1999. The Journal of Arachnology 27:604–662 RADIATION OF THE GENUS DYSDERA (ARANEAE, DYSDERIDAE) IN THE CANARY ISLANDS: THE ISLAND OF TENERIFE Miquel A. Arnedo1 and Carles Ribera: Departament de Biologia Animal, Universitat de Barcelona, Avinguda Diagonal 645, E-08028 Barcelona, Spain ABSTRACT. AnoverwhelmingnumberofendemicspeciesbelongingtothespidergenusDysderahave been reported from the oceanic archipelago of the Canary Islands. A complete taxonomic revision is currently being performed in order to assess the extent of this species’ radiation, as well as to supply enoughdatatoplaceitinaphylogeneticframework.ThepresentarticleisdevotedtotheDysderaspecies inhabitingtheislandofTenerife.Atotalof22speciesisrecognizedinTenerife,includingthecosmopolitan Dysdera crocota C.L. Koch 1839. Two new species are described: Dysdera guayota new species and Dysdera hernandezi new species. Ten new synonymies are reported: D. moquinalensis Wunderlich1991 andD.vilaflorensisWunderlich1991(cid:53)D.brevispinaWunderlich1991;D.medinaeWunderlich1991(cid:53) D.cribellataSimon1883;D.inaequuscapillataWunderlich1991(cid:53)D.crocota;D.pergradaWunderlich 1991, D. pseudopergrada Wunderlich 1991, D. tabaibaensis Wunderlich 1991, D. teideensis Wunderlich 1991 and D. teneriffensis Strand 1908 (cid:53) D. macra Simon 1883; D. obscuripes Wunderlich 1991 (cid:53) D. propinqua Ribera, Ferra´ndez & Blasco 1985. Sixteen species are redescribed: D. ambulotenta Ribera, Ferra´ndez & Blasco 1985; D. brevisetae Wunderlich 1991, D. brevispina Wunderlich 1991; D. chioensis Wunderlich1991;D.cribellataSimon1883;D.curvisetaeWunderlich1987;D.esquiveliRibera&Blasco 1986; D. gibbifera Wunderlich 1991; D. gollumi Ribera & Arnedo 1994; D. labradaensis Wunderlich 1991; D. macra Simon 1883; D. minutissima Wunderlich 1991; D. montanetensis Wunderlich 1991; D. propinqua Ribera, Ferra´ndez & Blasco 1985; D. unguimmanis Ribera, Ferra´ndez & Blasco 1985 and D. volcania Ribera, Ferra´ndez &Blasco 1985. The femalesoffourspecies: D.brevisetae,D.brevispina,D. minutissima and D. montanetensis are described for the first time. Females formerly assigned to both D. gibbifera and D. volcania are considered to be incorrect identifications. A neotype is designated for D. macra. The presence of D. rugichelis Simon 1907 in Tenerife is considered to be doubtful. Ecological and distributional patterns of the speciesare discussed. Species of the spider genus Dysdera La- volcanic archipelagos, the Canary Islands, treille1804areusuallyfoundinslightlydamp harbors about 50 endemic species (Simon but warm ground habitats. They arenocturnal 1883, 1907; Strand 1908; Schmidt 1973; Ri- wandering hunters that spend daytime in silk- bera etal.1985;Ribera&Blasco1986;Wun- en cocoons under stones, logs or bark (Rob- derlich 1987, 1991; Ribera & Arnedo 1994; erts 1995; pers. obs.). Dysdera specimens are Arnedo et al. 1996; Arnedo & Ribera 1997), notunusualincaves,whichcanbeconsidered which represent about one quarter of the de- as an expansion of their typical habitats; and scribed species in the genus to date. This fig- several cases of troglomorphic species have ure is even more remarkable when compared been reported (Ribera 1983, 1993; Ribera et withthenumberofendemicsintheremaining al. 1986). This species-rich genus includes archipelagos: one from the Azores (undescri- about 200 species with a circum-Mediterra- bed species), five from Madeira (Denis 1962; nean distribution, with the single exceptionof Wunderlich 1994) and one from Cape Verde the anthropophilous cosmopolitan D. crocota (Berland 1936). In addition, seven of these C.L. Koch 1839. The so-called Macaronesian species were troglobites with morphological archipelagos (Fig. 1) represent the western- adaptations to the hypogean environment. most limit of Dysdera’s range. One of these Nevertheless,thisoverwhelmingnumberof Dysdera species held by the Canaries could 1Division of Insect Biology, ESPM, University suggest a taxonomic artifact instead of a true of California–Berkeley, 201 Wellman Hall, Berke- speciesradiation.AdeeperlookintoCanarian ley, California 94720-3112 USA Dysdera taxonomy revealed some instances 604 ARNEDO& RIBERA—THEGENUS DYSDERA IN THE CANARYISLANDS 605 Figures 1–3.—Maps 1–3. 