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Journal of cave and karst studies

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Journal of cave and karst studies
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Journal of Cave & Karst Studies
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Continues NSS bulletin (OCLC: 2087737)
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National Speleological Society
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Article DOI: http://dx.doi.org/10.4311/2011LSC0252 Subterranean Aquatic Planarians of Sardinia, with a Discussion on the Penial Flagellum and the Bursal Canal Sphincter in the Genus Dendrocoelum (Platyhelminthes, Tricladida, Dendrocoelidae) Giacinta Angela Stocchino, Ronald Sluys, Palolo Marcia, and Renata Manconi Article DOI: http://dx.doi.org/10.4311/2011LSC0226 Occurrence of Troglobitic Clivinines in China (Insecta: Coleoptera: Carabidae) Mingyi Tian Article DOI: http://dx.doi.org/10.4311/2011LSC0256 The First Cavernicolous Nicoletiidae (Insecta: Zygentoma) from the United Arab Emirates Luis Espinasa and Luis F. Mendes Article DOI: http://dx.doi.org/10.4311/2011AN0233 The View of Maya Cave Ritual from the Overlook Rockshelter, Caves Branch River Valley, Central Belize Gabriel D. Wrobel, Rebecca Shelton, Shawn Morton, Joshua Lynch, and Christopher Andres Article DOI: http://dx.doi.org/10.4311/2011ES0262 Flow Characterization in the Santee Cave System in the Chapel Branch Creek Watershed, Upper Coastal Plain of South Carolina, USA Amy E. Edwards, Devendra M. Amatya, Thomas M. Williams, Daniel R. Hitchcock, and April L. James Article DOI: http://dx.doi.org/10.4311/2011LSC0257 New Species and New Records of Springtails (Hexapoda: Collembola) from Caves in the Salem Plateau of Illinois, USA Felipe N. Soto-Adames and Steven J. Taylor
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Vol. 75, no. 2 (2013)
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Journal of Cave and Karst Studies Volume 75 Number 2 August 2013Article 93Subterranean Aquatic Planarians of Sardinia, with a Discussion on the Penial Flagellum and the Bursal Canal Sphincter in the Genus Dendrocoelum (Platyhelminthes, Tricladida, Dendrocoelidae) Giacinta Angela Stocchino, Ronald Sluys, Palolo Marcia, and Renata ManconiArticle 113Occurrence of Troglobitic Clivinines in China (Insecta: Coleoptera: Carabidae) Mingyi TianArticle 121The First Cavernicolous Nicoletiidae (Insecta: Zygentoma) from the United Arab Emirates Luis Espinasa and Luis F. MendesArticle 126The View of Maya Cave Ritual from the Overlook Rockshelter, Caves Branch River Valley, Central Belize Gabriel D. Wrobel, Rebecca Shelton, Shawn Morton, Joshua Lynch, and Christopher AndresArticle 136Flow Characterization in the Santee Cave System in the Chapel Branch Creek Watershed, Upper Coastal Plain of South Carolina, USA Amy E. Edwards, Devendra M. Amatya, Thomas M. Williams, Daniel R. Hitchcock, and April L. JamesArticle 146New Species and New Records of Springtails (Hexapoda: Collembola) from Caves in the Salem Plateau of Illinois, USA Felipe N. Soto-Adames and Steven J. TaylorJournal of Cave and Karst StudiesVolume 75 Number 2 August 2013 August 2013 Volume 75, Number 2 ISSN 1090-6924 A Publication of the National Speleological Society JOUR NA L OF C AV E AN D K A R S T STUD I E S rf

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G U I DE TO A UT HOR S The Journal of Cave and Karst Studies is a multidisciplinary journal devoted to cave and karst research. The Journal is seeking original, unpublished manuscripts concerning the scientic study of caves or other karst features. Authors do not need to be members of the National Speleological Society, but preference is given to manuscripts of importance to North American speleology. LANGUAGES: The Journal of Cave and Karst Studies uses American-style English as its standard language and spelling style, with the exception of allowing a second abstract in another language when room allows. In the case of proper names, the Jour nal tries to accommodate other spellings and punctuation styles. In cases where the Editor-in-Chief nds it appropriate to use nonEnglish words outside of proper names (generally where no equivalent English word exists), the Journal italicizes them. However, the common abbreviations i.e., e.g., et al., and etc. should appear in roman text. Authors are encouraged to write for our combined professional and amateur readerships. CONTENT: Each paper will contain a title with the authors names and addresses, an abstract, and the text of the paper, including a summary or conclusions section. Acknowledgments and references follow the text. ABSTRACTS: An abstract stating the essential points and results must accompany all articles. An abstract is a summary, not a promise of what topics are covered in the paper. STYLE: The Journal consults The Chicago Manual of Style on most general style issues. REFERENCES: In the text, references to previously published work should be followed by the relevant authors name and date (and page number, when appropriate) in parentheses. All cited references are alphabetical at the end of the manuscript with senior authors last name rst, followed by date of publication, title, publisher, volume, and page numbers. Geological Society of America for mat should be used (see http://www.geosociety.org/pubs/geoguid5. htm). Please do not abbreviate periodical titles. Web references are acceptable when deemed appropriate. The references should follow the style of: Author (or publisher), year, Webpage title: Publisher (if a specic author is available), full URL (e.g., http://www. usgs.gov/citguide.html) and date when the web site was accessed in brackets; for example [accessed July 16, 2002]. If there are specic authors given, use their name and list the responsible organization as publisher. Because of the ephemeral nature of websites, please provide the specic date. Citations within the text should read: (Author, Year). SUBMISSION: Effective February 2011, all manuscripts are to be submitted via Peertrack, a web-based system for online submission. The web address is http://www.edmgr.com/jcks. Instructions are provided at that address. At your rst visit, you will be prompted to establish a login and password, after which you will enter information about your manuscript (e.g., authors and addresses, manuscript title, abstract, etc.). You will then enter your manuscript, tables, and gure les separately or all together as part of the manuscript. Manuscript les can be uploaded as DOC, WPD, RTF, TXT, or LaTeX. A DOC template with additional manuscript specications may be downloaded. (Note: LaTeX les should not use any unusual style les; a LaTeX template and BiBTeX le for the Journal may be downloaded or obtained from the Editor-inChief.) Table les can be uploaded as DOC, WPD, RTF, TXT, or LaTeX les, and gure les can be uploaded as TIFF, EPS, AI, or CDR les. Alternatively, authors may submit manuscripts as PDF or HTML les, but if the manuscript is accepted for publication, the manuscript will need to be submitted as one of the accepted le types listed above. Manuscripts must be typed, double spaced, and single-sided. Manuscripts should be no longer than 6,000 words plus tables and gures, but exceptions are permitted on a case-bycase basis. Authors of accepted papers exceeding this limit may have to pay a current page charge for the extra pages unless decided otherwise by the Editor-in-Chief. Extensive supporting data will be placed on the Journals website with a paper copy placed in the NSS archives and library. The data that are used within a paper must be made available. Authors may be required to provide supporting data in a fundamental format, such as ASCII for text data or comma-delimited ASCII for tabular data. DISCUSSIONS: Critical discussions of papers previously published in the Journal are welcome. Authors will be given an opportunity to reply. Discussions and replies must be limited to a maximum of 1000 words and discussions will be subject to review before publication. Discussions must be within 6 months after the original article appears. MEASUREMENTS: All measurements will be in Systeme Internationale (metric) except when quoting historical references. Other units will be allowed where necessary if placed in parentheses and following the SI units. FIGURES: Figures and lettering must be neat and legible. Figure captions should be on a separate sheet of paper and not within the gure. Figures should be numbered in sequence and referred to in the text by inserting (Fig. x). Most gures will be reduced, hence the lettering should be large. Photographs must be sharp and high contrast. Color will generally only be printed at authors expense. TABLES: See http://www.caves.org/pub/journal/PDF/Tables. pdf to get guidelines for table layout. COPYRIGHT AND AUTHORS RESPONSIBILITIES: It is the authors responsibility to clear any copyright or acknowledgement matters concerning text, tables, or gures used. Authors should also ensure adequate attention to sensitive or legal issues such as land owner and land manager concerns or policies. PROCESS: All submitted manuscripts are sent out to at least two experts in the eld. Reviewed manuscripts are then returned to the author for consideration of the referees remarks and revision, where appropriate. Revised manuscripts are returned to the appropriate Associate Editor who then recommends acceptance or rejection. The Editor-in-Chief makes nal decisions regarding publication. Upon acceptance, the senior author will be sent one set of PDF proofs for review. Examine the current issue for more information about the format used. ELECTRONIC FILES: The Journal is printed at high resolution. Illustrations must be a minimum of 300 dpi for acceptance.The Journal of Cave and Karst Studies (ISSN 1090-6924, CPM Number #40065056) is a multi-disciplinary, refereed journal published three times a year by the National Speleological Society, 2813 Cave Avenue, Huntsville, Alabama 35810-4431 USA; Phone (256) 852-1300; Fax (256) 851-9241, email: nss@caves.org; World Wide Web: http://www.caves.org/pub/journal/. Check the Journal website for subscripion rates. Back issues and cumulative indices are available from the NSS ofce. POSTMASTER: send address changes to the Journal of Cave and Karst Studies, 2813 Cave Avenue, Huntsville, Alabama 35810-4431 USA. The Journal of Cave and Karst Studies is covered by the following ISI Thomson Services Science Citation Index Expanded, ISI Alerting Services, and Current Contents/Physical, Chemical, and Earth Sciences. Copyright 2013 by the National Speleological Society, Inc. Front cover: A collection of planarians from Sardinia. See Stocchino et. al. in this issue.Published By The National Speleological SocietyEditor-in-Chief Malcolm S. FieldNational Center of Environmental Assessment (8623P) Ofce of Research and Development U.S. Environmental Protection Agency 1200 Pennsylvania Avenue NW Washington, DC 20460-0001 703-347-8601 Voice 703-347-8692 Fax eld.malcolm@epa.govProduction EditorScott A. EngelCH2M HILL 2095 Lakeside Centre Way, Suite 200 Knoxville, TN 37922 865-560-2954 scott.engel@ch2m.comJournal Copy EditorBill MixonJOURNAL AD VISOR Y BO ARD Penelope Boston Gareth Davies Luis Espinasa Derek F ord Louise Hose Leslie Melim Wil Orndorf Bill Shear Dorothy Vesper BO ARD OF EDITORS AnthropologyGeorge Crothers University of Kentucky211 Lafferty Hall george.crothers@uky.eduConservation-Life SciencesJulian J. Lewis & Salisa L. LewisLewis & Associates, LLC. lewisbioconsult@aol.comEarth SciencesBenjamin SchwartzDepartment of Biology Texas State University bs37@txstate.eduRobert BrinkmanDepartment of Geology, Environment, and Sustainability Hofstra University robert.brinkmann@hofstra.eduMario PariseNational Research Council, Italy m.parise@ba.irpi.cnr.itExplorationPaul BurgerCave Resources Ofce National Park Service Carlsbad, NM paul_burger@nps.govMicrobiologyKathleen H. LavoieDepartment of Biology State University of New York, Plattsburgh, lavoiekh@plattsburgh.eduPaleontologyGreg McDonaldPark Museum Management Program National Park Service, Fort Collins, CO greg_mcdonald@nps.govSocial SciencesJoseph C. DouglasHistory Department Volunteer State Community College joe.douglas@volstate.eduBook ReviewsArthur N. Palmer & Margaret V. PalmerDepartment of Earth Sciences State University of New York, Oneonta palmeran@oneonta.edu

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SUBTERRANEANAQUATICPLANARIANSOFSARDINIA, WITHADISCUSSIONONTHEPENIALFLAGELLUMAND THEBURSALCANALSPHINCTERINTHEGENUS DENDROCOELUM (PLATYHELMINTHES, TRICLADIDA,DENDROCOELIDAE) G IACINTA A NGELA S TOCCHINO 1 ,R ONALD S LUYS 2 ,P AOLO M ARCIA 1 AND R ENATA M ANCONI 1 Abstract: Thepaperprovidesthefirstdetailedaccountonthetaxonomicrichnessofthe subterraneanfreshwatertricladsfromSardinia,includingthedescriptionoffournew speciesforthegenera Dendrocoelum and Phagocata .Newrecordsfor Dugesiabenazzii Dugesia sp., Crenobiaalpina ,and Phagocata sp.arealsoreported.Thethreenewspecies of Dendrocoelum arethefirstreportedfortheislandofSardinia.Thesespeciesdisplaya bursalcanalsphincterandalargeadenodactylwithacharacteristicanatomywithazone offinecircularmusclefibersrunningthroughthemesenchymeofitspapilla.Adetailed analysisofthestructureofthepenialflagelluminthegenus Dendrocoelum highlightedsix mainconditions,someofwhichhavenotbeenpreviouslyreported,inregardtothe histologyofthetipofthepenispapillaandtheextentofitsinversion.Thenewspeciesof Phagocata representsthefirstspeciesrecordedfromItalyandthefirstanophtalmous speciesreportedfromEurope. I NTRODUCTION TheMediterraneanregionhasbeenidentifiedasa biodiversityhotspotonaglobalscale,withtheislandof Sardiniaasoneofitsregionalhotspots(Me dailand Que zel,1999;Myersetal.,2000).Togetherwithpartof coastalTuscany,theislandbelongstotheSardinian stygofaunisticprovince(Pesce,1985).Thepresenceofa greatlydiversifiedandancientkarst,datingtothe PalaeozoicandMesozoicperiodsoftheSardinian-Corsica plate,gaverisetoawiderangeofundergroundaquatic systems,rangingfrominlandandcoastalcavesto undergroundrivers(Pesce,1985).Althoughmorethan 3000terrestrialandmarinekarsticcavesareregisteredin theRegionalSpeleologicalRegister,thegroundwater biodiversityofSardiniaislargelyunknown.Theliterature onthetaxonomicrichnessinthecaves,springs,andwells ofSardiniaismainlyfocusedonCrustacea(Lindberg, 1956;Stella,1957;PudduandPirodda,1973;Ruffoand VignaTaglianti,1975;Cassola,1982;PesceandMaggi, 1983;Pesce,1985;Cottarellietal.,1996;Arganoetal., 1997).AfewpapersrefertofreshwaterOligochaeta (Martinez-AnsemilandSambugar,2008),Gastropoda (GiustiandCastagnolo,1983),Trichoptera(Morettiand Cianficconi,1983;Cianficconietal.,1998),andmarine cave-dwellingfauna(seeManconietal.,2009and referencestherein). FaunisticinvestigationsofSardinianepigeanwaters, promotedfirstbyBenazzi(1938)andsubsequentlyfurther developedbyLeporiandco-workers,highlighteda considerabletaxonomicrichnessandendemicityofthe Tricladida(Lepori,1951;Palaetal.,1980a,b,c,1981,1995, 1999,2000;Casuetal.,1982;Stocchinoetal.,2005).In contrast,veryfewdataareavailableontheSardinian groundwatertriclads,datingbackto1938whenBenazzi recorded Atrioplanaria sp.fromahydropetrichabitaton limestoneinnorth-westernSardinia(Benazzi,1938).Other specimens,discoveredin1952fromSuColoruCave (northernSardinia)andfromanoldmine(BaccuArrosu, southwesternSardinia),wereassignedbythesameauthor to Atrioplanaria sp.(Benazzi,1982).Palaetal.(1980c) reportedon Crenobiaalpina Dana,1776fromsomesprings ontheGennargentuMassif(central-easternSardinia). Apartfromthesedatanonewrecordsofsubterranean tricladsfromtheislandhavebecomeavailableforalong periodoftime.Onlyrecentlyhaveafewstygobiological studiesonSardiniantricladsbeenperformed(Stocchino, 2003;Stocchinoetal.,2008).Thepresentpaperprovidesa detailedaccountonthetaxonomicrichnessofthe subterraneantricladsfromSardinia,includingthedescriptionoffournewspeciesandnewrecordsforseveralothers. M ATERIALSAND M ETHODS Thespecimens,collectedduringtheperiod2000–2010, weretransferredtothelaboratory,rearedinshadedboxes insemi-darkconditionsat18 6 2 u Candfedwithfresh beefliver.Formorphologicalstudy,specimenswerefixed *Correspondingauthor:stocchin@uniss.it 1 DipartimentodiScienzedellaNaturaedelTerritorio,Universita `diSassari,Via Muroni25,I-07100,Sassari,Italy 2 NetherlandsCentreforBiodiversityNaturalis(sectionZMA),P.O.Box9514,2300 RALeiden,TheNetherlands,andInstituteforBiodiversityandEcosystem Dynamics,UniversityofAmsterdam G.A.Stocchino,R.Sluys,P.Marcia,andR.Manconi–SubterraneanaquaticplanariansofSardinia,withadiscussiononthepenial flagellumandthebursalcanalsphincterinthegenus Dendrocoelum (Platyhelminthes,Tricladida,Dendrocoelidae). JournalofCaveand KarstStudies, v.75,no.2,p.93–112.DOI:10.4311/2011LSC0252 JournalofCaveandKarstStudies, August2013 N 93

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for24hoursinBouin’sfluid,dehydratedinagraded ethanolseries,clearedintoluene,andembeddedin paraffin.SerialsectionsofspecimensofthePuntadella ScomunicaSpringweremadeatintervalsof5,6,or7 m m andstainedwithhaematoxylin-eosin(Harris),Hansen’s haematoxylinandeosin-orange,DaneHerman’stetrachrome,Mallory-Cason,orPasini’sreagent. SerialsectionsofspecimensfromtheMonteMajore Caveweremadeatintervalsof5or8 m mandstainedin DaneHerman’stetrachromeorMallory-Cason.Serial sectionsofspecimensfromSaUcca‘esuPeltusuCave weremadeatintervalsof8 m mandwerestainedwith Mallory-Cason.Serialsectionsofthespecimenfromthe CuccuruTiriaCaveweremadeatintervalsof5 m mand stainedinDaneHerman’stetrachrome.Serialsectionsof specimensfromtheSuCantaruSpringweremadeat intervalsof5 m mandstainedinhaematoxylin-eosin (Harris).SerialsectionsofspecimensfromtheGennargentuMassifspringweremadeatintervalsof8 m mand stainedinHarris’shaematoxylin-eosin.Thematerialis depositedintheNetherlandsCentreforBiodiversity Naturalis(sectionZMA)andintheGiacintaA.Stocchino collection(CGAS),UniversityofSassari. S YSTEMATIC A CCOUNT (AllabbreviationsforlabelsinFigures2–9 arelistedinTable1) OrderTRICLADIDALang,1884 SuborderCONTINENTICOLACarranza,Littlewood, Clough,Ruiz-Trillo,Bagun aandRiutort,1998 SuperfamilyPLANARIOIDEAStimpson,1857 FamilyDENDROCOELIDAEHallez,1892 Genus Dendrocoelum O ¨ rsted,1844 Dendrocoelummariae StocchinoandSluyssp.nov. (Table2;Figs.1,2) Materialexamined .Allindividuals(n 5 20)wereasexualat collection.Themajorityoftheanimalsunderwent asexualizationprocessafterayearofrearinginthe laboratory. Holotype:ZMAV.Pl.7100.1,sagittalsectionson17 slides,PuntadellaScomunica,AsinaraIsland(41 u 05 9 35 0 N, 8 u 18 9 4 0 E),December2000,coll.M.PirasandG.A. Stocchino. Paratypes:ZMAV.Pl.7100.2,ZMAV.Pl.7100.3,ZMA V.Pl.7100.4,ibid.,sagittalsectionson41slides,22slides, 22slides;ZMAV.Pl.7100.5,ibid.,onesetoftransverse sectionson61slides;CGASPla1.1,ibid.,sagittalsections on37slides,immaturespecimen;CGASPla1.2,ibid., horizontalsectionson27slides,notcompletelymature specimen. Othermaterial:ZMAV.Pl.7101.1,ZMAV.Pl.7101.2, twosetsofsagittalsectionson5slidesand19slides, respectively,notcompletelymaturespecimens.Punta dellaScomunica,AsinaraIsland(41 u 05 9 35 0 N8 u 18 9 4 0 E) May2005,coll.G.A.Stocchino;CGASPla1.3,ibid.one setofsagittalsectionson12slides;CGASPla1.4–12ibid., ninesetsofsagittalsectionson13slides,13slides,23slides, 14slides,16slides,18slides,22slides,21slides,37slides, respectively,notcompletelymaturespecimens.CGASPla 1.13–14,twosetsoftransversesectionson21slidesand46 slides,respectively,notcompletelymaturespecimens,Punta dellaScomunica,AsinaraIsland(41 u 05 9 35 0 N8 u 18 9 4 0 E) spring2000,coll.M.PirasandG.A.Stocchino. Etymology.ThespecificepithetreferstoProfessor emeritusMariaPala(UniversityofSassari),inrecognition ofhercontributionstothebiologyandsystematicsof freshwatertriclads. Table1.Abbreviationsusedinfigures. AbbreviationTerm a adenodactyl ab bulboftheadenodactyl ap papillaoftheadenodactyl bc bursalcanal bg bulbglands ca commonatrium cb copulatorybursa cm circularmuscles co copulatoryapparatus cod commonoviduct cvs commonvasdeferens e eyes ed ejaculatoryduct ep epithelium eep externalepithelium f flagellum fcm finecircularmuscles g gonopore h head ic intestinalcaecum iep internalepithelium l lumen lm longitudinalmuscles m musculature ma maleatrium od oviduct pb penisbulb pg penisglands ph pharynx pl plugofcells pp penispapilla ppl penispapillalumen s sphincter sg shellglands sp sperm spv spermiducalvesicles sv seminalvesicle vd vasdeferens vdv vesicleofthevasdeferens S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 94 N JournalofCaveandKarstStudies, August2013

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Habitat.From2000to2005(springandwinter) specimensof D.mariae werecollectedfromasmallperennial spring(atanaltitudeofca.350masl)ontheeasterncoastof AsinaraIsland(Fig.1).Theseasonalsamplingshighlighted theconstantpresenceofplanariansatthissite.Substrate rangesfromrockstolargebouldersandsand.Thespring ischaracterizedbyanextremelyfluctuatingflow,with minimumvaluesduringthesummer.Watertemperature rangesbetween13and21.4 u Cthroughouttheyear;pH valuesrangefrom7to7.3.Thewaterisrichinminerals (totaldissolvedsolidsca.600mgL 2 1 )andischaracterized byhighvaluesofchlorides,duetothesaltyaerosolblownin bywind.Dendrocoelidaewereassociatedwithadiverse invertebratefauna,consistingofGastropoda,Tubificidae, Ostracoda,Cyclopoida,Asellidae,Amphipoda,Hydracnidia,Chironomidae,andtheendemicanuran Discoglossus sardus Tschudi,1837. Geographicaldistribution.EndemictoAsinaraIsland andonlyknownfromthetypelocality. Diagnosis. Dendrocoelummariae ischaracterizedbyan adenodactylthatismuchlargerthanthepenispapilla,a copulatoryapparatusfarbehindthepharyngealpocket, thepresenceofasphincterintheterminaltractofthe bursalcanal,andbythepenisbeingindorsalpositionand theadenodactyllocatedventrally. Description.Liveanimalsareunpigmented,typically whitish,withabodysizeof8to10mminlengthanda widthrangingfromca.2mminthecentralpartofthebody tolessthan1mmatthelevelofthehead.Theanteriorend istruncated,withthemiddlepartofthefrontalmargin convex,andisprovidedwithapairofroundedlateral lobes.Justbehindtheeyesthereisaslightbutclear constrictionorneckthatsetsofftheheadfromtherestof thebody(Fig.2A). Table2.ChecklistofgroundwatertricladsfromSardinia. TaxaCavesSpringsLocality Coordinates Reference LatitudeLongitude DugesiidaeBall,1974 Dugesia Girard,1850 Dugesiabenazzii Lepori,19511Su CantaruSpring, MonteAlbo 41 u 34 9 30 0 N9 u 40 9 31 0 EPresentpaper Dugesia sp.1SaUcca‘esu PeltusuCave 40 u 27 9 00 0 N8 u 40 9 43 0 EPresentpaper DendrocoelidaeHallez,1892 Dendrocoelum O ¨ rsted,1844 Dendrocoelummariae 1PuntadellaScomunica Spring,AsinaraIsland 41 u 05 9 35 0 N8 u 18 9 4 0 EPresentpaper Dendrocoelumnuraghum 1MonteMajoreCave40 u 30 9 51 0 N8 u 36 9 37 0 EPresentpaper Dendrocoelumvesiculosum 1SaUcca‘esu PeltusuCave 40 u 27 9 00 0 N8 u 40 9 43 0 EPresentpaper PlanariidaeStimpson,1857 Atrioplanaria deBeauchamp,1932 Atrioplanaria sp.1Sassarisurroundings40 u 42 9 02 0 N8 u 35 9 47 0 EBenazzi,1938 Atrioplanaria sp.1SuColoruCave40 u 49 9 01 0 N8 u 48 9 46 0 EBenazzi,1982 Atrioplanaria sp.1BaccuArrosuMine39 u 9 9 36 0 N8 u 45 9 15 0 EBenazzi,1982 Phagocata Leidy,1857 Phagocataobscura 1CuccuruTiriaCave39 u 19 9 25 0 N8 u 34 9 29 0 EPresentpaper Phagocata sp.1MonteMajoreCave40 u 30 9 51 0 N8 u 36 9 37 0 EPresentpaper Phagocata sp.1EligheMannuSpring, AsinaraIsland 41 u 05 9 36 0 N8 u 18 9 21 0 EPresentpaper Phagocata sp.1Ala `deiSardi40 u 38 9 60 0 N9 u 19 9 45 0 EPresentpaper Crenobia Kenk,1930 Crenobiaalpina (Dana,1766)4EastGennargentu Massif 40 u 09 9 24 0 N9 u 15 9 16 0 EPalaetal., 1980c C.alpina 1SuSesseneSpring,West GennargentuMassif 40 u 01 9 40 0 N9 u 12 9 1 0 EPresentpaper G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 95

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Thetwoeyesaresituatedrathercloselytogether.The distancebetweentheeyesisslightlylessthan1/4ofthe widthoftheneck.Inter-oculardistanceissmallerthan thedistancefromeacheyecuptothelateralmargin, andthedistancefromthefrontalmargintotheeyesis greaterthanthedistancetothelateralmargin.Supernumeraryeyesareoftenpresent,usually2or3oneither side,sometimesarrangedinaslightlycrescent-shaped configuration. Thesubterminalanterioradhesiveorgan,withalength comparabletotheinteroculardistance,ismoderately developedandconsistsofashallowcup.Thesurfaceof theorganconsistsofawell-definedareaofinfranucleate epithelialcellspiercedbynumerousglandducts.Thecell bodiesoftheglandsarescatteredthroughoutthemesenchymeoftheanteriorpartofthebody.Twotypesofglands, stronglyorweaklyacidophilic,arerecognizablewithDane HermanÂ’stetrachrome,PasiniÂ’sreagent,andhaematoxylineosin(Harris)stains.Themusculatureassociatedwiththis organconsistsofamorestronglydevelopedsectionofthe ventrallongitudinalbodymusculature. Theintestineiswellvisible,theanteriorramusreaches toaleveljustbehindtheneckandbears13to15branches oneachside.Thetwoposteriorrami,eachwith16or17 branches,usuallyconvergebehindthecopulatoryapparatustoformacommonbranch.Thepharynxislocatedin theposteriorhalfofthebodyandmeasuresabout1/8of thebodylength(Fig.2A).Itsinternalmusclezoneconsists ofaverythicklayerofintermingledcircularand longitudinalfibers.Asubepitheliallayeroflongitudinal musclesfollowedbyalayerofcircularfibersformsthethin outerzoneofmuscles. Thetwoovariesoccurontheventralsideoftheanterior regionbehindthebrain.Theovariesarelocatedat1/5ofthe distancebetweenthebrainandtherootofthepharynx.The twooviductsoriginatefromthedorsalpartoftheovaries andareprovidedwithaslightexpansionattheiranterior end,thetuba.Theoviductsrunposteriorly,fusingbehind thecopulatoryapparatustoformaratherlongcommon oviduct.Thelatterrunsanteriorlytotherightsideofthe openingofthebursalcanalintotheatrium.Thecommon oviductreceivesnumerousopeningsofeosinophilicshell Figure1.GeographicdistributionofgroundwaterplanariansinSardinia.DatafromBenazzi(1938,1982),Palaetal. (1980c),andthisstudy. S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 96 N JournalofCaveandKarstStudies, August2013

