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Two new species of Manaosbiidae (Opiliones: Laniatores) from Panama, with comments on interspecific variation in penis morphology PDF

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Preview Two new species of Manaosbiidae (Opiliones: Laniatores) from Panama, with comments on interspecific variation in penis morphology

2011.TheJournalofArachnology39:92-101 Two newspecies ofManaosbiidae (Opiliones: Laniatores) from Panama, with commentson interspecific variation in penis morphology Victor R.Townsend,Jr.: Department ofBiology, VirginiaWesleyan College, 1584Wesleyan Drive, Norfolk, Virginia 23502 USA. E-mail: [email protected] MarcA. Milne: DepartmentofBiological Sciences, Old Dominion University, Norfolk, Virginia23529 USA DanielN. Proud: DepartmentofBiology, UniversityofLouisianaatLafayette, Box42451, Lafayette, Louisiana70504- 2451 USA Abstract. InCentral America, the family Manaosbiidaeisrecordedonly from PanamaandCosta Rica. Fourspecies occurinthisregion: BarromiwillicimsiGoodnight&Goodnight 1942,BugahitiatriacanthaRoewer 1915,Poassalimhata Roewer1943,andZygopachyhisalhonunginisChamberlin1925.Inthispaper,wedescribeBcirroncifelgenhaiwrinewspecies (Code Province, Panama) and Bugahitiaakininew species (Code Province, Panama) and report a new record for B. nilliaimi(CodeProvince,Panama).WeusedSEMtoexaminethepenismorphologyofBarronaGoodnight&Goodnight 1942 and theCaribbean species CrauelliisnwntgomeryiGoodnight &Goodnight 1947 and Rhopalocranausalhilineatus Roewer1932.WecomparedgenitalmorphologyofthesespecieswithpublisheddescriptionsforManaosbiidaefromSouth America. Withrespecttogenitalmorphology,wefoundthatthemostvariablecharacterswerethenumberandrelative sizesofthesetaethatoccuronthelateralmarginsoftheventralplate.Otherfeaturesthatexhibitedinterspecificvariation includedtheshapeoftheventralplate,theshapeofthedistalborderoftheventralplate,andtheshapeandarmatureofthe apexofthestylus. Keywords: CentralAmerica,morphology.Neotropics,taxonomy TheManaosbiidaeisamemberofthesuborderLaniatores.It tarsus I ofmales in most species are generally enlarged and belongsto thesuperfamilyGonyleptoidea,a lineagethatalso frequentlyfused(Kury2007).Observationsofthesesegments includestheCosmetidae,Cranaidae,andGonyleptidae(Kury with the aid of scanning electron microscopy (SEM) have 2007). Recently, a phylogeneticanalysis usingmoleculardata revealed that these tarsal segments have numerous pore (Giribetetal. 2009)supportedthemembershipofManaosbii- openings, which are hypothesized to beconnected to packed daewithin thisclade. However, this studyalso indicated that clusters ofexocrineglands that may function in intraspecific Manaosbiidae is polyphyletic, at least with respect to the communication (Willemart et al. 2010). The only known inclusion ofthegenusZygopachyhisChamberlin. Manaosbii- specieswithinthefamilyfromCentralAmericathatlacksthe dae was initially recognized as the subfamily Manaosbiinae enlargedsegmentsoftarsusIisZygopachylusalhomarginis.In withintheGonyleptidae(Roewer 1943). Kury(1997)elevated this species, malesconstructand defend mud nestsand mate thegrouptofamilystatus,refinedthecharactersdistinguishing with visiting females (Rodriquez & Guerrero 1976; Mora theManaosbiidaefromtheCranaidaeandGonyleptidae,and 1990). Following oviposition, the males remain in the nests provideddiagnosticcharactersforthefamily. and actively defend the eggs against ants and conspecifics Of the 47 species and 27 genera currently placed in the (Mora 1990). Currently, four manaosbiid species are known Manaosbiidae (Kury 2007), 12 species are known only from fromCentralAmerica,namelyBarronawilliamsiGoodnight& femaleholotypes(Kury2003). Themorphologyofthe penis, Goodnight 1942a (Panama), Bugahitia triacantha Roewer animportantstructureinmoderntaxonomicdescriptionsfor 1915 (Panama), ZygopachylusalhomarginisChamberlin 1925 Opiliones (Acosta et al. 2007), has only been described for (Panama) and Poassa limhata Roewer 1943 (Costa Rica). seven species, all from South America (Silhavy 1979; Kury Each of these monotypic genera is endemic to Central 1997,2007). America (Kury 2003). In this study, we describe two new Currently, the Manaosbiidaehasageographicdistribution species, Barronafelgenhaueriand Bugahitiaakini. To provide that includes the Caribbean islands of Trinidad and St. greater insights into the phylogenetic relationships ofmana- Vincent,northernSouthAmerica,andCentralAmerica(Kury osbiids, we used SEM to examine the penis morphology of 2003).Mostspeciesaresmall(3.5-10mminscutallength)and Barrona and two Caribbean species (Cranellus montgomeryi knownfromonlyrelativelyfewrecords.Thisisdue,atleastin and Rhopalocranausalhilineatus). We compared these obser- part, to undersampling or lack ofsampling the leaflitter, a vations to published descriptions of penis morphology for microhabitatin which theycan be relativelyabundant(Kury sevenspeciesfrom SouthAmerica(Silhavy 1979; Kury 1997, 2007). Little is known about the natural history of these 2007). harvestmen, although Townsend et al. (2008a) provided METHODS observations concerning activity, habitat use and geographic distribution for Cranellus montgomeryi Goodnight & Good- The specimensexamined in this studyaredeposited inthe night 1947aand RhopalocranausalhilineatusRoewer 1932on American Museum of Natural History, New York, USA theCaribbeanislandofTrinidad.Thetwobasalsegmentson (AMNH);SenckenbergMuseum,Frankfurt,Germany(SMF) 92 TOWNSENDETAL.