1, Macaronesian archipelagos; 2, the Canary Islands; 3, Tenerife, with its oldest areas encircled. that could, at least, call into question this isodes that formed the archipelago are prob- amazing number of endemics. On the one ably the result of a propagating fracture orig- hand, 22 out of the 50 recognized species inated in the Atlas formation during the weredescribedfromonlyoneofthesexes,19 Alpine orogeny, about 25 Mya ago (Anguita ofwhichwereknownfromasinglespecimen. & Herna´n 1975). This model would explain Moreover, some of the species lacked infor- both the reduction of the age of the islands mation regarding their locality, type material from east to west and the continuation of ac- was lost, or both. Finally, 27 specieswerede- tive volcanism in the older islands. The ap- scribed in a single publication together with proximate ages of the subaerial parts of the 106 new species from the Macaronesia(Wun- islands, as recovered using the K-Ar tech- derlich 1991). On the other hand, most of the nique, range from about 22 Mya to less than published descriptions of the Canarian Dys- 1 Mya. The estimated geological age for each dera were vague enough to correspond to islandis:Fuerteventura20-22Mya,Lanzarote more than one species, or failed to supply the 15-19 Mya, Gran Canaria 14–16 Mya, Ten- necessary information for the study of such erife 11.6–14 Mya, La Gomera 10–12 Mya, interesting spiders in a phylogenetic frame- LaPalma1.6–2MyaandElHierro0.8–1Mya work. (Cantagrel et al. 1984, Mitchell-Thome´ 1985, Withtheaimofconfirmingtheexistenceof Ancocheaetal.1990,Coelloetal.1992).The this radiation, completing the species descrip- island of Tenerife is located roughly at the tions as well as their geographical distribu- center of the line drawn through the archipel- tions and, finally, offering enoughdatatoper- ago. Tenerife is both the biggest (2058 km2) form a phylogenetic analysis of the group, a and the highest (3717 m) island in the archi- major revisionary work on Canarian Dysdera pelago. is currently being developed (Ribera & Ar- Elevation together with trade winds play nedo1994;Arnedo&Ribera1996;Arnedoet important ecological roles on oceanic islands, al.1996;Arnedo&Ribera1997).Thepresent especiallyattropicalandsubtropicallatitudes. article is devoted to thetaxonomicrevisionof Theyarebothresponsibleforthepresenceand the genus on the Island of Tenerife. distribution of the different ecological zones. The Canary Islands lie in the Atlantic IntheparticularcaseoftheCanaries,thejoint Ocean 100 km from the north-western coast effect of the humid and cool NE trade winds, of Africa (Fig. 2). The different volcanic ep- betweenaltitudesof400–1200m,andthedry 606 THE JOURNALOF ARACHNOLOGY trade winds from the NW, above 2000 m, 1883 ((cid:63),(cid:47); one locality); D. labradaensis cause a temperature inversion. In this area, a Wunderlich 1991 ((cid:47), one locality); D. levipes nearly permanent cloud belt is formed. Con- Wunderlich 1987 ((cid:63),(cid:47)); D.medinaeWunder- sequently,strongecologicalsegregationisob- lich 1991 ((cid:63),(cid:47)); D. minutissima Wunderlich served between northern, more humid, and 1991 ((cid:63), single specimen); D. montanetensis southern, dryer slopes. Five major ecological Wunderlich 1991 ((cid:63), single specimen); D. zonescanberecognizedonnorthernslopesof moquinalensis Wunderlich 1991 ((cid:63), single the islands. The first, from the seashore to up specimen); D. obscuripes Wunderlich 1991 to 250 m, is characterized by the presence of ((cid:63),(cid:47)); D. pergrada Wunderlich 1991 ((cid:63),(cid:47); dry-arid subtropical shrubs. The second, from one locality); D. propinqua Riberaet al.1985 250–600 m, features humid to semi-aridtrop- ((cid:63), single specimen); D. pseudopergrada ical shrubs and woods. The third, from 600– Wunderlich 1991((cid:63),(cid:47)); D. rugichelis Simon 1000 m, is covered by the cloud belt and fea- 1907 ((cid:63), single specimen in Tenerife); D. ta- tures a typical subtropical wood,theso-called baibaensis Wunderlich 1991 ((cid:63), single spec- laurel forest. In the fourth, from 1000–2000 imen); D. teideensis Wunderlich 1991 ((cid:63),(cid:47)); m, anendemicpineforestoccurs.Finally,dry D. teneriffensis Strand 1908 ((cid:47); single