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Reign of the Red Queen: the future of bats hangs in the balance

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Title:
Reign of the Red Queen: the future of bats hangs in the balance
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US National Park Service
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J. Judson Wynne ( suggested by )
Wynne, J. Judson
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Excerpt: "Maligned in old wives' tales for "getting in your hair" and wrongly considered a primary vector for rabies, bats are in fact among the most important contributors to Earth's ecosystems. They are a keystone species in cave ecosystems, their guano adding allochthonous nutrients to an otherwise energy-limited environment; they are important seed dispersers, particularly in tropical forests; and they provide critical ecological services to humankind. In this latter role, insectivorous bats are one of the most overlooked yet important animals. In the United States alone, bats provide between $4 and $50 billion annually in pest-control- related ecological services to agriculture. Other bat species are important pollinators for agave-the plant used for making tequila. Yet, North American bat colonies are facing a crisis with the westward advance of white- nose syndrome (WNS), a disease responsible for the deaths of nearly seven million bats since it was first detected in Howe Cave, near Albany, New York, during the winter of 2006- 2007. Since then, WNS has been confirmed in 23 states and 5 Canadian provinces. WNS is a disease caused by the cold-loving fungus, Pseudogymnoascus destructans, which infects the skin of the ears, snout, and wings of hibernating bats. During hibernation, a bat's immune response is in a suppressed state as are other metabolic functions, enabling the fungus to spread relatively unchecked. When fully expressed, WNS often presents as a prominent white fungal growth. The fungal hyphae penetrate deeply into the connective tissue and cause severe damage..."
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Description
Excerpt: "Maligned in old wives' tales for "getting in
your hair" and wrongly considered a primary vector for rabies,
bats are in fact among the most important contributors to
Earth's ecosystems. They are a keystone species in cave
ecosystems, their guano adding allochthonous nutrients to an
otherwise energy-limited environment; they are important seed
dispersers, particularly in tropical forests; and they provide
critical ecological services to humankind. In this latter role,
insectivorous bats are one of the most overlooked yet important
animals. In the United States alone, bats provide between $4
and $50 billion annually in pest-control- related ecological
services to agriculture. Other bat species are important
pollinators for agave-the plant used for making tequila. Yet,
North American bat colonies are facing a crisis with the
westward advance of white- nose syndrome (WNS), a disease
responsible for the deaths of nearly seven million bats since
it was first detected in Howe Cave, near Albany, New York,
during the winter of 2006- 2007. Since then, WNS has been
confirmed in 23 states and 5 Canadian provinces. WNS is a
disease caused by the cold-loving fungus, Pseudogymnoascus
destructans, which infects the skin of the ears, snout, and
wings of hibernating bats. During hibernation, a bat's immune
response is in a suppressed state as are other metabolic
functions, enabling the fungus to spread relatively unchecked.
When fully expressed, WNS often presents as a prominent white
fungal growth. The fungal hyphae penetrate deeply into the
connective tissue and cause severe damage..."



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Troglomorphicpseudoscorpions(Arachnida:Pseudoscorpiones)ofnorthernArizona,withthedescription oftwonewshort-rangeendemicspecies MarkS.Harvey 1,2,3,4,5 and J.JudsonWynne 6 : 1 DepartmentofTerrestrialZoology,WesternAustralianMuseum,Locked Bag49,WelshpoolDC,WesternAustralia6986,Australia; 2 ResearchAssociate,DivisionofInvertebrateZoology, AmericanMuseumofNaturalHistory,CentralParkWestat79thStreet,NewYork,NewYork10024-5192,U.S.A; 3 ResearchAssociate,CaliforniaAcademyofSciences,55MusicConcourseDrive,SanFrancisco,California94118, U.S.A; 4 Adjunct,SchoolofAnimalBiology,UniversityofWesternAustralia,Crawley,WesternAustralia6009, Australia; 5 Adjunct,SchoolofNaturalSciences,EdithCowanUniversity,Joondalup,WesternAustralia6027, Australia; 6 DepartmentofBiologicalSciences,ColoradoPlateauBiodiversityCenter,LandscapeConservation Initiative,NorthernArizonaUniversity,Box5640,Flagstaff,Arizona86011,U.S.A.E-mail: mark.harvey@museum.wa.gov.au Abstract. ThisstudyreportsonthepseudoscorpionfaunaofthesubterraneanecosystemsofnorthernArizona,U.S.A. Ourworkresultedinthedescriptionsoftwonewspecies, Hesperochernesbradybaughi sp.nov.and Tuberochernescohni sp. nov.(Chernetidae)andtherangeexpansionofonespecies, Larcacavicola (Muchmore1981)(Larcidae).Allofthesespecies werecave-adaptedandfoundwithincavesonGrandCanyon-ParashantNationalMonumentinnorthwesternArizona. Baseduponthiswork,thegenus Archeolarca HoffandClawsonisnewlysynonymizedwith Larca Chamberlin,andthe followingspeciesaretransferredfrom Archeolarca to Larca ,formingthenewcombinations L.aalbui (Muchmore1984), L.cavicola (Muchmore1981), L.guadalupensis (Muchmore1981)and L.welbourni (Muchmore1981).Despiteintensive samplingonthemonument,thetwonewspeciesweredetectedinonlyonecave.Thiscavesupportsthegreatestdiversityof troglomorphicarthropodspeciesonthemonumentallofwhichareshort-rangeendemicsoccurringinonlyonecave. MaintainingthemanagementrecommendationsprovidedbyPeckandWynne(2013)forthiscaveshouldaidinthelongtermpersistenceofthesenewpseudoscorpionspecies,aswellastheothertroglomorphicarthropods. Keywords: Nearctic,troglomorphy,troglobite,newsynonymy,cave urn:lsid:zoobank.org:pub:A15CB9DB-5B36-4A7C-8052-08E2EC1F4D34 ThepseudoscorpionfaunaofNorthAmericancavesis moderatelywellknown,thankslargelytotheeffortsofJ.C. Chamberlin,C.C.Hoff,E.M.Be nedict,D.R.MalcolmandW.B. Muchmorewhohavecharacterize danddescribedmanydifferent NorthAmericantroglobitesandtr oglophiles.Therearecurrently 144namedspeciesfoundincave habitatsacrosstheUnited StatesincludingsixspeciesinfivefamiliesfromArizona: Pseudogarypushypogeus Muchmore1981(Pseudogarypidae), Albiorixanophthalmus Muchmore1999(Ideoroncidae), Chitrellinachiricahuae Muchmore,1996(Syarinidae), Archeolarca cavicola Muchmore1981, A.welbourni Muchmore1981 (Larcidae)and Tuberochernesubicki Muchmore1997(Chernetidae)(Muchmore1996;Muchmore&Pape1999;Harvey& Muchmore2013).Only A.anophthalmus and C.chiricahuae had troglobiticmodificationsincludingthecompletelackofeyes andpallidbodycolor(Muchmore1996;Muchmore&Pape 1999;Harvey&Muchmore2013),whereastheothersareless obviouslymodifiedwithonlytheslightlyattenuatedappendageshintingatanobligatesubterraneanexistence(Muchmore 1981,1997). Priortothiswork,allofthesecavernicolousspeciesfrom ArizonaoccurredsouthoftheColoradoRiverwith P. hypogeus A.cavicola and A.welbourni fromnorthernArizona (CoconinoCounty)and A.anophthalmus C.chiricahuae and T.ubicki fromsouth-easternArizona(Pima,Cochiseand SantaCruzCounties,respectively).DuringbiologicalinventoriesofcavesontheGrandCanyon-ParashantNational Monument(hereafterreferredtoasParashant)innorthwesternArizona,oneofus(J.J.W.)andcolleaguesfound representativesofthreedifferentpseudoscorpionspecies, whicharethesubjectofthisstudy. Overthepastseveralyears,Parashantcaveshaveyielded othersignificantandinterestingarthropodspeciesmanyof whicharerestrictedtothecaveenvironment.Theseinclude twonewgenera(comprisingtwonewspecies)abooklouse (orderPsocoptera,familySphaeropsocidae: Troglosphaeropsocusvoylesi Mockford2009(Mockford2009),andacave cricket(familyRhaphidophoridae:cf. Ceuthophilus n.gen.n. sp.,CohnandSwanson,unpublisheddata).Thisworkalso resultedintheidentificationofseveralcave-adaptedandcavelimitedspeciesincludingaleiodidbeetle, Ptomaphagus parashant PeckandWynne2013(Peck&Wynne2013),an undescribedspeciesofcentipede(familyAnopsobiidae; Wynne,unpublisheddata),anundescribedIsopodspecies, Brackenridgia n.sp.(S.Taiti,inlitt.),andarecentlydescribed cavelimitedmillipede, Pratherodesmusvoylesi Shear2009 (Shearetal.2009).Additionally,threenewspeciesof trogloxenicbeetleswerereportedfromParashantcaves including Eleodeswynnei Aalbu,Smith,andTriplehorn2012 (Tenebrionidae;Aalbuetal.2012),anundescribedspeciesof thecarabidbeetlegenus Rhadine LeConte(Carabidae:the perlevis species-group;T.C.Barr,inlitt.),andanundescribed carabidbeetlespecies Pterostichus Stephens(Carabidae,K. Will,inlitt.). METHODS ThejuniorauthorandcolleaguessampledcavesonGrand Canyon-ParashantNationalMonumentduring414August 2014.TheJournalofArachnology42:205219 205

