PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Volume 53,No. 10.pp. 117-149, 63 figs. August22, 2002 A Revision ofthe African Spider Genus Raecius Simon, 1892 (Araneae, Zorocratidae) by Charles E. Griswold SchlingerCuratorofArachnida, DepartmentofEntomology. CaliforniaAcademyofSciences, Golden GatePark. SanFrancisco. California 94118 and Research ProfessorofBiology. SanFranciscoState University email: [email protected] ThetropicalAfricanspidergenusRaeciusSimon, 1892isdefinedandrevisedandakeyto thefivevalidspeciesisprovided.Raeciuscongoensis,R.jocqueiandR.scharffinewspecies aredescribed.MnesitheusvittatusSimon, 1907andM.zoropsidesStrand, 1916arenewju- niorsynonymsoiRaeciusasper(Thorell, 1899).Raeciusaculeatus Dahl, 1901 isexcluded from thegenusandisincertaesedisinZorocratidae. Raecius are cryptic spiders in theirhabits and history. They arerare in collections, usually col- lectedsinglyora fewatatime, andtheonly series incollectionshaveresultedfrom masscollecting throughpitfalltrappingorconcertedeffortstodigthem fromburrows.Raeciusoccurinthedisjunct, Afromontane forests (Griswold 1991a; White 1978, 1983) in tropical Africa (e.g., Mt. Cameroon, Bioko Island, the Eastern Arc mountains ofTanzania) but are notrestrictedto these habitats, occur- ringaswellinlowlandforestsandaridareas. Inthis,the 16thinaseriesofmonographsonspidersoc- curringintheAfromontaneregion(Griswold 1985, 1987b, 1987c, 1987d, 1990, 1991b, 1994, 1997a, 1997b, 1998a, 1998b,2000,2001;GriswoldandPlatnick 1987;GriswoldandLedford2001),Irevise the genus Raecius and discuss its taxonomic placement in the Zorocratidae. ThespidersplacedinRaeciushaveacomplexandconfiisedtaxonomichistory. Indeed,formuch ofthe 20th century synonyms ofthe same specieswereplaced indifferent families, the Tengellidae andZoropsidae.Koch(1875:32)describedthefirstRaeciusspeciesfromajuvenilefromEthiopia(in whatistodayEritrea). HecalleditAmaurobiuscrassipes,whichheplacedintheAgelenidae. Simon (1892:230)proposedthegenusRaeciusforKoch'sspeciesandplaceditintheZoropsidaealongwith thegeneraZoropsisandZorocrates.Thorell(1899:18)proposedMnesitheusasperforafemalespider from Mt. Cameroon that he placed in the Dictynidae. Simon (1903:975) listed Mnesitheus in the Dictynidae, andin 1907 describedanotherspeciesM. vittatus fromBioko Island(Simon 1907:227). In 1916 Strand described a male spider from Mt. Cameroon as Mnesitheus zoropsides (Strand 1916:140). Dahl (1901a, 1901b, 1913) treatedRaecius andMnesitheus in the Zorocratidae, but this placement was not followed by subsequent workers. Petrunkevitch (1928) listed Raecius in the ZoropsidaeandMnesitheus intheTengellidae, aplacement followedby Roewer(1954) andBonnet (1957, 1958).ItwasnotuntilLehtinen'scomprehensivestudyoftypespecimensofcribellatespiders thatthesynonymyofthesegenerawasrecognized(Lehtinen 1967:250,262).Lehtinensynonymized MnesitheuswithRaecius, andtransferredthe lattergenus intohisMiturgidae, Uliodoninae. I under- took the first quantitative phylogenetic analysis ofexemplars for taxa placed in the Tengellidae, 117 118 PROCEEDINGS OF THE CALIFORNIAACADEMYOF SCIENCES Volume 53,No. 10 Miturgidae, Zoropsidae and Lycosoidea (Griswold 1993). This analysis treatedtaxathathad one or more classical (i.e., grate-shaped tapetum) or newly discovered characters (e.g., male tibial crack, oval calamistrum, and/or interlocking lobes on the tegulum and subtegulum ofthe male pedipalpal bulb). SignificantresuhswerethatRaeciusformedacladealongwiththeotherformermiturgidgen- era Campostichomma, Zorocrates, Zorodictyma, and Udiiba. These taxa were not closely related to Miturgidae (represented in that study by Uliodon tarantulinus) but no family transfer was made. Within this clade the entire cribellum suggested that the Malagasy genus Zorodictyna and African Raecius are sister groups. Griswold, Coddington, Platnick, and Forster (1999) examined an even broaderarray ofaraneomorph taxa. Theirphylogenetic analysis suggested that the formermiturgid genera Raecius and Uduba (and by implication, Campostichomma, Zorocrates, and Zorodictyna) comprise the family Zorocratidae Dahl 1913. Synapomorphies ofZorocratidae are the male tibial crack,clumpedcribellarspigots(Figs. 19,21),andamalepedipalpaltibiaventroapicalprocess(Figs. 22,38). Comparison to Zorodictyna and moredistantlyrelatedzorocratidssuggestsseveralsynapo- morphies forRaecius. The long, narrow carapace (Fig. 1) contrastswith the oval carapaces ofother zorocratids.Theflattened,ribbon-likeembolus(Figs.37,51)andbluntembolictip(Figs.40,62)may be additional Raecius synapomorphies as other zorocratids have cylindrical or attenuate emboli (Griswold 1993, figs. 11, 13, 20, 34). Phylogenetic