The Placement of the Spider Genus Periegops and the Phylogeny of Scytodoidea (Araneae: Araneomorphae)

The placement of the spider genus Periegops and the phylogeny of Scytodoidea (Araneae: Araneomorphae)

FACUNDO M. LABARQUE1 & MARTÍN J. RAMÍREZ1 1Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Av. Ángel Gallardo 470, C1405DJR, Buenos Aires, Argentina. [email protected] / [email protected]

Abstract

The relationships of Scytodoidea, including the families Drymusidae, Periegopidae, Scytodidae and Sicariidae, have been con- tentious for a long time. Here we present a reviewed phylogenetic analysis of scytodoid spiders, emphasizing Periegops, the only genus in the family Periegopidae. In our analysis the Scytodoidea are united by the fusion of the third abdominal entapophyses into a median lobe, the presence of female palpal femoral thorns and associated cheliceral stridulatory ridges, a membranous lobe on the cheliceral promargin, and the loss of minor ampullate gland spigots. A basal split within Scytodoidea defines two monophyletic groups: Sicariidae and a group formed by Scytodidae as the sister group of Periegopidae plus Dry- musidae, all united by having bipectinate prolateral claws on tarsi I–II, one major ampullate spigot accompanied by a nubbin, and the posterior median spinnerets with a mesal field of spicules. Periegops is the sister group of Drymusidae, united by the regain of promarginal cheliceral teeth and a triangular cheliceral lamina, which is continuous with the paturon margin.

Key words: Drymusa, Drymusidae, Haplogyne, morphology, Scytodes, Stedocys, Scytodidae, Sicariidae, Sicarius, Loxosceles

Introduction

The family Periegopidae currently comprises only the genus Periegops, with two species: the type species Perie- gops suteri (Urquhart) from the Banks Peninsula on the South Island of New Zealand (Vink 2006), and Periegops australia Forster, from southeastern Queensland (Forster 1995). A single female of an unidentified Periegops species from the East Cape region of the North Island of New Zealand was also mentioned by Forster (1995); at present, the delimitation of species within Periegops is dependent on the detailed structure of the male copulatory bulb, hence the identification of this specimen should wait for new data, although mitochondrial COI molecular evidence suggests that it may be a distinct species (unpublished data mentioned in Vink 2006). The genus Periegops was first included in the subfamily Periegopinae by Simon (1893), and placed in the Sicariidae, along with Drymusinae, Loxoscelinae, Plectreurinae, Scytodinae and Sicariinae. Simon was the first to discover that Periegops and Scytodes had a bipectinate “external superior” tarsal claw on the anterior tarsi. Simon included Diguetia and Pertica (later synonymized with Segestrioides; Brescovit & Rheims 2005) in Periegopinae, but Petrunkevitch (1928) removed them and placed them into Diguetinae. Bryant (1935a, b) suggested a family rank for Periegops that included also Plectreurys and Diguetia, but she took no formal action. Lehtinen (1986) studied only the female holotype of P. suteri. He mentioned that Periegopidae, Drymusidae, Scytodidae and Ochyroceratidae share a particular dentition pattern in the tarsal claws, but he never explained or described this character in detail, so his observations were not conclusive. He also suggested that Periegops could be a strongly apomorphic derivative of the classical (non-monophyletic, Platnick et al. 1991), Scytodoidea, composed of the Caponiidae, Diguetidae, Loxoscelidae, Ochyroceratidae, Plectreuridae, Pholcidae, Scytodidae, Sicari- idae and Tetrablemmidae (Brignoli 1975, 1978). Platnick et al. (1991) investigated the spinneret morphology of haplogyne araneomorph spiders (Haplogynae), and produced a cladistic analysis with all haplogyne families except Periegopidae. They obtained a restricted Scyt- odoidea, composed of Sicariidae, Scytodidae, and Drymusidae, in which Scytodidae and Drymusidae were united

Accepted by W. Fannes: 29 Dec. 2011; published: 14 May 2012 1 by having a field of spicules on the median surface of the posterior median spinnerets. Ramírez (2000) added a few characters from the respiratory system to the same dataset, further strengthening Platnick et al.’s hypothesis. Gris- wold et al. (2005) reviewed most of the character systems used in those sources, but only scored two families of Haplogynae (Filistatidae and Segestriidae) as outgroups for their analysis. Forster (1995) was the first to examine the tracheal system of Periegops, and their cuticular structures using scanning electron microscopy. He hypothesized that the Periegopidae were related to the ecribellate, haplogyne families Diguetidae, Drymusidae, Plectreuridae, Scytodidae and Sicariidae (that is, Simon’s Sicariidae, plus Diguetidae). Within this group, Forster suggested that Periegopidae was most closely related to the Scytodidae, based on two characters: (a) posterior tracheal system composed of a short median apodemal lobe and a distinct pair of simple lateral tracheae, a configuration also shared with Drymusidae; and (b) a double row of teeth on the ventral surface of the proclaws, and a single row on the retroclaws of leg I and II (later found in Drymusa as well, see below). Lamy (1902) was the first to describe the tracheae of Scytodes and Loxosceles, which both share a fused median apodeme. Ramírez (2000) confirmed that at least in Loxosceles and Sicarius the median lobe retained the muscle insertions corresponding to the paired third abdominal entapophyses, proposing their fusion on a median structure as a further synapomorphy for a clade that included Drymusa, Loxosceles, Scytodes and Sicarius (plus Periegops, although this genus was not included in the analysis). In the cladistic analysis of Ramírez (2000), the elongation of the median apodeme is a synapomorphy for Sicariidae, thus a short median apodeme is plesiomorphic for Scytodes, Drymusa, and Periegops. Labarque & Ramírez (2007a) reported that the bipectinate proclaws are also present in Drymusa, and that Periegops also have a field of spicules on the median surface of the posterior median spinnerets (PMS), along with Drymusa and Scytodes. In summary, the more promising characters that may help resolve the closest relationships of Periegops (the bipectinate proclaws, the spicules on the PMS), are shared both with Dymusa and Scytodes. Here we intend to clar- ify the relationships of Periegops and its relatives performing a cladistic analysis of Scytodoid spiders, reviewing and expanding on the previous analyses (Platnick et al. 1991; Ramírez 2000; Griswold et al. 2005).

Material and methods

Specimen preparation Clean museum specimens were chosen when available. Specimens in 80% alcohol were imaged with a Nikon DXM1200 digital camera mounted on a stereoscopic microscope Nikon SMZ1500 and the focal planes composed with Helicon Focus 3.10.3 and 4.01 Pro (Khmelik et al. 2006). Preparations were carefully cleaned using fine brushes, a thin jet of alcohol from a thinned pipette, or ultrasonic cleaner; some setae were removed to expose structures, especially those on legs and palp, spinnerets and chelicerae. For scanning electron microscope (SEM), all preparations were dehydrated in a series of increasing concentrations of ethanol (80%, 90%, 95%, 100%), and critical-point dried. After drying and brushing, they were mounted on adhesive copper tape (Electron Microscopy Sciences, EMS 77802) affixed to a stub and secured with a conductive paint of colloidal graphite on isopropyl alcohol base (EMS 12660). Prior to SEM examination under high vacuum with a FEI XL30 TMP, the structures were sputter-coated with Au-Pd.