—NEWMANAOSBIIDAEFROMCENTRALAMERICA 93 and the Museo de Invertebrados G.B. Fairchild de la Manaosbiinae:Roewer1943:14,56;Soares&Soares1949:224. Universidad de Panama, Panama City, Panama (MIUP). Manausbiinae[misspelling]: Mello-Leitao 1949:12. Specimens were examined and photographed with a Leica Stygnoleptinae[part]: H. Soares 1972:68. Zoom stereomicroscope. Digital images of specimens were Manaosbiidae: Kury 1997:3; Kury2003:206; Kury2007:209. processedandthebodyandlegsegmentsweremeasuredwith Emended diagnosis.—Gonyleptoidea with abdominal scute theaidofaLeicaimagecapturingsystem. only slightly wider than carapace, ocularium small, without tbPChryaeaAnndCeRaule.mllniattiMrsmaiialmrnloaaennnsStddegaopofNtmofeBeramrrtrboyhremieorrnanPana2dR0rw0aiqR9lnuh.lgeioeapsmWaSslueioomfwcmeTrcirraotieln,nlaiecucodsPtlaaeldade,nlcabtWisemeldpasienitcneiPaImrttnohemdveniisfefnsriceoeoilm,ndf cdstsluetpebpriseron,renicsggflsroeiaarsorn;mnum,laataubtubrduenorrtaemcurilboemnesnarecldolaennssoycrauorstnweeuiagmtamhrIeIeunIat;n1sa-Ip;3regmdpeseienmdpdeaiarlploalaplrlulposywarilsstlmmhfaaoerlopgmlateeuihrrr,septcdwihyilnatitinunhfbdooterurrhimt-e- Jsuclaynn2i0n0g6ealencdtr2o0n08m.icPreonsisceospywe(rSeEMdi)s.seSctpeedciamnednsprweepraereddehfyo-r csapli;ncifooxramItVubbearrcelelsyavinsdibalermaebdovweitshcustpe,indiofrosramllaypiccoavlertuebderwcilteh; deGolmappvAPLiroalxgnhoiozMllaauyadonoatetcinmttdNa.eeesioonsgdd,fiH-ngeipFoadermiugirononatndaohraftt-putpoaemwRhd1naaroioe5aitrgt-rtc&daacrkehgphgtoLaedhwVanyaandaiootfpGieot(lisaetecodn2hnoeaydo0oglseta0edaimtiho2wtanatncepfH)iheniai,vota,MaldgithnuiBaalaho.lncUdcnutlgefrhddSmioseionfsAilcaes(Aeldrnt.S7r1licc.iuiu-g90goopreh43mets%eIspt7Pn,0isnty)ehso0mea,ndatniC0atirusuocdeheNsi,ebfdtrnaKeeriontSswudwiatsoehlireEic.lrwatwoyowiMhrene(aapta,v2mtrnedhs0ae(pe.dot0t1stlhh7u9htoTsae)ee9bdemth.nyx7ixlUoei)aesaepnrans,msdmeeiceesnidvcpctttieenKuetihewslrenescyuriesdikedlmrtrisoddiahtytfintayaawsoiptnnaabilenho&ddsn-y,-gef savendstsVGdleuiorxeoeuinnbroptngDypgetsscoemiahasrtahshzseit,arnlaetelurnlalanrlydeGtiyity,,lgprsobceahlblefuorw,taeootnisPi,tncfidasettaaoaWdcmb,thnlnaIiooaeo.o;a-)sndfn—uomImi,.ctdsItpetaatIBwehdadv,ydrinra—elioasisu,rzmartcsusiaPslndasloleaar,syrrlwIetstuIscrwess,owtatiCplrhiaoaitoaaganenhnlSvnhgvddeuotpdfulepsnarmlelataiai-barrortp(lnrieaacsStieacbaltceketxm.o,anelaoelpseloffri,idiVnooncoiCgclonsnaotmcedTraectulsasertlbeeetsldsaifle,neeatteyrnts,;rociw.olrdbiostenaRataafpsgpinsrhda;lacsadrpyvialdsie,i&lecoIntssluntIhtearralIetsEtlavT-aycecleeIoa,GubdVpbraso,ubgaptedrwlallognodibasaauoarteueantt,h,-;lrt,se SYSTEMATICS Included genera. Azulamus Roewer 1957, Barrona Good- nightandGoodnight 1942, BelenmodesStrand 1942, Belemu- ManaosbiidaeRoewer 1943 lusRoewer 1932,BugahitiaRoewer 1915, CameliamisRoewer Mitobatinae[part]: Simon 1879:226. 1912, Clavicranaus Roewer 1915, Cranelliis Roewer 1932, Prostygninae[part]:Roewer1913:140; 1923:449;Mello-Leitao Ciiciitacola Mello-Leitao 1940, Dihiinostra Roewer 1943, 1932:103;Goodnight&Goodnight 1942a:11. Gonogotm Roewer 1943, Manaoshia Roewer 1943, Mazar- Cranainae [part]: Roewer 1913:349; 1923:536; Mello-Leitao uniiis Roewer 1943, Meridia Roewer 1913, Paramicrocranaus 1931:118; 1932:111; 1941:440;Roewer1938:6;Goodnight& Soares 1970, Pentacranaus Roewer 1963, Poecilocranaus Goodnight 1942b:7; Soares&Soares 1948:583. Roewer 1943, RhopalocranausRoewer 1913, Rhopalocranellus Hernandariinae [part]: Roewer 1913:460; 1923:582; Mello- Roewer 1925, Sanvicenlia Roewer 1943, Saramacia Roewer Leitao 1932:129; Soares&Soares 1949:221. 1913, Semostriis Roewer 1943, Syncranaiis Roewer 1913, Heterocranainae[part]: Roewer 1913:417; 1923:567. TegyraSorensen 1932andZygopachylusChamberlin 1925. KEYTOTHEMANAOSBIIDAEOFCENTRALAMERICA 1. Secondfreetergitewithsinglespiniformtubercle (Bugahitia)...! Secondfreetergitewithpairedgranulartubercles 3 2. Oculariumwithpairedspiniformtubercles;pairedspiniformtuberclesonabdominalscutalareaIIIwithoutsmallerencircling granulartubercles;tarsalformula6:15:6:8 Bugahitiaakinine\NspQCxes{¥\g. 1) Oculariumwithpairedgranulartubercles;pairedspiniformtuberclesonabdominalscutalareaIIIencircledbysmallertubercles; tarsalformula6:12:6:? BugahitiatriacanthaRoewer 1915(Fig.2) 3. Marginsofabdominalscutumunarmed (Barrona)...4 Marginsofabdominalscutumwithsinglerowofgranulartubercleswithterminaltubercle(adjacenttoareasIIIorIV)enlarged 