The Goblin Spider Genera Prodysderina, Aschnaoonops, and Bidysderina (Araneae, Oonopidae)

ABSTRACT A new genus, Prodysderina, is established for a group of Neotropical oonopids belonging to the Dysderina complex and characterized by having a laterally incised, tuberculate, but unridged sternum, a groove connecting the posterior (but not the anterior) spiracles, and a male embolus with an elongated distal prong and a reduced proximal prong. Dysderina armata Simon is transferred to Prodysderina and selected as the type species; eight new species are described from Venezuela (P. megarmata, P. rollardae, P. janetae) and Colombia (P. piedecuesta, P. rasgon, P. santander, P. filandia, P. otun). The genus Aschnaoonops contains species that resemble those of Prodysderina but have a twisted (and usually basally widened) embolus in males, and a reduced genital atrium in females. That genus occurs in the Andes from Peru north to Colombia, east across northern South America, and north into the West Indies. Dysderina similis (Keyserling) and D. propinqua (Keyserling) from Colombia, and D. simla Chickering from Trinidad, are transferred to Aschnaoonops, and females of the two Keyserling species are described for the first time. One new species, A. silvae, has been taken by canopy fogging and appears to be widespread in the Amazonian portions of Peru, Ecuador, and Colombia. A total of 36 other new, ground-dwelling, microdistributed species are described: A. yasuni, A. tiputini, A. cosanga, A. ramirezi, A. jatun, and A. marshalli from Ecuador, A. leticia, A. orito, A. pira, A. paez, A. huila, A. meta, A. alban, A. chingaza, A. pamplona, A. pedro, and A. marta from Colombia, A. chorro, A. indio, A. tachira, A. tariba, A. teleferico, A. jaji, A. merida, A. aquada, A. masneri, A. trujillo, A. cristalina, A. bocono, A. simoni, and A. margaretae from Venezuela, A. malkini, A. caninde, and A. belem from Brazil, A. villalba from Puerto Rico, and A. gorda from the Virgin Islands. Another new genus, Bidysderina, is established for a group of species resembling those above in sternal structure but having differently constructed male palps; five new species (B. perdido, B. bifida, B. niarchos, B. wagra, B. cayambe) are described from Napo province, Ecuador.


INTRODUCTION
The goblin spiders detailed below have received relatively little attention in the literature. Only a few species have been described, mostly misplaced in the genus Dysderina Simon. However, these animals lack the transverse ridges on the sternum, and the groove connecting the anterior spiracles, that characterize the members of Dysderina (see Platnick and Dupérré, 2011a). The distinctiveness of these taxa was first recognized by Dumitrescu and Georgescu (1987), who described one of them as ''Prodysderina spinigera (Simon). '' In attempting to establish the new genus they called Prodysderina, Dumitrescu and Georgescu included in it two species described by Simon, but because they failed to designate either of them as the type species, their generic name is a nomen nudum, and is not available. Of the two species, the one detailed first in their paper was ''Prodysderina spinigera,'' but it is just as well that they did not designate it as the type species, because the specimens they studied and illustrated, from Venezuela, do not actually belong to Dysderina spinigera Simon, which was originally described from Saint Vincent in the Lesser Antilles. Those Saint Vincent specimens belong to a different genus, Simonoonops Harvey (see Platnick and Dupérré, 2011b). Dumitrescu and Georgescu (1987: pl. 5) nevertheless presented useful illustrations of the species they misidentified as ''Prodysderina spinigera'' as well as of the second species they attempted to assign to their genus, Dysderina armata Simon. Even though Dumitrescu and Georgescu's concept of Dysderina was erroneously based on a Mediterranean (rather than South American) species, their efforts have provided, for other systematists, the basis for a working concept of Prodysderina. In describing the genus as new below, we therefore retain their original generic name, and thereby make it available (with new authors and a new date). As the type species, we choose the one that was correctly identified by Dumitrescu and Georgescu, D. armata. That species shows a number of peculiarities, including a laterally incised, tuberculate, but unridged sternum, a groove connecting the posterior (but not the anterior) spiracles, and a male embolus with a greatly elongated distal prong and a much shorter, thinner proximal prong (figs. 61-73). Eight apparently related new species are described below from Venezuela and Colombia. The taxon that Dumitrescu and Georgescu misidentified as ''Prodysderina spinigera'' shares the sternal and spiracular features of these Prodysderina species but differs in having a basally widened and twisted embolus in males, and a reduced genital atrium in females. Those features are shared with a wide range of species, treated below, that are here considered congeneric with the misidentified Venezuelan species. Their distributions range from Peru northward; the group appears to be most speciose in Colombia and Venezuela, but also extends into northeastern Brazil and even the West Indies.
