Tegenaria achaea, BRIGNOLI, 1977

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 : 775-779

TEGENARIA ACHAEA BRIGNOLI, 1977

( FIG. 15A, B )

Tegenaria achaea Brignoli, 1977b: 945 , fig. 4, female.

Types

Holotype and paratype. Greece: North Aegean: Ikaria, ‘grotte Phutra to Nao , dans le massif Messaria près Petropoulion’ , 2 ♀ ( MHNG, Hel-75/27), 24.iv.1975, Hauser .

KEY TO EUROPEAN TEGENARIA SPECIES

Only European species are included in the key. The following species are excluded as material was unavailable for examination or insufficient: Tegenaria animata Kratochvíl & Miller, 1940, Tegenaria bayeri Kratochvíl, 1934, Tege- naria bosnica Kratochvíl & Miller, 1940, Tegenaria chumachenkoi Kovblyuk & Ponomarev, 2008, Tegenaria decolorata Kratochvíl & Miller, 1940, Tegenaria maelfaiti Bosmans, 2011, Tegenaria oribata Simon, 1916, Tegenaria podoprygorai (Kovblyuk, 2006), Tegenaria scopifera Barrientos, Ribera & Pons, 2002, and Tegenaria taurica Charitonov, 1947.

As material was unavailable, the female of Tegenaria lapicidinarum Spassky, 1934, and the male of Tegenaria levantina Barrientos, 1981, are not included in the key.

1. Male..............................................................................................................................................2

-. Female.........................................................................................................................................35

2. Row of several dorsal trichobothria on cymbium....................................................................................3

-. Trichobothria on cymbium absent......................................................................................................12

3 Tegular apophysis between tegulum and conductor present (e.g. Fig. 15G) ................................................. 4

-. Tegular apophysis absent .................................................................................................................. 6

4. Tegular apophysis protruding only retrolaterad, ventral branch of RTA with indistinct rim and moderately protruding bulge ..................................................................................................... Tegenaria eleonorae

-. Tegular apophysis protruding also distally, ventral branch of RTA with distinct longitudinal rim...................5

5. Ventral branch of RTA with long, drawn-out rim, almost as long as palp tibia, distal branch of conductor reaching two thirds of cymbium tip, distal segment of PMS shorter than basal segment.................. Tegenaria armigera

-. Long, drawn-out rim of RTA relatively shorter than palp tibia, distal branch of conductor reaching one third of cymbium tip, distal segment of PMS longer than basal segment .................................... Tegenaria tyrrhenica

6. Dorsal branch of RTA with two to three elongated spine-like points ......................................................... 7

-. Dorsal branch of RTA different...........................................................................................................8

7. Distal portion of conductor transversal to cymbium...................................................... Tegenaria dalmatica

-. Distal portion of conductor parallel to cymbium............................................................. Tegenaria ramblae

8. Dorsal branch of RTA strongly protruding and pointed...........................................................................9

-. Dorsal branch of RTA distally truncated and stepped ........................................................................... 10

9. Ventral branch of RTA distally not protruding, terminal end of conductor bifid (retrolateral view)................... ............................................................................................................................ Tegenaria femoralis

-. Ventral branch of RTA distally tusk-like elongated and protruding, terminal end of conductor indistinctly bifid (retrolateral view).................................................................................................... Tegenaria annulata

10 Conductor much shorter than cymbium....................................................................... Tegenaria parietina

-. Conductor longer than three quarters of cymbium length...................................................................... 11

11. Lateral branch of RTA distinctly truncated, flat, ventral portion of terminal end of conductor truncated........... .......................................................................................................................... Tegenaria ferruginea

-. Lateral branch of RTA moderately pointed, longer than broad, ventral portion of terminal end of conductor pointed ................................................................................................................... Tegenaria carensis

12. Median apophysis with distal sclerite plate-, spoon-, or pocket-like ........................................................ 13

-. Median apophysis with distal sclerite forming a finger-, thorn-, or hook-like structure (can be in combination with a plate) ........................................................................................................................................ 21

13. RTA with dorsal and lateral branch equally long, dorsal branch distally truncated (Figs 14N, 16K, X), median apophysis long, band-like, protruding, distally moderately bent ventrad...................................................14

-. RTA and median apophysis differently shaped.....................................................................................15

14. Embolus short and distally truncated, distal portion of conductor only moderately elongated.......................... ........................................................................................................................... Tegenaria domestica

-. Embolus longer, filiform, distal portion of conductor longer than wide, conductor with bifid terminal end with very distinctively formed dorsal part ...................................................................... Tegenaria annae sp. nov.