Material examined The following specimens were examined to score the phylogenetic dataset. See Acknowledgements for a list of institutional acronyms and curators. Hypochilus pococki Platnick: 1 ♀, USA, North Carolina, Swain Co., Great Smokey Mountains, Deep Creek Campground, 0.3 Km along Stone Pile Trial, 585 m, N 35°27.848', W 83°26.078', J. Bond, F. Coyle coll., 19.X.2002, MACN-Ar 28173 (ARAMR-000641). 1 ♂, same data, MACN-Ar 29171 (ARAMR-000642). Kukulcania hibernalis (Hentz): 1 ♀, ARGENTINA, Capital Federal, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, M. Ramírez, C. Grismado, A. Ojanguren-Afilastro coll., 6.I.2004, MACN-Ar 10427 (ARAMR-000701). 1 ♂, same locality, M. Ramírez col., 16.I.2003, MACN-Ar 10422 (ARAMR-000075). Ariadna boesenbergi Keyserling: 1 ♀, ARGENTINA, Buenos Aires, Ruta Pergamino, Arrecifes Km 210, S. Rodríguez Gil col., XI.2007, MACN-Ar 28170 (ARAMR-000966). 1 ♂, ARGENTINA, Capital Federal, M. Miranda col., I.1989, MACN-Ar 10177 (ARAMR-000965).

2 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE Pholcus phalangioides (Fuesslin): 1 ♂, 1 ♀ and 3 immatures, ARGENTINA, Misiones, Santa María, De Carlo, Schiapelli, Viana, Galiano, coll., XI.1954, MACN-Ar 4337. Sicarius rupestris (Holmberg): 2 ♀, ARGENTINA, San Juan, Ruta 150, camino a Paso de Agua Negra, 6 km Oeste de Arrequintín, Maury col., 23.I.1982, MACN-Ar 28169 (ARAMR-000881) and MACN-Ar 28167 (ARAMR-000889), respectively. 1 ♂, same data, MACN-Ar 28168 (ARAMR-000882). Loxosceles rufescens (Dufour): 1 ♀, SPAIN, Granada, Cueva GL-10, Calahonda, M. Ramírez col., 8.XII.1983, UB, Col Aràcnids Reg. Nº 2067 Fl. 83 (ARAMR-000618). 1 ♂, same data (ARAMR-000617). Scytodes globula Nicolet: 2 ♂, ARGENTINA, Buenos Aires, Isla Martín García, A.Ojanguren, J.Barneche coll., 4.I.2006, MACN-Ar 28174 (ARAMR-000558) and MACN-Ar 28172 (ARAMR-000813), respectively. 1 ♀, ARGENTINA, Buenos Aires, Villa Madero, in a house, C. Scioscia col., 13.X.2006, MACN-Ar 10789 (ARAMR- 000520). 1 ♀, URUGUAY, Camino a Artigas Ruta 30, A. Ojanguren, C. Mattoni, F. Labarque coll., 13.XII.2005, MACN-Ar 28175 (ARAMR-000944). Stedocys leopoldi (Giltay): 1 ♀, THAILAND, Chiang Mai, Doi Chiang Dao WS, Amphen Chiangdao, Mae Ta Man forest, field station, 1500 m, N 19°19'13.2'', E 98°49'47.0'', M. Ramírez col., 1.X.2003, MACN-Ar 23714 (ARAMR-000637). 1 ♂, THAILAND, Chiang Mai, Doi Inthanon National Park, nr. intersect. rd. to Mae Chaem and checkpoint, wet primary forest, 1800 m, N 18°31'33.2'', E 98°29'57.7'', M. Ramírez col., 3.X.2003, MACN-Ar 23713 (ARAMR-000199). Periegops suteri (Urquhart): 1 ♂, NEW ZEALAND, Banks Peninsula, Hinewai Reserve, near "Big Beech", S 43˚48,59', W 173˚01,28', C.J.Vink col., 2.X.2001, AMNH (ARAMR-000615). 1 ♀, same data, AMNH (ARAMR- 000616). Drymusa capensis Simon: 1 ♀, SOUTH AFRICA, Western Cape, Table Mountain National Park, Newland´s Forest, 145 m, S 33º58,440', W 18º26,648', J. Miller, H. Wood, N. Larsen coll., 25.II.2006, CASENT 9023625 (ARAMR-000919). Drymusa rengan Labarque & Ramírez, 2007a: 1 ♀, CHILE, Maule (RegionVII), Curicó, Los Queñes, S 35°01', W 70°48', M. Ramírez, F. Labarque coll., 16.II.2005, MACN-Ar 11000 (ARAMR-000522). 1 ♂, same data, MACN-Ar 11007. Drymusa serrana Goloboff & Ramírez: 1 ♀, ARGENTINA, Buenos Aires, Sierra de la Ventana, Parque Pro- vincial Ernesto Tornquist, S 38°1,142', W 62°0,97', G. Binford, J. Miller, F. Labarque, C. Ellison coll., 10.I.2005, MACN-Ar 11018.

Terminals and scorings The present study concentrates on Scytodoidea as restricted by Platnick et al. (1991). The data matrix (Table 1) includes Scytodes globula and Stedocys leopoldi (Scytodidae), Drymusa capensis, D. serrana and D. rengan (Drymusidae), Sicarius rupestris, Loxosceles reclusa and L. rufescens (Sicariidae). Four representatives of the main clades of haplogynes and basal araneomorphs were chosen to root the analysis, according to the previous analyses of Platnick et al. (1991), Ramírez (2000) and Griswold et al. (2005); these are Hypochilus pococki (Hypochilidae), Kukulcania hibernalis (Filistatidae), Ariadna boesenbergi (Segestriidae) and Pholcus phalangioides (Pholcidae). All character scorings were critically evaluated and reexamined from voucher specimens (see Material examined); a few scorings in previous studies from predicted rather than observed occurrences (e.g., for internal anatomy) are represented here as missing entries.

Cladistic analysis Analyses were performed with TNT 1.1 (Goloboff et al. 2008a). The complete dataset is available for down- load from http://aracnologia.macn.gov.ar/biblio/suppl/, and the dataset is deposited in TreeBase with accession number 12227. Multistate characters with states showing clear intermediacy (characters 3, 48, 51–53, 70 and 79) were considered ordered (additive). Tree searches. This matrix is small enough to allow for exact searches (TNT command ienum;). We analyzed the dataset under weighting regimes against homoplasy, using implied weighting (Goloboff 1993), and under equal weights. We assessed the sensitivity of the results to variations in the strength of the weighting function, with inte- ger values of the constant of concavity k = 1 to 9. Bremer support. Bremer support values (BS; Bremer 1994) were calculated with an exact search retaining sub- optimal trees under equal weights (string of commands hold 2000; subopt 5; ienum; bsupport;) Resampling measures. We also calculated support values using group frequencies under jackknifing (see Goloboff et al. 2003), with the usual probability of alteration of p = 0.36. The absolute frequencies are reported

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 3 over the optimal preferred tree, thus all values are reported (even below 0.5). We performed 10,000 pseudorepli- cates, each with an exact search keeping up to 1000 trees (string of commands hold 1000; resample jak repl 10000 freq from 0 [ienum]; where Tree 0 is the optimal tree). In addition to jackknifing, we similarly calculated group frequencies under bootstrap (not shown); because both values were highly correlated, we report only the jackknifing values here.