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2005,16May2007,1625August2007,1221May2008, and512March2009.Theysampledallcavesidentifiedas havingdeepzonelikeconditions( n 5 10).Giventheshort durationofstudy(betweentwotofoursitevisits),and potentialseasonaleffects,confidentlyidentifyingthiszonal environmentwasnotpossible.Thecavedeepzoneisrequired habitatforcave-adaptedarthropodsandischaracterizedby completedarkness,stabletemperature,anear-watersaturated atmosphereandlimitedtonoairflow(asinHowarth1980, 1982).ParashantislocatedinnorthwesternArizona,encompassesapproximately4,451km 2 ,andischaracterizedby ruggedterraincontainingdeeplyincisedcanyons,mesas,and mountains.VegetationzonesincludeMojaveDesertcontainingcreosotebush( Larreatridentata )andJoshuatrees( Yucca brevifolia )atlowerelevations,gradatingthroughGreatBasin pinyon( Pinusedulis )andjuniper( Juniperus spp.)woodlands toColoradoPlateaugrasslandsandPonderosapine( Pinus ponderosa) forestwithaspen( Populustremuloides )groveson Mt.Trumbull(elevation2,447m).Allofthecavesreferredto inthispaperwerelocatedwithintheSupai,Kaibab,or Redwalllimestoneformations.Elevationforthecavesthat werestudiedrangesfrom736to1,590m. Althoughweinventoried10Parashantcaves,weprovide descriptionsforonlythethreecaves(PARA-1001,PARA2204andPARA-3503)wherepseudoscorpionsweredetected. PARA-1001Cavewasthesecondmostbiologicallydiverse caveonParashant(Wynne,unpublisheddata),andsupports thelargestknowncricketroostinnorthernArizona(Wynne& Voyles2014).AsmallsolutioncavewithintheKaibab limestone,ithadatotalsurveyedlengthanddepthof76.2m and10.4m,respectively.Thiscavehadasmallsouth-facing verticalentrance(135 u aspect)atbottomcenterofalarge sinkhole.Vegetationwascharacterizedasjuniperscrublands at1,585melevation,andwaslocatedonthenorthsideofthe lowerColoradoRiveralongthewesternextentoftheGrand Canyon.PARA-2204Cavewasthemostbiologicallydiverse caveonthemonument(Wynne,unpublisheddata).The deepestextentofthiscavecontainedactivespeleothem formationsandsupportedanear-saturatedwateratmosphere year-round.LocatedwithintheSupaiformation,thislarge solutioncave(totalsurveyedlength175m)wascomprisedof severalsinuousphreaticpassages.Thiscavehasonehorizontalentrance(330 u aspect)andwassituatedwithinacanyon nearthebaseofthecanyon'snorth-face.Locatedat1,272m elevation,thiscaveoccurredwithinthevegetationtransition zoneofMojaveDesertscrubandjuniperwoodlands.PARA3503Cavewasadrycavewithnoevidenceofrecent speleothemactivity,andsupportedasummerroostofbats, Myotis sp.(Wynne,unpublisheddata).Thecavehadalarge horizontalentrance(135 u aspect)situateduponahighbench (1,102melevationonanexposedcliffface).Thiscavewas situatedalongthesouth-faceofoneofthelargestcanyons drainingintotheColoradoRiverfromthenorth.Occurring withintheRedwallformation,thislargesolutioncave contained540mofsurveyedlengthwithanestimatedsurvey depthof14.2m.VegetationwascharacterizedasMojave Desertscrub. Theworkconductedin2005waspartofabiological baselinestudy[refertoWynne&Voyles(2014)fora descriptionofsamplingmethods].Later(between2007and 2009),thesecavesweresystematicallysampledtocharacterize thecave-dwellingarthropodcommunities.Intervalsampling usingbaitedpitfalltraps,timedsearches,andopportunistic samplingtechniqueswereused.Toapplythesetechniques, detailedmapsforeachcavewererequired.Forinterval sampling,weestablishedupto10samplingintervals(which includedasamplingstationateitherwallandoneatcave centerlineforatotalof # 3samplingstationsperinterval).We used10 % ofthetotalcavelengthtoestablishthesampling interval(e.g.,fora1,000mlongcave,thesamplinginterval wasevery100m). Ateachsamplingstation,wedeployedlivecapturebaited pitfalltrapsandconductedtimedsearches.Forpitfalltraps, weusedtwo907gstackedplasticcontainers(13.5cmhigh, 10.8cmdiameterrimand8.9cmbase).Ateaspoonofpeanut butterwasusedasbaitandplacedinthebottomofthe exteriorcontainer.Atthebottomoftheinteriorcontainer,we madeseveraldozenholessothebaitcouldbreathe''to attractarthropods(e.g.,Barber1931).Attemptsweremadeto counter-sinkeachpitfalltrapwithinthecavesedimentor rockfall.Whenthiswasnotpossible,webuiltrampsaround eachtrapusinglocalmaterials(e.g.,rocks,woodendebris, etc.)soarthropodscouldaccessthetrapandfallin(e.g., Ashmoleetal.1992).Todiscouragesmallmammals,we placedsmallrocksaroundtheedgesofthetrapandthen coveredthemouthofthetrapwithacaprock.Pitfalltraps weredeployedforthreetofourdays(athreedaydeployment occurredonceduetoschedulingconstraints).Fortimed searches,weestablisheda1mradiusaroundeachsampling station(wherethepitfalltrapwouldbedeployed)and searchedforarthropodswithinthat 3mcircle.Aoneto threeminutetimedsearch(oneminuteifnoarthropodswere observed,threeminutesifarthropodsweredetected)was conductedbeforepitfalltrapdeploymentandpriortotrap removal. Opportunisticcollectingwasexecutedbytwotothreetrained observersastheytraversedthelengthofeachcave.This techniquewasappliedastheobserverswereintransitbetween samplingintervalswhiledeployingandremovingpitfalltraps andconductingtimedsearches.Opportunisticcollectingwas notconductedwhileatsamplingstationsandwasresumedonly whentheobserverswereintransitonceagain.Thistechnique wasusedatleasttwicepercave(bothduringpitfalltrap deploymentandretrievaltrips).Forexample,acavecontaining 10samplestationarrays,therewere27individualrandom walks''persitevisit(i.e.,ninerandomwalksamplestimesthree observerscollectingalongtheirbetweenstations).Becausewe conductedtwositevisitspercave,therewouldbeatotalof54 samples.Foronecave,PARA1001Cave,wehadtwoobservers conducttheopportunisticcollecting. Analpha-numericcodingsystemdevelopedbytheNational ParkService(NPS)wasusedtosafeguardthelocationofboth cavesandtheirresources.Weonlyprovidegeneralizedlatitude andlongitudecoordinatesoftheareatokeeptheprecise locationofthecaveconfidential.ParashantNationalMonumentheadquartersinSaintGeorge,Utahhastheciphertable withcavecodes.Acopyofthispaperwithactualcavenames isonfileatbothmonumentheadquarters,NationalPark ServiceandtheNationalCaveandKarstResearchInstitute, Carlsbad,NewMexico. 206 THEJOURNALOFARACHNOLOGY