15. Distal portion of conductor almost parallel to cymbium.........................................................................16

-. Distal portion of conductor transversal to cymbium, elongated posteriad, RTA with a row of small denticles..... ............................................................................................................................. Tegenaria argaeica

16. RTA with lobe-, bulge-, or rim-like dorsal branch.................................................................................17

-. RTA with pointed branch/branches .................................................................................................... 18

17. RTA with strongly protruding lobe- or rim-like dorsal branch, very distinctly formed terminal end of conductor, tegular apophysis absent, cymbium inconspicuous........................................................... Tegenaria hauseri

-. RTA with bulge-like branches, terminal end of conductor inconspicuously pointed, lamelliform tegular apophysis present, cymbium conspicuously modified and distally elongated...................................... Tegenaria ariadnae

18. Lateral branch of RTA shifted posteriad (originating median of the tibia), distant, strongly protruding, and pointed............................................................................................. Tegenaria vankeerorum sp. nov.

-. Lateral branch of RTA different........................................................................................................19

19. Conductor reduced to transparent lamelliform appendage.............................................. Tegenaria racovitzai

-. Conductor stronger sclerotized, not only lamelliform............................................................................20

20. RTA with two strongly sclerotized points, tegular apophysis absent.................................... Tegenaria hasperi

-. RTA with only one strongly sclerotized point, tegular apophysis expressed as pocket-like structure ................ .......................................................................................................................... Tegenaria rhodiensis

21. Ventral part of terminal end of conductor distinctly elongated and pointed (Fig. 18I, J, N, O), dorsal part complex...................................................................................................................................................22

-. Terminal end of conductor different................................................................................................... 24

22. RTA with massive protruding bulge, clearly visible in ventral view................ Tegenaria circeoensis sp. nov.

-. RTA lobe- or moderately bulge-like, may be distally bent ...................................................................... 23

23. RTA distally not bent, dorsal part of terminal end of conductor convex ‘rounded’............. Tegenaria parmenidis

-. RTA distally bent, dorsal part of terminal end of conductor concave or straight ................ Tegenaria sbordonii

24. Dorsal branch of RTA broadly elongated and protruding, distally bifid ( Fig. 17P )............................. Tegenaria campestris, Tegenaria bozhkovi, Tegenaria montana, Tegenaria rilaensis (see comment sub Teg. rilaensis)

-. Dorsal branch of RTA differently shaped or absent...............................................................................25

25. Lateral branch of RTA broad and strongly protruding, distally with a straight, truncated ventral part and a strongly elongated, finger-shaped dorsal protuberance, conductor and median apophysis strongly elongated and narrowly pointed, hook-like (Figs 14K–L, 15V–W)..................................... Tegenaria schoenhoferi sp. nov.

-. Lateral branch of RTA different, conductor and/or median apophysis not strongly elongated and narrowly pointed, hook-like.......................................................................................................................................26

26. Tegulum medially strongly protruding (Figs 19B–C, 20B, K) ..................................................................27

-. No distinct protuberance medially at the tegulum................................................................................28

27. Dorsal branch of RTA truncated, median protrusion of tegulum more or less constantly rounded (Fig. 20B), distal end of conductor moderately bent ventrad (lateral view)................................................. Tegenaria silvestris

-. Dorsal branch of RTA hook-like, pointed, and distally bent ventrad, median protrusion of tegulum bent distally (Fig. 20K), distal portion of conductor straight (lateral view)............................................. Tegenaria parvula

28. Embolus relatively short, not exceeding cymbium width or truncated......................................................29

-. Embolus long, filiform.....................................................................................................................31

29. Terminal end of embolus truncated .............................................................................. Tegenaria mirifica

-. Terminal end of embolus pointed ...................................................................................................... 30

30. Conductor with distinctly protruding dorsal portion, terminal end of conductor bifid, RTA with two protruding points............................................................................................................ Tegenaria mercanturensis

-. Dorsal portion of conductor not protruding, terminal end of conductor with only one point, RTA different. Tegenaria percuriosa

31. Conductor parallel to cymbium .........................................................................................................32

-. Dorsal portion of conductor transversal to cymbium......................................................... Tegenaria pagana

32. Distal branch of RTA distinctly leaf-shaped ............................................................... Tegenaria regispyrrhi

-. Distal branch of RTA different..........................................................................................................33

33. Dorsal branch of RTA hook-like, pointing anteriad, distinct rim at conductor present............. Tegenaria henroti

-. Dorsal branch of RTA pointed, straight, or distally bent ventroposteriad..................................................34

34. Dorsal branch of RTA pointed and distally bent ventroposteriad, terminal end of conductor bifid .................... ........................................................................................................................... Tegenaria tridentina

