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The Neotropical orb-weaver genera Edricus and Wagneriana (Araneae: Araneidae) PDF

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Preview The Neotropical orb-weaver genera Edricus and Wagneriana (Araneae: Araneidae)

THE NEOTROPICAL ORB-WEAVER GENERA EDRICUS AND WAGNERIANA (ARANEAE: ARANEIDAE) HERBERTW. LEVI1 Abstract. Edricus and Wagneriana species are females were described under different Neotropical, nocturnal orb weavers. Edricus species names. A future goal of these studies is the arefoundfromMexicoandPanamatoEcuador. Most aWnadgnseorutihaenaastesrpnecSieosutahreAmfeoruincda.inBetchaeusAemaofzothnesairme-a earnaa.lyTshiissofcainntbeerrdeloanteioonnshliypsbyameoxanmgingienng- ilarity of the structure of male and female genitalia and comparing all species of each genus and the presence of a paramedian apophysis in the and only after most of the genera are re- male palpus, they are related toAlpaida, Eriophora, vised. ParTahweirxeiaa,reantdwoVesrpreuccieossao.f Edricus, both previously Revised herearethegeneraEdricusand known from only one sex, and 39 species of Wag- Wagneriana which share a similar abdo- neriana, 26 of them new. That is, only one-third of men shape and similar genitalia, both syn- thespecieswerepreviously known. Anawixia Cham- apomorphies. The species of the two gen- sbneyarnmloeinnsyaamnrsdePosfayrntaohvneeyrmnriuazcmeodes.aWMaeglnleor-iLaeintaa.oaErieghstubjsepcetciivees deirsatinhcatvieve cbaereanpacceonafnudsesdtedrensupmitsehaptheesi.r Only one common species of Wagner- INTRODUCTION iana was previously readily recognized, W. tauricornis, common in Florida. Roewer This is oneof a series of papers authored (1942) and Bonnet (1959) list four other bteynLdeevdittohmraokugehitthpoespsiebrlieodto1i9d6e8nt-i1f9y9N0eion-- 3sp9ecdiiefs.feTrehnetcsoplelceicetsi,onssoamveailoafblwehiincchluhdaedd tropical orb weavers. The araneid orb been placed in other genera of the family. weavers are the third largest spider family with 2,600 named species world-wide METHODS AND MATERIALS (Platnick, personal communication, 1989). About one-third of the Neotropical orb- The methods were the same used in my weaverspecieshavenow been revised. De- previous papers: careful comparison of the spite the popularity of orb weavers as re- structures of the genitalia of males and search subjects in studiesof behavior,ecol- females made by illustrating them to sep- ogy, and silk production, identification has arate species. The features that make the been severely hampered because the de- genusdistinct are also illustrated,although scriptions of species are scattered through a careful comparison of genera has to wait old literature and were made (as is unfor- until the species of more of them are tunately still sometimes done) without ref- known. Species descriptions cite the dis- erence to or comparison with previously tinguishing features and the localities described species. Some had been placed where specimens have been found. De- in wrong genera and frequently malesand spite this, problems invariably remain. Some of them can be resolved by studying 'Museum of Comparative Zoology, Harvard Uni- specimens, others by using more sophisti- versity, Cambridge, Massachusetts 02138. cated methods. (But molecular studies Bull. Mus. Comp. Zool., 152(6): 363-415, July, 1991 363 364 Bulletin Must iparative Zoology, Vol. 152, No. 6 would be inappropriate for this study, as ier to examine than the epigyna of species the) would not facilitate recognition or of other genera. riacement ofspecimens intothespecies or Paratypesandholotypes. Otherthanthe mI atching of names