Abbreviations used throughout text and figures

3e third opisthosomal entapophyses AC aciniform spigot ALS anterior lateral spinneret Bm blunt macroseta EF epigastric furrow GD gland ductules IC inferior claw ITrC inter-tracheal canal IS inner spermatheca Lt lateral trachea MAP major ampullate gland spigot mAP minor ampullate gland spigot mS median spicules N nubbin Ony onychium OS outer spermatheca PBc prolateral bipectinated claw PI piriform spigot pL cheliceral promarginal lobe pT cheliceral promarginal tooth pF posepigastric foveae PLS posterior lateral spinneret PMS posterior median spinneret S spermatheca Sr spaced stridulatory ridges Vo venom outlet

CHARACTERS

The characters from the Haplogynae data matrix (Platnick et al. 1991; Ramírez 2000) were organized by body region following the Atlas of Griswold et al. (2005). All taxa were scored for a total of 102 characters. Of these, 56 are from Platnick et al. (1991) plus the complementary 16 from Ramírez (2000), and 15 from Griswold et al. (2005); the remaining 15 are new characters proposed here. Character provenance and equivalences between studies are listed as (Study: Character number), using the abbreviations P (Platnick et al. 1991), R (Ramírez 2000), and G (Griswold et al. 2005). Legacy characters that have been reformulated are marked with an asterisk. We have included several characters used in previous analyses of haplogynes which are not informative for the parsimony analysis. This has been done with the intention of allowing the integration of the present dataset into more integra- tive studies (Ramírez et al. 2007).

LEGS Figs. 2–12 Character 0: Legs, autospasy line configuration: (0) coxa-trochanter; (1) patella–tibia (R: 76). Character 1: Metatarsal trichobothria, number: (0) one or two; (1) three or more (G: 5). Character 2: Trichobothrial base hood, texture: (0) smooth; (1) with transverse ridges (P: 64, G: 8).

4 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE Character 3: Trichobothrial base margin, shape (except for first tibial trichobothria): (0) entire; (1) broadly notched; (2) crenulate (P: 13). States are ordered. Character 4: Tibial Emerit´s glands: (0) absent; (1) present (P: 31). Character 5: Metatarsus and tarsus I–II, conspicuous prolateral scopula of spatulate setae: (0) absent; (1) present (P: 26). Character 6: Tarsus, basal third, cuticle sclerotization: (0) sclerotized; (1) unsclerotized ring (P: 50). Character 7: Tarsus, segmentation: (0) entire; (1) pseudosegmented (P: 41). Character 8: Tarsal organ, conformation: (0) exposed; (1) capsulate (P: 65, G: 2). Tarsal organ conformation in Pholcus phalangioides was scored as capsulated, instead of exposed (contra Platnick et al. 1991). The exposed condition was presumed to be plesiomorphic for the family by Platnick et al.; however the capsullate condition resulted primitive for pholcids (Huber 2000). Character 9: Tarsi, proprioceptor bristles: (0) absent; (1) present (R: 81). Character 10: Tarsi, distal end, segmentation: (0) entire; (1) distal articulation, delimiting an onychium. The onychium is an apical portion of the tarsus, bearing the pretarsus and the claws, and articulated through a soft membrane. An onychium is present in Periegops, Drymusa (Figs. 3, 6 A–D; Labarque & Ramírez 2007a, b), Scytodidae (Figs. 4, 7 A–D), and Loxosceles (Figs. 5, 8 A–B). Character 11: Inferior tarsal claw: (0) present; (1) absent (P: 36). Character 12: Proclaws I–II, dentition: (0) single; (1) double (R: 80). Members of Scytodoidea (Figs. 3, 4) have the ventral surfaces of the proclaws I and II clearly bipectinate, with longer teeth on the prolateral row (state 1; pg. 266, Simon 1893; Forster 1995; Labarque & Ramírez 2007a, b). Character 13: Paired tarsal claws I–II, length: (0) similar in size; (1) proclaw enlarged (P: 11). Character 14: Tarsal cuticle texture: (0) fingerprint; (1) smooth; (2) discrete cells (G: 10). Character 15: Female palp, femoral thorns: (0) absent; (1) present (G: 16). Character 16: Female palpal tarsi, prolateral-apical-dorsal setae tip, shape: (0) acute; (1) clavate. Scytodes (Fig. 12 C; Simon 1893, fig. 235) and Stedocys (Fig. 12 D; Labarque et al. 2009, fig. 27) have clavated setae (state 1). These macrosetae have a pore-bearing, widened, blunt tip. The presence of a pore at the tip is suggestive of a chemosensory function (Barth 2001). This character is also present on male palpal cymbium of the genus Scy- todes (Rheims & Brescovit 2000, 2001, 2004; Brescovit & Rheims 2000, 2001; Rheims et al. 2005, 2007) and Dictis (Ramírez pers. obs.; see also Paik 1978: fig. 93 4; Schenkel 1963: fig. 10), while it is absent in Stedocys males (Giltay 1935: fig. 3; Lehtinen 1986: fig. 7; Ono 1995: fig. 4; Labarque et al. 2009: fig. 10–11, 29). Character 17: Female palpal claw: (0) present; (1) absent (P: 37). Character 18: Calamistrum, setae rows, number: (0) two; (1) one; (3) three (G: 26).

PROSOMA Figs. 13–18 Character 19: Thoracic region, height relative to height at posterior lateral eyes: (0) similarly elevated (Fig. 13 A, B, E, F); (1) at least twice as high, forming a dome (Fig. 13 C, D). Scytodidae have the carapace posteriorly domed, with the thoracic region much higher than the cephalic area (state 1). Character 20: Clypeal hood: (0) absent; (1) present (P: 17 and G: 30). Character 21: Cephalic region, anterior margins, configuration: (0) separated; (1) prolonged between clypeus and endites, meeting in midline (cheliceral foramen) (P: 22, G: 32). Character 22: Cheliceral paturons, interaction: (0) free; (1) fused (P: 34, G: 38). Character 23: Paturon, mesal surface next to furrows margin ends: (0) concave (median cheliceral concavity); (1) straight or convex (P: 4, G: 37). Character 24: Chelicerae, promarginal teeth: (0) present; (1) absent (G: 39). Character 25: Chelicerae, peg teeth: (0) absent; (1) present (P: 19, G: 41). Character 26: Chelicerae, medial lamina: (0) absent; (1) present (P: 2, G: 40). Character 27: Chelicerae, medial lamina, distal end shape: (0) triangular and continuous; (1) acute and separated by deep depression. A triangular or subtriangular lamina continuous with the paturon occurs in Periegops, Drymusa (Figs. 15, 18 A, B) and Pholcus. An acute lamina occurs in Scytodidae (Figs. 15, 18 C, D), Sicariidae (Figs. 15, 18 E, F) and Kukulcania. This character is only applicable for taxa with cheliceral lamina. Character 28: Chelicerae, promarginal lobe: (0) absent; (1) present. A promarginal membranous lobe is present in

