Tegenaria pagana, C. L. KOCH, 1840

Bolzern, Angelo, Burckhardt, Daniel & Hänggi, Ambros, 2013, Phylogeny and taxonomy of European funnel-web spiders of the Tegenaria-Malthonica complex (Araneae: Agelenidae) based upon morphological and molecular data, Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 168 (4), pp. 723-848 : 805-806

TEGENARIA PAGANA C. L. KOCH, 1840 , STAT. REV.

( FIG. 23K–W )

Tegenaria pagana C. L. Koch, 1841: 31 , 32, pl. 262, figs 612, 613.

Tegenaria subtilis Simon, 1870: 275–277 ; Simon, 1873: 170, pl. 1, figs 9, 10, male; Thorell, 1875a: 77.

Tegenaria testacea Simon, 1870: 278–280 , male; female belongs to Teg. domestica (see Machado, 1941).

Tegenaria proxima Pickard-Cambridge, 1873: 217 , 218, male.

Tegenaria variata Thorell, 1875c: 74 , 75, female.

Tegenaria urbana Simon, 1875: 67–69 .

Tegenaria bidentata Keyserling, 1878: 597–599 , pl. 14, fig. 19, male.

Tegenaria modesta Keyserling, 1878: 594–597 , pl. 14, fig. 18, female (male, fig. 17, is a synonym of Teg. domestica ), syn. nov.

Tegenaria obscura Banks, 1898: 230 , pl. 14, fig. 26; synonymized by Roth (1956: 176).

Tegenaria pagana cavernicola Simon, 1907: 547 , 548.

Tegenaria pagana proxima: Strand, 1909: 585 , 586.

Tegenaria antrias Crosby, 1936: 2 , pl. 1, fig. 3, female; Roth, 1952: 284, 285, synonymized by Roth (1956: 176).

Tegenaria simplex Bryant, 1936: 90 , 91, pl. 3, fig. 9, female.

Tegenaria pagana urbana: Simon, 1937: 1010, 1041 , syn. nov.

Tegenaria castro Chamberlin & Ivie, 1942: 21 , 22, pl. 3, figs 27–29.

Philoicides pallidus de Mello-Leitão, 1944: 335 , fig. 21, female; Roth, 1967: 314, pl. 51, fig. 3; synonymized by Ramirez, Grismado & Blick (2004).

Tegenaria cerrutii Roewer, 1960: 91–93 , fig. 2a–f, female; Brignoli, 1971a: 119, 120, figs 78, 79, syn. nov.

Tegenaria marinae Brignoli, 1971a: 120 , 121, figs 80, 81, female; Brignoli, 1977a: 50, figs 29, 30, syn. nov.

Tegenaria baronii Brignoli, 1977a: 47–50 , figs 25, 26, female, syn. nov.

Malthonica pagana: Guseinov et al., 2005: 164 .

Types

Syntypes. Greece: ‘aus der Gegend von Nauplia’, 1 ♂, 1 ♀ ( NHML, BM 1917.1.4.98), unknown .

Sub Tegenaria pagana urbana: Probable syntypes. France: Gallia, 9 ♂, 16 ♀ ( MNHN, 1982) .

Sub Tegenaria castro : Paratypes. USA: California: Los Angeles , 1 ♂, 1 ♀ ( AMNH), 3.v.1936 , Grant; Friant , 1 ♀ ( AMNH), 1.iii.1913 .

Sub Tegenaria cerrutii : Holotype. Italy: Sicily: Palermo, Mt. Pelegrino , Grotta Addaure (Adura?), ♀ ( SMF, 13374), Cerruti.

Sub Tegenaria marinae : Holotype. Italy: Lazio: Latina, Grotta Valmarino , ♀ ( MCSN, 543), 31.iii.1970, Sbordoni.

Sub Tegenaria baronii : Holotype. Italy: Marche: Fabriano, Grotta Frasassi , ♀ ( MCSN, 102), ix.1964, Baroni.

Other material examined

Croatia (2 ♀) ; France (2 ♂, 9 ♀) ; Greece (1 ♂, 7 ♀) ; Italy (9 ♂, 49 ♀) ; Malta (3 ♀) ; Portugal (1 ♂, 10 ♀) ; Spain (5 ♂, 19 ♀) ; Switzerland (1 ♀) . Africa: Algeria (4 ♂, 17 ♀) ; Cape Verde (1 ♂, 1 ♀) ; Egypt (3 ♂, 16 ♀) ; Libya (1 ♀) ; Morocco (2 ♂, 3 ♀) ; Tunisia (2 ♂, 2 ♀) . Asia: Syria (1 ♂, 4 ♀) . South America : Chile (6 ♂, 6 ♀) . North America : USA (1 ♂, 1 ♀) .

Description

Several descriptions under different names, some including measurements, were provided by Roth (1952), Brignoli (1971a, 1977a), and Levy (1996). Additional information is provided here.

Measurements: Female (N = 1): CL 3.41, CW 2.66, STL 1.76, STW 1.59. Leg I (3.97, 1.38, 3.63, 3.72, 2.17), II (3.48, 1.29, 2.85, 3.29, 1.62), III (3.17, 1.10, 2.38, 3.12, 1.42), IV (4.09, 1.29, 3.64, 4.44, 1.68).

Pedipalp (1.48, 0.62, 0.88, 1.36). EPL 0.22, EPW 0.54, ATL 0.14, ATW 0.39. Eyes: PME 0.17, PLE 0.18, AME 0.14, ALE 0.19. Eye distances: PME–PME 0.5–1 x PME, PME–AME 1 x PME, PME–PLE 0.5–1 x PME, PME–ALE 0.5–1 x PME, AME–AME <0.5 x AME, AME–ALE <0.5 x AME. CLY1 1.5–2 x AME, CLY2 0.5–1 x ALE.