5 4. Scutum with4white patches; smallerpatches on abdominal scutal area I, largerpatches on area II; tarsal formula 6:12:6:6 Barronafelgenhauerinewspecies(Fig. 3) Scutumwithoutwhitepatches; carapaceblack with lightermottling; tarsal formula6:16:6:7 BarronawilliamsiGoodnight&Goodnight 1942(Fig.4) 5. Terminalconicaltubercleonmarginofscutummuchlargerthanothertuberclesonscutalmargin;anteriorregionofcarapace with lighter mottling; more than 12 small tubercles on margins ofabdominal scutum Zygopachylusalhomarginis 1925Chamberlin(Fig. 5) Terminal tubercle on scutal margin only slightly larger than othertubercles on margin; anterior region ofcarapace without lightermottling;lessthan 12smalltuberclesonmarginsofabdominalscutum Poas.saliinhataRoewer 1943(Fig.6) — — — 94 THEJOURNALOFARACHNOLOGY — Figures 1-6. The Manaosbiidae ofCentral America: 1. Biigahitia cikini, new species, holotype, female; 2. B. triacanthci, Roewer 1915, holotype,female;3. Banomifelgenliuueri,newspecies,holotype,female;4. B. williamsi,malefromColonProvince,Panama;5.Zygopachyhts alhoniarginis,femalefrom BarroColoradoIsland,Panama;6.Poussaliinhata,(Roewer 1943),holotype,female.Scalebars = 2mm. BananaGoodnight&Goodnight 1942 withconcavedistalmargin;stylusunarmed,bentwithafolded Banana Goodnight & Goodnight 1942:11; Goodnight & apex. Basitarsus I of male spindle-like; 2 basal segments Goodnight 1947:11;Soaresetal. 1992:4;Kury1997:4;Kury enlarged. 2003:207. Type species.—Banana williamsi Goodnight & Goodnight Bananafe(lFgiegsn.ha3u,er7i-,15n)ewspecies 194E2m,ebnydeodridgiianganlosdiess.ignaAtnitoenr.iormarginofcarapacewith4-5 Materialexamined. PANAMA: CadePravince: Holotype granular tubercles on each side. Eye mound with 2 granular female, ParqueNacionalGeneral DivisionOmarTorrijosH., tubercles on each side, anterior tubercle smaller. Abdominal ElCope(08°49.2'80"N,80°05'45.7"W),23-28 February2007, socnutsaclutaarleaasre1aan11d11m1u1cwhitlhaprgaeirretdhatunbegrrcalneusl;asrptiunbiefrocrlmestuobneracrleeas hVi.kTiongwnsteranidls,Aa.tSanviigthztkyianndmJo.nRtaayn,ecorlaliencftoerdebstyh(aAndMNaHlo)n.g II;Va-rVeau1n1aurnmaerdmeadn,deixncdeipsttinfcotr,amfaerwgignrsanouflasrcututbuemrcluensa;ramreeda.s mPaalreat,yspeasm:el1o—cafteimoanl,e,1^colNloevcteemdbewrith198h0o,lDo.tyMpeosl(eAyM(NMHI)U;P).1 Lateralmarginsofscutumunarmed. Freetergiteswithpaired Etymology. Thisspecies isapatronym in honorofBruce granulartubercles,lateraledgeswithorwithoutsingletubercle Felgenhauerwhohasmademanycontributionstothestudyof on each side. Anal operculum with scattered granular themorphologyandnatural historyoftropicalarthropods. tubercles. Pedipalpal femur and patella unarmed; tibia with Diagnosis. Dorsal scutum attenuate pyriform with scutal 4ectal(lili)and4or5mesal(liliorliili)spines;tarsuswith4 areas poorlydefined, area I with 3 granularand 1 spiniform ectal (lili) and 5 mesal (lili) spines. Coxa IV with 2 dorsal tubercleeachside,areaIIwith 1 granulartubercleandalarge tubercles; posterior spiniform tubercle larger than anterior white patch each side, area III with 2 granular and 1 large granular tubercle; femora 111 IV with paired, dorsal apical spiniform tubercle each side, areas IV-V indistinct and tubercles; tarsal formula: 6:12-16:6:6-7; tarsal claws unpecti- unarmed (Fig. 7). Ocularium with large spiniform tubercle nate. Color ofscutum black to dark brown with orwithout and a smaller anterior granular tubercle each side (Fig. 8). white patches. Ventral plate of penis rectangular, elongate Anterior margin ofcarapace with 4 small granular tubercles — TOWNSENDETAL.—NEWMANAOSBIIDAEFROMCENTRALAMERICA 95 13 14 15 i'; — Figures 13-15. Barronafelgenhaueri,newspecies,male,paratype: v1i3.ew.TaSrcsaulseIb,arlsate=ral2mvimew;(Fi1g4.. P1e3)n;is2,50doprsmal(Fviigesw.;141,5.15P)e.nis, lateral Dorsum (Fig. 7): anterior margin of carapace with 4 granulartubercles on eachside; eyemound with a spiniform — tubercle and an anterior granular tubercle on each side typFei:gu7.reHsab7i-t1u2s.,doBrasrarlonvaiewf;el8g.enOhcauulerair,iunm,ewlatsepreaclievsi,ewf;em9.alTea,rshuosloI-, (pFaiigr.ed8);laarbgdeormignraalnuslcarututmubwerictlhes4dainstdinc6t asrmeaalsl;eraregaraInwuiltahr Tl1aatmresrmauls(vIFiiVeg,ws;.la19t0-e.r1a2Tl)a.rviseuws.IIS,callateerbaalrsvi=ew;21m1.mTa(rFsigu.s7I)I;I,0l.a3temramlv(iFeiwg;.182).; tVpuabierirnecddliessstp;iinncaitrfeoaarnmdIItusbwmeiortcohltehs2;agngrdraan4nuulglarararnuttulbuaebrrecrtlcuelbsee;srclaiernse;aaarreIeIaIasswIIiV-tV-h oneach side (Fig. 7). Cheliceral sockets ofcarapace shallow bearing small spines; lateral margins of abdominal scutum (Fig. 7). Cheliceral bulla smooth. Basal tarsal segments I of without tubercles. Free tergite I with pair of granular themaleswollenandspindle-like(Fig. 13). FreetergiteIwith tubercles; II with pair of median granular tubercles and 1 paired granular tubercles; II with paired spiniform tubercles granular tubercle on the margin each side; III with