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glandsalongitsentirelength.Thevitellariaarelocated betweentheintestinaldiverticulaandthetestes. Thewell-developedtestesarenumerousandessentially ventralinposition(inonespecimensomefolliclesare situatedbetweentheintestinaldiverticula).Thetestes extendfromashortdistancebehindtheovariestothefar posteriorendofthebodyandarearrangedoneithersideof themidlineintwotothreelongitudinalzones.Thesperm ductsformwell-developedspermiducalvesicles,packed withsperm,betweenthemouthandtheanteriorlevelofthe penisbulb(Fig.2B). Thecopulatoryapparatusoccupiestheposteriorhalfof thepostpharyngealregion.Theelongated,smallcopulatory bursaissac-shapedandsituatedinthedorsalpartofthe body.Thebursaislinedwithahighglandularepithelium andissurroundedbyalayeroflongitudinalmuscles.The bursalcanalrunsposteriorlytotheleftofthepenisand graduallywidens,thenturnsventrallyandafternarrowing againopensintothecommonatrium.Thewallofthebursal canalconsistsofanucleatedepitheliumwithciliatedcells thatgraduallyrangefromcuboidalintheproximaltractto cylindricalinthemoredistalsectionofthecanal.Thecanal issurroundedbyasubepitheliallayerofcircularmuscles followedbyalayeroflongitudinalfibers.Justbeforethe openingintotheatriumthelayerofcircularmuscles becomesthickerandthusformsasphincter(Fig.2B). Figure2. Dendrocoelummariae .(A)habitusofalivingspecimen,(B)holotypeZMAV.Pl.7100.1,sagittalreconstructionof thecopulatoryapparatus(anteriortotheright). G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 97

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Inthemalecopulatoryapparatusthesmall,muscular penisbulbhousesaseminalvesicle,devoidofaproper epithelium,thatissurroundedbyseveralinterwovenlayers ofmuscle.Thevasadeferentiaentertheseminalvesicle closelytogetherfromtheantero-lateralsidesandopen symmetricallyinitsanteriorpart.Theseminalvesicle communicateswiththeejaculatoryductthatopensatthe tipofthesmallpenispapilla.Thepenispapillaisslightly asymmetrical,withtheventralpartsomewhatlargerthan thedorsalonebecauseofthedorsallydisplacedpositionof theejaculatoryduct.Thepenispapillashowsadivision intotwoparts,abasalpartwithathinepithelium underlainbyastronglayerofcircularmusclesandadistal partcoveredwithclaviformepithelialcellswithbasal nuclei,butdevoidofmusculature.Thetwopartsare separatedbyaconstriction.Thepenispapillahasthesame lengthasthepenisbulb(Fig.2B).Inonlyonespecimen (ZMAV.Pl.7100.4)thepenispapillaistwiceaslargeasthe penisbulb.Inallexaminedspecimensnopenisglandswere evident.Thepenisislocateddorsallytotheadenodactyl, whilethebursalcanalissituatedtotheleftofthemidline. Thepenislengthisabout1/3to1/4ofthelengthofthe adenodactyldependingonthestateofcontractionofthe latterorgan. Theadenodactylisverylargeandconsistsofafree papillaandawell-developedbulbarpart.Thebulbis dorsal-ventralinorientation,andtheadenodactylmakesa sharpposteriorbendonthetransitionofthebulbtothe papilla.Inthemajorityoftheexaminedmaturespecimens theadenodactylisthrustoutoftheplanarianbody;in theholotypeZMAV.Pl.7100.1itstipispushedinto theparenchymacaudallytothecopulatoryapparatus (Fig.2B).Alloftheseconditionsareverylikelydueto preservationartifacts.Thebulbconsistsofrowsof alternatinglongitudinalandcircularmusclesfibers.The lumenoftheadenodactylislinedbyalayerofciliatedcells, anditissurroundedbyawell-developedzoneofglandular mesenchymatictissue.Throughthissectionofthemesenchymerunsalayerofcircularmusclethatisparticularly evidentinspecimensZMAV.Pl.7100.4,ZMAV.Pl. 7100.5,andZMAV.Pl.7101.2.Inthelast-mentioned specimen,stainedinMallory-Cason,thislayerofcircular muscleispaleblue,whereastheintermingledmusclesstain brightblue.Ectallytothiszoneofcircularmusclesrunsa layeroflongitudinalmusclesfibersthatstainsred. Themaleatriumislinedbyacolumnarepithelium,and itissurroundedbyasubepitheliallayerofcircularmuscles followedbythreelayersoflongitudinalfibers.Themale atriumcommunicatesviaaconstrictionwiththecommon atrium.Inthispartitreceivestheopeningofthecommon oviduct(Fig.2B). Discussion.Theexternalmorphologyof D.mariae resemblesthatof D.adenodactylosum (Stankovic and Koma rek,1927),bothintheshapeofthebodyandthe frequentpresenceofsupernumeraryeyes.However,in D. mariae theeyesarearrangedintwolongitudinal,almost straightrows,whereasin D.adenodactylosum theyform twocurvedrows(seeKenk,1978,p.47). Dendrocoelummariae sharesthepresenceofaverylarge adenodactyl,largerthanthepenis,with D.adenodactylosum D.maculatum (Stankovic andKoma rek,1927), D. lacustre (Stankovic ,1938), D.lychnidicum (Stankovic 1969), D.ochridense (Stankovic andKoma rek,1927), D. sanctinaumi (Stankovic andKoma rek,1927), D.minimum Kenk,1978, D.nausicaae (Stankovic andKoma rek,1927), and D.dani Bromley,1982.Allofthesespeciesareendemic totheLakeOhridarea,withtheexceptionof D.nausicaae and D.dani Dendrocoelumnausicaae isoneofthemost widelydistributedspeciesofthegenus,togetherwith D. lacteum Mu ¨ller,1774and D.album (Steinmann,1910), while D.dani isonlyknownfromnorthernIsrael (Gourbault,1972;Kenk,1978;Bromley,1982).Furthermore, D.mariae issimilarto D.adenodactylosum and D. maculatum withrespecttotheanatomyoftheadenodactyl. Inthelatterspeciestheadenodactylischaracterizedbythe presenceofazoneoffinecircularmusclefibersthatruns throughthemesenchymeofthepapilla.Thisparticular adenodactylanatomy,calledtheBalkantype,isalso reportedfor D.lacteum, threerecentlydescribednew speciesfromTunisia(Harrathetal.,2012),andtheother twonewspeciesfromSardiniareportedinthepresent paper.However, D.mariae differsfrom D.adenodactylosum D.maculatum D.lacustre D.lychnidicum D. ochridense D.sanctinaumi D.nausicaae, and D.minimum intheabsenceofthepenisglandsandtheglandularfieldof tallepidermalcellssurroundingthegonoporethatis characteristicforthesespecies.Theglandularfieldaround thegonoporeandpeculiaritiesinthehistologyofthe oviducts(presenceofnumerouspear-shapecells)were consideredbyDeBeauchamp(1931,1932)tobedistinguishingcharacteristicsofthesubgenus Neodendrocoelum Koma rek,1926.Thissubgenuscomprisesahomogeneous groupofspecieswitheyes,restrictedtotheDinaricarea andrecordedchieflyfromLakeOhridanditstributary streamsandsprings.AccordingtoKoma rek(1926)allof theseBalkanspeciesarecharacterizedbyalargepenisand alargeadenodactyl,thelatterbeingbiggerthanthepenis, andbythepresenceoftwoatrialcompartments. Dendrocoelummariae differsfrom D.dani intheabsenceofa copulatorybursainthelatterthatisreplacedbybursalintestinalcommunications(Bromley,1982). Inmostoftheknownspeciesofthegenusthe copulatoryapparatusispositionedjustbehindthepharyngealpocket,whilethepenisandtheadenodactylinthe majorityofspeciesarelocalizedontheoppositesidesofthe body.Incontrast,in D.mariae thecopulatoryapparatusis situatedfarbehindthepharyngealpocket,whilethepenis andtheadenodactylarealwayslocateddorsallyand ventrally,respectively. Thenewspeciesbearsatruesphincterintheterminal tractofthebursalcanal.Thisprominentstructureis mainlyformedbyseverallayersofcircularmuscle S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 98 N JournalofCaveandKarstStudies, August2013

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surroundingadistinctnarrowingofthebursalcanallumen (seeseparatediscussionbelow). D ENDROCOELUMNURAGHUM S TOCCHINOAND S LUYSSP NOV (Table2;Figs.1,3) Materialexamined. Holotype:ZMAV.Pl.7102.1,Monte MajoreCave(40 u 30 9 51 0 N,8 u 36 9 37 0 E),13May2009,coll. B.CadedduandP.Marcia,sagittalsectionson4slides. Othermaterial:CGASPla2.1,MonteMajoreCave (40 u 30 9 51 0 N,8 u 36 9 37 0 E),April2005,coll.P.MarciaandG. Tomasin,sagittalsectionson24slides. Etymology.Thespecificepithetreferstonuraghe,the typicalBronzeAgemegalithicedificesymbolofSardinia. Geographicaldistribution.EndemictotheMonte MajoreCaveandonlyknownfromthetypelocality. Habitat.Specimensof D.nuraghum werefoundinthe MonteMajoreCaveatca.530masl,underpebblesina smallstreamletatca.130metersfromthecaveentrancein completelydarkconditions(Fig.1).Planarianswere associatedwith Proasellus sp.(Asellidae).Althoughseveral samplingcampaigns(20intotal)wereheldinaperiodof fiveyears(2005to2010)only4planarianswerefoundon threeoccasionsduringthespring(April,May).Onlythree ofthefourindividualscollectedweresexual;oneofthese sexualspecimensdidnotsurviveunderlaboratoryconditions.Theimmature,asexualspecimenwasprocessedfor karyologicalanalyses(notreportedinthispaper). Diagnosis. Dendrocoelumnuraghum ischaracterizedby thepresenceofasinglecommonvasdeferens,alargepenis papillawithintrovertedapicalpart,averylargeadenodactyllocatedtotheleftofthepenispapilla,thepresence ofasphincterintheterminaltractofthebursalcanal,and bytheposteriorextensionofthetestestoonlythelevelof therootofthepharynx. Description.Liveadultanimalshadabodysizeofca. 8mminlengthandawidthrangingfromca.2mminthe centralpartofthebodytoca.0.6mmatthelevelofthe head.Theyareunpigmentedandfairlytransparent,sothat theintestinalbranches,pharynx,andcopulatoryapparatus areclearlyvisible.Theanteriorendistruncated,witha convexmid-frontalmargin,andisprovidedwithapairof roundedlaterallobes(Fig.3A). Thetwoeyesaresituatedrathercloselytogether.There isaslightneck-likeconstrictionjustbehindtheeyes,after whichthebodygraduallywidens,attainingitsmaximum widthatthelevelofthepharynxandthecopulatory apparatus.Thedistancebetweentheeyesis1/3to1/4ofthe widthoftheneck.Inter-oculardistanceissmallerthanthe distancefromeacheyecuptothelateralmargin,andthe distancefromthefrontalmargintotheeyesisgreaterthan thedistancetothelateralmargin. Thesubterminalanterioradhesiveorganhasalength slightlygreaterthantheinter-oculardistanceandconsists ofashallowcup.Thesurfaceoftheorganconsistsofan areaofinfranucleatedepithelialcellspiercedbynumerous glandducts.Thecellbodiesoftheveryabundantglands arescatteredthroughoutthemesenchymeoftheanterior partofthebody.Theypassabovethebrainandthen aboveandundertheintestinaldiverticula,reachingup totheanteriorsurfaceoftheovaries.Themusculature oftheadhesiveorganconsistsofamorestronglydevelopedsectionoftheusualventrallongitudinalbody musculature. Theintestineiswellvisible,theanteriorramusreaches toaleveljustbehindtheneckandbears7or8brancheson eachside.Thetwoposteriorramibear16or17branches. Thepharynxislocatedintheposteriorhalfofthebodyand measuresabout1/7ofthebodylength(Fig.3A).Its internalmusclezoneconsistsofathicklayerofintermingledcircularandlongitudinalfibers.Asubepitheliallayer oflongitudinalmusclesfollowedbyalayerofcircular fibersformsthethinouterzoneofmuscles. Thetwoovariesoccurontheventralsideoftheanterior regionbehindthebrain.Theovariesarelocatedatlessthan 1/3ofthedistancebetweenthebrainandtherootofthe pharynx.Theoviductsoriginatefromthelatero-dorsalpart oftheovariesandareprovidedwithaslightexpansionat theiranteriorend,thetuba.Theoviductsrunposteriorly, convergingbehindthecopulatoryapparatustoforma singlecommonoviduct.Thelatterrunsanteriorlytothe rightsideoftheopeningofthebursalcanalandopensinto theendpartofthemaleatrium.Thedistalpartofthe oviductsandthefirsttractofthecommonoviductreceive theopeningsoftheshellglands(Fig.3B). Thenumerouswell-developedtestesareessentially ventralinposition,butsomearesituatedinadorsal positionorinthemiddleofthebody.Somefolliclesare ovalinshapeandoccupyalmosttheentiredorso-ventral spaceofthebody.Thetestesextendfromashortdistance behindtheovariestotheleveloftherootofthepharynx. Thespermductsformverylargespermiducalvesicles, packedwithsperm,betweenthemouthandtheanterior levelofthepenisbulb(Fig.3B). Intheholotype(ZMAV.Pl.7102.1)thecopulatory apparatusislocalizedjustbehindthepharyngealpocket, whileinCGASPla2.1specimenthecopulatoryapparatus occupiestheposteriorhalfofthepostpharyngealregion. Thepenisislocatedtotheright,thebursalcanalandthe adenodactyltotheleftofthemidline.Thecopulatory bursahastheshapeofalargesackthatoccupiesalmostthe entiredorso-ventraldiameterofthebody.Thebursais linedwithatallglandularepitheliumandissurroundedby alayeroflongitudinalmuscle.Thebursalcanalruns posteriorlytotheleftofthepenisandgraduallywidens, thenturnsventrallytoopenintothecommonatrium.The wallofthebursalcanalconsistsofanucleatedepithelium withciliatedcells.Thecanalissurroundedbyalayerof circularmuscle,followedbyalayeroflongitudinalfibers. Atashortdistancefromtheopeninginthecommon atriumthecanallumenreduces,whilethecircularmuscle layerbecomesverythick;thistractofthebursalcanalis envelopedbyasphincter(Fig.3C). G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 99

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Figure3. Dendrocoelumnuraghum .(A)habitusofalivingspecimen,(B-C),holotypeZMAV.Pl.7102.1,sagittal reconstructionsofthecopulatoryapparatus(anteriortotheright). S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 100 N JournalofCaveandKarstStudies, August2013

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Inthemalecopulatoryapparatusthemuscularpenis bulbisofmoderatesize.Thevasadeferentiapenetratethe penisbulbseparately,butveryclosetoeachother,atits anteriorpart,andthentheyconvergetowardthemidlineof thepenisbulbtoformacommonvasdeferensthatopens intotheanteriorpartofthepenispapillalumen(Fig.3B). Thelongpenispapillameasures3/4ofthetotalpenis length.Thepenislengthisca.1/2oftheadenodactyl length.Thepenispapillaisbarrel-shapedintheholotype andfinger-shapedinspecimenCGASPla2.1;thelatter conditionmaybeduetoapreservationartifact. Thepenispapillaiscoveredbyaverythinepitheliumat thebasalpartthatbecomesthickerattheapicalpart.The epitheliumisunderlinedbyathicklayerofcircularmuscle thatisthickerintheventralpartandatthebaseofthe papilla.Inbothspecimensexaminedtheapicalpartofthe penispapillalacksmusclelayersandappearsintroverted intotheverylargepapillalumenlikeapseudoflagellum (Fig.3B;seediscussiononflagellumbelow). Inthepenispapillalumenoftheholotypeapackof spermispresent.Thepenisisrichinglands,whichinthe holotypearelocatedinthepenisbulbandinthedorsal partofthepapilla.Theseglandsopenintothelargepenis lumen,whichisfullofsecretion. Theadenodactylisverylargeandconsistsofafree papillaandawell-developed,verymuscularbulbarpart (Fig.3C).Thefreepapillaisabout1/2ofthetotal adenodactyllength.Thebulbstartsinthedorsalpartof thebodyandhasadorsal-ventralorientation.Itconsistsof rowsofintermingledlongitudinalandcircularmuscles fiberscoveredbyathinlayeroflongitudinalfibers.Ectally tothisthinlongitudinalmusclelayerrunsathicklayerof finecircularmusclefibersthatcontinuesitscoursethrough themesenchymeoftheadenodactylpapilla(Fig.3C).In specimenCGASPla2.1,stainedinDaneHermanÂ’s tetrachrome,thisthicklayerisgreyandcontrastswith theothermuscularfibers,whichstainbrightyellow.Inthe holotype,stainedinMallory-Cason,thislayerispaleblue, thelongitudinalfibersstainred,andtheintermingled musclesstainbrightblue.Thefreepapilla,whichprotrudes posteriorlyanddorsallyintotheatrium,ischaracterizedby athinliningepitheliumunderlainbythreelayersof muscles:asubepitheliallayeroflongitudinalmuscle,a thicklayeroffinecircularmusclefibers,andathininner layeroflongitudinalmuscle. Themaleatriumconsistsofananteriorpartthathouses thepenispapillaandacanalleadingobliquely,ina postero-ventraldirection,intothecommongenitalatrium. Thiscanalislinedbyathickepitheliumandissurrounded byastronglayerofcircularandlongitudinalmusclefibers. Thecommonatriumreceivestheopeningofthebursal canal. Discussion. Dendrocoelumnuraghum ischaracterizedby thefusionofthetwovasadeferentiaintoanintrapenial commonduct.Withinthegenus Dendrocoelum this characterisreportedforonlyfourotherspecies: D. jablanicense (Stankovic andKoma rek,1927), D.puteale Kenk,1930, D.kenki DeBeauchamp,1937,and D. constrictum HarrathandSluys,2012. Dendrocoelum nuraghum differsfrom D.puteale and D.kenki because thesearebothanophtalmousspecies. Dendrocoelumputeale ischaracterizedbyanextrabulbarseminalvesiclethat receivesinitsrightpartacommonvasdeferens,originating fromthefusionofthetwovasadeferentiaataleveljust behindthecopulatorybursa(DeBeauchamp,1932).In D. kenki thevasadeferentiaunitejustbeforeenteringthe penisbulb.Moreover,inthisspeciesthepenisisinthe middle,whiletheadenodactylisontherightsideof thebody(DeBeauchamp,1937),incontrastto D. nuraghum inwhichthepenisislocalizedontherightand theadenodactylontheleftofthemidlineofthebody. Dendrocoelumnuraghum issimilarto D.jablanicense becauseitsvasadeferentiaalsoseparatelypenetratethe penisbulbandfusetoacommonductbeforeopeninginto thepenispapillalumen.However, D.nuraghum differs from D.jablanicense intheabsenceoftheglandularfieldof tallepidermalcellsthatinthelattersurroundsthe gonopore.Moreover,theadenodactylin D.nuraghum is ontheleftofthemidlineofthebody,whereasin D. jablanicense itisontheright.Thelast-mentionedspeciesis alsocharacterizedbyadoublecommunicationbetweenthe bursalcanalandacommonatrium(seeKenk,1978),which isabsentin D.nuraghum Dendrocoelumnuraghum ischaracterizedbyanirregular distributionofthetestes(ventral,dorsal,andbetweenthe intestinaldiverticula),whichextendposteriorlytothelevel oftherootofthepharynx.Incontrast,in D.puteale andin D.jablanicense thetestesextendtothetailendandare ratherdorsalandventral,respectively.In D.kenki the testesaremainlyventralandextendposteriorlytothelevel ofthecopulatorybursa. Dendrocoelumnuraghum hasadistinctnarrowinginits bursalcanal,surroundedbyasphincter.Thischaracteris sharedwithtwootherSardiniandendrocoelids(seebelow). Thelargepenisandlargeadenodactylof D.nuraghum are sharedwith D.adenodactylosum D.maculatum D.lacustre D.lychnidicum D.ochridense D.sanctinaumi D.minimum D.nausicaae and D.dani.Dendrocoelumnuraghum differs from D.adenodactylosum D.maculatum D.lacustre D. lychnidicum D.ochridense D.sanctinaumi D.minimum ,and D.nausicaae intheabsenceoftheglandularfieldaroundthe genitalporethatispresentinallofthesespecies. D ENDROCOELUMVESICULOSUM S TOCCHINOAND S LUYSSP NOV (Table2;Figs.1,4,5,6) Materialexamined. Thepopulationischaracterizedbythe coexistenceoftwomorphotypes:anophtalmous(morphotype1)andtwo-eyed(morphotype2)individuals. Holotype:ZMAV.Pl.7103.1,sagittalsectionson4 slidesofamorphotype-2animal,SaUccae `suPeltusu Cave(40 u 27 9 00 0 N,8 u 40 9 43 0 E),6May2009,coll.P.Marcia, B.Cadeddu,F.StochandG.Tomasin. G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 101

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Paratypes:CGASPla3.1,ibid.,sagittalsectionson3 slidesofamorphotype-2animal;ZMAV.Pl.7103.2,ibid., sagittalsectionson3slidesofamorphotype-2animal, ZMAV.Pl.7103.3,ibid.,sagittalsectionson4slidesofa morphotype-1animal. Othermaterial:CGASPla3.2,sagittalsectionson6 slidesofamorphotype-2animal,notcompletelymature, SaUcca‘esuPeltusuCave(40 u 27 9 00 0 N,8 u 40 9 43 0 E)3 September2007,coll.P.MarciaandM.Fois;CGASPla 3.3,ibid.,sagittalsectionson13slides,ofamorphotype-1 animal,notcompletelymature;ZMAV.Pl.7104.1,sagittal sectionson5slidesofamorphotype-1animal,F1 offspring;ZMAV.Pl.7105.1,onenotcompletelymature morphotype-1animal,resultingfromtheF2offspring, sagittalsectionson3slides. Etymology.Thespecificepithetisderivedfromthe Latinadjective vesiculosus ,fullofvesicles,andalludesto thepresenceofanexpansionorvesicleineachvasa deferens. Lifecycle.Specimensof Dendrocoelumvesiculosum were collectedinSeptember2007andMay2009.Onthefirst occasiontwojuvenileswerecollected,onespecimenbeing anophtalmousandtheotherwithtwoverysmalleyes.The lifecycleoftheseindividualswasobservedforaboutthree yearsunderlaboratoryconditions.Inthisspeciesthe breedingperiodfollowsaseasonalcycle.FromMaytoJuly thespecimensdevelopedacopulatoryapparatus,whilein summer-autumntheylaidcocoons.InMay2008,after 7monthsofrearing,asexualizationprocesswasdisplayedin bothspecimens.InthefollowingJunethetwoanimals produced2cocoons.Fromtheonlyfertilecocoonfiveyoung anophtalmousplanarians(F1)hatchedafterfiveweeksof development.Onlythreeofthesewormssurvivedafter 7monthsofrearing(January2009),andtheybecamesexual inJuly2009andproducedthreecocoonsduringOctober 2009.Fromtheonlyfertilecocoonfouryounganophtalmousplanarians(F2)hatchedafter4weeks.InJune2010the onlysurvivingplanarianbecamesexuallymature. Figure4. Dendrocoelumvesiculosum. Habitusoflivingspecimens,(A)morphotype1characterizedbytheabsenceofeyes,(B) morphotype2witheyes. S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 102 N JournalofCaveandKarstStudies, August2013

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Thefirsttwoworms,i.e.,theparents,werekilledand preservedinJuly2008,afterthefirstoffspring(F1)had beenproduced,onemonthaftercocoondeposition.Atthat time,bothgonadsandthecopulatoryapparatusappeared tobeinastateofregression,althoughabundantsperm werestillpresentinthevasadeferentia.Thisconditionof regressionwasparticularlyevidentinthetwo-eyed specimen. Duringthesecondcollection,fivespecimenswere captured,onesexualanophtalmousanimalandfour witheyes,ofwhichonlytwoweresexuallymature andeventuallyonlyonesurvived.Theothertwoanimals becamesexuallymatureinJanuary2010,aftereight monthsofrearing.Allspecimenswerepreservedatthat time. Habitat.Specimensof Dendrocoelumvesiculosum were foundintotallydarkconditionsunderpebblesinasmall streamintheSaUccaÂ’esuPeltusuCaveatanaltitudeof ca.560masl(Fig.1).Thedendrocoelidswereassociated with Dugesia sp. Figure5. Dendrocoelumvesiculosum .(A-B)holotypeZMAV.Pl.7103.1,sagittalreconstructionsofthecopulatoryapparatus (anteriortotheleft),(C)CGASPla3.1sagittalreconstructionsofthepenispapilla(anteriortotheright),(D)ZMAV.Pl. 7103.2,sagittalreconstructionsofthepenispapilla(anteriortotheright). G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 103