resolutionwithinRaecius is difficultbecauseR. congoensisisknownonlyfromfemalesandR.scharfjionlyfrommales,butatleasttheenlargedRTA may be a synapomorphy unitingR. asperandR.jocquei. The sistergroup relationshipoiRaecius and the MalagasyZorodictyna is ofparticularinterest, beingoneofatleastfivedocumentedcasesofMadagascar-tropicalAfricanmontane,vicariantsister grouprelationships(Griswold2000). FourotherparallelpatternsoccurwithinCyatholipidae(within Ulwembua and betweenAlaranea and Scharffia) and Phyxelididae (withinPhyxelida and between Ambohima and Kulalania). This corroborates the hypothesis in Griswold (1991a) suggesting that Madagascar is related to areas in tropical East and South Africa. This tropical African-Madagascar vicariance patternparallels that suggested for lemurs (Yoderet al. 1996) and tenrecs (Asher 1997). Recent molecular clock evidence suggests that the mammalian disjunctions may be Eocene in age (Yoderetal. 1996),andalandbridgealongtheDavieFractureZonehasbeenpostulatedasapathfor dispersalfromAfricatoMadagascaratthistime(McCall 1997).Thereplicatedbiogeographicpattern among spiders may also result from dispersal during the early Tertiary. METHODS Speciesdescriptionsrepresentasingleindividualofeachsexthatisnotedattheheadofeachde- scription. Thesectiononvariationrepresentsthevariabilityinseveral individualsofeachsexchosen torepresentthe full rangeofoverall size. Allmeasurementsareinmillimeters. The lengthandwidth ofthe cephalothorax and abdomen were measured from above and represent maximum values, chelicerae were excluded from cephalothorax length but included in measurements oftotal length; carapaceheightwasmeasuredfromthesideandrepresentsthemaximumheight;theocularareawas measured from above with the dimensions ofthe ocularquadrangle being measured from the outer marginoftheeyes;theposterioreyerowwasmeasuredfromaboveandtheanterioreyerowwasmea- suredfrom infront; eyeinterdistancesareabbreviated,e.g.,AME-AME=distancebetweenanterior medianeyes, ALE-PLE =distancebetween lateral eyes; clypeal heightrepresents thedistance from themedianmarginoftheclypeustothelowermarginoftheAME;chelicerallengthcomprisesthedis- tance from the median margin ofthe clypeus to the apex ofthe paturon (not including the fang); lengthsoflegsegmentsweremeasuredalongthedorsalmarginasinGriswold(1987a). Spinesarere- ported for the dorsal (d), prolateral (p), ventral (v), and retrolateral (r) surfaces of the legs and pedipalpiandarenotedfromtheproximaltodistalendsofeachsegment.Abbreviationsfortheocular GRISWOLD: AFRICANRAECIUSSPECIES 19 region(AME,ALE,PME,PLE,AER,PER)andlegsarestandardfortheAraneae.Alistofabbrevia- tionsusedtorefertospecialmorphologicalfeaturesispresentedinTable 1.Labeldataarequotedver- batim in the listings ofmaterial examined. Tracheaewereexaminedbyfirstdissectingoffthedorsalcuticleoftheabdomen,digestingaway nonchitinoustissuein 1 percentpotassiumhydroxide(KOH)eitherwarmedbeneathadesklampfor 72-120hoursorboiledfor5-1 minutes,andthenstainingthetracheaeandotherchitinousstructures in Chlorazol Black. Male pedipalpi were expandedby immersingthemovernight in aweak, watery solutionofKOHandtransferringthemtodistilledwaterwhereexpansioncontinued. Pedipalpiwere KOH transferredbackandforthbetween anddistilledwateruntilexpansionstopped.Femalegenitalia wereexcisedfromtheabdomen, cleanedinasolutionoftrypsinandwater,andillustrated. Pedipalpi andvulvaewereexaminedinalcohol andinlacticacid. Subsequently,toallowexaminationofinter- nalstructures,vulvaewerebleachedthroughbriefimmersionatroomtemperatureina5.25%sodium hypochlorite solution (CLOROX® householdbleach). Bleachingprovidedrapid, excellentclearing ofeven the most heavily sclerotized genitalia with no apparent distortion offeatures. Vulvae were then stained lightly with Chlorazol Black, temporarily mounted in lactic acid, and examinedwith a compoundmicroscope. Tenninology forparts ofthevulva follows Sierwald(1989). Priorto exami- nationwithaHitachiS-520ScanningElectronMicroscopeallstructureswerecleanedinanultrasonic cleaner: pedipalpi and female genitalia were critical point dried, other structures were air dried. Spinneretswerepreparedbyfirstcausingfullextensionbysqueezingtheabdomenwithforceps,lock- ingtheforcepsinpositionwithapaperclip,andtransferringthespiderandforcepstoabsoluteethanol for24hours.Thewholeabdomenwasthencleanedwithanultrasoniccleaner,criticalpointdried,and mountedonastubusingElmer'sGlue. Tapetainthe indirecteyeswereexaminedinlivingorfreshly deadspecimensas inGriswold(1993). SpecimenswereexaminedwithaWildM5A, Olympus SZH or Leica MZAPO stereomicroscope, and/or a Nikon compound microscope modified for 3 dimen- sional imagingby