spec- subalpine shrubispresentfrom2000mtothe imen, lost); D. unguimmanis Ribera et al. top. Southern slopes lack a laurel forest zone 1985 ((cid:47), single specimen); D. vilaflorensis andtransitionbetweensub-aridshrubsandthe Wunderlich 1991 ((cid:63),singlespecimen)andD. pine forest takes place at higher elevation. volcania Ribera et al. 1985 ((cid:63),(cid:47); one locali- Apart from these climatic-related ecosys- ty) (Bo¨senberg 1895; Strand 1908; Denis tems, an additional ecological zone is present 1941,1953;Schmidt1975;Riberaetal.1985; in volcanic islands: the hypogean environ- Wunderlich 1987, 1991; Ribera & Arnedo ment. The subterranean environment in the 1994; Arnedo et al. 1996; Arnedo & Ribera Canariesisrepresentedbybothlavatubesand 1997).Sixofthesespeciesdisplayedmorpho- the MSS (mesocavernous shallow stratum) logical adaptations to the hypogean environ- (Juberthie et al. 1980, 1981; Orom´ı et al. ment and were considered to be true troglob- 1986; Medina 1991). Due to their short life- ites. With a single exception (Dysdera span,lavatubesarefoundonlyinareasofthe ratonensis Wunderlich 1991 from La Palma), islands with a relatively recent pahoehoe-like the lava tubes of Tenerife hold all troglo- basaltic volcanism. This explains the lack of morphic Dysdera documented so far in the tubes in the islands of La Gomera and their Canaries. scarcity in Gran Canaria and most of Fuerte- METHODS ventura. However, even in the absence of caves, a very rich underground environment, The current study was based on the adop- in the form of shallow, intermediate-sized,in- tion of the so-called ‘diagnosability’ (Baum terconnected voids, is present in all the is- 1992) phylogenetic species concept (Nixon & lands. Wheeler 1990, 1992; Wheeler & Nixon 1990, Before the present study 29 endemic spe- Davis & Nixon 1992). Species arerecognized cies of Dysdera had been reported from Ten- as the most exclusive set of populations that erife,byfarthemostspecies-richislandinthe display a unique combination of character- archipelago. These species were: D. ambulo- states, when semaphoronts arecompared(Da- tenta Ribera et al. 1985 ((cid:63),(cid:47); one locality); vis&Nixon1992).Thisconceptwasselected D. brevisetae Wunderlich 1991 ((cid:63), single because it is easily applicable in practice, it specimen); D. brevispina Wunderlich 1991 avoids any referencetoprocesses,andisfully ((cid:63), single specimen); D. chioensis Wunder- compatible with a phylogenetic framework. lich 1991 ((cid:47), one locality); D. cribellata Si- However, this definition is not free of theo- mon 1883 ((cid:63),(cid:47)); D. curvisetae Wunderlich reticalproblems(Frost&Kluge1994)andhas 1987 ((cid:63), single specimen); D. esquiveli Ri- been considered to be excessively restrictive. bera&Blasco1986((cid:63),(cid:47));D.gibbiferaWun- Inaddition,inthepresentapproximation,only derlich 1991 ((cid:63),(cid:47)); D. gollumi Ribera & Ar- morphological characters were taken into ac- nedo 1994 ((cid:47), one locality); D. iguanensis count, which has probably resulted in an un- Wunderlich1987((cid:63),(cid:47));D.inaequuscapillata derestimation of the total number of species. Wunderlich 1991 ((cid:63),(cid:47)); D. insulana Simon Additionalstudiesconsideringmolecular,eco- ARNEDO& RIBERA—THEGENUS DYSDERA IN THE CANARYISLANDS 607 logicalorbehavioralcharacterswouldbenec- by a slash). In patellae, the number of spines essary in order to recover the total amount of and their position (ventral or dorsal) wascod- diversity of the genus. ed.Tibiaeusuallyshowthemostcomplexspi- The first stage in the assessment of the tax- nation pattern. For each tibia, the number of onomic status of the Tenerifean species was spines was recorded from four zones (here- to gather a large number of specimens (350), after referred as ‘bands’): proximal, medial- which were made available from scientificin- proximal, medial-distal and distal. For coding stitutions, various personal collections, and purposes, the number of spines on thefrontal, three collection expeditions to the island by medial and posterior regions were separated theauthors.Thefollowingcolleaguesandmu- by points. In the descriptions(intra-individual seum kindly supplied material for the present variation), hyphens