Nemastomatidae Simon, 1872 Jürgen Gruber Etymology: Nemastoma, from Greek Nema (Thread) and Stoma (Mouth), in Allusion to the Thin Pedipalps

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a closer relationship between Ischyropsalis and Ceratolasma (alone) were in part in- terpreted as plesiomorphies (e.g., form of sternum and endites, simple penis); but other supporting characters, such as structure of pedipalps (where independent loss of plumose setae occurred twice in the hypothesis) and ovipositor, or midgut anatomy (Dumitrescu, 1975a, 1980), were disregarded. In the combined morpho- logical and molecular analysis of Giribet et al. (2002) and in the molecular analysis of Shultz and Regier (2001) Ischyropsalis is sister to Ceratolasma, but not to Hesper- onemastoma. So the earlier hypothesis that Ceratolasma and Acuclavella from western North America are the closest relations of Ischyropsalis is still tenable. Taracus with its likewise elongated chelicerae and similar chelae dentition is not closely related. Main references: • Systematics: Dresco (1966), Martens (1969a, 1978b), Martens et al. (1981), Prieto (1990a,b), Luque (1991). • Natural history: Juberthie (1961a, 1965, 1974), Martens (1969a,c, 1975a,b, 1978b).

Nemastomatidae Simon, 1872 Jürgen Gruber Etymology: Nemastoma, from Greek nema (thread) and stoma (mouth), in allusion to the thin pedipalps. Characterization: • Size: Body length between 1.2 and 5.6 mm. • Dorsum: Flattened to strongly arched. With scutum magnum (Figures 4.16a–c) or scutum parvum with tendency to fusion (Ortholasmatinae, Figure 4.16d). In some species a scutum compositum develops by secondary sclerotization of membranes posterior and lateral of the primary scutum (in the adult stage) to include also at least one of the free tergites. Ocularium near anterior border of scutum, in some forms with processes, elaborated into a hood (Ortholas- matinae, Figure 4.16d) or divided into two columns; reduced in troglobites. Three laminae suprachelicerales generally well developed and delimited from carapace. Ozopores not freely visible. Scutum ﬁnely or coarsely granulated, variously sculptured or armed with macrosculptural elements—nearly ﬂat, with low segmental humps on areas (Figure 4.16c), with cone-shaped tubercles (Figure 4.16a), with low or high pegs or spines, in some groups with rows or a network of connected anvil-shaped tubercles (Figures 4.16b,d). • Venter: Corona analis well developed, as are free sternites; anterior sternite narrowed to base of genital operculum. Genital operculum with well-developed basal suture in females, without it in males. Spiracles not visible. Prosomal s– sternal area fused with coxae. Leg coxae well delimited but immovable, with o– marginal rows of tubercles, often anvil shaped (Figure 4.16c). l– 29859_U04.qxd 8/18/06 12:28 PM Page 149

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ab c

g e

f Figure 4.16. Nemastomatidae. Nemastomatinae, males, habitus: (a) Paranemastoma superbum, dorsal; (b) d h Mitostoma gracile, dorsal (body length about 1.5 mm); (c) Nemastoma bidentatum sparsum, lateral. (d): Ortholasmatinae: Dendrolasma mirabile, male, habitus dorsal (body length about 2.9 mm); (e) Ortholasma rugosum, right chelicera of male, prolateral view; (f) same for Gilarovia turcica. (g–h): Right ij k l m n pedipalps of males, prolateral view: (g) O. rugosum; (h) G. turcica, showing stridulatory area. Penes: (i) Mitostoma gracile; (j) Nemastoma triste; (k) “Pyza” taurica; (l) Paranemastoma superbum, glans penis, ventral; (m) Dendrolasma mirabile; (n) D. mirabile, ovipositor. Scale bars: e–h, k, m–n = 0.5 mm. Sources: a–c, i–j, l, unpublished from J. Martens; d–e, g, i, m, n, from Shear and Gruber (1983); f, h from Gruber (1976); k from Gruber (1979).

