Notes on Mediterranean Theridiidae (Araneae) – II

Taxonomic and faunistic amendments are provided for 15 species and one subspecies of comb-footed spiders (Th eridiidae) of the Mediterranean region, in the genera Anatolidion, Episinus, Heterotheridion, Th eridion and Th eridula. Th e following taxonomic changes are proposed: Anatolidion osmani Wunderlich, 2008 and Th eridion crinigerum Simon, 1881 are synonymised with T. gentile Simon, 1881, making it the type species of the monotypic genus Anatolidion Wunderlich, 2008. Episinus albescens Denis, 1965 is synonymised with E. algiricus Lucas, 1846, Th eridion xinjiangense (Hu & Wu, 1989) with Heterotheridion nigrovariegatum (Simon, 1873). Th eridion aelleni Hubert, 1970 is removed from synonymy of Th eridion spinitarse O. P.-Cambridge, 1876 and transferred to Th eridula. Th e recent transfer of Th eridion pinicola Simon, 1873 and T. genistae Simon, 1873 into Paidiscura has to be rejected. Th eridion genistae turanicum Charitonov, 1946 from Uzbekistan is raised to species level. New faunistic records are presented for Th eridion pinicola from North Africa, Anatolidion gentile, Th eridion genistae and T. hemerobium from Greece. Several poorly known (sub-)species are redescribed: Anatolidion gentile, Episinus maculipes numidicus Kulczyński, 1905, Th eridion genistae, T. glaucinum Simon, 1881, T. musivum Simon, 1873, T. pinicola, T. pyrenaeum Denis, 1944, T. semitinctum Simon, 1914 and T. spinitarse O.


Introduction *
In the Mediterranean region spider taxonomy has been far less thoroughly considered and revised than in central and northern European countries, mainly owing to a lack of continuous research (Th aler 2000).Similar defi ciencies in the current knowledge of spider diversity are also still substantial among the Th eridiidae (Knofl ach and Th aler 2000).About half of the roughly 280 comb-footed spider species described from Europe and the Mediterranean, including North Africa and Macaronesia, are known from one sex only and many of them have not been found again since their fi rst discovery (Knofl ach and Th aler 2000).Further problems arise in the long list of "species inquirendae", which are diffi cult to interpret.As many theridiids show large areas of distribution, numerous synonyms may be expected.Th is contribution is the continuation of a stepwise approach and concept to improve taxonomic and faunistic insights on comb-footed spiders in this region.We re-examined miscellaneous representatives of the genera Anatolidion, Episinus, Heterotheridion, Th eridion and Th eridula, with most species having been described by the French arachnologists Eugène Simon, Jacques Denis and Hippolyte Lucas, and we present new records and comparative remarks.As major taxonomic changes, four new synonymies are here proposed and a previous synonym is revalidated.

Material and methods
Specimens were examined using a Leica Wild M8 stereoscopic microscope with a micrometer eyepiece.Male and female genitalia were dissected and studied as temporary mounts by submerging them in glycerine and Hoyer's compound solution (Kraus 1984) on half-covered slides under a Wild M20 microscope with a drawing tube.Living spiders were photographed with a Nikon F3, Medical-Nikkor 120 mm lens, ring fl ash and a teleconverter.All measurements are in mm.
strong, as indicated by the naming of Simon's female as T. crinigerum.For further details see Simon (1881) and Wunderlich (2008).Surprisingly, Dalmas (1922: 87, sub T. crinigerum) mentions an apical spur on femur IV in the male "L'éperon apical du femur IV du mâle est une particularité très remarquable de cette espèce.".Th is particular spur was not visible in the males examined and was not mentioned by Simon (1881: 107).
Somatic features, colouration (Figs 22,(25)(26).Habitus, colour and pattern similar to E. maculipes.Sternum dark, its light median stripe narrower than in E. maculipes, sometimes even missing.Carapace with brown median band and reticulate pattern at margins.Th ese and leg markings more diff use than in E. maculipes.Venter and epigaster usually dark or greyish.
