A tamerican museum Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3556, 26 pp., 14 figures, 1 table March 8, 2007 Spinneret Spigot Morphology in Synaphrid Spiders (Araneae, Synaphridae), with Comments on the Systematics of the Family and Description of a New Species of Synaphris Simon 1894 from Spain LARA LOPARDO,1 GUSTAVO HORMIGA,2 AND ANTONIO MELIC3 ABSTRACT We describe for the first time the spigot morphology of two synaphrid species (one of each of two synaphrid genera, Synaphris and Cepheia) as well as the morphology of the respiratory system of Synaphris. We also provide a taxonomic description of a new species of Synaphris from Spain, including detailed information about its morphology. This new species is known only from males, and it might belong to the so-called letourneuxi species group. Some morphological features proposed as synapomorphies for the genus Synaphris and/or the Synaphridae are questioned and discussed. Putative synapomorphies proposed here include a distinct constriction on the tarsus- metatarsus joints; a cheliceral keel ending in a strong promarginal cheliceral tooth; scarce number of maxillary setae; distal maxillary setae clavate; and a characteristic palpal morphology, comprising a distinctive tibial morphology, a modified cymbium with two separate areas, a palpal dorsal translucent expansion of the embolar base, a retrolateral paracymbium, a reduced furrow separating the major ampullate field from the piriform field, and the retention of at least one triad spigot in males. Refuted synapomorphies are the metatarsal subdistal anastomosed lyriform organ, the notched tibial trichobothrial base, and the tarsal pseudosegmentation. We also discuss the phylogenetic placement of the family, suggesting a close relationship to the araneoid Cyatholipidae. 1 Department of Biological Sciences, The George Washington University, 2023 G St. NW, Washington, D.C. 20052, USA ([email protected]). 2 Department of Biological Sciences, The George Washington University), 2023 G St. NW, Washington, D.C. 20052, USA ([email protected]). 3 Sociedad Entomologica Aragonesa (SEA), Avenida Radio Juventud 37, 50012 Zaragoza, Spain ([email protected]). Copyright © American Museum of Natural History 2007 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3556 Fig. 1. Map showing the geographic distribution of the described species of Cepheia and Synaphris around the Mediterranean region in Europe. For the geographic distribution of the recent species from Madagascar, see Miller (in press). For Synaphris, each location represents one described species. For Cepheia, all locations represent the geographical distribution of Cepheia longiseta. INTRODUCTION metatarsi (all features above from Marusik and Lehtinen, 2003); embolar and cymbial The recently erected araneoid spider family conductor; and pseudosegmentation of tarsi Synaphridae Wunderlich 1986 (Marusik and (Marusik et al., 2005). Most of the features Lehtinen, 2003, see also Schiitt, 2003) com¬ listed above were observed in only one or two prises two genera of minute spiders: the Synaphris species and generalized for the monotypic Cepheia Simon 1894, from south¬ family. No detailed study of the morphology ern Europe, and Synaphris Simon 1894, with of Cepheia longiseta and of most Synaphris seven species described from the Canary species has ever been done, and their spinneret Islands, Croatia, Ukraine, Turkmenistan, spigot morphology remains undescribed. and Egypt (fig. 1; Platnick, 2006). Two new Given the importance of spigot morphology Synaphris species and a third synaphrid genus for reconstructing the cladistic relationships of from Madagascar are currently being de¬ araneoid families (e.g., Griswold et al., 1998, scribed (Miller, in press). Recent studies 2005) the study of this character system should (Miller, in press; Marusik et al., 2005; contribute to understanding the phylogenetic Marusik and