Ineffective as Dumitrescu and Georgescu's attempt at establishing a new generic name may have been, their paper nevertheless contains careful descriptions and useful illustrations of the morphology of the specimens they studied. The same cannot be said for a more recent effort by Makhan and Ezzatpanah (2011) that also established a new generic name within the Oonopidae, Aschnaoonops, based on a single included species, Aschnaoonops aschnae, from Suriname. Their ''paper'' is perhaps best characterized as the taxonomic equivalent of medical malpractice. Those authors provided no differential diagnosis (i.e., no information identifying the putatively most closely related genus or genera, and indicating how their genus can be distinguished from those putative relatives). Their entire generic ''description'' reads as follows: Small brown species. Carapace round. Palp with a C-shaped projection. Underside of the projection strongly sclerotized and upper side soft, open and seed-like inside. Legs with large thick spines and hairs.
Their species ''description'' merely repeats this information, adding only a measurement of the single known specimen and a few uninformative comments on its coloration and setation. Six extremely low-quality photographs were presented; from the two habitus views, one can determine only that their male specimen is a hard-bodied oonopid. There is a ventral view that is so out of focus that one cannot determine anything at all about the structure of the sternum. A lateral view of a first leg shows that the femur, tibia, and metatarsus are each heavily spined. In concert, these figures suggest that the specimen belongs to the Dysderina complex, a placement confirmed by two photographs of the male palp.
Those authors claimed that the ''holotype will be deposited in the collection of the University of Suriname, Department of Entomology, Paramaribo, Suriname.'' As documented by workers on other groups of arthropods who have had the misfortune of having to deal with the ''contributions'' of Makhan, this claim appears to be science fiction (see the comments by Bolton et al., 2008, who charitably considered a work by Makhan to be ''one of the most inadequate papers that has ever been produced in ant taxonomy''). Our repeated requests to examine the holotype of A. aschnae have, unsurprisingly, been declined (see Jä ch, 2006, for details on Makhan's prior attempts to extort large sums from investigators seeking to examine his types).
Our first inclination was simply to treat Aschnaoonops as a nomen dubium, on the grounds that nothing in the published description or illustrations suffices to identify the genus, and that the type is unlikely ever to be available for examination. Although the name clearly refers to some member of the Dysderina complex, the generic description could easily apply to members of that complex as disparate as Paradysderina Platnick and Dupérré (cf. Platnick and Dupérré, 2011c: figs. 122-125) and Costarina Platnick and Dupérré (cf. Platnick and Dupérré, 2012: figs. 80-84). However, males of one of the new species described below from Pará, Brazil, have palps that appear to resemble those of A. aschnae, and on that basis we have decided to use the generic name to refer to the group of species that seem to be most closely related to A. margaretae, new species (which is the taxon that was misidentified as ''Prodysderina spinigera'' by Dumitrescu and Georgescu, 1987). Although the palpal photographs provided by Makhan and Ezzatpanah do not show a basally widened embolus, the Brazilian specimens, when seen in ventral or retrolateral views, clearly have that type of embolus, even though few traces of the widened basal portions are apparent in prolateral view (figs. 560-565).