Selected comparative remarks, diagnosis.E. maculipes is well characterised by numerous authors, for details of morphology and distribution see Knofl ach and Th aler (2000).Some features are given here for identifi cation and diff erentiation compared to E. algiricus: Prosoma larger than that of E. algiricus.Sternum dark, with marked light median stripe, which widens anteriorly.Carapace with clear brown median band and reticulate pattern at margins .Legs conspicuously speckled.Venter pale.Male palp larger than that of E. algiricus (Figs 21 vs. 22; see also Kulczyński 1905), in ventral view 0.6-0.7 mm broad, 0.9-1.0mm long (distance between distal rim of tibia and tip of cymbium).Palpal tibia 1.5 times wider Distribution.Th is widespread expansive Mediterranean species (for details see Knofl ach and Th aler 2000) has recently been reported from Crimea, which is the easternmost record (Kovblyuk et al. 2008).Episinus maculipes numidicus Kulczyński, 1905Figs 19-20 E. m. numidica Kulczyński, 1905: 437, pl. 11, fi gs 4, 17, male, female, type region North Africa.
Description, identifi cation.Kulczyński (1905).Taxonomic status.Considerations on the subspecifi c rank have to be postponed owing to lack of material, especially males.
Somatic features, colouration.Th e single female shows a similar colour pattern to E. maculipes, except for the sternum, which lacks a light median stripe.
Epigynum/vulva .Epigynal cavity considerably wider than long, 1.8 times in the examined female.Cavity 0.24 mm wide, about as long as receptacula seminis.Median septum broad and rather short.
Diff erentiation from E. maculipes.As indicated by Kulczyński (1905) Episinus m. numidicus is smaller than E. maculipes.Th e following features agree well with the description of Kulczyński (1905): Sternum almost completely dark, without light median stripe.Median septum of female epigynum considerably broader and shorter than in E. maculipes (Fig. 19 vs. 14; see also Kulczyński 1905, fi gs 17 vs. 18).Ratio width to length of epigynal cavity larger in E. m. numidicus.Cavity about as long as receptacula, while in E. maculipes it is 1.2-1.3times longer than receptacula.

Episinus theridioides
Synonymy.Th e synonymisation of E. pyrenaeus (Simon, 1914) with E. theridioides by Bosmans and de Castro (2002) is supported by the present material from the Pyr-enees and from Corsica, though only females have been compared.General appearance (Fig. 27), shape of epigynum and overall course of copulatory ducts (Figs 28-29) are not or barely diff erent from E. theridioides .
Distribution.Episinus theridioides occurs discontinuously in the Mediterranean (Knofl ach and Th aler 2000) and is known only from a few localities in the French and Spanish Pyrenees, Cantabria, Corsica and Sardinia; for a distribution map see Bosmans and de Castro (2002).Unlike members of the E. truncatus-group it appears to be an exclusively epigeic species.
Generic placement.Wunderlich (2008) established a new monotypic genus Heterotheridion for this strikingly coloured species  owing to its genital characters, such as the palpal tibia being elongated into a slender outgrowth, the cymbium with an apical, scaly bulge and a distal embolus directed clockwise (Fig. 59).Furthermore, the anterior median eyes are smaller than the posterior medians, the posterior eye row is procurved and the legs are rather long (Wunderlich 2008).According to its mating behaviour, H. nigrovariegatum does not correspond to representatives of the genus Th eridion (Knofl ach 2004).Time and duration of sperm induction and number of insertions are clearly diff erent.Sperm induction takes place independently of copulation and lasts correspondingly longer.Copulation usually consists of two insertions only (Knofl ach 2004).
