Eratigena atrica, COMB. NOV.

ERATIGENA ATRICA (C. L. KOCH, 1843) COMB. NOV.

(FIGS 1J, 2H–J, 8A–B, 9I–O, 10A–G)

Tegenaria atrica C. L. Koch, 1843: 105–107, fig. 825; Blackwall, 1861: 165, pl. 11, fig. 106; Simon, 1875: 81, pl. 5, fig. 5; Locket & Millidge, 1953: 10, figs 6A, 7A, 9A, 11B.

Tegenaria saeva Blackwall, 1844: 179–182, syn. nov.; Simon, 1937: 1003, 1039, figs 1545–1546; Roth, 1968: 29, figs 36–39 (synonymized with Tegenaria gigantea, rejected by Brignoli, 1978a: 273); Locket, Millidge & Merrett, 1974: 42, figs 23A, E; Locket, 1975: 85–90, figs 1, 4, 6–11, 13–16.

Tegenaria duellica Simon, 1875: 83–85, pl. V, fig. 6, syn. nov.

Tegenaria larva Simon, 1875: 86, 87, Planche V: fig. 8.

Tegenaria nervosa Simon, 1870: 273–275, synonymized by Simon (1937), reactivated by Brignoli (1978a) but only based on drawings.

Tegenaria hibernica Pickard-Cambridge, 1891: 86, fig. 4, male; Pickard-Cambridge, 1893: 150, fig. 6, female.

Tegenaria gigantea Chamberlin & Ivie, 1935: 31, pl. XIII, fig. 106; synonymized with Teg. duellica by Brignoli, 1978a: 271, 273; Merrett, 1980; Oxford & Smith, 1987; Oxford & Plowman, 1991; Heimer & Nentwig, 1991; Croucher, Oxford & Searle, 2004; Croucher et al., 2007; Oxford, 2008.

Tegenaria praegrandis Fox, 1937: 176–177, fig. 3.

Tegenaria deroueti Dresco, 1957: 212–215, figs 2, 13–14.

Tegenaria derouetae: Denis, 1959: 173.

Tegenaria propinqua Locket, 1975: 85–90, figs 2, 3, 5, 17–19; synonymized with Teg. gigantea by Crawford & Locket (1976).

Types

No type material available for Teg. atrica, Teg. deroueti, Teg. duellica, Teg. larva, Teg. nervosa, and Teg. saeva in the collection of either the MNHN or NHML. Sub Tegenaria gigantea: Paratypes. Canada: British Columbia: Vancouver Island, Sidney, 2 ♂, 2 ♀ (AMNH), 16.ix.1935, Chamberlin & Ivie; 4 ♂, 4 ♀ (AMNH), same data as previous; Vancouver Island, South Saanich, 1 ♂ (AMNH), 1922.

Other material examined

Austria (1 ♂, 4 ♀); Belgium (1 ♂, 18 ♀); Czech Republic (2 ♀); Germany (23 ♂, 34 ♀); France (2 ♂, 6 ♀); Italy (1 ♂); Luxemburg (1 ♀); Poland (1 ♂, 1 ♀); Spain (2 ♂, 3 ♀); Sweden (1 ♀); Switzerland (21 ♂, 17 ♀). Asia: Lebanon (1 ♀).

Sub Tegenaria duellica / gigantea: France (1 ♀); United Kingdom (5 ♂, 3 ♀); Spain (3 ♀); Portugal (3 ♀). North America: USA (1 ♀).

KEY TO EUROPEAN SPECIES OF ERATIGENA GEN. NOV.

Eratigena vidua (Cárdenas & Barrientos, 2011) comb. nov. (from Tegenaria) is not treated here as material was not available for examination. The new combination is based on the information provided in the original description (Cárdenas & Barrientos, 2011).