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 5 Scytodoidea (Figs. 15, 18 A–F) and in Kukulcania (Griswold et al. 2005: fig. 126 B, behind the lamina). Character 29: Chelicerae, promarginal lobe setae, shape: (0) filiform, long; (1) conical, small. Long and filiform setae lean against the promarginal lobe of Periegops (Fig. 15 A), Drymusa capensis (Fig. 15 B), and Scytodi- dae (Fig. 15 C, D). Smooth conical setae appear in Drymusa rengan (Labarque & Ramírez 2007a, fig. 25), Drymusa serrana (Labarque, pers. obs.) and Sicariidae (Fig. 15 E, F). This character is only applicable for taxa with a cheliceral promarginal lobe. Character 30: Chelicerae, promarginal lobe setae, distribution: (0) dispersed; (1) in a basal row. In Scytodidae the promarginal lobe setae are dispersed (Fig. 15 C, D). In all the other members of Scytodoidea the setae are arranged in a row basal to the lobe (Fig. 15 A, B, E, F; Labarque & Ramírez 2007a, fig. 25). This character is only applicable for taxa with a cheliceral promarginal lobe. Character 31: Cheliceral gland, elevation: (0) not elevated; (1) on a mound (P: 20, G: 42). Character 32: Female chelicerae, stridulatory ridges: (0) absent; (1) present (G: 45). Character 33: Female chelicerae, stridulatory ridges, configuration: (0) a few large and well-spaced ridges; (1) long series of minute ridges; (2) shallow multiple scales. In Scytodidae females (Fig. 16 C, D) the stridulatory ridges are conspicuous and well-spaced (state 0; Rheims & Brescovit 2000, 2001, 2004; Rheims et al. 2005, 2007). Sicarius females (Fig. 16 F) have a long series of minute ridges (state 1). Females of Periegops, Dry- musa and Loxosceles (Fig. 16 A, B, E) have shallow multiple ridges or scales (state 2) (e.g., Fischer et al. 2009). This character is only applicable for taxa with stridulatory striae on the chelicerae. Character 34: Cheliceral fang, venom outlet, position: (0) distal; (1) basal. Spider poison is injected through a tiny opening at the tip of the cheliceral fang (state 0; Foelix 1996). Scytodes has specialized venom glands, with an anterior part that produces the venom and a posterior part that produces a glue-like substance (Kovoor & Zyl- berberg 1972). The secretory epithelia of these glands extend into the chelicerae, and the orifice through which the spit is ejected is located near the base of the fang (state 1; Fig. 18 C; Suter & Stratton 2009: fig. 7). The outlet forms an open channel from the base of the fang towards its distal end (Fig. 18 C, D). Stedocys has a similar conformation (Fig. 18 D; Labarque et al. 2009, fig. 17, 18). Character 35: Cheliceral fang, venom outlet orientation: (0) posterior; (1) anterior. In Periegops and Drymusa (Fig. 15 A, B) the venom gland outlet opens towards the cheliceral promargin (state 1). In Sicariidae (Fig. 18 E, F) and in the outgroup taxa examined here the orifice opens towards the retromargin. Scytodidae is coded as not applicable (see character 35). Character 36: Anterior median eyes: (0) present; (1) absent (P: 0). Character 37: Labrum, lateral posterior protuberances: (0) absent; (1) present (P: 49). Character 38: Serrula: (0) absent; (1) present. The serrula is absent in Sicarius (Simon 1893: 269; Lehtinen 1986: fig. 3). Character 39: Serrula teeth rows: (0) multiple; (1) single (G: 49). Sicarius is coded as inapplicable (see character 39). Character 40: Sternum, sigilla: (0) present; (1) absent (P: 8, G: 50). Character 41: Labium and sternum junction configuration: (0) free or separated by partially membranous suture; (1) fused (P: 38*). After observing many intermediate cases of partially fused labium, we have redefined this character to limit state 1 to the totally fused condition, connected to the sternum by hard cuticle and lacking any movement. Character 42: Venom gland, position: (0) endocheliceral; (1) extending into carapace (P: 7, G: 52). Character 43: Coxal glands duct, shape: (0) convoluted; (1) simple (P: 9, G: 53). Character 44: Midgut, diverticula on chelicerae: (0) present; (1) absent (P: 4, G: 54). Character 45: Dorsal dilator muscle M1 of pharynx, insertion: (0) on carapace; (1) on rostrum (P: 6, G: 55).

OPHISTOSOMA Figs. 19, 20 Character 46: Fifth median opisthosomal endosternite: (0) present; (1) absent (P: 10, G: 56). Character 47: Intestine, shape: (0) M-shaped; (1) straight or only curved (P: 14, G: 57). Character 48: Heart ostia, pair number: (0) four (1st to 4th); (1) three (1st to 3rd); (2) two (1st to 2nd). (P: 15*, G: 58*). Hypochilus has four, Kukulcania, Pholcus and Loxcosceles reclusa three, and Ariadna two. All the remaining scytodoids are unknown for this character, although are expected to have three. States are ordered.

6 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE Character 49: Third dorso-ventral opisthosomal muscles: (0) present; (1) absent (R: 75, G: 59). Character 50: Anterior respiratory system, conformation: (0) book-lungs; (1) tracheae (R: 74). Character 51: Posterior respiratory system, lateral structures conformation: (0) book-lungs; (1) lamellae; (2) lateral tracheae (R: 16, G: 61). States are ordered. Character 52: Posterior respiratory system, opening or 3rd opisthosomal entapophyses position, relative to epigastric furrow and spinnerets: (0) about midway between both; (1) just behind epigastric furrow; (2) just anterior to spinnerets (R: 18). States are ordered. Character 53: Posterior respiratory system, spiracles or 3rd opisthosomal entapophyses, configuration: (0) separate; (1) contiguous; (2) fused (R: 32, G: 60). States are ordered. Character 54: Posterior respiratory system, 3rd opisthosomal entapophyses, shape: (0) short, flat or absent; (1) elongated (R: 67). Character 55: Posterior respiratory system, fused 3rd opisthosomal entapophyses, shape: (0) short, slender; (1) elongated, broad (R: 68). Character 56: Posterior respiratory system, 3rd opisthosomal entapophyses, conformation: (0) short muscle apodemes; (1) long median tracheae (R: 69, G: 63). Character 57: Posterior respiratory system, transverse furrow between 3rd opisthosomal entapophyses: (0) present; (1) absent. (R: 70). Character 58: Posterior respiratory system, inter-tracheal canal: (0) absent; (1) present (R: 71). Character 59: Posterior respiratory system, lateral structures: (0) present (as booklungs or lateral tracheae); (1) absent (lost). Character 60: Posterior respiratory system, lateral tracheae, shape: (0) simple; (1) branched (R: 72*, G: 62*). Character 61: Anal tubercle, supra-anal organ: (0) absent; (1) present. (R: 79). Character 62: Male epiandrous spigots: (0) present; (1) absent (G: 66).