Synonymy.Th eridion xinjiangensis (Hu & Wu, 1989) is synonymised with H. nigrovariegatum from the literature (fi gs 121 in Zhu 1998) because of the following distinct characters: Conformation of male palp and epigynum very similar (Figs 44,59).Tibial and cymbial outgrowth typical, bulbus genitalis fully corresponding; conductor characteristically pointed, embolus clockwise, sperm duct highly convoluted within voluminous subtegulum and tegulum, median apophysis inconspicuous and hidden by cymbium (Fig. 59).Epigynum and vulva of Th eridion xinjiangensis largely resemble those of H. nigrovariegatum in shape of copulatory orifi ces, coiled course of copulatory ducts and relatively large receptacula (Fig. 44).Th e only apparent morphological diff erence concerns the opisthosoma, which ends in a small tubercle in T. xinjiangensis (see fi g. 121B in Zhu 1998).Specimens examined from Uzbekistan strengthen this synonymy.Also the type province Xinjiang is not so far distant from the hitherto known eastern records in Kyrgyzstan and Tajikistan.
Somatic features, colouration .Small Th eridion species, carapace and sternum dark brown, legs annulated and abdomen with numerous dark dots and characteristic whitish folium, which widens at midline and fades within whitish area (Fig. 34).Female epigastric region bulging.For details see Simon (1873a).
Male palp .Tibia rather small and short, with one retrolateral trichobothrium.Bulbus largely developed (see also Fig. 35).Embolus forms a long, thread-like spiral.Embolar base with knob-like locking device.Conductor a large, membranous, folded structure with furrow guiding embolus.Median apophysis inconspicuous, with a few scales, hidden within cymbium.

Th eridion glaucinum
Somatic features, colouration.Carapace light, with narrow dark margins and dark central patch.Sternum dark.Legs light yellow with dark annulations.Abdomen in this faded specimen beige, a few faint dots on dorsum and a small dark longitudinal patch on venter.Spinnerets light, surrounded by dark pigmentation.For details see Simon (1881).
Epigynum/vulva .Epigynal cavity wider than long, its width 0.09 mm at midline; its overall shape trapezoidal, as anterior and posterior borders unevenly wide.Copulatory ducts rather short, ca.0.2 mm long.Th ey diverge sideways and form a short inwards coil towards the receptacula thereby gradually narrowing.Receptacula seminis rather elongated.
Affi nities.Th eridion glaucinum appears to be closely related to T. petraeum L. Koch, 1872.General appearance and epigynal characters are rather similar.Females of T. petraeum are comparatively larger, show an evenly transverse-oval epigynal cavity, copulatory ducts of 0.5 mm length and globular receptacula.As in other representatives of this species group, the epigynum of T. glaucinum is fi lled with mating plug secretions (see Knofl ach 1998).
Distribution.Known only from the type locality.No further reference noted since its description.and present data from Tuscany and Emilia-Romagna).In northern Italy it may almost meet its eastern vicariant Th eridion refugum, another representative of this still insuffi ciently known, ground-living, lapidicolous species complex.Th e allopatric occurrence of these closely allied species indicates diff erent glacial refugia and respective reinvading processes.In contrast to its sibling T. pyrenaeum, T. hannoniae appears to be  restricted to lower altitudes; highest localities being about 800-1100 m.T. hannoniae occurs under stones in natural boulder fi elds and scree, but also in human debris, such as quarries, dikes and railway constructions (Bosmans et al. 1994;Staudt 2003).
Male palp .Palp less elongated and considerably smaller than in T. pictum, length of tibia and tarsus ca.0.4 mm (versus ca.0.7 in T. pictum).Tibia rather broad at base as compared with distal rim, about 0.7 of distal width in ventral view, thus little constricted.Shape of conductor and median apophysis diagnostic.Prolateral tip of median apophysis closer to tip of conductor than in T. pictum.Embolus short, distal part 0.12 mm long (T.pictum 0.3 mm, see Knofl ach 1998).