-. Dorsal branch of RTA pointed but not bent, terminal end of conductor with one point only............................ ..................................................................................................................... Tegenaria lapicidinarum

35. Row of several dorsal trichobothria present on palp tarsus....................................................................36

-. Trichobothria on palp tarsus absent...................................................................................................45

36. Epigyne with distinct band- or bar-like posterior sclerite with anterior margin concave.............................37

-. Epigyne without posterior sclerite or with sclerite with anterior margin convex........................................ 40

37. Posterior sclerite posteriorly protruding along its whole width (Fig. 16Q)............................ Tegenaria carensis

-. Posterior sclerite with straight posterior margin or only medially protruding............................................38

38. Posterior sclerite medially almost half as long as wide.................................................... Tegenaria ramblae

-. Posterior sclerite medially much shorter than wide.............................................................................. 39

39. Lateral margins of the median region of the epigyne follows distinctly the run of the posterior sclerite (especially anteriorly, Fig. 21J, white arrow), vulva strongly convoluted (in two spirals), separated by less than two duct diameters, more than three in Teg. parietina) ............................................................. Tegenaria ferruginea

-. Lateral margins of the median region of the epigyne runs toward the middle of the epigyne (Fig. 21P, white arrow), vulva less convoluted (especially first spiral), separated by more than three duct diameters ........................... ............................................................................................................................Tegenaria parietina

40. Epigyne with well-separated posterior sclerite, expressed as large plate or bulge.......................................41

-. Epigyne without posterior sclerite or with median plate distinctly connected to epigynal plate (anteriorly) or with broad opening posteriorly.................................................................................................................43

41. Posterior sclerite expressed as a large plate, copulatory openings laterally of this plate (Fig. 20Q)................... ............................................................................................................................ Tegenaria eleonorae

-. Posterior sclerite bulge-like..............................................................................................................42

42. Posterior sclerite globular bulge-like, copulatory openings laterally, vulva with strongly convoluted ducts, almost touching each other................................................................................................ Tegenaria tyrrhenica

-. Posterior sclerite trapezoidal bulge-like, copulatory openings anteriorly, vulva distinctly convoluted, well sepa- rated..................................................................................................................... Tegenaria annulata

43. Epigyne with broad posterior opening (Fig. 15I, white arrow).......................................... Tegenaria armigera

-. Epigyne median plate distinctly connected to epigynal plate (anteriorly).................................................. 44

44. Median plate moderately protruding, suboval, distinctly round, or oval copulatory openings anteriorly of median plate.................................................................................................................... Tegenaria dalmatica

-. Median plate not protruding, subrectangular, pocket-like copulatory openings visible anteriorly of median plate..................................................................................................................... Tegenaria femoralis

45. Very distinct epigyne, anteriomedially with a narrow rectangular septum, posteriorly with a semicircle-shaped pocket, opening anteriad ( Fig. 17D )............................................................................... Tegenaria hauseri

-. Different.......................................................................................................................................46

46. Epigyne with a distinct, strongly sclerotized transversal rim forming a pocket, opening posteriad, copulatory opening anteriolaterally of this structure (Fig. 24A, E, G, I)............ Tegenaria regispyrrhi (incl. aff. regispyrrhi)

-. Different.......................................................................................................................................47

47. Epigyne with protruding median plate, suboval, anteriolaterally with distinct pockets, opening medioposteriad (Fig. 23P, R, T, V) ...................................................................................................... Tegenaria pagana

-. Different.......................................................................................................................................48

48 Epigyne with a distinct posterior sclerite, expressed as bar-, bulge-, or plate-like structure.........................49

-. Posterior sclerite absent or strongly fused with epigynal plate ............................................................... 59

49. Very complex vulva with two receptaculi and convoluted duct, indistinct small epigynal teeth at the anteriolateral margin of median region sometimes present (Figs 15E–F, 18G–H) .................................... Tegenaria argaeica

-. Vulva less complex, no epigynal teeth or only ‘pseudo teeth’ present ....................................................... 50

50. Posterior sclerite expressed as plate-like structure reaching the anterior margin of median region or bulge-like with anterior margin convex, ‘pseudo teeth’ present ............................................................................. 51

-. Posterior sclerite expressed as a bar-like structure with anterior margin straight or concave ...................... 52

51. Posterior sclerite expressed as plate-like structure reaching the anterior margin of median region, ‘pseudo teeth’ absent...................................................................................................................... Tegenaria henroti

-. Posterior sclerite bulge-like with anterior margin convex, large, reddish receptaculi visible through epigynal plate, ‘pseudo teeth’ present ............................................................................................. Tegenaria racovitzai

52. ‘Pseudo teeth’ present ..................................................................................................................... 53

-. ‘Pseudo teeth’ absent.......................................................................................................................54

53. CL longer than 3.5 mm, vulva as long as wide............................................................ Tegenaria domestica

-. CL shorter than 3.5 mm, vulva twice as long as wide......................................... Tegenaria annae sp. nov.

