A new ladybird spider from Hungary (Araneae, Eresidae)

Abstract According to the most recent taxonomic literature, three species of the genus Eresus are known in Central Europe, Eresus kollari, Eresus sandaliatus and Eresus moravicus. We recognized a fourth distinctive species from Hungary, which is described as Eresus hermani sp. n. Eresus hermani has an early spring copulation period, females have a light grey (grizzled) cephalothorax due to a heavy cover of lightly colored setae, and an epigyne with large flat areas posterior to the epigynal pit, while males are distinguished by a broad and blunt terminal tooth of the conductor. An updated and modified comparative table of Řezáč et al. (2008) to include all four Central European Eresus species, and a simple key to the species group’s species are given. Habitus, epigyne, vulva and conductor of Eresus kollari, Eresus moravicus and Eresus sandaliatus are also illustrated. An annotated list of papers illustrating Eresus hermani due to misidentifications is presented.


Introduction
The velvet spiders (family Eresidae) are among the most attractive spiders in Europe. The family contains nine genera and 96 described species worldwide. The genus Eresus All specimens of the new species examined, including holotype and four paratypes, have been deposited in the Soil Zoological Collection (former Arachnoidea Collection) of the Department of Zoology, Hungarian Natural History Museum (HNHM) Budapest (curator Dr. László Dányi).
Specimens and copulatory organs were studied using a Leica MZ FL III stereomicroscope and photographed by Canon Q Imaging Micro 5.0 RTV at the Institute of Genetics, BRC. Scanning electron micrographs were taken with a Hitachi S-4700 microscope at the Department of Applied and Environmental Chemistry, University of Szeged, Hungary.

Taxonomy
Remark. The genus Eresus in Central Europe has a long and difficult nomenclatural history. Some available "old names" were examined, such as Eresus illustris (presently considered nomen dubium, specimens are irretraceable), which is marked as possibly Hungarian (despite the fact Koch himself wrote "Vaterland: Unbekannt" [trans. Locality: Unknown]), but discarded it on the basis of the description and color image (Koch 1838, fig. 317), where the male has six black dots on the opisthosoma and only the dorsal side of hind femora as red, whereas E. hermani males have only four dots and clearly red hind legs patellae and tibiae, without any black, and tarsi and metatarsi are brownish grey (Fig. 1B). The female of E. illustris is unknown. The other possible candidate, E. fulvus Rossi 1846 (type specimens can no longer be found in NHMW), described by female specimens only, can also be excluded as a potential synonym, since they all have a large area covered by yellow/orange setae on the cephalothorax ["nitide fulvus" in the description of Rossi (1846)], whereas E. hermani females have no truly yellow setae on the prosoma at all; instead, its dorsal cephalothorax is light brownishgrey overall. According to Řezáč et al. 2008 (page 275.) E. fulvus Rossi differs from E. moravicus by "having spermatheca that are less lobed, and having copulatory ducts that are almost horizontal in the centre of the vulva." By contrast, spermathecae of E. hermani are rather conspicuously lobed, at least as much as in Eresus moravicus (Figs 4C,F and 5B,D).
Etymology. Dedicated to Ottó Herman (1835-1914, the Hungarian arachnologist and polymath, who first recognized color variants within Hungarian Eresus forms, to commemorate the 100 th anniversary of his passing. Diagnosis. Females of this species differ from all other Eresus females by the carapace's short, off-white to light brown hairs, intermingled with small clumps of long, black hairs, giving a light, grizzled appearance to the prosoma, and by an epigyne with a pair of flat plateaus adjoining the sides of the broad median lobe laterally. Males are characterized by the narrow groove and blunt, broad terminal tooth of the conductor, and distinguished from other Eresus species, except E. moravicus, by having almost entirely red hind legs. They differ from E. moravicus males by having red color on the thoracic dorsum only laterally, having a less prominent cephalic region with an almost flat area between PLE and PME, and by narrower strips of white setae on L I. This species has an early spring copulation period, and exhibits a marked difference in the sizes of the sexes: males are relatively small, while females are comparatively large among Central European Eresus spp. (Table 1).
Description. Male. Prosoma (Fig. 1B): Length 2.9-4.1 (mean 3.4, N = 15) Prominent, color dark ferruginous brown, covered by long, black hairs intermingled with scattered, short, white ones. Cephalic region barely broader than thoracic part, weakly broadening towards the front, steeply raised posteriorly, but area between PME Color of prosoma black, sparsely sprinkled with off-white to light brown setae, more heavily anteriorly ( Fig. 1E) black, sparsely sprinkled with off-white to light brown setae, more heavily anteriorly (