SPINNERETS Figs. 21–28 Character 63: Tartipores: (0) absent; (1) present (P: 63). Character 64: Cribellum: (0) present; (1) absent (P: 1, G: 71). Character 65: Cribellum, configuration: (0) entire; (1) divided (P: 45, G: 72). Character 66: Cribellum spigots, shape: (0) strobilate; (1) claviform (P: 30 and G: 74). Character 67: ALS articles, number: (0) three; (1) two, intermediate article missing (P: 40, G: 75). Character 68: ALS distal article, anterior hook: (0) absent; (1) present (P: 24). Character 69: ALS basal article, anteromedian setae: (0) dispersed; (1) in a single row (P: 66). Character 70: Female ALS, MAP gland spigots, number: (0) more than three; (1) three; (2) two; (3) one plus nubbin; (4) one (P: 33, G: 78). States are ordered. Sicarius rupestris was coded as missing, because the MAP spigots are either undifferentiated from PI spigots, or missing (Fig. 26 C, D). Character 71: ALS, MAP gland spigots, arrangement: (0) clustered; (1) dispersed (P: 53, G: 76). Character 72: ALS, piriform gland spigots shaft, shape: (0) narrow; (1) broadened (P: 59). Character 73: ALS, MAP gland spigots, aperture: (0) constricted; (1) elongated. Loxosceles has a special type of MAP, with elongated aperture and flexible base (Figs. 27 D, 28 B, F; Platnick et al. 1991, figs. 93, 98) which produces a ribbon-like silk strand (Coddington et al. 2002). The MAP cuticle presents transverse basal ridges, seemingly to allow bending, and longitudinal apical ridges. Character 74: PMS, shape: (0) rounded or digitiform: (1) tetrahedral. (P: 61). Character 75: PMS gland spigots bases, interaction: (0) free; (1) fused, forming a ring (P: 60). This character is only applicable for taxa with two or more spigots on the PMS; Loxosceles is coded as inapplicable, because it lacks spigots on the PMS (Figs. 26 E, 27 C, D). Character 76: PMS base, setae on anteromedian surface, shape: (0) straight, plumose; (1) bent, thick, with elevated rims (P: 25). Character 77: PMS, mesal surface, spicules: (0) absent; (1) present (P: 58). Character 78: PMS, mAP gland spigots: (0) absent; (1) present (P: 56*, G: 81). Character 79: Female, PMS, mAP gland spigots, number: (0) more than two; (1) two; (2) one plus nubbin; (3) one (P: 56, G: 82). States are ordered.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 7 Character 80: Cylindrical gland spigots: (0) absent; (1) present (P: 23, G: 86). Character 81: Paracribellar spigots: (0) present; (1) absent (P: 44, G: 88). Character 82: PLS, setal pit: (0) absent; (1) present (P: 57). Character 83: Female PLS, aciniform gland spigots: (0) present; (1) absent (P: 62*). Character 84: Female PLS, aciniform gland spigots, number: (0) more than one; (1) one (P: 62*). One AC spigot occurs in Scytodidae (Figs. 23 C, F, 24 D, F) and Drymusa rengan (Labarque & Ramírez 2007a: figs. 42, 43). Character 85: PLS, aciniform gland spigots, arrangement: (0) dispersed; (1) in a single, closely packed line (P: 43). Character 86: Female PLS, modified gland spigots: (0) absent; (1) present (P: 54, G: 96). Character 87: Female PLS, aggregate gland spigots: (0) absent; (1) present (P: 27, G: 95).

MALE GENITALIA Character 88: Tibia, retrolateral process (RTA): (0) absent; (1) present (P: 48, G: 105). Character 89: Cymbium (tarsi), paracymbium: (0) absent; (1) present (P: 47, G: 112). Character 90: Bulb, tegulum and subtegulum, interaction: (0) separate; (1) fused (P: 35). Character 91: Bulb, conductor: (0) present; (1) absent (P: 39, G: 118). Character 92: Bulb, median apophysis: (0) present; (1) absent (G: 123).

FEMALE GENITALIA Figs. 29–31 Character 93: Fertilization duct: (0) absent; (1) present. See Figs. 29, 30 (P: 46, G: 130). Character 94: Gonopore, position relative to epigastric fold: (0) gonopore on epigastric fold; (1) gonopore advanced from epigastric fold (G: 135). Character 95: Posepigastric foveae: (0) absent; (1) present (Fig. 31 C). Foveae, also called “copulatory pockets” (Saaristo 1997) or “scutulae” (Brignoli 1976), are a pair of depressed sclerotized areas located laterally and posterior to the epigastric furrow, present in females of some Scytodidae. In most Scytodes species, the foveae have definite “mesal borders” or position ridges (Gertsch 1958b, 1967; Valerio 1981). Scytodes (Fig. 31 C; Valerio 1981; Rheims & Brescovit 2000, 2001, 2004; Rheims et al. 2005, 2007) and Dictis (Paik 1978; Sarristo 1997) females have foveae with well defined position ridges (except for Scytodes opoxtli Rheims et al., 2007, which lacks foveae). Soeuria Saaristo have only squamous areas (Sarristo 1997), and Stedocys lacks both structures (Fig. 31 D; Lehtinen 1986; Labarque et al. 2009: fig. 9).

SILK Character 96: Cribellate band, axial lines: (0) present; (1) absent (G: 136). Character 97: Cribellate band, reserve warp: (0) present; (1) absent (G: 137). Character 98: Cribellate band, profile: (0) uniform; (1) puffed (G: 139). Character 99: Cribellate fibrils, nodules: (0) absent; (1) present (G: 138).

BEHAVIOR Character 100: Posture on web: (0) inverted; (1) erect (G: 140). Character 101: Cribellate combing movements, leg support: (0) fixed leg III; (1) mobile, braced leg IV (G: 141).

Results and Discussion

The analysis under equal weights resulted in a single tree (Fig. 32) of 97 steps, which was also found under implied weighting with constant of concavity k from 4 to 9. Stronger weighting against homoplasy (k from 1 to 3) resulted in an alterative tree with Scytodidae nested whithin Drymusidae; such a rearrangement involves groups with low support values (Bremer support = 1, jackknifing < 50%). Our analysis corroborates the placement of Periegopidae in Scytodoidea, and more specifically, in a derived clade with bipectinated proclaws on tarsi I–II, together with Scytodidae and Drymusidae (Clade A in Fig. 32). Scytodoidea is here well supported by one non-homoplastic character, the fused 3rd ophistosomical apodemes (ch. 53; Fig. 19; Lamy 1902; Forster 1995; Ramírez 2000), and four homoplastic ones. The presence of female che-