SL3D, Inc., and drawings were made with a mounted phototube (camera lucida). Table 1.Abbreviations formorphologicaltermsusedintextand figures. AC-aciniform gland spigot(s) ML - median lobe ofepigynum AER-anterioreye row MS - median sectorofepigynum ALE-anteriorlateral eyes N-nubbin ALS - anterior lateral spinneret OAL- length ofoculararea AME- anteriormedian eyes OQA- ocularquadrangle, anterior AN-anelli OQL -ocularquadrangle, length C -conductor OQP ocularquadrangle, posterior BS -base ofspennatheca P-petiole CD-copulatory duct PER-posterioreye row CY-cylindrical gland spigot(s) PI-pirifonTi gland spigot(s) DTA-dorsal apophysis oftibia PLE-posterior lateral eyes E-embolus PLS -posteriorlateral spinneret EF-epigynal fold PME -posteriormedian eyes FD-fertilization duct PMS -posteriormedian spinneret HS -head ofspennatheca PsFl-pseudoflagelliform gland spigot(s) ITC - inferiortarsal claw RTA-retrolateral apophysis oftibia LL- lateral lobes ofepigynum ST- subtegulum MA - median apophysis oftegulum STP - sclerotizedtegularprocess MAP-majorampullate gland spigot(s) T- tegulum mAP-minorampullate gland spigot(s) VTA-ventral apophysis oftibia 120 PROCEEDINGS OFTHECALIFORNIAACADEMY OF SCIENCES Volume 53,No. 10 Throughoutthetextreferencestofiguresfromotherpapersarenotedinlowercase(fig.),whereasref- erences to figures accompanyingthispaperare capitaHzed(Fig.). Natural History RaeciushavebeenobservedinthefieldinCameroon,onBiokoIsland, EquatorialGuineaandin Tanzania.Theyoccurinlowlandandmontaneforest(Fig. 26)andinmontanegrassland(Fig.27)and at leastR. asperoccurs inboth montane forestandmontane grassland. The type of7?. crassipeswas taken in a dry riverbed in northern Eritrea, a habitat much drier than that ofotherRaecius species. Theyareburrowingortubebuildingspidersthatlinetheirburrowswithsilkandprovidetheentrance withacollarofcribellate silk (Fig. 25). BurrowsoiR. asperfrom Mt. Cameroonwere6-8 cmdeep, and straight or slightly curved. In forest, burrows are typically made in rotting fallen logs, orsilken tubesortunnelsmaybemadeinmossontreetrunks. Ingrasslandtheburrowsare inturfandmaybe abundant. In burned grassland in January on Mt. Cameroon (Fig. 27), when theburrows were most conspicuous, 3 person-hoursofcollectingrevealedmorethan 70 individuals. Manyadultfemalesat this localityhadeggsacsoryoungintheburrow. Theseeggsacswerethicklywrappedwithsilkand coveredwithdirtorotherdebris. PopulationsfromforestandgrasslandonMt. Cameroondifferedin phenology. Two egg sacs collected in mist forest at 1425 m had only eggs within (37 and41 eggs), whereas sixteenegg sacs collectedduringthe sameweek from grasslandat 2050 mhadjuveniles or werealreadyempty.Ofthesesixteen,ninewereemptyandsevenhad23 to55 (average42)juveniles. A female from Mt. Kupe had an egg sac containing approximately 1 10 eggs. One to two egg sacs couldbefoundinafemale'stube,withonesacalwaysmoreadvancedphenologicallythantheother. A captive individual of R. scharffi captured prey with a grab and bite. Prey was carried in the cheliceraeandwasnotwrapped.CaptiveindividualsofbothR. asperandR.scharfficardedcribellate silkusinglegsIVbracedagainsteachotherandmovedsimultaneously.This"mobilelegIV"carding istypicalofentelegynes(Eberhard 1988,Griswoldetal. 1999).Cribellatesilkhasbothaxiallinesand reserve warp. Taxonomy Raecius Simon, 1892 RaeciusSimon, 1892:230(typespeciesAmaurobiuscrassipesL. Koch 1875:32,fromEritrea,inBMNH,exam- ined,described inAgelenidae). Petrunkevitch 1928:146. Roewer 1954:1283. Bonnet 1958:3846. Lehtinen 1967:250 (transferred to Miturgidae). Brignoli 1983:544. Platnick 1989:432; 1993:598; 1997:682; 2002. Griswoldetal. 1999:59 (transferredtoZorocratidae). MnesitlieiisThorell, 1899:18 (type speciesMnesit/ieiisasperThorell, fromCameroon, in RMS,examined, de- scribed in Dictynidae). Simon 1903:975. Petmnkevitch 1928:91. Roewer 1954:1379. Bonnet 1957:2968. SynonymizedbyLehtinen 1967:250. — Diagnosis. Zorocratid spiders with entire cribellum (Figs. 18-19) and oval calamistrum (Fig. 8)thatdifferfromZorodictynainhavingfeatheryhairs(Figs. 6,9,43)andlackingadorsobasal projection ofthe cymbium, and that differ from all other zorocratids in the flattened, ribbon-like embolus(Figs.37,51),bluntembolictip(Figs.40,62),andelongatecarapacewithlength 1.47to 1.90 times width (Fig. 1)—. Description. Medium sized to large spiders, total length 4.68-15.73; sexual dimorphism slight, male with relatively longer legs. Carapace elongate-oval in dorsal view (Figs. 1, 24), length 1.47to 1.90 timeswidth, domed(Fig. 2), height0.38 to0.74timeswidth; fovea linear,deep. Ocular area broad, PER width 2.34^.87 times OA length; ocularquadrangle rectangularto trapezoidal, in