Male palp: RTA with two branches, lateral branch leaf-shaped and distinctly stepped, distally protruding, dorsal branch strongly sclerotized and distally pointed, broad. Filiform embolus shorter than 2 x CB, originating at 8 o’clock position, distal tip at 3 o’clock position. Conductor almost orthogonal to cymbium with distal portion elongated and bent posteriad, lateral margin completely folded. Terminal end bifid, ventral part massive, pointed, dorsal part indistinctly pointed. Connection of conductor to tegulum moderately sclerotized. MA originating at 4–5 o’clock position, strongly protruding, distally with finger-shaped, distally spoon-like, sclerite. MA membranously connected to tegulum. Basal part of tegulum visible with strongly sclerotized band-like structure.

Epigyne and vulva: Epigynal plate strongly sclerotized and oval or kidney-like shaped, protruding. Posterior sclerite absent. Laterally with crescent-shaped pockets. CO are situated in these pockets, inside each of which an additional orthogonal pocket originates. Epigynal teeth absent (indistinct denticles present in the holotypes of Teg. marinae and Teg. baronii ). Vulva consists of CBD, no distinct RC recognizable. First half (CD) of CBD convoluted around second half, both parts strongly sclerotized. FD only represented by small, leaf-shaped appendages.

Other important characters: Cheliceral promargin with four, retromargin with four to seven teeth. Colulus developed as trapezoidal plate with the distal margin medially notched. Same pattern of distal spigots on PMS (in females) as described for the type species. PLS with distal segment as long as basal segment. Tarsal trichobothria on cymbium and palp tarsus absent. Tarsal trichobothria six to nine. Small teeth on paired claws of leg I nine to 11. Leg spination: male palp (2–0–0–0 or 3–0–0–0, 2–0–0, 1–2–0– 0), female palp (3–0–0–0, 2–0–0, 2–2–0–0), leg femora (2–2–2–0 or 2–3–1–0 or 2–3–2–0 or 2–4–3–0 or 2–5– 3–0 or 2–5–4–0, 2–2–1–0 or 2–2–2–0 or 2–3–2–0 or 2–3–3–0 or 2–4–2–0, 1–1–1–0 or 1–1–2–0 or 2–1–1–0 or 2–2–2–0 or 2–3–2–0, 1–1–1–0 or 2–1–1–0), patellae (all 2–0–0), tibiae [0–2–0–2p or 0–2–0–2p+1 or 0–2– 1–2p or 0–2–1–3p or 0–2–2–3p or 0–2–1–2p+1 (indistinct dorsal spines possible), 0–2–0–1p or 0–2–1– 1+1p+1 or 0–2–2–1+1p+1 or 0–2–2–1+2p or 0–2–2– 1p+1 or 0–2–2–3p (indistinct dorsal spines possible), 2–2–1–1 or 2–2–2–1+1p or 2–2–2–2+1p or 2–2–2–3 or 2–2–2–3p, 2–2–2–2 or 2–2–2–2+1p or 2–2–2–1+2p], metatarsi (0–0–0–3p+1, 0–1–0–3p+1 or 0–1–1–3p+1 or 0–2–1–3p+1, 0–3–2–3p+1 or 0–3–3–3p+1, 0–3–3– 3p+1), tarsi (all 0).

Coloration: Carapace margin narrow, continuous, indistinctly darkened; dorsally with two indistinct symmetrical longitudinal bands, head region darkened. Sternum with distinct pale median band and three symmetrical pairs of pale dots laterally, the last pair strongly fused with the median band. Opisthosoma brown-yellowish, anteriorly with pale median band, continuing in broad chevrons posteriad, laterally with dark band or moderately mottled. Legs moderately annulated (in the type specimens of Teg. baronii and Teg. marinae not visible, may be a result of alcohol preservation). ALS indistinctly darkened or pale, PLS with basal segment darkened, distal segment pale.

Distribution

Reported from southern and Central Europe, Mediterranean region, and introduced into North and South America.

Discussion

As in other Tegenaria species , Teg. pagana shows great variation in somatic and genital characters (e.g. Fig. 23P–W ), which is reflected in the long list of synonyms. This phenomenon was well illustrated by Levy (1996: 97, 98, figs 45–48). He mentioned that the Tegenaria pagana -complex of Brignoli (1971a, 1977a) may represent this variation. The small posterior denticles on the epigyne of Teg. marinae are also found in some specimens of otherwise characteristic Teg. pagana and may reflect this variation. Here, we follow Levy and regard Teg. cerrutii Brignoli, 1971 , Teg. marinae Brignoli, 1971 , and Teg. baronii Brignoli, 1977 , as junior synonyms of Teg. pagana C. L. Koch, 1940 .

Figures

Figure 23. A–D, Tegenaria tridentina; E-H, Tegenaria mirifica; I, J, Tegenaria levantina; K-W, Tegenaria pagana, including the type specimens of Tegenaria cerrutii (R, S), Tegenaria marinae (T, U), and Tegenaria baronii (V, W). Left male palp in ventral (A, E, K) and retrolateral views (B, F, L); epigyne in ventral (C, G, I, P, R, T, V) and vulva in dorsal view (D, H, J, Q, S, U, W); chelicerae in ventral view (O); face of female in frontal (N) and sternum in ventral view (M). Scale bars = 0.5 mm (T–W without scale).