair of light brown weakly bulging areas in front of epigynum and short but wide posterior plate. Internal genitalia with very large pore plates at slightly variable distances ( Figs 205 , 208, 211 , 221 ), median pouch of uterus externus folded toward anteriorly producing duplicature of cuticle ( Figs 204 , 207, 210 , 220 ); lateral pouches of uterus externus with sclerotized lateral margins, apparently forming curved ridges rather than pockets.

Distribution. Widespread, ranging from eastern Central Europe to Turkey ( Fig. 443 ). Most old records whose specimens I could not study are credible. Almost certainly erroneous are the records from Crete, Turkmenistan, and Iraq shown in the map of Drensky (1939: 249). Drensky never specified the sources for these dots in the map, and all available data strongly suggest that the species does not occur in these places. Dubious are also the records from the Adriatic Coast ( Kolosváry 1938, 1939; Caporiacco 1949); large recent collections from the Adriatic Coast include the similar genus Stygopholcus , but never Hoplopholcus (B.A. Huber, unpublished data). Other records known to me, of specimens I have not seen, are listed in Table 1 . All coordinates in this table are approximate, derived from Google Earth and other online sources.

Natural history. In eastern Central Europe, this species is mostly found in and around buildings, but it seems to be less tolerant than Pholcus phalangioides against the conditions in rooms constantly occupied by humans ( Kovács & Szinetár 2016). At the same time it seems to tolerate higher luminance and lower temperature ( Kovács & Szinetár 2016). The same authors also reported that males and females sometimes share webs; copulation takes place between April and August; egg-laying takes place from May to September; egg-sacs contain about 30– 36 eggs; the time between copulation and egg-laying is approximately 29 days; the dome-shaped web has a diameter of about 55 cm and is more stable than the web of P. phalangioides .

In Turkey, all specimens were collected in natural habitats, among and under rocks near the ground. They built large webs with a diameter of up to 40 cm, part of which protruded from under the rock. Even at light disturbance, the spiders fled toward the back and were very difficult to catch unless the rock was small enough to be moved.

Figures

FIGURE 443. Known distribution of Hoplopholcus forskali. B = the historical region “Banat” (type locality). Green dots: records confirmed. Red dots: credible records from the literature.

FIGURES 194–199. Hoplopholcus spp., live specimens. 194–197. H. forskali (Thorell, 1871), males from Taşkent, females from Zindan Mağarası (before and after egg-laying). 198. H. konya sp. n., female from Körükini Mağarası. 199. H. longipes (Spassky, 1934), male from Kintrishi Protected Area.

FIGURES 200–205. Hoplopholcus forskali (Thorell, 1871) from Hungary, Molnár János Cave (ZFMK Ar 20936). 200–202. Left male palp in prolateral, dorsal, and retrolateral views. 203–205. Epigynum, ventral view, and cleared female genitalia, ventral and dorsal views.

FIGURES 206–211. Hoplopholcus forskali (Thorell, 1871); epigyna, ventral views, and cleared female genitalia, ventral and dorsal views. 206–208. Bulgaria, Fetenci (NMNHS). 209–211. Turkey, Taşkent (ZFMK, Ar 20942).

FIGURES 212–221. Hoplopholcus forskali (Thorell, 1871) from Hungary, Molnár János Cave (ZFMK Ar 20936). 212–213. Left procursus, prolateral and retrolateral views. 214–215. Male chelicerae, frontal and lateral views. 216–219. Left genital bulb, dorsal, prolateral, ventral, and retrolateral views. 220–221. Cleared female genitalia, ventral and dorsal views. mp, median pouch of uterus externus; lp, lateral pouch of uterus externus. Scale lines: 0.5 mm.

FIGURES 222–227. Hoplopholcus forskali (Thorell, 1871) (ZFMK Ar 20942). 222. Female carapace and ocular area, frontal (slightly dorsal) view. 223. Male eyes and ‘pseudo-eyes’ (arrows). 224–225. Male and female ALS. 226–227. Male and female palpal tarsal organs. Scale lines: 10 µm (224–227), 20 µm (223), 100 µm (222).

FIGURES 228–233. Hoplopholcus forskali (Thorell, 1871) (ZFMK Ar 20942). 228–229. Right and left male cheliceral apophyses (of same specimen—note asymmetry of modified hairs). 230–231. Distal and subdistal modified hairs on left male cheliceral apophysis. 232. Male gonopore with epiandrous spigots. 233. Stridulatory ridges on female chelicera. Scale lines: 2 µm (231), 10 µm (228–230, 233), 20 µm (232).

FIGURES 234–239. Hoplopholcus forskali (Thorell, 1871) (ZFMK Ar 20942). 234. Right procursus tip, retrolateral view. 235–236. Details of Fig. 234: hair-like processes and detail of transparent process. 237. Left procursus tip, distal view. 238. Left procursus tip, ventral view (arrows in Figs 237–238 point at process between ventral spine and transparent process). 239. Prolateral spiny membrane (arrows point at possible gland openings). psm, prolateral spiny membrane; tp, transparent process; vs, ventral spine. Scale lines: 10 µm (235–236, 239), 100 µm (234, 237–238).