most specimens widest behind, OQP/OQA 0.93-1.36; eight eyes in 2 nearly straight rows, indirect GRISWOLD: AFRICANRAECIUSSPECIES 121 AME eyes with canoe-shaped tapetum. Clypeus low, height 0.84-2.06 times diameter. Chelicerae stoutwith largeboss (Fig. 2), length 6.31-12.00 timesclypeal height, pro- andretromarginsoffang furrowinmostspecimenswith3teeth. Sternumlength 1.31-1.88timeswidth;labiumlong,withbasal notch, length 1.19-1.67 times width; pedipalpal coxal endites nearly parallel (Fig. 3), length 1.94-3.00timeswidth,serrulateethinsinglerowalongoutermargin.Legformula 1423or4123,ratio offemurIlength/carapacewidth 1.32-2.84inmale,0.97-1.87infemale;integumentfinelywrinkled, withplumoseandfeatherysetae(sensuLehtinen 1975)onlegsandbody(Figs. 6,9,43^4);maleleg tibiaewithbasal crack(Griswold 1993, figs. 3,4); trochanterswithbroad, shallownotch. Spination, male basic pattern, comprises: pedipalpus: femur dO-1-1-1, rO-0-0-1; leg I: femur d1-0-0, tibia V2-2-2-2, pO-1-1-0, rO-1-1-0, metatarsus v2-2-l, pl-1-1, rl-1-1; leg II: femur d1-0-0, tibia v2-2-2, pO-1-1-0, rO-1-0, metatarsus v2-2-l, pi-1-2, rO-1-0; leg 111: femur: d1-0-1, tibia: vl-1-2, pi-1-0, dO-1-0, rl-1-0, metatarsus vl-1-2, pl-1-2, dO-1-0, rl-1-2; leg IV: femur: dl-0-0, tibia vl-2-2, rO-1-1-0, metatarsus: vl-1-2, pO-0-2, rO-0-2. Raeciiisjocqiiei has more dorsal spines on femur and tibiaIandR.scharffihasfewerlateralspinesontibiaandmetatarsusIandfewerventralspinesontibia II.Femalebasicspinationpattern:pedipalpus: femurd1-1-1,tibiapO-1-0,tarsusp1-1,vO-1; legI: fe- mur: d1-0-1,tibiav2-2-2-2,metatarsuspO-0-2,v2-2-2,rO-0-2; legII: femur: dl-0-1,patellapi,tibia pl-0-1, vl-1-2, metatarsus pi-0-2, v2-2-l, rO-0-2; leg III: femur dl-0-0, patella pi, tibia dO-1-0, pl-O-l, vl-1-2, rl-0-1, metatarsus p 0-1-1-2, v2-2-2, rl-1-2; leg IV: femur dl-0-0, tibia pl-0-0, vl-1-2,metatarsuspO-0-2,vl-1-2,rO-0-2.Femoraldorsal,patellarprolateral,andlateralspinesofthe tibiaeandmetatarsimaybefewerormorethanthispattern.Calamistrumrectangular-oval,arisingba- sallyandextendingfor2/3 length segment, attenuatedistally(Figs. 6, 8). Trichobothrialpattern: ab- sent from leg femora and patellae, tibiae with adorsobasal group, few prolaterals, and a retrolateral rowextendingtoapex,metatarsiwithadorsal,irregularrow,thisrowrestrictedtoapicalhalfof1Vth, tarsi with dorsobasal row that divides into 2-3 rows apically, pedipalpal tibiae with pro- and retrolateralrows, inmalesextendingdorsadandretrolateradofRTA, apparentlyabsentfromtarsi of mostspecimens,althoughafemaleofR.jocqueihasasingledorsaltrichobothriumononepedipalpal tarsus; trichobothrial base with transversely-ridged hood (Figs. 4, 7). Tarsal organ median to basal, capsulate,orificesimple,ovaltoround(Figs.5,7).MalescopulaebeneathtarsiI-IV,femalescopulae beneathalltarsiandatleastmetatarsiIandII(Fig. 1),scopulaeextenddistadoftarsalapices,mayob- scure claws; dorsal scopulate patch on cymbium absent (Fig. 53). Preening combs absent. Superior tarsalclawspectinate,thoseoflegs 1 andIIwith 8-12 teeth,thoseoflegsIII-IVwith3-7teeth, infe- rior tarsal claws (ITC) simple, large or reduced to nubbins or absent; female pedipalpus claw pectinate,with3-10teeth;clawtuftsabsent.Malepedipalpaltibiawithretroapical(RTA)andventral (VTA) apophyses (Figs. 22, 38), dorsal apophysis (DTA) present {R. scharffi: Fig. 55) or absent. Pedipalpal tarsus (Figs. 22, 23) with large, triangular petiole (P) attached to alveolus; subtegulum (ST)cup-shaped,with4-5 anelli (AN),prolateralmarginwith lobethatarticulateswithcorrespond- ing lobe on tegulum in unexpanded bulb (Fig. 23); tegulum (T) U-shaped, simple; firmly attached embolus (E) arising on prolateral side oftegulum, embolus flattened and ribbon-like, may have grooveorfoldalongoutermargin(Fig. 51);medianapophysis(MA)flexiblyattached;conductor(C) hyaline, originafing from retroapex ofT, cup or fan-shaped (Figs. 29, 45, 55); tegulum with no (R. scharffi),one{R.asper)ortwo(R.jocquei)sclerotizedprocesses(STP);fundusinsubtegulum,reser- voirandejaculatory duct simple, without loops orswitchbacks, spiraling aroundoutermargin ofT. Abdomenoval,withoutscuta(Figs. 1-2).Respiratorysystemcomprisingtwoanteriorbooklungsand a single, small spiraclejust anteriadofcribellum leading into foursimpletracheal tubes confinedto abdomen (examined in juvenile R. asper from Mann's Spring). Epigynum (Fig. 41) divided by epigynal folds (EF) into median sector (MS) and lateral lobes (LL); median sector with transverse lobe(ML),variouslyshaped;LLtoothpresentorabsent;vulva(Figs.42,58;Griswold 1993,figs.29, 30)withcopulatoryducts(CD)anteromedian, short, slender,enteringspermathecalhead(HS) later- allyorventrolaterally;HSsmall,hemispherical;baseofspermatheca(BS)simple,short, fertilizafion 