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Geographicaldistribution.EndemictotheSaUcca‘e suPeltusuCaveandonlyknownfromthetypelocality. Diagnosis. Dendrocoelumvesiculosum ischaracterized by:thepresenceinthepopulationofbothanophtalmous andtwo-eyedindividuals;apeculiarpenishistology,with vacuolatedcellsliningthetipofthepapilla;aconvoluted adenodactyl;asphincterintheterminaltractofthebursal canal;averynarrowhorizontaltractofbursalcanal;and thepresenceofapeculiarexpansioninthevasadeferentia, differentfromthespermiducalvesicles. Description.Liveanimalsareunpigmented,typically whitish,withabodysizeof8to10mminlengthanda widthrangingfromca.2mminthecentralpartofthebody to1mmatthelevelofthehead.Theanteriorendis truncatedwithapairofroundedlaterallobes(Fig.4).In theocularanimalstheeyesaresmallandplacednotfar fromthefrontalmargin.Theirdistancefromthefrontal marginisslightlygreaterthanthedistancetothelateral margin.Inter-oculardistanceisequaltothedistancefrom eacheyecuptothelateralmargin.Justatlevelofeyesthere isaslightconstrictionorneckthatsetsofftheheadfrom therestofthebody(Fig.4B). Theanterior,subterminaladhesiveorgan,withawidth slightlymorethantheinter-oculardistance,isfeebly developedandconsistsofasmallpatchofepidermiswith infranucleatedcellsthatispiercedbynumerousopenings ofglandsducts. Thepharynxislocatedintheposteriorhalfofthebody andmeasuressomewhatlessthan1/6ofthebodylength. Thethinouterzoneofmusclesconsistsofasubepithelial layeroflongitudinalmuscle,followedbyalayerofcircular muscles.Theinnerepitheliumisunderlainbyathicklayer ofintermingledcircularandlongitudinalfibers. Thetwoventral,pairedovariesarelocatedatabout1/3 ofthedistancebetweenthebrainandtherootofthe pharynx.Theanterior-mostsectionsoftheoviductsare expandedtoformatuba.Theoviductsrunposteriorly, convergebehindthecopulatoryapparatus,anduniteto formacommonoviductthatopensintotheterminaltract ofthemaleatrium.Thecommonoviductreceivesthe openingsofeosinophilicshellglands. Well-developedroundedresorptivevesiclesarepresent alongtheoviductsofallspecimensexamined.Eachvesicle, characterizedbythepresenceofvacuoles,communicates withtheoviductthroughashort,narrowduct(cf.Sluys, 1989).Spermispresent,bothintheresorptivevesiclesand intheshortinterconnectingductules. Thewell-developed,numeroustestesarelocalizedin bothventralandindorsalposition,whilesomefolliclesare situatedinthemiddleofthebody.Thetestesextend throughoutthebodyfromdirectlybehindtheovariesto thetailend. Thesac-shapedcopulatorybursaissituatedjustbehind thepharynx.Thebursaislinedwithatallglandular epitheliumandissurroundedbyalayeroflongitudinal muscle.Fromthecopulatorybursathebursalcanalruns posteriorlyandhorizontallytotheleftofthepenis.This horizontaltractofthebursalcanalisverynarrowandis linedwithanucleated,ciliatedepithelium,surroundedby onlyathinlayeroflongitudinalmusclefibers.Atthelevelof theinsertionoftheadenodactylpapillathebursalcanal widens,andfromthispointitissurroundedbyalayerof circularmuscle,followedbyalayeroflongitudinalfibers. Beforeopeningintotheatriumthebursalcanalnarrows again,whilethecircularmusclesbecomethicker,thus formingasphincter(Fig.5B).Thisstructureismostevident inthosespecimensinwhichthetipoftheadenodactyl papillaisnotdeeplyinsertedintothebursalcanal. Thewell-developedpenisbulb,formedbyintermingled circularandlongitudinalmusclefibers,housesalarge Figure6. Dendrocoelumvesiculosum .(A)photomicrographofthepenispapillatipshowingthevacuolatesecretory epithelium,(B)photomicrographoftheexpansion/vesicleinthevasdeferens. S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 104 N JournalofCaveandKarstStudies, August2013

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seminalvesiclethatislinedbyavacuolatedepithelium thecellsofwhichprotrudeirregularlyintothelumen (Fig.5A). Inallspecimensexaminedthetwovasadeferentiarun ventrally,andwhentheductsarriveattheleveloforjust beforethecopulatorybursatheyshowapeculiarexpansion oftheirdorsalwall,thusformingalargevesiclethatcanbe eitheremptyorfullofsperm.Thewallofthisvesicleis linedbytall,claviform,vacuolatedcells,verydifferent fromtherestoftheducts,whicharelinedwithathin epithelium(Figs5A,6B). InspecimenCGASPla3.1abundantspermispresentin thelumenofthetwoposteriorintestinalbranches.Inthe samespecimentherightvasdeferensshowsatthelevelof theexpansionaconnectionwiththelumenofaposterior intestinalbranch,mostlikelyduetoapreservationartifact. Afterhavinggivenrisetoanexpansion,thevasa deferentiaformwell-developedspermiducalvesicles,packed withsperm.Thereaftertheductscurvetothedorsalside,in somecases(ZMAV.Pl.7103.1,ZMAV.Pl.7103.2,CGAS Pla3.1)almostreachingthedorsalpartofthebody,and penetratethepenisbulbtoopen,closelytogether,intothe antero-lateralsectionoftheseminalvesicle.Inspecimens ZMAV.Pl.7103.2andZMAV.Pl.7103.3therightvas deferensentersthepenisbulbdorsallytotheleftone.The penisisrichinglandsthatopenintotheseminalvesicleand intothepenispapillalumen(Fig.5A). Thepenispapillaischaracterizedbyathickbasalpart andathinnerdistalpart.Thebasalpartiscoveredwitha flatepitheliumthatisunderlainbyathicklayerofcircular muscle.Theepitheliumontheapicalpartconsistsoflarge, rounded,andvacuolatedcells,withbasalnuclei,veryrich insecretiongranules.Thisepitheliumcoversboththe externalandtheinternaldistalpartofthepenispapilla (Figs5A,6A).Intheholotype(ZMAV.Pl.7103.1)sperm ispresentinthemaleatrium. Thepenispapillamayassumedifferentshapes.Inthe holotypeandinZMAV.Pl.7103.3itisconsiderably elongatedandprovidedwithatunnel-shapedlumen (Fig.5A).InCGASPla3.1thepapillaismorecontracted andthereforeprovidedwithawiderlumen(Fig.5C).In ZMAV.Pl.7103.2thecontractionandinversionofthetip ismoreadvancedandthusformsapseudo-flagellum (Fig.5D). Theadenodactylconsistsofawell-developedbulbar partandafreepapilla.Thebulbisverylargeandoccupies almosttheentiredorso-ventralspaceofthebody(Fig.5B). Inallexaminedspecimenstheshapeoftheentire adenodactylissinuous,inthatthebulboriginatesatthe ventralregionofthebodyandthengraduallyextendsto theleftwithadorso-lateralorientation.Theadenodactyl papillafirsthasadorso-ventralorientationandthenmakes aposteriorlydirectedbend,withthetippointingintothe terminaltractofthebursalcanal.Duetothelargesize,the particularorientation,andthedifferentstatesofcontractionoftheadenodactyl,thepenismaybelocalizedat differentpositions.InspecimensZMAV.Pl.7103.2,ZMA V.Pl.7103.3,andZMAV.Pl.7104.1thepenisislocatedon therightsideoftheadenodactyl,whereasinCGASPla3.1 andZMAV.Pl.7103.1thepenisislocatedontheleftside ofanddorsallytotheadenodactyl,respectively. Thebulboftheadenodactylconsistsofintermingled rowsoflongitudinalandci rcularmuscle,boundedbya thinlayeroflongitudinalfibers.Ectallytothisthin longitudinalmusclelayerrunsathicklayeroffine circularmusclefibersthatcontinuesitscoursethrough themesenchymeoftheadenodactylpapilla.Ectallyto thiszoneofcircularmusclesrunsathinlayerof longitudinalmusclefibers(Fig.5B).Inallexamined specimens,stainedinMallory-Cason,thislayeroffine circularfibersispaleblue,whilethelongitudinalfibers stainredandtheintermingledmusclesstainbrightblue. Noglandswereobservedtodischargeintothelumenof theadenodactyl. Themaleatriumislinedbyanucleatedepithelium surroundedbyasubepitheliallayerofcircularmuscles, followedbyalayeroflongitudinalfibers.Justbefore openinginthecommonatriumthemaleatriumreceivesthe openingofthecommonoviduct. Discussion.Inthecourseofourlong-termstudyofthe lifecycle,duringtwogenerations(3years),onlyanophtalmousoffspringwasobserved.Partialortotallossofeyes mayresultfromadaptationtothesubterraneanenvironment.Aseasonallifecycle,asobservedin D.vesiculosum hasbeenreportedforonlyafewhypogeanspecies,suchas Atrioplanariadelamarei (Gourbault,1972).Themajorityof subterraneantricladsproducecocoonsduringtheentire year(Gourbault,1972). Thenumberofcocoonsin D.vesiculosum isverylow(2 or3).Generally,thenumberofcocoonsproducedby hypogeanspeciesvariesbetweenspecies,butitisalways considerablylowerthaninepigeanspecies(Gourbault, 1972).Thefertilityofthecocoonsisinhypogeantriclads alsoveryvariablebetweenspecies(Gourbault,1972).In D. vesiculosum thefertilitywas30to50%andonly4or5 younghatchedfromeachcocoon,inagreementwiththe generalconditionofhypogeantriclads,inwhichgenerally fewerthaneightyounghatchfromacocoon. Thejuvenilesof D.vesiculosum becamesexuallymature 8to12monthsafterhatching,incontrastto D.collini (De Beauchamp,1919),whichneeds4to5months,and Phagocatavitta juvenilesthatbecomesexualafter9or 10months(Gourbault,1972).However,thelownumberof younghatchingfromthecocoonsiscounterbalancedbyan increaseoflongevity.Aspecimenof D.mariae wasstill aliveafter6years,althoughlaboratoryconditionsareless harshthanthoseinsitu,forexample,withrespecttofood availability.Ourobservationsonthelifehistoryillustrate considerableslowingdownofthecocoondeposition rhythm,aswellasembryonicandpost-embryonicdevelopment(Kstrategy)andareinagreementwiththe adaptivestrategiestypicalofsubterraneanfauna(Juberthie G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 105

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andDecu,1994)andinparticularofstygobionttriclads (Gourbault,1994).Thisstrategyensuresamoreefficient managementofalowenergybudget. Dendrocoelumvesiculosum showsapeculiarpenis papillahistology,characterizedbythepresenceoflarge, vacuolatedcellsthatsurroundboththeinternalandthe externalwallofthetipofthepenispapilla.Thischaracter isuniqueforthisspecies(seediscussiononflagellum below). Anotheruniquecharacterforthisnewspeciesisthe presenceofapeculiardorsalexpansioninthevasa deferentia.Thisexpansiondiffersfromtheusualspermiducalvesiclesandhasnotbeendescribedearlierforany otherspeciesofthegenus Dendrocoelum Dendrocoelumvesiculosum ischaracterizedbyalarge adenodactylandbyasphincterintheterminaltractofthe bursalcanal.Thesecharactersaresharedwithtwoother Sardinianspecies(seediscussiononsphincterbelow). FamilyPLANARIIDAEStimpson,1857 Genus Phagocata Leidy,1847 Phagocataobscura StocchinoandSluyssp.nov. (Table2,Figs.1,7) Materialexamined. Holotype:ZMAV.Pl.7106.1,sagittal sectionson34slides,CuccuruTiriaCave(39 u 19 9 25 0 N, 8 u 34 9 29 0 E),28March2006,coll.P.Marcia. Habitat.Specimensof Phagocataobscura werefound underpebblesinastreamrunningintotallydarkconditionintheCuccuruTiriaCave(Fig.1).Planarians wereassociatedwith Stenasellus sp.(Stenasellidae)and Oligochaeta. Etymology.Thespecificepithet obscura, fromtheLatin adjective obscurus (dark,somber,hidden,invisible),alludes toboththecavehabitatofthespeciesandthefactthat somedetailsofitsanatomycouldnotbediscernedinthe materialavailable. Geographicaldistribution.EndemictotheCuccuru TiriaCaveandonlyknownfromthetypelocality. Diagnosis. Phagocataobscura ischaracterizedbythe absenceofeyes,aconicalhead,thepresenceofawelldevelopedpenisbulbcontainingaspaciousseminalvesicle, aplugofcellsintheejaculatoryduct,acommonvas deferens,andtesteslocatedmainlyinprepharyngeal position. Description.Liveanimalsareunpigmented,typically whitish,withabodysizeinanelongatedstateofca. 6mminlengthandawidthrangingfrom1mminthe centralpartofthebodyto0.5mmatthelevelofthe head.Theanteriorendisobtuselypointedanddevoidof eyes(Fig.7A).Thepharynxislocatedintheposterior halfofthebodyandmeasuresabout1/5ofthebody length. Thetwoovariesarelocatedatashortdistancebehind thebrain;theyaresmallandpoorlydeveloped.The oviductsrundorsallytotheventralcordsandcurve dorsallyatthelevelofthepenisbulb.Theoviductsarestill visibleatthispoint,buttheircontinuationandsubsequent communicationwiththefemalecopulatoryapparatus couldnotbediscerned. Thenumerousbutpoorlydevelopedtestesareclosely packedtogetherinventralposition.Theybeginata considerabledistancebehindtheovariesandextend posteriorlytotheleveloftherootofthepharynx.Some folliclesmayextendtohalfwayalongthepharyngeal pocket. Thecopulatoryapparatusislocatedatsomedistance behindthepharyngealpocket.Thesmallsac-shaped copulatorybursaislinedwithatallepitheliumandis surroundedbyalayeroflongitudinalmuscle.Fromthe bursa,anarrowbursalcanal,linedwithasquamousto cuboidal,nucleatedepithelium,runsdorsallytothemale atrium.Neartheposteriorendofthemaleatriumthe canalcurvesventrallytoopenintothecommonatrium (Fig.7B). Thepenisbulbiswell-developed,withalength approximatelyequaltothatofthepenispapilla;its musculatureisformedbyintermingledlongitudinaland circularmusclesfibers. Asinglevasdeferensopensintotheproximal,anterior sectionoftheintrabulbarseminalvesicle.Thisductmay representacommonvasdeferens,originatingfromthe unionofthetwovasadeferentiaintheanteriorpartof thepenisbulb,butsuchcouldnotbediscernedinthe sections. Theepitheliumoftheelongated,finger-shapedpenis papillaisunderlainbyathinlayerofcircularmusclefibers. Theejaculatoryductrunscentrallythroughthepenisand opensatthetipofthepapilla.Theejaculatoryductis histologicallydifferentiatedintotwosections.Ashort anteriorportionisenlargedtoformarelativelyspacious seminalvesicleorintrabulbarcavitythatislinedwithtall secretorycellswithbasalnucleiandsurroundedbyalayer oflongitudinalmuscle.Theposteriorpartofthiscavityis linedwithmuchtallercells,formingaplugthatfillsthe entirelumenofthecanal(Fig.7B).Thesecond,distal sectionoftheejaculatoryductislinedbyanepithelium withcuboidalcells. Themaleatriumislinedbytallnucleatedcells, surroundedbyathicklayerofcircularmuscle. Discussion. Phagocataobscura isthefirstanophtalmous speciesofthegenusreportedinEurope.Accordingto Kenk(1978)allEuropeanspeciesof Phagocata are‘‘two eyedplanariidswithatruncatedhead.’’Thistraitappears tobeconsistentlypresentinallspeciesthatwere subsequentlydescribed(cf.Sluysetal.,1995;Vila-Farre ` etal.,2011). Theplugofcellsintheejaculatoryductissharedwitha groupofelevenspecies(cf.Sluysetal.,1995;Vila-Farre ` etal.,2011): P.albissima (Vejdovsky ,1883), P.armeniaca P.undulata P.bosniaca P.macedonica (Stankovic ,1926), P. illyrica (Koma rek,1919), P.maculata P.dalmatica (Stankovic andKoma rek,1927), P.ochridana P.stankovici S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 106 N JournalofCaveandKarstStudies, August2013

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(Reisinger,1960),and P.hellenica (Vila-Farre `andSluys, 2011).However,allofthesespecieshavetwoeyes. Phagocataobscura sharesitsprincipallyprepharyngeal testeswiththeNorthAmericanspeciesof Phagocata .In contrastto P.obscura ,Palaearcticspeciesof Phagocata usuallyhavetheirtestesdistributedthroughtheentirebody. AlthoughStocchinoetal.(2008)tentativelyascribed thismaterialto Atrioplanaria ,thepresentdetailedmorphologicalstudyhighlightedthatthisisthefirstandonly speciesof Phagocata recordedforItaly.Only Phagocata sp. wasreportedbeforenowfromTuscanyandSardinia (Benazzi,1982;Stocchino,2003).Thefewrecordsof P. vitta reportedfortheItalianfauna(Belloetal.,1995; Ferreri,1995)arenotsupportedbyamorphological accountandshouldthereforebeconsidereddoubtful. P HAGOCATA SP (Table2;Figs.1,8A,B,D). Asexualindividualsfromthreepopulationswere collectedfromthefollowingthreesites;absenceofafully developedcopulatoryapparatuspreventedamoredetailed assessmentoftheirtaxonomicstatus: Figure7. Phagocataobscura .(A)habitusofalivingspecimen,(B)holotypeZMAV.Pl.7106.1,sagittalreconstructionofthe copulatoryapparatus(anteriortotheleft). G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 107

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1)MonteMajoreCave(40 u 30 9 51 0 N,8 u 36 9 37 0 E),coll. G.A.StocchinoandP.Marcia(Figs1,8A).Twentyfivespecimenswerecollectedduringtwooccasionsin JuneandJuly2010.Thespecimenswerefoundina smallpoolwithmuddybottom,originatingfromdrip water.Theanimalsarewhitish,verysmallandslender. Theheadistruncated,withtwosmalleyes,situated closetogetherandlocatedfarfromthefrontalmargin. Aftermorethanoneyearofrearingtheystilldidnot showanysignofasexualizationprocess. Figure8.(A) Phagocata sp.,habitusofalivingspecimenfromtheMonteMajoreCave,(B) Phagocata sp.,habitusofaliving specimenfromtheEligheMannuSpring,AsinaraIsland,(C) Dugesiabenazzii ,habitusofalivingspecimenfromtheSu CantaruSpring(MonteAlbo),(D) Phagocata sp.,habitusofalivingspecimenfromaspringatAla `deiSardi,(E) Dugesia sp., habitusoflivespecimensfromtheSaUcca‘esuPeltusuCave. S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 108 N JournalofCaveandKarstStudies, August2013

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2)EligheMannuSpring,AsinaraIsland(41 u 05 9 36 0 N, 8 u 18 9 21 0 E),December2000–June2002,coll.M.Piras andG.A.Stocchino(Figs1,8B).Eighteenasexual specimenswerecollectedduringeightcollectionsmade fromDecember2000toJune2002.Noneofthe specimenscollectedsurvivedunderlaboratoryconditions.Theanimalsareunpigmented,withtruncated headandwithtwosmalleyeslocatedfarfromthefrontal marginandsituatedclosetogether.Planarianswere associatedwithadiverseinvertebratefauna,including Microdalyellia sp.(Platyhelminthes,Dalyelliidae), Ancylus sp.(Gastropoda,Ancylidae),Planorbidae(Gastropoda,Pulmonata),Nematoda,Oligochaeta,Ostracoda, Cladocera,Cyclopoida,Hydracnidia(Arachnida), Ephemeroptera,Dytiscidae,Hydrophilidae,Chironomidae,Culicidae,Ceratopogonidae,Polycentropodidae, andRhyacophilidae. 3)Ala `deiSardi(40 u 38 9 60 0 N,9 u 19 9 45 0 E),February2010, coll.M.Fois(Figs1,8D).Justoneindividualwas collectedfromaspring.Theanimalisunpigmented, whitish,withabodysizeinelongatestateofca.7mm inlengthand1mminwidth.Theheadistruncated, withtwoeyeslocatedfarfromthefrontalmargin.Each eyeisaccompaniedbyonesmallsupernumeraryeye. G ENUS C RENOBIA K ENK ,1930 C RENOBIAALPINA (D ANA ,1766) (Table2,Fig.1) Materialexamined. CGASPla4.1-4,sagittalsectionson11 slides,13slides,13slides,16slides,respectively,immature specimens,SuSesseneSpring,1368masl,westernfaceofthe GennargentuMassif(40 u 01 9 40 0 N,9 u 12 9 1 0 E),February2004, coll.R.FaddaandM.Deiana(watertemperature7 u C). Comparativediscussion.Unfortunately,allspecimens collectedwereimmature,i.e.withoutcopulatoryapparatus.Therefore,theanimalswereidentifiedsolelyonthe basisofexternalappearanceandpresenceofonlyone pharynx.Thelatterconditionrulesoutthepolypharyngeal forms Crenobiaalpinamontenigrina (Mra zek,1904), C. alpinaanophtalma (Mra zek,1907),and C.alpinateratophila (Steinmann,1908).Thisnewrecordindicatesawider geographicdistributionthanpreviouslyknown,although thespeciesisrestrictedexclusivelytotheGennargentu Massifsprings,asreportedearlierbyPalaetal.(1980c). FamilyDUGESIIDAEBall,1974 Genus Dugesia Girard,1850 Dugesiabenazzii Lepori,1951 (Table2;Figs.1,8C) Materialexamined. ZMAV.Pl.7107.1,onesetofsagittal sectionson21slides,ZMAV.Pl.7107.2,onesetof transversesectionson37slides,ZMAV.Pl.7107.3,oneset ofhorizontalsectionson7slides,SuCantaruSpring, MonteAlbo(40 u 34 9 30 0 N,9 u 40 9 31 0 E),5December1998, coll.G.A.StocchinoandD.Salaris. Comparativediscussion.Allspecimensweresexualat collection.Identificationofthespecieswasbasedon histologicalstudyandkaryologicalanalysis. Dugesia benazzii ischaracterizedbyafinger-shapedpenispapilla, apointeddiaphragm,acentralcourseoftheejaculatory ductopeningatthetipofthepenispapilla,and symmetricalopeningsoftheoviductsintothebursalcanal. Specimensof D.benazzii fromtheMonteAlboSpringhave afoldontheleftofthepenispapilla,likethemajorityof Sardinianpopulations(Lepori,1951;Stocchino,unpublisheddata).Thechromosomalnumberis2n 5 16;n 5 8. Thisisthefirstpopulationof D.benazzii recordedfroma Sardinianspring. D UGESIA SP (Table2;Figs.1,8E) Comparativediscussion.Threefissiparousspecimens werefoundunderpebblesinasmallstreamrunninginthe darkSaUcca‘esuPeltusuCave(40 u 27 9 00 0 N,8 u 40 9 43 0 E) Figure9. Dendrocoelumlacteum fromtheEbrodelta,Spain.(A)photomicrographofaninverted,trueflagellum(specimenRS 266fromthecollectionsofNCBNaturalis-sectionZMA),(B)photomicrographofaneverted,trueflagellum(specimenRS219). G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 109

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on3September2007(coll.P.MarciaandM.Fois),atan altitudeofca.560masl(Fig.1).Dugesiidswereassociated with Dendrocoelumvesiculosum (seeabove).Aftertwo monthsofrearing,5individualswerepresent,buttheydid notsurviveunderlaboratoryconditions.Thisisthefirst recordofdugesiidsfromaSardiniancave. G ENERAL D ISCUSSION T HE F LAGELLUMIN D ENDROCOELID F LATWORMS Theterm flagellum isoftenused sensulato forthetipof apenispapillathatcanbeinverted,thusprojectinginto thelumenofthepapilla.Duringthehistoryoftriclad taxonomyseveralattemptshavebeenmadetodefinethe variousstatesofinversionofthepenis,notablyhowto differentiatebetweenaso-calledtrueflagellumanda pseudoflagellum.Koma rek(1926)definedatrueflagellumasafoldoftheepitheliumliningtheinternalsurface ofthepenispapillaandmentionedasexamples Dendrocoelumlacteum and D.infernale (Steinmann,1907). Further,hementioned D.mrazeki (Vejdovsky ,1895)as anexampleofaflagellumthatismerelyformedbythe invaginateddistalpartoft hepenispapilla.Thesame distinctionhadalreadybeenmadebySteinmann(1909). Herecognizedthatinmostspeciestheflagellumis‘‘ nach innengestu ¨lpt ’’(foldedinwards,inflected),whereasin D.lacteum and D.infernale theflagellumis‘‘ umgestu ¨lpt ’’ (turnedinsideout).Healsonotedthatintheflagellumof thesetwospeciesthemusculatureiswell-developed, notablythelongitudinalmuscles ;thecircularmusculature isonlyweak. Despitethestudiesoftheseworkers,Hyman(1931, p.318)consideredpresenceorabsenceofaneversible flagellumtobeanunreliabletaxonomiccharactersince‘‘it hasprovedimpossibletoconstructaprecisemorphological definitionoftheflagellum.’’ Followingupontheworkoftheseearlierworkers, however,wedistinguishfortheflagellum s.l .inthegenus Dendrocoelum sixmainconditionsinregardtothe histologyofthetipofthepenispapillaandtheextent ofitsinversionorinvagination.Amongoursixconditions,thenumbers2,3,and4correspondtothethree mainconditionsdistinguishe dearlierbyDeBeauchamp (1932). (1)Completelyinvertedpenispapilla.Inthiscasethe entirepenispapillamaybefullyinvertedor invaginated,e.g., D.adenodactylosum D.albidum Kenk,1978, D.magnum (Stankovic ,1969),and D. cf. beauchampi (cf.SluysandBenazzi,1992). (2)Invertedtipofpenispapilla.Thisconditionrefersto ahistologically homogeneous penis(i.e.,withno stronghistologicaldifferencesbetweenthebasalpart andtheapicalpartofthepenispapilla)withanapical partthatmaybetemporarilyinvaginated,asisthe casein,forexample, D.mrazeki. (3)Pseudoflagellum.Thiscaseispresentinspecieswitha penispapillathatcanbedividedintotwoparts,abasal partwithastronglayerofcircularmuscleandathin distalpartwithafewornomusclesandwithan externalepithelium(whenitiseverted)thatisthicker thanthebasalpart.Inthiscaseonlythedistalpartof thepapillaisinvertible,whilethebasalpartprotrudes intotheatrium.Thepseudo-flagellumismorefrequent whentheapicalpartofthepenispapillaisverythin withrespecttothebasalpart.Itispresent,for example,in D.maculatum D.sanctinaumi D.albidum, D.lacustre D.minimum,D.translucidum Kenk,1978, D.komareki (Stankovic ,1969), D.nausicaae D.remyi DeBeauchamp,1926, D.caspicum Porfirjevaand Dyganova,1973, D.longipenis ,Komarek,1916,andin theSardinian D.nuraghum (thispaper).Apeculiar caseisfoundin D.tubuliferum DeBeauchamp,1919, inwhichtheapicalpartofthepenispapillahasthe shapeofatube,whichmaybeconsideredtorepresent apseudo-flagellum.Asimilarconditionoccursin D. hercynicum Flo ¨sser,1959(cf.Sluys2012). (4)Trueflagellum.Thisdesignatesatall,pleatedsection oftheliningepitheliumofthedistallumenofthe penispapillathatmaybeevertedorinverted, matchingthedefinitionofKoma rek(1926)(Fig.7). Moreover,thetrueflagellumiseasilydistinguished bythearrangementofitsmusculature,inthatthe circularfibersofthepenispapillareachonlytothe baseoftheflagellumanddonotextendontoit. Therefore,theflagellumshowsonlylongitudinal musclesunderneathitstall,vacuolatedepithelium. AccordingtoDeBeauchamp(1932)andGourbault (1972)the‘‘trueflagellum’’isacharacterthatonly occursin D.lacteum and D.infernale .Later,Kenk (1978)reporteditalsofor D.cruciferum (Stankovic 1969).In D.lacteum theflagellummaybecome extraordinarilylong,reachingthecopulatorybursa ofthepartnerandenvelopingthespermsduring copulation(cf.DeBeauchamp,1932,Fig.36). (5)Inflatedflagellum.Thepenispapillaischaracterized byabasalthickpart,withasquamousepitheliumthat isunderlainbyastronglayerofcircularmuscle,anda thinnerdistalpart.Theepitheliumofthisapicalpartis formedbylargeroundedvacuolatedcells,withbasal nuclei,veryrichinsecretiongranules.Thisepithelium coversboththeexternalandtheinternalapicalpartof thepenispapilla.Thepenispapillamayassume differentshapes,considerablyelongatedwithatunnel-shapedlumen,sac-shapeddelimitatingawide lumen,orglobularwithitsdistalpartinvertedintoa widerpenispapillalumen(likeapseudo-flagellum). ThisconditionispresentintheSardinianspecies D. vesiculosum (thispaper)anddiffersfromthetrue flagellumbecausethevacuolatedepitheliumispresent bothontheouterandinnersurfaceofthetipofthe penispapilla(Fig.6A). S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 110 N JournalofCaveandKarstStudies, August2013