54. Posterior sclerite broad, pocket-like or narrow, semicircularly shaped, protruding posteriad.........................55

-. Posterior sclerite broad, bar- or bulge-like, only inconspicuously protruding..............................................56

55. Posterior sclerite broad, pocket-like, protruding, vulva globularly shaped............................. Tegenaria achaea

-. Posterior sclerite narrow, semicircularly shaped, protruding posteriad, vulva with distinct appendages, visible through the epigynal plate ...................................................................................... Tegenaria percuriosa

56. Distal margin of posterior sclerite straight.............................................................. Tegenaria faniapollinis

-. Distal margin of posterior sclerite concave..........................................................................................57

57. Distal margin of posterior sclerite only moderately concave, as wide as deep, vulva spherical......................... ............................................................................................................................... Tegenaria pieperi

-. Distal margin of posterior sclerite distinctly concave, vulvae without distinguishable RC, duct-like .............. 58

58. Distal margin of posterior sclerite distinctly concave, semicircularly shaped, anteriorly of posterior sclerite without septum (Fig. 23C)................................................................................................... Tegenaria tridentina

-. Anteriorly of posterior sclerite with a very distinct, tongue-like shaped median septum (Fig. 23I).................... ........................................................................................................................... Tegenaria levantina

59. Epigyne with distinct epigynal teeth, originating at the posterior margin of the median plate or ‘pseudo teeth’ (distally pointed projections of lateral margin of median region).............................................................60

-. Epigyne without epigynal teeth ........................................................................................................ 61

60. Epigynal teeth, originating at the posterior margin of the median plate..........................Tegenaria rhodiensis

-. ‘Pseudo teeth’ (distally pointed projections of lateral margin of median region) present.................................. ........................................................................................................ Tegenaria pindosiensis sp. nov.

61. Epigyne distinctly ‘half-mask’-shaped (Fig. 16A), vulva as in Figures 14I, 16B.............................................. .................................................................................................... Tegenaria montiszasensis sp. nov.

-. Character combination different........................................................................................................62

62. Epigyne very distinct ( Fig. 17U ), distal segment of PLS almost two x longer than basal segment .................... ........................................................................................................................ Tegenaria schmalfussi

-. Character combination different........................................................................................................63

63. Epigynal median region anteriorly continuously separated from the epigynal plate by a distinctly sclerotized rim ( Fig. 22Q , white arrow) ................................................................................................................... 64

-. Distinct separation of the median region by a sclerotized rim absent ...................................................... 68

64. Median region oval, no pocket expressed, vulva subrectangular shaped .............................. Tegenaria mirifica

-. Median region rectangular or trapezoidal shaped, vulva globular or suboval shaped .................................. 65

65. Median region with a distinct pocket in the middle, opening anteriad ..................................................... 66

-. Median region without median pocket................................................................................................67

66. Copulatory openings are located at the lateral sides of the median pocket ........................ Tegenaria sbordonii

-. Copulatory openings are much more anteriorly located (Fig. 18P).................................. Tegenaria parmenidis

67. Copulatory openings originating distant from the lateral rim, no lateral pockets developed ............................ .......................................................................................................................... Tegenaria capolongoi

-. Copulatory openings originating right at the lateral rim, distinct pockets posteriorly of the copulatory openings at the lateral rim present........................................................................... Tegenaria circeoensis sp. nov.

68. Distinct median plate, smoothly sclerotized, vulva developed as a strongly convoluted duct.........................69

-. Median plate distinct or indistinct, mostly irregularly sclerotized, vulva globular or irregularly shaped, never only duct-like .......................................................................................................................................71

69. Epigynal median plate almost as long as wide, epigyne and vulva as in Figures 19D–I, 20C–E, H, I, L–M....70

-. Epigynal median plate wider than long ( Fig. 17H, I ) ....... Tegenaria campestris complex (Deltshev, 1993, 2008b)

70. Epigyne and vulva as in Figure 20L, M, genital openings distinct ..................................... Tegenaria parvula

-. Epigyne and vulva as in Figures 19D–I, 20C–E, H, I ..................................................... Tegenaria silvestris