122 PROCEEDINGS OFTHE CALIFORNIAACADEMY OF SCIENCES Volume 53, No. 10 duct (FD) posterior. Cribellum wide, short, width greater than 3 times length, spinning field entire (Figs. 18-19), cribellar spigots arranged in longitudinal lineargroups (Figs. 19, 21). Six spinnerets (Fig. 18), anteriors (ALS) and posterior laterals (PLS) two-segmented, posterior medians (PMS) one-segmented; ALS conical, PMS and PLS cylindrical, distal segment ofPLS domed, short, less than 1/3 lengthbasalsegment.Female(examinedinR. asperandR.Jocqiiei)withALS(Fig. 1 1)hav- ingtwo largemajorampullategland spigots (MAP) on mesal margin andoval field ofmorethan 30 piriformgland spigots (PI); PMS (Fig. 12)withone large minorampullategland spigot(mAP)near anteriormarginandlargecentralnubbin, several acinifomiglandspigots(AC),with4-5 largecylin- drical gland spigots (CY) with long conical bases and long slender shafts on posterior surface of spinneretextendingtobase;PLS(Figs. 13,20)with4-6largeCYonmarginofACspinningfield,ap- parent apical pseudoflagellifomi gland spigot (PsFl) present (/?. asper. Fig. 20) ornot apparent {R. Jocqiiei: Fig. 13). Male (examined in R.Jocqiiei: Figs. 14-17) lacking CY, other spigots reduced in number:onlyoneMAPonALSaccompaniedbyanubbin,andapicalnubbinonPLS(Fig. 17). Sucha PLSapicalnubbinmayreplacethePsFlspigotinthemale,suggestingthataPsFlmaybepresent(but undistinguished) in—the female ofR.Jocqiiei. Male epiandrous spigots absent (Fig. 44). Composition. —Five species. Distribution. Tropical Africa from Cote d'lvoire on the west across the Congo basin to EritreaandTanzania inthe east(Fig. 63). Keyto Speciesof Raecius 1 Dorsalabdominalmarkingsuniform(Koch 1875,pi. 3,fig.4); legfomiula4123;occumnginsemiaridEritrea crassipes(L. Koch) Dorsalabdominalmarkingswithatleastafaintlightlongitudinalband(Fig. 1); legfomiula 1423;occurringinmoist equatorialAfrica 2 2(1)Males 3 Females 5 3(2)PedipalpalbulbwithSTP;medianapophysisnotproducedintodistalhook; DTAabsent; RTAlarge,lengthgreater than0.5timeslengthoftibia 4 PedipalpalbulblackingSTP,medianapophysisproducedintodistalhook (Figs. 54-56);pedipalpaltibiawithDTA; RTAsmall,attenuateatapex, lessthan0.4timeslengthoftibia(Fig. 55) sx/fi://;///newspecies 4(3)PedipalpalbulbhavingmedianapophysisexpandedandbilobateatapexandSTPcurvedandnarrow(Fig.47), pedipalpaltibiawithRTA expandedandbilobateatapex,lengthgreaterthan0.85timeslengthoftibia(Fig.46) jocqiieinewspecies Pedipalpalbulbhavingmedianapophysiscup-shapedandSTPbroad(Fig.40),pedipalpaltibiawith RTAtruncateat apex, length0.5-0.67times lengthoftibia(Fig.29) (75/;e/-(Thorell) 5(2)Epigynalmedianlobeconcaveposteriorly,lengthlessthan 1.0timeswidth 6 Epigynalmedianlobeconvexposteriorly,narrow, length 1.2-1.7timeswidth(Figs. 31,33,35). . . . a5/j£'r(Thorell) 6(5)Epigynalmedianlobedeeplyconcaveposteriorly,U-shaped,length0.80-0.85 timeswidth, lateral lobeswithteeth (Fig.48) jocqiieinewspecies Epigynalmedian lobeweaklyconcaveposteriorly,verybroad, lengthlessthan0.55timeswidth,laterallobeswithout teeth(Fig.57) cowgoenswnewspecies Raeciusasper(Thorell, 1899) Figures 1-3, 6, 8, 18, 20, 24-42, 63 MnesitheiisasperIhoxQW, 1899:18 (femaletype specimen from Mapanja, Cameroon, 1884-1885, Valdauand Knutson, in RMS, examined). Simon 1903:975. Roewer 1954:1379. Bonnet 1957:2968. Raeciusasper, Lehtinen, 1967:262. Platnick 1997:682; 2002. , GRISWOLD: AFRICANRAECIUSSPECIES 23 1 Mnesitheus vittatiis Simon, 1907:227 (threejuvenile syntypes from Moka (sic), Fernando Poo [Bioko Island, Equatorial Guinea], in MNHN, AR817, examined). Roewer 1954: 1379. Bonnet 1957: 2968. NEW SYN- ONYMY. Raecius vittatiis, Lehtinen 1967:262. Platnick2002. Mnesitheuszoropsides Strand, 1916:140 (male type specimen from Buea, Cameroon, in NHMV, examined). Roewer 1954:1379. Bonnet 1957:2968. NEW SYNONYMY. Raecius zoropsides, Lehtinen, 1967:262. Griswold 1993:7. Platnick 1997:682; 2002. Griswold and Ubick 1999:31. — Notes. Males, females andjuveniles have been collected in association on Bioko Island. EquatorialGuinea. MalescorrespondindetailtoR.zoropsidesStrand,juvenilestoR. vittatiisSimon, and females to R. asperThorell. The lattername is the seniorsynonyin. This specieswas discussed andillustratedinGriswold(1993:14, figs. 25-27). DatafromtheholotypeofMnesitheuszoropsides providedthemaleRaeciuscharactersinthephylogeneticmatrix.Thisspeciesalsoprovidedexemplar data forRaecius—in Griswold et al. (1999). Diagnosis. Malewithpedipalpalbulbhavingmedianapophysiscup-shapedandSTPpromi- nent and broad (Figs. 30, 37-40), pedipalpal tibia with RTA truncate at apex, 0.5-0.67 times the length ofthe tibia (Fig. 29). Female with epigynal