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Specimensrepresentingthreespeciescollectedbyoneofus (J.J.W.)andcolleaguesformthebasisofthisstudy.All specimenswerecollectedandstoredin70 % ethanol.The holotypesofbothnewspeciesandspecimensoftheknown speciesaredepositedintheMuseumofNorthernArizona, Flagstaff,Arizona(MNA).Temporaryslidemountswere preparedbymountingthemonmicroscopeslideswith10or 12mmcoverslipssupportedbysmallsectionsof0.25,0.35or 0.50mmdiameternylonfishinglineinadropoflacticacidat roomtemperaturefortwoormoredays.Afterstudythe specimenswererinsedinwaterandreturnedto75 % ethanol withthedissectedportionsplacedin12 3 3mmglassgenitalia microvials(BioQuipProducts,Inc.).Allspecimenswere studiedusingaLeicaDM2500compoundmicroscopeand illustratedwiththeaidofadrawingtube.Measurementswere takenatthehighestpossiblemagnificationusinganocular graticule.Terminologyandmensurationmostlyfollow Chamberlin(1931),withtheexceptionofthenomenclature ofthepedipalps,legsandwithsomeminormodificationsto theterminologyofthetrichobothria(Harvey1992),cheliceral setation(Harvey&Edward2007),cheliceralrallum(Judson 2007)andfacesoftheappendages(Harveyetal.2012). TAXONOMY FamilyLarcidaeHarvey1992 Larca Chamberlin1930 Larca Chamberlin1930:616. Archeolarca HoffandClawson1952:23. Syn.nov. Typespecies Larca:Garypuslatus Hansen1884,by originaldesignation. Archeolarca:Archeolarcarotunda HoffandClawson1952, byoriginaldesignation. Remarks. Thegenus Larca wascreatedbyChamberlin (1930)forthetypespecies L.lata (Hansen)fromEuropeand L.granulata (Banks1891)fromeasternU.S.A.Sincethen, otherspecieshavebeenaddedfromEurope(Beier1939a; Gardini1983;Henderickx&Vets2002;Zaragoza2005)and NorthAmerica(Hoff1961;Benedict&Malcolm1978; Muchmore1981). Archeolarca wasdescribedforthetype species A.rotunda whichwascollectedfrompackratmiddens andporcupinenestsinUtah(Hoff&Clawson1952).Since then,fouradditionalspecieshavebeendescribedfromother partsofwesternNorthAmerica,allfromcaveecosystems (Muchmore1981,1984),and A.rotunda hasbeenfoundin NewMexicoandOregon(Hoff1956a;Benedict&Malcolm 1978). Archeolarca onlydiffersfrom Larca inthepossessionof fourtrichobothriaonthemovablechelalfingerofadults, whereasspeciesof Larca haveonlytwoorthreetrichobothria (e.g.Hoff1961;Benedict&Malcolm1978;Muchmore1981; Gardini1983;Muchmore1984,1990;Henderickx&Vets 2002;Zaragoza2005).Mostadultspecimensfromthe Parashanthavefourtrichobothriaonthemovablechelal finger(Fig.12),consistentwithbeingaspeciesof Archeolarca butonemalehasfourontherightchelaandthreeontheleft (Fig.11)raisingtheissueofwhetherthegenerashouldbe retained. Themaintenanceofgarypoidgenerabasedsolelyon trichobothrialnumberhasbeenabandonedforseveralother groupsincludingthegarypidgenera Anagarypus Chamberlin 1930withseventrichobothriaonthefixedfingerandoneor twoonthemovablefingerformingapatternof7/12 (Muchmore1982), Eremogarypus Beier1955,withapattern of58/13(e.g.,Beier1962;Beier1973), Synsphyronus Chamberlin1930,withapatternof58/13(e.g.,Chamberlin 1943;Harvey1987b,2011)and Thaumastogarypus Beier1947, withapatternof78/34(e.g.Beier1947;Mahnert1982),and thegeogarypidgenus Geogarypus Chamberlin1930inwhich adultsnormallyhavean8/4pattern,but G.bucculentus Beier 1955and G.connatus Harvey1987havea7/4pattern(Harvey 1986,1987a).Intra-specificvariationinthenumberof trichobothriaofthemovablechelalfingerhasbeenreported inthegenus Serianus Chamberlin1930(Garypinidae).Hoff (1950)foundthatasmallseriesofspecimensof S.minutus Hoff1950(nowknownas S.argentinae Muchmore1981due tosecondaryhomonymyoftheoriginalname)includedadults withthenormalfourtrichobothriaonthemovablechelal finger,aswellassomewithonlytwoorthreetrichobothria. Similarly,Mahnert(1988)foundthatthetypeseriesof Paraserianusbolivianus Beier1939possessedthreeorfour trichobothriaonthemovablechelalfinger.Giventhatthe mainfeatureusedtosubstantiatethegenus Paraserianus by Beier(1939b)wasthepresenceofonlythreesuchtrichobothria(asopposedtofourin Serianus ),Mahnert(1988)placed Paraserianus asasynonymof Serianus Comparisonofspecimensofmanyspeciesof Larca and Archeolarca byoneofus(M.S.H.),includingthetypespecies ofbothgenera,hasrevealednoothersignificantdifferences thatcouldbeconsideredtomaintaindistinctgenera,and Archeolarca ishereregardedasasynonymof Larca ,resulting inthefollowingnewcombinations: L.aalbui (Muchmore 1984), comb.nov. L.cavicola (Muchmore1981), comb.nov. L. guadalupensis (Muchmore1981) comb.nov. and L.welbourni (Muchmore1981) comb.nov Larcacavicola (Muchmore)comb.nov. (Figs.114) Archeolarcacavicola Muchmore1981:5556,Figs.11,12. Materialexamined. U.S.A.: Arizona :MohaveCounty:1 male,PARA-3503Cave,GrandCanyon-ParashantNationalMonument,ca.UTM0247400N,4020000E,Zone12S, baitedpitfalltrap1A,20May2008,J.J.Wynne(MNA);1 female,samedataexceptbaitedpitfalltrap1C,6March 2009,J.JWynne(MNA);1tritonymph,1deutonymph, samedataexcepttrap2B,10March2009,J.J.Wynne (MNA);1tritonymph,samedataexcepttrap7A(MNA);1 tritonymph,samedataexcepto pportunisticcollectingina possibledeepzone(MNA);1male,PARA-2204Cave, GrandCanyon-ParashantNationalMonument,ca.UTM 025100N,4041000E,Zone12S,M,baitedpitfalltrap2B, 17May2008,J.J.Wynne(MNA);1female,samedata except20May2008(MNA);1tritonymph,samedata excepttrap1A(MNA);1male,samedataexcepttrap1B (MNA). Diagnosis. Larcacavicola resemblestheotherspecies previouslyincludedinthegenus Archeolarca inpossessing fourtrichobothriaonthemovablechelalfinger,butoccasionallythisisreducedtothreetrichobothria.Itdiffersfromthese speciesbyhavingreducedeyes,especiallytheposteriorpair, whicharenoticeablysmallerthantheanteriorpair. HARVEY&WYNNETROGLOMORPHICPSEUDOSCORPIONSFROMARIZONA 207