71. Median plate strongly and irregularly sclerotized, very large and distinct copulatory openings........................ .................................................................................................................... Tegenaria mercanturensis

-. Character combination different........................................................................................................72

72. Strongly sclerotized epigyne, copulatory openings triangular or almond-like shaped, vulva suboval shaped ....... ............................................................................................................................ Tegenaria ariadnae

-. Epigyne less sclerotized without distinct median plate, vulva irregularly formed or with convoluted fertilization duct.............................................................................................................................................73

73. Indistinct median region rectangular shaped, copulatory openings at the anterior border of this area, most often plugged, vulva very distinctly shaped (Fig. 15N).............................................................. Tegenaria hasperi

-. Median region trapezoidal shaped, copulatory opening laterally of this area, ducts and receptaculi clearly visible through the epigynal plate, vulva with broad copulatory ducts, small globular receptaculum, and convoluted fertilization duct ( Figs 17X–Z , 19N, O) ........................................................... Tegenaria croatia sp. nov.

Description

General description was provided by Brignoli (1977b). Following his statement on page 945, ‘Misura: (in mm, leggermente approssimate):...’ (Measurement: (in mm, approximately):...) (1977b: 945), the two females were remeasured and the values given here: Female (N = 2): CL 2.03–2.60, CW 1.46–1.91, STL 1.01–1.33, STW 1.01–1.15, OL 3.72, OW 2.69. Leg I (3.41, 1.04, 3.22, 3.3, 1.9), II (2.54–2.75, 0.78–0.92, 2.22–2.68, 2.33–3.05, 1.36–1.53), III (2.15, 0.66, 1.89, 2.3, 1.15), IV (2.68–3.56, 0.76–0.96, 2.48–3.26, 3.14– 4.08, 1.37–1.57). Palp (0.99–1.31, 0.35–0.51, 0.79– 0.92, 0.88–1.19). Eyes: PME 0.08, PLE 0.09, AME 0.06, ALE: 0.09. Eye distances: PME–PME 1.5 x PME, PME–AME 1.5 x PME, PME–PLE 1.5 x PME,

PME–ALE 1–1.5 x PME, AME–AME 1 x AME, AME– ALE 1 x AME.

Distribution

Reported from the island of Ikaria, Greece.

Figures

Figure 15. A, B, Tegenaria achaea; C-F, Tegenaria argaeica; G-J, Tegenaria armigera; K, L, O-Q, Tegenaria dalmatica; M, N, Tegenaria hasperi; R-U, Tegenaria annulata; and V, X, Tegenaria schoenhoferi sp. nov. Left male palp in lateral (C, H, L, U, W), and ventral views (D, G, K, T, V); epigyne in ventral (A, E, I, M, O, R) and vulva in dorsal view (B, F, J, N, P, S); comparison of vulva of small and large ‘forms’ of Teg. armigera (J); Prosoma in frontal view (Q), abdomen in dorsal view (X). Scale bars = 0.5 mm (epigyne and vulva of Teg. argaeica, E–F without scale; photos from syntype taken with digital camera through the oculars).

Figure 17. A, B, Tegenaria domestica; C, F, G, M, N, Tegenaria racovitzai; D, E, Q, R, Tegenaria hauseri; H-J, Tegenaria bozhkovi; K, L, O, P, Tegenaria campestris; S, T, Tegenaria pieperi; U, Tegenaria schmalfussi; V, Tegenaria faniapollinis; W, Tegenaria percuriosa (holotype of Tegenaria bithyniae); X-Z, Tegenaria croatica sp. nov. Left male palp in ventral (M, O, Q) and retrolateral views (N, P, R); epigyne in ventral (D, F, H, K, S, U–W, Y) and posterioventral views (T), vulva in dorsal (E, G, I, L, U, Z) and lateral views (J); face of female Teg. domestica (A) and Teg. racovitzai (C); cheliceral margins of female Teg. domestica in posterioventral view (B); habitus, dorsal view (X). Scale bars = 0.5 mm (no scale for Teg. faniapollinis, V, and Teg. schmalfussi, U).

› Figure 22. A, B, Tegenaria capolongoi; C-K, Tegenaria parmenidis; L-S, Tegenaria circeoensis sp. nov. Left male palp in ventral (E, N), retrolateral (F, O), and dorsal views (G, P); epigyne in ventral (A, C, Q, S) and vulva in dorsal view (B, D, R); intraspecific epigynal morphological variation (S); face of male in frontal view (J); spinnerets in ventral view (K); habitus (H) of male in dorsal and sternum in ventral view (I); carapace and abdomen of male in dorsal view (L, M). Scale bars = 0.5 mm (except 1 mm for H).