Distribution.Th eridion hemerobium is widespread in North America (Levi 1957a) and Europe (Bosmans et al. 1994;Anthes 2000).From Greece it was hitherto not known (Bosmans et al. 1994;Anthes 2000;Bosmans and Chatzaki 2005).Th e present fi nding bridges a distribution gap in southeast Europe, though its occurrence is not surprising.A further record comes from Bulgaria (Deltshev 1992); the easternmost one from Israel (Levy 1998).Interestingly, the species has so far not been mentioned from Russia (Mikhailov 1997(Mikhailov , 1998)).T. hemerobium occurs stenotopically on the vegetation in wetlands and on banks of lakes, ponds and running water (Anthes 2000) and also colonises human-made structures in these habitats, e.g.bridges and fences (Jones 1994;Daws 2003).
Somatic features, colouration .Overall colouration of carapace, sternum, gnathocoxae and chelicerae bright red in living specimens.Th is striking colour fades in specimens preserved in alcohol.Legs and palps lighter, with distal femora, patellae and distal tibiae reddish, but no dark annulations.Abdomen uniformly dark brown to ruby coloured.Male epigaster sclerotised, seminal vesicle (part of male genital system) conspicuously dark, translucent.
Male palp .Tibia very short, wider than long, with one retrolateral trichobothrium.Embolar base with anchoring process for tegular notch, as in other Th eridion species.Distal part of embolus rather short (0.2 mm long) and stout.Conductor covered with minute scales.Th eridiid tegular apophysis straight and pointed, closely adjoining conductor.Median apophysis sickle-shaped.
Male palp .Conformation of male palp as in other representatives of the Th eridion varians group (see Knofl ach 1998).Tibia short, with one retrolateral trichobothrium.Conductor pointed and curved.Prolateral part of median apophysis triangular and strongly protruding.Th eridiid tegular apophysis pointed.Embolus slender.Distal part of embolus 0.25 mm long.
Epigynum/vulva .Epigynal cavity rounded, its posterior border strongly sclerotised; with central copulatory orifi ces fused to common, rounded opening.Copulatory ducts ca.0.4 mm long.Th ey diverge laterally, forming a few coils, turn inwards and enter the receptacula by a fi nal wide coil.In mated females the epigynal cavity is fi lled by plug secretions.All parts of vulva clearly translucent through integument.
Copulatory behaviour.Copulation follows the pattern of the Th eridion varians group, with 6-8 sperm inductions, a presumed initial pseudocopulation and a fi nal mating plug sequence (Knofl ach 2004).
Generic placement: Th e generic placement of Th eridion pinicola in Paidiscura by Wunderlich (2008) has to be rejected, since there is no convincing argument for such a combination.T. pinicola is clearly a member of the T. varians group, see above and Knofl ach (2004), for Paidiscura see Knofl ach and Th aler (2000).

Th eridion pyrenaeum
Description, identifi cation.Denis (1944), Bosmans et al. (1994).Taxonomic status.Th e statement of Bosmans et al. (1994: 238) ".. further material of T. pyrenaeum is needed to allow a biometric study of some parts of the palp and epigyne to decide about the status of this species" indicates its close similarity to T. hannoniae and the arising problem of interspecifi c/intraspecifi c classifi cation.Th is requirement remains.Judging from the present specimens, T. pyrenaeum is larger in body and leg size, although in the females this was not always clearly reliable (Fig. 56).However, the dimensions of the male and female genitalia (cymbium length and width of epigynal cavity) from various locations show consistent diff erences.
Epigynum/vulva .Overall genital morphology as in T. hannoniae, distinguished by size and proportion of following parts: Epigynal cavity larger than in T. pyrenaeum, 0.12-0.15mm wide (n=11), its width exceeding the distance to outside of ducts (x, Fig. 53).Receptacula seminis as long as or shorter than width of epigynal cavity.