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Description Adults: Color:carapace,pedipalpsandcoxae deepred-brown,abdomenpalered-brownandlegspale yellow-brown. Chelicera:with4setaeonhand,with sbs absent,and1 subdistalsetaonmovablefinger(Fig.7);allsetaeacuminate; seta bs slightlyshorterthanothers;with2dorsallyrifissures and1ventrallyrifissure;galeaof L and K verylongwith3 terminalrami,ramiofmalesmallerthanonfemale;rallumof 4blades,themostdistalbladewithseveralserrationson leadingedge,otherbladessmooth;serrulaexteriorwith14( L ), 16( K )blades;laminaexteriorpresent. Pedipalp(Fig.9):mostsurfacesoftrochanter,femur, patellaandchelalhandlightlyandsparselygranulate,chelal fingerssmooth;trochanter,femur,patellaandchelalhand withprominent,curved,slightlydenticulatesetaearranged sparsely;patellawith3smallsub-basallyrifissures;trochanter 1.831.99( L ),1.901.93( K ),femur4.745.94( L ),4.574.95( K ), patella3.634.47( L ),3.693.94( K ),chela(withpedicel)4.47 5.28( L ),4.084.54( K ),chela(withoutpedicel)4.225.02( L ), 3.854.26( K ),hand(withpedicel)2.172.49( L ),1.942.08( K ) 3 longerthanbroad,movablefinger(withpedicel)0.961.01 ( L ),0.991.00( K ) 3 longerthanhand.Fixedchelalfingerwith 8trichobothria,movablechelalfingerwith4trichobothria (Fig.12),although sb absentfromleftchelaofonemale (Fig.11): eb esb ib and ist situatedsubbasally, est,isb and it submedially, et subdistally,and est opposite it ; b and sb situatedsubbasally,and st and t situatedsubmedially,with st situatedverycloseto t ;patchofmicrosetaenotpresenton externalmarginoffixedchelalfingernear et .Venom apparatuspresentinbothchelalfingers,venomductsfairly short,terminatinginnodusramosusslightlydistalto et in fixedfinger(Figs.11,12).Chelalteethpointed,slightly retrorse,becomingroundedbasally;fixedfingerwith32( L K )teeth;movablefingerwith32( L ),33( K )teeth;accessory teethabsent. Carapace(Figs.3,6):0.770.86( L ),0.74( K ) 3 longerthan broad;anteriormarginstraight;with2pairsofrounded corneateeyes,tapetumpresent;with31( L ),32( K )setae, arrangedwith4( L K )nearanteriormarginand4( L K )near posteriormargin;with1deep,broadmedianfurrow. Coxalregion:manducatoryprocessroundedwith1small sub-oralseta,and9( L ),12( K )additionalsetae;median maxillarylyrifissurelarge,roundedandsituatedsubmedially; posteriormaxillarylyrifissurerounded.CoxaeItoIV becomingprogressivelywider.ChaetotaxyofcoxaeIIV: L 6:6:6:14; K ,6:7:9:16. Legs:femoraIandIIlongerthanpatellae;junctionbetween femoraandpatellaeIIIandIVveryangulate;femoraIIIand IVmuchsmallerthanpatellaeIIIandIV;femur + patellaof legIV5.92( L ),5.27( K ) 3 longerthanbroad(Fig.10); metatarsiandtarsinotfused;tarsusIVwithouttactileseta; subterminaltarsalsetaearcuateandacuminate;clawssimple; aroliummuchlongerthanclaws,notdivided. Abdomen:tergitesIIXandsternitesIVVIIIofmaleand femalewithmedialsuturelinefullydividingeachsclerite, sterniteIXpartiallydivided.Tergalchaetotaxy: L ,4:6:10:10: 11:12:11:10:10:6(arrangedT4T):7:2; K ,6:5:7:9:10:11: 11:13:9:6(arrangedT4T):8:2;tergitesIXuniseriate. Sternalchaetotaxy: L ,19:(0)19[3 + 3](0):(0)6(0):7:9:7:8: 8:6:3:2; K ,14:(0)8(0):(0)4(0):6:7:6:8:9:6:4:2;sternites IVXuniseriate; L and K sterniteIIandIIIwithallsetae situatednearposteriormargin.Spiracleswithhelix.Anal plates(tergiteXIIandsterniteXII)situatedbetweentergiteXI andsterniteXI,andsurroundedbydesclerotizedregionof Figures15. Larcacavicola (Muchmore),malefromPARA-2004Cave:1.Body,dorsal;2.Body,ventral;3.Carapace,dorsal;4.Leftchela, lateral;5.Analregion,posterior. 208 THEJOURNALOFARACHNOLOGY

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Figures614. Larcacavicola (Muchmore),specimensfromPARA-3503Cave:6.Carapace,dorsal,male;7.Chelicera,dorsal,male;8. Rallum,lateral,male;9.Rightpedipalp,dorsal,male;10.LeftlegIV,male;11.Leftchela,lateral,male;12.Leftchela,lateral,female;13.Left chela,tritonymph;14.Leftchela,deutonymph.Scalelines 5 0.1mm(Figs.7,8),0.2mm(Figs.1114),0.5mm(Figs.6,9,10). HARVEY&WYNNETROGLOMORPHICPSEUDOSCORPIONSFROMARIZONA 209

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tergiteXIandsterniteXI;sterniteXIwithca.18( L ),24( K ) smalllyrifissures.Pleuralmembranefinelywrinkled-plicate; withoutanysetae. Genitalia:male:verysimilartothatdescribedfor L.laceyi Muchmore,1981byMuchmore(1981).Femalewith1pairof lateralcribriformplatesand2pairsofmediancribriform plates;spermathecaeabsent. Dimensions:male(PARA-3503Cave)followedbyother males(whereapplicable):Bodylength2.40(2.142.42). Pedipalps:trochanter0.371/0.186(0.3510.387/0.1920.207), femur1.021/0.172(0.9230.976/0.1870.206),patella0.859/ 0.192(0.7680.832/0.2000.229),chela(withpedicel)1.220/ 0.231(1.1731.286/0.2620.272),chela(withoutpedicel)1.160 (1.1061.216),handlength0.576(0.5690.622),movablefinger length0.582(0.5470.595).Chelicera0.200/0.115,movable fingerlength0.130.Carapace0.605/0.784(0.6210.656/0.763 0.772);anterioreyediameter0.059,posterioreyediameter 0.043.LegI:femur0.382/0.090,patella0.249/0.092,tibia0.350/ 0.067,metatarsus0.252/0.042,tarsus0.218/0.042.LegIV:femur + patella0.740/0.125,tibia0.605/0.079,metatarsus0.285/0.055, tarsus0.270/0.048. Female(PARA-3503Cave)followedbyotherfemale(where applicable):Bodylength2.85(2.72).Pedipalps:trochanter 0.422/0.219(0.408/0.215),femur1.108/0.224(0.978/0.214), patella0.992/0.252(0.822/0.223),chela(withpedicel)1.394/ 0.307(1.304/0.320),chela(withoutpedicel)1.309(1.232),hand length0.640(0.621),movablefingerlength0.643(0.616). Chelicera0.240/0.131,movablefingerlength0.150.Carapace 0.708/0.960);anterioreyediameter0.049,posterioreye diameter0.048.LegI:femur0.410/0.103,patella0.289/0.117, tibia0.382/0.075,metatarsus0.261/0.059,tarsus0.237/0.048. LegIV:femur + patella0.828/0.157,tibia0.660/0.095, metatarsus0.300/0.067,tarsus0.282/0.058. Tritonymph: Color:carapace,pedipalpsandcoxaeredbrown,abdomenpalered-brownandlegspaleyellow-brown. Chelicera:with4setaeonhandand1onmovablefinger; galealongandslenderwith3terminalrami. Pedipalp:trochanter1.97,femur5.05,patella3.90,chela (withpedicel)4.58,chela(withoutpedicel)4.32,hand(without pedicel)2.17 3 longerthanbroad,andmovablefinger1.02 3 longerthanhand(withoutpedicel).Fixedchelalfingerwith7 trichobothria,movablechelalfingerwith3trichobothria (Fig.13): eb esb ist and ib situatedbasally; est and it medially; et distally, isb absent; b subbasally, st and t submedially, sb absent.Fixedchelalfingerwith26teeth; movablefingerwith22teeth. Carapace:0.85 3 longerthanbroad;with2pairsofsmall roundedcorneateeyes;with4setaenearanteriormarginand3 nearposteriormargin;withdeepmedianfurrow. Legs:muchasinadults. Abdomen:tergalchaetotaxy:4:4:6:7:8:7:8:6:6:6 (arrangedT4T):7:2.Sternalchaetotaxy:2:(0)7(0):(0)3(0): 4:4:4:5:6:4:2:2. Dimensions(mm)(PARA-3503Cave):Bodylength1.75. Pedipalps:trochanter0.314/0.159,femur0.768/0.152,patella 0.643/0.165,chela(withpedicel)1.040/0.227,chela(without pedicel)0.981,handlength0.493,movablefingerlength0.501. Carapace0.544/0.640. Deutonymph: Color:carapace,pedipalpsandcoxaepale red-brown,abdomenandlegspaleyellow-brown. Chelicera:with4setaeonhandand1onmovablefinger; galealongandslenderwith3terminalrami. Pedipalp:trochanter2.11,femur5.16,patella3.50,chela (withpedicel)4.19,chela(withoutpedicel)3.94,hand(without pedicel)2.02 3 longerthanbroad,andmovablefinger0.97 3 longerthanhand(withoutpedicel).Fixedchelalfingerwith6 trichobothria,movablechelalfingerwith2trichobothria (Fig.14): eb ist and ib situatedbasally; est and it medially; et distally; it subdistally, esb and isb absent; b subbasally, t submedially, sb and st absent.Fixedchelalfingerwith24 teeth;movablefingerwith21teeth. Carapace:0.82 3 longerthanbroad;with2pairsofsmall roundedcorneateeyes;with4setaenearanteriormarginand4 nearposteriormargin;withdeepmedianfurrow. Legs:muchasinadults. Abdomen:tergalchaetotaxy:4:4:4:6:6:6:6:6:6:6 (arrangedT4T):4:2.Sternalchaetotaxy:0:(0)2(0):(0)2(0): 3:2:4:4:4:4:4:2. Dimensions(mm)(PARA-3503Cave):Bodylength1.49. Pedipalps:trochanter0.278/0.132,femur0.629/0.122,patella 0.514/0.147,chela(withpedicel)0.850/0.203,chela(without pedicel)0.800,handlength0.410,movablefingerlength0.397. Carapace0.490/0.600. Remarks. Larcacavicola wasdescribedfromasingle femalecollectedinCaveoftheDomes,GrandCanyon NationalPark,CoconinoCounty,Arizona(Muchmore 1981).Thenewspecimensweretakenfromtwodifferent caveswithintheParashant,PARA-3503CaveandPARA2204Cave,expandingtheknownrangeofthisspeciessome 160kmwestofthetypelocality.Specimensfrombothcave localitieshaveshorterandslightlythinnerpedipalpalsegments thanthefemaleholotype.Inaddition,thePARA-3503Cave specimenshaveslightlylongerandthinnerpedipalpsthan thosefromPARA-2204Cave.Theredonotappeartobeany othermorphologicalfeaturesthatwouldwarranttherecognitionofmorethanonespeciesamongstthesespecimens whichareallhereattributedto L.cavicola .Asnotedby Muchmore(1981),thisspeciesshowssomeobvioustroglomorphicfeaturesconsistentwithanobligatesubterranean existenceincludinglong,slenderpedipalpsandlegs,reduced posterioreyes,andfewersetaeonthecarapace.Giventhe findingsofbothMuchmore(1981)andthepresentstudy,we considerthisspeciestobetroglobitic.Ausefulmeasureof troglomorphicadaptationinlarcidpseudoscorpionswas proposedbyGardini(1983),whofoundthattheratio pedipalpalfemurlength/carapacelengthwaslowerinepigean speciesof Larca thanincavernicolousspecies.Thispattern wasalsoobservedintwonewSpanishspeciesof Larca (Zaragoza2005).Asimilarconditionisfoundinthespecies formerlydescribedin Archeolarca .Theepigean L.rotunda has alowratioof1.20(male),1.36(female)(Hoff&Clawson 1952),whereasthecavernicolousspeciesgenerallyhavehigher ratios: L.aalbui 1.57(male), L.cavicola 1.44(female), L. guadalupensis 1.34(female)and L.welbourni 1.47(female) (Muchmore1981,1984).Theratiosofthenewspecimensof L. cavicola recordedhere[1.69(male),1.56(female)]arehigher thanthefemaleholotype,butweascribethistoindividual variation. Twoofthethreepost-embryonicnymphalstages(deutonymphandtritonymph)arepresentinthesamples,andthey 210 THEJOURNALOFARACHNOLOGY