pair of eapaanicdrheds1igdgerraa(nnFuiulgla.arr7)t.tuubFbeerecrmlcuelrseaaonnnddtt1ihgberiaamnIauVrlgasirtnrtauiebgaehtrc.chlTeasirodsnea;lmaIfrIoIgrimwnuiltoahf msmheaardpgieiannaengadrcahbneusalidraeir;ngttuusbbpeeirrnccellsee.ssoAannnadflre1oeptgeerrragcniutulelsuamrsimwtiuilbtaehrrci8lnegsrioaznenualtnahdre 6:13:6:6. Tarsal claws III-IV unpectinate (Figs. 9-12). Penis: tuberclesbearingspines. ventral platewith lateral borders straight and parallel, distal Venter: coxae I-III with 1-2 rows of granular tubercles border slightly concave, uncleft; with third and fourth distal bearing spines, IV with scattered granular tubercles bearing curved spines flattened; glans without dorsal or ventral spines. process;stylusbentwithfoldedapex(Figs. 14, 15). Chelicera: smoothwithsparsesetae. Description. Female: Measurements (paratype, in mm): Pedipalp: trochanter length: 0.51 mm; femur length: dorsal scute length4.17; cephalothorax length 1.35; mesoter- 1.47 mm; patella length: 0.93 mm; tibia length: 1.21 mm; gum width 3.73; cephalothorax width 2.64; leg segments tarsus length: 1.22; total length: 5.34; coxa with one ventral (length):trochanterI:0.48;femurI:2.88;patellaI:0.83;tibia tubercle bearing a spine; trochanter with one mesal tubercle I: 1.59; metatarsus I: 2.95; tarsus I: 2.38; total leg I: 11.11; bearing a spine; femur and patella smooth; tibia ectal lili, trochanter II: 0.57; femur II: 6.15; patella II: 1.28; tibia II: mesal lili; tarsusectal lili, mesal lili. 4.25; metatarsus II: 5.46; tarsus II: 5.26; total leg II: 22.97; Legs (Figs. 9-12): coxa IV with 2 spiniform tubercles; trochanterIII:0.61;femurIII:4.58;patellaIII: 1.24;tibiaIII: trochanters witha retrolateral granulartubercle; femora I-II 2.30;metatarsusIII:4.54;tarsusIII: 2.35;totallegIII: 15.62; smooth, femora III IV with dorsal, apical spine on retro- trochanterIV:0.61;femurIV: 6.12;patellaIV: 1.39;tibiaIV: lateral surface, patellae-tarsi I-IV smoothwith sparsespines; 3.06;metatarsus IV: 6.61;tarsusIV: 3.14;totalleg IV: 20.93. tarsal formula: 6:12:6:6. ———— —— 96 THEJOURNALOFARACHNOLOGY Color:dorsumdarkbrown-black,withpairedwhitepatches on scutal groove and paired white patches on scutal area II, posterior patches larger than anterior ones; trochanters, patellae and chelicerae darker than pedipalps and femora; metatarsiannulate. Male: Measurements (in mm): dorsal scute length 4.39; cephalothorax length 1.57; mesotergum width 3.78; cephalo- thoraxwidth2.85;totallengthpedipalp:5.65;totallengthleg I: 12.49; total length leg II: 24.34; total length leg III: 16.58; total length leg IV: 21.84. Leg I: similar to female with the exceptionthatthe2mostbasalsegmentsareswollen(Fig. 13). LegsII-IVsimilartofemale. Tarsal formula: 6:13:6:6. Color: similartofemale. Genitalia (Figs. 14, 15): truncus long and slender; ventral plateelongate subrectanglar, tapering towardsdistal margin, withadistal borderentire, slightlyconcave(Figs. 14); lateral borderswith5straight-t-2recurvedsetae(Figs. 14, 15).Stylus straightwithapexfoldedand unarmed(Fig. 14). Habitat. Specimens were collected from vegetation and spaces beneath logs from hiking trails in montane rainforest on a moderate slope. They were found after dark between 2100-2300 hr in the dry season during light to moderate periodsofrainfall. BarranawilliamsiGoodnight&Goodnight 1942 (Figs. 4, 16-19) Barrona williamsi Goodnight & Goodnight 1942:11, fig. 26; Goodnight & Goodnight 1947:11, figs. 1, 2; Soares et al. 1992:4; Kury2003:207. Material examined. PANAMA: Code Province: Male, Parque Nacional General Division Omar Torrijos H., El Cope(08°49.2'80"N,80°05'45.7"W),23-28 February2007,V. Townsend, A. Savitzky and J. Ray, captured by hand along trails at night in montane rainforest (AMNH); Colon Prov- ince: male, Parque Nacional Soberania (09°07'55.3"N, 79°43T4.2"W), 1983, L. Sorkin (AMNH); Panama Prov- i7hn9ac°Dne3:ed8sc'br5emi5anp.let7eia5o,t"nhW.)lP,oagMrSsaqelaupenetdegmrebNnoeaicrctkaislo2idn0au0a9rl(,iFniggRSs.u.thmMe1mi6im-r1toa9rn).nd-ia(,n0T9gr°cu0(a3npA'ctM4uu1Nsr.eH0d)8l"o.bNny,g dpLieasntFtieaisrlg,aulprSoeEvrsiMtei:1wo6n-o116fo.9fd.—DitsohtrBeasalaprleprnooirvnstia;ieoww1ni9lolofLifaattmthesherieadGliposetvonaiidlsen;wipog1orh8f.ttitVo&henenGtodrofiaoslttdahnleviigtpeiehwpntioosf;1f94tt12hh7,ee. .: andslender;ventralplatedefinedasanelongatesubrectangle, stylus.Scalebars=50pm.Abbreviations:g=glanspenis,s=stylus, tapering towards distal margin, with a distal border entire, vp= ventralplate. slightlyconcave(Figs. 16, 17);lateralborderswith4straight-i- 3recurvedsetae(Fig. 18).Stylusstraightwithapexfoldedand 3 smallgranulartubercles,similarinsize. Abdominalscutum ' unarmed(Fig. 19). with 4 distinct areas; areas I and II unarmed orarmed with Remarks. This species was previously known only from paired granular tubercles; area III with paired spiniform I three specimens (female holotype, two male paratypes) tuberclesthatmayormaynotbeencircledbyaringofsmaller i collected at Barro Colorado Island, Canal Zone, Panama granulartubercles; areaIV-Vunarmed;anteriormarginwith (Goodnight & Goodnight 1942, Goodnight & Goodnight asinglemedianprocess. Lateralmarginsofscutumunarmed. 