Both genera include some species with anomalous characters. For example, females of six species of Prodysderina (P. armata, P. megarmata, P. piedecuesta, P. rasgon, P. filandia, and P. otun) have the dorsal scutum fused to the epigastric scutum (figs. 63, 130); based on the similar male palps of P. rasgon and P. santander, we predict that the unknown female of the latter species will also show this character, which is not found in females of the other two species of the genus (P. rollardae and P. janetae). In the males, these scuta appear to have fused only in P. rasgon and P. santander. Even odder is that the females of P. piedecuesta and P. otun have the postepigastric scutum fused to the epigastric scutum (figs. 108, 109, 674), a feature not found in any of the other females treated below (although it is ubiquitous in the males). Within this group of species, even some individual specimens are anomalous; one of the females of P. filandia has normal female genitalia and a normal right palp, but the left palp bears, at its tip, a reduced male bulb subtending an apparently normal embolus (figs. 140-143). Males of P. filandia are unknown, but the embolus on the teratological female (figs. 144, 145) corresponds well to those of the other known species in the genus! Similarly perplexing diversity occurs within Aschnaoonops. Although most species resemble those of Prodysderina in having a substantial spinneret scutum and a groove connecting the posterior spiracles, two species (A pedro, fig. 380; A. cristalina, fig. 508) have seemingly lost the spinneret scutum, and in the females of two species (A. paez, figs. 299, 301; A. cristalina, fig. 509) the groove that normally connects the posterior spiracles seems to have moved anteriorly and lost its spiracular connections (males of A. paez are unknown, but those of A. cristalina have only a weak, almost obsolete, groove connecting those spiracles). Some reduction of the spinneret scutum also appears in A. jaji; although females of that species have a distinct scutum ( fig. 456), males show only a slightly sclerotized rim, bearing the usual elongate setae. All these somatically anomalous species nevertheless have genitalia that seem typical for the genus.
In Aschnaoonops, the dorsal and epigastric scuta appear to have fused only in the females of A. cosanga, A. indio, A. teleferico, A. trujillo, A. simla, and A. villalba, and in both sexes of A. tariba. The two known males of A. marta resemble those of some species of Paradysderina in showing asymmetry between the left and right palps; the right palpal bulb is much less ''inflated'' than the left, and there are also some slight but consistent differences in the shape of the embolus on the right and left palps (figs. 394-399). The single known female of A. villalba is also anomalous; the female genitalia are greatly reduced (figs. 588, 589) and the posterior legs have spines on the prolateral sides of tibiae and metatarsi III and IV that have not been detected on any other specimens. However, the right and left posterior legs show different spination patterns, and we suspect that this specimen is teratological.
Some species groups can be recognized within the large genus Aschnaoonops. A group of seven Venezuelan species (A. tachira, A. tariba, A. teleferico, A. merida, A. aquada, A. masneri, and A. trujillo) are united by having an extremely complex embolus, with multiple processes, accompanied by a triangular projection on the retrolateral surface of the cymbium (as in figs. 430, 441). Interestingly, A. villalba,from Puerto Rico,shares these characters (figs. 586,587) and seems to belong to the tachira group. Four species (A. meta, A. similis, and A. chingaza from Colombia, plus A. belem from Brazil) share a more apically situated projection on the male palpal cymbium (as in figs. 320, 332). Two species (A. ramirezi and A. marshalli from Ecuador) share a deep notch on the ventral pedicel margin of females (figs. 254, 270), but that feature has apparently been acquired independently by two members of the tachira group ( A. merida and A. trujillo, figs. 468, 497), as well as by one species of Prodysderina (P. otun, fig. 674).
A third genus, Bidysderina, is established below for a small group of species with similar sternal structure, but with very different male genitalia. The palpal bulb is small, not expanded as in more typical members of the Dysderina complex, and the embolus is almost entirely bifid (figs. 627, 660), producing the appearance of a separate embolus (presumably corresponding to the distal prong) and conductor (presumably corresponding to the proximal prong; this conductor is sometimes bifid as well). The male endites are distinctively modified, with a complex distal process ( fig. 595). The female genitalia still retain an atrium, but it is short and small (figs. 630, 652). The five species assigned to this genus are known only from Napo province, Ecuador; although they have been taken on the ground (e.g., in pitfall traps), they have also been collected by beating foliage and even by canopy fogging, and at elevations as high as 3850 m.
ETYMOLOGY: The specific name is a noun in apposition taken from the type locality. DIAGNOSIS: Males resemble those of A.