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150 Taxonomy

• Chelicerae: Normal proportions, often sexually dimorphic; dentition of chela with narrow, diaphanous teeth (Figures 4.16e,f). • Pedipalps: Length and girth variable, from short and stout ones to strongly elongate and gracile ones (Figures 4.16a,b). Femora with “basal bend.” Tibiae and tarsi with narrowed bases and somewhat inﬂated, less markedly so in elongated, thin pedipalp. Tarsal claw absent, but with “nipplelike” rudiment. With clavate glandular setae especially on distal segments, sometimes reduced in adults (Figures 4.16g,h). • Legs: Generally moderately short to long, segments of various shapes—femora to tibiae may be incrassate or thin, distal segments generally thin. Femora often with pseudoarticulations. Surface of proximal segments with various microsculptural elements; metatarsi without calcanei. Tarsi moderately pluri- articulated. • Genitalia: Penis (Figures 4.16i–m) long, shaft with incrassate bulb-shaped base (its walls often cleaved medially) containing the two intrinsic penis muscles, their tendons therefore long. Shaft varying in shape from stout with extended bulbous base (Figure 4.16i) to slender and straight with relatively shorter basal portion (Figure 4.16k) or extremely slender and elongate with basal bulb sharply set off and bent against main portion of shaft (Figure 4.16j). The shaft may bear lateral wings below the glans in certain groups (Figure 4.16k). Glans variously shaped—symmetrical (Figure 4.16k) or more or less asymmetrical (Figure 4.16l), with mostly short stylus (exceptions especially in Ortholasmatinae, Figure 4.16m) and variable setation. Ovipositor unsegmented, truncus soft skinned, short to very short, with nonglandular setae, apical portion (furca) with two weakly delimited segments, sometimes slightly sclerotized (Figure 4.16n). • Color: Mostly pale brown to black, sometimes patterning with paler or darker parts; a common and striking pattern in several genera consists of silvery white (nacreous) or golden spots on a generally black dorsum (Figure 4.16a). • Sexual dimorphism: Apart from size and proportions, males may show a more prominent dorsal armature with cones or spines, and generally a different shape of genital operculum (longer and narrower, without basal suture). Male chelicerae in most species with glandular organs opening on the basal segment, often associated with dorsal apophyses; toothlike apophyses sometimes also on second segment (Mitostoma, ortholasmatines). Pedipalps of males may bear distal apophyses on patellae; glandular areas on femur, tibia, and especially patella may occur in ortholasmatines.

Key to subfamilies: 1. Ocularium with median process extending anteriorly and forming a hood to- gether with lateral carapace processes (Figure 4.16d)...... Ortholasmatinae 2. Ocularium without median process extending anteriorly or forming a hood (Fig- s– ures 4.16a–c; a median erect spine or a bipartite ocularium may occur)...... o– ...... Nemastomatinae l– 29859_U04.qxd 8/18/06 12:28 PM Page 151

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Distribution: The family shows a disjunct Holarctic distribution: Nemastomatinae are a western Palaearctic group, occurring in nearly all of Europe, in the north to Scandinavia and Iceland, in the east to the Urals and the Caucasus; in North Africa in the northern part of the Atlas region; in southwestern Asia from Anatolia and the Levant to Caucasia and northern Iran, with outliers in the Pamirs, the Tien Shan, and the northwestern Himalayas (J. Martens, pers. comm.). Most species are re- stricted to rather small areas in the mountains of the southern part of the whole area; only a few species extend to the far north and northeast. One western European species has been introduced into eastern North America. Ortholasmatinae is an Am- phipaciﬁc group—western North America (Mexico, California, Oregon, Washington, British Columbia) and eastern Asia (southern Japan and northern Thailand). Relationships: Nemastomatidae represent the largest and most diverse group of troguloids. Interfamilial relationships in this superfamily are still somewhat contro- versial (see discussion under Dicranolasmatidae). Nemastomatidae and Dicranolas- matidae show several (but mainly probably symplesiomorphic) similarities (Ortho- lasmatinae show convergent similarity to the latter); some allegedly synapomorphic characters proving monophyly of Nemastomatidae + Dicranolasmatidae (Martens, 1978b: clavate hairs of pedipalps; Shear & Gruber, 1983: penis muscles with long tendons) are either symplesiomorphies (also in Nipponopsalididae) or erroneous (short tendons in Dicranolasma). A recent phylogenetic analysis (Giribet et al., 2002) supported monophyly of Nemastomatidae as a sister to Dicranolasma and Trogulus. Main references: • Systematics: Gruber & Martens (1968), Gruber (1976, 1979), Martens (1978b), Rambla (1980a, 1983), Shear & Gruber (1983), Starega (1986), Schwendinger & Gruber (1992), Prieto (2004). • Natural history: Immel (1954), Juberthie (1964), Starega (1986), Thaler & Knoﬂach (2001).

Nipponopsalididae Martens, 1976 Jürgen Gruber Etymology: Nipponopsalis, combination of Nippon (Japan) with the ending of Ischy- ropsalis. Characterization: • Size: Body length between 2.3 and 4.1 mm. • Dorsum: Integument generally poorly sclerotized, with limits of sclerites sometimes indistinct. Carapace domed with large low ocularium, eyes rather large, no laminae suprachelicerales (pièces épimériennes), anterior border with median infolding and slight emarginations above chelicerae; ozopores very small, visible on carapace border. Mesopeltidium fused to carapace, metapel- –s tidium free. Opisthosoma with scutum parvum in males (with some tendency to –o –l