Lehtinen, 2003; Schiitt, 2003) position of synaphrids. In this paper we have proposed putative synapomorphies in describe for the first time the spigot morphol¬ support of the monophyly of the family and/or ogy of two synaphrid species (one of each of of the genus Synaphris. The familial synapo¬ two synaphrid genera; see also Miller, in press, morphies include a cheliceral keel that ends in for the spinneret spigot morphology of three a single strong prolateral tooth (Wunderlich, 1986, 1995; “tooth-like projection”, Schiitt, recently described synaphrid species). We also 2003; also “process with pointed tip”, provide a taxonomic description of a new Marusik and Lehtinen, 2003); presence of species of Synaphris from Spain, including paracymbium; cymbium modified into detailed information about the external mor¬ a rounded lobe; palpal tibia pressed against phology and the tracheal system. We discuss the cymbial base and with two pits (Marusik some of the morphological features of sy¬ and Lehtinen, 2003: 147, fig. 17; see naphrids and their potential value as system¬ Discussion below); a flattened and transparent atic characters. The new species is the first extension of the embolic base called lamella; record of the genus Synaphris for the Iberian absence of leg spines; tarsi longer than Peninsula. In addition, we report the first metatarsi; a notched tibial trichobothrial base; records of Cepheia from Portugal and the anastomosing lyriform organs on subdistal Baleares Islands. 2007 LOPARDO ET AL.: SYNAPHRID SPIGOT MORPHOLOGY 3 TABLE 1 Length of Individual Segments and Total Length of Left Legs, for Seven Specimens of Synaphris saphiynis, n.sp. Measurements are in millimeters, range in parentheses. Lemur Patella Tibia Metatarsus Tarsus Total Leg I 0.30 (0.30-0.34) 0.12 (0.12-0.13) 0.25 (0.25-0.28) 0.19 (0.18-0.20) 0.20 (0.20-0.22) 1.05 (1.05-1.15) Leg II 0.29 (0.28 0.30) 0.12 (0.11-0.12) 0.22 (0.22-0.25) 0.17 (0.17-0.19) 0.20 (0.20-0.22) 1.00 (0.99-1.07) Leg III 0.25 (0.25-0.29) 0.11 (0.09-0.13) 0.18 (0.17-0.22) 0.16 (0.16-0.18) 0.18 (0.17-0.20) 0.88 (0.88-0.97) Leg IV 0.30 (0.30-0.35) 0.12 (0.11-0.13) 0.27 (0.27-0.29) 0.19 (0.19-0.20) 0.20 (0.19-0.22) 1.09 (1.09-1.17) MATERIAL AND METHODS averages are slightly different. We comment on and propose some changes in the system of Methods of study follow Hormiga (2003). names used by Marusik et al. (2005). Study Specimens were studied in 75% ethanol using specimens will be deposited in the American a Leica MZAPO stereomicroscope. For ob¬ Museum of Natural History (AMNH, New servation of respiratory structures, the abdo¬ York), the California Academy of Sciences mens of two specimens were bisected horizon¬ (CAS, San Francisco), the Museum of tally and digested with SIGMA Pancreatin LP Comparative Zoology (MCZ, Harvard 1750 enzyme complex in a solution of sodium University), and the Museo Nacional de borate, prepared following the concentrations Ciencias Naturales de Madrid (MNCN, described by Dingerkus and Uhler (1977) as Spain). For abbreviations used through fig¬ modified in Alvarez Padilla and Hormiga (in ures and text see appendix 1. prep.). Some of the tracheal system prepara¬ tions were stained with an aqueous solution of chlorazol black as described in Hormiga RESULTS (1994). The bisected abdomen was left in the Synaphris saphrynis, new species enzymatic solution overnight and at room figures 1-62 temperature. After the enzymatic digestion, the specimens were transferred to distilled Types: 1 $ holotype and 7 $ paratypes water for observation. All measurements are from SPAIN: Toledo, Huecas, 29.V.2003, in millimeters. Carapace height was measured Kleijn et al. col. (30T-395937) holotype at the highest point, from the carapace lateral in MNCN-20.02/16523; paratypes in edge, not from the sternum. Abdominal MNCN-20.02/16524, \$ paratype in height, length, and width were measured at AMNH, 1 $ paratype in MCZ, 1 $ paratype