1. Legs distinctly annulated, at least two femora with more than two dorsal spines, male bulb with distinct and massive transversal ridge of conductor base (Fig. 14A), vulva with distinct and long appendages at the CD................................................................................................................................................2

-. Legs either pale, completely darkened or only coxa and proximal part of femora darkened (exception: E. sicana, legs may be annulated), other characters different.................................................................................3

2. Basal part of MA very strongly sclerotized, transversal ridge of conductor base with distinct border line of sclerotization, long appendix anteriorly of CD shorter than RC height................................ Eratigena inermis

-. Basal part of MA less sclerotized, transversal ridge of conductor base without special border line of sclerotization, long appendix anteriorly of CD reaches at least to the top of the RC................................. Eratigena vomeroi

3. Trichobothria on female tarsal tibia and on male cymbium present, length of male tibia I shorter than or equal to the length of carapace..................................................................................................................4

-. Trichobothria on female tarsal tibia and on male cymbium absent, male tibia I longer than carapace (exceptions: Eratigena fuesslini and Eratigena bucculenta sensu Barrientos, 1991) .......................................................7

4. Patellar apophysis on male palp present, RTA strongly reduced, epigyne with special posterior plate, protruding ventroposteriad (Figs 8K, M, 11G, I, K)................................................................................................5

-. Patellar apophysis absent, RTA with two simple branches, distally truncated, epigyne without posterior sclerite .................................................................................................................................................... 6

5. Patellar apophysis with two well-separated points, conductor with straight terminal end and as long as the alveolus, epigynal posterior sclerite rectangularly shaped and copulatory openings lateral ..... Eratigena feminea

-. Patellar apophysis with three moderately separated points, conductor shorter than the alveolus with terminal end bent ventrad, epigynal posterior sclerite more triangular, copulatory opening anterior................................... .......................................................................................... Eratigena bucculenta (sensu Machado, 1941)

6. Male palp with pyramidal-shaped structure at conductor connection, terminal end of conductor pointing orthogo- nally away from cymbium (in lateral view), vulva with small diverticula at copulatory duct originating laterally, egg-shaped receptacula irregularly sclerotized ........................................................... Eratigena barrientosi

-. Male palp without conspicuous structure at connection of conductor, terminal end of conductor pointing dorsad or posteriad (in lateral view), vulva with small diverticula at copulatory duct originating dorsally, globular recep- tacula smoothly sclerotized ....................................................................................... Eratigena incognita

7. Legs with only coxa and proximal part of femora darkened (exception: Eratigena fuesslini, completely darkened), distal segment of PLS as long as or only marginally longer than basal segment, median apophysis on male left palp originating at 7–9 o’clock position, basal portion of tegulum not visible between embolus and conductor in ventral view (Fig. 9A), epigyne with ‘pseudo teeth’ (Fig. 10A–C) or with distinct atrial cavity in combination with a posterior sclerite expressed as a large bulge ...................................................................................... 8

-. Legs either pale or completely darkened (exception: Eratigena sicana, legs may be annulated), distal segment of PLS longer than basal segment, median apophysis on male left palp originating at 5–7 o’clock position, basal portion of tegulum visible in ventral view (Fig. 10L), epigyne different .................................................... 10

8. Sternum with a pale median band and symmetrical pale dots laterally, female tibia I longer than carapace length, origin of embolus at 10–11 o’clock position, terminal end of conductor simple, epigynal ‘pseudo teeth’ present, atrium without a distinct cavity..................................................................................... Eratigena atrica

-. Sternum with a pale median band, female tibia I shorter than or as long as carapace length, origin of embolus at 8–10 o’clock position, terminal end of conductor complex, epigynal teeth present, atrium forms a distinct cavity in combination with a posterior sclerite expressed as a large bulge .......................................................... 9

9. Carapace longer than 4.8 mm, conductor very broad in retrolateral view with a very complex terminal end, epigyne with a posterior sclerite expressed as a large bulge, not protruding posteriad, copulatory ducts relatively short....................................................................................................................... Eratigena agrestis

-. Carapace shorter than 4.5 mm, conductor relatively slender in retrolateral view with a bifid terminal end, epigyne with a posterior sclerite expressed as a large bulge protruding posteriad, copulatory ducts relatively long ........ ............................................................................................................................. Eratigena fuesslini

10. Distal portion of conductor not elongated (shorter than broad), lateral margin of conductor folded only at the terminal half, transversal ridge massive and moderately protruding, epigyne and vulva as in Figs 10M–N and 11N–P..........................................................................................................................................11