8 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE liceral stridulatory ridges (ch. 32; Fig. 16) and palpal femoral thorns (ch. 15; Fig. 11), although not homoplastic in this dataset, also occur in several entelegynes (Griswold et al. 2005), as well as in other haplogynes such as Diguetidae (Gertsch 1958a: 2; Brescovit & Rheims 2005: fig. 7), Plectreuridae (Gertsch 1958c: 6, 32) and Lep- tonetidae (Ledford & Griswold 2010). The presence of a cheliceral promarginal lobe (ch. 28; Fig. 15) is convergent with Filistatidae (Griswold et al. 2005: fig. 126 B, C). Finally, the absence of a minor ampullate gland spigots on the PMS (ch. 78; Figs. 21 D, F, 22–23 E, 24 C, F, 25 D, F, 26 E, F, 27 E, 28 C, D; Platnick et al. 1991) is shared with Hypochilus pococki (Griswold et al. 2005: fig. 1 C), although in scytodoids it is certainly a loss, while in Hypochilus it might be a primitive absence. Sicariidae, the sister group of all the remaining scytodoids, is well supported by the loss of the inferior tarsal claw in all tarsi (ch. 11; Figs. 5, 8; Platnick et al. 1991), the fused 3rd abdominal entapophyses forming a broad, elongated structure (ch. 56; Fig. 19 E, F; Ramírez 2000) and a row of conical setae against the cheliceral lobe (ch. 29; Fig. 15 E, F). Homoplasy of the last character is due the convergence in some Drymusa species (Labarque & Ramírez 2007a: fig. 25). Within Sicariidae, Sicarius is supported by the loss of several structures such as the onychium (ch. 10; Figs. 5 C, 8 C), the serrula (ch. 38), and the posterior respiratory system (ch. 59; Fig. 19 F; Ramírez 2000), and also by the presence of long series of minute stridulatory ridges on chelicerae (ch. 33; Fig. 16 F; Ger- schman & Schiapelli 1979, fig. 3) and the rounded PMS (ch. 74; Fig. 26 E, F). In turn, Loxosceles is well supported by a MAP spigot with elongated aperture (ch. 73; Figs. 27 D, 28 B, F) and the presence of a setal pit on the PLS (ch. 82; Figs. 27 F, 28 E; Platnick et al. 1991), both non-homoplastic. Drymusa, Periegops and Scytodidae are grouped together (Clade A), well supported by three non homoplastic characters: the bipectinated proclaws on tarsi I–II (ch. 12; Figs. 3, 4; Simon 1893; Forster 1995: figs. 11, 12; Labarque & Ramírez 2007a, b: figs. 28, 29), the regain of a nubbin next to the single MAP on the ALS (ch. 70; Figs. 21 C, 22 D, 24 B, E), and the presence of a median field of spicules on the PMS (ch .77; Figs. 21 D, F, 22–23 E, 24 C, 25 D, F; Platnick et al. 1991: figs. 74, 76, 84, 87; Labarque & Ramírez 2007a, b: fig. 40, 41; Labarque et al. 2009: figs. 34, 36). Simon (1893) already mentioned the bipectinated condition of one of the paired claws of the anterior legs of Scytodes and Periegops. As we report here (Figs. 3, 4), the double row of teeth are on the ventral face of the proclaw (not the “external” claw as mentioned by Simon). Forster (1995) noted that the single row of teeth on the retroclaw of Periegops is placed on the mesal side, similar to the inner row on the proclaw (Fig. 3 A, B), thus suggesting that Periegops is derived from an ancestor with bipectinate claws. A similar arrangement occurs in Drymusa species (Fig. 3 D; Labarque & Ramírez 2007a: fig. 29), however, the arrangement in Scytodidae is different. In Scytodes (Fig. 4 B) and Stedocys (Fig. 4 C) the single row of teeth on the retroclaw is sinuous, ectal at the base and medial to mesal at the tip. This, together with the single row found in outgroups suggests that the ancestral condition for the retroclaw of scytodoids is unipectinate. Forster (1995) also suggested that Drymusa, Periegops and Scytodidae (Clade A) were united by the reduction in number of aciniform gland spigots on the female PLS to only one. In order to test Forster’s hypothesis, we scored this condition as character 84. According to our results, the reduction of female AC gland spigots in PLS occurred in Scytodidae (Figs. 24 C, F, 25 D, F; Plat- nick et al. 1991: fig. 74; Labarque et al. 2009: figs. 34, 36). Females of other scytodoid families (P. suteri, Fig. 21 E; D. capensis, Fig. 23 F; Drymusa sp. from South Africa, Platnick et al. 1991: fig. 85, 88; and D. serrana, Labarque & Ramírez 2007b) have many AC gland spigots on the PLS. However, within Drymusa and Periegops this character is homoplasious, because an immature female of P. australia (Forster 1995, fig. 34) and females of D. rengan (Labarque & Ramírez 2007a: fig. 42, 43) have only one AC gland spigot on the PLS. The family Scytodidae arises as the sister group of Periegops plus Drymusa and is well supported by four non homoplastic characters: an apical group of blunt macrosetae on the female palpal tarsus (ch. 16; Fig. 12 C, D; Simon 1893, fig. 253; Labarque et al. 2009: fig. 27), the thoracic region of the prosoma forming a dome (ch. 19; Fig. 13 C, D), a cheliceral stridulatory file composed by few, large, well spaced ridges (ch. 33; Fig. 16 C, D; Labarque et al. 2009: fig. 19), and the venom outlet placed at the base of the fang (ch. 34; Figs. 15 C, D, 18 C, D; Labarque et al. 2009: figs. 16, 17, 42, 43). The clade is also supported by two homoplasious characters, the fingerprint tarsal cuticular texture (ch. 14; right area of Fig. 9 C) and the reduction to one AC gland spigot on female PLS (ch. 84; Figs. 24 C, F, 25 D, F). On close examination, the apical palpal macrosetae of scytodids have an apical pore (Fig. 12 C), suggestive of a chemosensory function (Barth 2001), which may be additional to a tactile function as suggested by Simon (1893). These apical macrosetae also occur on the male palpal cymbium in species of Scytodes (Rheims & Brescovit 2000, 2001, 2004; Brescovit & Rheims 2000, 2001; Rheims et al. 2005, 2007) and Dictis (see drawings on Paik 1978: fig. 93 4; Schenkel 1963: fig. 10; also on females, Ramírez pers. obs.). However, Stedocys

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 9 males lack these macrosetae (Giltay 1935: fig. 3; Lehtinen 1986: fig. 7; Ono 1995: fig. 4; Labarque et al. 2009: figs. 10, 11, 29). We were not able to examine Soeuria and the literature is not conclusive on this subject (Saaristo 1997). Scytodes species are well known for their ability to spit a gluey substance produced by modified venom glands. The glue-like substance is ejected from the venom outlet, located near the base of the fang (Fig. 15 C, 18 C; Kovoor & Zylberberg 1972; Foelix 1996; Labarque et al. 2009, fig. 43). Stedocys (Fig. 15 D, 18 D; Labarque et al. 2009, figs. 16, 17, 42) have the venom outlet and the fang similarly modified as in Scytodes, which suggests a similar spitting mechanism. The affiliation of Periegops suteri with Drymusa, as well as the internal relationships among Drymusa species are weakly supported in this analysis, as evidenced by the low support values and the few, homoplasious synapomorphies on the branches involved. Periegops and Drymusa are here joined by the triangular cheliceral lamina, continuous with paturon margin (ch. 27; Figs. 17–18 A–B), and by the regain of cheliceral promarginal teeth (ch. 24; Fig. 15 A, B). This last result differs from Forster’s (1995) idea, where all families closely related to Periegops (his Sicarioidea) have lost all their cheliceral teeth. Drymusa (Figs. 15, 18 B; Valerio 1971, fig. 10; Alayón Garcia 1981, 1987; Brescovit et al. 2004; Bonaldo et al. 2006; Labarque & Ramírez 2007a, b, figs. 24–27; Rheims et al. 2008), Diguetidae (Gertsch 1958a: 9) and Plectreuridae (Gertsch 1958c: 6, 32), all considered members of Sicari- oidea by Forster, have promarginal cheliceral teeth. Furthermore, Periegops and Drymusa have the venom outlet facing anteriorly (ch. 35; Fig. 15 B), rather than posteriorly. This character did not arise as a synapomorphy of these taxa because of the inapplicable scoring in Scytodidae, where the venom outlet is modified for the spitting function. We were unable to find autapomorphies for Periegops. Forster interpreted that Periegops chelicerae were free, with seven or eigth small denticles on the promargin, and three teeth on retromargin (1995: figs. 8–10). We found that in Periegops the cheliceral bases are fused with a stiff articulation (Figs. 14, 17 A) and have two teeth on both promargin and retromargin (Figs. 15, 18 A), similarly as occurs in Drymusa (Figs. 14, 15, 17–18 B). Periegops has six broad setae against the cheliceral promarginal lobe, similarly as in Drymusa capensis (Figs. 15, 18 A, B). These modified setae in Periegops were considered homologous with the row of small setae in D. rengan (see character 30; Labarque & Ramírez 2007a, fig. 25), D. serrana (Labarque, pers. obs.) and Sicariidae (Fig. 15 E, F). Lehtinen (1986) suggested that the spigots in the male gonopore region (epiandrious spigots) are totally reduced in all haplogyne spiders with unpaired vulvae. We found however that P. suteri males have epiandrous spigots (char. 62; Fig. 20 A) and females have unpaired vulvae (Fig. 29 A, B, Forster 1995: fig. 23). Both families Drymusidae and Periegopidae contain a single genus each, and given their apparently close affiliation one wonders if they should be better synonymized. However, the genus Drymusa, widely distributed in America and South Africa, is much more diverse than currently known (Abel Perez Gonzalez, pers. comm.), and hence we believe that the taxonomic rank of both groups will be better evaluated when such diversity is properly studied. Given the close relationship and morphological similarity of Periegops and Drymusa, a reassessment of the classification of Drymusidae should consider the placement of Periegops as well.