1947). Firstandthirdfreetergitesunarmed;secondfreetergitewitha Bugahilia Roewer 1915 median spiniformtubercle. Analoperculumsmooth. Pedipal- pal femurand patella unarmed; tibiawith4ectal (lili)and5 Bii&1g9ah2Si6ot:ai3ra5e7sR;o1Re9ow4ee9w:r2e3r119;1195K3:u11r:019y0;71;R99oM7ee:wl4;leorK-Lu1er9iy2t3a2:o05101839;:3220M:7e4.l0l4o;-LSeoiatraeos msfpeeismnaoelsr.a(lCiIioIlIxi)aIsVIpiVnwewisit;thhtap5rasiourrsedmw,oirtdheor4ssamelacltlaapltiuc(blaeillric)glreaasn,ndusli4amrimletasurabeilnrc(slliiezlsei;;) Type species. Biigahitia triacantha Roewer 1915, by tarsalformula:6:14-16:7:8;tarsalclawsunpectinate.Colorof originaldesignation. scutum dark brown, with yellow legs mottled with black; Emended diagnosis. Anterior margin ofcarapace with or spiniform tubercles on abdominal scutal area III and second withoutmedianspiniformtubercle.Oculariumwith2ormore freetergiteyelloworwhite,contrastingstronglywithdorsum. ' tubercleseachside,anteriorgranulartuberclesmallerorwith Metatarsus 1 with distal expansion near joint with tarsus. — TOWNSENDETAL.—NEWMANAOSBIIDAEFROMCENTRALAMERICA 97 maleswollenandspindle-like.FreetergitesIandIIIunarmed; IIwith 1 medianspiniformtubercle(Fig. 20).Femurandtibia IV straight. Tarsal formula 6:14-16:7:8. Tarsal claws III-IV unpectinate(Figs. 22-25). Penis: unknown. Description. Female: Measurements (holotype, in mm): dorsal scute length 3.53; cephalothorax length 1.32; mesoter- gum width 3.59; cephalothorax width 2.36; leg segments (length):trochanterI:0.50;femur1:4.43;patellaI:0.89;tibia I: 2.84; metatarsus I: 5.30; tarsus 1: 2.00; total leg I: 15.96; trochanter II; 0.72; femur II: 11.41; patella II: 1.17; tibia II: 8.81; metatarsus II: 10.67; tarsus II: 5.67; total leg II: 38.45; trochanterIII:0.80;femurIII: 7.59;patella111; 1.38;tibiaIII: 3.69;metatarsusIII: 6.98;tarsus III: 3.34;totallegIII: 23.78; trochanterIV;0.90;femurIV: 10.36;patellaIV: 1.59;tibiaIV: 5.16;metatarsus IV: 9.99; tarsus IV:4.23; total legIV: 32.23. Dorsum(Fig. 20):anteriormarginofcarapacewithmedian spiniformtubercle;eyemoundwithalargerspiniformtubercle and a smaller, anteriorgranular tubercleeach side (Fig. 21); abdominalscutumwith4distinctareas;area Ismoothwitha fewsparsespines;areaIIsmoothwithafewsparsespines;area III with paired spiniform tubercles not encircled by smaller tubercles at the base; areas IV-V smooth; lateral margins of abdominalscutumwithouttubercles.Freetergite1smooth;II withamedianspiniformtubercle;HIsmooth.Analoperculum smooth. Venter:coxaeI-IIIwithrowsofgranulartuberclesbearing spines, IVwithscatteredgranulartuberclesbearingspines. Chelicera: smoothwithmanysetae. Pedipalp: trochanter length; 0.37 mm; femur length: 1.55 mm; patella length: 0.70 mm; tibia length; 0.96 mm; tarsus length; 1.03 mm; total length: 4.61 mm; coxa, — Figures 20-25. Bugahitiaakini,newspecies,female,holotype:20. trochanter, femur, and patella smooth; tibia ectal lili, mesal Habitus,dorsalview;21.Ocularium,lateralview;22.TarsusI,lateral liili; tarsusectal lili,mesal lili. T(vFiaiergws.;us2213).I;V,ITamrlasmtuesr(aFlIiIg,sv.ilea2wt2.e-r2aS5lc)a.vlieew;bar2s4.=Tar2sumsmIII,(Filga.te2r0a)l;vi0e.2w;m25m. bbeeaaLrreiignnsggss(ppFiiinngeesss.;;2ft2er-om2co5h)ra:antIec-roIsIxaswmiotIhoVtfhew,wi,tfhesmmao5lrlaggrIraIanInu-ulIlaVarwritttuuhbbeearrccpllaeeissr ofdorsal,apicalgranulartubercles;patellae-tarsi1-IVsmooth BMaaslietagresnuistaIlioafumnaklneowsnp.indle-like; basal 3 segments swollen. witCholsopra:rsdeorspsiunmes;datrakrsablrofwonr,muwliat:h6l:i1g4h-t1e6r:,7y:8e.llowishmargins on abdominal scutum and free tergites; ocularium, paired Bugahitiaakini, newspecies tubercles on area HI, and single tubercle on free tergite H —(Figs. 1,20-25) yellow,contrastingstronglywithdorsum;legs,cheliceraeand Materialexamined. PANAMA; CodeProvince: Holotype pedipalpsyellowmottledwithblack. female,ParqueNacionalGeneralDivisionOmarTorrijosH., Male: Un—known. ElCope(08°49.2'80"N,80°05'45.7"W),23-28 February2007, Habitat, —SameasBarronafelgenhaueri. V.Townsend,A.SavitzkyandJ.Ray,collectedbyhandalong Remarks. Theholotypeandparatypedifferslightlyinsize hiking trails at night in montane rainforest (AMNH). and with respect to the morphology ofmetatarsus 1. In the ParEattyympoel:og1y.fe—maTlhei,scolslpeecciteesdwiisthahoplaottyrpoeny(mAMiNnH)h.onor of ehoxlpoatnydpee,d tihnecdiosmtpalarriesgoinonwoifththitshatlegofsegthmeentpariastnyoptei.ceTahbilsy Jonathan Akin who has made many contributions to the morphology resembles that of the male holotype of B. studyofnatural history and for his invaluable assistance on triacantha, which also has a spindled basitarsus. The priorfieldtr—ips. basitarsusofthefemaleholotypeofB. akiniisnotexpanded. Diagnosis. Dorsal scutum pyriform with scutal areas CranellusmontgomeryiGoodnight&Goodnight 1947a poorlydefined,areasI-II unarmed,area III with 1 spiniform (Figs. 26-29) aatursbeeaprsicnlIieVfo-erVamcihtnudsbiiesdrtecilnnecottaannedndcaiurnscmlaaerldmleebrdya(rnFitinegg.rio2of0r).sgmrOaaclnluuellraarrtiutubuembrecwrlicetslh,e Cra1n1e,ll1u2s;mKounrtygo2m0e0r3y:i20G7o;oTdonwignhsten&dGeotoadl.ni2g0h0t8a1:95497-a6:06,,ffiiggss.. each side (Fig. 21). Anterior margin of carapace unarmed 2h,j;Townsendet—al. 2008b:1027. (Fig.20). Cheliceral sockets of carapace very shallow Materialexamined. TRINIDAD,W.I.:6males,6females, (Fig.20). Cheliceral bulla smooth. Basal tarsal segments I of Lalaja Trace (10°44'28.3"N, 6ri6'17.3''W), July 2007, D. — — THEJOURNALOFARACHNOLOGY Figures 26-29.