Note:
Without exception, the epigyne of Eresus moravicus specimens that we studied match those in Fig. 2L-P in Řezáč et al. (2008), but differ slightly from that shown in Fig. 2K (Řezáč et al. 2008), which seems to be depicted also as a drawing in Fig. 4H (Řezáč et al. 2008). The main difference is the direction of the anterior portion of fissures, which are typically directed slightly laterally, instead of medially. To aid differentiation of E. moravicus, we provide comparative photographs and a drawing of E. moravicus epigyne in Fig. 4D-E and Fig. 5C, which we believe to be typical of the species. Table 1.

Řezáč, 2008
Females Prosoma length 6.6-9.9 (mean 8.2) 5.9-9.9 (mean 7.5) Color of prosoma entire prosoma grizzled light brown due to a heavy cover of offwhite to light brown setae ( Fig. 1A) black, except orange anterior ( Color of hind legs red, tarsal joints brownish grey (Fig. 1B) red, tarsal joints brownish grey (Fig. 1D) White transverse stripes on Leg I-II narrow, covering only the distal edge of segments (Fig. 1B) broad at the distal part of segments, usually extending to the proximal end of next segment (Fig. 1D) Red color on thoracic dorsum only on flanks, at most a few red hairs posteriorly (Fig. 1B) extends to the middle, at least posteriorly (Fig. 1D) Conductor in lateral view wrinkled, clearly wider than long (Fig. 3B) wrinkled, somewhat longer than wide (Fig. 3E) Terminal tooth of conductor strongly incurvated, broad and blunt (Fig. 3B, C) strongly incurvated, narrows to a relatively pointed tip ( Fig. 3E, F) Groove of conductor in lateral view deep, narrow, ν (Greek nu) or narrow U shaped (Fig. 3B.) round (Fig. 3E.) and PLE nearly flat. Thoracic part bordered laterally by narrow red stripes, never extending to posterior dorsum. Chelicerae: Blackish-brown, covered by long, nearly adpressed black hairs; basal half with scattered white hairs on the front.
Legs: Legs I-II dark orange-brown with black hairs; Fe II and Pt II orange with red hairs, Ti II often with a dorsal patch of red hairs. Distal edges of Fe, Pt, Ti and Ta with narrow, white stripe dorsally, usually not extending to the proximal part of the next distal segment. Legs III and IV largely orange, covered with red hairs, Ta and Mt dull grayish-brown due to a mixture of reddish and black hairs, except for a proximodorsal patch of red on Mt.
Opisthosoma (Fig. 1B): Dorsally red with scattered white hairs except for two pairs of black spots. Red area and black spots seamed by a more-or less continuous line of white hairs. Ventral side of opisthosoma black with the exception of some red hairs on the branchial opercula.
Palps (Fig. 3A-C): Conductor broad, strongly wrinkled. Terminal tooth broad and blunt, somewhat longer than the lamella, with a strong, sudden bend at the base or somewhat more distally. Groove deep, narrow, ν (Greek nu) or narrow U shaped at the base in lateral view. Inner, spiny lamella high, about as high as terminal tooth.
Female. Prosoma (Fig. 1A): Length 6.6-9.9 (mean 8.2, n = 21), prominent, especially the cephalic region, dark orange-brown with a heavy cover of short, off-white to light brown hairs and with scattered, small clumps of long, black hairs giving a grizzled appearance.
Chelicerae: Dark orange brown, front of basal 1/3-3/4 same color as prosoma. Legs: Rusty red, Fe, Pt, Ti and Mt of all legs covered by black hairs with pale brown hairs scattered among them, the latter gradually decreasing in number from L I to L IV, usually clustering to form indistinct cross bands dorsally at the distal edge of each segments. Ta usually black, except for a small cluster of pale hairs basally.
Palps: Similar in color to L I. Opisthosoma (Fig. 1A): Brownish-black, covered by long black hairs with a scattering of short pale hairs at its anterior.
Epigyne (Figs 4A, 5A, 6A): Moderately deep, median lobe broad (ratio between the greatest width of ML to the greatest width of epigyne: 6:10), considerably flared posteriorly, reaching well over the posterior margin of the epigynal pit. Posterior edge of the epigynal depression not reaching posterior epigyne, but followed by a pair of flat, somewhat wrinkled plateaus adjoining the fissures laterally. Posterior part of fissures inclined towards the midline, turning parallel to the longitudinal axis before the short, incurved anterior tips.
Habitat. Edges of a local variety of downy oak scrub woodland (Ceraso mahaleb-Quercetum pubescentis) and the interim zone between calcareous open rocky grasslands (Seselio leucospermi-Festucetum pallentis) and degraded scrubland. Phenology. Eresus hermani matures in August-September, wandering males can be found from the end of March to the end of April (inferred copulation period) and females lay eggs in June. This phenology clearly sets Eresus hermani apart from the other Hungarian Eresus species: E. moravicus matures in late spring and mates in early summer, while E. kollari matures in late summer -early autumn, immediately followed by a copulation period in autumn. The phenology of Eresus hermani is essentially the same as that of E. sandaliatus (Řezač et al. 2008), which, however, does not occur in Hungary or within the Carpathian Basin.       Cs. Szinetár (2006): p. 23. Fig. 3 The caption of this figure says "Female Eresus cinnaberinus", but, in fact, the picture shows a female Eresus hermani sp. n., as is evident from the heavy cover of light setae on the prosoma and the base of chelicerae.