-. Distal portion of conductor moderately to strongly elongated, lateral margin of conductor completely folded, transversal ridge (if present) only expressed as a membranous ridge, epigyne and vulva not as above ..........12

11. RTA strongly protruding ventrad, terminal end of conductor bifid and with additional spur (Fig. 10O, black arrow), strongly sclerotized epigynal plate with two symmetrically arranged reniform depressions with copulatory openings (Fig. 10M)................................................................................................ Eratigena montigena

-. RTA simple, terminal end indistinct with only one somewhat elongated point, epigyne with posterior sclerite expressed as strongly sclerotized and protruding plate (median of the atrium)............................................. ........................................................................................ Eratigena bucculenta (sensu Barrientos, 1991)

12. Dorsal branch of RTA dorsally bent posteriad (Fig. 12R), short dorsal spike at male palp tibia present, terminal end of conductor bent or convoluted, complex, median apophysis only moderately protruding, epigyne with a distinct atrial cavity ....................................................................................................................... 13

-. Dorsal branch of RTA not as above, short dorsal spike at male palp tibia absent, terminal end of conductor simple with elongated point, median apophysis protruding, epigyne with a distinct posterior sclerite expressed as a protruding sclerotized plate or with a portion protruding strongly posteriad (Fig. 12A, F, I) ........................ 14

13. Distal portion of conductor strongly elongated, terminal end of conductor convoluted, relatively short male palp tibia, CD barely visible through epigynal plate, vulva with very long and convoluted CD (Fig. 13H)................ ................................................................................................................................. Eratigena picta

-. Distal portion of conductor only moderately elongated, terminal end of conductor bent, relatively long male palp tibia, CD distinctly visible through epigynal plate, vulva with short CD........................... Eratigena balearica

14. Eyes very small, clypeus higher than three x AME, at least one pair of spines prolateral at palp tibia, seven or more tarsal trichobothria, dorsal branch of RTA distinct (Fig. 12L, M), female tibia I longer than carapace, appendages at copulatory ducts long.................................................................................................15

-. Eyes larger, clypeus lower than three x AME, two (not paired) prolateral spines at palp tibia, up to six tarsal trichobothria, dorsal branch of RTA different, female tibia I shorter than or equal to the length of carapace, appendages at copulatory ducts short or absent..................................................................................16

15. Carapace shorter than 3.5 mm, distal tip of conductor not reaching the distal margin of the alveolus, epigyne lacking distinct membranous region anteriorly of posterior sclerite, vulva with short, convoluted appendages.... ............................................................................................................................. Eratigena herculea

-. Carapace longer than 4 mm, RTA as in Figure 12L, M, distal tip of conductor reaching distal margin of alveolus, epigyne with distinct membranous region basal of posterior sclerite, vulva with long, convoluted appendages.... ........................................................................................................................... Eratigena hispanica

16. Dorsal branch of RTA three-pointed (Fig. 12O), vulva with long oval and irregularly sclerotized (enclosing convoluted ducts) receptacula ...................................................................................... Eratigena arganoi

-. Dorsal branch of RTA with one point, vulva with globular and smoothly sclerotized receptacula .................. 17

17. Ratio of bulb to cymbium length larger than 0.65, terminal end of conductor expressed as a strongly posteriad elongated point, epigyne with a strongly posteriad protruding portion, vulva with very long convoluted ducts ............................................................................................................................... Eratigena sardoa

-. Ratio bulb to cymbium length smaller than 0.60, terminal end of conductor inconspicuous, epigyne with a portion protruding posteriad ...................................................................................................Eratigena sicana

Sub Tegenaria saeva: France (7 ♂, 9 ♀); United Kingdom (7 ♂, 3 ♀); Portugal (5 ♀); Spain (3 ♂, 3 ♀).