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exhibitthesametrichobothrialpatternasillustratedfor L. aalbui (underthename Archeolarcaaalbui )byHarvey(1992). FamilyChernetidaeMenge1855 SubfamilyChernetinaeMenge1855 Hesperochernes Chamberlin1924 Hesperochernes Chamberlin1924:8990. Typespecies. Hesperocherneslaurae Chamberlin1924,by originaldesignation. Remarks. Thegenus Hesperochernes currentlycomprises 19NorthAmericanspecies,rangingasfarsouthasthe DominicanRepublicandMexico(e.g.,Ellingsen1910; Chamberlin1924;Beier1933,1976)andasfarnorthas Canada(Hoff1945),andasingleJapanesespecies(Sato1983). Muchmore(1974)provideddetailsonhowtoseparate Hesperochernes fromthemorphologicallysimilargenera Chernes Menge1855and Dinocheirus Chamberlin1929,but admittedthatthecompositionofthegenuswasnotfully resolvedduetouncertaintiesinthemorphologyofseveral species. Hesperochernes iscurrentlydiagnosedbythefollowingcombinationofcharacters:rallumcomposedof4blades; tarsusIIIandIVwithoutconspicuoustactileseta;setaeof pedipalpsandtergitesnotlargeandleaf-like;female spermathecaewithlongpairedductsterminatinginrounded sacs;andcheliceralsetae bs and sbs usuallydentateor denticulate.Ofthesecharacters,Muchmore(1974)wasonly abletonominatethespermathecalmorphologyandthe denticulate bs and sbs asfeaturesthatdistinguishitfrom Chernes .Itappears,however,thatsomespeciescurrently assignedto Hesperochernes haveanacuminate bs ,including H.canadensis H.holsingeri H.molestus H.montanus H. occidentalis and H.riograndensis (Chamberlin1935;Hoff 1945;Hoff&Clawson1952;Hoff1956b;Hoff&Bolsterli 1956;Muchmore1994).Moreover,thenewspeciesdescribed belowclearlydemonstratesthelabilenatureofthisfeature, withthemalehavingastronglydenticulate bs onboth chelicerae,butthetwofemaleshavinganacuminate bs .It wouldseemthatthisfeatureshouldbeusedwithconsiderable caution,andthatthenatureofthespermathecaeistheonly featurethatcanbereliablyusedtoseparate Hesperochernes from Chernes AlthoughMuchmore(1974)wasabletoconfirmthegeneric placementofseveralspeciesfromtheU.S.A.andCanada [ H.laurae,H.mimulus Chamberlin1952, H.mirabilis (Banks 1895), H.molestus Hoff1956, H.occidentalis (Hoffand Bolsterli1956). H.riograndensis HoffandClawson1952, H. tamiae Beier1930,and H.utahensis HoffandClawson1952], hewasnotabletoascertainwhetherotherswerecorrectly placed[ H.canadensis Hoff1945, H.montanus Chamberlin 1935, H.pallipes (Banks1893), H.paludis (Moles1914), H. thomomysi Hoff1948,and H.unicolor (Banks1908)].The samecanbesaidoftheCentralAmericanandAsianspecies currentlyincludedin Hesperochernes H.globosus (Ellingsen 1910), H.tumidus Beier1933and H.inusitatus Hoff1946from Mexico, H.vespertilionis Beier1976fromDominicanRepublic,and H.shinjoensis Sato1983fromJapan,asthemorphology ofthespermathecaehasnotyetbeenascertained(Ellingsen 1910;Beier1933;Hoff1946a;Beier1976;Sato1983). Speciesof Hesperochernes arefrequentlycollectedincaves orareassociatedwithotheranimals.Thecave-dwellingspecies includethreeeyelessspeciesthathavelongslenderpedipalps consistentwithstrongtroglomorphisms, H.holsingeri H. mirabilis and H.occidentalis ,aswellastheneweyelessspecies describedbelowthathaslonglegsbuthasrobustpedipalps. Thespeciesassociatedwithrodentsinclude H.mimulus H. molestus H.riograndensis H.tamiae H.thomomysi and H. utahensis (Beier1930;Hoff1945,1946b;Chamberlin1952; Hoff&Clawson1952;Hoff1956b),while H.vespertilionis was collectedwithinabatroost(Beier1976). Hesperochernes laurae and H.unicolor werefoundwithinbothwasp'sand ant'snests(Banks1908;Chamberlin1924;Hoff1947), respectively, H.montanus wasfoundinabird'snest (Chamberlin1935),and H.tumidus wascollectedlyingon thegroundinpodsof Inga sp.''(translatedfromtheoriginal German)(Beier1933).Thepoorlydescribedandmostlikely misplaced H.paludis wastakenfrombothrottenpoplartree logsonthegroundandlivestandingpoplartrees(Moles 1914),andtheonlyspeciesrecordedfromoutsideofNorth America, H.shinjoensis fromnorthernJapan,wascollected fromundertreebark(Sato1983).Theotherspecieslackany habitatdata. Hesperochernesbradybaughi sp.nov. urn:lsid:zoobank.org:act:5419D319-EF22-4722-926F-1F8EC 080400B Figs.1526 Materialexamined. Types. U.S.A.: Arizona :Mohave County:holotypemale,PARA-1001Cave,GrandCanyonParashantNationalMonument,ca.UTM0264500N,4060700 E,Zone12S,baitedpitfalltrap3B,20August2007,J.J.Wynne (MNA);1female,samedataasholotypeexceptbaitedpitfall trap5A(MNA);1female,samedataasholotypeexcept opportunistic,midcave,13August2005(NMA). Etymology. ThisspeciesisnamedforJeffBradybaugh, formersuperintendentofGrandCanyon-ParashantNational Monumentandanadvocateforcaveresearch,conservation andmanagementbothonParashantandwithintheNational ParkService. Diagnosis. Hesperochernesbradybaughi mostcloselyresemblesthreeotherspeciesofthegenusthatarealso completelyeyelessandhavelongslenderlegs[e.g.femur + patellaIV5.19(male),5.375.56(female) 3 longerthan broad], H.mirabilis H.holsingeri and H.riograndensis Hesperochernesbradybaughi lackstheslenderpedipalps characteristicof H.mirabilis and H.holsingeri ,andthemale chelaof H.bradybaughi ismarkedlyswollen,especiallyonthe dorsalface(Fig.21),unlikethemaleof H.riograndensis which isnotswollen.Itisalsosubstantiallylargerthan H. riograndensis ,e.g.,chela(withoutpedicel)of H.riograndensis is0.956(male),0.970(female)mm,whereas H.bradybaughi is 1.434(male),1.5021.510(female)mm. Description Adults: Color:pedipalpsandcarapacedark red-brown,legslightred-brown,tergitesyellow-brown, sternitespaleyellow-brown. Chelicera:with5setaeonhandand1subdistalsetaon movablefinger(Fig.23);setae ls and is acuminate, es and bs dentate, sbs denticulateinfemale,acuminateinmale;with2 dorsallyrifissuresand1ventrallyrifissure;galeaof L and K with6rami;rallumof4blades,the2distalbladeswithseveral HARVEY&WYNNETROGLOMORPHICPSEUDOSCORPIONSFROMARIZONA 211