—CranelliismoiilgoiueryiRoewer1932,penis,SEM: Figures 30-33.—RliopalocranausalhilineatusGoodnight&Good- 26.Dorsalviewofthedistalportionofthepenis;27.Lateralviewof night 1947,penis,SEM: 30. Dorsalviewofthedistalportionofthe distalportionofthepenis;28.Ventralviewofthedistalportionofthe penis;31.Lateralviewofdistalportionofthepenis;32.Ventralview penis; 29. Lateral view ofthe distal tip ofthestylus. Scalebars = ofthedistalportionofthepenis;33.Lateralviewofthedistaltipof 50pm(Figs.26-28);25pm(Fig.29). thestylus.Sealebars= 50pm(Figs.30-32);25pm(Fig.33). hPMirokoriunndge,tcraaiBplltseuuirnedelRfbiiydngwhe,aonoddlNdaounrrdtihn(egrAntMhNeHRd)aa;nyg3ienmalle(ea1sf0,°l4i5t3t'efr5e2ma.al5lo"enNsg,, 2fi0g0s,8al:e5,9-f;60G,irifibgest. e2tf,alg.;2T0o0w9:n1s8e.nd etal. 2008b;1027-1029, 61°15.7'0.7"W),July2006, D. Proudand P. Resslar,captured Material examined. TRINIDAD, W.I.: 10 males, 10 byhand in leaflitteralonghikingtrail inmontanerainforest females, Mt. Tamana, Central Range (10°28T5.5"N, (AMNH). 61°11'50.5"W),July2008, M. MooreandJ.Toraya,captured Description.- A/rt/c genitalia (Figs. 26-29).- Truncus long by hand late in the afternoon in leaf litter from tropical and slender; ventral plate defined as an elongate rectangle, seasonal forest—(AMNH). tapering towards distal margin, with a distal border entire, Description. Male genitalia: Truncus long and slender; slightlyconcave(Figs. 26,27);lateralborderswith3straight+ ventral plate defined as an elongate rectangle, tapering 3recurvedsetae(Fig. 28).Stylusstraightwithapexfoldedand towards distal margin, with a distal border entire, slightly unarmed(Fig. 29). concave (Figs. 30, 31); lateral borders with 4 straight -f- 3 recurved setae(Fig. 32). Stylusstraightwithapexfoldedand RliopalocranausalhilineatusRoewer 1932 unarmed (Fig. 33). (Figs. 30-33) Remarks. Thisspeciesisverycommon intheleaflitterin Rliopalocranaus alhilineatus Roewer 1932:285, fig. 3; Good- most forested habitats island-wide. Individuals have been night & Goodnight 1947a:8; Gonzalez-Sponga 1991:205, capturedfromleaflitter,treebuttressesandfrombeneathlogs figs. 29-36; Burns et al. 2007:140; Townsend et al. and rocks. TOWNSENDETAL.—NEWMANAOSBIIDAEFROMCENTRALAMERICA 99 — Table 1. Interspecific variation in penis morphology among Manaosbiidae. Data for the South American species are based upon examinationsofpublishedfigures,micrographsordescriptions(Silhavy 1979,Kury 1997,2007). Shapeofthedistal Setaeonlateral Shapeofthe Species Shapeoftheventralplate borderoftheventralplate borderofventralplate apexofstylus Barronafelgenliaueri Elongate,rectangular Slightlyconcave 5straight+2recurved Folded Barronawilliamsi Elongate,rectangular Slightlyconcave 4straight+3recurved Folded Cranellusmontgomeryi Elongate,rectangular Slightlyconcave 3straight+3recurved Folded “Isocranaus”strinatii Elongate,rectangular Substraight 6straight Folded Manaosbiascopulata Veryelongate,rectangular Paraboliccleft 4straight+3recurved Folded Rhopalocranausalbilineatus Elongate,rectangular Slightlyconcave 4straight+3recurved Folded Rhopalocranausbordoni Elongate,rectangular Slightlyconcave 4straight+3recurved Folded Saramaciaalvarengai Elongate,rectangular Paraboliccleft 9straight Folded Saramaciaanmdata Elongate,rectangular Paraboliccleft 8straight Folded Saramacialuca.sae Elongate,rectangular Paraboliccleft 8straight Folded Syncranauscribrum Elongate,rectangular Substraight 4straight+3recurved Papillate — Naturalhistory. Littleisknownaboutthenatural history paredourobservationswithpublisheddescriptionsofgenital ofharvestmen from the family Manaosbiidae. In Trinidad, morphology for seven species from South America (Silhavy Townsend et al. (2008a) reported that Rhopalocranaus 1979; Kury 1997, 2007). With respect to overall appearance, albilineatusisahabitatgeneralistandexhibitsanisland-wide the penis morphology exhibited by Barrona spp. was most distribution. In contrast, Cranellus montgomeryi is a habitat similar to that of Rhopalocranaus spp. and Cranellus specialist, with a distribution limited to montane rainforest montgomeryi. and elfin woodland in the Northern Range. In montane However,weobservedrelativelylittleintragenericvariation rainforest, R. albilineatuswaspresent, butnotascommon as (Table 1) inpenismorphology. The only featuresthatvaried C. montgomeryi. In Panama, only the natural history of within the genera Barrona, Rhopalocranaus and Saramacia Zygopachylusalbomarginis has been examined (Rodriquez& were the numberand shape ofsetaeon the lateral borderof Guerrero 1976; Mora 1990). Duringthecourseofthisstudy, the ventral plate. Other characters associated with the penis wehadopportunitiestoobservemanaosbiids fromtwosites: includingtheshapeoftheventralplate,theshapeofthedistal Parque Summit, a lowland seasonal forest near the Canal borderofthe ventral plate, and the shapeoftheapexofthe Zone;andParqueGeneralDivisionOmarTorrijos,amontane styluswereconservativewithinagenus,butvariedamongthe rainforestnearElCope.AtParqueSummit,Barronawilliamsi genera that we compared (Table 1). Most taxa possess an is syntopic with Z. albomarginis. Individuals of both sexes elongate, rectangular ventral plate, with the exception of from each species were observed occupying refugia beneath Manaosbiascopulata, which hasa veryelongateventralplate logsduringtheday. Duringabriefone-daysurvey,twoadult (Kury 2007). Most species also have a stylus with a folded maleZ.albomarginiswereobservedresidingwithinmudnests, apex, with theexception ofSyncranaiiscribriim, which has a butnoeggs,nymphsorfemaleswereobservedinthesearenas. papillate stylar tip (Kury 1997). The penises of Manaosbia MaleB. williamsiwerefoundnearby,beneathadjacentlogsor scopulata and Saramacia spp. have a parabolic cleft in the inspacesbetweenthebarkandwoodoffallentrees.AtParque distalmarginoftheventralplate,incontrasttoothertaxa,in General Division Omar Torrijos, sampling occurred over a whichthemarginmaybeslightlyconcaveorevensubstraight periodofseveraldays,mostlyatnightbetween2000-2400h. (Table 1).Withrespecttootherfamiliesofharvestmenwithin Individualsoffourspecies,includingBarronafelgenliaueri,B. the Laniatores, variation in penis morphology within the williamsi, Bugabitia akini, and an undescribed species of Manaosbiidaeappearstoberelativelyconservative,similarto Zygopachyluswerecollectedfromthelitterandfrombeneath levels reported for the Cosmetidae (Kury et al. 2007; Town- logsandsmallrocks.Noindividualswereobservedoccupying send et al. 2010), and considerably less diverse than that mudnests;however,amale-femalepairofB.felgenliaueriwas observedfortheGonyleptidae(Kury&Pinto-da-Rocha2007) collectedfrombeneaththesamelog.Wedidnotobserveany orOncopodidae(Schwendinger& Martens2002). instances offeeding, reproductive behavior, or ectoparasites The functional significance of genital morphology has for the manaosbiids at Parque GD Omar Torrijos. All four received relatively little attention within the Gonyleptoidea species were found in the same microhabitat along either orthesuborder Laniatores. Currently, the functional aspects walGekniintgaltramiolsrpohrolfoorgeys.t—edWgiets.h respect to penis morphology, oOfncogepnoidtaildaemo(rSpchhowleongdyinghearve& Moanrlytenbsee2n002e)x.plored in the harvestmenofthefamilyManaosbiidaepossessamoderately ACKNOWLEDGMENTS long truncus, which is distally divided into a rectangular ventral plate and a dorsal distal-oriented glans that lacks This research was made possible with the support of a dorsal orventral processes and terminates in a stylus with a Battenendow'edprofessorship(VRT),VWCFacultySummer fthoeldpeedniosrmpoapriplhloatleogaypeoxf(BTaarbrloena1)f.elIgnentlhiisausetruiday,ndweB.dweislclriiabmesdi rDeesveealrochpmefnutnd.G,ranatnsd(VsRuTp)p,ortthefVrWomCAscNieAncMeunadnedrgrIaNdBuiaot.e from Central America and Cranellus montgomeryi and Assistance in the field was provided by A. Savitzky, J. Ray, Rhopalocranaus albilineatus from the Caribbean and com- C.Viquez,M.Gibbons,R.Miranda,S.Bermudez,andP.Van 100 THEJOURNALOFARACHNOLOGY Zandt. We thank H. Ring for help with the Leica image Kury,A.B.&R.Pinto-da-Rocha.2007.Gonyleptidae. Pp. 196-203. capturingsystemandthecollectionofmorphometricdata.We InHarvestmen:TheBiologyofOpiliones.(R.Pinto-da-Rocha,G. are especially grateful to J. Huff (AMNH), N. Platnick Machado & G. Giribet, eds.). Harvard University Press, Cam- J(aanAeoMgneNyrHm)(oS,uMsFL).r,evPJir.eeAwneldritsnmiafno(rnAiM(nSNsiMHgFh))t,;fulaDn.cdoQmAum.ienTntoteusrroionn(hMoaInUaPne)da,rtliwPeo.r Kurtbyrh,iedgAte.y,Bp.eM,assOps.eacVci.ehsuMsaoenftztaCsn.yinlolrata&(CA.raScahnmipdaai,o.Op2i0l0i7o.neRse,deCsocsrmiepttiidoane)o.f edrxapfotrtoefdthuinsdmearnussccirenitpitf.icSppeercmiimtennsucmoblelerctSeEd/iAn-1P1a-n07a.maSpweecrie- MelJalomoue-rrLnieacilatnaooofs,.ACrR.aeFcv.hisndtoea.lod1go9y26M3.u5s:N3eo2ut5a-Ps3a3u3sl.oibsrteaO1p4i:l3i2o7n-e3s83.Laniatoressul- mensfromCostaRicawereexportedunderscientificpassport Mello-Leitao,C.F.de. 1931.Opilioesnovosoucriticos.Archivesdo no. 01607. MuseudoNacionalRiodeJaneiro33:117-148. LITERATURECITED Mello-Leitao, C.F. de. 1932. Opilioesdo Brasil. Revistado Museu Paulista 17:1-505. Acosta, L.E., A. Perez-G & A.L. Tourinho. 