separate the number of study:Dr.E.EnghofffromtheZoologiskMu- spines on each side of the body if different. seumofCopenhagen(ZMK),R.Garc´ıa‘Felo’ In the tables (intraspecific variation) hyphens (S/C de la Palma, Canary Islands) (RG), F. separate the minimum and maximum number Gasparo (Trieste, Italy) (FG), Mr. P.D. Hill- of spines observed in any specimen. Between yard from the Natural History Museum of 6–10 individuals were examined from each London (BMNH), Dr. M. Grasshoff from the species whenever possible. Forschungsinstitut und Naturmuseum Senck- Structures of male bulb and female vulva enberg (SMF), Dr. P. Orom´ı from the Univ- were mostly named after Deeleman-Reinhold ersidad de La Laguna (UL), Dr. G. Ortega & Deeleman (1988), with the addition of sev- fromtheMuseodeCienciasNaturalesdeSan- eral features particular to Canarian Dysdera ta Cruz de Tenerife (MCNT), Dr. C. Rollard (Arnedo et al. 1996; Arnedo & Ribera 1997). from the Muse´um National d’Histoire Natu- Nevertheless, some new characters from the relle de Paris (MNHN) and J. Wunderlich female vulva have been added or have been (Straubenhardt, Germany) (JW). The material redefined and deserve further considerations. provided by the authors’ expeditions is stored The vulva of the genus Dysdera is divided at the collection of Arachnids of the Univer- into two major diverticles: an anterior diver- sity of Barcelona, Spain (UB). ticle and a posterior one. They are separated Characters were investigated under a Wild by the epigastric furrow at the ventral side, Heerbrugg (12–100(cid:51)) dissecting microscope. andtheoviductopeningatthedorsalone.The Female vulva ((cid:53) endogyne, Mcheidze 1972) posterior diverticle is mostly membranous, was removed and muscle tissues digested us- with the single exception of the transversal ingaKOH(35%)solutionbeforeobservation. bar. This is located at the anterior dorsal side Male bulbi and spinnerets were removed, and holds a frontal projection (‘bursal valve’, cleanedbymeansofultrasoundandexamined V) that closes the oviduct openings. On the using a HITACHI S-2300 scanning electron other hand, most of the anterior diverticle is microscope at 10–15 Kv. All measurements usually sclerotized. The most conspicuous are in millimeters. Somatic morphology mea- structure is a T-shaped spermatheca (S) locat- surements were taken using an ocular micro- edattheventralsideofthemostanteriormar- meterinthedissectingmicroscope.Characters gin.Themediallateralmarginsoftheanterior examined together with their diagnostic reso- diverticle are invaginated forming two differ- lutionhavebeendiscussedelsewhere(Arnedo ent pouches: a dorsal pouch, which corre- et al. 1996). All characters were recorded in sponds to the so-called ‘dorsal arch’ (DA) DELTAformat(Dallwitz1980;Dallwitzetal. (Deeleman-Reinhold & Deeleman 1988) and 1993). a ventral one, hereafter referred as ‘ventral Terminology.—Leg spination was record- arch’ (VA). The DA is usually completely ed for each segment using the format of Ar- sclerotized. The dorsal side of the DA locks nedo et al. (1996). Only spines present on fe- the V and is called the ‘dorsalfold’(DF)(Ar- morae, patellae and tibiaewereconsidered.In nedo etal.1996). Thefoldthatseparatesboth femora, spines are usually arranged in one or diverticles is named the ‘major fold’ (MF), to two (anterior and posterior) rows parallel to differentiate it from several additional folds the segment. The number of rows and spines that are sometimes found on the DA lateral per row were recorded (In Tables 2–16, the borders. The development and sclerotization number of spines in each row were separated degreeoftheMFarehighlyvariable.Insome 608 THE JOURNALOF ARACHNOLOGY Table 1.