median lobe convex posteriorly, narrow, length 1.2-1.7 times width (Figs. 3—1, 33, 35, 41 ). Male (Moca, Bioko). Total length 10.00. Carapace, chelicerae, legs and pedipalpi yel- low-brown, labium and pedipalpal coxae yellow-brown, lighter at tips, sternum, leg coxae and tro- chantersyellow-white; carapacewith sidesofcaput andparsthoracicadusky andcoveredwith dark setae, center ofcarapace bare and pale; caput dark at anterolateral margins above cheliceral bases; dark pigment surrounding each eye and extending between ALE and PLE; retrolateral process of pedipalpal tibia (RTA) darkbrown to black; dorsum ofabdomen dark gray with narrowcentral yel- low-gray longitudinal band, simple anteriorlyand scallopedandbroken into spotsposteriorly; sides and venter yellow-gray mottled with dark spots; spinnerets yellow-gray. Carapace 5.33 long, 3.24 AME wide, 1.81 high;clypeus0.24high,height 1.04times diameter;oculararea0.57long, 1.47wide, OAL 1.30 times OQL; ratio ofeyes AME:ALE:PME:PLE, 1.14:1.07:1.14:1.00, diameter ofPME 0.23; AME interdistance0.5 times AME diameter, AME-ALE 0.44times AME; PME interdistance 0.56timesPME, PME-PLE 1.44times PME; ALE-PLE0.57 timesPLE. Chelicerae0.98 long. Ster- num2.75 long, 1.69wide;labium 1.20long,0.78wide;pedipalpalcoxae 1.73 long,0.78wide. Femur Ilength 1.49timescarapacewidth. Spination:basicpattern.ITCIandIIabsent.IIIandIVreducedto nubbins. Legmeasurements(Femur+Patella+Tibia+Metatarsus+Tarsus=[Total]): I:4.83 +2.05 +4.83+4.15 +2.18-[18.04]; 11:3.85+ 1.80+3.37+3.02+2.10=[14.14]; 111:3.07+ 1.46+ 1.80+ 2.44+ 1.61 =[10.38]; IV:4.39+ 1.80+3.66+3.71 +2.00=[15.56];pedipalpus:2.10+0.93+0.88+ [absent]+ 1.80=[5.71].PedipalpaltibiawithRTAcurved,widenedattruncate,serratedapex,length 0.5-0.67timesthatoftibia,VTAcylindrical,taperingtoapex,pro-andretroventralmarginoftibiaat tibia-tarsusjointproducedintosmall processes. Pedipalpalbulbwithmedianapophysiscup-shaped, STPprominentandbroad,narrowedapically,embolusshortandribbon-like,andconductorsmalland narrow (Figs. 28-30, 37^0; Griswold 1993, figs. 25-27). Variation (N = 5). Total length 8.71-12.14; ratios of carapace length/width 1.59-1.65, height/width0.44-0.56,ratiosofPER/OQP2.37-2.90,PER/OAL2.59-3.10,OQP/OQA 1.06-1.21 ratiosofclypealheight/diameterAME0.88-1.50, cheliceral length/clypealheight7.68-10.73, ratio oflengthfemurI/carapacewidth 1.34-1.40.Markingsofabdomenrangefromafaintanteriorlongitu- dinallightmarkandunifonnlypalesidestoaboldlongitudinallightmarkbreakingintochevronspos- teriorlywith sides mottled. ITC I and II ma—ybe reduced to nubbins orabsent. Female(Mann'sSpring,Cameroon). Total length 12.27. Carapacered-brown,darkestante- riorly andon sides ofpars cephalica, palest aroundthoracic fovea, sides covered sparselywith dark scales; chelicerae dark brown except boss red; labium and pedipalpal coxae red-brown, lighter at 124 PROCEEDINGSOFTHE CALIFORNIAACADEMYOF SCIENCES Volume 53,No. 10 bases andtips; sternumandlegcoxaeyellow-brown, legs I and II shadingto duskyred-brownfrom basalthirdoftibiaetoapexoftarsi;abdomenmarkedasinmale(Figs. 1-3).Carapace6.29long,3.90 AME wide,2.19high;clypeus0.35high,height 1.75times diameter;oculararea0.62long, 1.88wide, OAL 1.32 times OQL; ratio ofeyes AME:ALE:PME:PLE, 1.00:1.28:1.21:1.07, diameter ofPME 0.24; AME-AME interdistance0.86timesAME diameter, AME-ALE 1.07 times AME; PME-PME interdistance 0.70 times PME, PME-PLE 1.82 times PME; ALE-PLE 0.93 times PLE. Chelicerae 2.94 long. Sternum 2.82 long, 1.92 wide; labium 1.41 long, 0.98 wide; pedipalpal coxae 2.16 long, 0.90wide.Femur1length 1.05timescarapacewidth. Spination:basicpattern. ITCIandIIreducedto nubbins.IIIsmall,IVlarge.Legmeasurements(Femur+Patella+Tibia+Metatarsus+Tarsus=[To- tal]): I: 4.10+2.05 +3.51 +2.88+2.20= [14.74]; II: 3.80+1.85 +2.73 +2.29+ 1.95 = [12.62]; III: 2.93+1.61 + 1.46+ 1.85+ 1.32= [9.17]; IV: 4.00+2.00+2.93 +2.88+ 1.56=[13.37];pedipalpus: 2.10+1.07+ 1.12+[absent]+ 1.61 =[5.90].Epigynumwithmedianlobeconvexposteriorly,narrow, length 1.3 times width, lateral lobes contiguous posteriorly, without teeth (Fig. 31). Vulva with spennathecal head oval (Fig. 32). Variation (N=ll). Total length 10.85-14.42; ratios of carapace length/width 1.47-1.53, height/width0.53-0.73,ratiosofPER/OQP2.56-3.64,PER/OAL2.89-4.87,OQP/OQA 1.08-1.36, AME ratiosofclypealheight/diameter 1.48-2.06,chelicerallengtli/clypealheight7.32-9.41,ratioof lengthfemurI/carapacewidth0.97-1.14.Markingsofabdomenrangefromafaintanteriorlongitudi- nal lightmarkanduniformsidestoaboldlongitudinal lightmarkbreakingintochevronsposteriorly with sides mottled (Figs. 1, 2). Patellar spines present or absent. Scopulae restricted to tarsus and metatarsus I andtarsus