Diagnosis

Eratigena atrica can be separated from all other Eratigena gen. nov. species by having leg coxa and proximal parts of femora darkened (spotted, as in E. agrestis, all other Eratigena gen. nov. species with different patterns), short dorsal spike at male palp tibia present (as in E. agrestis, E. fuesslini, E. barri- entosi, E. montigena, E. picta, and E. balearica, absent in all other species), very massive conductor, laterally folded only at the terminal half with the very distinct shape of the conductor in retrolateral view (Fig. 9J–L), MA broad pocket-like, strongly attached to the tegulum, originating at 7–9 o’clock position (as in E. agrestis and E. fuesslini, in all other species more basal, 5–7 o’clock position), basal portion of tegulum almost completely hidden by upper part (embolic division) of tegulum (as in E. agrestis, E. fuesslini, and to some extent also in E. sardoa, in all other species of Eratigena gen. nov. well visible), strongly expressed epigynal ‘pseudo teeth’ (Fig. 10A–C, white arrows), which are missing in all other Eratigena gen. nov. species.

Description

Measurements: Male (N = 2): CL 4.87–6.78, CW 3.65– 5.12, STL 2.35–3.23, STW 2.12–3.05. Leg I (7.66– 11.12, 1.91–2.85, 7.40–11.40, 7.79–11.31, 3.52–4.43), II (6.17–8.69, 1.76–2.69, 5.42–7.93, 6.29–9.38, 2.79– 3.43), III (5.42–7.52, 1.62–2.30, 4.47–6.32, 6.23–8.94, 2.48–3.36), IV (6.95–9.32, 1.69–2.49, 6.33–8.60, 8.58– 12.56, 3.19–4.21). Pedipalp (2.50–3.24, 0.92–1.26, 1.10–1.42, 2.22–2.81), bulbL 1.26–1.30. Female (N = 1): CL 5.84, CW 4.10, STL 2.82, STW 2.49, OL 6.27, OW 4.13. Leg I (6.94, 2.28, 6.61, 3.21), II (6.12, 2.03, 5.08, 5.83, 2.85), III (5.60, 1.94, 4.37, 6.05, 2.55), IV (7.10, 1.99, 6.43, 8.67, 2.96). Pedipalp (2.78, 1.10, 1.80, 2.68). EPL 1.19, EPW 1.41, ATL 0.95, ATW 0.60. Eyes: PME 0.22–0.24, PLE 0.23–0.25, AME 0.25– 0.27, ALE 0.26–0.27. Eye distances: PME- PME 1 x PME, PME- AME 0.5 x PME, PME- PLE 1–1.5 x PME, PME- ALE 1–1.5 x PME, AME- AME 0.5 x AME, AME- ALE <0.5 x AME. CLY1 2–2.5 x AME, CLY2 1.5–2 x ALE.

Male palp: RTA with two branches, lateral branch simple and pointed, dorsal branch a large knoll with a strongly sclerotized and pointed protrusion. Short dorsal spike on palp tibia present. Embolus length about 1–1.25 x CB, originating at 10–12 o’clock position, distal tip at 4 o’clock position. Conductor massive, distal portion not elongated, folded only at the terminal half. Terminal end of conductor expressed as strongly sclerotized and elongated point directing ventrad. Transversal ridge of conductor expressed as membranous lamella. Conductor membranously connected to tegulum. MA originating at 7–9 o’clock position, moderately protruding, wider than long, distally with pocket-like sclerite. Connection of MA to tegulum at least partly sclerotized.

Epigyne and vulva: Epigyne medially with strongly sclerotized, long rectangular plate, anteriorly of which the CO are located. Posterior sclerite absent. Epigynal teeth absent, but ‘pseudo teeth’ present. Vulva consists of distinguishable CD, RC, and FD. CD short and straight without long appendages. RC irregularly oblong and sclerotized, enclosing convoluted ducts, separated by more than two times their diameter. FD only represented by small, leaf-shaped appendages.