Kovács et al. (2010): figure 1C-F figure 2D
According to captions, fig. 1C-F of this paper depict the genital organs of female Eresus kollari. However, the anterior part of fissures of the epigyna are nearly parallel, epigynal pits are followed by large flat plateaus at the sides of median lobes, anterior copulatory ducts are round and weakly sclerotized, spermathecae strongly lobed, all features that distinguish Eresus hermani sp. n. unambiguously. Additionally, the epigyne shown in fig. 1E is grossly malformed, having supernumerary rudiments of fissures, a kind of abnormality frequent among females raised in captivity. Figure 2D is labeled as female Eresus kollari. Again, this figure shows a female Eresus hermani sp. n., as evidenced by the dense cover of lightly colored setae on the cephalic region and basal segments of chelicerae. The reason for these misidentifications is that at the time of writing, the authors (including the corresponding author of the present paper) considered females of Eresus hermani sp. n. as merely an extreme local variant of Eresus kollari. (Note: by contrast, fig. 2F. indeed shows a female Eresus kollari next to a male of the same species, as can be judged by the sparsely distributed light setae on the prosoma.) Miller et al. (2012): figure 2A Figure 2. A. of this paper is mislabeled as Eresus kollari, whereas in fact it depicts a female Eresus hermani sp. n. Again, the true identity of the specimen shown in this picture is revealed by the light color of the prosoma and basal chelicerae. The obvious reason for the misidentification is that at the time of the completion of this Atlas, the concept of Eresus hermani sp. n. as a discreet species was not yet formed. In this paper, figure 6. shows a female Eresus hermani sp. n. mislabeled as Eresus kollari. Heavy cover of the prosoma by lightly colored hairs gives away the identity of the depicted specimen.