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serrationsonleadingedge,otherbladessmooth;serrula exteriorwith18( L ),17( K )blades;laminaexteriorpresent. Pedipalp(Fig.24):surfacesoftrochanter,femur,patella andchelalhandcoarselygranulate,chelafingersmostly smooth;patellawith5smallsub-basallyrifissures;trochanter 1.84( L ),1.861.88( K ),femur3.17( L ),2.953.09( K ),patella 2.62( L ),2.542.66( K ),chela(withpedicel)3.07( L ),3.233.34 ( K ),chela(withoutpedicel)2.83( L ),2.983.09( K ),hand1.49 ( L ),1.341.64( K ) 3 longerthanbroad,movablefinger0.93 ( L ),0.860.96( K ) 3 longerthanhand.Fixedchelalfingerwith 8trichobothria,movablechelalfingerwith4trichobothria (Figs.21,22): eb and esb situatedbasally, ib and ist subbasally, est and isb submedially, et and it subdistally, isb situatedmidwaybetween ist and it ,and et slightlydistalto it ; t situatedsubdistally, st situatedcloserto t thanto sb .Venom apparatusonlypresentinmovablechelalfinger,venomducts long,terminatinginnodusramosusdistalto st (Figs.21,22). Fixedfingerwith2largesensillaeonretrolateralface,and2on prolateralface;movablechelalfingerwithsensillaslightly proximalto sb inmaleandslightlydistalto sb infemale,with 2receptors.Chelaofmalewithoutmound.Chelalteeth pointedandslightlyretrorse,basalteethmorerounded;fixed fingerwith44( L ),48( K )teeth,plus11( L ),9( K )retrolateral and10( L ),7( K )prolateralaccessoryteeth;movablefinger with46( L ),50( K )teeth,plus9( L K )retrolateraland6( L ),4( K ) prolateralaccessoryteeth. Carapace(Fig.17):coarselygranulate,1.15( L ),0.981.10 ( K ) 3 longerthanbroad;withouteyesoreyespots;with100 ( L ),83( K )setae,arrangedwith61( L ),42( K )(including6near anteriormargin)inanteriorzone,25( L ),34( K )inmedian zone,and14( L ),17( K )inposteriorzone;with2deepfurrows, posteriorfurrowsituatedslightlyclosertoposteriorcarapace marginthantoanteriorfurrow. Coxalregion:maxillaegranulate;manducatoryprocess somewhatacute,with2apicalacuminatesetae,1smallsuboralsetaand37( L ),32( K )additionalsetae;medianmaxillary lyrifissureroundedandsituatedsubmedially;posteriormaxillarylyrifissurerounded.Legcoxaesmooth;chaetotaxyof coxaeIIV: L ,18:19:23:ca.60; K ,18:21:25:ca.65. Legs:veryslender;junctionbetweenfemoraandpatellaeI andIIstronglyobliquetolongaxis;junctionbetweenfemora andpatellaeIIIandIVveryangulate;femoraIIIandIVmuch smallerthanpatellaeIIIandIV;femur + patellaoflegIV5.19 ( L ),5.375.56( K ) 3 longerthanbroad;alltarsiwithslit sensillumonraisedmound;malelegInotmodified;tarsiIII andIVwithouttactileseta,butwithpairedsubdistalsetae; subterminaltarsalsetaearcuateandacute;clawssimple; aroliumaboutsamelengthasclaws,notdivided. Abdomen:tergitesIXandsternitesIVXofmaleand femalewithmediansuturelinefullydividingeachsegment. Tergalchaetotaxy: L ,11:12:11:18:19:18:20:18:17:18:13:2; K ,12:13:13:17:17:18:19:19:21:16:14:2;uniseriate,except Figures1520. Hesperochernesbradybaughi ,sp.nov.:15.Body,dorsal,maleholotype;16.Body,ventral,maleholotype;17.Carapace, dorsal,maleholotype;18.Body,dorsal,femaleparatype;19.Body,ventral,femaleparatype;20.Leftchela,lateral,maleholotype. 212 THEJOURNALOFARACHNOLOGY

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formedialandlateraldiscalsetaontergitesIVIX;setae thickenedandstronglydentate.Sternalchaetotaxy: L ,30:(3)22 [2 + 2](3):(1)8(1):19:21:21:20:20:16:8(arrangedT6T):2; K ca.40:(3)10(3):(1)5(1):14:21:20:20:18:16:11(arranged T9T):2;uniseriate,exceptforlateraldiscalsetaonsternites VIIX;setaeofanteriorsternitesacicular,becomingprogressivelymoredenticulateonposteriorsternites.Spiracleswith helix.Analplates(tergiteXIIandsterniteXII)situatedbetween tergiteXIandsterniteXI,analsetaenotdenticulate.Pleural membranewrinkledandsomewhatstellate;withoutanysetae. Genitalia:maleofthechernetidtype.Female(Fig.26):with apairoflongthin-walledspermathecaeterminatingin roundedsacs. Dimensions:Maleholotype:Bodylength3.11.Pedipalps: trochanter0.518/0.282,femur974/0.307,patella0.824/0.314, chela(withpedicel)1.552/0.506,chela(withoutpedicel)1.434, handlength0.756,movablefingerlength0.704.Chelicera 0.322/0.165,movablefingerlength0.252.Carapace0.956/ 0.830.LegI:femur0.268/0.161,patella0.495/0.136,tibia 0.503/0.102,tarsus0.495/0.079.LegIV:femur + patella0.883/ 0.170,tibia0.778/0.107,tarsus0.557/0.085. Female(paratypelodgedinMNA)followedbyother female(whereapplicable):Bodylength2.82(4.21).Pedipalps: trochanter0.552/0.294(0.566/0.304),femur1.034/0.335(1.042/ 0.353),patella0.904/0.340(0.942/0.371),chela(withpedicel) 1.624/0.486(1.635/0.506),chela(withoutpedicel)1.502 Figures2126. Hesperochernesbradybaughi ,sp.nov.:21.Leftchela,lateral,maleholotype;22.Leftchela,lateral,femaleparatype;23. Chelicera,dorsal,femaleparatype;24.Rightpedipalp,dorsal,maleholotype;25.LeftlegIV,maleholotype;26.Spermathecae,femaleparatype. Scalelines 5 0.1mm(Fig.23),0.2mm(Fig.26),0.5mm(Figs.21,22,24,25). HARVEY&WYNNETROGLOMORPHICPSEUDOSCORPIONSFROMARIZONA 213