Acknowledgements

We wish to thank Charles Griswold, Lara Lopardo and Norman Platnick for comments on early versions of this manuscript. We also wish to thank Matías Izquierdo, for providing SEM images to elucidate character scorings in the outgroups. This work was supported by grants from ANPCyT (1393), CONICET (PIP 3209) and NSF (No. EAR-0228699), and by a doctoral grant from CONICET to Facundo Labarque. Cor Vink collected and made available the specimens of Periegops used for this study. Nikolaj Scharff, Chaweewan Hutacharern and the Department of National Parks, Wildlife and Plant Conservation of Thailand provided support for the ATOL expedition to Thai- land in 2003. Permits for collection of specimens were issued by Corporación Nacional Forestal (Chile), Departa- mento de Áreas Protegidas, Dirección Administración de Áreas Protegidas y Conservación de la Biodiversidad de la Provincia de Buenos Aires, and Administración de Parques Nacionales (Argentina). The following institutions and curators provided the specimens used for this study: American Museum of Natural History, New York (AMNH, Norman Platnick), California Academy of Sciences, San Francisco (CASENT, Charles Griswold), Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires (MACN, Cristina Scioscia), National Museum of Natural History, Universitat de Barcelona (UB, Carles Ribera). We also wish to thank Wouter Fannes (Editor), Joel Ledford and one anonymous reviewer for their useful suggestions and corrections.

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12 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 1. Periegops suteri, preserved specimens. A–C. Female. D–F. Male. A, D. Dorsal view. B, E. Lateral. C, F. Ventral. (B, image mirrored). Scales: A–F 2 cm.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 13 FIGURE 2. Female tarsus I. A. Periegops suteri, right tarsus. B. Drymusa capensis, left. C. Scytodes globula, left. D. Stedocys leopoldi, right. E. Loxosceles rufescens, left. F. Sicarius rupestris, left. (B–D, images mirrored). Scales: A, C, E 200 µm, B, D, F 500 µm.

14 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 3. Female right tarsal claws I. A–B. Periegops suteri. C–D. Drymusa capensis. A, C. Retrolateral-apical view. B, D. Apical view. Scales: A–B 50 µm, C–D 100 µm. Abbreviations: IC, inferior claw (character 11); Ony, onychium (character 10); PBc, prolateral bipectinated claw (character 12, state 1).

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 15 FIGURE 4. Female tarsal claws I. A–B. Scytodes globula, left tarsus. C–D. Stedocys leopoldi, right tarsus. A, C. Retrolateral-apical view. B. Apical view. D. Retrolateral view. Scales: A–D 50 µm. Abbreviations: IC, inferior claw (character 11); Ony, onychium (character 10); PBc, prolateral bipectinated claw (character 12, state 1).

16 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 5. Female left tarsal claws I. A–B. Loxosceles rufescens. C–D. Sicarius rupestris. A. Retrolateral-apical view. B, D. Apical view. C. Retrolateral view. Scales: A 50 µm, B–D 100 µm. Abbreviations: Ony, onychium (character 10).

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 17 FIGURE 6. Female right tarsal claws IV. A–B. Periegops suteri. C–D. Drymusa capensis. A, C. Retrolateral-apical view. B, D. Apical view. Scales: A–D 50 µm. Abbreviations: IC, inferior claw (character 11); Ony, onychium (character 10).

18 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 7. Female tarsal claws IV. A–B. Scytodes globula, left tarsus (retrolateral claw lost). C–D. Stedocys leopoldi, right tarsus. A, C. Retrolateral-apical view. B, D. Apical view. Scales: A–D 50 µm. Abbreviations: IC, inferior claw (character 11); Ony, onychium (character 10).

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 19 FIGURE 8. Female left tarsal claws IV. A–B. Loxosceles rufescens. C–D. Sicarius rupestris. A. Retrolateral-apical view. B, D. Apical view. C. Retrolateral view. Scales: A 50 µm, B 100 µm, C–D 200 µm. Abbreviations: Ony, onychium (character 10).

20 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 9. Female leg trichobothria. A. Periegops suteri, tibia I. B. Drymusa capensis, metatarsus I. C. Scytodes globula, metatarsus IV. D. Stedocys leopoldi, metatarsus I. E. Loxosceles rufescens, metatarsus I. F. Sicarius rupestris, metatarsus IV. Scales: A 10 µm, B 50 µm, C–F 20 µm.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 21 FIGURE 10. Female leg tarsal organ. A. Periegops suteri, tarsus I. B. Drymusa capensis, tarsus IV. C. Scytodes globula, tarsus I. D. Stedocys leopoldi, tarsus IV. E. Loxosceles rufescens, tarsus I. F. Sicarius rupestris, tarsus IV. Scales: A, C–D 20 µm, B, E–F 10 µm.

22 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 11. Female left palpal femoral thorns. A. Periegops suteri, arrow to thorns. B. Drymusa capensis. C. Scytodes globula. D. Stedocys leopoldi. E. Loxosceles rufescens. F. Sicarius rupestris. Scales: A 200 µm, B 50 µm, C–D, F 100 µm, E 20 µm.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 23 FIGURE 12. Female left palpal tarsus, apical view. A. Periegops suteri. B. Drymusa capensis. C. Scytodes globula, inset showing pore on blunt macroseta. D. Stedocys leopoldi. E. Loxosceles rufescens. F. Sicarius rupestris. Scales: A, C–E 50 µm, B, F 100 µm. Abbreviations: Bm, blunt macroseta (character 16).

24 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 13. Female carapace, lateral view. A. Periegops suteri. B. Drymusa capensis. C. Scytodes globula, right side, mirrored. D. Stedocys leopoldi. E. Loxosceles rufescens. F. Sicarius rupestres. Scales: A–F 1 mm.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 25 FIGURE 14. Chelicerae, anterior view. A. Periegops suteri. B. Drymusa capensis. C. Scytodes globula. D. Stedocys leopoldi. E. Loxosceles rufescens. F. Sicarius rupestris, mirrored. Scales: A–B, F 500 µm, C–E 200 µm.

26 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 15. Cheliceral promargin, anterior view. A. Periegops suteri. B. Drymusa capensis. C. Scytodes globula. D. Stedocys leopoldi. E. Loxosceles rufescens. F. Sicarius rupestris, mirrored. A–B, E–F. Inset showing row of modified setae against promarginal lobe base (character 30). Scales: A–B, F 500 µm, C 50 µm, D–E 100 µm. Abbreviations: pL, promarginal lobe (character 28); pT, cheliceral promarginal tooth (character 24); Vo, venom outlet (character 35).

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 27 FIGURE 16. Chelicerae, ectal view, inset to stridulatory field. A. Periegops suteri. B. Drymusa capensis. C. Scytodes globula. D. Ste- docys leopoldi. E. Loxosceles rufescens. F. Sicarius rupestris. D, F, Mirrored. Scales: A–B, F 500 µm, C–E 200 µm. Abbreviations: Sr, spaced stridulatory ridges (character 33, state 1).