II orextendingonto tibia I andmetatarsus II. Epigynawith lateral lobeteeth present(Figs. 33, 35)orabsent(Fig. 31),medianlobelength 1.2-1.7timeswidth,spennathecalhead round (Fig. 34) to ov—al (Figs. 32, 36) Distribution. South—western Cameroon and Bioko Island in the GulfofGuinea (Fig. 63). MaterialExamined. CAMEROON: Southwest: FakoDivision: Mt. Cameroon: mistfor- est on south side, elev. 1425 m, 4°06'28"N, 9°07'10"E, 26-28 January 1992, C. Griswold, J. Coddington, G. Homiiga, S. Larcher (3? CAS, 2? USNM); grassland near Mann's Spring, elev. 2050 m,4°08'N, 9°07'E, 21-25 January 1992, C. Griswold,J. Coddington, G. Hormiga, S. Larcher (17? CAS, 54? USNM); Buea, type of Muesitheus zoropsides Strand (IcTNHMV); Mapanja, 1884-1885, KnutsonandValdau,typeoiMnesitheusasperThorell(1 ? RMS). MemeDivision: Mt. Kupe above Nyassosso, forest, elev. 800-1200 m, 4°50'N, 9°4rE, 16-19 February 1992, C. Griswold,N. Scharff,C.Wanzie,S.Larcher,P.Masongo(1? CAS).Northwest:Bali,BaflichuMbu, Shum Laka, elev. 1600 m, pitfall, December 1991-February 1992, H. Doutrelepont (1 ? MRAC). EQUATORIAL GUINEA: Bioko: Moca, 3°22'0"N, 8°39'57"E, elev. ca. 1400 m, 6-10 October 1998,K.DabneyandD.Ubick(3o"1 ? CAS), 1500 m,"ex:silk-linedburrowontreefern,"3-10Octo- ber 1998, D. Ubick(1o^ CAS); PicoBasile, 3°37'38"N, 8°48'15"E, elev. ca. 1750 m, inrottinglogs, 27-29 September 1998, K. Dabney and D. Ubick (lo"2? CAS); Moka (sic), types ofMnesitheiis MNHN vittatus Simon (3juveniles, AR817). Raeciuscongoensis, new species Figures 7, 57, 58, 63 — Type. Femaleholotype fromCongo, Lulimbi, embouchurede lariv. Ishashadans le lac Ed- ward(Sud-Est)baledeKyangiro,"danslesolpreleve,"collectedJuly-August 1976,M. Lejeune,de- positedin MRAC (168.827). Paratypefemale, samedataanddeposition except"terreausous litiere, (MRAC galerie forestiere" — 168.862). Etymology. Refers to occurrence in Congo. GRJSWOLD: AFRICANRAECIUSSPECIES 125 — Note. This species was discussed and illustrated in Griswold (1993:14, figs. 28-30). Data fromthespeciesprovidedthefemaleRaeciuscharactersinthephylogeneticmatrix.Thisspeciesalso provided exempla—rdata forRaecius in Griswold et al. (1999). Diagnosis. Female with epigynal median lobe weakly concave posteriorly, very broad, length less t—han 0.55 times width, lateral lobes without teeth (Fig. 57). Maleunknown. Male. Unknown. — Female (Holotype). Total length 15.73. Carapace orange-brown dorsally, shadingto dusky red-brown laterally and on pars cephalica; sides sparsely coveredwith dark setaethatoverlayblack reticulate patterns that radiate from thoracic fovea; thoracic fovea dusky; each eye narrowly sur- rounded by black pigment; chelicerae dark red-brown except boss orange; labium and pedipalpal coxae orange-brown with dusky maculations; sternum dusky gray-brown; leg coxae yellow-brown with dusky markings, these mostpronouncedon I and II, reticulatebasallybecoming soliddistally; legsandpedipalpi yellow-brown, legs I andIIwithdarkgray forminglongitudinal lateral anddorsal mottlingonfemoraandtibiae;abdomenyellow-gray,faded,markingsappeartohavecomprisednar- rowdorsal longitudinal lightbandand lightanddarkspots laterally. Carapace 7.52 long,4.38 wide, AME OAL 2.95high;clypeus0.39high,height2.05times diameter;oculararea0.64long,2.05wide, 1.36 times OQL; ratio ofeyes AME:ALE:PME:PLE, 1.00:1.18:1.04:1.33, diameter ofPME 0.20; AME-AME interdistance 1.11 times AME diameter, AME-ALE 1.26 times AME; PME-PME interdistance 1.57 times PME, PME-PLE 2.71 times PME; ALE-PLE equals PLE. Chelicerae 3.10 long. Sternum 3.84 long, 2.04 wide; labium 1.53 long, 1.06 wide; pedipalpal coxae 2.39 long, 0.98 wide.FemurIlength 1.10timescarapacewidth. Spination:typical. ITC1andIIabsent.IIIverysmall, IVsmall.Legmeasurements(Femur+Patella+Tibia+Metatarsus+Tarsus=[Total]): 1:4.83 +2.49 +3.61 +2.83+2.34=[16.10]; II:4.05+2.20+2.78+2.44+2.00= [13.47]; III: 3.27+ 1.80+ 1.51 + 2.15+1.56=[10.29]; IV:4.59+2.29+3.32+3.41 +2.00= [15.61];pedipalpus: 2.39+ 1.22+ 1.15+ [absent]+2.07=[6.83]. Epigynumwithmedianlobeweaklyconcaveposteriorly,verybroad, length lessthan0.55timeswidth,laterallobescontiguousposteriorly,withoutteeth(Fig.57;Griswold 1993, fig.28).Vulvawithcopulatoryductbroadwithlateraldiverticulum,spemiathecalheadnarrowwitha few pores, spemiathecal base short, extending mesad to fertilization duct (Fig. 58; Griswold 1993, figs. 29, 30). Variation (N = 2). Total length 12.53-15.73; ratios of carapace length/width 1.51-1.72, height/width0.69-0.71,PER/0AL3.22-3.45,OQP/OQA 1.10-1.29;ratiosofclypealheight/diame- terAME 1.86-2.05,chelicerallength/clypealheight7.95-8.33,ratiooflengthfemurI/carapacewidth 