with specimens.) Re- holotypeand doubtful specimens, all spec- maining problems are that some species imens examined of a new species are ran be clearly recognized in one sex only marked and reported as paratypes. This (e.g., thereare difficulties in separating fe- permits wide distribution of paratypes to males of W. jelskii, W. maseta, and W. museums in Central and South America transitoria, also females of hassleri and and will facilitate future identification of silvae). Another persistent problem is specimens. Holotypes are deposited in a whether differences between specimens museum with a professional arachnologist from different placesrepresentspeciesdif- as curator or may have to be returned to ferencesor geographical variation (e.g., W. the country where collected. Rarely is the huanca, Figs. 149, 150). In some cases the specimen illustrated or described not the association of males with females may re- holotype, but if the holotype is in poor main uncertain (e.g., W. taim). condition, a better specimen was used for Eye ratios. The diameters of the pos- description and illustration. Ifan adequate terior median eyes and lateral eyes as seen illustrationwasmadeearlier Idid not make in profilearemeasuredbycomparison with another even if better specimens of the the diameter of the anterior median eye same species were found later. The holo- as seen in profile. The distances between types of previously named species have the borders of the anterior median eyes been examined and illustrated over a pe- and between the anterior median eyesand riod of twenty years because many had anterior lateral eyesare measured by com- been misplaced. (Misplaced specimens are parison with the diameter of the anterior a unique problem of revising a very large median eyes in profile. The distances be- family.) Some specimens for which my in- tween the posterior median eyes and be- formation was incomplete could not be tween the posterior median eyes and pos- borrowed a second time (those of the Ca- terior lateral eyes are measured by the racas and Rio de Janeiro museums). comparison with the diameter of the pos- terior median eyes as seen in profile. The method was first suggested by H. Homann ACKNOWLEDGMENTS (personal communication) as an alterna- tive to giving absolute measurements or The specimens used for this revision be- reporting the fractions as read in a mi- long to or are deposited in the following crometer. However, the measurements collections, whose curators I thank for here and those in my other araneid papers making the specimens available to me: are only rough calculations as araneid AMNH American Museum of Natural specimens of the same species are quite History, New York, United variable in their eye diameter and place- States; N. Platnick, L. Sorkin ment, and commonly one or two eyes are BMNH British Museum (Natural His- <l< formedor missing (e.g.,theanteriorme- tory), London, England; P. dian eyes of the holotype of W. Janeiro Hillyard, F. Wanless are absent) CAS California Academy of Sci- Internal female genitalia. Illustrations ences, San Francisco, United tweemrpeormaraidley ibnyHmoyoeurn'tsinmgedtiheumepoingaynmuim- SUtbaitecsk; W. J. Pulawski, D. cope slide, the dorsal side facing up. CNC Canadian National Collec- Sin.. Wagneriana epigyna are lightly tions, Ottawa, Canada; C. sclerotiz* d and relatively flat they are eas- Dondale Edricus and Wagneriana • Levi 365 CUC Cornell University Collection, de San Marco's, Lima, Peru; kept in the AMNH; N. Plat- D. Silva D. nick MIUP Museo de Invertebrados, DU D. Ubick Universidad de Panama, Pan- EMUCB Essig Museum, University of ama City, Panama; D. Quin- California