the maximum points. To account for length in CAS). variation, measurements are expressed first as Etymology: The species epithet is an the length of the described specimen, then as arbitrary combination of letters. the range of all observed specimens (in Diagnosis: Males of Synaphris saphrynis, parentheses, table 1). After dissection, palps n.sp. can be distinguished from other were cleared in clove oil. Palp drawings were Synaphris species by the following combina¬ made with a camera lucida attached to a Leica tion of palpal characters: conductor with DMRM compound microscope. For SEM tegular groove accompanying the distal por¬ images, the specimens were critical-point dried tion of the embolus; with two distal apophy¬ and sputter-coated with gold-palladium. ses, the ventral one (hereafter “Cap”, “sub- Images were taken with a LEO 1430VP terminal apophysis” of Marusik et al. microscope at the Department of Biological 2005) clearly bipartite with both tips round¬ Sciences (GWU) SEM facility. Species de¬ ed (i.e., without an irregular border); an scriptions and measurements follow Lopardo embolar membranous expansion (“lamella” (2005) and Marusik et al. (2005). Leg formula of Marusik et al. 2005); width of embolar refers to the relative length of legs. Two legs expansion base 2/5 the width of the expansion are considered equally long when their range (i.e., base of expansion width/expansion of variation highly overlaps, even if their width: 0.40). 4 AMERICAN MUSEUM NOVITATES NO. 3556 Figs. 2-6. Synaphris saphrynis, n.sp. Male paratype. 2. Ventral view; 3. Frontal view; 4. Cephalothorax, lateral view; 5. Detail of mouthparts, frontal view, arrow to clavate maxillary setae; 6. Detail of cheliceral prolateral short hairs. Description: Male: Total length 1.00 setae located between LE (one pair), and (0.95-1.08). Carapace length 0.44 (0.43-0.47), slightly posterior to central one behind PME. width 0.40 (0.38-0.43), height 0.20 (0.19-0.23). Chelicerae with median keel ending in single Carapace with four setae (bases referred to as strong promarginal tooth; retromarginal teeth “tubercles” in Marusik et al., 2005) along absent (figs. 3, 5, 8, 9). Promargin of chelicera midline and four laterally, two on each side with three lateral short hairs with larger bases (figs. 3, 4). Midline setae located on clypeus and rounded strong scale on dorsal surface (one), slightly posterior to PME (one), and on (figs. 3, 6). Labrum globose, with minute dorsalmost carapace surface (two). Lateral denticles grouped on its ventral surface, 2007 LOPARDO ET AL.: SYNAPHRID SPIGOT MORPHOLOGY 5 Figs. 7-10. Synaphris saphrynis, n.sp. Male paratype. 7. Sternum, ventral view; 8. Mouthparts, ventral . . view; 9 Mouthparts, posterior view; 10 Detail of mouthparts, showing labrum and labium, ventral view. globose expansion arising from anterior sur¬ dorsal band of anastomosed ridges (figs. 17, face (figs. 5, 8, 10). Maxillary setae scarce 18; the “subdistal anastomosed lyriform or¬ (figs. 5, 8, 10; compare to symphytognathid gan” of Marusik and Lehtinen, 2003). Legs maxillary setae in Griswold et al. 1998: without spines, tarsal organ located in middle fig. 21C), distal maxillary setae clavate (arrow dorsal region of tarsus, capsulate, with round¬ in fig. 5). Clypeus slightly convex, height 0.14 ed orifice (figs. 19, 20). Three claws, serrate (0.12-0.15), ca. 5-6 AME diameters. Sternum accessory (false claw) setae present. Claw teeth length 0.27 (0.24-0.28), width 0.29 (0.26-0.30), (paired claws/ inferior claw): Leg I, paired length/width 0.96 (0.89-1.00), cuticle squa- claws with five teeth/ inferior claw with two mate (fig. 7), posterior margin truncated, teeth and one dorsal denticles (figs. 23-25); II wide, about two times width of coxa IV and III, four teeth/ as leg I; IV, two teeth/ two (figs. 2, 7). Abdomen oval, length 0.50 (0.47- teeth and one dorsal denticle (figs. 28, arrow 0.64), width 0.44 (0.40-0.47), height 0.36 in 29). Distal tooth two times longer than (0.36-0.43). Three epiandrous spigots cen¬ other teeth in paired claws. Leg hairs serrate. trally distributed along the epigastric furrow Cuticular surface of appendages squamate (figs. 31, 32). Legs: leg formula 4=123. Leg (fig. 20). Tarsus slightly longer than meta¬ measurements: see table 1. Femoral spot and tarsus on legs I and II, same length on legs III prolateral clasping spine absent (figs. 11, 13, and IV (see table 1, figs. 11, 12, 16, 27). 