Eratigena, Bolzern & Burckhardt & Hänggi, 2013

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 : 738-743

ERATIGENA GEN. NOV.

Type species

Tegenaria atrica C. L. Koch, 1843 , present designation.

Etymology

Anagram of Tegenaria , gender feminine.

Diagnosis

Agelenid spiders bearing the following character combination: plumose hairs present (absent in Lycosoides , Maimuna , and Textrix ). AER and PER straight or only slightly pro- or recurved in dorsal view (both rows recurved in Lycosoides , Maimuna , and Textrix ; both rows procurved in Agelena , Agelescape , Allagelena , and Benoitia ) and moderately procurved in frontal view (AER strongly procurved in Agelena , Agelescape , Allagelena , Benoitia , and Malthonica ; AER recurved in Lycosoides , Maimuna , and Textrix ). Cheliceral retromargin with six or more teeth (fewer than three teeth in Lycosoides , Maimuna , and Textrix ; six or fewer teeth in Tegenaria ) that decrease in size from distal to proximal (all equal in Tegenaria ). Trochanter straight or slightly curved (notched in Aterigena , Histopona , and Malthonica ). Patellae with

KEY TO EUROPEAN AGELENID GENERA (COELOTINAE NOT INCLUDED)

Pseudotegenaria Caporiacco, 1934 , is excluded here because the only remaining species, P. parva Caporiacco, 1934 , is mentioned only from Libya and no specimen was available for examination.

1. Trochanters III and IV notched .......................................................................................................... 2

-. All trochanters straight or only slightly curved .....................................................................................5

2. Dorsal and lateral spines present on patellae III and IV. Tarsus IV with one to two ventral spines................. ........................................................................................................................... Aterigena (five spp.)

-. Only dorsal spines present on all patellae............................................................................................3

3. Colulus developed as medially slightly divided plate, sometimes only two hairy plates visible, patellar apophysis on male palps sometimes present, median apophysis absent ........................................... Histopona (20 spp.)

-. Colulus strongly reduced, only hairs present ( Fig. 1L ). Patellar apophysis on male palps absent, median apophysis present .......................................................................................................................................... 4

4. Eyes well developed. Tarsi with fewer than seven dorsal trichobothria. Cheliceral retromargin with one or two larger and several smaller teeth ............................................................................. Malthonica (two spp.)

-. Eyes very small or lacking, tarsi with seven or more dorsal trichobothria. Several equally large teeth on the cheliceral retromargin.................................................................................................. Hadites (one sp.)

5. PER in dorsal view considerably recurved or procurved. AER in frontal view either considerably procurved or slightly recurved. Patellae I and II with dorsal and prolateral spines. Colulus clearly divided into two hairy plates ............................................................................................................................................ 6

-. Both eye rows in frontal and dorsal view more or less straight (eyes may be reduced). Patellae with dorsal spines only. Colulus trapezoidal plate with distal margin straight, w-shaped, or with a notch...............................12

6. Eye rows in dorsal view recurved; PME largest. Feathery hairs absent.....................................................7

-. Eye rows in dorsal view procurved; PME never largest. Feathery hairs present ( Fig. 2A ) .............................9

7. Conductor on male palp with conspicuous laterodorsal projection; femur of male palp with flat but distinct prominence. No patellar apophysis. Epigyne without raised median portion reaching anteriorly into large atrium ............................................................................................................... Maimuna (seven spp.)

-. Conductor on male palp without laterodorsal projection; femoral apophysis absent; patellar apophysis sometimes present. Epigyne with posteriorly protruding pocket or slightly raised median portion .................................8

8. Conductor simple; median apophysis absent; patellar apophysis absent. Epigyne with posteriorly protruding pocket.................................................................................................................... Textrix (seven spp.)

-. Conductor more complex; median apophysis present; patellar apophysis present. Epigyne with slightly raised median portion reaching anteriorly into atrium... ..................................................... Lycosoides (ten spp.)

9. Distinct spikes absent on the anal tubercle. Conductor helical, strongly protruding. Epigynal plate with two clearly divided copulatory openings ...................................................................................... Benoitia (nine spp.)

-. Anal tubercle with distinct long and dark spikes. Conductor never helical and strongly protruding. Epigynal plate longitudinally not completely divided.................................................................................................10

10. Embolus thin and filamentous. Conductor simple. Anterior margin of epigynal atrium with protruding scapus ....................................................................................................................... Agelescape (seven spp.)