Berkeley, Califor- tero A. MLP nia, United States; E. I. Museo, Universidad Nacional Schlinger, C. E. Griswold de La Plata, La Plata, Argen- FSCA FloridaStateCollectionof Ar- tina; R. F. Arrozpide thropods, Gainesville, Flori- MNRJ Museu Nacional, Rio de Ja- da, United States; G. B. Ed- neiro, Brazil; A. Timotheo da wards Costa IBNP Inventario Biologico Nacion- MNHN Museum National d'Histoire al, San Lorenzo, Paraguay; J. Naturelle, Paris, France; J. A. Kochalka Heurtault IMPR I. M. P. Rinaldi, Rio Claro, MNSD Museo Nacional de Historia Est. Sao Paulo, Brazil Natural, Santo Domingo, Do- INPA Instituto Nacional de Pesquis- minican Republic; B. C. Rey- as da Amazonia, Manaus, Est. noso S. MPM Amazonas, Brazil; J. A. Ra- Milwaukee Public Museum, phael, H. Hofer Milwaukee, Wisconsin, Unit- IRSNB Institut Royal des Sciences ed States; J. P. Jass Naturelles de Belgique, Brus- MZSP Museu de Zoologia da Univ- sels, Belgium; L. Baert ersidade de Sao Paulo, Sao JAK John A. Kochalka Paulo, Brazil; P. Vanzolini, L. MACN Museo Argentino de Ciencias Neme, J. L. M. Leme Naturales, Buenos Aires, Ar- MZUF Museo Zoologico, Universita, MCN gMeunsteinua;deE.CAi.enMcaiausryNaturais, NHMW FNlaotruernhcies,toIrtailsyc;hSe.sMaMsucsheeruimn,i SPuolr,tBoraAzlielg;rAe.,LRiisoe,GEr.aBnudcekudpo NMB WNaiteunr,hiAsusttorriias;chJ.esGrMuubesreum, MCZ Museum of Comparative Zo- Basel, Switzerland; E. Sutter ology, Cambridge, Massachu- NRMS Naturhistoriska Riksmuseet, setts, United States Stockholm, Sweden; T. Krone- MECN Museo Ecuatoriano de Cien- stedt cias Naturales, Quito, Ecua- PAN Polska Akademia Nauk, War- dor; L. Aviles szawa, Poland; A. Riedel, W. MEG M. E. Galiano Starega, J. Proszynski, A. Slo- MHNC Museu de Historia Natural, jewska, E. Kierych Capao da Imbuia, Curitiba, REL R. E. Leech Parana, Brazil; L. Bittencourt, SJ Steve Johnson S. de Fatima Caron SMF Natur-Museum und For- MHNM Museo de Historia Natural de schungs-Institut, Sencken- Montevideo, Uruguay; R. M. berg, Frankfurt am Main, Capocasale Germany; M. Grasshoff MHNMC Museo de Historia Natural, UCR University of California, Riv- Medellin, Colombia; Marco A. erside, California, United Frommer Serna D. States; S. I. MHNSM Museo de Historia Natural, USNM National Museum of Natural Universidad Nacional Mayor History, Smithsonian Institu- Museum of Comparative Zoology, Vol. 152, No. 6 tion, Washington, D.C., Unit- of Edricus are almost 1.5 to 2 times the ed States; J. Coddington diameter of the anterior medians (Fig. 5). YI)L Y. D. Lubin The height of the clypeus equals three to ZMK Zoologisk Museum, Copen- fourdiametersof theanterior median eyes hagen, Denmark; H. Enghoff (Fig. 5). The prosomal modifications, the ZSM Zoologische Staatssammlung, high clypeus and the large posterior me- Munich, Germany dian eyes are synapomorphies for the two species of Edricus. Unlike that of Wag- would also like to thank Renner L. C. neriana the palpus has a paramedian 1 Baptista for his help in locating obscure apophysis in the shape of a toad-stool on Brazilian collecting sites, Allen Dean for a its side (PM in Figs. 6, 13). The fourth leg gift of specimens, W. Eberhard, J. A. Ko- isaslongasorlongerthan the first,another chalka, and Yael Lubin for ecological in- synapomorphy for the two species. (The formation, Dee Woessner for word pro- fourth leg is also longer than the first in cessingthe keys, Lorna Levi forrewording the unrelated Micrathena species, Levi, sentences and carefully reading the whole 1985.) manuscript, and Diana Sherry for con- Description. Carapace, chelicerae, en- verting the manuscript into an electronic dites, sternum orange-brown. Coxae or- version. Laura