28 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 17. Chelicerae, posterior view. A. Periegops suteri. B. Drymusa capensis. C. Scytodes globula. D. Stedocys leopoldi. E. Loxosceles rufescens. F. Sicarius rupestris. D, F, mirrored. C–F. Inset showing deep incision between cheliceral lamina and paturon (character 27, state 1). Scales: A–B, F 500 µm, C–E 200 µm.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 29 FIGURE 18. Cheliceral retromargin, posterior view. A. Periegops suteri. B. Drymusa capensis. C. Scytodes globula. D. Stedocys leopoldi. E. Loxosceles rufescens. F. Sicarius rupestris, mirrored. Scales: A–B, F 500 µm, C, E 100 µm, D 50 µm. Abbreviation: Vo, venom outlet (character 35).

30 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 19. Female tracheae, dorsal view. A. Drymusa capensis. B. Drymusa rengan. C. Drymusa serrana. D. Scytodes globula. E. Loxosceles laeta. F. Sicarius rupestris. Scales: F, 500 µm, A–E 200 µm. Abbreviation: 3e, third opisthosomal entapophyses; ITrC, inter-tracheal canal; Lt, lateral trachea.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 31 FIGURE 20. Male epiandrium. A. Periegops suteri. B. Scytodes globula. C. Stedocys leopoldi. D–E. Loxosceles rufescens. F. Sicarius rupestris. A–D. Ventral view. E. Anterior. F. Lateral, right. Scales: A–C, F 100 µm, D–E 50 µm.

32 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 21. Periegops suteri, spinnerets. A–E. Female. F. Male. A. Spinnerets, posterior-ventral view. B. Same, lateral view (right), numbers on ALS articles (character 67). C. Right ALS. D. PMS, posterior view. E. Right PLS. F. PMS, ventral view. Scales: A–B 100 µm, C–D, F 20 µm, E 50 µm. Abbreviations: AC, aciniform gland spigot; MAP, major ampullate gland spigot; mS, PMS median spicules (character 77); N, nubbin.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 33 FIGURE 22. Periegops suteri, male spinnerets. A. Spinnerets, posterior-ventral view, numbers on ALS articles (character 67). B. Same, lateral view (left). C. Right ALS. D. ALS, MAP area. E. PMS, anterior view. F. Left PLS. Scales: A–B 100 µm, C, E–F 20 µm, D 10 µm. Abbreviations: AC, aciniform gland spigot; MAP, major ampullate gland spigot; mS, PMS median spicules (character 77); N, nubbin; PI, piriform gland spigot.

34 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 23. Drymusa capensis, female spinnerets. A. Spinnerets, posterior-ventral view, numbers on ALS articles (character 67). B. Same, lateral view (right). C. Left ALS. D. Left ALS, MAP area. E. PMS right, posterior view. F. Right PLS. Scales: A–B 200 µm, C, F 50 µm, D–E 20 µm. Abbreviations: AC, aciniform gland spigot; MAP, major ampullate gland spigot; mS, PMS median spicules (character 77); PI, piriform gland spigot.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 35 FIGURE 24. Scytodes globula, spinnerets. A–C. Female. D–F. Male. A. Spinnerets, lateral view (left), numbers on ALS articles (character 67). B. Left ALS. C. PMS–PLS, mesal view. D. Spinnerets, posterior-ventral view. E. Left ALS. F. Left PMS–PLS. Scales: A 200 µm, B–C 50 µm, D 100 µm, E–F 20 µm. Abbreviations: AC, aciniform gland spigot; MAP, major ampullate gland spigot; mS, PMS median spicules (character 77); N, nubbin; PI, piriform gland spigot.

36 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 25. Stedocys leopoldi, spinnerets. A–D. Male. E–F. Female. A. Spinnerets, posterior-ventral view, numbers on ALS articles (character 67). B. Left ALS, MAP area. C. Right ALS. D. Right PLS–PMS. E. Right ALS. F. Left PMS–PLS. Scales: A 200 µm, B 20 µm, C–F 50 µm. Abbreviations: AC, aciniform gland spigot; MAP, major ampullate gland spigot; mS, PMS median spicules (character 77); PI, piriform gland spigot.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 37 FIGURE 26. Sicarius rupestris, spinnerets. A, C, E. Female. B, D, F. Male. A. Spinnerets, lateral view (right), numbers on ALS articles (character 67). B. Spinnerets, posterior-ventral view. C. Right ALS, lateral view. D. Left ALS. E. PMS, posterior view. F. Left PMS. Scales: A–B 200 µm, C 100 µm, D, F 20 µm, E 50 µm. Abbreviations: AC, aciniform gland spigot; PI, piriform gland spigot.

38 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 27. Spinnerets. A–B. Sicarius rupestris. C–F. Loxosceles rufescens. A. Female. B–F. Male. A. Right PLS, mesal view. B. Left PLS. C. Spinnerets, posterior-ventral view, numbers on ALS articles (character 67). D. Right ALS. E. PMS, anterior view. F. Left PLS, arrow to setal pit. Scales: A–B 50 µm, C 200 µm, D–F 20 µm. Abbreviations: AC, aciniform gland spigot; MAP, major ampullate gland spigot; PI, piriform gland spigot.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 39 FIGURE 28. Loxosceles rufescens, female spinnerets. A. Spinnerets, posterior-ventral view, numbers on ALS articles (character 67). B. Right ALS. C. PMS, posterior view. D. PMS, anterior view. E. Left PLS, arrow to setal pit. F. Left ALS, MAP area. Scales: A 200 µm, B–E 20 µm, F 10 µm. Abbreviations: AC, aciniform gland spigot; MAP, major ampullate gland spigot; PI, piriform gland spigot.

40 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 29. Female genitalia. A–B. Periegops suteri. C–D. Drymusa capensis. E–F. Scytodes globula. A, E. Dorsal view. B. Closeup to receptacules. C. Right lateral view. D. Left posterior receptaculum, closeup. F. Right receptaculum, closeup. Scales: A, C, F 100 µm, B 20 µm, D 50 µm, E 200 µm. Abbreviations: GD, gland ductules; IS, inner spermatheca; S, spermatheca.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 41 FIGURE 30. Female genitalia. A–B. Stedocys leopoldi. C–D. Loxosceles rufescens. E. B. Sicarius rupestris. A, C, E. Dorsal view. B– D. Right receptaculum, closeup, lateral view. (B, mirrored). Scales: A, E 200 µm, B, D 100 µm, C 500 µm. Abbreviations: IS, inner spermatheca; OS; outer spermatheca; S, spermatheca.

42 · Zootaxa 3312 © 2012 Magnolia Press FACUNDO M. LABARQUE FIGURE 31. Female external genitalia. A. Periegops suteri. B. Drymusa capensis. C. Scytodes globula. D. Stedocys leopoldi. E. Loxosceles rufescens. F. Drymusa rengan. A–F. Ventral view. Scales: A–F 500 µm. Abbreviations: EF, epigastric furrow; pF, posepigastric foveae.

PHYLOGENY OF SCYTODOIDEA Zootaxa 3312 © 2012 Magnolia Press · 43 FIGURE 32. The single most parsimonious tree under equal weights, and implied weights with concavity values k = 4–9. Unambigu- ous synapomorphies are mapped on branches. Empty and filled hashmarks represent homoplasious and nonhomoplasious transforma- tions respectively. Large circles on each branch display Bremer support values / jackknife frequencies.