—Abbreviationsused in text and figures (Figs. 4–12). Female genitalia Male copulatory bulb DA (cid:53) dorsal arch T (cid:53) tegulum DF (cid:53) dorsal fold DD (cid:53) distal division MF (cid:53) major fold IS (cid:53) internal sclerite S (cid:53) spermatheca ES (cid:53) external sclerite TB (cid:53) transversalbar DH (cid:53) distal haematodoca V (cid:53) bursal valve C (cid:53) crest VA (cid:53) ventral arch AC (cid:53) additional crest AVD (cid:53) additional ventral diverticle LF (cid:53) lateral fold over L, betweeninternal and exter- nal sclerites Eyes L (cid:53) lateral sheet AME (cid:53) anterior medial eyes AL (cid:53) additional lateral sheet at back internal border PME (cid:53) posterior medial eyes P (cid:53) posterior apophysis PLE (cid:53) posterior lateral eyes Spinnerets Cheliceral teeth ALS (cid:53) anterior lateral spinnerets B (cid:53) basal tooth PMS (cid:53) posterior medial spinnerets M (cid:53) medial tooth PLS (cid:53) posterior lateral spinnerets D (cid:53) distal tooth MS (cid:53) major ampulate gland spigot PS (cid:53) polar pyriform gland spigot continental Dysdera species, the MF almost found in Deeleman-Reinhold & Deeleman entirely isolates the DA from the VA. The (1988). marginsoftheMFmaybemarkedlyseparated Dysdera ambulotenta Ribera, Ferra´ndez & from each other or stuck together forming an Blasco 1985 internal rim. The VA exhibits a widerangeof Figs. 13–24, Table 2 sclerotization levels, from mostly sclerotized to completely membranous. In most of the Dysdera ambulotenta Ribera, Ferra´ndez & Blasco Canarianrepresentatives,anadditionalventral 1985: 54–57, fig. 1 [(cid:63),(cid:47)]. Holotype male and diverticle (AVD) in the VA has been ob- paratype female (allotype) from CuevadelVien- to-Sobrado, El Amparo, Icod de los Vinos, Ten- served. The AVDisrecognizedbyaninternal erife, Canary Islands; 14 May 1981, J.L. Mart´ın rim ventral to the MF and by its own external leg.; (cid:63) (T-CS-17), (cid:47) (T-CS-18). Stored at UL. sclerotization, usually tooth shaped. Spinner- Examined. Wunderlich 1991: 284–287. etsandtheirassociatedspigotglandswereas- signed after Platnick et al. (1991). See Table Diagnosis.—Dysdera ambulotenta can be 1 for a complete list of abbreviations. distinguishedfromallotherCanarianDysdera species, except D. labradaensis, by its large FAMILY DYSDERIDAE size (carapace(cid:46) 6.5mm)andremarkableeye Genus Dysdera Latreille 1804 reduction. It differs from D. labradaensis Note: An excellent and thorough diagnosis (male unknown) by complete absence of both and description of the genus Dysdera can be the posterior lateral (PLE) and posterior me- Table 2.—Intraspecificspination variability of Dysderaambulotenta. Proximal Medial-proximal Medial-distal Distal Tibia 3 dorsal 1.0-2.1 1.0-2.0-1 0 1.0.1 Tibia 4 dorsal 0-1.0.0-1 1.0-3.1 0-1.0-3.0-1 1.0.1 Tibia 3 ventral 1.1-3.1 1.1-2.1 0 1.0.0-1 Tibia 4 ventral 1.1-2.1 1.1-2.1 0-2.0-2.1-2 0-1.0-2.1 Number of rows Number of spines Femur 3 dorsal 2 0-3/0-1 Femur 4 dorsal 2 0-4/0-4 ARNEDO& RIBERA—THEGENUS DYSDERA IN THE CANARYISLANDS 609 Figures 4–12.—Diagrams showing the characters included in the abbreviations list (Table 1). 4, Cara- pacefrontalregion,dorsalview;5,Leftchelicera,ventralview;6,Rightbulb,frontalview;7,Rightbulb distaltip,frontalview;8,Bulb,externalview;9,Bulb,posteriorview;10,Vulva,ventralview;11,Vulva, dorsal view; 12, Vulva, transversesection. dial eyes (PME), lack of spines on legs 1 and straight PME, PLE lost; AME markedly re- 2 and absence of an additional ventral diver- duced; AME diameter 0.09 mm; AME sepa- ticle (AVD) in the vulva (Fig. 17). In both ration 0.936 mm. Labium trapezoid-shaped, sexes absence of a polar spigot (PS) in the base wider than distal part; longer than wide anterior lateral spinnerets (ALS) is unique to at base; semicircular groove at tip. Sternum this species (Fig. 23). dark orange, frontally darker, becoming ligh- Description.—Holotypemale:Figs.13–15, ter towards back; very slightly wrinkled, 19–22. Carapace (Fig. 13) 7.35 mm long; mainly between legs, frontal border; uniform- maximum width 5.6 mm; minimum width ly covered in slender black hairs. 3.78 mm. Reddish-orange, frontally darker, Chelicerae (Fig. 14) 4.41 mm long, about 2⁄ 5 becoming lighter towards back; slightly fo- of carapace length in dorsal view; fang me- veateatborders,slightlywrinkledwithafine- dium-sized, 2.8 mm; basal segment dorsal textured granular surface primarily at the an- side completely covered with piligerousgran- terior end. Frontal border roughly straight, ulations (small, densely), ventral sidesmooth. from 1⁄ to 3⁄ carapace length; anterior lateral Chelicera inner groove medium-size, about 2⁄ 2 5 5 bordersconvergent(backwardslong,parallel); cheliceral length; armed with three teeth and