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(1.510),handlength0.797(0.698),movablefingerlength0.768 (0.816).Chelicera0.327/0.152,movablefingerlength0.244. Carapace1.040/0.944(1.000/1.021).LegI:femur0.300/0.182, patella0.540/0.146,tibia0.56/0.108,tarsus0.520/0.079.Leg IV:femur + patella1.010/0.188(1.000/0.180),tibia0.830/ 0.121,tarsus0.580/0.084. Remarks. Asstatedinthediagnosis, H.bradybaughi appearstobemostsimilarto H.riograndensis butdiffersin beingsubstantiallylargerandwithamarkedlyswollenmale chela,especiallyonthedorsalface.Theonlyknownlocation of H.riograndensis islocated670kmESEofParashant,and themicrohabitatofbothspeciesdifferswith H.bradybaughi beingfoundinacaveand H.riograndensis collectedfromthe nestofakangaroorat(Heteromyidae: Dipodomys )(Hoff& Clawson1952).Giventhelackofeyesandeyespots,we consider H.bradybaughi tobeatroglobite. Tuberochernes Muchmore Tuberochernes Muchmore1997:206207. Typespecies. Tuberochernesaalbui Muchmore1997,by originaldesignation. Diagnosis. Tuberochernes differsfromallotherchernetid generabythecombinedpresenceofadistinctmedium-sized moundontheprolateralfaceofthepedipalpalchelaofmales, andfourbladesinthecheliceralrallum. Remarks. Thegenus Tuberochernes wasdescribedby Muchmore(1997)fortwospeciesofcave-dwellingpseudoscorpionsfromsouthwesternU.S.A., T.aalbui and T.ubicki butthediscoveryofathirdspecies,alsofromacavein southwesternU.S.A.,doesnotnecessitateanalterationofthe originaldescriptionapartfromthenatureofthetactilesetaof legIV.Muchmore(1997)observedthatthetactilesetaofleg IVwasshort,distallylocated''andvariablyacuminateor finelydenticulate''.Closeexaminationoftheposteriortarsiof thenewspeciesdescribedbelowdoesnotrevealatactileseta ofthisnature,andwesuggestthisfeatureappearstobe variablewithinthegenus. Themostobviousfeaturethatdistinguishes Tuberochernes isthepresenceofamedium-sizedmoundontheprolateral marginofthechelalhandinmales(Muchmore1997).Inthis respect,itresemblesseveralotherchernetidgenera,including malesof Mirochernes Beier1930and Bituberochernes Muchmore1974,andbothmalesandfemalesof Interchernes Muchmore1980and Petterchernes Heurtault1986,which weredistinguishedfrom Tuberochernes byMuchmore(1997). Bituberochernes furtherdiffersfrom Tuberochernes bya moundbeingalsopresentonthepedipalpalpatella.The functionofthemoundhasnotbeenascertained,butthe moundof T.cohni has5smallpores,whichmayberesponsible fordischargingfluids,possiblyduringsexualinteractionswith females. Tuberochernescohni sp.nov. urn:lsid:zoobank.org:act:12896B35-DD1C-4E0B-B66F-F9B 30170D476 Figs.2737 Materialexamined Type: U.S.A.: Arizona :Mohave County:holotypemale,PARA-1001Cave,GrandCanyonParashantNationalMonument,ca.UTM0264500N, 4060700E,Zone12S,thedeeperextentofthetwilightzone (nearthedarkzone),opportunisticcollecting,13August2005, J.J.Wynne(MNA). Etymology. ThisspeciesisnamedforthelateDr. TheodoreTed''Cohn.CohnwasanOrthopteristandthe leadingauthoritywhoidentifiedthenewgenusofrhaphidophoridcricketknownfromPARA-1001Cave.Dr.Cohn passedawayinNovember2013atage82.Hewasapassionate educatorandentomologist. Diagnosis. Tuberochernescohni differsfromtheothertwo speciesofthegenus, T.aalbui and T.ubicki ,bythemore anteriorlypositionedmoundonthepedipalpalchela. Description Adultmale: Color:pedipalpsandcarapace darkred-brown,legslightred-brown,tergitesyellow-brown, sternitespaleyellow-brown. Chelicera:with6setaeonhandand1subdistalsetaon movablefinger(Fig.32);setae es sbs and bs dentate, ls and is acuminate;with2dorsallyrifissuresand1ventrallyrifissure; galeabroken;rallumof4blades,themostdistalbladewith severalserrationsonleadingedge,otherbladessmooth; serrulaexteriorwith17blades;laminaexteriorpresent. Pedipalp(Fig.33):surfacesoftrochanter,femur,patella andchelalhandcoarselygranulate,chelafingersmostly smooth;patellawith5smallsub-basallyrifissures;trochanter 1.73,femur2.83,patella2.88,chela(withpedicel)3.39,chela (withoutpedicel)3.11,hand1.40 3 longerthanbroad, movablefinger1.23 3 longerthanhand.Fixedchelalfinger with8trichobothria,movablechelalfingerwith4trichobothria(Fig.31): eb and esb situatedbasally, ib and ist subbasally, est and isb submedially, et and it subdistally, isb situatedmidwaybetween ist and it ,and et slightlydistalto it ; t situatedsubdistally, st situatedmuchcloserto t thanto sb Venomapparatusonlypresentinmovablechelalfinger, venomductslong,terminatinginnodusramosusmidwayat levelof st (Fig.31).Fixedfingerwith3sensillaeonretrolateral face,and1onprolateralface;movablechelalfingerwith sensillaslightlydistaltosb,with2receptors.Chelawith prominent,medium-sizedmoundonprolateralface(Figs.30, 34),with5smallpores.Chelalteethpointedandslightly retrorse,basalteethmorerounded;fixedfingerwith37teeth, plus7retrolateraland3prolateralaccessoryteeth;movable fingerwith42teeth,plus4retrolateraland0prolateral accessoryteeth. Carapace(Fig.29):coarselygranulate,1.19 3 longerthan broad;withouteyesoreyespots;with96setae,arrangedwith 54(including6nearanteriormargin)inanteriorzone,28in medianzone,and14inposteriorzone;with2deepfurrows, posteriorfurrowsituatedclosertoposteriorcarapacemargin thantoanteriorfurrow. Coxalregion:maxillaegranulate;manducatoryprocess somewhatacute,with2apicalacuminatesetae,1smallsuboralsetaand25additionalsetae;medianmaxillarylyrifissure roundedandsituatedsubmedially;posteriormaxillarylyrifissurerounded.Legcoxaesmooth;chaetotaxyofcoxaeIIV: 13:12:14:34. Legs(Figs.3537):junctionbetweenfemoraandpatellaeI andIIstronglyobliquetolongaxis;junctionbetweenfemora andpatellaeIIIandIVveryangulate;femoraIIIandIVmuch smallerthanpatellaeIIIandIV;femur + patellaoflegIV4.03 3 longerthanbroad;alltarsiwithslitsensillumonraised 214 THEJOURNALOFARACHNOLOGY

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mound;legImodifiedwithtibiathickened,tarsusslightly curvedandventralmarginsofpatellaandtibiawithcoarse granulation;tarsiIIIandIVwithouttactileseta,butwith pairedsubdistalsetae;subterminaltarsalsetaearcuateand acute;clawssimple;aroliumaboutsamelengthasclaws,not divided. Abdomen:tergitesIIXandsternitesVXofwithmedian suturelinefullydividingeachsegment.Tergalchaetotaxy:15: 20:20:20:22:22:21:21:22:17:10:2;uniseriate,exceptfor medialandlateraldiscalsetaontergitesIVIX;setae thickenedandstronglydentate.Sternalchaetotaxy:51:(0)8 [2 + 2](0):(1)8(1):12:16:17:18:17:14:8(arrangedT6T):2; uniseriate,exceptforlateraldiscalsetaonsternitesIVXI; setaeofanteriorsternitesacicular,becomingprogressively moredenticulateonposteriorsternites.Spiracleswithhelix. Analplates(tergiteXIIandsterniteXII)situatedbetween tergiteXIandsterniteXI,analsetaedenticulate.Pleural membranelongitudinallystriate;withoutanysetae. Genitalia:ofthechernetidtype. Dimensions:maleholotype:Bodylength3.38.Pedipalps: trochanter0.576/0.332,femur0.944/0.334,patella0.910/0.316, chela(withpedicel)1.390/0.410,chela(withoutpedicel)1.276, handlength0.573,movablefingerlength0.704.Chelicera 0.333/0.134,movablefingerlength0.240.Carapace1.009/ 0.848.LegI:femur0.305/0.249,patella0.560/0.253,tibia 0.621/0.174,tarsus0.442/0.089.LegIV:femur + patella0.859/ 0.213,tibia0.692/0.134,tarsus0.533/0.954. Remarks. Tuberochernescohni possessessomeveryslight modificationsconsistentwithtroglomorphicadaptationsof whichthemostprominentisthecompletelackofeyes (Fig.29)andtheslightlyelongatedlegsegments.Thus,this animalisconsideredatroglobite.Itappearstobearacloser resemblanceto T.ubicki fromacaveintheSantaRita Mountains,Arizona(610km),thanto T.aalbui fromacavein theInyoNationalForest,California(415km),duetothe similarlyexpandedtibiaIinmalesofthetwoArizonaspecies. DISCUSSION Ourreviewofthepseudoscorpionsdetectedwithinthecaves ofGrandCanyon-ParashantNationalMonumenthasrevealedamodestfaunaofthreespecies: Larcacavicola (family Larcidae), Hesperochernesbradybaughi and Tuberochernes cohni (bothinthefamilyChernetidae).Allshowmodifications consistentwithobligateexistenceincaveenvironments,but noneshowtheclassicsignsofextremetroglomorphismfound inmanycave-adaptedpseudoscorpions(e.g.Heurtault1994; Harveyetal.2000).BothspeciesofChernetidaelackeyesand havelongslenderlegs,whichappeartobetroglomorphic modificationsduetotheirsubterraneanexistence,although theirpedipalpsdonotappeartobemodifiedcomparedto epigeanspeciesofthegenus.Othersubterraneanspeciesof Hesperochernes withthinlegsandnoeyespots H.holsingeri fromIndiana, H.mirabilis fromAlabama,Georgia,Indiana, Kentucky,Ohio,TennesseeandVirginia,and H.occidentalis fromArkansas,Missouri,OklahomaandTexasappeartobe morehighlymodifiedastheyhaveelongatepedipalps.Both newspeciesdescribedfromtheParashantmayrepresentshortrangeendemicspeciesasdefinedbyHarvey(2002)andHarvey Figures2730. Tuberochernescohni ,sp.nov.,maleholotype:27.Body,dorsal;28.Body,ventral;29.Carapace,dorsal;30.Right chela,dorsal. HARVEY&WYNNETROGLOMORPHICPSEUDOSCORPIONSFROMARIZONA 215