Leibensperger helped angetoorange-brown. Legsorange-brown. throughout with all researches. Two re- Abdomen with some black and white pig- viewers made valuable suggestions to im- ment. Carapace without macrosetae. Ab- prove the manuscript. domen with four pairs of lateral tubercles The beginning of this revision was sup- and three posterior median ones. In males ported by National Science Foundation the anterior lateral tubercles are promi- grants B-5133, GB-36161, BMS 75-05719, nent spines (Figs. 7, 14). The shape of the DEB 76-15568, DEB 79-23004, DEB 80- abdomen with pairs of tubercles and pos- 19732,and BSR 83-12771. Publicationcosts terior median tubercles (Fig. 3, 11) resem- for this study were covered in part by the bles that of Wagneriana. The epigynum Wetmore-Colles Fund. has a median lobe (Figs. 1, 9) also resem- bling that of Wagneriana. Both these sim- Edricus 0. P.-Cambridge ilarities are regarded as synapomorphies. Epigynum small with a posterior me- EdricusO. P.-Cambridge, 1890:57. TypespeciesEd- dian plate that differs in shape in the two irsumnasscsupilniingeer(uBsonbnyet,mo1n9o5t6y:py1.647T)h.e generic name swpietchieosne(Fmiagsc.ro2s,et1a0.).ThMaelmealpealopfalE.paptreoldl-a Note. The literature isconfused because uctus has a tooth on the endite and a hook females of the common Witica crassicau- on the first coxa (Fig. 8), E. spinigerus has d(i (Keyserling) wereerroneously matched lost both. Edricus productus has a mac- with Edricus males and placed in the ge- roseta on the fourth trochanter, E. spini- nus Edricus. Also several Alpaida (see list gerusdoesnot. Thesecondtibia hasasmall below) species had been described in Ed- spur with one or two macrosetae in males runs. of E. productus (Fig. 7), with one mac- Diagnosis. Unlike those of related gen- roseta in E. spinigerus (Fig. 14). The pal- era and most other araneids, the carapace pus is similar in structure to that of Wag- and sternum of Edricus are modified. The neriana species. The conductor is on the carapace in both sexes is elongated and ventral face of the palpus as in Wagner- narrowed in the thoracic region (Figs. 3, iana, Parawixia,andAlpaida (not nearthe < . II. 14) and the sternum is posteriorly rim as in Araneus). narrowed and elongated (Figs. 4, 8, 12, Distribution. There are only two spe- 15 l nlike those of most other araneids, cies, one in Mexico, the other in Central epl Pronous, the posterior median eyes' America to Ecuador (Map 1). Edricus and Wagneriana • Levi 367 productus Map 1. Distributionof Edricusspecies. Misplaced Species dianapophysisroundedlaterally(Fig.6); Mexico (Map 1) productus Edricusatomarius (Simon, 1895); Roewer, 1942: 761 Third coxa adjacent to fourth (Fig. 15); is Alpaida atomaria (Simon). See Levi, 1988: 458. palpus with median apophysis angular E. cayana (Taczanowski, 1873); Roewer, 1942: 762 laterally (Fig. 13); Panama to Ecuador is Witica cayana (Taczanowski). See Levi, 1986: (Map 1) spinigerus 44. E.crassicauda (Keyserling, 1865); Roewer, 1942: 762 isWiticacrassicauda (Keyserling). See Levi, 1986: Edricus productus O.P.-Cambridge 41. Figures 1-8; Map 1 E.ensiferdiCaporiacco, 1947:25isAlpaidatruncata (Keyserling). See Levi, 1988: 472. Edricus productus O. P.-Cambridge, 1896: 186, pi. E. eupalaestris Mello-Leitao, 1943: 177 is Wagner- 23, fig. 5, 8. Male holotype from Cuernavaca, Mo- iana eupalaestris (Mello-Leitao). relos State, Mexico, in BMNH, examined. F. P.- E. tricuspis (Getaz, 1893); Roewer, 1942: 762 is Wi- Cambridge, 1904: 500, pi. 49, fig. 26, S. Roewer, tica crassicauda (Keyserling). See Levi, 1986: 41. 