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(6)Inflexibleflagellum.Theshapeofthisflagellumis independentofthedegreeofextensionorcontraction ofthepenispapillaandcannotbeinvaginatedor inverted.Itispresentinthespecies D.stenophallus CodreanuandBa lcesco,1967and D.nekoum Sluys, 2012(cf.Sluys,2012). S PHINCTER AllthreenewSardinian Dendrocoelum speciesshow aroundtheterminaltractofthebursalcanalawelldevelopedsphincter,i.e.,anincreaseinthethicknessofthe circularmusclelayer.Suchasphincterwasfirstmentioned byDeBeauchamp(1932)and,subsequently,byKenk(1978) for D.adenodactylosum .However,examinationof D. adenodactylosum materialfromthecollectionsoftheNCB Naturalis(sectionZMA;specimensnumberRS157-1,1572),revealedthatawell-developedcoatofmusclesispresent ontheverticallyrunningpartofthecanal(i.e.,afterithas curveddownwardstoopenintotheatrium)consistingof alternatingrowsoflongitudinalandcircularmuscle;the horizontalpartofthebursalcanalismerelysurroundedbya layeroflongitudinalmusclefibers.Thiswasalsocorrectly observedbyDeBeauchamp(1932,1937),whodescribedthe verticalsectiontoconstituteasphincter.However,itlooks differentfromapropersphincter,whichprobablyinduced Kenk(1978,p.13)torefertoitas‘‘sphincter’’.Kenk(1978) reportedthe‘‘sphincter’’in D.adenodactylosum tobe formedprincipallybycircularmuscles,contrarytoour observationsandthoseofDeBeauchamp(1932).Inour opinion,thedifferencebetweenthemusculatureonthe horizontalandverticalsectionsofthebursalcanalmerely givestheimpressionofthepresenceofatruesphincter. Furthermore,thatthisisnotatruesphincterisalso suggestedbythefactthatthereisnodistinctconstriction ofthelumenofthecanal.Allofthiscontrastswiththe situationintheSardinianspecies,whichclearlyshowatrue sphinctertobepresentinallthreespecies,whiletherestof thebursalcanalissurroundedbyasubepitheliallayerof circularmuscles,followedbyalayeroflongitudinalfibers. Atruesphincterisalsopresentin,forexample, D.agile DeBeauchamp,1932, D.barbei DeBeauchamp,1956, D. atriostrictum (CodreanuandBa lcesco,1967), D.caspicum D.clujanum Codreanu,1943, D.collini,D.cruciferum (Stankovic ,1969), D.grimmi Dyganova,1983, D.remyi and D.romanodanubialis (Codreanu,1949). A CKNOWLEDGEMENTS WearegratefultoR.Salaris,D.Salaris,M.Piras,B. Cadeddu,R.Fadda,M.Deiana,G.Tomasin,andM.Foisfor theirkindhelpinthefield.WethankG.Corsoforherkind support.Thisresearchwasf undedbytheMinisterodell’Universita `edellaRicercaScientificaeTecnologica(MIURPRIN2008‘L’endemismonellafaunaitaliana:dallaconoscenzasistematicaebiogeograf icaallaconservazione’),the EuropeanProgramINTERREGSardinia-Corsica-Tuscanyon Biodiversity,andbytheregioneAutonomaSardegna(CRP60215).P.MarciawassupportedbyagrantfromRAS (‘‘Promozionedellaricercascie ntificaedell’innovazionetecnologicainSardegna’’,POSardegnaFSE2007-13,L.R.7/2007). G.A.StocchinoacknowledgesfinancialsupportbyFondazione BancodiSardegnaandSYNTHESYS,aprogrammeofthe EuropeanCommissionunderthe6thResearchandTechnologicalDevelopmentFrameworkProgramme‘‘Structuringthe EuropeanResearchArea’’,whichenabledhertoworkatthe ZoologicalMuseumAmsterdaminNovemberandDecember 2008(grantnumber:NL-TAF4717).Completionofthisstudy wasmadepossiblebyagrantfromtheNetherlandsCentrefor BiodiversityNaturalistoR.Sluys. R EFERENCES Argano,R.,Messana,G.,DeMatthaeis,E.,Cobolli,M.,andKetmaier, V.,1997,Usodimarcatoribiochimiciperlostudiodelpopolamento stigobiodiSardegna,gliAsellota(CrustaceaIsopoda):Biogeographia,v.19,p.157–171. Bello,G.,Falleni,A.,Fredj,G.,Gremigni,V.,Hochberg,F.G.,andVernet, G.,1995,‘‘Turbellaria’’,Gnathostomulida,Orthonectida,Dicyemida, Nemertea, in Minelli,A.,Ruffo,S.,andLaPosta,S.,eds.,Checklist delleSpeciedellaFaunaItaliana,v.4:Bologna,Calderini,p.1–35. Benazzi,M.,1938,TricladiPaludicolidellaSardegna:ArchivioZoologico Italiano,v.25,p.85–94. Benazzi,M.,1982,Tricladicavernicoliitaliani:LavoridellaSocieta ` ItalianadiBiogeografia,n.s.,v.7,p.7–14. Bromley,H.J.,1982,Themorphology,karyologyandreproductionofa newspeciesof Dendrocoelum O ¨ rsted(Turbellaria,Tricladida)fromthe headwatersoftheriverJordan:IsraelJournalofZoology,v.31, p.119–136. Cassola,F.,1982,IlpopolamentocavernicolodellaSardegna:Lavoridella Societa `italianadiBiogeografia,n.s.,v.7,p.615–755. Casu,S.,Pala,M.,andVacca,R.A.,1982,Distribuzionegeograficain Sardegnadiplanaried’acquadolceappartenentiallaspecie Dugesia ( S .) polychroa e Dugesia ( S .) mediterranea (Turbellaria,Tricladida): BollettinodellaSocieta `SardadiScienzeNaturali,v.21,p.177–184. Cianficconi,F.,Corallini,C.,andMoretti,G.P.,1998,Trichopteranfauna oftheItaliansprings, in Botosaneanu,L.,ed.,StudiesinCrenobiology:TheBiologyofSpringsandSpringbrooks:Leiden,Backhuys Publisher,p.125–140. Codreanu,R.,1949,SurunnouveauTricladeocule dude file duDanube: Palaeodendrocoelumromanodanubialis n.g.,n.sp.:Bulletinbiologique delaFranceetdelaBelgique,v.83,p.284–287. Codreanu,R.,andBa lcesco,D.,1967,Surtrois Dendrocoelides aveugles nouveauxdessourcesduBanat(Roumanie):RevueRoumainede Biologie,Se riedeZoologie,v.12,p.287–294. Cottarelli,V.,Bruno,M.C.,andForniz,C.,1996,CopepodiArpacticoidi eSincaridi(Crustacea)diacquesotterraneedelleisolecircumsarde: Biogeographia,v.18,p.261–272. DeBeauchamp,P.,1931,NouvellesdiagnosesdeTricladesobscuricoles, IV.Essaid’uneclassificationdesDendrocoelidae:Bulletindela Socie te ZoologiquedeFrance,v.55,p.155–163. DeBeauchamp,P.,1932,Turbellarie s,Hirudine es,Branchiobdellide s: Biospeleologica58,ser.2:ArchivesdeZoologieExpe rimentaleet Ge ne rale,v.73,p.113–380. DeBeauchamp,P.,1937,Turbellarie sTricladesdeYougoslaviere colte s parMM.RemyetHubault:BulletindelaSocie te Zoologiquede France,v.62,p.351–365. Ferreri,D.,1995,Molluschi,IrudineieTurbellariTricladidelleacque dolcidellaprovinciadiLecce:ThalassiaSalentina,v.21,p.29–49. doi:10.1285/i15910725v21p29. Giusti,F.,andCastagnolo,L.,1983,NotulaemalacologicaeXXX:I molluschiviventi,terrestried’acquadolce,nellostudiobiogeografico dell’IsoladiSardegna:LavoridellaSocieta `ItalianadiBiogeografia,n. s.,v.8,p.227–249. G.A.S TOCCHINO ,R.S LUYS ,P.M ARCIA AND R.M ANCONI JournalofCaveandKarstStudies, August2013 N 111

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Gourbault,N.,1972,RecherchessurlesTricladesPaludicoleshypoge s: Me moiresduMuse umNationald’HistoireNaturelle,ser.AZoologie no.73,249p. Gourbault,N.,1994,Turbellaria,Tricladida, in Juberthie,C.,andDecu, V.,eds.,EncyclopaediaBiospeologica,v.I:Moulis,France,Socie te deBiospe ologie,p.41–44. Harrath,H.A.,Sluys,R.,Ghlala,A.,andAlwasel,S.,2012,Thefirst subterraneanfreshwaterplanariansfromNorthAfrica,withan analysisofadenodactylstructureinthegenus Dendrocoelum (Platyhelminthes,Tricladida,Dendrocoelidae):JournalofCaveand KarstStudies,v.74,p.48–57.doi:10.4311/2011LSC0215. Hyman,L.H.,1931,Studiesonthemorphology,taxonomy,and distributionofNorthAmericantricladTurbellaria.IV.Recent Europeanrevisionsofthetriclads,andtheirapplicationtothe Americanforms,withakeytothelatterandnewnotesondistribution: TransactionsoftheAmericanMicroscopicalSociety,v.50,p.316–335. Juberthie,C.,andDecu,V.,1994,Structureetdiversite dudomaine souterrain;particularite sdeshabitatsetadaptationsdesespe `ces, in Juberthie,C.,andDecu,V.,eds.,EncyclopaediaBiospeologica,v.I: Moulis,France,Socie te deBiospe ologie,p.5–22. Kenk,R.,1978,Theplanarians(Turbellaria:TricladidaPaludicola)oftheLake OhridinMacedonia:SmithsonianC ontributionstoZoologyno.280,56p. Koma rek,J.,1926,Contributiona `larevisiondesTricladestche `quesd’eau douce,19p.[French,condensedtranslationthatappearstobepartof Koma rek,SupplementtotheAccountofFresh-WaterTricladidea FoundinBohemiabyVejdovsky ,Ve stn kKra lovske C eske Spolec nostiNauk,Tr idamatematicko-pr rodove decka ,Sborn kVejdovske ho,for1925,article7,32pages,1plate[inCzech],buttheFrench summarydoesnotappearinthejournal]. Lepori,N.G.,1951,Sullecaratteristichemorfologicheesullaposizione sistematicadellaplanariadiSardegnaeCorsicagia `ascrittaa Dugesia (Euplanaria) gonocephala (Duge `s):AttidellaSocieta `Toscanadi ScienzeNaturali,v.58,p.28–47. Lindberg,K.,1956,Cyclopides(Crustace sCope podes)delaSardaigne: MemoriedellaSocieta `EntomologicaItaliana,v.35,p.71–79. Manconi,R.,Ledda,F.D.,Serusi,A.,Corso,G.,andStocchino,G.A., 2009,Spongesofmarinecaves:Notesonthestatusofthe Mediterraneanpalaeoendemic Petrobionamassiliana (Porifera:Calcarea:Lithonida)withnewrecordsfromSardinia:ItalianJournalof Zoology,v.76,no.3,p.306–315.doi:10.1080/11250000802629471. Mart nez-Ansemil,R.,andSambugar,B.,2008,Anellididiambienti acquaticisotterraneidiSardegna:Memoriedell’IstitutoItalianodi Speleologia,ser.II,v.21,p.110–112. Me dail,F.,andQue zel,P.,1999,BiodiversityhotspotsintheMediterraneanBasin:Settingglobalconservationpriorities:Conservation Biology,v.13,p.1510–1513.doi:10.1046/j.1523-1739.1999.98467.x. Myers,N.,Mittermeier,R.A.,Mittermeier,C.G.,daFonseca,G.A.B., andKent,J.,2000,Biodiversityhotspotsforconservationpriorities: Nature,v.403,p.853–858.doi:10.1038/35002501. Moretti,G.P.,andCianficconi,F.,1983,Leattualiconoscenzesui TricotteridellaSardegna:LavoridellaSocieta `ItalianadiBiogeografia,v.8,p.593–639. Pala,M.,Casu,S.,andVacca,R.A.,1980a,Primidatisulladistribuzione geograficadiplanariedelgruppo Dugesialugubris-polychroa (Turbellaria,Tricladida)inSardegna,conparticolareriguardoallapresenza diunamutazionetetraploidenelfiumeCixerri(Siliqua,Cagliari): BollettinodellaSocieta `SardadiScienzeNaturali,v.19,p.183–188. Pala,M.,Casu,S.,andVacca,R.A.,1980b,RinvenimentodiunaPlanaria ascrivibilea Dugesiaetruscamonoadenodactyla Lepori(Turbellaria, Tricladida)nell’IsoladiMolara(Sardegna):BollettinodellaSocieta ` SardadiScienzeNaturali,v.19,p.177–181. Pala,M.,Casu,S.,andVacca,R.A.,1980c,Sullapresenzadi Crenobia alpina (Dana)(Turbellaria,Tricladida)inSardegna:Bollettinodella Societa `SardadiScienzeNaturali,v.19,p.171–175. Pala,M.,Casu,S.,andStocchino,G.A.,1999,Karyologyandkaryotype analysisofdiploidfreshwaterplanarianpopulationsofthe Dugesia gonocephala group(Platyhelminthes,Tricladida)foundinSardinia: Hydrobiologia,v.392,p.113–119. Pala,M.,Casu,S.,andVacca,R.A.,1981, Dugesiahepta ,nuovaspeciedi planariadiacquadolcediSardegnaappartenenteallasuperspecie Dugesiagonocephala (Duge `s)(Turbellaria,Tricladida):Bollettino dellaSocieta `SardadiScienzeNaturali,v.20,p.97–107. Pala,M.,Stocchino,G.A.,Corso,G.,andCasu,S.,2000,Anewspeciesof freshwaterplanarianbelongingtothegenus Dugesia (Platyhelminthes, Tricladida)fromSardinia:ItalianJournalofZoology,v.67, p.379–383.doi:10.1080/11250000009356343. Pala,M.,Vacca,R.A.,Casu,S.,andStocchino,G.A.,1995,The freshwaterplanarian Dugesiasicula LeporifromSardinia(Platyhelminthes,Tricladida):Hydrobiologia,v.310,p.151–156. Pesce,G.L.,1985,ThegroundwaterfaunaofItaly:Asynthesis: Stygologia,v.1,no.2,p.129–159. Pesce,G.L.,andMaggi,D.,1983,Primidatisullacomposizionedelle biocenosifreatichediSardegna:LavoridellaSocieta `Italianadi Biogeografia,n.s.,v.8,p.813–818. Puddu,S.,andPirodda,G.,1973,Catalogosistematicoragionatodella faunacavernicoladellaSardegna:RendicontideiSeminaridella Facolta `diScienzedell’Universita `diCagliari,v.43,p.151–205. Reisinger,E.,1960,VitaleNervenfa ¨rbungenbeiPlathelminthenundihre Abha ¨ngigkeitvomphysiologischenZustanddesOrganismus:Zeitschriftfu ¨rwissenschaftlicheZoologie,v.164,p.271–293. Ruffo,S.,andVignaTaglianti,A.,1975,Unanuova Bogidiella di Sardegna(CrustaceaAmphipoda,Gammaridae):FragmentaEntomologica,v.11,p.73–82. Sluys,R.,1989,Spermresorptionintriclads(Platyhelminthes,Tricladida): InvertebrateReproductionandDevelopment,v.15,p.89–95.doi: 10.1080/07924259.1989.9672028. Sluys,R.,2012,Anew,siblingspeciesofcaveflatwormfromSwitzerland (Platyhelminthes,Tricladida,Dendrocoelidae):RevueSuissede Zoologie,v.119,p.181–188. Sluys,R.,andBenazzi,M.,1992,Anewfindingofasubterranean dendrocoelidflatwormfromItaly(Platyhelminthes,Tricladida, Paludicola):Stygologia,v.7,p.213–217. Sluys,R.,Ribas,M.,andBagun a `,J.,1995,Taxonomy,ecology,and karyologyofanewspeciesof Phagocata fromSpain,withadiscussion onthephylogeneticsystematicsofthegenus Phagocata s.l. (Platyhelminthes,Tricladida,Paludicola):CanadianJournalof Zoology,v.73,p.557–568.doi:10.1139/z95-064. Stankovic ,S.,1926,U ¨ berzweineuePlanarienartenderBalkanhalbinsel nebstBemerkungenu ¨berVerbreitungder Planariaolivacea O. Schmidt:ZoologischerAnzeiger,v.66,p.231–240. Stankovic ,S.,1938,Noviprilozipoznavanjuendemic nihtrikladaOhridskog jezera(Nouvellecontributiona `laconnaissancedesTricladesende miques dulacd’Ohrid):GlasnikSkopskogNauc nogDrus tva,v.18,p.1–12. Stankovic ,S.,1969,Turbellarie sTricladesende miquesnouveauxdulac d’Ohrid:Archivfu ¨rHydrobiologie,v.65,p.413–435. Stankovic ,S.,andKoma rek,J.,1927,DieSu ¨ b wasser-Tricladendes WestbalkansunddiezoogeographischenProblemedieserGegend: ZoologischeJahrbu ¨cher,Abteilungfu ¨rSystematik,O ¨ kologieund GeographiederTiere,v.53,p.591–674. Steinmann,P.,1909,UntersuchungenanneuenTricladen:Zeitschriftfu ¨r wissenschaftlicheZoologie,v.93,p.157–184 + 1pl. Steinmann,P.,1910,EineneueGattungderpaludicolenTricladenausder UmgebungvonBasel( Polycladodesalba n.g.n.sp.):Verhandlungen derNaturforschendenGesellschaftinBasel,v.21,p.186–196. Stella,E.,1957,IlPlanctondelleacqued iunagrottadellaSardegna:Bollettino diZoologia,v.24,no.1,p.39–44.doi:10.1080/11250005709438235. Stocchino,G.A.,2003,Ecosistemisorgentizidellepiccoleisolemediterranee:Biodiversita `etipologieambientalinelParcoNazionaledell’ Asinara.PhDThesis,UniversityofSassari,169p. Stocchino,G.A.,Corso,G.,Manconi,R.,Casu,S.,andPala,M.,2005, EndemicfreshwaterplanariansofSardinia:Redescriptionof Dugesia hepta (Platyhelminthes,Tricladida)withacomparisonofthe Mediterraneanspeciesofthegenus:JournalofNaturalHistory, v.39,no.22,p.1947–1960.doi:10.1080/00222930500060025. Stocchino,G.A.,Marcia,P.,Grafitti,G.,Manconi,R.,Corso,G.,Casu, S.,andPala,M.,2008,TricladicavernicolidiSardegna:Memorie dell’IstitutoItalianodiSpeleologia,ser.II,v.21,p.121–124. Vejdovsky ,F.,1883,Exkrec n appara tPlanari [TheexcretoryApparatusof Planarians]:Zpra vyozaseda n C eske Spolec nostiNaukvPraze,p.273–280. Vejdovsky ,F.,1895,ZurvergleichendenAnatomiederTurbellarien (ZugleicheinBeitragzurTurbellarien-FaunaBo ¨hmens):Zeitschrift fu ¨rwissenschaftlicheZoologie,v.60,p.163–214. Vila-Farre ,M.,Sluys,R.,Almagro,I .,Handberg-Thorsager,M.,and Romero,R.,2011,Freshwaterplanarians(Platyhelminthes,Tricladida)fromtheIberianPeninsulaandGreece:diversityandnoteson ecology:Zootaxa,v.2779,p.1–38. S UBTERRANEANAQUATICPLANARIANSOFSARDINIA,WITHADISCUSSIONONTHEPENIALFLAGELLUMANDTHEBURSALCANALSPHINCTERINTHEGENUS D ENDROCOELUM (P LATYHELMINTHES ,T RICLADIDA ,D ENDROCOELIDAE ) 112 N JournalofCaveandKarstStudies, August2013

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OCCURRENCEOFTROGLOBITICCLIVININESINCHINA (INSECTA:COLEOPTERA:CARABIDAE) M INGYI T IAN DepartmentofEntomology,CollegeofNaturalResourcesandEnvironment,SouthChinaAgriculturalUniversity,Wushanlu483,Guangzhou,Guangdong 510640,China,mytian168@yahoo.com.cn Abstract: Anewgenus, Guiodytes gen.nov.,andtwonewspecies, G.cavicola sp.nov. and G.bedosae sp.nov.,ofthegroundbeetletribeClivinini(Coleoptera:Carabidae)are describedfromtwocavesofGuangxi,southernChina. Guiodytes gen.nov.is characterized.ItisthefirstgenusoftroglobiticclivininesdiscoveredinAsia,andits relationshipswithotherclivininesremainunclear. I NTRODUCTION Therearenumerouscavernicolouscarabidbeetles recordedintheworld(Casaleetal.,1998).ManyanophtalmousspeciesbelongtothetribeClivininiofScaritinae, especiallytothehumicolousgenus Reicheia Saulcy,1863. Butthereareafeweyelesstroglobiticclivininespecies: Italodytesstammeri Mu ¨ller,1938(fromItaly), Typhloreicheia monacha CasaleandMarcia,2011,and T.ilianae Casaleand Marcia,2011(fromSardinia,Italy), Spelaeodytesmirabilis Miller,1863(fromCroatia), Antroforcepsbolivari Barr,1967 (fromMexico),and Trogloclivinabrehieri Deuve,2003(from NewBritainIsland,PapuaNewGuinea).Anothertroglobiticspecies, Clivinasubterranea Decu,NitzuandJuberthie, 1994(fromRomania),isnotblind. InChina,thefirsthighlymodifiedtroglobiticcarabid species, Sinaphaenopsmirabilissimus UenoandWang,was describedin1991(UenoandWang,1991).AllChinese troglobiticgroundbeetlesaremembersofeitherthetribe TrechiniorthetribePlatynini.Trechiniisthedominantgroup, representedsofarbyabouteightyspeciesofmorethanthirty genera(Chenetal.,2001;Tian,2008).Althoughseveralspecies oftheplatyninegenera Jujiroa Ue no,1952and Xestagonum Habu,1978havebeenrecorded,allaremicrophthalmous,with reduced,butevident,eyes(VignaTaglianti,1995;Uenoand Kishimoto,2001;Deuve,2001,2004;Ueno,2007). Recently,incollaborationwithDr.LouisDeharvengof theMuse umNationald’HistorieNaturelleinParis,a biologicalsurveyoncavefaunawasconductedinGuangxi, southernChina,aspartofresearchactivitiesoftheWorld BankGEF-financedprojectGuangxiIntegratedForestry DevelopmentandConservation.Oneofthemostimportant discoveriesduringthesurveywasthatoftwointeresting specimensofeyelessclivinines.Thisisthefirstrecordof blindcarabidclivininesinChina.Detailedstudyshowed thattheycorrespondtotwodifferentspeciesofanewgenus. M ATERIALSAND M ETHODS Thespecimenswerecollectedbyhandincavesof Guangxiandpreservedin60%ethanolsolutionbefore study.Othersurfaceorsoilspeciesofclivinines,dryand mountedspecimens,wereexaminedforcomparison.All specimensaredepositedintheinsectcollectionsofSouth ChinaAgriculturalUniversity,Guangzhou,Guangdong Province. Dissectionandobservationwereconductedundera LeicaS8AP0microscope.Genitaliaandrelatedpiecesof smallstructureswereremoved,putin10%potassium hydroxidefor24hoursforcleaning,thenstuckonpaperboardandpinnedbeneaththeassociatedspecimen. PicturesweretakenbymeansofaLeicaMicrosystem LASV3.6,andthenprocessedusingAdobePhotoshop CS5software. Abbreviationsusedinthetextareasthesameas describedinTian(2009):HW 5 headwidth;HL 5 head length;PW 5 pronotalwidth;PL 5 pronotallength;EW 5 elytralwidth;EL 5 elytrallength.Thetermsforfemale genitaliaareasinDeuve(1993). R ESULTS G UIODYTES GEN NOV Typespecies: Guiodytescavicola sp.nov.(Guangxi) Genericcharacteristics: Descriptionisbasedentirelyon females.HabitusasinFigures1and2.Bodymoderately elongate;uppersidefinelyandsparselypunctate.Head stout,eyeseffaced;wingofclypeusnarrow,sharply projectedanteriorly;bothsupraorbitalsetiferouspores present,locatedfarfromeachother;supraorbitalfurrows wideandlong;supra-antennalplateswelldefined,broad andstronglyconvex,muchwiderthanclypealwings;frons andvertexconvex;fronswithoutcarinaortubercle;orbits welldeveloped,genaewelldeveloped;labrumcomplete, mandiblesquitestout,leftmandiblewithoutmediantooth; maxillarypalpiensiform;neckconstrictedbypunctures. Antennomeres1and2glabrous,withalongerpreapical setaonantennomere1andashortoneonbaseof antennomere2;antennomere2normallyjointedto antennomere1;antennapubescentfromantennomere3; antennomeres5–10cylindrical( G.cavicola sp.nov.),or distinctlysub-moniliform( G.bedosae sp.nov.). Pronotum(Figs.8and9)peltate,medianlinenot bifurcateanteriorly,withtwonormalsetaeateachside, M.Tian–OccurrenceoftroglobiticclivininesinChina(Insect:Coleoptera:Carabidae). JournalofCaveandKarstStudies, v.75,no.2, p.113–120.DOI:10.4311/2011LSC0226 JournalofCaveandKarstStudies, August2013 N 113

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oneatanteriorthird,onenearhindangle;medianlinewell marked,moderatelydeep,notreachingbase,butjoining basalconstriction;baseofpronotumconspicuouslydepressed;sidewiththreedistinctteethorprojectionsbefore hindangle;hindanglenearlyrectangular;basalborder closetopeduncle;anteriortransverseimpressionfreefrom medianline,veryclosetoanteriormargin. Elytra(Figs.1and2)elongate-ovate,withbaseand shouldersserrate,lateralmarginslightlyundulate( G. cavicola sp.nov.)ordistinctlycrenulate( G.bedosae sp. nov.);striaedeep,intervalsofsubequalwidth,strongly convex;stria3withfourfoveolatesetiferouspores; umbilicalsetiferouspuncturescompletethroughout,with severalmuchlongersetaeapartfromtheshortones;base ofstria6evidentlycarinate,striae7and8jointedatbase, carinateatbaseandapexrespectively.Hindwingsreduced. Proepisternumnottumidlaterally,invisiblefrom above;protarsomere1notenlarged;mesotibiawitha conspicuoustuber-likespuratsubapex. Femalegenitalia (Figs.12and13) : Bothgonosubcoxite IXandgonocoxiteIXstronglyelongate,gonocoxiteIX deeply( G.cavicola sp.nov.)orslightlycurved( G.bedosae sp.nov.),sharp( G.cavicola sp.nov.)orblunt( G.bedosae sp.nov.)atapex,withensiformsetaeonoutermargin. Male: Unknown.Inspiteofourefforts,wewereunable tocatchanotherspecimeninthecaves,andnomaleis currentlyavailable.Itispresumedthat Guiodytes isasrare asmanyothercavedwellingspeciesofCarabidaeinChina. Relationshipof Guiodytes withinClivinini: Sofarthe knowngeneraofanophthalmousclivininesareisolated. Italodytes Mu ¨llerisendemictoApulia,southernItaly, Spelaeodytes Miller,1863toCroatia, Antroforceps Barr, 1967toMexico, Trogloclivina toNewBritain,PapuaNew Guinea,and Guiodytes toGuangxi,southernChina. Jeannel(1957)wasprobablyrighttoseparate Italodytes fromClivinini,consideringitspeculiarmorphological characteristics. Guiodytes isevidentlycloserto Trogloclivina thanto Italodytes inappearance,buttherelationships arenotclear.Intheabsenceofmales,itisdifficultto Figure1. Guiodytescavicola sp.nov.,habitusofholotype. Figure2. Guiodytesbedosae sp.nov.,habitusofholotype. O CCURRENCEOFTROGLOBITICCLIVININESIN C HINA( I NSECT: C OLEOPTERA: C ARABIDAE) 114 N JournalofCaveandKarstStudies, August2013

