Troglomorphic pseudoscorpions (Arachnida: Pseudoscorpiones) of northern Arizona, with the description of two new short-range endemic species

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Troglomorphic pseudoscorpions (Arachnida: Pseudoscorpiones) of northern Arizona, with the description of two new short-range endemic species

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Title:
Troglomorphic pseudoscorpions (Arachnida: Pseudoscorpiones) of northern Arizona, with the description of two new short-range endemic species
Creator:
J. Judson Wynne, Ph.D. ( suggested by )
Language:
English

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Mark S. Harvey 1,2,3,4,5 and J. Judson Wynne 6 : 1 Department of Terrestrial Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Western Australia 6986, Australia; 2 Research Associate, Division of Invertebrate Zoology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024-5192, U.S.A; 3 Research Associate, California Academy of Sciences, 55 Music Concourse Drive, San Francisco, California 94118, U.S.A; 4 Adjunct, School of Animal Biology, University of Western Australia, Crawley, Western Australia 6009, Australia; 5 Adjunct, School of Natural Sciences, Edith Cowan University, Joondalup, Western Australia 6027, Australia; 6 Department of Biological Sciences, Colorado Plateau Biodiversity Center, Landscape Conservation Initiative, Northern Arizona University, Box 5640, Flagstaff, Arizona 86011, U.S.A. E-mail: mark.harvey@museum.wa.gov.au Abstract. This study reports on the pseudoscorpion fauna of the subterranean ecosystems of northern Arizona, U.S.A. Our work resulted in the descriptions of two new species, Hesperochernes bradybaughi sp. nov. and Tuberochernes cohni sp. nov. (Chernetidae) and the range expansion of one species, Larca cavicola (Muchmore 1981) (Larcidae). All of these species were cave-adapted and found within caves on Grand Canyon-Parashant National Monument in northwestern Arizona. Based upon this work, the genus Archeolarca Hoff and Clawson is newly synonymized with Larca Chamberlin, and the following species are transferred from Archeolarca to Larca , forming the new combinations L. aalbui (Muchmore 1984), L. cavicola (Muchmore 1981), L. guadalupensis (Muchmore 1981) and L. welbourni (Muchmore 1981). Despite intensive sampling on the monument, the two new species were detected in only one cave. This cave supports the greatest diversity of troglomorphic arthropod species on the monument-all of which are short-range endemics occurring in only one cave. Maintaining the management recommendations provided by Peck and Wynne (2013) for this cave should aid in the long- term persistence of these new pseudoscorpion species, as well as the other troglomorphic arthropods. Keywords: Nearctic, troglomorphy, troglobite, new synonymy, cave urn:lsid:zoobank.org:pub:A15CB9DB-5B36-4A7C-8052-08E2EC1F4D34
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Description
Mark S. Harvey
1,2,3,4,5
and
J. Judson Wynne
6
:
1
Department of Terrestrial Zoology, Western Australian
Museum, Locked Bag 49, Welshpool DC, Western
Australia 6986, Australia;
2
Research Associate, Division of Invertebrate Zoology,
American Museum of Natural History, Central Park West
at 79th Street, New York, New York 10024-5192,
U.S.A;
3
Research Associate, California Academy of Sciences,
55 Music Concourse Drive, San Francisco, California
94118, U.S.A;
4
Adjunct, School of Animal Biology, University of
Western Australia, Crawley, Western Australia 6009,
Australia;
5
Adjunct, School of Natural Sciences, Edith Cowan
University, Joondalup, Western Australia 6027,
Australia;
6
Department of Biological Sciences, Colorado Plateau
Biodiversity Center, Landscape Conservation
Initiative, Northern Arizona University, Box 5640,
Flagstaff, Arizona 86011, U.S.A. E-mail:
mark.harvey@museum.wa.gov.au
Abstract.
This study reports on the pseudoscorpion fauna of the
subterranean ecosystems of northern Arizona, U.S.A. Our
work resulted in the descriptions of two new
species,
Hesperochernes bradybaughi
sp. nov. and
Tuberochernes cohni
sp. nov. (Chernetidae) and the range expansion of one
species,
Larca cavicola
(Muchmore 1981) (Larcidae). All of these species were
cave-adapted and found within caves on Grand
Canyon-Parashant National Monument in northwestern
Arizona. Based upon this work, the genus
Archeolarca
Hoff and Clawson is newly synonymized with
Larca
Chamberlin, and the following species are transferred
from
Archeolarca
to
Larca
, forming the new combinations
L. aalbui
(Muchmore 1984),
L. cavicola
(Muchmore 1981),
L. guadalupensis
(Muchmore 1981) and
L. welbourni
(Muchmore 1981). Despite intensive sampling on the
monument, the two new species were detected in only one
cave. This cave supports the greatest diversity of
troglomorphic arthropod species on the monument-all of
which are short-range endemics occurring in only one
cave. Maintaining the management recommendations
provided by Peck and Wynne (2013) for this cave should
aid in the long- term persistence of these new
pseudoscorpion species, as well as the other
troglomorphic arthropods.
Keywords:
Nearctic, troglomorphy, troglobite, new synonymy,
cave
urn:lsid:zoobank.org:pub:A15CB9DB-5B36-4A7C-8052-08E2EC1F4D34