1942: 762. Bonnet, 1956: 1648. E.istrAulnpcaaitduas t(rKeuynsceartlain(gK,ey1s8e6r5l)i;ng)R.oeSweeer,Le1v9i4,2:1978682: Description. Female from Escuintla, 472. Chiapas. Sternum orange with median white streak. Venter of abdomen with a Unrecognizable species pair of white blotches framed by black. Posterior median eyes 1.5diameters of an- Edricus rubricornis Mello-Leitao, 1940: 204. Female terior medians, anterior laterals 0.9 di- h(oMlNoRtJy)p,e lofsrt.om Colatina, Espirito Santo, Brazil ameter, posterior laterals 1 diameter. An- terior median eyes 1.2 diameters apart. Key to Edricus species Posterior median eyes2.5 diameters apart. 1. FMaelmeasles 32 Ambmd.oCmaernapaascein6.F3igmumrel3o.ngT,ot2a.8lmlemngwtihde1.5 2-(1). PPooss2ttt)e;eerrriMoiireoorrxvimmiceeeoddwii(aaaMlnnamopppsllata1tt)eeasooffloeenppgiiggasyynnwuuimmdeiipnnr(ppFoooidsgsu.--ctus mSFiemrcs;otnfmdeetmpauattraerl3ls.au5sam2nm.d6;tmpiabmit;aell3ta.a5rasnumsdm1;t.i1bitahmim4r.d.1, terior view about twice as wide as long 2.5mm. Fourth femur 4.5 mm; patella and (Fig. 10); Panama to Ecuador (Map 1) . tibia4.4mm; metatarsus3.1 mm,tarsus 1.0 spinigerus mm. 3(1). Thtihredircodxiaamseetpearra(tFeidg.fr8)o;mpfaoluprutshwbiythabmoeu-t Malefrom Tepic,Nayarit. Posteriorme- Comparative Zoology, Vol. 152, No. 6 dun eves 1 5 diameters of anterior me- poneta, 29 Nov. 1939, imm. (H. Bogert, dians anterior laterals 0.8 diameter, pos- N. E. Voices, AMNH); Tepic, 6 (MCZ), 15 e,tp>eaxrleeipssoar2l1l8daftideeairmmaauelmrsteet1rwedsiritsaahpmaaepriattnre.dtri..EstnPAidonnsictttteeerrtiiwoooiorrtthhmm.eetddoFoiiitraahsnn,t SAM1eau9pl6gt3k..,in11,9i965Am33mM,,.N2iH<m5)(mW;(..B.8(kJMC.ma,lGkNMeir.nWt,sGTcAoehopM,diNcnHW,ig).1h,3t,MI2vai-Bey7., coxa with hook (Fig 8). Fourth trochanter AMNH).Jalisco: LagodeChapala, 16July with one macroseta. Second tibia thicker 1976, 9 (J. Harp, R. Mitchell, SJ). Colima: 7tm)haacnrAobfsidresottamaeonnndrlwiiigkthethtldheiagst,tatolfwsofpeoumnralbleeefatrbliuentgg(wFoiintgeh. c(1a9Wn.:kJm.1G6EerktMsmacnhzS,anWiU.lrluoIav,piea1,n1,AMM6aNyHJ)u1l.9y6M31,i9c8ih5mo,md.-6 a5(n.F4itge.rmi7om)r. plTaoointrga,lof2l.se3pnigmntemhs9sw.l6iidghemt.mly.FisrcCslaterrfaoetpimazucered k((EWm.ooEIl.leTSyec,hpleZixonclgone,re,r1o,vE2iM5cU0h,CmBM,)C2.Z4).CAhuPigua.epba1ls9a:7:7E0,s.-86 3.5 mm; patella and tibia 4.0 mm; meta- cuintla, 2 (Crawford, MCZ). tarsus 2.8 mm; tarsus 1.1 mm. Second pa- tella and tibia 3.4 mm; third, 2.3 mm. Edncus sp.mgerusO. P.-Cambr.dge Fourth femur 3.7 mm; patella and tibia Figures 9-15; Map 1 3.9mm;metatarsus2.9 mm;tarsus 1.8 mm. Edricusspinigerus O. P.-Cambridge, 1890: 58, pi. 4, Illustrations. The illustrations were fig. 1, <5. Male holotype from Bugaba, Panama, in made from a female from Escuintla, Chia- BMNH, examined. Keyserling, 1892: 33, pi 2, fig. *^$£l£*^J£&?£gi pas, and fromamalefrom Tepic, Nayarit. Variation. The abdomen shape is quite variable in females, especially its width Description. Female from Ecuador, and the size of the spines. Total length of Legs orange-brown with indistinct, dusky females 11.7 to 18.5 mm, of males 9.5 to longitudinal streaks. Venter of abdomen 9.6. The male from Puebla had only one black between epigynum and spinnerets macroseta on the spur of the tibia. with a white line on each side, white bor- Diagnosis. Females differ from E. spi- dered withblack on each side behind spin- nigerus by the posterior median plate of nerets. Posterior median eyes 2.2 diame- the epigynum being as wide as long (Fig. ters of anterior medians, anterior laterals 2) and the outline of the carapace (Fig. 3). 