roundedatmaximumdorsalwidthpoint,back lamina at base; D (cid:46) B (cid:46) M (large teeth; D, lateral borders rounded; back margin wide, Bnotverydifferent);Dround,locatedrough- 610 THE JOURNALOF ARACHNOLOGY densely covered with short hairs. Claws with more than 15 teeth, slender, length twiceclaw width. Abdomen 10.5 mm long; whitish; cy- lindrical. Abdominal dorsal hairs 0.027 mm long (short); medium-sized, roughly straight, not compressed, blunt, tip enlarged; uniform- ly, thickly distributed. Male bulb (Fig. 15): T as long as DD; ex- ternal distal border straight; internal sloped backwards.DDbentabout45(cid:56)inlateralview; internal distal border not expanded. ES wider, moresclerotizedthanIS;IScontinuoustotip. DD tip (Figs. 19–21) straight in lateral view. C present, long; distal endonDDinternaltip; well-developed; located far from DD distal tip; proximal border continuously decreasing; distal border markedly sloped, upper tip not projected,pointed;externalsidehollowed.AC absent. LF present; distally not projected; poorly developed. L well-developed; external border not sclerotized, laterally slightly fold- ed, distal border divergent, continuous. AL Figures 13–18.—Dysdera ambulotenta. 13, Car- present, well-developed; proximal border in apace, dorsal; 14, Left chelicera, ventral; 15, Left posterior view fused with DH. P (Fig. 22) male bulb, external; 16, Vulva, dorsal; 17 Vulva, fused to T; markedly sloped on its proximal ventral; 18, Vulva, lateral. Scale bars in mm. part, perpendicular on distal; lateral length as long as or longer than T width; ridge present, lyatcenterofgroove;Bclosetobasallamina; perpendicular to T, not expanded; upper mar- MatmiddleofBandD.Legsorange.Lengths gin smooth; not distally projected; back mar- ofmaledescribedabove:fe16.6mm(allmea- gin not folded. surements in mm); pa1 4.3; ti1 6.3; me1 5.4; Paratype female: Figs. 16–18, 23, 24. All ta11.2;total23.8;fe26.3;pa24;ti26.2;me2 characters as in male except: Carapace 7.21 5.2; ta2 1.2; total 22.9; fe3 5.2; pa3 2.9; ti3 mm long;maximumwidth5.6mm;minimum 3.8; me3 4.5; ta3 1.3; total 17.7; fe4 6.3; pa4 width 3.64 mm. AME separation 1.16 mm. 3.4;ti45.3;me46;ta41.4;total22.4;relative Chelicerae 4 mm long; fang 3.22 mm. Legs length: 1 (cid:46) 2 (cid:46) 4 (cid:46) 3; palp: fe 3.5; pa 2; ti orange. Lengths of female described above: 2; ta 1.9; total 9.4. Spination: palp, leg1, leg2 fe1 6.6 mm (all measurements in mm); pa1 spineless. Fe3 dorsal spines in two rows: an- 4.3; ti1 6.2; me1 5.4; ta1 1.3; total 23.8; fe2 terior 3 (distal); posterior 1 (proximal); ti3 6.1; pa2 4.1; ti2 6.1; me2 5.4; ta2 1.3; total dorsal spines arranged in three bands: proxi- 23; fe3 5.2; pa3 3; ti3 3.9; me3 4.8; ta3 1.2; mal 1.2.1; medial-proximal 1.1-2.1; distal total 18.1; fe4 6.5; pa4 3.5; ti4 5.2; me4 6.3; 1.0.1; ti3 ventral spines arranged in three ta4 1.5; total 23; relative length 1 (cid:46) 2 (cid:53) 4(cid:46) bands: proximal 1-0.3-2.1; medial-proximal 3; palp: fe 4; pa 2; ti 1.9; ta 2.5; total 10.4. 1.3-2.1; distal 1.0.1;withtwoterminalspines. Spination: palp, leg1, leg2 spineless. Fe3 dor- Fe4 dorsal spines in two rows: anterior 4-2; sal spines in tworows:anterior2;posterior1; posterior 3-2; ti4 dorsal spines arranged in ti3 dorsal spines arranged in three bands: four bands: proximal 1-0.0.1-0; medial-prox- proximal 1.0.1; medial-proximal 1.0.1; distal imal 1.2.1; medial-distal 1.3-2.1; distal 1.0.1; 1.0.1; ti3 ventral spines arranged in three ti4 ventral spines arranged in four bands: bands: proximal 2-1.2.2-1; medial-proximal proximal 1.2-1.1; medial-proximal 1.2.1; me- 1.3-1.0-1; distal 1.0.1; with two terminal dial-distal1.2-1.1;distal1.1-0.1;withtwoter- spines.Fe4dorsalspinesintworows:anterior minal spines. Dorsal side of frontal legs with 1; posterior 2; ti4 dorsal spines arranged in a fine-textured piligerous granular surface; four bands: proximal 1-0.0.1-0; medial-prox- ventral side of palp smooth; posterior legs imal 1.2-0.1; medial-distal 1.3-1.1; distal ARNEDO& RIBERA—THEGENUS DYSDERA IN THE CANARYISLANDS 611 Figures19–24.—Dysdera ambulotenta, right malebulbandfemalespinnerets.19,DDfrontal;20,DD external; 21, DD internal; 22, P internal view; 23, Right ALS; 24, Right PLS. 1.0.0; ti4 ventral spines arranged in four ness, roughly straight, not compressed, blunt, bands: proximal 1.1.1; medial-proximal 1.1- tip enlarged; uniformly, thickly distributed. 