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Figures3–7. Guiodytes spp.nov.:3.Headof G.cavicola ,dorsalview.4.Headof G.bedosae ,dorsalview.5.Headof G. cavicola ,ventralview.6.Rightantennaof G.cavicola. 7.Rightantennaof G.bedosae M.T IAN JournalofCaveandKarstStudies, August2013 N 115

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discussthephylogeneticpositionof Guiodytes withinthe ChineseorOrientalfaunaofClivinini(Putzeys,1866; Baehr,1999;Balkenohl,1999,2001;YangandTian,2008). Thenewgenusmighthaveaffinitieswith Clivina Latreille, 1802.Itdiffersfrom Clivina byitseyescompletelyreduced, anteriorandposteriorsupraorbitalsetiferousporeslocated farfromeachother,orbitsandgenaewelldeveloped,each ofantennomeres5–10longerthanwide,pronotumwith threetoothedprojectionsjustbeforehindangle,andhind wingsreduced. Adaptationof Guiodytes tosubterraneanenvironments: Apartfromeffacedeyesandreducedhindwings,noother evidentmorphologicalmodificationswerefoundin Guiodytes speciesforadaptingtoasubterraneanenvironment. Theappendages,suchaspalpi,antennae,andlegs,arenot moreslenderthanthoseinsurfacespecies,though antennomeres5–10aremoreorlesselongate.Thebody of Guiodytes ismoderatelyelongateforClivinini,andthe elytralchaetotaxalpatternisnotsospecial,astrichobothriaarenotverylong.Althoughtheylookslightlylighter incolorthansurfaceorsoilspeciesofClivinini,both speciesof Guiodytes arenotdepigmented. Etymology: ‘‘ Gui ’’istheabbreviationofGuangxi ZhuangMinorityAutonomousRegioninChinese.The genericname, Guiodytes ,referstothedistributionofthe newgenus. Distribution: SouthernChina(Guangxi). G UIODYTESCAVICOLA SP NOV Holotype: Female,NongshuiCave,ShangjiaCun,Disu Xiang,Du’anXian,HechiCity,Guangxi,23 u 84.066 9 N/ 108 u 00.201 9 E,2010-IV-24,MingyiTianleg. Description: Length(fromapexofleftmandibletoend ofelytra)7.0mm;width2.2mm. Lightdarkbrown;tarsiandantennaelightbrown,palpi yellow,apexofmandiblesdarkbrown.HabitusasinFig.1. Head(Figs.3–5)stout,slightlylongerthanwide,HL/ HW 5 1.17;narrowerthanpronotum(HW/PW 5 0.71); clypeuswide,ratherflat,bisetose,slightlyemarginateat anteriormargin;lateralwingsnarrow,projectedanteriorly, extendeddistinctlybeyondanteriormarginofclypeus, separatedfromclypeusbyanindistinctnotch;supraantennalplateswelldefined,clearlyseparatedfromclypeal wingsbyobviousnotch,moreorlessrounded,strongly convex,smoothandglabrous;fronsandvertexconvex; supraorbitalfurrowsverydeepandwide,extendedtothe levelofhindsupraorbitalpore,subparallelalongfrons, stronglydivergentafterwards;supraorbitalcarinaabsent; fronsseparatedfromclypeusbyinconspicuousfrontoclypealsuture;eyescompletelyreducedanddisappeared;both anteriorandposteriorsupraorbitalporespresent,located farfromeachother;genaewelldeveloped;neckconstrictiondistinct,beginningatlevelofhindsupraorbitalpore; labrumconspicuouslywide,aswideasclypeusexcluding clypealwings,5-setose,andciliateonbothsides,almost straightatanteriormargin;mandiblestout,moreorless widened,rightmandiblewithoutmediantooth;palpi ensiform,glabrous,apicalsegmentsofbothmaxillary andlabialpalpimuchlongerthanpenultimateones respectively;labialpalpomere2bisetose;ligulathin, unisetoseatapex;mentumwelldeveloped,withtwopairs ofsetae,onepairsituatedbeneathmentaltooth,otherat base,closetosidemarginsofmentum;mediantooth pointed,laterallobeswide,almosttruncateatapex; submentumdistinctlyseparatedfrommentum,quadrisetose.Antennaefiliform(Fig.6),comparativelylongfor Clivinini,extendedtoscutellum;antennomeres1and2 glabrousandsmooth,antennapubescentfromantennomere3;antennomeres2and3subequalinlength; antennomeres5–10subcylindrical,somewhatflat,distinctlylongerthanwide. Pronotum(Fig.8)peltateinform,muchwiderthan head;discsmooth,moderatelyconvex,slightlylongerthan wide(PL/PW 5 1.09);apexdeeplyconcave,marginatedin medianportion,anterioranglesstronglyprotruded,nearly rectangular,widestataboutmiddle;sidestronglyserrated justbeforehindangle,withthreeconspicuoustoothed projections;marginalchannelwideanduneven,widest nearanteriorlateralsetiferouspore,endedbeforeposterior lateralsetiferouspore;anteriorsetiferousporesituatedat aboutapicalfourth,posterioroneataboutbasalthird,just beforehindangle;baseunbordered;medianlineclearand deep;basaltransverseimpressiondeepandwide,with severaltransversestriaebehindandasmallmarginal denticle;anterortransverselinefreefrommedianline,very closetoanteriormargin.Peduncledistinct. Elytrawiderthanpronotum(EW/PW 5 1.10),elongate-ovate,EL/EW 5 1.85;discstronglyconvex;lateral marginslightlyexpanded,widestataboutmiddle;base moreorlessstraight;shouldersbroadlyobtuse,withthree largeserratedteeth;lateralmarginmoreorlesssmoothbut undulatenearbase;striaedeep,intervalsstronglyconvex; interval1withasmallbutdistincttubercleatbase,just beforescutellarpore;intervals5–9borderedatbase; intervals6–8carinateatbase,baseofintervals7and8 joined;interval3withfoursetiferouspores,setaemoderate inlengthforclivinines;marginalchannelwithuninterruptedseriesofsmallsetiferousporesandseverallargepores withmuchlongersetae;scutellarstriaandscutellarpore present;hindwingsreduced. Undersideofheadandproepisternumwithdenseisodiametricpunctures,withoutwrink les;prosternumsmooth,with sparsepunctures;abdomenalsternamoreorlesspunctured; sternumVIIoffemalewithtwopairsofsubapicalsetae; epipleuronwithafewcoarserpuncturesnearbase. Legsnotmodifiedcomparedwithsurfaceclivinines; forelegstout,profemurmoderatelydilated,smooth, withoutcarinaorrugaventrally;protibia(Fig.10)well developed,quadridentate,withdistinctandcomplete carinadorsally,sulcusindistinct;lateralupperspine elongate-ensiform,bluntatapex,muchlongerandstouter thansubapicalspur;protarsislender,tarsomere1very O CCURRENCEOFTROGLOBITICCLIVININESIN C HINA( I NSECT: C OLEOPTERA: C ARABIDAE) 116 N JournalofCaveandKarstStudies, August2013

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Figures8–13. Guiodytes spp.nov.:8.Pronotumof G.cavicola. 9.Pronotumof G.bedosae. 10.Protibiaof G.cavicola ,dorsal view.11.Protibiaof G.bedosae ,dorsalview.12.Femalegenitaliaof G.cavicola ,ventralview.13.Femalegenitaliaof G. bedosae ,ventralview. M.T IAN JournalofCaveandKarstStudies, August2013 N 117

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long,muchlongerthantarsomeres2–4combined;middle andhindlegsslender,mesotibiagraduallydilatedtowards apex,withaconspicuoustuber-likespuratsubapex. Femalegenitalia (Fig.12) : LateralotergiteIXwell developed,nearlyreversetriangularinform,withfourlong setaeatsubapexandapex;gonosubcoxiteIXtriangular, withonelongsetaatmedianportionofoutermargin; gonocoxiteIXverylongandslender,graduallyandevenly curved,verysharpatapex,withonelongsetanearbase,and twostrongandlongensiformsetaeonoutermargin. Male: Unknown. Etymology: Thenameofthisnewspeciesreferstoits cavehabitat. Distribution: China(Guangxi:Du’anXian).Known onlyfromthetypelocality.Itisacavewithtorrential floodingjustpasttheentrance(Figs.14and15).Nearthe entrancethereisapumpingstation. G UIODYTESBEDOSAE SP NOV Holotype: Female,PaomaCave,XiadongXiang,LongzhouXian,SouternGuangxi,106 u 86.903 9 E/22 u 37.422 9 N,2010-IV-15,MingyiTianleg. Description: Length(fromapexofleftmandibletoend ofelytra)5.2mm;width1.8mm. Lightdarkbrown;tarsiandantennaelightbrown,palpi yellow,apexofmandiblesdarkbrown.Habitusasin Figure2. Head(Fig.4)stout,slightlylongerthanwide,HL/LW 5 1.20,slightlynarrowerthanpronotum(HW/PW 5 0.74);clypeuswide,ratherflat,bisetose,distinctlyemarginatedatanteriormargin;lateralwingsnarrow,projected anteriorly,extendedclearlybeyondanteriormarginof clypeus,separatedfromclypeusbyanindistinctnotch; supra-antennalplatewelldefined,clearlyseparatedfrom clypealwingbyanobviousnotch,elongate-rounded, stronglyconvex,smoothandglabrous;fronsandvertex convex;supraorbitalfurrowdeepandwide,extendedto levelofhindsupraorbitalpore,slightlydivergentalong frons,posteriorlystronglydivergent;supraorbitalcarina absent;fronsseparatedfromclypeusbyinconspicuous frontoclypealsuture;eyescompletelyreduced;bothanteriorandposteriorsupraorbitalporespresent,locatedfar fromeachother;genaewelldeveloped;neckbroad, constrictiondistinct,beginningatlevelofhindsupraorbital Figures14–17.Habitatcavesof Guiodyte :14and15.EntranceofNongshuiCave.16and17.EntranceofPaomaCave. O CCURRENCEOFTROGLOBITICCLIVININESIN C HINA( I NSECT: C OLEOPTERA: C ARABIDAE) 118 N JournalofCaveandKarstStudies, August2013

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pore;labrumconspicuouslywide,aswideasclypeus excludingclypealwings,7-setose,andciliateonbothsides, almoststraightatanteriormargin;mandiblestoutand widened,rightmandiblewithoutmediantooth;palpi ensiform,glabrous,stout,dilatedataboutbasalthird; apicalsegmentsofbothmaxillaryandlabialpalpimuch longerthanpenultimateonesrespectively;labialpalpomere 2bisetose;ligulathin,unisetoseatapex;mentumwell developed,withtwopairsofsetae,onepairsituated beneathmentaltooth,otheratbaseclosetosidemarginsof mentum;mediantoothverysharp,laterallobeswide, almosttruncateatapex;submentumdistinctlyseparated frommentum,quadrisetose.Antennaesub-moniliform (Fig.7),moderateinlengthforClivinini,shorterthan abovespecies,extendedtobaseofpronotum;antennomeres1and2glabrousandsmooth,antennapubescent fromantennomere3;antennomeres2and3subequalin length;antennomeres5–10somewhatflat,slightlylonger thanwide. Pronotum(Fig.9)peltateinform,muchwiderthan head;discsmooth,moderatelyconvex;slightlylongerthan wide(PL/PW 5 1.04);anteriormargindeeplyconcave, anterioranglesprotruded,nearlyrectangular;widestat aboutmiddle;sidestronglyserratedjustbeforehindangle, withthreeconspicuoustoothedprojections;marginal channelwide,endedbeforeposteriorsetiferouspore; anteriorsetiferousporesituatedataboutapicalfourth, posterioroneataboutbasalthird,justbeforehindangle, whichismoreobtusethanthatof G.cavicola ;base unbordered,almoststraight;medianlinedeepanddistinct; basaltransversesulcuswideanddeep,narrowerthan G. cavicola ,withseveraltransversestriaebehind;anterior transverseimpressionfreefrommedianline,verycloseto anteriormargin.Peduncledistinct. Elytrawiderthanpronotum(EW/PW 5 1.21),elongate-ovate,EL/EW 5 1.73;discstronglyconvex;lateral margingentlyexpanded,widestataboutmiddle;basemore orlessstraight,littleindentationscorrespondingtothe intervals;shoulderswithtwoevidentserratedteeth,lateral margindistinctlycrenulatedthroughout;striaepunctate, deep,intervalsstronglyconvex;intervals6–8carinateat base,baseofintervals7and8separate;interval3withfour setiferouspores,setaemoderateinlengthforclivinines; marginalchannelwithuninterruptedseriesofsmall setiferousporesandseverallargeporeswithmuchlonger setae;scutellarstrialonganddistinct,scutellarpore present.Hindwingsreduced. Undersideofheadandproepisternumwithdense isodiametricpunctures,withoutwrinkles;prosternum smooth;abdomenalsternamoreorlesspunctured;sternum VIIoffemalewithtwopairsofsubapicalsetae;epipleuron withafewcoarserpuncturesnearbase. Legsnormalforclivinines;forelegstout,profemur moderatelydilated,smooth;protibia(Fig.11)welldeveloped,quadridentate,withdistinctandcompletecarina dorsally,butwithoutsulcus;lateralupperspineelongateensiform,bluntatapex,muchlongerandstouterthan subapicalspur;protarsislender,tarsomere1long,slightly longerthantarsomeres2–4combined;middleandhindlegs slender,mesotibiagraduallydilatedtowardsapex,witha conspicuoustuber-likespuratsubapex. Femalegenitalia (Fig.13) : LateralotergiteIXweakly sclerotized,withfourlongsetaeatinnermargin;gonosubcoxiteIXtriangular,withonelongsetaeatmedian portionofoutermargin;gonocoxiteIXlongandslender, slightlycurved,apexsomewhatblunt,eachsideofbase withonelongseta,andtwosmallandshortensiformsetae onoutermargin. Male: Unknown. Remarks: Thisnewspeciesissimilarto G.cavicola sp. nov.,butitissmaller,headstouter,labrum7-setose,palpi slightlystouter,antennaeshorterandsub-moniliform, lateralmarginofelytradistinctlycrenulatethroughout, andthefemalegenitaliawithgonocoxiteIXslightly curved,andbluntatapex. Etymology: ThisnewspeciesisnamedinhonorofDr. AnneBedos(Muse umNationald’HistorieNaturelle),a wellknownbiospeleologist. Distribution: China(Guangxi:LongzhouXian).The uniquespecimenwascaughtbyhandbeneathapieceof soilinPaomaCave,atabout150mfromtheentrance.The caveisaslongas4km,accordingtolocalvillagers,butwe hadonlyexploredasmallpartofitbecauseofflooding (Figs.15and16). A CKNOWLEDGEMENTS IamparticularlyindebtedtoDrs.LouisDeharveng, ThierryDeuve,andAnneBedos(MNHN)fortheir encouragementduringthestudy,especiallytheircritical reading,correctionofthetextandsuggestionstoimprove themanuscript.MythanksalsotoDr.TonyWhitten (FaunaandFloraInternational),Ms.LiuJin(World Bank),Mrs.FengBin,LiGuiyu(BiodiversityOffice, GuangxiForestryBureau,Nanning),andDr.LiYoubang (CollegeofLifeSciences,GuangxiNormalUniversity, Guilin)fortheirvariousassistances.Inaddition,Ithank theJournalofCaveandKarstStudiesAssociateEditors, AdvisoryBoardandreviewersfortheirencouragementand constructivecriticism.Thisstudywassponsoredbythe WorldBankGEF-financedprojectGuangxiIntegrated ForestryDevelopmentandConservation. R EFERENCES Baehr,M.,1999,TwonewgeneraofclivinineScaritinaefromtheOriental Region(Coleoptera:Carabidae), in Zamotajlov,A.,andSciaky,R., eds.,AdvancesinCarabidology,Krasnodar,Russia,MuisoPublishers,p.115–126. Balkenohl,M.,1999,NewClivininifromtheOrientalregion2. Clivina rugosofemoralis nov.spec.and Rugiluclivinaleonina nov.spec.from Laos(Coleoptera,Carabidae,Scaritinae):LinzerBiologischeBeitra ¨ge, v.31,no.1,p.337–344. M.T IAN JournalofCaveandKarstStudies, August2013 N 119

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Balkenohl,M.,2001,KeyandCatalogueoftheTribeClivininifromthe OrientalRealm,WithRevisionsoftheGenera Thliboclivina Kult,and Trilophidius Jeannel(Insecta,Coleptera,Carabidae,Scarititae,Clivinini):Sofia,PensoftPublishers,SeriesFaunisticano.21,83p. Barr,T.C.Jr.,1967, Antroforceps ,aneyelesscavescaritinefromMexico (Coleoptera:Carabidae):TheColeopterists’Bulletin,v.21,p.65–70. Casale,A.,andMarcia,P.,2011,Twonew Typhloreicheia speciesfrom Sardiniaandtheirbiogeographicalsignificance(Coleoptera,Carabidae, Scaritinae):ZooKeys,v.134,p.15–31.doi:10.3897/zookeys.134.1707. Casale,A.,VignaTaglianti,A.,andJuberthie,C.,1998,Coleoptera Carabidae, in Juberthie,C.,andDecu,V.,eds.,Encyclopaedia Biospeologicavol.II,MoulisandBucarest,Socie te Internationale deBiospe ologie,p.1047–1081. ChenZhiping,Decu,V.,Juberthie,C.,andUe no,S.-I.,2001,Chine, in Juberthie,C.,andDecu,V.,eds.,EncyclopaediaBiospeologica vol.III,MoulisandBucarest,Socie te InternationaledeBiospe ologie, p.1763–1781. Decu,V.,Nitzu,E.,andJuberthie,C.,1994, Clivinasubterranea (Caraboidea,Scaritidae),nouvelleespe `cedelagrotte‘‘Pes teradela Movile’’,DobrogeaMe ridionale,Roumanie:Me moiresdeBiospe ologie,v.21,p.41–45. Deuve,T.,1993,L’AbdomenetlesGenitaliadesFemellesdeCole opte `res Adephaga:Me moiresduMuse umNationald’HistoireNaturelle, no.155,184p. Deuve,T.,2001,Deuxnouveaux Semiaphaenops deChine,cavernicoles dansunkarstdunord-estYunnan(Coleoptera,Trechidae):Nouvelle Revued’Entomologie,v.18,p.187–192. Deuve,T.,2003,UnClivininitroglobieanophtalmed’unkarstdeNouvelleBretagne,enPapouasie-NouvelleGuine e(Coleoptera:Caraboidea: Scaritidae):NouvelleRevued’Entomologie,v.20,p.231–235. Deuve,T.,2004,Deuxnouvelles Jujiroa cavernicolesdesuddelaChineet dunordduVietnam(Coleoptera,Caraboidea):BulletindelaSociete entomologiquedeFrance,v.109,p.361–366. Jeannel,R.,1957,Re visiondespetitsScaritidesendoge svoisinde Reicheia Saulcy:RevueFranc aised’Entomologie,v.24,p.129–212. Miller,L.,1863,EinneuerGrottenka ¨ferausderGruppederScaritiden: WienerentomologischeMonatsschrift,v.7,p.28–30. Mu ¨ller,G.,1938, Italodytesstammeri ,nuovegenereenuovaspeciedi carabidicavernicolidell’Italiameridionale:AttidelMuseoCivicodi StoriaNaturalediTrieste,v.13,p.135–139. Putzeys,J.,1866,Re visionge ne raledesClivinides:AnnalesdelaSocie te EntomologiquedeBelgique,Bruxelles,v.10,242p. TianMingyi,2008,Anoverviewtocave-dwellingcarabid(Insecta: Coleoptera)inChina:Proceedingsofthe14thNationalConference ofSpeleology,Wulong,Chongqing(inChinese),p.333–342. TianMingyi,2009,Newrecordsandnewspeciesofcave-dwellingtrechine beetlesfromMulunNatureReserve,northernGuangxi,China (Insecta:Coleoptera:Carabidae:Trechinae):SubterraneanBiology, v.7,p.69–73. Ue no,S.-I.,2007,OccurrenceofaNewCaveSpeciesof Jujiroa (Coleoptera,Carabidae,Platyninae)fromCentralSichuan,Southwest China:Elytra,v.35,no.1,p.21–26. Ue no,S.-I.,andKishimoto,T.,2001,Anewcavespeciesofthegenus Jujiroa (Coleoptera,Carabidae,Platyninae)fromsouthernSichuan, SouthwestChina:JournalofSpeleologicalSocietyofJapan,v.26, p.30–36. Ue no,S.-I.,andF.X.Wang,1991,Discoveryofahighlyspecializedcave trechine(Carabidae:Trechinae)inSouthwestChina:Elytra,v.19, p.127–135. VignaTaglianti,A.,1995,Anew Jujiroa fromSichuan,China (Coleoptera,Carabidae):InternationalJournalofSpeleology,v.23, no.3–4,p.179–189. YangDefang,andTianMingyi,2008,TwonewrecordgeneraofClivinini inChina(Coleoptera:Carabidae): Entomotaxonomia ,v.30,no.4, p.255–258. O CCURRENCEOFTROGLOBITICCLIVININESIN C HINA( I NSECT: C OLEOPTERA: C ARABIDAE) 120 N JournalofCaveandKarstStudies, August2013

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THEFIRSTCAVERNICOLOUSNICOLETIIDAE(INSECTA: ZYGENTOMA)FROMTHEUNITEDARABEMIRATES L UIS E SPINASA 1 AND L UIS F.M ENDES 2 Abstract: Lepidosporamazyadi ,n.sp.,isdescribedanddifferentiatedfromalltheother known Lepidosporas.s .ThenewspecieswascollectedfromacaveinJebelHafeetinthe UnitedArabEmirates.Morphologyandpreliminaryanalysesof16SrRNADNA sequencesaredescribed.Thenewspeciesisuniquebecause,unlikeotherspeciesinthe genus,itlackssexuallydimorphicterminalfilaments.Itisalsothefirstnicoletiid reportedfromtheUnitedArabEmiratesandthesecondcave-adaptedspeciesfrom Arabia. I NTRODUCTION In1997,acavewasencounteredduringconstruction workonthetopofJebelHafeet,nearthevillageofMazyad intheUnitedArabEmirates.Subsequentexplorationof thecaveshowedittohave450metersofpassageanda depthof96meters,becomingthelongestcaveintheUAE. Duringtheoriginalexplorations,nicoletiidinsectswere observedandcollected,butsampleswheretoodegradedto beidentified.In2011,anotherexplorationtripwas conductedduringwhichsixspecimensweresuccessfully collected.Adescriptionofthisnewspeciesofnicoletiidand preliminaryDNAsequencedataareprovidedhere. M ATERIALSAND M ETHODS Dissectionsweremadewiththeaidofastereo microscope.Illustrationsweremadewiththeaidofa cameralucidaattachedtoamicroscope.Specimenswillbe depositedinthecollectionoftheAmericanMuseumof NaturalHistory,NewYork. GenomicDNAfromtheholotypeandafemale paratypewereextractedusingQiagen’sDNEasyTissue Kitbydigestingaleginlysisbuffer.Amplificationand sequencingofthe16SrRNAfragmentwasdoneasin EspinasaandGiribet(2009)followingstandardprotocols withprimersusedinthepastfornicoletiids.Chromatogramsobtainedfromtheautomatedsequencerwereread andcontigsmadeusingthesequenceeditingsoftware Sequencher3.0.Externalprimerswereexcludedfromthe analyses.Sequenceswerealignedandneighbor-joining analysiswasperformedwithClustalW2. R ESULTSAND D ISCUSSION Moleculardatawereobtainedfromtwoindividuals.A partial16SrRNAfragment486bpinlengthwasobtained (Fig.1).Bothspecimenshadidenticalsequences.NucleotidealignmentoftheUAEspecimenwithotherspeciesof Nicoletiidsfromwhich16SrRNAsequenceareavailable showedconsiderablesequencedifferences.Aneighborjoiningtreeshowedthenewspeciesdistantlyrelatedto Australiatelura, aninquilinespeciesofthesubfamily AtelurinaefromAustraliaandNewZealand(111bp differences;22.8%).Nevertheless,theseresultsshouldbe consideredaspreliminary.Withtheexceptionofthe AmericannicoletiidsofthesubfamilyCubacubaninae, veryfewstudiesontheOldWorldnicoletiidshaveused DNAsequencing,anditislikelythatotherunsequenced speciesaremorecloselyrelated. L EPIDOSPORAMAZYADI ,E SPINASAAND M ENDES N EWSPECIES (Figs.2A–H,3A–G) M ATERIAL :Holotypemale,body10mm,tarsusthirdleg 1.1mm.Paratypes:onemale(9mm),threefemales(13,13, and12mm),andonejuvenile(5.5mm).Inthevicinityof 40R375360E/3661550N,MagharetQasirHafeetCave, JebelHafeet,nearthevillageofMazyadandthecityofAl Ain,EmirateofAbuDhabi,UnitedArabEmirates.6/17/ 2011and8/8/2011.CollectedbyTimFogg,PamFogg, AngusTillotson,andAbdulRaheemAbdulRahman IqtedarAlMarzooqi. D ESCRIPTION :BodyproportionsasinFigure2A.Maximumbodylengthofmale10mm,offemale13mm. Maximumconservedlengthofantennaeandcaudal appendages8mm.Headandbodywithtypicalscales (Figs.2B,2G,3B–C,and3F).Generalcolorlightyellowto white.Headwithscales,macrochaetae,andmicrochaetaeas showninFigure2B.Pedicellusofmaleslightlyshorterthan firstarticleandwithadistinctprotuberantapophysisthat extendsinwardsandventrally(Fig.2B–C).Ontheapophysis basallytherearemultiplemicrochaetaeanddistallyacouple ofmacrochaetaeandadistinctsmallspiniformchaetaonits apex(Fig.2C).Basalarticlesofantennaeoffemalesimple. Appendageslong,asinothercave-adaptednicoletiid species.Thisismostevidentinthemaxillarypalp,whichis particularlylong(Figs.2Aand2E).Mandiblechaetotaxyas inFigure2H,withaboutfivemacrochaetaeoncentral-distal 1 SchoolofScience,MaristCollege,3399NorthRoad,Poughkeepsie,NY12601. luis.espinasa@marist.edu 2 InstitutodeInvestigac a oCient ficaTropical-JBT/CZ.R.daJunqueira,141300343Lisboa,Portugal.luis.mendes@iict.pt L.EspinasaandL.F.Mendes–ThefirstcavernicolousNicoletiidae(Insecta:Zygentoma)fromTheUnitedArabEmirates. Journalof CaveandKarstStudies, v.75,no.2,p.121–125.DOI:10.4311/2011LSC0256 JournalofCaveandKarstStudies, August2013 N 121