-. Embolus either broad and short or spiral, elongated with an attached membrane. Conductor complex. Anterior margin of epigynal atrium without scapus.......................................................................................... 11

11. Patellar protuberance absent. Two tibia apophyses developed. Embolus short and broad. Vulva with an interiorly originating spermathecal head; spermathecal apophyses present......................................... Agelena (70 spp.)

-. Patellar protuberance with a long spine. One tibial apophysis developed. Embolus long and spiral with attached membrane. Vulva with a laterally or medially originating spermathecal head; spermathecal apophyses absent.. .......................................................................................................................... Allagelena (five spp.)

12. Cheliceral retromargin with six or more teeth (more proximal teeth decrease in size). RTA with mostly two branches; lateroventral ridge absent. Conductor with a membranous or massive (not strongly sclerotized) transverse ridge, terminal end not bifid (only in E. montigena ) with ventral apex either forming elongated process or being more complex (several points or spiral). Median apophysis strongly attached to the tegulum (sometimes only membranous), distally with a simple, pocket-like sclerite. Vulva irregularly sclerotized (enclosed convoluted duct) and/or with diverticula attached to the copulatory duct (exception: E. sicana with two ST)..................... ............................................................................................................ Eratigena gen. nov. (19 spp.)

-. Cheliceral retromargin with three to six large, subequal teeth. RTA mostly with three branches or strongly protruding bulge and lateroventral ridge. Conductor lamelliform, sometimes with sclerotized transverse ridge, apex often bifid with ventral ending simple; median apophysis strongly protruding with distal sclerite plate-like or more complex. Vulva only convoluted duct or with more or less regularly sclerotized, globular spermathecae ............................................................................................................................ Tegenaria (56 spp.)

dorsal but without lateral spines (as in Histopona , Malthonica , and Tegenaria ; all other European genera with lateral patellar spines). Lacking ventral spines on all tarsi (as in Malthonica and Tegenaria ; all other European genera with ventral spines). Colulus forming rectangular or trapezoidal plate with distal margin straight or w-shaped (in Tegenaria trapezoidal and notched medially; colulus strongly reduced in Hadites and Malthonica ; two separated plates in all other European agelenids). Females with one minor ampullate gland spigot, very prominent, and two to four cylindrical gland spigots distally on PMS (as in some species of Aterigena and Malthonica , all other European genera with different patterns). Male palp: RTA with one or two branches, may also be reduced (complex in most Tegenaria ), palp tibia often with short dorsal spike (absent in all other European genera), filiform embolus, mostly with massive conductor (lamelliform in Tegenaria ) with a membranous or massive transverse ridge (as in Lycosoides , Maimuna , and Textrix ; in other genera absent or only moderately sclerotized) and a complex or strongly elongated terminal ending (as in Allagelena and Maimuna ; simple in Tegenaria ), only moderately elongated median apophysis with distal plate-like sclerite (absent in Histopona and Textrix ; without sclerite in Agelena , Agelescape , and Benoitia ). Female: epigyne without separated or strongly fused median area (as in Agelena , Agelescape , Allagelena , Benoitia , Lycosoides , and Maimuna ; clearly separated in Tegenaria ). Vulvae either with an irregularly sclerotized RC with enclosed convoluted ducts or with distinct appendages at the copulatory duct (exception: E. sicana ).