14). Setae on legs with large elevated, striated Trichobothria: Trichobothrial bases simple bases (figs. 17, 18, 20, 22). Leg tarsi without and smooth, with proximal hood bearing pseudosegmentation (figs. 12, 19, 23, 24). two lateral ridges, similar on all legs and Tarsal-metatarsal joint constricted (figs. 11, segments (figs. 15, 21, 22). Tarsal trichobo¬ 12, 16-18, 27). Distal area of metatarsi with thria absent. Legs I and II, tibia 1-1-0; 6 AMERICAN MUSEUM NOVITATES NO. 3556 2007 LOPARDO ET AL.: SYNAPHRID SPIGOT MORPHOLOGY 7 metatarsus 0-1-0. Legs III and IV, tibia pl-2- located dorsal and distally (fig. 56). Cymbium 0-0; metatarsal trichobothria absent. Color: capsulated, comprising two distinct areas: one Carapace brown with undefined lighter radii, rounded, dorsal, retrolateral, with hairs, one surface slightly wrinkled; sternum brown, membranous (indistinguishable under light darker than carapace, not homogeneous but microscope), prolateral, with no setae without definite pattern, border with thick (figs. 49, 54). Retrolateral margin of cymbium orange stripe. Legs orange. Abdomen greenish with notch delimiting basal paracymbium black, with pattern of irregular transversal (figs. 47, 52, 53, 60). Measurements: femur lighter bands. Eyes: AME black, other eyes 0.12 (0.11-0.14), patella 0.05 (0.05-0.06), tibia pearly white. Diameter: AME 0.02, PME 0.02, length 0.07(0.06-0.07), tibia width 0.12 (0.09- PLE 0.03, ALE 0.04. Respiratory system: 0.12), tibia length/width 0.55 (0.55-0.67). Anterior booklungs transformed into trache¬ Embolus filiform, long, thin with thickened ae, connected by a transverse duct (figs. 30, tip (figs. 46, 51, 59, 62). Embolar base flat, 35-37, 39). Anterior spiracles connected to ventral, with dorsal flat translucent, membra¬ epigastric furrow (figs. 2, 31). Five tracheal nous expansion (figs. 46, 49, 50; “lamella”, tubes arise from each anterior spiracle, four Marusik et al., 2005; Marusik and Lehtinen, oriented anteriorly toward cephalothorax, one 2003). Base of embolar expansion about 2/5 its oriented laterally first, then turning poster¬ maximum width. Embolus running clockwise iorly. Posterior tracheal system (figs. 30, (in left palp), exteriorly surrounding junction 34, 37-39) with two distant spiracular open¬ of two areas of cymbium, which areas may act ings (figs. 2, 33) exteriorly connected by thin as cymbial conductor (figs. 46, 54). Tegular ridge (i.e., one wide spiracular opening). Thin groove also present, accompanying embolus ridge leading to deep, flat, membranous toward tip of bulb, terminating in pointed atrium, anteriorly ending in sclerotized U- apophysis (figs. 50, 59; “terminal apophysis”, shaped duct that connects tracheal ducts Marusik et al., 2005; Marusik and Lehtinen, arising from spiracles. Two main tracheal 2003). This apophysis may act as conductor, bundles arise from the junction of tracheal and presents small pore opening close to tip ducts and U-shaped atrial duct, one on each (fig. 61).Dorsal edge of embolar expansion side, directing tracheoles mainly anteriorly. with weak furrow, which might also be related Smaller clumps diverge laterally, some tra¬ to embolus (figs. 50, 51). Ventral to pointed cheoles seem to branch off clumps and apophysis is another apophysis, with two disperse irregularly around abdominal space. pointed tips (figs. 50, 57, 58; “subterminal Both tracheal systems seem