0.9 diameter, posterior laterals 0.9 diam- The male differs by the folding of the dis- eter. Anterior median eyes their diameter tal end of the median apophysis (Fig. 6) apart. Posterior median eyes 1.1 diameters afonudrtthhetrsohcahpaentoefrthheassatemrancurmos(eFtiag,. l8)a.ckTihneg alpeanrgtt.h 8A.b0dmomm.enCaraaspianceF3i.g0urmem11l.ongT,ot1a.l6 mm in /.. spinigerus. wide. First femur2.5 mm; patellaand Records. MEXICOSinaloa: Mazatlan,6 tibia 2.7 mm; metatarsus 1.9 mm; tarsus Sept. 1956, 2 (A. F. Archer, AMNH), 2 1.0 mm. Second patella and tibia 2.3 mm; Sept. 1977, 2 (C. E. Griswold, EMUCB); third, 1.7 mm. Fourth femur 3.0 mm; pa- La Concordia, N Copala, 610 m, 10 Sept. tellaand tibia3.1 mm; metatarsus2.1 mm; 1979, 2 (D. E., J. A. Breedlove, CAS); 32 tarsus 0.9 mm. km E Villa Union, 19 Sept. 1964, imm. (E. Male from Depto. Cauca, Colombia. I. Schlinger, UCR). Nayarit: San Bias, 17- Coxae brown; legs dusky orange. Posterior 21 Sept 1953, 2 (B. Malkin, AMNH); Aca- median eyes 1.6 diameters of anterior me- :dricus productus(O. P.-Cambridge). 1-5. Female. 1. Epigynum, ventral. 2. Epigynum, posterior. 3. Dorsal. 4. idcoxae. 5. Eyeareaandchelicerae. 6-8. Male. 6. Leftpalpus. 7. Dorsalwithsecond leg. 8. Sternumandcoxae. Sternumandcoxae. i1i3g-e1r5u.s(M0a.leP..-1C3a.mbPrailpdugse.).149.-1D2o.rsFaelmwailteh.s9e.cEopnidgylengu.m1,5.veSnttrearln.u1m0.anEdpicgoyxnauem., posterior. 11. Dorsal. 12. Edricus and Wacneriana • Levi 369 radix Figures16-23. Wagnerianaanatomy. 16-18. W. tayosn.sp.,female.16.Carapace,dorsal. 17.Carapace,chehcera,lateral. 18. Eyeareaandchelicerae. 19-20. W. transitoria, left palpus. 21-23. IV. tayosn. sp., male. 21. Carapace,dorsal. 22. Carapace andchelicerae, lateral. 23. Eyearea,chelicera, andrightpalpus. Scalelines. 1.0mm,genitalia, 0.1 mm. irative Zoology, Vol. 152, No. 6 ddtmieieraadinnosir,aenlyaaentesteyerera1isl.so2r10.d.l4i8atadedmirieaaatlmmeseertt0se.er8ras.pdaiaArpantam.treettrP,eioros1,rt.e3prmoidesoi--r- lw71aa52tr0okdns,mm,,2E(F1GSS1.Ca-WAn1t).7o;PMDr4eao7smycioktn1mt9g,8oSM6,CdSZea2),nl;to8soT(CiDGon.olamolBrai.annddgEioodas-,,, ametersfrom laterals. Enditewithouttooth. Rio Palenque Sta., 18-30 May 1975, 6 (S., First coxa without hook. Fourth trochanter J. Peck, CNC). Bolivar: Balzapampa, 700 without macroseta. Second tibia thicker m, 28 May 1938, 2 (W. Clarke-Macintyre, than first, with a distal spur and one mac- AMNH). roseta (Fig.m1m4). Total lengmthm6.5 mm. Car- afpeamcuer 32..04 mm;lpoantge,lla1.5and tibwiaid2e..7 Fmimrs;t Wagneriana F. P.-Cambridge metatarsus 2.0 mm; tarsus 1.0 mm. Second Wagneria McCook, 1894: 203. Type species by 2pF.ao7tuermltlmah;afmneedmtuattriabri2as.u72s.m12.m1m;mmm;p;attehtlialrrads,usa1n0..d68mtmimmbi..a mnvaooaminmdooyelt,lyius1ps8kpy3r0E(e,pNoeecfiaocrrvuaepa,itdea1idu9pr4tib0ecy:roarW6nna5,i0sg)anO.ne.drPib.ay-CGRaiomsbtblir,nied1ag8ue4.-8,DTefhsoe-r Illustrations. The illustrations were WagnerianaF. P.-Cambridge, 1904:497. Newname made from specimens from Pichincha for Wagneriana McCook, preoccupied. The name Province, Ecuador. is of feminine gender. Anawixia Chamberlin, 1916: 258. Type species by Variation. Total length of females 8.0 monotypyandoriginaldesignationA.atopaCham- to 10.2 mm, of males 6.5 to 7.1. berlin, 1916: 258 [= W. transitoria (C. L. Koch)]. Diagnosis. The femalecan be separated NEW SYNONYMY. from that of E. productus by the shape of Paraverrucosa Mello-Leitao, 1939a: 64. Typespecies theepigynum,widerthanlongin posterior bLeyitmaoo,no1t9y3p9ay:a6n5.dNdeEsiWgnaStYioNnOPN.YnMeYg.lecta Mello- view (Fig. 10), andby thedifferent outline of the carapace (Fig. 11). The male is sep- Diagnosis. The carapace is high, the ce- arated by the shape of the distal end of phalic region slightly swollen behind the the medianapophysis (Fig. 13),bytheout- eyes (Figs. 18, 23), and in the female the line of the sternum (Fig. 15), and by the sidesof the thoracic region are usually gla- lack of a macroseta on the fourth trochan- brous, often dark (Figs. 16, 27, 38, 63). ter. The carapace of the female may have a Xataral History. The specimens from pair of macrosetae or more in the thoracic Tinalandia were collected by beating veg- groove (Figs. 32, 38; Levi, 1976, figs. 62, etation. Lubin (personal communication, 63, 64, 67). The glabrous often dark sides 1989) found an immature in a nearly ver- of the carapace as well as the macrosetae tical web, 10 cm in diameter, 5 cm above in the thoracic groove aresynapomorphies ground. The adult female was a meter of the species of Wagneriana. The ma- away, an immature male in a similar-sized crosetae may be absent, perhaps second- web 5 to 10 cm above ground and 10 cm arily lost. The paramedian apophysis of in diameter. The web was symmetrical the male palpus is an L-shaped rod (Fig. with an open hub, the female sat cephalic 19), lying on its side, rarely rounded (U- region down in the hub. Another maleand shaped) or withan acuteangle (V-shaped), female were collected sitting together un- a synapomorphy shared by all species of der a leaf about 10 cm from the female's Wagneriana but not so in Edricus species. web The terminal apophysis and embolus are COLOMBIA Records. NW Cauca: Pacific fused (Figs. 19, 20). Another synapomor- istal plain Guapi, Jan. 1973, 6 (W. phy of Wagneriana species is the modi- Eberhard, MCZ). ECUADOR Pichincha: fication of the base of the median apoph- Rio Palenque, km 47 on Santo Domingo- ysis above the radix; it may have a small Quevedo Road, 150 m, 15 Mar. 1982,9,3, depression (Fig. 28) or teeth (Fig. 19), but imm. (Y. Lubin YDL 383, MCZ); Rio Pi- lacks the large concavity of the median Edricus and Wagneriana • Levi 371 apophysis of males of Eriophora and Par- median anterior notch (or pocket) as in awixia. In most genera of araneids the at- Alpaida (a pocket ispresentin W. yacuma, tachment of the median apophysis is not Fig. 172, and the epigynum is rebordered modified; it may be a sclerotized bar or in W. gavensis, Fig. 134). Posteriorly the may not be sclerotized. epigynum has a median plate and two lat- Description. The coloration of all spe- eral plates, the lateral plates continue ven- cies is about the same: carapace, sternum, trally and form the wide median lobe in legs yellowish to orange-brown. Carapace ventral view (Figs. 25, 30). The seminal often with dusky marks and sides of tho- receptacles of all females were illustrated, racic region usually, but not always, dark although it is not known whether this will and shiny, and cephalic region with some be useful. They areeasily examined unlike white setae (Fig. 63). Sternum always those of some other araneid genera. The darker than coxae. Legs usually with in- receptaclesofsomespeciesareconsistently distinct dusky rings. Abdomenspotted with thin-walled (Fig. 125), others thick-walled shades of brown, usually without folium (Fig. 118). often with dark median band (Figs. 57, In males the carapace is much narrower 86);notwospecimensofaspeciesareiden- in front than in females, high in the tho- tical. Venter gray to black with indistinct racic region, with few short setae and lat- white marks. All Wagnerianaspecieshave eral eyes not on tubercles (Figs. 21-23). a narrow, soft abdomen, longer than wide All males have one patellar macroseta on with nine to 15 tubercles (Figs. 27, 70, the palpus; endite has a large tooth (Figs. 153), three pairs anterior and middle on 22, 23) facing a tubercle on proximal end sides, one pair posterior on sides, and two of palpal femur. First coxa with a distal, or three