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portion.MaxillaasshowninFigure2E.Lastarticleof subequallengthtopenultimatearticle.Apexofgaleawith twosubequalconulesandtwoteethonlacinia(Fig.2F). LabialpalpasinFigure2D,withapicalarticledistinctly longerthanwideandlongerthanthepenultimatearticle. Pro-,meso-,andmetanotawithseveralmacrochaetaeon postero-lateralmargins,apartfromseveralsetaeofvaried sizes(Fig.2G).Legslong;hindtibiaabout8 3 longerthan wideandonefifthshorterthantarsus(Fig.3A). AbdominaltergitesasinFigure3F,withrobust macrochaetaeontheirposterioredges.UrotergiteX protruding,verydeeplyemarginatedinbothsexes,but moresoinmales.Inmalesmultiplemacrochaetaeplustwo rowsofscleritizedpegs,onelateral(Fig.3F)andtheother ventral(Fig.3G).ThetipsofurotergiteXalsowith scleritizedpegs(Fig.3F).Depthofemarginationalmostas deepasthemacrochaetaeandslightlywider(Fig.3F–G). InfemalesurotergiteXslightlylessdeep,withoutlateral andventralpegs,butinsteadofthepairofpegsatthetips theyhaveadistinctlargemacrochaetae. AbdominalsternitesasinFigures3B–D.UrosterniteI dividedintoamediansterniteand1 + 1lateralcoxites. UrosternitesI–VIIIentire.StyletsinurosternitesII–IX. Styletshaveaterminalspinewithsmallteethandventrally withtwomacrochaetaeplusanotherdistalpair(Fig.3B– D).StyletsIXlargerthanothers(Fig.3C–D).Posterior marginofurosterniteVIIIofmaleprominent,straight,and withacoupleofmacrochaetae(Fig.3C).Penisand parameraasinFigure3C.Parameralong,almost8 3 longerthanwide.Parameraattainabouttwothirdsthe lengthofstyletsIXinthelargestmale(10mm).Subgenital plateoffemaletriangulartosub-parabolic(Fig.3D).In thelargestfemale(13mm),theovipositorsurpassesapexof styletsIXbytwothirdsthelengthofstylets(Fig.3D). Gonapophyseswithabout14to15annuli.Maleand femalecerciandappendixdorsalissimple.Cerciwith trichobothria.(Fig.3E). P OSTEMBRYONIC D EVELOPMENT :Onlypartiallyunderstoodduetosamplesize.Bothmales10and9mmshare equalcharacteristics:pedicelluswithdistinctprotuberant apophysisandparameresattainabouttwothirdsthe lengthofstyletsIX.Inthethreefemales(13,13,and 12mm)theovipositorsurpassesapexofstyletsIXbytwo thirdstoonehalfthelengthofstyletsandgonapophyses withabout14to15annuli.Thespeciesappearstoshare withothercavernicolousspeciesaratherlatesexual development.Thesinglejuvenileisquitelarge,andat 5.5mmlongithasyettodevelopeithermaleorfemale character,andparameraorovipositorareabsent. E TYMOLOGY : mazyadi .Fromthenearbyvillageof Mazyad,wherethecaveislocated. R EMARKS :Thenewspeciesbelongswithinthesubfamily ColetiniinaebecauseithasurosterniteIdivided,II–VIII entire,andcoxitesIXofmalefree,malepedicelluswithan apophysiswithoutapparentglands,andparameresnot divided(Mendes,1988).Itbelongswithinthegenus Lepidospora becausestyletsarepresentonurosternites II–IXandtheyhavetypicalscales(Wygodzinsky,1980).It belongstothenominatesubgenusbecausetherearescales onboththebodyandonthecephaliccapsule.Itcanbe differentiatedfromotherspecieswithinthegenusbythe completelackofsclerotizedpegsonallthreeterminal filamentsinthemale(cerciandparacercus).Itis immediatelydistinguishablefromtheonlyother Lepidospora knownfromtheArabianPeninsula, L.(L.) angustotergum, atroglobiontordeepedaphicspeciesfrom southeasternOman(Mendes,2002a),bythequitedifferent urotergiteX,shorterandlessdelicateparamera,andlack ofsubmedianmacrochaetaeonurosterniteVIIIinthe male.Femalesaredifferentiatedbythedistinctsubgenital plateandmuchlongerovipositor. Thenewspeciesenterstheidentificationkeyspreviously proposedforthegenus(Mendes,2002b)atpoint3forthe males,asallremainingknown Lepidospora havepegsat leastonthecerci,asreportedearlier.Thefemalerunsto couplet21,andisquitedifferentfrom L.(L.)angustotergum ,asmentionedbefore.Itisnotpossibletoenterthe otheroption(21 9 )duetotheadultsizeandtheelongation oftheappendages.Thetwomostrecentlydescribed species,bothepigeanfromtheSocotraIslands(Mendes, 2004),weredescribedafterthekeywasproposed.Theycan bedifferentiatedfromthenewspeciesbecausemaleshave sclerotizedpegsonthecerciandparacercus,bythe elongationoflegs,maxillaryandlabialpalps,bythesize andslendernessofparamera,andbytheshapeofboththe antennalpedicellusandofurotergiteX.Femalesare unknowninthesespecies. D ISTRIBUTION :Knownonlyfromtypelocality. A CKNOWLEDGMENTS ThePresidentoftheUnitedArabEmirates,His HighnessSheikhKhalifabinZayedAlNahyan,isthanked forhissupportfortheresearchintotheMagharetJebel Figure1.16SrRNAfragmentsequenceof Lepidosporamazyadi n.sp. T HEFIRSTCAVERNICOLOUS N ICOLETIIDAE (I NSECTA :Z YGENTOMA ) FROMTHE U NITED A RAB E MIRATES 122 N JournalofCaveandKarstStudies, August2013

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Figure2. Lepidosporamazyadi n.sp.Scalesandmicrochaetaepartiallyshown.A,Body.B,Headandantennae.C,Ventral viewofbasalportionofantennae.D,Labium.E,Maxilla.F,Apexofmaxilla.G,Thoracicnota.H,Mandible. L.E SPINASAAND L.F.M ENDES JournalofCaveandKarstStudies, August2013 N 123

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Figure3. Lepidosporamazyadi n.sp.Scalesandmicrochaetaepartiallyshown.A,Hindleg.B,UrosternaIandII.C,Genital areaofmale.D,Genitalareaoffemale.E,Caudalappendages.F,DorsalviewofurotergaVIII-X(ventralpegsrepresented withdotsononesingleside).G,VentralviewofurotegiteX. T HEFIRSTCAVERNICOLOUS N ICOLETIIDAE (I NSECTA :Z YGENTOMA ) FROMTHE U NITED A RAB E MIRATES 124 N JournalofCaveandKarstStudies, August2013

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Hafeetcavefromwhichthespecimenwasidentified.H.E. MubarakSa’adal-Ahbabi,ChairmanoftheDepartment ofThePresident’sAffairs,NickCochrane-Dyet,andPeter Hellyerforfacilitatingthestudyofthenicoletiidspecimens.PamandTimFoggwerebothcollectorsandour liaisonforsourcesofinformation.DNAwassequencedby studentsofthegeneticscourse(BIOL320)atMarist College.WethankGraemeSmithandJulianJ.Lewisfor reviewingthemanuscript. R EFERENCES Espinasa,L.,andGiribet,G.,2009,Livinginthedark—species delimitationbasedoncombinedmolecularandmorphological evidenceinthenicoletiidgenus Texoreddellia Wygodzinsky,1973 (Hexapoda:Zygentoma:Nicoletiidae)inTexasandMexico, in Colkendolpher,J.C.,andReddell,J.R.,eds.,StudiesontheCave andEndogeanFaunaofNorthAmericaV,Austin,TexasMemorial MuseumSpeleologicalMonographs7,p.87–110. Mendes,L.F.,1988,SurdeuxnouvellesNicoletiidae(Zygentoma) cavernicolesdeGre `ceetdeTurquieetremarquessurlasyste matique delafamille:RevueSuissedeZoologie,v.95,p.751–772. Mendes,L.F.,2002a,Onanew Lepidospora s.s.(Zygentoma:Nicoletiidae)fromOman:RevistaBrasileiradeZoologia,v.24,no.1/2, p.107–112. Mendes,L.F.,2002b,Somenewdataanddescriptionsofthysanurans (Zygentoma:Nicoletiidae)fromCentralandEasternAfrica:Annales duMuse eroyaldel’AfriqueCentrale,Zoologie,v.290,p.87–127. Mendes,L.F.,2004,Zygentoma(Insecta)fromtheSocotraArchipelago: FaunaofArabia,v.20,p.357–398. Wygodzinsky,P.W.,1980,AsurveyoftheNicoletiinaeofEurope (Nicoletiidae,Thysanura,Insecta),AmericanMuseumNovitates no.2695,24p. L.E SPINASAAND L.F.M ENDES JournalofCaveandKarstStudies, August2013 N 125

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THEVIEWOFMAYACAVERITUALFROMTHE OVERLOOKROCKSHELTER,CAVESBRANCHRIVER VALLEY,CENTRALBELIZE G ABRIEL D.W ROBEL 1 ,R EBECCA S HELTON 2 ,S HAWN M ORTON 3 ,J OSHUA L YNCH 4 AND C HRISTOPHER A NDRES 5 Abstract: ArchaeologicalinvestigationsoftheOverlookRockshelterintheCaves BranchRiverValleyofcentralBelizeofferauniqueviewofancientMayacaveritual throughthecompleterecoveryandanalysisofallartifactswithinthesite’stwosmall activityareas.Ingeneral,theassemblagecontainsmanyofthesametypesofobjects documentedfromothernearbycavesandrockshelters.However,thenearly1700 ceramicssherdsshowedalmostnorefits,demonstratingthatsherdsweredepositedatthe siteindividually,ratherthanascompletevessels.Thehumanboneassemblagerepresents threeorfourindividuals,withthemajorityofthebonescomprisingasingleindividual, andalloftheseweredepositedasincompletesecondaryinterments.Analogiesforthis depositionalbehaviorbasedonarchaeologicalandethnographicstudiessuggestthatthis rocksheltermayrepresentawaypointwithinaritualcircuitcomposedofmultiple locationsoverwhichfragmentsofcompleteitemssuchasceramicvesselsandsecondary burialswerespread. I NTRODUCTION Mayacavearchaeologyhasgainedmuchmomentum withinthelasttwodecades,supplementingtraditional archaeologicalapproachesbyexpandingdiscussionsofthe rolesofideologyandritualwithinMayasociopolitical structure.Whileethnographicandethnohistoricstudies linkcaves,rockshelters,andsinkholestorainandfertility rituals,iconographicandarchaeologicaldatafromprecontactcontextsoftendemonstratethefurtheruseofthese naturalfeaturesinritesofpoliticalaccession,aggrandizement,legitimization,andsocialincorporation(BassieSweet,1996;PruferandBrady,2005;VogtandStuart, 2005).Thus,animportantaspectofMayaspeleoarchaeologyisthecontextualizationofcaveuse,whichhasbeen accomplishedbyidentifyingthenatureandtimingofritual activitiesperformedincavesthroughtheanalysisand interpretationofartifactassemblages.Thecurrentstudy focusesonasingle,smallrocksheltersite,theOverlook Rockshelter(OVR),thatwasinvestigatedasonecomponentofabroadregionalcavesurveycurrentlybeing conductedbytheCentralBelizeArchaeologicalSurvey project. Increasingly,cavestudiesintheMayaareahavetaken theformofbroadregionalsurveysthathavesoughtto articulatedatafromavarietyofsubterraneansitesinto moretraditionalarchaeologicalresearchdesigns,actively incorporatingdatafromsitesofdifferentsize,morphology, andproximitytootherarchaeologicalcontexts(Awe,1998; BonorVillarejo,1987;Bradyetal.,1997;Hardy,2009; Peterson,2006;Prufer,2002;Rissolo,2003;Wrobeletal., 2009).Incontrasttothegeneralizedpictureofancient Mayacaveusepreviouslyavailable,whichwasgenerally basedonsinglesites,theseregionalapproachesare designedtopaintaholisticpictureoftherangeofcave ritualspracticedbyspecificgroupslivinginaparticular area.Furthermore,suchstudiescanhelpilluminate broaderpatternsofcaveusethatarenotnecessarily restrictedtoritualsperformedinasinglelocale.Such patternswouldbeinvisibleincaseswhereresearchers treatedcavecontextsasisolatedandboundedritual contexts.Thus,thedatafromOVRcanbecomparedto andcontrastedwithotherrelatedsites,providingcontextualizationforidentifiedritualbehaviors.Furthermore, ethnographicanalogyisoftenutilizedasameansoflinking artifactassemblagestospecifictypesofrituals,aswellas thenatureoftheideologicalsignificancecavesheldwithin Mayabeliefsystems(seeBrady,1989,andBradyand Prufer,2005forextensivereviewsandexamples).Asan example,ourdiscussionofOVRasacavestemsfrom ethnographicandethnolinguisticstudiessuggestingthat rockshelterswereconceptualizedandappropriatedby Mesoamericanpeoplesascaves(seeRisollo,2005,p.354– 356).However,ethnographicanalogydoesnotprovideall theanswers,andasRisollo(2005,p.354)pointsout, ‘‘whilesemanticanalysesamongthemodernMayaare importantinsteeringthedirectionofresearch,itremains forarchaeologiststodocumentthepalpableevidenceof undergroundutilizationinordertodefineboundariesof theancientMayaconceptof‘cave’.’’ 1 DepartmentofAnthropology,MichiganStateUniversity,EastLansing,MI48824, USA,wrobelg@msu.edu 2 ARConsultants,Inc.,Dallas,TX 3 DepartmentofArchaeology,UniversityofCalgary,Alberta,Canada 4 DepartmentofAnthropology,TexasA&MUniversity,CollegeStation,TX778434352,USA 5 DepartmentofSociologyandAnthropology,UniversityofMississippi,Oxford, MS38677-1848,USA G.D.Wrobel,R.Shelton,S.Morton,J.Lynch,andC.Andres–TheviewofMayacaveritualfromtheOverlookRockshelter,Caves BranchRiverValley,CentralBelize. JournalofCaveandKarstStudies, v.75,no.2,p.126–135.DOI:10.4311/2011AN0233 126 N JournalofCaveandKarstStudies, August2013

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OverlookRockshelterwastargetedforexcavationin 2009becauseitpossessedarelativelyuniquecombination ofattributesnotfoundatothersitesinoursurvey.OVRis asmalloverhangwithnoassociateddarkzoneandlocated onasheerlimestoneclifffacehiddenbyvegetationhigh abovetheCavesBranchValley(Fig.1).Surfacecollection hadrevealedanassemblageconsistingprimarilyofheavily fragmentedceramics.Inparticular,theunexpectedpresenceofhumanremainsatsuchasmallandisolatedsitewas intriguing;rockshelterburialshavebeennotedatother nearbysites,suchasCavesBranchRockshelter(Glassman andBonor,2005;Wrobeletal.,2007),SapodillaRockshelter(WrobelandShelton,2011),ActunNakBeh (Halperin,2005),andUayazbaKab(Gibbs,2000),but thesesitespossessgenerallylargeractivityareaswithan associateddark-zonecave(Fig.2).Furthermore,unlikethe OVR,theseotherexamplesareeasilyaccessibleandseem tobedirectlyassociatedwithnearbysettlement.Becauseof thesmallsizeoftheactivityareasandthefactthatthissite isnotactivelyusedfortourism,wedecidedtoexcavatethe culturalfeaturescompletely,collectingallartifacts.While taphonomicprocesseshavelikelyplayedtheirpart,wefeel Figure1.Elmer‘‘Neko’’MedranoattheOverlookRockshelter.Viewtowardthenorth. Figure2.Mapofstudyarea,showingnearbysites. G.D.W ROBEL ,R.S HELTON ,S.M ORTON ,J.L YNCH AND C.A NDRES JournalofCaveandKarstStudies, August2013 N 127

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reasonablyconfidentinstatingthatthevastmajorityofthe artifactsdepositedinthepastremainedincontext.There arenopreviousattemptstocollectandstudyacomplete artifactassemblagefromarockshelter,andforthisreason thisstudyrepresentsauniqueandinvaluablesourceof datawithwhichtointerprettheritualactivitiesperformed atsuchsites. D ESCRIPTIONOFTHE S ITEANDTHE E XCAVATIONS MembersoftheBelizeValleyArchaeologicalReconnaissanceprojectandguidesfromIanAnderson’sCaves BranchAdventureCompanyandJungleLodgefirst reportedOverlookRockshelterafterabriefreconnaissanceinthesummerof2008.Thegroundsurfaceis uneven,withonlytwodiscreteareasthatcontained scatteredartifactsandhumanbone.Inbothareas,bones andsherdshadbeenstacked(probablyrecently)onsome oftheloosestonesonthesurface(Fig.3).Ceramicsherds accountedformostoftheartifactsvisibleonthesurface, thoughfreshwater jute shellsandseveralrivercobbles werealsonoted.Typicalofmostothercaveandrocksheltersitesinthearea,the jute hadbroken,orspirelopped,tipstofacilitateconsumptionoftheanimalinside (seeHalperinetal.,2003fordiscussionoftheritual significanceof jute amongtheMaya).Therestofthe surfacedoesnotcontainsoiloranytracesofhuman activityandiscomposedofflowstoneandlargeboulders ofbreakdownfromtheclifffaceabove.Therockshelteris orientedroughlynorthtosouth,facingeastwitha spectacularviewofthevalleyfloor.Bylateafternoon thesunclearstheoverhangingroofandtheentire rockshelterisexposedtodirectsunlight.Thewallofthe OVRcurvesslightlyoutwardtowardthenorthernmost endoftheshelter.Ashortclimbupthelimestonewallof theelevatednortherncorneroftheOVRrevealsanother smallrockshelter;ithasnoartifactsvisibleonthesurface. Excavationunitsweresetupintheonlytwoareaswith noticeableculturalmaterial(Fig.4).Theseareaswerethe onlyareaswithintherockshelterthatcontainedloosesoil, whileallothersurfaceswerecomposedeitherofsolidor erodinglimestone.Furthermore,theboundariesofboth werenaturallylinedbylargestonesandthuswererelatively easytodetermine.Thesoilwithintheseareasmatchedthe matrixfoundinotherrocksheltersinthesurrounding area—loose,dark,andcomposedmainlyoflimestone breakdownfromthecliffwalls(Hardy,2009,p.62).Allof thisloosesoilfrombothareascomprisedamatrix containingceramicsherds,aswellasafewotherpieces ofmaterialcultural,thatweremixedthroughoutit.The surfaceoftheseareascontainedrecentbreakdownfromthe rockshelterceilingintheformofseverallargerocksthat wereremovedpriortoexcavation.Allvisiblescatteredor stackedartifactsandboneswerecollectedasasingle surfacecontext.Basedonpriorexperiencewithrockshelters,wedidnotanticipatefindingclearstratigraphy,so arbitrary20cmlevelswereused. AreaAislocatedinthesouthernmostportionofthe rockshelterandwasdefinedbyrockcollapse(northand Figure3.AreaA,showingrecentlypiledbonesandartifacts totheleft,andtheareaofdisturbancetotheright. Figure4.TopographicmapoftheOverlookRockshelter. T HEVIEWOF M AYACAVERITUALFROMTHE O VERLOOK R OCKSHELTER ,C AVES B RANCH R IVER V ALLEY ,C ENTRAL B ELIZE 128 N JournalofCaveandKarstStudies, August2013

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south),thenaturalcavewall(west),andacrudely constructedretainingwall(east).Itmeasured202cm(EW)by384cm(N-S)atitslargest.Therewasanarrow trench,initiallyhypothesizedtobeasmalllooterÂ’spit,that uponfurtherinvestigationturnedouttobeanareaof naturalcollapseintoanarrowfissurebeneaththesoil. AreaBwastheothermainfocusofexcavationefforts.It measured220cm(E-W)by302cm(N-S)atitslargestand wasseparatedfromAreaAbyaloosepileofcollapsed rock.Excavationswithinbothareas,asexpected,didnot revealanyclearstratigraphy(i.e.,thesoilmatrixwas relativelyuniformincolorandartifactdensitythroughout) andsterilesoilwasreachedatamaximumdepthof approximately60cmwithinmostofbothunits.Artifacts foundintheexcavationsfrombothareasincluded numerouspotterysherds, jute shells,andafewpiecesof obsidian.AreaAcontainedasmall,drilledslatediskanda carvedmarineshellbead,aswellasahigherconcentration ofhumanbone(Fig.5).AreaBcontainedafish-netweight andarelativelyhigherconcentrationof jute shells. Asmallflatareaofgroundthatrunsfromthebaseof theeasternstonewallboundaryofAreaAtotheedgeof therockshelteralsocontainedafewartifacts,including smallceramicsherdsandafewfaunalremains.These remainswerecollectedasAreaC.Theshallowsoilwas verydense,anditisverylikelythattheculturalremains spilledoverfromAreaA.Also,someofthesoiland artifactsfromAreaBwerewashingoutbetweenthelarge bouldersdefiningtheeasternedgeofthecontext.These artifacts,whichincludedsomeofthelargestsherdsfrom theassemblage,werecollectedasAreaD,thoughthese clearlyoriginatefromAreaB. O STEOLOGICAL A NALYSIS Theinventoryoftheskeletalremainsshowsthatmost boneslikelycomefromasingleincompleteindividual, basedonsimilartaphonomiccharacteristics,thegenerally consistentsizeandrobusticityofelements,andinsome casescorrespondingarticularsurfacesofadjacentbones(in theleftarmandthefeet).However,themajorityofthe skeletonisabsent,andthesmallsizeofthefissureinArea Aisnotsufficienttoexplainthemissingelements.Mostof thebonesarecompleteand,apartfromtheribs,no fragmentswerefound.Whereavailable,sexandage indicatorsconsistentlyindicatethattheprimaryindividual isanolderadultfemale.Theageisbasedonthepresenceof osteophytesvisibleonthelumbarvertebraandondiscrete patchesofarthritisnotedontheglenoidsurfaceandonthe pelvisaroundtheauricularsurface.Inaddition,the individualdisplaysapronouncedpreauricularsulcusand pubicpitting,whicharetypicallyinterpretedasstress resultingfromchildbirth. Eachsegmentofthespinalcolumnisrepresentedbyat leastonevertebra.Fewoftheseappeartobeconsecutive, however,andthusthevertebralbonesincludedinthe assemblagewerelikelyselectedindividuallyratherthanas segments(i.e.,upperorlowerbody).Thepelvisincludes smallfragmentsofthesacrumandsegmentsoftheleftos coxae,includingtheischium,pubis,acetabulum,anda Figure5.Photographof(a)slatefragmentandnetweight, (b)drilledslatefragment,and(c)shellbeadfromLots4 and5,OverlookRockshelter,Belize. G.D.W ROBEL ,R.S HELTON ,S.M ORTON ,J.L YNCH AND C.A NDRES JournalofCaveandKarstStudies, August2013 N 129

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portionoftheleftauricularsurface.Approximatelyhalf oftheribswerepresent,thoughmanyofthesehad fragmented.Themostconsistentlyrepresentedportionof theskeletonistheleftarm,whichincludestheclavicle, humerus,ulna,radius,andmuchofthescapula.Theleft handisrepresentedbyallfivemetacarpals,aswellasthe lunate,capitate,andhamate.Therighthandhasonly metacarpals1,3,and4,thoughfivephalangeswere identifiedthatmaybelongtoeitherhand.Footbones includeseveraltarsalsfromeachfoot(left 5 navicular, talus,andcalcaneus;right 5 navicular,talus,and3rd cuneiform)andonlyafragmentofasinglemetatarsal.Toe phalangespresentincludedallproximalphalangesexcept thefirstleft,nomiddlephalanges,andonlyasingledistal phalangeoftherightfirsttoe.Nofragmentsoftheleg boneswerepresent,thoughbothpatellaswerefound.An AMSanalysis(Beta-279539;1640 6 40 BP )ofaribgavea calibratedtwo-sigmarangeof AD 330–540,placingthe individualsecurelywithintheEarlyClassic(AD300–600) period. Withintheboneassemblagearealsoatleastafew randomelementsfromotherindividuals,includingan unfusedcoracoidprocessfromajuvenile’sscapula.Also, theonlyfourteethfoundatthesite,allofwhichwere permanent,representatleasttwodifferentindividuals basedondifferencesinattrition.Anupperleftfirst premolarhadanearlycompleteroot,suggestingan approximateagearound10to12years.Alowerleftfirst premolarhadacompleteroot,butdisplayedasimilar amountofwear,andthuscouldrepresentthesame individual.Theremainingtwoteeth,alowerrightlateral incisorandcanine,werefromaslightlyolderindividual anddisplayedminimalamountsofattritionandsimilarly polishedsurfaces.Sincenocranialelementswerefound, andallteethseemtocomefromrelativelyyoung individuals,itislikelythatnonebelongtotheprimary individual.Wehavecalculatedaminimumnumberofthree individualsonthebasisofthedifferentagesrepresentedby theteethandbones,thoughitiscertainlyconceivablethat someoftheteethorsmallerbonefragmentsbelongto additionalindividuals. C ERAMIC A NALYSIS Excavationsrevealedapproximately1700ceramic sherds,whichdemonstrateddiagnosticattributesspanning theLatePreclassic(300BCtoAD250)throughtheLate Classic(AD600–900)periods;identificationwasbasedon thetype-varietymethodofGifford,1976.Therelatively smallassemblagecontainedawiderangeoftypeswithin fourwares:UaxactunUnslipped,PetenGloss,PineRidge Carbonate,andPasoCaballoWaxyWare.TheLate PreclassicdiagnosticspecimenswerelimitedtotwoSierra RedrimsherdsandpossiblyoneSapoteStriated: Unspecified(redrim)variety.EarlyClassicsherdsincluded ringbasesofunspecifiedtypesandoneActuncanOrange Polychrome:ActuncanVarietyrimsherd.Inaddition,a numberofdecoratedbodysherdsappeartobeofthe Protoclassic–EarlyClassicCocayApplique dtype,whichis alsoknownasTziminApplique d(Reents,1980,p.168– 186,60).TheLateClassicisrepresentedbyseveralDolphin HeadRed:DolphinVarietyrimsherds,whichdatetothe earlyfacetSpanishLookoutphase.Over86%ofthesherds wereunslipped,with30%ofthesebeingstriated.Lessthan 10%wereslipped,andonlythreepolychromesherdswere present.Therewerenoapparentdifferencesintheceramics betweenthesub-assemblagesfromAreasAandB. Furthermore,comparisonswithintheunitsdidnotreveal expectedtemporaldifferencesintheceramicsfoundinthe upperandlowerlevelsofthedeposits.Thishasimplicationsforthesite’sformationprocesses.Mixingofsherds throughoutverticallevelswasalsonotedatothernearby rockshelters(Hardy,2009;Wrobeletal.,2007).AtCaves BranchRockshelter,mixingwasexplainedbytheconstant intrusionofnewburials.Asthisbehaviorisnotevidentat OverlookRockshelter,itispossiblethatthedepositional patternobservedistheresultofextensivebioturbationin therockshelter’sunconsolidatedmatrix(seeHaynesand Agogino,1986).Alternativeculturalhypothesesinclude thatthisdepositionalpatternreflectsanasyetunidentified culturalprocessthatwouldhaveactedtochurnthematrix orthatthesesitesmayhavebeenusedoverarelatively protractedperiodintheLateClassic,inwhichcase depositswouldincludeoldsherdsmovedfromother locationsonthelandscape.Inreferencetothissecond possibility,JoelPalka(personalcommunication,March 2012)hasnotedthatamongthemodernLacandonMaya atMensabak,Chiapas,sherdsandotherancientobjects thatarefoundinfieldsarethoughttobeassociatedwith thepastorwithgodsorancestorsandarepickedupand redepositedwithinrockshelters.Thispossibilityisespeciallyintriguingbecauseitwouldalsoexplainthelackof stratigraphyinthedepositsatOVR,anditthusdeserves furtherconsiderationinfuturestudies. Overall,thebroadtimescaleandthetypesofvessel formsrepresentedatOverlookRockshelterfitrelatively wellwithinestablishedregionalpatternsforrockshelters andcaves(Aweetal.,1998;Grahametal.,1980;Hardy, 2009;Helmke,2009;Reents,1980).Asexpected,OVR(like otherrockshelters)doesnotshowthefullvarietyof ceramicformsoftennotedindark-zonecaves,which,in additiontoutilitarianvessels,oftenincludevases,shoeshapedollas,censordrums,ocarinas,comales,anddisksor lids(Helmke,2009,p.494–95;Reents,1980).Similarlyto bothcavesandotherrockshelters,OVR’sassemblagehasa relativelylargepercentageofsherdsfromunslippedand striatedjars(ollas)andbowls.Theformsoftheplainware vesselsfromwhichthesherdsoriginateareconsistentwith thosetypicallyidentifiedasutilitarianwhentheyarefound inhouseholdorothernon-cavecontexts.Unfortunately, wecannotmakespecificinferencesaboutwhichofthe manypossibleritualordomesticactivitiesthesevessels T HEVIEWOF M AYACAVERITUALFROMTHE O VERLOOK R OCKSHELTER ,C AVES B RANCH R IVER V ALLEY ,C ENTRAL B ELIZE 130 N JournalofCaveandKarstStudies, August2013