Description

Body size medium to large (carapace length between 2 and 7 mm). Margin of carapace narrowly and continuously darkened (for cave-living species, pigmentation absent); two symmetrical longitudinal dark bands present dorsally on carapace, serrated, continuous, or reduced to three to four conspicuous triangles. Sternum slightly longer than wide with a distinct pattern of pale median region, sometimes additionally with three to four lateral spots; plumose hairs present on carapace, legs, and opisthosoma. Chelicerae with three promarginal teeth and six or more retromar- ginal teeth; retromarginal teeth decrease in size proximally. Labium as wide as long. AER and PER straight or only slightly pro- or recurved in dorsal view and moderately procurved in frontal view. Anterior eyes larger than posterior ones or lateral eyes larger than median ones or all eyes equal, except AME slightly smaller. All trochanters straight or slightly curved. Leg I or IV longest, III shortest. Legs sometimes without pattern, only coxa and proximal part of femur darkened, annulated, or completely darkened. Palp and leg spination: palp femora with one to two dorsal and sometimes one pro- and/or one retrolateral spines, female palp tibia with two dorsal and either one + one pair, two, or two paired prolateral spines; male palp tibia dorsodistally often with a short spike and prolateral with one + one pair, two, or two pairs of spines; all leg femora with one to two dorsal spines and variable number of lateral spines; patellae with two dorsal and no lateral spines; number of dorsal spines on metatarsi variable; metatarsus I without or with one prolateral spine, others variable; metatarsi III and IV ventrodistally with one pair + one spines; tarsi I and II lacking spines, III and IV with zero to several prolateral and one to several retrolateral spines, without ventral spines. Spinnerets: colulus rectangular or trapezoidal plate with the distal margin straight or w-shaped. ALS onesegmented, with a field of several pyriform spigots distally and with two major ampullate spigots medially (present in all agelenids). PMS as long as or slightly shorter than ALS, bearing one conspicuously prominent spigot. PMS with one minor ampullated and two to four cylindrical gland spigots, one medially located and two to four posteriolaterally located and several aciniform gland spigots. PLS longer than all others with distal segment as long as or longer than basal segment. PLS bearing typically one basal and one medial cylindrical gland spigot. Male palp without femoral apophysis, patellar apophysis can be present. RTA mostly with two branches and simple (in some species several short or bent points can be expressed), sometimes reduced to one branch or entirely lacking. Embolus filiform, getting thinner to apex. Conductor mostly massive (thick and broad, if different then always median with a white membranous part), transversally with a membranous or massive ridge, mostly indistinct, elongated distal portion (exception: E. picta ) and lateral margin folded along the terminal half or the whole length, terminal end consists mostly of one elongated peak or a more complex structure (spiral, several points) but dorsally only rarely with rounded bulge. Median apophysis usually not protruding (as wide as or wider than long, exception e.g. E. sardoa ), consisting of membranous base and distal sclerite, which is pocket- or spoonshaped and can be fixed to tegulum by strong sclerotization. Epigynal plate strongly sclerotized without or with strongly fused median plate; epigynal plate sometimes with distinct, cave-like atrium. Posterior sclerite either absent or forming a large bulge or a strongly sclerotized transverse plate protruding ventrally (posteriorly of the copulatory opening). Epigynal teeth mostly present, originating posteriorly of the genital openings, but sometimes reduced or forming ‘pseudo teeth’ (= elongation of lateral margin of atrial region). Vulvae at least partly with irregularly sclerotized structure enclosing a convoluted duct or with appendages at the copulatory duct, possibly homologous with spermathecal head of Bennett (2006) and Sierwald (1989) (exception: E. sicana ). Fertilization ducts only represented by short, leafshaped appendages.

Comment

Comprising 17 species of which most are limited to Italy, France, and the Iberian Peninsula. Eratigena agrestis and E. atrica also occur in Central Europe. These species may have been introduced to North America and the UK.

Figures

Figure 1. Scanning electron microscopy pictures of spinnerets of female and other structures. A–D, Tegenaria domestica; E, F, Tegenaria parietina; G, H, Tegenaria dalmatica; I, Eratigena agrestis; J, Eratigena atrica; K, Aterigena ligurica; L-N, Malthonica lusithanica; O, Allagelena gracilens. A, L, overview; B, D, F, I, K, N, PMS; E, H, J, ALS; M, distal segment of PLS; C, G, colulus; O, tarsal trichobothrium. Abbreviations: ALS, anterior lateral spinnerets; CGS, cylindrical gland spigot; COL, colulus; mAS, minor ampullate gland spigot; MAS, major ampullate gland spigot; N, nubbin; PGS, piriform gland spigots; PLS, posterior lateral spinnerets; PMS, posterior median spinnerets; T, tartipore; TS, tracheal stigma.

Figure 2. Microscope and scanning electron microscopy pictures of character details. Plumose hairs (A) and claws (B) on tarsus I of Tegenaria dalmatica; C, embolus tip of Tegenaria ferruginea; D, male palp of Tegenaria dalmatica in ventroretrolateral view; E, retrolateral tibial apophysis in retrolateral view of Tegenaria ferruginea; F, epigyne in ventral view of Tegenaria domestica; G, long and dark spikes on the anal tubercle of Agelena labyrinthica; H, face and chelicerae, frontal view; I, cheliceral margins with teeth, ventral view; and J, sternum, ventral view, of female Eratigena atrica. Abbreviations: BT, basal portion of tegulum; C, conductor; CO, copulatory opening; CON, connection of tegulum and conductor; DB, dorsal branch; DC, distal portion of conductor; DP, dorsal part of terminal end of conductor; E, embolus; EP, epigynal plate; LB, lateral branch; LM, lateral margin of median region; MA, median apophysis; MC, lateral margin of conductor (here entirely folded); MP, median plate; MR, median region; PS, posterior sclerite (here bar- or band like); PT, ‘pseudo teeth’; TEC, terminal end of conductor (here bifid); VB, ventral branch; VP, ventral part of terminal end of conductor. Scale bars for F–J = 0.5 mm.