2.1; medial-distal 0-1.1-0.1; distal 0-1.1-0.1; Vulva (Figs. 16–18) rectangle-like in dorsal with two terminal spines. view, frontally rounded; slightly wider than Abdomen 7 mm long. Abdominal dorsal long; DF wide. MFwell-developed;markedly hairs 0.036 mm long (short); medium thick- sclerotized along its extent. VA frontalregion 612 THE JOURNALOF ARACHNOLOGY completely sclerotized; posterior region scler- poorly developed but complete bulb lateral otized exceptforinternalarea.AVDabsent.S sheet (L) (Fig. 31), and both males and fe- attachment projected under VA; arms as long males have anterior medial eyes separated by as DA, slightly curved; tips dorsally project- less than ⅔ of its diameter from each other, ed; neck as wide as arms. TB usual shape. longer chelicera inner groove and cheliceral ALS(Figs.23–24)withoutPS;remainingpir- distaltooth(D)markedlylargerthanbasalone iform spigots no more external than MS, ar- (B) (Fig. 26). ranged in three rows; 18 piriform gland spig- Description.—Holotypemale:Figs.25–27, ots; PMS, PLS with 5–10 aciniform gland 31–34. Carapace (Fig. 25) 3.62 mm long; spigots. maximum width 2.51 mm; minimum width Intraspecific variation.—Male cephalo- 1.84 mm. Dark red, darkened at borders; thorax ranges in length from 7.00–7.35 mm, heavilywrinkled,foveate,withafine-textured female from 6.51–7.00 mm. Sometimes cara- granular surface. Frontal border roughly pacelateralmarginangledatmaximumwitdth round,from½–3⁄ carapacelength;anteriorlat- 5 point. AME reduction variable, from tiny eral borders parallel or silghtly divergent; bright spots to absent. Chelicera relative roundedatmaximumdorsalwidthpoint,back lengthfrom0.43–0.48.Daslargeasorslight- lateral borders rounded; back margin narrow, ly larger than B. One male from Los Roques straight;slightlysteppedinlateralview.AME with M distinctly closer to D. Ingeneral,che- diameter 0.21 mm; PLE 0.21 mm; PME 0.16 liceral teeth are large. Spination variability in mm; AME on edge of frontal border, separat- Table 2. ed one from another about ˙1 of diameter, Additional material examined.—TENERIFE: close to PLE; PME very close to each other, El Sauzal: Cueva de Labrada-Mechas, 13 March less than ¼ PME diameter from PLE.Labium 1982, 1(cid:63) subadult (J.L. Mart´ın, num. 2520 UL). trapezoid-shaped, base wider than distal part; Icod de los Vinos: Cueva de Felipe Revento´n, 17 as long as wide at base (triangle-like); semi- March 1984, some remains (J.J. Herna`ndez, num. circular groove at tip. Sternum brownish-red, 2534UL).CuevadelViento-Sobrado,10December darkened on borders; heavily wrinkled; uni- 1982,1juv.(J.L.Mart´ın,num.2517UL);2Novem- formly covered in slender black hairs. ber 1991, 1juv. (J.L. Mart´ın, num. 2518 UL). La Chelicerae (Fig. 26) 1.58 mm long, about 2⁄ Orotava: Cueva del Bucio, 27 November 1984, 5 some remains (J.L. Mart´ın & A. Machado, num. of carapace length in dorsal view; fang long, 2794 UL); 4 March 1985, 1juv. (J.L. Mart´ın & A. 1.35 mm; basal segment dorsal, ventral side Machado,num.2532UL).1April1991,1(cid:47)(Lucas completely covered with piligerous granula- & Rando, num. 2511 UL). Cueva de los Roques, tions. Chelicera inner groove long, about ½ 11 August 1986, 1(cid:63) (J.L. Mart´ın, num. 2512 UL); cheliceral length; armed with three teeth and 27October1991,?(onechelicera)(C.Ribera,num. lamina at base; D (cid:46) B (cid:53) M (B, M small); D 2568UL);25September1996,1(cid:47)(P.Orom´ı,num. trapezoid,locatednearsegmenttip;Bcloseto 3184 UB). basal lamina; M close to D. Legs orange. Distribution.—Tenerifean endemic. Exclu- Lengths of male described above: fe12.4mm sively known from lava tubes. It is the most (all measurements in mm); pa1 1.44;ti1 2.14; widespread of troglomorphic Dysdera. me12;ta10.51;total8.49;fe22.23;pa21.35; ti2 1.91; me2 1.98; ta2 0.51; total 7.98; fe3 Dysdera brevisetae Wunderlich 1991 1.79; pa3 1.07; ti3 1.26; me3 1.68; ta3 0.51; Figs. 25–36 total 6.31; fe4 2.19; pa4 1.12; ti4 1.82; me4 DysderabrevisetaeWunderlich1991:289–290,fig. 2; ta4 0.53; total 7.66; relative length: 1 (cid:46) 2 14–16 [(cid:63)]. Holotype male from Monte de las (cid:46)4(cid:46)3;palp:fe1.4;pa0.74;ti0.74;ta0.74; Mercedes, La Laguna, Tenerife, Canary Islands; total 3.62. Spination: spineless. Dorsalsideof inII,M.Kno¨selleg.;num.37166.StoredatSMF. frontallegscoveredwithhairs,lackingagran- Examined. -Wunderlich 1991: 284–287. ularsurface;ventralsideofpalpsmooth;long, Diagnosis.