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wereactuallyusedinpriortotheirfragmentationandthe subsequentdepositionoftheirpieces.However,the consistencyofOVR’sassemblagedoesappeartobe somewhatdistinctfromothernearbydocumentedrocksheltersthatgenerallycontainaverylargepercentageof thesameplainwares,butalsotendtoincludeindividual sherdsfromamuchwidervarietyofspecializedvessel forms,includingpolychromedishes,columnorpyriform vases,miniatures,ceramicspindlewhorls,andnetweights (Hardy,2009).Additionally,otherassemblagesoften containalargervarietyofdecoratedsherdsfromspecializedvesselssuchasTerminalClassicmodel-carvedsherds andDaylight-Darknightvessels(Hardy,2009;Wrobeland Shelton,2011).Thus,itappearsthatthisaspectof depositionalbehaviortypicalofcaveandrockshelterritual wasabsentatOVR. Theceramicanalysisimpliessomespecificdepositional behaviors,whichalsocanbeusedtoinformreconstructionsofritualuseoftherockshelter.Perhapsthemost significantdiscoveryduringtheanalysisoftheceramics wasthattheassemblagecontainedveryfewrefitsbetween sherds,withnoevidenceforthepresenceofpartialor completevessels.Only35ofthenearly1700sherdswere rimsherdsrepresentingover30vesselswithnorefits.These vesselsrepresentavarietyofforms,includingjars,dishes, andbowls.DespitethenumberofPetenGlossbodysherds, therewerenorimsherdsthatrefittotheincisedPeten Glossnecksherdsintheassemblage.Theapplique decorationsonthelimitednumberofdecoratedbody sherdsaretypicallyfoundaroundthecircumferenceof CocayApplique djars,yetiftherewerewholevesselsinthe assemblage,therewouldhavebeenamuchlargernumber ofdecoratedsherds.Basedontheoverallsmallsizeofthe sherds,theirerodedcondition,andthelackofrefits,it appearsthattheceramicsdepositedatOVRwereleftas individualsherdsinsteadofwholevessels. InherregionalrockshelterstudyoftheCavesBranch RiverValley,Hardy(2009)identifieddistinctlydifferent depositionalpatternsofceramics.Forinstance,thesixteen wholevesselsfoundattheCavesBranchRockshelter (CBR),allofwhichdatedtoaroundtheProtoclassic period,appearedtohavebeeninterredintactwithburials. However,thebroadertemporaldiversityandvariationin vesseltypesandthegenerallackofrefitsofthesherds foundthroughoutthematrixattheCBRandtheother rocksheltersinthestudysuggestedtoHardy(2009,p.103) thattheywereinsteaddepositedasfragments.Because thesesiteswereonlysampledusingtest-pits,thelackof refitsinhersamplecouldpossiblybeexplainedbyritual behaviorinwhichcompletevesselsweresmashedandthen spreadacrossanentiresinglesite.However,thedatafrom OVRsuggestotherwise.IntheneighboringSibunRiver ValleyandtheMayaMountains,depositionofcomplete vesselsleftwholeorrituallykilled/smashedandofscatters ofindividualsherds,havebeenrecorded(Peterson,2006, p.57,64,126;Prufer,2002).AtActunTohinQuintana Roo,Rissolo(2003,p.47–54)notesthatitappearsasif ceramicswerebroughtintothecaveassherdsandwere scatteredalongthesurfacessoastomarkthemaspaths (seealsoMoyes,2005,p.281).Recentsalvageoperationsat theSapodillaRockshelterfoundsimilarpatternstoCaves BranchRockshelter,withwholevesselslikelyassociated withburials,whilethesurfaceandmatrixwerelitteredwith seeminglyrandomceramicsherds(WrobelandShelton, 2011). D ISCUSSION Asmentionedabove,therockshelterwasexcavated completely,andthustheanalysesoftheartifactsandbones areuniqueinthattheyarebasedonanearlycomplete assemblage,assumingthatonlyasmallpercentageofthe assemblagewashedoutovertheedgeofthesteepcliffface. Theimportanceofthisapproachliesinbeingabletorule outthepossibilitythatmissingelementsfromtheskeleton orfromceramicvesselswereinitiallydepositedatthesite butareburiedelsewhereinit.Whilelootingmayalso accountforsomemissingelementsoftheassemblage,we suspectthisisminimal,giventhegenerallackofsurface disturbance.Inourexperience,artifactsworthremoving foundlyingonthesurfaceofasitegenerateatleastenough enthusiasmbylooterstodigsmallpits,noneofwhichwere evidentatOVR. Twootherritualbehaviorsthatareoftendocumented ethnographicallymayalsoberesponsibleforthespecific compositionofceramicswithintheOverlookRockshelter assemblage.Thefirstrelatestothecollectionofrituallychargedobjects(Brown,2000;Scott,2009,p.91),aswellas relocationofobjectsascachesinotherrituallocations (Brady,1989,p.109).Inthiscase,itisconceivablethat individualsvisitingOVRremovedceramicsfrompilesof completesmashedvesselsaskeepsakesorforuseinrituals performedelsewhere.Thesecondritualbehavioris sweeping,whichhasbeendocumentedextensivelyin ethnographiccontexts(Tozzer,1941,p.151–2;Vogt, 1976,p.99;BrownandEmery,2008,p.317;Scott,2009, p.97)andhasbeenhypothesizedinanumberof archaeologicalsettingsatbothcave(Halperin,2005; Prufer,2002,p.620)andsurfacesites(Coggins,1987; Glassmanetal.,1995,p.60;MathewsandGarber,2004, p.52–53).Indeed,itispossiblethatthedepositsatOVR mayhaveaccumulatedastheresultofsweepingcleana nearbyactivityarea.Iftheritualarearequiringsweeping waslocatedwithinOVR,itisalsopossiblethatthe majorityofceramicsinitiallydepositedtherewereswept overtheedgeofthecliffandthusnotrecovered.While theseremainpossibilities,wesuggestthattheyareunlikely, sincebothoftheseprocesseswouldresultinarather randomassortmentofceramicsherds,andamongthe 1700,wewouldexpectthatifeachbeganasawholevessel thatatleastsomeofthosevesselswouldberepresentedby morethantwopieces.Carefulexaminationofthe G.D.W ROBEL ,R.S HELTON ,S.M ORTON ,J.L YNCH AND C.A NDRES JournalofCaveandKarstStudies, August2013 N 131

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assemblagedemonstratedthroughcomparisonsofvariableslikethickness,color,shape,decorations,inclusions (temper),andformthatalmostnoneofthepiecescould belongtothesamevessel.Itseemsunlikelythatpostdepositionalcollectionofobjectsorsweepingwouldresult insuchaconsistentpattern.Furthermore,periodic cleaningoveranextendedperiodoftime,presumablyat theinitiationofanewrite,hasbeendemonstrated elsewheretoproduceadistinctivedistributionalpattern (Helmkeetal.,inpress).Thispatternischaracterizedby markeddisparityinsherddensitybetweenactivityareas andtheperipheralareasofsecondarydepositionthatsuch ‘‘sweeping’’produces.Nosuchpatternwasobservedat OVR.Thus,weareconvincedthatthisassemblage representsanon-randomassortmentofsherds,andthat individualvesselsareonlyrepresentedbyoneortwo sherds,whichismoreconsistentwiththehypothesisthat sherdsweredepositedatOVRindividually. ThesinglepartialsecondaryburialfoundatOverlook Rockshelterisunusual,butdoeshavesomecorrelatesthat mayhelptoexplainitssignificance.Forinstance,thethree burialsdocumentedatActunNakBeharealsothoughtto besecondarypartialburials.Halperin(2005)suggeststhat theymaybetheremainsofeliteswhowereincorporated intoaspecifictypeofcaveritualintendedtolegitimize authorityandpoweroftheirrelatives.UnlikeOVR, however,therockshelterentranceofActunNakBehlikely wouldhaveservedasanareaforpublicritual,becauseitis aflatopenspace,itisdirectlyconnectedtothe monumentalcenterofCahalUitzNabyasacbe,and thereisasmallandrestrictivedarkzonethatcouldhave beenusedforprivateceremonies.Furthermore,theOVR artifactassemblageconsistedprimarilyofdensedepositsof individualceramicsherds,whileActunNakBehhadvery fewsherds,perhapsasaresultofritualsweepingor periodiclooting(Halperin,2005,p.80).So,whiletheform oftheburialsatActunNakBeh(i.e.,secondaryand partial)issimilartothatatOVR,thecontextsatthetwo sitesarequitedifferent,especiallyinregardtotherelative isolationanddifficultaccessofOVR,likelysuggestingthat theritualsperformedatthetwositesweredistinctive. ArecentreportonasolitarywalledburialfromActun Bats’ubinsouthernBelizealsoshowssomebasiccorrelates withOVR,aswellasasimilarEarlyClassicdate(Prufer andDunham,2009).Basedonitsisolatedlocationandthe likelypostmortemmanipulationofskeletalelements,the authorsarguethattheartifactswithinActunBats’ub reflectasinglespecializedritualspecifictotheindividual basedontheimportanceofhisorhersocialroleasa shaman.WhilethesolitarynatureoftheOVRburialmay alsobearguedtoimplyindividualizedtreatment,thecase fortheBats’ubburialisbasedonmanycontextualaspects thatarenotevidentintheOVRinterment.Bats’ubisa disturbedprimaryburial,itcontainselaborategrave goods,anditissealedwithinadark-zoneenvironment. Inotherwords,theBats’ubburialclearlyparallelsthe reverentialmortuarybehaviorcharacteristicoftombs, whilethescatteredandalmostrandombonesoftheOVR burialaretreatedinawaysimilartotherestoftheartifact assemblagewithinthegeneralmatrix.Furthermore,unlike atOVR,thediagnosticartifactsandradiocarbondates fromtheActunBats’ubarealltemporallyhomogenous, supportingtheargumentthatthisassemblagerepresentsa single,thoughpossiblyprotracted,ritualevent. TheinventoryandanalysisoftheremainsatOVR providessomecluesfordecipheringthemortuarypathway leadingtotheinterment.Whiletheleftarmisrelatively complete,thealmostrandomassortmentofotherelements, suchasnon-continuouselementsofthespinalcolumnand feet,forinstance,andthecompletelackofcutmarksrules outperi-mortemmutilationofthebodyasanexplanation foritsincompletestate.Clearly,elementsoftheinitial primaryburialwereremovedafterdecomposition.Furthermore,whilethepresenceorabsenceofsomeelements appearsrandom(e.g.,carpals),thesuspiciouslycomplete absenceofleglongbonesandtheskullseemsdeliberate. (SeeTiesler,2004foradiscussionofnaturalandcultural influencesondecompositionleadingtodifferentialpreservationandrepresentationofbonesinMayaburials.)What islesscleariswhethertheelementsoftheOVRinterment weremovedtotherockshelterasasecondaryburial,or whethertheOVRwasthesiteoftheprimaryburialandthe missingelementsweresubsequentlyremoved.(SeeFitzsimmons,1998foradescriptionofprolongedmortuary practicesamongtheClassicperiodMayaelite.)The completelackofarticulationsupportstheformerhypothesis,whilethepresenceofmanyofthesmallestbones wouldtentativelysupportthelatter.(SeePrufer,2002, p.613,whodiscusseshowassemblagescanbeaffectedby thebehaviorofritualspecialists,whopickuppreviously discardedobjectsforreuseinceremonies.)However,the extrabones(teethandinfantclavicle)certainlymusthave beenmovedfromelsewhereasindividualelements,orone wouldexpecttohavefoundmoreelementsofthose individuals. ThenatureoftheartifactassemblageatOverlook Rocksheltermayofferthebestanalogyforexplainingthe presenceofthescatteredbonesofapartialskeleton.As noted,fewofthesherdscouldbefittedtogether,suggesting thatalmostallcamefromdifferentvessels.Furthermore, thepresenceofdiagnosticformsfromaspanofnearlya millenniumiscertainlycurious,sinceitmayimplyrepeated andcontinuoususe.Whilethispatternhasbeendocumentedatanumberofsiteslikethelargerrocksheltersand cavesintheregion(Hardy,2009),itissurprisingtofind suchconsistentdevotiontosuchasmall,hard-to-reach, andgenerallyunimpressivesiteastheOVR.Itmaybe, however,thatthesinglecavewasnotthefocusofthe ritual.Severalresearchershavearguedthatsitesandeven specificareaswithinsitesweresometimesseenasconnected partswithinarituallandscapeandformedaseriesofway pointsalongaritualcircuit(Garzaetal.,2001,p.22; T HEVIEWOF M AYACAVERITUALFROMTHE O VERLOOK R OCKSHELTER ,C AVES B RANCH R IVER V ALLEY ,C ENTRAL B ELIZE 132 N JournalofCaveandKarstStudies, August2013

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Kenward,2005;Moyes,2005;Peterson,2006,p.120; Prufer,2002,p.639;seeAdamsandBrady,2005foran ethnographicdiscussionofritualcircuits).Ethnographically,thisbehavioriscommontonearubiquitousandcan beseenasanactinritualsofthehunt(Brown,2005),ofthe house,ofthefield,andofthehealingofthesick(Vogt, 1976,p.9–10,54–55,75),aswellasinritualsofsociopoliticalinvestment(Tedlock,1982;seealsoFreideletal., 1993,p.419)andboundarymaintenance(Tozzer,1941; Vogt,1969,p.391).Ateachofthesesites,individuals wouldleaveasmalloffering. Intheethnographicrecord,offeringsaretypically composedofwhatwouldbeconsideredwholeobjectsin amaterialsense.Thatis,ritualistsdepositwholeandintact arrangementsofcandles,plants,andotherparaphernalia (seeTedlock,1982;Vogt,1976,p.9–10,54,55).Inour archaeologicalhypothesis,thedifferenceisthusstriking asweproposethatthelackofanentirevesselinour assemblagemaysuggestthatitsindividualpieceswere dispersedacrossthelandscapeinsteadofbeingdeposited together.However,whilethematerialsignatureofthe assemblagevariesfromourethnographicmodel,itremains conceptuallyanalogous:Theritualfunctionsbecause offeringsandlocationsareinexorablyboundasoneobject opentomanipulation(asintheZinacantecohealingritual describedbyVogt,1976,p.9–10);inthisway,theritualis nolessthanthesumofitsparts,andindividualofferings arebutfragmentsofthisgreaterwhole. Thepaucityofrimsherdsandtheabsenceoflarger sherdsandpartialvessels,aswellastherathermundane natureofmostofthenon-ceramicofferings,mayreflectthe smallandmarginalnatureofthesitecomparedtoothers onthecircuit.Ifthisscenarioisaccurate,theincomplete skeletonmayrepresentananalogoussituationinwhichthe partsoftheindividualwerealsosimilarlydispersed.The largestandmostdistinctivebonesoftheprimaryindividual—thelegbonesandthecranium—mayhavebeen reservedforanothermoremeaningfulorsignificant locationalongthecircuit.Inothercontexts,craniaand longbonesareoftenmissingfromeliteburials,likelyasa resultofremovalforuseinvenerationrituals(McAnany, 1995,p.60–63).Finally,theEarlyClassicdateofthebones fallssecurelybetweentheearliest( AD1)andlatest ( AD800)ceramicdates.Whilethissuggeststhatthe intermentlikelydoesnotrepresentadedicationritual,the secondarynatureoftheburialleavesopenthepossibility thatitwasplacedaspartofaterminationritualdespitethe nearly400yearinterval(seeKunenetal.,2002). C ONCLUSIONS TheOverlookRockshelterwasexcavatedcompletely, resultinginthecollectionofacompleteartifactassemblage.AnEarlyClassicpartialsecondaryburialofanadult femalehadbeenplacedwithintherockshelter,andwas foundscatteredwithinthematrix.Theartifactassemblage wasgenerallytypicalofthosefromotherrockshelters excavatedinthearea.Diagnosticceramicsshoweda varietyofformssimilartothosereportedinothercave contextsandalsoshowedarangeofdatesspanningthe LatePreclassicthroughLateClassicperiods,whichwere mixedthroughoutalllevelsofthematrix.Thelackoffits betweenthesherdsshowedthatnowholevesselshadbeen placedintherockshelter,thoughitiscertainlypossiblethat lootersremovedwholevesselsleftonthesurface.However, thegeneralnatureofthedepositsuggeststhatthe assemblagewascreatedoveralongperiodoftimethrough therepeateddepositionofsinglesherds,andoccasionally othersmall,common,andinexpensive(thoughperhaps symbolicallymeaningful)objects,suchasrivercobbles, smallusedobsidianbladefragments,and jute shells.The documentationofthecompleteartifactassemblageatOVR hasimportantimplicationsfortheinterpretationofother cavecontexts.Mostcavesitescontainubiquitoussingle sherdsinisolationorwithinclustersthatareoften hypothesizedtobetheremnantsofcompletesmashed vesselswhosefragmentsweresubsequentlymovedbywater orhumanactivity.Instead,siteslikethesemayrepresent partofaritualcircuitcomposedofmanysuchsites,where anindividualleavessmallofferingsateach.Similarly,this patternmayexplaintheincompletenessoftheskeleton, which,likethevessels,couldhavebeenspreadacrossthe landscape.Futureresearchfocusedonansweringquestions relatedtothenatureofsuchdepositswouldbenefitfrom detailedcomparisonsofassemblagesatmultiplesites acrossthelandscape. A CKNOWLEDGEMENTS Wewouldliketothankthefollowingindividualsand institutionsfortheirsupportandassistance:JaimeAwe, JohnMorris,RafaelGuerra,andtheBelizeInstituteof Archaeology;DavidHayles,IanAnderson,andallthestaff attheCavesBranchAdventureLodge;theBelizeArchaeologicalResearchandEducationFoundation;theUniversityofMississippi’sBarksdaleHonorsCollegeandDepartmentofSociologyandAnthropology;themanystudents participatingonthe2009CBASsummerfieldschool; ChristopheHelmke,JessicaHardyHaley,CameronHowell, LeahJaworskyj,BruceMinkin,andAlexMinkinwhoalso providedtheFigure5photos.Thismanuscriptwas strengthenedconsiderablybycommentsandsuggestions fromJamesBrady,DominiqueRissolo,JoelPalka,andan anonymousreviewer—manythankstothem. R EFERENCES C ITED Adams,A.E.,andBrady,J.E.,2005,EthnographicnotesonMaya Q’eqchi’caverites:Implicationsforarchaeologicalinterpretation, in Brady,J.E.,andPrufer,K.M.,eds.,IntheMawoftheEarthMonster: MesoamericanRitualCaveUse:Austin,UniversityofTexasPress, p.301–327. G.D.W ROBEL ,R.S HELTON ,S.M ORTON ,J.L YNCH AND C.A NDRES JournalofCaveandKarstStudies, August2013 N 133

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Awe,J.J.,1998,WesternBelizeRegionalCaveProject:Objectives, context,andproblemorientation, in Awe,J.J.TheWesternBelize RegionalCaveProject,aReportofthe1997FieldSeason:Durham, UniversityofNewHampshire,DepartmentofAnthropologyOccasionalPaperNo.1,p.1–22. Awe,J.J.,Helmke,C.G.B.,andGriffith,C.S.,1998,Archaeological reconnaissanceintheRoaringCreekValley:Caves,rockshelters,and settlementarchitecture, in Awe,J.J.,ed.,TheWesternBelizeRegional CaveProject,aReportofthe1997FieldSeason:Durham,University ofNewHampshire,DepartmentofAnthropologyOccasionalPaper No.1,p.223–244. Bassie-Sweet,K.,1996,AttheEdgeoftheWorld:CavesandLateClassic MayaWorldView:Norman,UniversityofOklahomaPress,245p. BonorVillarejo,J.L.,1987,ExploracionesenlasgrutasdeCalcehtoky Oxkintok,Yucata n,Mexico :Mayab,v.3,p.24–32. Brady,J.E.,1989,AninvestigationofMayaritualcave-usewithspecial referencetoNajTunich,Peten,Guatemala[Ph.D.thesis]:Los Angeles,UniversityofCalifornia,478p. Brady,J.E.,andPrufer,K.M.,eds.,2005,IntheMawoftheEarth Monster:MesoamericanRitualCaveUse:Austin,UniversityofTexas Press,438p. Brady,J.E.,Scott,A.,Cobb,A.,Rodas,I.,Fogarty,J.,andUrquizu Sa nchez,M.,1997,GlimpsesofthedarksideofthePetexbatun project:ThePetexbatunregionalcavesurvey:AncientMesoamerica, v.8,p.353–364.doi:10.1017/S0956536100001784. Brown,L.A.,2000,Fromdiscardedtodivination:Demarcatingthesacred throughthecollectionandcurationofdiscardedobjects:Latin AmericanAntiquity,v.11,no.4,p.319–333. Brown,L.A.,2005,Plantingthebones:Huntingceremonialismat contemporaryandnineteenth-centuryshrinesintheGuatemalan Highlands:LatinAmericanAntiquity,v.16,no.2,p.131–146. Brown,L.A.,andEmery,K.F.,2008,Negotiationswiththeanimate forest:HuntingshrinesintheGuatemalanHighlands:Journalof ArchaeologicalMethodandTheory,v.15,p.300–337.doi:10.1007/ s10816-008-9055-7. Coggins,C.,1987,NewfireatChichenItza, in Memoriasdelprimer coloquiointernacionaldeMayistas,5–10Agosto,1985:Mexico, UniversidadNacionalAuto nomadeMe xico,p.427–484. Fitzsimmons,J.L.,1998,ClassicMayamortuaryanniversariesatPiedras Negras,Guatemala:AncientMesoamerica,v.9,no.2,p.271–278. doi:10.1017/S095653610000198X. Freidel,D.,Schele,L.,andParker,J.,1993,MayaCosmos:Three ThousandYearsontheShaman’sPath:NewYork,WilliamMorrow, 544p. Garza,S.,Brady,J.E.,andChristensen,C.,2001,BalamNaCave4: ImplicationsforunderstandingPreclassiccavemortuarypractices: CaliforniaAnthropologist,v.28,no.1,p.15–21. Gibbs,S.A.,2000,Aninterpretationofthesignificanceofhumanremains fromthecavesofthesouthernMayalowlands[M.A.thesis]: Peterborough,TrentUniversity,202p. Gifford,J.C.,1976,PrehistoricPotteryAnalysisandtheCeramicsof BartonRamieintheBelizeValley:Cambridge,HarvardUniversity, PeabodyMuseumMemoirsno.18,360p. Glassman,D.M.,andBonorVillarejo,J.L.,2005,Mortuarypracticesof theprehistoricMayafromCavesBranchRockshelter,Belize, in Prufer,K.M.,andBrady,J.E.,eds.,StoneHousesandEarthLords: MayaReligionintheCaveContext:Boulder,UniversityPressof Colorado,p.285–296. Glassman,D.M.,Conlon,J.M.,andGarber,J.F.,1995,Surveyandinitial excavationsatFloralPark, in TheBelizeValleyArchaeologyProject: Resultsofthe1994fieldseason:Ms.onfile,InstituteofArchaeology, Belmopan,Belize. Graham,E.,McNatt,L.,andGutchen,M.A.,1980,Excavationsat FootprintCave,CavesBranch,Belize:JournalofFieldArchaeology, v.7,no.2,p.153–172.doi:10.1179/009346980791505518. Halperin,C.T.,2005,SocialpowerandsacredspaceatActunNakBeh, in Prufer,K.M.,andBrady,J.E.,eds.,StoneHousesandEarthLords: MayaReligionintheCaveContext:Boulder,UniversityPressof Colorado,p.71–90. Halperin,C.T.,Garza,S.,Prufer,K.M.,andBrady,J.E.,2003,Cavesand ancientMayaritualuseof jute :LatinAmericanAntiquity,v.14, p.207–219. Hardy,J.L.,2009,Understandingfunctionalandsymbolicvariationin rocksheltersoftheCavesBranchRiverValleyofwesternBelize, CentralAmerica[M.A.thesis]:Oxford,UniversityofMississippi, 172p. Haynes,C.V.,Jr.,andAgogino,G.A.,1986,GeochronologyofSandia Cave:Washington,SmithsonianInstitutionPress,Smithsonian ContributionstoAnthropologyno.32,39p. Helmke,C.G.B.,2009,AncientMayacaveusageasattestedintheglyphic corpusoftheMayalowlandsandthecavesoftheRoaringCreek Valley,Belize[Ph.D.thesis]:London,UniversityCollege,701p. Helmke,C.,Awe,J.J.,Morton,S.G.,andIannone,G.,Thearchaeology andepigraphyoftheCuychenVase,MacalValley,Belize:Maya Archaeology,v.3[inpress]. Kenward,A.,2005,Showingtheway:Thefunctionofthreesmallcavesin theSibun-ManateeKarst, in Prufer,K.M.,andBrady,J.E.,eds., StoneHousesandEarthLords:MayaReligionintheCaveContext: Boulder,UniversityPressofColorado,p.249–259. Kunen,J.L.,Galindo,M.J.,andChase,E.,2002,Pitsandbones: IdentifyingMayaritualbehaviorinthearchaeologicalrecord:Ancient Mesoamerica,v.13,p.197–211.doi:10.1017/S0956536102132032. Mathews,J.P.,andGarber,J.F.,2004,Modelsofcosmicorder:Physical expressionofsacredspaceamongtheancientMaya:Ancient Mesoamerica,v.15,p.49–59.doi:10.1017/S0956536104151031. McAnany,P.A.,1995,LivingwiththeAncestors:KinshipandKingshipin AncientMayaSociety:Austin,UniversityofTexasPress,229p. Moyes,H.,2005,Clusterconcentrations,boundarymarkers,andritual pathways:AGISanalysisofartifactclusterpatternsatActun TunichilMuknal,Belize, in Brady,J.E.,andPrufer,K.M.,eds.,Inthe MawoftheEarthMonster:MesoamericanRitualCaveUse:Austin, UniversityofTexasPress,p.269–300. Peterson,P.,2006,AncientMayaRitualCaveUseintheSibunValley, Belize:Austin,AssociationforMexicanCaveStudies,Bulletin16, 148p. Prufer,K.M.,2002,Communities,caves,andritualspecialists:Astudyof sacredspacesintheMayaMountainsofsouthernBelize[Ph.D. thesis]:Carbondale,SouthernIllinoisUniversity,755p. Prufer,K.M.,andBrady,J.E.,2005,Introduction:Religionandtherole ofcavesinLowlandMayaarchaeology, in Prufer,K.M.,andBrady, J.E.,eds.,StoneHousesandEarthLords:MayaReligionintheCave Context:Boulder,UniversityPressofColorado,p.1–24. Prufer,K.M.,andDunham,P.S.,2009,Ashaman’sburialfromanEarly ClassiccaveintheMayaMountainso fBelize,CentralAmerica:World Archaeology,v.41,no.2,p.295–320.doi:10.1080/00438240902844236. Reents,D.,1980,TheprehistoricpotteryfromPetroglyphCave,Caves BranchValley,ElCayoDistrict,Belize,CentralAmerica[M.A. thesis]:Austin,UniversityofTexas,323p. Rissolo,D.,2003,AncientMayaCaveUseintheYalahauRegion, NorthernQuintanaRoo,Mexico:Austin,AssociationforMexican CaveStudies,Bulletin12,151p. Rissolo,D.,2005,BeneaththeYalahau:Emergingpatternsofancient MayaritualcaveusefromnorthernQuintanaRoo,Mexico, in Brady, J.E.,andPrufer,K.M.,eds.,IntheMawoftheEarthMonster: MesoamericanRitualCaveUse:Austin,UniversityofTexasPress, p.342–372. Scott,A.M.,2009,Communicatingwiththesacredearthscape:An ethnoarchaeologicalinvestigationofKaqchikelMayaceremoniesin highlandGuatemala[Ph.D.thesis]:Austin,UniversityofTexas, 227p. Tedlock,B.,1982,TimeandtheHighlandMaya:Albuquerque,University ofNewMexicoPress,309p. Tiesler,V.,2004,Mayamortuarytreatmentsoftheelite:Anosteotaphonomicperspective, in Gran aBehrens,D.,Grube,N.,Prager, C.M.,Sachse,F.,Teufel,S.,andWagner,E.,eds.,Continuity andChange.MayaReligiousPracticesinTemporalPerspective: 5thEuropeanMayaConference,UniversityofBonn,December 2000:Munich,VerlagAntonSaurwein,ActaMesoamericana14, p.143–156. Tozzer,A.M.,1941,Landa’sRelacio ndelasCosasdeYucata n:Atranslation: Cambridge,HarvardUniversity,PapersofthePeabodyMuseumof ArchaeologyandEthnology,vol.18,394p. Vogt,E.Z.,1969,Zinacantan:AMayaCommunityintheHighlands ofChiapas:Cambridge,BelknapPressofHarvardUniversity, 733p. Vogt,E.Z.,1976,TortillasfortheGods:ASymbolicAnalysisof ZinacantecoRituals:Cambridge,HarvardUniversityPress,234p. T HEVIEWOF M AYACAVERITUALFROMTHE O VERLOOK R OCKSHELTER ,C AVES B RANCH R IVER V ALLEY ,C ENTRAL B ELIZE 134 N JournalofCaveandKarstStudies, August2013