1.09-1.10;numberofventroapicalspinesonmetatarsiIandIIrangingfrom2to5;medianandproxi- maldorsalfemoralspinespresentorabsent;scopulaeweaktostrongbeneathmetatarsusII,extending from Vi to entire leng—th ofsegment. Distribution. South—eastern Congo basin (Fig. 63). Material Examined. Only the holotype andparatype. Raecius crassipes (L. Koch, 1875) Figure 63 Amawobius crassipes L. Koch, 1875:32, pi. 3, f 4 (holotype juvenile from "Ethiopia" "in dem trocken StrombettedesAnseba" [fromthedrystreambedoftheAnsebaRiver,inwhatistodaynorthern Eritrea], in BMNH, examined.) Raeciuscrassipes, Simon, 1892:230. Roewer 1954:1283. Bonnet 1958:3847. Platnick2002. — Note. Thejuvenile femaletype specimen is faded, shriveledanddisarticulated, butall parts arepresent:Therelativelylongcarapaceandentirecribellumwithlinearbunchesofcribellarspigots leavenodoubtthatit iscongeneric withtheotherspecieshereplacedinRaecius. Becausethe speci- 26 PROCEEDINGS OF THE CALIFORNIA ACADEMYOF SCIENCES 1 Volume 53,No. 10 menisimmature,diagnosisisdifficult,butbecausethetypelocalityisgeographicallydistantandeco- logically differen—t (i.e., arid) from those ofotherRaecius species, I considerR. crassipes distinct. Diagnosis. The juvenile specimen lacks diagnostic genitalia and is faded. Koch's figure (1875,pi. 3,fig.4)depictsaspiderwithdarkcarapaceandpaleparscephalicaanduniformlypaleab- domen.0\hQrRaeciusspecieshavedarkabdomenswithatleastafaintlongitudinalmedianband.The 4123 leg fomiula differs from—the 1423 ofotherRaecius. JuvenileTypeFemale. Totallengthapproximately8.0 mm.Specimenfaded,Koch'sfigure (1875,pi.3,fig.4)depictsadarkcarapaceandpaleparscephalicaandunifAonMnElypaleabdomen.Cara- pace4.80long,3.15wide, 1.55high;clypeus0.25high,height2.27times diameter;oculararea 0.45 long, 1.35wide,OAL 1.28timesOQL;radoofeyesAME:ALE:PME:PLE, 1.00:1.72:1.54:1.72, diameterofPME 0.17; AME-AME interdistance 1.18 times AME diameter, AME-ALE 1.45 times AME; PME-PME interdistance 0.68 times PME, PME-PLE 1.88 times PME; ALE-PLE 0.26times PLE. Chelicerae 2.35 long. Sternum 2.25 long, 1.40 wide, apex forming blunt angle; labium 1.00 long, 0.80 wide; pedipalpal coxae 1.70 long, 0.75 wide. Femur I length 1.06 times carapace width. SpinationtypicalforfemaleRaeciusexceptfemoralackingspines. Scopulae:dense,extendingbelow apical 1/2 ofmetatarsi I andII,restrictedtotarsi III andIV. ITCpresentonall legs, ITC III smallest. Legmeasurements(Femur+Patella+Tibia+Metatarsus+Tarsus= [Total]): I: 3.35+ 1.70+2.50+ 1.70+ 1.65=[10.90];II:2.85+1.50+2.00+ 1.60+ 1.60=[9.55];III:2.20+ 1.40+1.25+ 1.65+1.25 = [7.75]; IV: 3.25 + 1.65 +2.65 +2.50+ 1.60= [11.65];pedipalpus: 1.50+0.85 +0.75 + [absent] + = 1.40 [4.50]. — Distribution. Knownonly fromthetype locality, inanaridregionofnorthernEritrea(Fig. 63). — Material Examined. Only the type. Raeciusjocquei, new species Figures 10-17, 19, 21-23, 43-53, 63 — Type. Male holotype from Cote d'lvoire, Appouesso, foret classee de la Bossematie, pitfall traps, 31 October 1994,R. JocqueandN. Seabe,depositedinMRAC (202.373). Paratypes, samelo- cality, 180/4? MRA—C, Icfl? CAS. Etymology. Named in honor ofRudy Jocque, who collected these specimens as well as manyother—rare and interestingAfricanArachnida. Note. Thi—s species provided partial exemplardata forRaecius in Griswold et al. (1999). Diagnosis. Male with pedipalpal bulb (Figs. 45^7, 50-53) having median apophysis ex- pandedandbilobateatapexandSTPprominent,curvedandnarrow,pedipalpaltibiawithRTAlarge, expanded andbilobate at apex, greaterthan 0.85 times the length ofthetibia. Femalewith epigynal median lobe deeply con—cave posteriorly, U-shaped, length 0.80-0.85 times width (Fig. 48). Male (Holotype). Total length 6.76. Spider yellow-gray, unmarked except carapace dusky laterallyoverlainsparselywithdarksetae,leavingmedianpaleband,broadonparscephalicaandnar- row on pars thoracica; each eye surrounded with black pigment; dorsal process ofpedipalpal tibia darkbrown; dorsumofcymbiumdusky; abdomendarkgraydorsallyexcept fornarrowmedianpale band, sides mottled. Carapace4.00 long, 2.63 wide, 1.27 high; clypeus 0.18 high, height 1.13 dmes AME diameter; ocular area 0.40 long, 0.97 wide, OAL 1.21 times OQL; ratio of eyes AME:ALE:PME:PLE, 1.10:1.30:1.20:1.00, diameter ofPME 0.17; AME interdistance 0.45 times AME diameter, AME-ALE 0.27 times AME; PME interdistance 0.33 times PME, PME-PLE 1.08 times PME; ALE-PLE0.50timesPLE. Chelicerae 1.39 long. Sternum 1.98 long, 1.35 wide; labium 0.73 long, 0.55 wide; pedipalpal coxae 1.25 long, 0.42 wide. Femur 1 length 1.33 times carapace width. Spination: basicpatternexceptfemoraI-IVd1-1-1,tibiaId1-1-1,metatarsiIIIandIVp1-1-2. ITC I small, slender, II small, stout. 111 and IV large. Legmeasurements (Femur+ Patella + Tibia +