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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. LITERATURECITED Aalbu,R.L.,A.D.Smith&C.A.Triplehorn.2012.Arevisionofthe Eleodes (subgenus Caverneleodes )withnewspeciesandnoteson cavebreeding Eleodes (Tenebrionidae:Amphidorini).Annales Zoologica62:199216. Ashmole,N.P.,P.Orom ,M.J.Ashmole&J.L.Mart n.1992. Primaryfaunalsuccessioninvolcanicterrain:lavaandcavestudies ontheCanaryIslands.BiologicalJournaloftheLinneanSociety 46:207234. Banks,N.1908.ThepseudoscorpionsofTexas.Bulletinofthe WisconsinNaturalHistorySociety6:3942. Barber,H.S.1931.Trapsforcaveinhabitinginsects.Journalofthe MitchellSociety46:259266. Barr,T.C.Jr.1967.Observationsontheecologyofcaves.American Naturalist101:475491. Beier,M.1930.DiePseudoskorpionedesWienerNaturhistorischen Museums.III.AnnalendesNaturhistorischenMuseumsinWien 44:199222. Beier,M.1933.PseudoskorpioneausMexiko.ZoologischerAnzeiger 104:91101. Beier,M.1939a.DiePseudoscorpioniden-Faunaderiberischen Halbinsel.ZoologischeJahrbu cher,Abteilungfu rSystematik, O kologieundGeographiederTiere72:157202. Beier,M.1939b.ThePseudoscorpionideacollectedbythePercy SladenTrustExpeditiontoLakeTiticaca.AnnalsandMagazineof NaturalHistory(11)3:288290. Beier,M.1947.ZurKenntnisderPseudoscorpionidenfaunades su dlichenAfrika,insbesonderedersu dwest-undsu dafrikanischen Trockengebiete.Eos,Madrid23:285339. Beier,M.1962.PseudoscorpionidenausderNamib-Wu ste.Annalsof theTransvaalMuseum24:223230. Beier,M.1973.WeitereszurKenntnisderPseudoscorpioniden Su dwestafrikas.Cimbebasia,A2:97101. Beier,M.1976.PseudoscorpionevonderDominicanischenRepublik (InselHaiti).RevueSuissedeZoologie83:4558. Benedict,E.M.&D.R.Malcolm.1978.Somegarypoidfalse scorpionsfromwesternNorthAmerica(Pseudoscorpionida: GarypidaeandOlpiidae).JournalofArachnology5:113132. Chamberlin,J.C.1924. Hesperocherneslaurae ,anewspeciesoffalse scorpionfromCaliforniainhabitingthenestof Vespa .Pan-Pacific Entomologist1:8992. 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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Chamberlin,J.C.1935.Anewspeciesoffalsescorpion( Hesperochernes )fromabird'snestinMontana(ArachnidaChelonethida).Pan-PacificEntomologist11:3739. Chamberlin,J.C.1943.Thetaxonomyofthefalsescorpiongenus Synsphyronus withremarksofthesporadiclossofstabilityingenerally constantmorphologicalcharacters(Arachnida:Chelonethida).AnnalsoftheEntomologicalSocietyofAmerica36:486500. Chamberlin,J.C.1952.Newandlittle-knownfalsescorpions (Arachnida,Chelonethida)fromMontereyCounty,California. BulletinoftheAmericanMuseumofNaturalHistory99:259312. Culver,D.C.2005.Speciesinteractions.Pp.539543. In Encyclopedia ofCaves.