2007. Methods for Mello-Leitao, C.F. de. 1935. Algumas notas sobre os Laniatores. taxonomic study. Pp. 494-510. In Harvestmen: The Biology of ArchivesdoMuseudoNacionalRiodeJaneiro36:87-116. Opiliones. (R. Pinto-da-Rocha,G. Machado&G. Giribet,eds.). Mello-Leitao, C.F. de. 1940. Un solifugo da Argentina e alguns HarvardUniversityPress,Cambridge, Massachusetts. opilioesdaColombia.AnnaisdaAcademiaBrasileiradeSciencias Burns,J.A., R.K. Hunter& V.R. Townsend, Jr. 2007. Treeuseby 12:301-311. harvestmen(Arachnida:Opiliones)intherainforestsofTrinidad, Mora, G. 1990. Paternal care in a Neotropical harvestman, W.I.CaribbeanJournalofScience43:138-142. Zygopachyhtsalhomarginis(Arachnida,Opiliones: Gonyleptidae). Chamberlin, R.V. 1925. Diagnoses of new American Arachnida. AnimalBehaviour39:582-593. GiriBbulelte,tiGn.of&theA.MBu.seKuumryo.fC2o00m7p.araPthiyvteogZeonoyloagnyd67b:i2o1g1e-o2g4r8a.phy. Rodcrogmupeozr,taCm.iAe.nto& dS.e GZuyegrorpearcoh.yli1i9s76.alLhaomahrigsitnoirsia (nCahtaumrbalerlyine)l Pp. 62-87. In Harvestmen: The Biology ofOpiliones. (R. Pinto- (Arachnida:Opiliones:Gonyleptidae). 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GooCdonliogrhatd,oC.IJs.la&nd,M.LC.anGaolodZnoingeh.t.A1m9e4r2ia.caPnhalMaunsgieduamfrNoomviBtaartreos RoeTMwouersrien,io,Cd.iNFS.Zoo314l9:o21g5-.i3a4O.epdiliAonnaitdoenmiaausCoSmipiadr-aAmtearidkeal.laBUonlilveetrtisniotadedii GGGooooooGf1dddua1nnniu9iii8naggg:anhhh1a-ttto.1,,,f8A.TCCCmr...eJJiJ...nrii&&&dcaadMMMn....LLLAM...mueGsGGrooeoioooucdddmannnniiiNgggoMhhhvttuti...stae111tu999e44m4s772abbN1...o1v6SSP7itth:utua1dadl-iita1eee3snss.gooi1nnd3a5tt1hhf:ee1r-opp1hhm3aa.llBraainntggiiisddh RRooeeWsWwweeeeenbbsrreec,,rrhkkaCCfnn..teeFlFcc.i.hhctth1e1ee9n933d21dVe..eerrrWEeWerieEidnirtetde,zeerur,e1e9B2Wr13We9e)eb2.mb3ee)eAr.nrrkckn2hnAe8iebc:vch1hh0atf1tinei-edr1lVN6VuIa4.ntE.ugr(e(5g7n.e.sEcEdhrreigrgcaahnntNzzeau,utnunBgrgewriddlseei-rnr JimfP,1-aruo2Rs1n.ta.Ly.goSnf.inCa&eentdHro.alEB.rMAam.seilrSio(caOarp.eisl.iAomne1e9rs9,i1c.GaonnNyoMlveuopstiegdueamnee)r.NooRveeivtiaeststepasecBi1re3a4sd0i:e- RRooees1ww:mee2urr7s,,5e-3uCC5.m.0FF..i.n1S19t94o338.c.kWheoOipltimel.rieoAnreWkseibvaeurfsokrndeZcoeohmltoegNiaX,tLuSrthUiobscetkorhroiGlsomcnhy3el0neBp:tRlie-di8ec.nh.- Jubleeritrhaied,eECn.tom1o9l7o0.giaIX3.5:7O9p9i-l8i0o2n.s des Galapagos: Galanonuna RoeSweenrc,kenCb.eFr.gi1a9n5a7.26A:r1a2c-6h8n.ida arthrogastra aus Peru III. Senck- mPipc.ro1p3h7t-l1ia5l3n.iaMigsesni.onnZovo.olosgp.iquneov.Bei(gGeonayulxepitliesdaGea)l.apVaoglousmeete2n. Roeewnebre,rgiCa.nFa.B1i9o6l3o.giOcpail3i8o:n6e7s-94a.us Peru und Colombia (Arachnida REocyuaaldoMru(sNe.uemtJf.orLeCleenutpr,al19Af6r4i-c1a9,65B)r.us(sNel.s.Leleup,ed.).Tervuren SchAwretnhdrionggaesrt,raPa.uJs.P&eruJ.V).MaSretnecnkse.nbe2r0g0i2a.naPeBniioslogmiocrap4h4o:l4o5-g7y2.in Kury, A.B. 1997. The genera Sanunacki and Syncranaiis Roewer, Oncopodidae (Opiliones, Laniatores): evolutionary trends and with notes on the status of the Manaosbiidae (Opiliones, relationships.JournalofArachnology30:425^34. Laniatores,Gonyleptoidea). BoletimdoMuseoNacional,Zoolo- Silhavy. 1979.OpilionidsofthesuborderGonyleptomorphifromthe gia374:1-22. Americancaves,collectedbyDr. PierreStrinati. RevueSuissede Kury,A.B.2003.AnnotatedcatalogueoftheLaniatoresoftheNew Zoologie86:321-334. World (Arachnida, Opiliones). Revista Iberica de Aracnologia, Simon, E. 1879. 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(Opiliones, Gonyleptidae, KeytotheCosmetidae(Arachnida,Opiliones)ofCentralAmerica, Prostygninae).RevistaBrasileiradeEntomologia36:1-14. withnotesonpenismorphologyandsexualdimorphisms.Zootaxa Soares, H.E.M. 1970. Novas especies de opilioes da Regiao 2414:1-26. Amazonica (Opiliones, Cosmetidae, Gonyleptidae, Phalangiidae, Townsend,V.R.Jr.,D.N.Proud&M.K.Moore.2008a.Harvestmen Stygnidae). RevistaBrasileiraBiologia30:323-338. (Opiliones)ofTrinidad,West Indies. LivingWorld2008:53-65. SoaGroens,yleHp.tiEd.aMe., P1h9a72l.angOipidearea, OPphialliaonlgoogdiicdaae)V.ariRaeviIIs.ta(OBprialsiiolneeisr:a TowBnusrensn,d,R.VK..R.HuJnrt.,erD,.NS..R.PrLoauzda,roMw.iKtz.&MoBo.rEe.,FeJl.gA.enThiabubeert.ts2,00J8.bA.. deBiologia32:65-74. Parasiticand phoretic mitesassociated with neotropical harvest- Sorensen, W.E. 1932. Descriptiones Laniatorum (Arachnidorum menfromTrinidad,W.1.AnnalsoftheEntomologicalSocietyof OKpaiilLi.onHeunmriSkusbeonr.diDneits)K.onOgpeluisgepDoasntshkuemVuimdenresckoagbneorvnietsSetelsekdaibdst WilAlmeemarritc,aR1.0H1.:,1A0.26P-e1r0e3z2-.G,J.Farine&P.Gnaspini.2010.Sexually skrifter, Kobenhavn, Naturvidenskabeligog MathematiskAfdel- dimorphic tegumental gland openings in Laniatores (Arachnida, ing,ser.93:197-422. Opiliones), with newdataon 23species. JournalofMorphology Strand,E. 1942.MiscellaneanomenclatoriazoologicaetPaleontolo- 271:641-653. gicaX.FoliaZoologicaetHydrobiologica 11:386-402. Manuscriptreceived27March2009, revised3October2010.

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