FIGURES 240–245. Hoplopholcus forskali (Thorell, 1871) (ZFMK Ar 20942). 240. Right bulbal processes, distal view (dmp, dorsal membranous process; es, embolar sclerite;vs, ventral sclerite;arrow points at sperm duct opening).241.Tarsal organ on female leg 4.242–243.Spines on male femur 1. 244. Detail of male tarsus 1. 245. Claws on male tarsus 4. Scale lines:10 µm (241, 244–245), 100 µm (240, 242–243).

Tables

TABLE 1. Credible but unchecked records of Hoplopholcus forskali shown in Fig. 443. Locality Latitude Longitude Reference

Locality	Latitude	Longitude	Reference
Hungary: Kecskemét	46.892	19.690	Chyzer & Kulczyński 1891
Hungary: Körmend	47.014	16.595	Szinetár 1992
Hungary: Szobathely	47.230	16.620	Szinetár 1992
Hungary: Györ	47.680	17.650	Szinetár 1992
Hungary: Balatongyörök	46.760	17.350	Szinetár et al. 1999
Hungary: Szentbékkálla	46.887	17.564	Szinetár et al. 1999
Hungary: Tapolca	46.883	17.438	Szinetár et al. 1999
Hungary: Badacsonylábdihegy	46.790	17.475	Szinetár et al. 1999
Hungary: Révfülöp	46.829	17.627	Szinetár et al. 1999
Hungary: Szigliget	46.793	17.438	Szinetár et al. 1999
Hungary: Bezedek	45.864	18.585	Kovács & Szinetár 2016
Hungary: Dorog	47.725	18.729	Kovács & Szinetár 2016
Hungary: Komló	46.194	18.252	Kovács & Szinetár 2016
Hungary: Pálköve	46.825	17.594	Kovács & Szinetár 2016
Hungary: Pécs	46.073	18.230	Kovács & Szinetár 2016
Hungary: Salföld	46.834	17.551	Kovács & Szinetár 2016
Hungary: Sikondá	46.181	18.215	Kovács & Szinetár 2016
Hungary: Paloznak	46.983	17.942	Kovács & Szinetár 2016
Hungary: Csorna	47.610	17.245	Kovács & Szinetár 2016
Hungary: Tarcal	48.131	21.343	Kovács & Szinetár 2016
Slovakia: Tekovské Lužany (“Nagysalló”)	48.100	18.540	Dudich 1933
Slovakia: Zombor	48.136	19.448	Herman 1879 in Kovács & Szinetár 2016
Serbia: Jasenak1	44.620	20.252	Chyzer & Kulczyński 1891, Damin 1900
Serbia: Zemun	45.840	20.400	Damin 1900
Serbia: Potporanj	45.020	21.245	Chyzer & Kulczyński 1891
Serbia: Vranić	44.600	20.325	Ćurčić et al. 2004
Serbia: Monastery Caves, near Minićevo	43.700	22.352	Deltshev et al. 1997, Ćurčić et al. 1999
Serbia: Bela Palanka	43.220	22.310	Stojićević 1929
Serbia: Belgrade	44.792	20.439	Stojićević 1929
Serbia: Leskovac	43.000	21.940	Stojićević 1929
Serbia: Niš	43.323	21.900	Stojićević 1929
Serbia: Šabac	44.749	19.693	Stojićević 1929
Serbia: Senjak	44.792	20.440	Stojićević 1929
Serbia: Valjevo (“Walewo”)	44.275	19.885	Stojićević 1929
Serbia: Vranje	42.550	21.900	Stojićević 1929
Serbia: Stara Pazova	44.990	20.160	Chyzer & Kulczyński 1891
Croatia: Osijek (“Eszék”)	45.555	18.695	Chyzer & Kulczyński 1896, Damin 1900
Macedonia: Štip	41.740	22.200	Stojićević 1929
Macedonia: Prilep	41.345	21.553	Drensky 1929, Stojićević 1929
Macedonia: Skopje	41.995	21.430	Drensky 1929
Macedonia: Bitola	41.030	21.330	Drensky 1929
Macedonia: Veles	41.714	21.771	Drensky 1929, Stojićević 1929
Macedonia: Babuna	41.680	21.802	Stojićević 1929
Romania: Rekaş	45.800	21.510	Chyzer & Kulczyński 1891
Romania: Oradea (“Nagyvárad”)	47.060	21.920	Chyzer & Kulczyński 1891
Romania: Reșița	45.300	21.880	Chyzer & Kulczyński 1891
Romania: Drenkova	44.650	21.950	Chyzer & Kulczyński 1891
Romania: Buzad	45.940	21.547	Chyzer & Kulczyński 1891
Romania: Orşova	44.726	22.392	Herman 1879 in Kovács & Szinetár 2016
Romania: Sviniţa-Orşova	44.600	22.250	Fuhn & Niculescu-Burlacu 1970
Bulgaria: Vratsa distr., Cherepish vill., Studenata cave	43.101	23.615	Brignoli 1976