(D.C.Culver&W.B.White,eds.).Elsevier,Burlington, Massachusetts. Ellingsen,E.1910.DiePseudoskorpionedesBerlinerMuseums. MitteilungausdemZoologischenMuseuminBerlin4:357423. Gardini,G.1983. Larcaitalica n.sp.cavernicoladell'Appennino Abruzzese(Pseudoscorpionida,Garypidae)(Pseudoscorpionid'ItaliaXV).BollettinodellaSocieta `EntomologicaItaliana115:6369. Harvey,M.S.1986.TheAustralianGeogarypidae,newstatus,witha reviewofthegenericclassification(Arachnida:Pseudoscorpionida).AustralianJournalofZoology34:753778. Harvey,M.S.1987a.Redescriptionsof Geogarypusbucculentus Beier and G.pustulatus Beier(Geogarypidae:Pseudoscorpionida). BulletinoftheBritishArachnologicalSociety7:137141. Harvey,M.S.1987b.Arevisionofthegenus Synsphyronus Chamberlin(Garypidae:Pseudoscorpionida:Arachnida).AustralianJournalofZoology,SupplementarySeries126:199. Harvey,M.S.1992.Thephylogenyandclassificationofthe Pseudoscorpionida(Chelicerata:Arachnida).InvertebrateTaxonomy6:13731435. Harvey,M.S.2002.Short-rangeendemismintheAustralianfauna: someexamplesfromnon-marineenvironments.Invertebrate Systematics16:555570. Harvey,M.S.2011.Twonewspeciesof Synsphyronus (Pseudoscorpiones:Garypidae)fromsouthernWesternAustraliangranite landforms.RecordsoftheWesternAustralianMuseum26:1122. Harvey,M.S.&K.L.Edward.2007.Areviewofthepseudoscorpion genus Ideoblothrus (Pseudoscorpiones,Syarinidae)fromwestern andnorthernAustralia.JournalofNaturalHistory41:445472. Harvey,M.S.&W.B.Muchmore.2013.Thesystematicsofthe pseudoscorpionfamilyIdeoroncidae(Pseudoscorpiones,Neobisioidea)intheNewWorld.JournalofArachnology41:229290. Harvey,M.S.,P.B.Ratnaweera,P.V.Randeniya&M.R.Wijesinghe. 2012.Anewspeciesofthepseudoscorpiongenus Megachernes (Pseudoscorpiones:Chernetidae)associatedwithathreatenedSri Lankanrainforestrodent,withareviewofhostassociationsof Megachernes .JournalofNaturalHistory46:25192535. Harvey,M.S.,M.G.Rix,V.W.Framenau,Z.R.Hamilton,M.S. Johnson,R.J.Teale,G.Humphreys&W.F.Humphreys.2011. Protectingtheinnocent:studyingshort-rangeendemictaxa enhancesconservationoutcomes.InvertebrateSystematics25: 110. Harvey,M.S.,W.A.Shear&H.Hoch.2000.Onychophora, Arachnida,myriapodsandInsecta.Pp.7994. In Subterranean ecosystems.(H.Wilkens,D.C.Culver&W.F.Humphreys,eds.). Elsevier,Amsterdam. Henderickx,H.&V.Vets.2002.Anew Larca (Arachnida: Pseudoscorpiones:Larcidae)fromCrete.BulletinoftheBritish ArachnologicalSociety12:280282. Heurtault,J.1994.Pseudoscorpions.Pp.185196. In Encyclopaedia biospeologica.(C.Juberthie&V.Decu,eds.).Vol.1.Socie te de Biospeologie,MoulisandBucarest. Hoff,C.C.1945. Hesperochernescanadensis ,anewchernetidpseudoscorpionfromCanada.AmericanMuseumNovitates1273:14. Hoff,C.C.1946a.Newpseudoscorpions,chieflyneotropical,ofthe suborderMonosphyronida.AmericanMuseumNovitates1318:132. Hoff,C.C.1946b.Astudyofthetypecollectionsofsome pseudoscorpionsoriginallydescribedbyNathanBanks.Journal oftheWashingtonAcademyofSciences36:195205. Hoff,C.C.1947.Thespeciesofthepseudoscorpiongenus Chelanops describedbyBanks.BulletinoftheMuseumofComparative Zoology98:471550. Hoff,C.C.1950.Pseudoescorpionidosnuevosopococonocidosdela Argentina(Arachnida,Pseudoscorpionida).Arthropoda,Buenos Aires1:225237. Hoff,C.C.1956a.DiplosphyronidpseudoscorpionsfromNew Mexico.AmericanMuseumNovitates1780:149. Hoff,C.C.1956b.PseudoscorpionsofthefamilyChernetidaefrom NewMexico.AmericanMuseumNovitates1800:166. Hoff,C.C.1961.PseudoscorpionsfromColorado.Bulletinofthe AmericanMuseumofNaturalHistory122:409464. Hoff,C.C.