Other important characters: Cheliceral promargin with three, retromargin with seven to nine teeth. Colulus rectangular with distal margin w-shaped. PMS with one prominent minor ampullate gland spigot and three to four cylindrical gland spigots laterally. Trichobothria on cymbium and palp tarsus absent. Seven to ten tarsal trichobothria. Small teeth on paired claws of leg I: 13–14. Leg spination: male palp (2–0–0–0 or 2–1–0–0 or 2–1–1–0, 2–0–0, 1–2p– 0–0), female palp (2–0–0–0 or 2–0–1–0 or 2–1–0–0, 2–0–0, 2–2p–0–0), leg femora (2–3–3–0 or 2–3–4–0 or 3–3–2–0 or 3–3–3–0 or 3–3–4–0 or 3–4–4–0, 2–3–2–0 or 2–3–3–0 or 3–3–3–0, 2–2–2–0 or 2–3–2–0, or 2–3– 3–0 or 2–3–4–0 or 2–4–2–0, 2–2–1–0 or 2–3–1–0 or 2–3–2–0), patellae (all 2–0–0), tibiae (0–0–0–1p+1+1p or 0–0–0–2p or 0–1–0–1p+1, 0–1–0–2p or 0–2–0–2p or 2–2–0–2p, 2–1–1–1p+2+1p or 2–2–2–1p+1+1p or 2–2– 2–2p or 2–2–3–2p, 2–2–2–1p+1+1p+1 or 2–2–2–2p or 2–2–2–3p), metatarsi (0–0–0–1p+2+1p+1 or 0–0–0– 4p+1, 0–2–0–4p+1 or 0–3–0–3p+1+2p or 0–3–0–5p, 1+1p–4–3–5p or 2–4–4–5p, 3–4–4–1p+1+1p+2+2p or 3–4–4–5p+1), tarsi (0, 0 or 0–1–0–0 or 0–1–1–0 or 0–1–3–0 or 0–2–2–0, 0–1–3–0 or 0–2–3–0, 0–2–3–0 or 0–2–4–0).

Coloration: Two symmetrical longitudinal dark bands dorsally on carapace, sometimes serrated or reduced to triangular dots. Sternum with distinct pattern of pale median band and three symmetrical pairs of pale dots laterally. Opisthosoma darkened with one pale band anteriorly in the middle, laterally with pale dots, continuing to the posterior in chevrons. Legs not annulated, darkened, sometimes only coxa and proximal part of femora with dark spots. ALS slightly darkened, PLS with both segments darkened dorsally, distal segment moderately paler.

Distribution

Reported from most west, Central, and north European countries. Introduced into North America (first mentioned sub Teg. praegrandis Fox, 1937). Additionally, there was a specimen from Lebanon in the collection at SMF, unfortunately with insufficient locality information written on the related label.

Discussion

As mentioned by Locket (1975) and Brignoli (1978a), no type material is available for Teg. duellica. Here, the argumentation of Brignoli (1978a) is followed. The examined type material of Teg. gigantea and the examined specimens sub E. duellica are not separable. Therefore, Teg. gigantea Chamberlin & Ivie has to be recognized as a junior synonym of E. duellica (Simon) (sub Tegenaria) and is therefore also a synonym of E. atrica.

As in some other species of Tegenaria and Eratigena gen. nov., some morphological characters are highly variable in, e.g. the size of different body parts (e.g. Simon, 1937: 1003) or the patterns of leg spination. This variation has been reflected upon and discussed in many publications (Blackwall, 1861; Locket & Millidge, 1951, 1953; Denis, 1959; Roth, 1968; Locket et al., 1974; Crawford & Locket, 1976; Brignoli, 1978a; Barrientos & Ribera, 1988; Croucher et al., 2004, 2007). Whereas some authors considered this variation of intraspecific nature, others used it to recognize three species (E. atrica, E. saeva, E. duellica). Barrientos & Ribera (1988) were proponents of the former idea and their view is supported as follows: (1) as shown by Croucher et al. (2004), Bolzern et al. (2010), and the present paper, the three species are not recovered with CO1 and NADH1 gene sequences (the genetic distances are very small); (2) as suggested by Barrientos & Ribera (1988), seemingly major morphological differences arise when structures are not observed in the same perspectives. This is particularly relevant if published drawings are compared, as in the case of Teg. nervosa, which was synonymized with Teg. atrica by Simon (1937) and later again recognized as a valid species by Brignoli (1978a) based only on the drawing of Simon (1875).

Members of the British research group of Geoff Oxford (Oxford & Smith, 1987; Oxford & Plowman, 1991; Croucher et al., 2007; Anderson et al., 2009) suggested that in Great Britain the three species are mostly allopatric with narrow hybrid zones. The major problem with this concept is that it is not applicable to the continental populations in which transitional morphs exist.