Distribution.According to the current state of knowledge Th eridion pyrenaeum appears to be endemic to western European mountain systems, in allopatry with the widespread T. hannoniae.Up to the present, T. pyrenaeum has been found only in the French and Spanish Pyrenees and Sierra Nevada (Spain).Th ere it is confi ned to the higher zones, between 2500 and 3130 m altitude in Sierrra Nevada, about 2000 m in the type region Andorra (Denis 1957) and between 1600 and 2400 in the French Pyrenees (present paper).Ecological preference and altitudinal zoning resemble that of Th eridion petraeum in the Alps.Simon, 1914 Figs 68, 76-77 Material examined.Spain: El Pardal (CM), 3 ♀ (MHNP AR 2298).Italy: Lombardia, Lake Lugano, Porlezza, 1 ♀ (CTh ), 30.6.1962(Th aler 1966, Abb. 2d, sub Th eridion sp.).

Th eridion semitinctum
Description, identifi cation.Th is species has not been reported since the description of Simon (1914: 270, 297).A more detailed redescription including that of the unknown male is planned in comparison with the other members of the Th eridion melanurum group (Knofl ach, in preparation).For comparative analysis and illustration of females see Th aler (1966).
Somatic features, colouration.Th e species is clearly a representative of the Th eridion melanurum-group (Th aler 1966) and shares its general appearance, colour pat-tern and overall conformation of genital organs with Th eridion betteni, T. melanurum, T. mystaceum, etc. Abdomen with clear folium.
Somatic features, colouration.Specimens from Tunisia uniformly pale yellow, probably faded (but see Hubert 1970).Th e female from Cadiz shows an indistinct pattern: Carapace light brown with faint dusky margins and median band.Sternum dusky grey.Legs uniformly pale yellowish.Abdomen grey, with three pairs of pale transverse, oval patches, arranged in diagonal paramedian rows.Venter uniformly grey.Spinnerets contrastingly light.Male chelicerae longer than in female.Abdomen globular, a little longer than wide, without protuberances.For further details see Hubert (1970).
Male palp .Tibia rather asymmetrical, retrolaterally fairly extended (Fig. 80), but prolaterally excavated (Fig. 82), its distal rim in ventral-prolateral view oblique and almost reaching patella (Fig. 81).Two trichobothria present on retrolateral side of tibia.Cymbium elongate and slender, its base occupying prolateral side of palp, where it is markedly incised.Cymbial hood in dorsal-median position.Subtegulum and tegulum constitute the main part of genital bulb and are densely crossed by wide loops of sperm duct.Th e embolus forms a straight element inserted deeply within the tegulum.Subtegulum with strongly protruding sclerotised basal shaft, which is embedded within a cavity and surrounded by a noticeably developed basal haematodocha.Distal part of embolus conspicuously screwed.Embolar base encircled by a membranous apophysis, which interlocks with cymbial hood and thus is interpreted as the median apophysis.
Epigynum/vulva .Copulatory orifi ces roughly circular, ca.0.6 mm wide, with marked, sclerotised outlines.Th ey are clearly separate, but less than their diameter apart.Th eir distance to epigastric furrow equals about their diameter.Copulatory ducts rather short, forming a small coil, at entrance rather wide, narrowing towards receptacula.Receptacula seminis barely larger than copulatory orifi ces.
Taxonomic remarks, generic placement.Th e former synonymy of Th eridion aelleni with T. spinitarse by Brignoli (1984: 301) was concluded from illustrations only and cannot be supported here.Along with the present analysis of type material the resurrection of T. aelleni has to be confi rmed (for comparison see Th eridion spinitarse above).However, the generic placement among Th eridion evidently implies some diffi culty.Th e new allocation off ered here may be surprising and not fully satisfactory when considering the general appearance of hitherto known Th eridula species, which is known to be striking, mainly through a tuberculate and wide abdominal shape, often combined with bright colour.Especially the unmodifi ed abdomen of T. aelleni at fi rst glance argues against placement within Th eridula, but would suggest resemblance with the only hitherto known species of Paratheridula (see Levi 1957b).However, a transfer membrane) is reported to be completely missing, which appears to be the main criterion for its separation from Th eridula (Levi and Levi 1962).
Distribution.Th eridula aelleni is so far known only from the type locality in Tunisia and from Spain.