&J.E.Bolsterli.1956.PseudoscorpionsoftheMississippi Riverdrainagebasinarea.TransactionsoftheAmerican MicroscopicalSociety75:155179. Hoff,C.C.&D.L.Clawson.1952.Pseudoscorpionsfromrodent nests.AmericanMuseumNovitates1585:138. Howarth,F.G.1980.Thezoogeographyofspecializedcaveanimals:a bioclimaticmodel.Evolution34:394406. Howarth,F.G.1982.Bioclimaticandgeologicalfactorsgoverningthe evolutionanddistributionofHawaiiancaveinsects.Entomologia Generalis8:1726. Howarth,F.G.1983.Ecologyofcavearthropods.AnnualReviewof Entomology28:365389. 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Muchmore,W.B.1981.Cavernicolousspeciesof Larca Archeolarca and Pseudogarypus withnotesonthegenera,(Pseudoscorpionida, GarypidaeandPseudogarypidae).JournalofArachnology9:47 60. Muchmore,W.B.1982.Thegenus Anagarypus (Pseudoscorpionida: Garypidae).PacificInsects24:159163. Muchmore,W.B.1984.Newcavernicolouspseudoscorpionsfrom California(Pseudoscorpionida,ChthoniidaeandGarypidae). JournalofArachnology12:171175. Muchmore,W.B.1990.Pseudoscorpionida.Pp.503527. In Soil biologyguide.(D.L.Dindal,ed.).JohnWileyandSons,NewYork. Muchmore,W.B.1994.Somepseudoscorpions(Arachnida:Pseudoscorpionida)fromcavesinOhioandIndiana,U.S.A.Transactions oftheAmericanMicroscopicalSociety113:316324. Muchmore,W.B.1996.Aremarkablenewgenusandspeciesof Pseudoscorpionida(Syarinidae)fromacaveinArizona.SouthwesternNaturalist41:145148. Muchmore,W.B.1997. Tuberochernes (Pseudoscorpionida,Chernetidae),anewgenuswithspeciesincavesinCaliforniaandArizona. JournalofArachnology25:206212. 218 THEJOURNALOFARACHNOLOGY

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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


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Cras ut cursus ante, a fringilla nunc. Mauris lorem nunc, cursus sit amet enim ac, vehicula vestibulum mi. Mauris viverra nisl vel enim faucibus porta. Praesent sit amet ornare diam, non finibus nulla.

MLA

Cras efficitur magna et sapien varius, luctus ullamcorper dolor convallis. Orci varius natoque penatibus et magnis dis parturient montes, nascetur ridiculus mus. Fusce sit amet justo ut erat laoreet congue sed a ante.

CHICAGO

Phasellus ornare in augue eu imperdiet. Donec malesuada sapien ante, at vehicula orci tempor molestie. Proin vitae urna elit. Pellentesque vitae nisi et diam euismod malesuada aliquet non erat.

WIKIPEDIA

Nunc fringilla dolor ut dictum placerat. Proin ac neque rutrum, consectetur ligula id, laoreet ligula. Nulla lorem massa, consectetur vitae consequat in, lobortis at dolor. Nunc sed leo odio.