—Dysdera brevisetae is distin- spine-like hairs on posterior ti, fe. Clawswith guishedfromanyothermarkedlyfoveatespe- 10–14 teeth, length twice claw width. cies by its wider carapace frontal border and Abdomen 3.73 mm long; whitish; cylindri- spinelessfemoraandtibiae.Malesdifferfrom cal. Abdominal dorsal hairs 0.045 mm long; the morphologically similar D. macra by a medium thickness, roughly straight, not com- ARNEDO& RIBERA—THEGENUS DYSDERA IN THE CANARYISLANDS 613 pendicular to T in lateral view; lateral length from ⅓–2⁄ of T width; ridge present, perpen- 5 dicular to T, not expanded; upper margin smooth; not distally projected; back margin slightly folded towards internal side. Female: (from El Moquinal, La Orotava, Tenerife;num.2935,UB)Figs.28–30,32,33. All characters as in male except: Carapace 3.66 mm long; maximum width 2.65 mm; minimumwidth1.98mm.AMEdiameter0.23 mm; PLE 0.2 mm; PME 0.16 mm. Sternum dark red. Chelicerae 1.72 mm long; fang 1.4 mm; basal segment proximal dorsal, ventral side scantly covered with piligerous granula- tions. Legs orange. Lengths of female de- scribedabove:fe12.42mm(allmeasurements in mm); pa1 1.54; ti1 2.1;me1 2.05;ta10.53; total 8.64; fe2 2.25; pa2 1.44; ti2 1.91; me2 2; ta2 0.53; total 8.13; fe3 1.91; pa3 1.26; ti3 1.38; me3 1.68; ta3 0.51; total 6.56; fe4 2.28; pa4 1.26; ti4 1.91; me4 2.1; ta4 0.53; total 8.08; relative length 1 (cid:46) 2 (cid:46) 4 (cid:46) 3; palp: fe 1.4; pa 0.79; ti 0.56; ta 0.88; total 3.63. Abdomen 4.66 mm long; whitish; cylindri- cal. Abdominal dorsal hairs 0.16–0.18 mm long; thin, curved, compressed, pointed; uni- Figures 25–30.—Dysdera brevisetae. 25, Cara- formly, thickly distributed. Vulva (Figs. 28– pace, dorsal; 26, Left chelicera, ventral; 27, Right 30) arch-like in dorsal view, frontally round- male bulb, external; 28, Vulva, dorsal; 29, Vulva, ed; slightly wider than long; DF wide. MF ventral; 30, Vulva, lateral. Scale bars in mm. poorly developed. VA frontal region com- pletely sclerotized; posterior region sclero- tized at anterior area. AVD hardly visible. S attachment not projected under VA; arms as pressed,blunt,tipenlarged;uniformly,thickly longasDA,slightlycurved;tipsdorsallypro- distributed. jected; neck as wide as arms. TBusualshape. Malebulb (Fig.27):Tslightlysmallerthan ALS (Figs. 32–33) with PS; remaining piri- DD; external, internal distal border sloped formspigotsmoreexternalthanMS,arranged backwards. DD slightly bent in lateral view, in two rows; 7 (cid:49) 1 piriform gland spigots; clearlylessthan45(cid:56);internaldistalbordernot PMS,PLSwith5–10aciniformglandspigots. expanded. IS, ES equally developed; IS trun- Intraspecific variation.—Male cephalo- cated at DD middle part; ES bend markedly thorax ranges in length from 3.62–3.54 mm, sclerotized. DD tip (Figs. 31–33) straight in female from 3.40–3.66 mm. PLE-PME from lateral view. C present, short; distal end on 1⁄ –2⁄ diameter. Sternum ornamentation some- 5 5 DDinternal tip;well-developed;locatedclose what reduced. B may be larger than M. M to DD distal tip; proximal border sharply de- sometimes closer to B. creasing; distal border rounded, hardly Additional material examined.—TENERIFE: stepped, upper tip not projected, rounded; ex- La Laguna: Cocomoto, ? February 1989, 1(cid:63) (C. ternal side hollowed. AC present. LF absent. Deniz, num. 2680 UL). El Moquinal, under a bark L poorly developed; externalbordernotscler- ofEricascoparia,18October1994,1(cid:47)(P.Orom´ı, otized, laterally slightly folded; distal border num. 4001 UB). Monte de las Mercedes, 30 Janu- divergent, not continuous; upper sheet strong- ary 1989, 1(cid:63) (H. Enghoff, num. 2640 ZMK). Los ly folded at middle. AL present, very poorly Silos: Monte del Agua, 14 March 1987, 1juv. (H. developed; proximal border in posterior view Enghoff, num. 2669 ZMK); 1 February 1988, 1(cid:47) fused with DH. P (Fig. 34) fused to T; per- (J.J. Naranjo, num. 2598 UB); 3 March 1989, 1(cid:47)