to reach into apophysis”, Marusik et al., 2005; Marusik and prosoma. Spinnerets (fig. 40): Colulus large Lehtinen, 2003). Spermatic duct seems to and fleshy, triangular, about half length and undergo one transverse loop before reaching width of ALS, with three setae (fig. 41). ALS embolar base. Diameter of spermatic duct (fig. 42) with one MAP spigot, accompanied gradually decreases throughout its length, by nubbin and tartipore, separated by weak except for a sudden widening before entering furrow from PI field, which contains four base of embolus for fraction of loop length piriform spigots with reduced bases, inter¬ (arrow in fig. 48). spersed with few tartipores. PMS (fig. 43) with Female: Unknown. only one spigot, no nubbin. PLS (figs. 44, 45) Natural History: The specimens were with only two spigots of different morphology collected in pitfall traps from dry wheat and (see Discussion below). Palp (figs. 46-62): barley fields. Tibia rounded retrolaterally, without apophy¬ Distribution: Known only from the type ses (figs. 47, 53, 55). One tibial trichobothrium locality. Figs. 11-18. Synaphris saphrynis, n.sp. Male paratype, right leg I. 11. Prolateral view; 12. Tarsus and metatarsus, retrolateral view; 13. Femur, ventral view; 14. Same, detail at femoral spot area; 15. Tibia and patella, dorsal view; 16. Tarsus and metatarsus, dorsal view; 17. Detail of tarsus-metatarsus joint, dorsal view; 18. Same, prolateral view. AMERICAN MUSEUM NOVITATES NO. 3556 2007 LOPARDO ET AL.: SYNAPHRID SPIGOT MORPHOLOGY 9 Other Material Examined: SPAIN: Toledo: Huecas, same locality, 29.V.2003, 6 <?; 15.v.2003, 1 $, Kleijn et al. col. (A. Melic 5739-A). Cepheia longiseta (Simon, 1894) figures 63-80 Description of Spigot Morphology: Male: Colulus large, fleshy, triangular, about half length and width of ALS, with three setae (figs. 63, 64). ALS (figs. 65, 66) with one MAP spigot, accompanied by a nubbin and a tarti- pore, separated by weak (almost nonexistent) furrow from PI field. PI field, on external side of ALS, contains three PI spigots with reduced bases, posterior PI spigot base larger. PMS (figs. 67, 68) with two spigots of similar morphology, chemosensory seta (can be con¬ fused with spigot) located anteriorly; external¬ ly, its base deepens around shaft. PLS (figs. 69-71) with two spigots of slightly different morphology clumped in same field. Internal one with rounded, larger base and more cylindrical shaft, external one with oval base and tapering shaft. Short, thick chemo¬ sensory seta (can be confused with a small spigot) located more basally on internal side of distal PLS segment. Female: Colulus large, fleshy, triangular, about half length and width of ALS, with four setae (figs. 72, 73). Spinnerets as in male, except: four PI spigots (instead of three) on ALS (figs. 74, 75); one external CY spigot on PMS (figs. 76-78); and one internal CY on PLS (figs. 79, 80). Material Examined: 1 $ and 1 ? para- lectotypes from FRANCE (“Gallia”) coll. Figs. 27-29. Synaphris saphrynis, n.sp. Male paratype, right leg IV. 27. Tarsus and metatarsus, Simon 4538, b.849 (MNHN-AR 1059). dorsal view; 28. Tarsal claws, frontal-retrolateral Distribution: Cepheia longiseta has been view; 29. Same, retrolateral view, arrow to dorsal collected from dry regions and coastal areas of denticle on inferior claw. the western Mediterranean region in southern Figs. 19-26. Synaphris saphrynis, n.sp. Male paratype, right leg I. 19. Tarsus, dorsal view; 20. Same, detail of tarsal organ; 21. Tibia, dorsal view, detail of trichobothrial base; 22. Metatarsus, dorsal view, detail of trichobothrial base; 23. Tarsal claws, prolateral view (prolateral superior claw broken); 24. Same, retrolateral view; 25. Same, dorsal view; 26. Tibia-metatarsus joint, prolateral view. 10 AMERICAN MUSEUM NOVITATES NO. 3556 Fig. 30. Synaphris saphrynis, n.sp. Male paratype, schematic drawing showing tracheal system, dorsal view.