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Figures3137. Tuberochernescohni ,sp.nov.,maleholotype:31.Leftchela,lateral;32.Chelicera,dorsal;33.Rightpedipalp,dorsal;34.Left chela,detailofmound,ventral;35.LefttarsusIV;36.LeftI;37.LeftlegIV.Scalelines 5 0.2mm(Figs.32,34,35),0.5mm(Figs.31,33,36,37). 216 THEJOURNALOFARACHNOLOGY

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etal.(2011)duetotheirhighlyrestricteddistributions. Althoughthejuniorauthorandcolleaguessampledallknown cavesonParashant,theydetectedthesenewspeciesinonly onecave(PARA-1001Cave). Larcacavicola appearstobelesscave-adaptedthanthe others,asitretainseyes.However,thepedipalpsare noticeablythinnerthanepigeanspeciesofthegenus, suggestingmoderatemorphologicalmodificationstothecave environment. Larcacavicola wasfoundinPARA-3503and PARA-2204CavesandhasbeenfoundinCaveoftheDomes, asmallcavesituatedwithinGrandCanyonNationalPark, CoconinoCounty(Muchmore1981).Althoughthiscaveis alsolocatedintheGrandCanyonregion,itliesonthesouth sideoftheColoradoRiversome160kmfromtheParashant caves,andwesuggestthesepopulationsaregenetically isolatedfromeachother. Theonlyknownlocalityof Hesperochernesbradybaughi and Tuberochernescohni isPARA-1001.Thisisthesecondmost biologicallydiversecave,andthemostbiologicallysignificant caveonthemonument.Itsupportsthelargestknowncricket roostinArizona,whichrepresentsanundescribedgenusof rhaphidophoridcavecricket,cf Ceuthophilus n.gen.n.sp., Cohn&Swanson,unpublisheddata;(Wynne&Voyles2014). Itspopulationcontributessignificantlytothenutrientloading viacricketguano,cricketeggsandnymphs,aswellasdeceased individualsatvariouslifestages.Inotherregions,the ecologicalimportanceofcricketsoncaveecosystemsiswell documented(e.g.,Barr1967;Howarth1983;Taylor2003; Culver2005;Poulson2005).Giventhesizeoftheroost,we suggestthatcf Ceuthophilus n.gen.n.sp.representsakeystone specieswiththepresenceofthisanimalsupportingatleast fourcave-adaptedspeciesincludingashort-rangeendemicand troglomorphicleiodidbeetle, Ptomaphagusparashant (Peck& Wynne2013),anundescribedspeciesoftroglomorphic centipede(familyAnopsobiidae;Wynne,unpublisheddata), andthetwopseudoscorpionspeciesdescribedhere.Todate, P.parashant ,theanopsobiidcentipede,andthetwonew pseudoscorpionspecieshavebeendetectedonlyinPARA1001Cave.Twoothercavesonthemonument,withsimilar deepzonelikeconditions,weresampledusingthesame systematicsamplingdesignarewithina9.7kmradiusof PARA-1001;neitherofthesenewpseudoscorpionsspecies weredetectedatthesecaves. ManagementImplications. WerecommendthesamemanagementstrategiesproposedbyPeck&Wynne(2013)be maintainedforPARA-1001Cave.Thiscaveshouldnotbe gatedgivenitssouth-facingentranceandentrancestructure, anditshouldremainclosedtorecreationaluse.PARA-1001is consideredthesecondmostbiologicallydiversecaveonthe monumentandsupportsthegreatestdiversityoftroglomorphicarthropodspecies.Presently,alloftheseanimals (includingthetwonewpseudoscorpionspeciesdescribedhere) areknowntooccuronlywithinPARA-1001Cave.MaintainingthemanagementstrategiessuggestedbyPeck&Wynne (2013)shouldaidinthelong-termpersistenceofthese presumedshort-rangeendemicarthropods. ACKNOWLEDGMENTS SpecialthankstoJenniferFox,EathanMcIntyre,Ray KleinandRosiePepitoofGrandCanyon-ParashantNational Monument,DanielleNelsonandMattJohnsonwiththe ColoradoPlateauResearchStation,andNeilCobbofthe ColoradoPlateauMuseumofArthropodBiodiversityfor administrativeandlogisticalsupport.TamaandJohnCassidy, MichaelGowan,JohnKalman,TySpattaandKyleVoyles assistedwithfieldwork.TheSanBernardinoCaveSearchand RescueTeam,JonJasperandKyleVoyles,remainedon emergencystand-byduringfieldoperations.DaveDeckerand KyleVoylesprovideddescriptionsregardingthegeological andstructuralcharacteristicsofthestudycaves.DalePateand twoanonymousreviewersprovidedsuggestionsleadingtothe improvementofthismanuscript.TheExplorersClubrecognizedtwooftheseresearchtripsasflagexpeditions.Fieldwork wasfundedthroughaColoradoPlateauCESUcooperative agreementbetweentheNationalParkServiceandNorthern ArizonaUniversity. 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Chamberlin,J.C.1930.Asynopticclassificationofthefalsescorpions orchela-spinners,withareportonacosmopolitancollectionofthe same.PartII.TheDiplosphyronida(Arachnida-Chelonethida). AnnalsandMagazineofNaturalHistory(10)5:148,585620. Chamberlin,J.C.1931.ThearachnidorderChelonethida.Stanford UniversityPublications,BiologicalSciences7(1):1284. HARVEY&WYNNETROGLOMORPHICPSEUDOSCORPIONSFROMARIZONA 217

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Muchmore,W.B.&R.B.Pape.1999.Descriptionofaneyeless, cavernicolous Albiorix (Pseudoscorpionida:Ideoroncidae)inArizona,withobservationsonitsbiologyandecology.Southwestern Naturalist44:138147. Peck,S.B.&J.J.Wynne.2013. Ptomaphagusparashant Peckand Wynne,newspecies(Coleoptera:Leiodidae:Cholevinae:Ptomaphagini):themosttroglomorphiccholevinebeetleknownfrom westernNorthAmerica.TheColeopteristsBulletin67:309317. Poulson,T.L.2005.Foodsources.Pp.255264. In Encyclopediaof Caves.(D.C.Culver&W.B.White,eds.).Elsevier,Burlington,MA. Sato,H.1983. Hesperochernesshinjoensis ,anewpseudoscorpion (Chernetidae)fromJapan.BulletinoftheBiogeographicalSociety ofJapan38:3134. Shear,W.A.,S.J.Taylor,J.J.Wynne&J.K.Krejca.2009.Cave millipedsoftheUnitedStates.VIII.Newgeneraandspeciesof polydesmidanmillipedsfromcavesinthesouthwesternUnited States(Diplopoda,Polydesmida,PolydesmidaeandMacrosternodesmidae).Zootaxa2151:4765. Taylor,S.J.2003.America,North:Biospeleology.Pp.4549. In EncyclopediaofCavesandKarstScience.(J.Gunn,ed.).Fitzroy Dearborn,NewYork,NY. Wynne,J.J.&K.D.Voyles.2014.Cave-dwellingarthropodsand vertebratesofNorthRimGrandCanyon,withnotesonecology andmanagement.WesternNorthAmericanNaturalist74:117. Zaragoza,J.A.2005.Twonewcave-dwelling Larca speciesfromthe south-eastofSpain(Arachnida,Pseudoscorpiones,Larcidae). RevueSuissedeZoologie112:195213. Manuscriptreceived25May2014,revised10July2014. HARVEY&WYNNETROGLOMORPHICPSEUDOSCORPIONSFROMARIZONA 219