in a posterior, median line (the ventral hook (Figs. 22, 23) and in larger most posterior tubercle above the spinner- species with a dorsal tubercle that fits ets, which may face posteriorly or ven- against the carapace rim. As in Parawixia trally, may be lacking). The most anterior and Wixia some species have one or more pair of tubercles may be double in W. macrosetae on fourth trochanter. Second uzaga, W. spicata, W. gavensis, and W. femur (sometimes also third and fourth, iguape and this is consistent in all speci- rarely first) with ventral row of macrose- mens of several species (Figs. 126, 128, tae. In all species second tibia thicker than 132). The tubercles of some specimens of first, moreor lessswollen, with short,strong several species may be sclerotized spines, macrosetae. Abdomen similar to that of There is no median anterior tubercle (ex- female but smaller (Fig. 128). cept in W. turrigera[Figs. 203,204],which Theterminalapophysisandembolusare may not belong here). The position of the fused (Figs. 19, 20). The terminal division, tubercles of the abdomen is a synapomor- the terminal apophysis and embolus, is a phy shared with Edricus and Parawixia triangular structure, fan-shaped, its outer species. Theabdomen may betruncatebe- edge modified in various species. The fan- hind the spinnerets (Fig. 64) or may have shaped terminal division is a synapomor- a postanal tail (Fig. 66); both shapes may phy shared with Edricus and Parawixia befound indifferentspecimensofthesame species. Some female individuals have lost species. Posterior median eyes usually the macrosetae on the carapace and some slightly smaller than anterior medians, lat- males have lost the macrosetae on the erals the smallest (Figs. 17, 18). fourth trochanter (in species that normally Another apomorphy of Wagneriana have them). In all species there is consid- species shared with Edricus is the shape erable individual variation in markingsand oftheepigynum,awidemedian lobeoften in color, size, length, and prominence of with a minute, light scape at its tip (Figs, tubercles, and the length of the postanal 24, 45). It is not rebordered and lacks a tail. Twospecies always have the tubercles Bulletin Museum of Comparative Zoology, Vol. 152, No. 6 Plate1. Webof Wagnerianaundecimtuberculata,horizontaldiameteroforb26cm(photoW. Eberhard). drawn out: W. grandicornis (Figs. 65, 66) the tips of thin branches with their legs and W. heteracantha (Fig. 108). Only one pressedtotheirbodies. Theirirregularout- species,W. neglecta,consistentlyhasalong lines make them hard to recognize as a tail (Figs. 119, 122). Most species are sur- spider. When they have an orb they gen- prisingly similar in appearance and can erally hang in the hub or sit on one of the not be separated by color pattern or body frame or anchor lines facing away with a shape; they have to be separated by the line to the hub held by leg IV. genitalia. Distribution. All species are Neotropi- Natural History. All species make a cal. Only W. tauricornis extends its range complete orb web and hang cephalic re- into more temperate North America, and gion down in the middle. There is no re- W. spicata is found in Mexico. Most spe- treat (Plate 1). ciesarefound in theAmazondrainageand W. Fberhard (personal communication) southeastern South America (Maps 2-4). found that W. tauricornis and W. unde- cimtuberculata taketheirwebdown when Doubtful placement not in use (usually but not always in the One species, W. turrigera, has been placed in Wag- daytime) and then sit at exposed sites like neriana forconvenience; it may have tobe placed

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