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Vogt,E.Z.,andStuart,D.,2005,Somenotesonritualcavesamongthe ancientandmodernMaya, in Brady,J.E.,andPrufer,K.M.,eds.,In theMawoftheEarthMonster:MesoamericanRitualCaveUse: Austin,UniversityofTexasPress,p.155–185. Wrobel,G.D.,Jordan,J.,andHardy,J.,2009,Socialandpolitical transformationsintheCavesBranchRiverValley:Evidencefrom naturalandconstructedritualenvironments:ResearchReportsin BelizeanArchaeology,v.6,p.199–207. Wrobel,G.D.,andShelton,R.,2011,Preliminarysalvageoperationsat SapodillaRockshelter, in Andres,C.R.,andWrobel,G.D.,eds.,Report ontheCavesBranchArchaeologicalSurveyProject,Summer2010 FieldSeason:Oxford,Mississippi,BelizeArchaeologicalResearchand EducationFoundationOccasionalReport # 2,p.18–40. Wrobel,G.D.,Tyler,J.,andHardy,J.,2007,Rockshelterexcavationsin theCavesBranchRiverValley:ResearchReportsinBelizean Archaeology,v.4,p.187–196. G.D.W ROBEL ,R.S HELTON ,S.M ORTON ,J.L YNCH AND C.A NDRES JournalofCaveandKarstStudies, August2013 N 135

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FLOWCHARACTERIZATIONINTHESANTEECAVE SYSTEMINTHECHAPELBRANCHCREEKWATERSHED, UPPERCOASTALPLAINOFSOUTHCAROLINA,USA A MY E.E DWARDS 1 ,D EVENDRA M.A MATYA 2 ,T HOMAS M.W ILLIAMS 3 ,D ANIEL R.H ITCHCOCK 3 AND A PRIL L.J AMES 4 Abstract: Karstwatershedspossessbothdiffuseandconduitflowandvaryingdegrees ofconnectivitybetweensurfaceandgroundwateroverspatialscalesthatresultin complexhydrologyandcontaminanttransportprocesses.Theflowregimeandsurfacegroundwaterconnectionmustbeproperlyidentifiedandcharacterizedtoimprove managementinkarstwatershedswithimpairedwaterbodies,suchastheChapelBranch Creek(CBC),SouthCarolinawatershed,whichhasalong-termsamplingstation presentlylistedonanEPA303(d)listforphosphorous,pH,andnitrogen.Waterfrom thecarbonatelimestoneaquiferoftheSanteeCavesystemandspringseepsintheCBC watershedweremonitoredtocharacterizedominantflowtypeandsurface-groundwater connectionbymeasuringdissolvedcalciumandmagnesium,totalsuspendedsolids, volatilesuspendedsolids,alkalinity,pH,specificconductance,andstableisotopes( d 18 O, d 2 H).ThesemeasurementsindicatedthattheconduitflowtoSanteeCavespringwas rechargedpredominantlyfromdiffuseflow,withaslowresponseofsurfacewater infiltrationtotheconduit.QualitativedyetracesandstageelevationatSanteeCave springandtheadjacentLakeMarion(equaltotheelevationofthefloodedportionof CBC)alsoindicatedarelationbetweenfluctuatingbaseleveloftheCBCreservoir-like embaymentandelevationoftheSanteeLimestonekarstaquiferatthespring.Methods describedhereintocharacterizetheflowtypeandsurface-groundwaterconnectioninthe SanteeCavesystemcanbeappliednotonlytowatershedmanagementintheChapel BranchCreekwatershed,butalsotothegreaterregionwherethiscarbonatelimestone aquiferexists. I NTRODUCTION Thisstudyusesmeasurementsofdissolvedcalciumand magnesium,totalsuspendedsolids,volatilesuspendedsolids, alkalinity,pH,specificconductance,stableisotopes( d 18 O, d 2 H),dyetracing,andstageelevationmeasurementsfrom boththeSanteeCavespringandLakeMariontocharacterizetheflowregimeandsurface-groundwaterconnectionin theSanteeLimestone(SL)aquiferintheChapelBranch Creek(CBC)watershedinSouthCarolina.Thehydrology andwaterqualityoftheCBCwatershedissignificantly impactedbyitskarstterrainandtheshallowcarbonate aquiferoftheregionalSLformation.Thekarstterrain providesapotentialforrapidandsubstantiallossofsurface watertothegroundwaterintheaquifer.Theanisotropic porosityofthecarbonatebedrock,createdbybedding planes,fractures,andongoingdissolutionwithinthematrix ofthecarbonaterock,producesgroundwatermovement fromzonesofrechargetodischargethroughanetworkof voidsofdiversesizesandvariousdegreesofverticaland horizontalconnectivity.Theresultisawatersheddominated bysubsurfaceflow,inwhichsurfacewateriscommonlylost tothesubsurface,andgroundwaterisstoredordischarged viaspringstoCBCatdifferentratesandflowregimes throughouttheaquifer(Williamsetal.,2012). Thetwomaintypesofflowappliedtokarstaquifersare diffuseandconduit(ShusterandWhite,1971;Atkinson, 1977;Gunn,1986),alsoreferredintheliteratureasslow andfastflow,respectively.Diffuseflowoccurswhenthe voidsintheaquifermatrixarepoorlyconnectedand typicallyresultinasystemwithslowgroundwater movementandlongerperiodsofwaterstorage(Fiorillo, 2009).Conduitflowresultswhenvoidsintheaquifer becomewelldevelopedandpossesshighconnectivity,thus havinglesswaterstorageandamorerapidresponseof rechargetodischarge(Fiorillo,2009).Atkinson(1977) concludedthatmostflowinkarstaquifersoccursin conduits,whereasmostwaterisstoredinthematrix.A studybyWorthingtonetal.(2000)analyzedspringand welldatafromlimestoneanddolostoneaquifersandfound that96%ofstorageoccurredinmatrix,while94%offlow occurredinconduits.TheSLunit,intheCBCwatershed,is 1 FloridaA&MUniversity,SchooloftheEnvironment,Tallahassee,FL,32301, USA,amy.edwards@famu.edu 2 USDAForestService,CenterforForestedWetlandsResearch,Cordesville,SC, 29434,USA,damatya@fs.fed.us 3 BelleW.BaruchInstituteofCoastalEcologyandForestScience,Clemson University,Georgetown,SC,29442,USA,tmwllms@clemson.edudhitchc@clemson. edu 4 DepartmentofGeography,NipissingUniversity,NorthBay,ON,P1B8L7, Canada,aprilj@nipissingu.ca A.E.Edwards,D.M.Amatya,T.M.Williams,D.R.Hitchcock,andA.L.James–FlowcharacterizationintheSanteeCavesysteminthe ChapelBranchCreekwatershed,uppercoastalplainofSouthCarolina,USA. JournalofCaveandKarstStudies, v.75,no.2,p.136– 145.DOI:10.4311/2011ES0262 136 N JournalofCaveandKarstStudies, August2013

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thenorthernportionoftheFloridanaquifer.Several studiesoftheFloridanaquiferincludingthesouthernpart oftheFloridanaquiferinFloridafoundspringdischarge asprimarilydiffuserechargefromaquifermatrix(Martin andGordon,2000;MartinandDean,2001;Katz,2004; FloreaandVacher,2006;Mooreetal.,2009;Ritortoetal., 2009).EveninconduitdominatedsystemsoftheFloridan aquifer,thematrixcontributesheavilytoaquiferdischarge (Mooreetal.,2009). Geochemicalandisotopicdatafromspringshavebeen usedinnumerousstudiesindifferentkarstregionsto characterizeflowincarbonateaquifersanddegreeof surface-groundwaterconnec tions(e.g.Jakucs,1959; Ashton,1966;ShusterandWhite,1971;Jacobsonand Langmuir,1974;Atkinson,1977).Thedischargefrom springsrepresentsacompositeofalltheupbasinwatersin anaquifer,whichshowvariationsinhydrologyandwater chemistrydependingoncontributionsfromdiffuseand conduitflowcomponents(White,2003).Springsdominatedbydiffuseflowrechar gehavefewervariationsin flowandwaterchemistry(ShusterandWhite,1971).The wateristypicallysaturatedwithdissolvedions(e.g. calciumandmagnesium),bec ausediffuseflowhaslonger residencetimeintheaquifer;andtherefore,longer contactwiththebedrock(White,1988).Incontrast, systemsdominatedbyconduitflowrespondmorequickly toaquiferrechargeandproducegreatervariationsin springwaterchemistry(Shus terandWhite,1971;White, 1988).Thismorerapidinfiltrationandtransitofrechargewatersalsoleavesconduitspringsunder-saturated (White,1998),andhighsuspendedsolidsatsprings indicatearrivalofsurfacewaterscomingfromsinkholes anddirectsurfacerecharge(MahlerandLynch,1999; Masseietal.,2002;VesperandWhite,2004).Stable isotopicanalysisiscommonlyemployedtoinferthe flowpathsandsourcesofspringwatersbyusingendmembermixinganalysis(Katzetal.,1997;Plummeretal., 1998;Crissetal.,2000). Dyetracingisanestablishedmethodusedtodetermine connectionsbetweenrechargeanddischargeareasinkarst aquifersandwasusedinthisstudytoobtaininformation abouttheresponseoftheSLaquiferattheSanteeCave systemduringtimesoflowandhighlakeelevations. Theflowtypeandsurface-groundwaterconnectionsin theSLcarbonateaquiferareclearlyimportantfactorsin watermovement;andthus,alsocontaminanttransportand waterqualityoftheCBCwatershed.Thegoalofthisstudy istocharacterizeflowinthewatershedusinghydrologic, waterchemistry,dyetracer,andstableisotopedata collectedintheSanteeCavesystemandnearbyspring seepsduring2008–2009.Thespecificobjectivesofthis studyaretotesttwohypotheses:(1)thedischargefromSL toCBCispredominantlyadiffuseflowcomponentfrom thematrixwithslowsurface-groundwaterconnectionand (2)theaquiferwatertableatSanteeCaveisinfluencedby stageelevationsofthelowerfloodedportionofCBC. S ITE D ESCRIPTION Theregion,southofLakeMarion,SCisunderlainby theSanteeLimestone(SL),amiddleEocene(40ma), microfossiliferouslimemudstonetoslightlyshellywackestone(Willoughby,2002).TheSLisblanketedbythe UpperDuplinFormation,acombinationofclayand quartzpebblebeds,withsinkhole,modernalluviumand swampdepositsthroughouttheregion(Willoughby,2002). TheSLispartofthenorthernendofthemulti-state Floridanaquifersystemandamajorgroundwateraquifer incoastalSouthCarolinaforindustrial,agricultural,and publicpurposes(Hockensmith,2009).RechargetoSLisby precipitationandsurfacerunoffthatinfiltratesintothe subsurfacevoidsviasoilandkarsttopography(e.g. sinkholes,disappearingstreams)anddischargesviasprings liketheSanteeCavespringinSantee,SCalongtheCBC andthesouthernshoreofLakeMarion.Tritiumand radiocarbondatingfromaspringnorthwestoftheCBC watershedandfromregionalwellsforaSCDepartmentof HealthandEnvironmentalControl(DHEC)rechargeand aquifervulnerabilitystudyfoundthatgroundwaterwasa combinationofyoungerwatersfromrechargealongthe southernshoreofLakeMarionandolderwatersfrom lateralflowfromaquiferupdipareas,suchasthecityof Orangeburg,SC(Stone,per.comm.,2009). ThesurfacewatershedforChapelBranchCreekis 1,555hectare(3,840acres)andcomprisesmultipleland uses.TheSanteeCavesystemliesintheforestedSantee StateParkinthelowerpartofthewatershed.Theupper watershedlandusesarecropagriculture,roadwaysandthe majorinterstateI-95,acommercialstripofftheI-95exit, lowandmedium-densityresidentialareas,twogolfcourses, andforest.ThesurfacesoilsintheCBCwatershedaresand orsandyloamandvaryfromsandtoclayinthesubsurface (SCS,1988).Theupperwatershedisrelativelyflatand around36.6m(120ft)abovemeansealevel(a.m.s.l.), whilethelowerwatershedattheedgesofCBCandLake Marionhavesteepertopographyofaround25.9m(85ft) a.m.s.l. TheCBCwatersheddrainsdirectlytoLakeMarion (Fig.1).LakeMarionandalowerportionoftheCBCare shallowreservoirscreatedwhentheSanteeRiverwas dammedbytheCorpsofEngineersforproductionof hydroelectricpowerin1941.TheCBCwasaspring-fed surfacestreambeforeconstructionofthedam,but afterwards,thelowersectionbecameapartoftheLake Marionreservoir.Astretchoftheuppermostsurfacereach ofCBCwasdammedin1989(ERC,1999)tocreateapond foragolfcoursethatreleaseswaterdownstreambyan overflowculvertatlocationSL7(Fig.1).Waterfromthe culvertflowsinachannelandremainssurfacewateruntilit becomesadisappearingstreamwhereitenterstheSantee CavesystematapointlabeledSS(surfacestream)in Figures1and2.Duringperiodsofheavyrainfall,the surfacewaterexceedsthecapacityoftheSSconduitanda A.E.E DWARDS ,D.M.A MATYA ,T.M.W ILLIAMS ,D.R.H ITCHCOCK AND A.L.J AMES JournalofCaveandKarstStudies, August2013 N 137

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Figure1.GeneralizedflowdirectionsforthestudysitesamplinglocationsintheChapelBranchCreekwatershed. F LOWCHARACTERIZATIONINTHE S ANTEE C AVESYSTEMINTHE C HAPEL B RANCH C REEKWATERSHED UPPERCOASTALPLAINOF S OUTH C AROLINA ,USA 138 N JournalofCaveandKarstStudies, August2013

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portionofflowcontinuesassurfacewaterviaaby-pass channel(seenasdashedbluelineinFigure1)tocombine withCS(SanteeCavespring)dischargeabout175m downstream.Theflowthenentersthelowersectionof CBC,ashallowwaterbodyshownin2009bathymetrydata tohaveanaveragewidthof200metersanddepthof 3meters(Amatyaetal.,2011). TheSanteeCavesystemisastreampassagecavewith approximately300metersofsurveyedpassage(Holler, 2000).TheflowatCSoccursaftertheconvergenceoftwo mainstreampassages,oneoriginatingatGWandtheother atSS(Figs.1and2).Basedonthecavesurveymap,the streampassagestartingattheSSlocationdrops3m (10feet)alongthepassagetoCS.TheGWpassageis steeper,dropping6m(20feet)toCS.Thesetwopassages combinebeforeemergingatCSat26m(85feet)MSL.The SSlocationwassampledwherethesurfacestream disappearsintothecave.TheGWlocationwassampled wherethegroundwaterconduitemergesatthebottomofa sinkhole.Theinaccessibleupstreamconduitpassage feedingGWmostlikelycontinuesinasoutheastern direction,inlinewitha2009sinkholecollapse,asinkhole pond,andtwoothersinkholesdetectedinLiDARimages (EdwardsandAmatya,2009;2010).Akarstwindow betweenGWandCSexposesthecavestreamandthecave systemlieswithinaforestednaturepreservemanagedby theSouthCarolinaDepartmentofNaturalResources. Severalsmallspringseeps(SPS)(Figs.1and2)around 30mmdiameterathigherelevationsthanCS,andnot connectedtotheSanteeCavesystem,werenotmonitored forflow,butweresampledinAugust2009fordissolved calciumandmagnesium,pH,alkalinity,andspecific conductanceforcomparisontoGW,CS,andSS.Several ofthehigherelevationseepsdriedupduringaperiodof lowprecipitationandlowlakelevelsin2009(Amatyaetal., 2011). AlthoughwaterisdischargedfromtheSLaquiferinto theCBCatnumeroussmallspringsalongtheshoreline,the onlyspringabovetheshorelinewithsubstantialflowisCS. Awaterbudgetusingrainfalldatafromthewatershedand flowdataatCSindicatedthatthesourceofCSdischarge isnotsolelyaccountedforbyrunoffcollectedbythe upstreamsurface-definedwatershed,butalsoincludes groundwatercontributionsfromalargersubsurface watershed(Williamsetal.,2012).Theestimatedbaseflow fortheregion,usingdatacollectedatCS,is0.09m 3 /s (90l/s).Usingthesedatawithareferencevalueof2.3L/ (s/km 2 )assumedfromthemethodbyQuinlanandRay (1995),theapproximatesubsurfacedrainageareaforCSis 39.1km 2 .Thiswaterbudget-basedcalculatedareais 3.5 timesthesurfacedrainageareaof10.9km 2 (1090ha) delineatedattheCSlocationintheCBCwatershedstudy. Thisresultwasplausiblegiventhatsurfaceandsubsurface watershedsoftendonotoverlapinkarstwatersheds(White, 1988).Thisconclusionwasfurthersubstantiatedbythe resultsfromhydrologiccomputermodelingfortheCBC watershedstudy,inwhichmodelingresultspredictedmore surfaceflowatCBCsubwatershedoutletmonitoring stationsthanmeasured(Amatyaetal.,2011).This ‘‘missing’’flowatthesubwatershedoutletswasassumed tobelostfromthesurfacetothegroundwateraquifer matrix,eventuallydischargingdownstreamintotheCBCvia CSorothersprings(Williamsetal.,2012).Alsoduringthe watershedstudy,flowatCSwasfoundtobeyielded predominantlybytheGWcontribution,withpeakflowsin thehydrographoccurringfromraineventsduetoinfluxof surfacewaterfromthedisappearingstreamatSS(Williams etal.,2012).Awaterbalanceapproachindicatedflowfrom theGWpassagetoCSwasnearlycontinuousat0.08m 3 /s; over60%oftheflow,nearly50%ofTPloading,andover 70%oftotalnitrogenloadingintotheCBCreservoir (Williamsetal.,2012). Thesehydrologicmeasurementsfromthewatershed studyindicateadominantsubsurfacesourceintheCS dischargeandlikelyotherspringsdischargingintothe CBC.Thenatureofthiscomplexwatershedhydrology, combinedwiththekarsttopographyandmultipleland uses,makethiswatershedsusceptibletowaterquality impairment.Along-termsamplingstationintheCBChas beenontheU.S.EPA303(d)listforimpairedwaterbodies forvariouscombinationsofparametersforyears,andis presentlylistedforphosphorous,pHandnitrogen.Past incidencesofpollutionincludeawastelagoonatasewage treatmentplantintheCBCsurfacewatershedthat disappearedovernightinthe1990s,andthefumigant EDBfoundindischargefromaspringoutsidetheCBC surfacewatershed(Stone,per.comm.,2009).Considering thispastandongoingwaterqualityimpairment,characterizingthecarbonateaquiferforflowtypeandsurfacegroundwaterconnectivityisrelevantforimprovingwater Figure2.ImagesofsamplinglocationsSS,GW,CS,and SPS,providingtypicalconduitpatternandrelativescale. Researchersprovidescaleforthesekarstfeatures. A.E.E DWARDS ,D.M.A MATYA ,T.M.W ILLIAMS ,D.R.H ITCHCOCK AND A.L.J AMES JournalofCaveandKarstStudies, August2013 N 139

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qualityintheCBCwatershedandtheregionsouthofLake Marion. S ITE M ONITORING DyeTraces Threequalitativedyetracesbasedonvisualinspection wereconductedintheSanteeCavesystem.Thefirsttwo traceswerecarriedoutonAugust28,2008.A16FL.OZ. bottleofBrightDyesLiquidConcentrateFluorescentRed wasinjectedintothedisappearingstreamatlocationSS, anda16FL.OZ.bottleofBrightDyesLiquidConcentrate FluorescentGreen/YellowwasinjectedatlocationGW. ArrivaltimefromGWandSSlocationstolocationatCS (Fig.1)andthepredictedflowpathaccordingtothecave surveywererecorded.Sinceavisibleamountoforganics entersthedisappearingstream,andreddyereactswith organicstodecompose,athirdtracewasconductedon April2,2009atlocationSSusinga16FL.OZ.bottleof BrightDyesLiquidConcentrateFluorescentGreen/Yellow dye. HydrologicData StagedataatCSwerecollectedusinganInfinities WaterLevelDataLoggerat15-minuteintervals.Proceduresfordetailedinstrumentcalibration,datacollection andcalculationsarefoundinAmatyaetal.(2010).The elevationoftheCSwaterlevelwasdeterminedbyadding thestageinfeetmeasuredforstagedischargerelationship tothecavesurveyelevationforcomparisonwithLake Marionelevation,whichwasusedasaproxyforCBC elevation.LakeMarionmaximumelevationdata(datum NGVD1927)wasobtainedfromUSGSgagingstation # 02169921atLakeMarionnearElloree,SC(USGS, 2009). WaterQualityandIsotopeSamplingandAnalysis Waterqualitymeasurementsweretakenonapredominantlymonthlybasisduringvariousflowconditions. MeasurementsforpHandspecificconductanceweretaken inthefieldusingOaktonpH/conductivity/ u C10series handheldmeters,whichwerecalibratedwithOakton buffers(4.01,7.00,10.0,84 m Sand447 m S)duringevery fieldvisit.ALaMotteAlkalinitykit,ModelDR-Awas usedinthefieldtotitrateforalkalinity.Grabsampleswere collectedusingprotocolsandcontainersdependentonthe desiredanalysisaccordingtoStandardMethods(Clesceri etal.,1998).Samplesfordissolvedcalciumandmagnesium concentrationswereanalyzedbytheSanteeCooper AnalyticalandBiologicalLaboratorylocatedinMoncks Corner,SC.AdetaileddescriptionofStandardOperating ProceduresisincludedintheTMDLQualityAssurance ProjectPlanandFinalDraftReport(Amatya etal,2010).SamplesforTotal(TSS)andVolatile(VSS) SuspendedSolidswereanalyzedbyprojectpersonnelat theUSDAForestServiceCenterforForestedWetlands ResearchusingStandardMethods2540DandE(1988). WaterchemistrydataonTSS,VSS,pH,specificconductance,andalkalinitywerestatisticallyanalyzedfortheir meansanddistributionsandplottedusingaprogramin MATLABsoftware(Mathworks,2009).TheTukey’stest forsignificantdifferenceinmultiplemeanconcentrations ofeachoftheparametersbetweenthelocationswasusedat a 5 0.05levelusingastandarderrorbarrepresenting95% confidenceintervals. Samplescollectedforstableisotope( d 18 O, d 2 H) analyseswerefilteredandstoredawayfromlightinglass vialswithnoheadspace.Analysiswasperformedwitha PicarroL2120-ihigh-precisionisotopicwateranalyzer, usingCavityRing-DownSpectroscopy(CRDS),inthe DepartmentofGeographyatNipissingUniversity,Ontario,Canada.Sampleprecision(maximumstandard deviationofallinjectedsamples)for d 18 Oand d Dwas # 0.12 % and # 0.26 % ,respectively.Accuracy(average deviationfromtruevalueor‘BigDelta’)for d 18 Oand d D was # 0.24 % and # 0.81 % ,respectively.Accuracywas evaluatedusingsecondarylabstandardsandIAEA standardVSMOW2.Samplesweretestedforchemical contaminationwithpotentialtoaffectCRDSresultsusing Picarro’sChemCorrect TM software. R ESULTS A.H YDROLOGICALAND D YE T RACE D ATA Theimpactoffluctuatinglakelevelsonthelimestone aquifer,asobservedatCS,isshowninFigure3.Thedaily maximumelevationofCShadanoveralltrendthat matchedtheriseandfallofLakeMariondailymaximum elevationin2009. Twoqualitativedyetracingeventswereconductedat SSduringlowandhighflowconditions.Approximate arrivalofdyeatCSwasobservedusingvisualinspection only.ThereddyeinjectedatSSonAugust28,2008was expected,butnotobserved,atCS,eventhoughdepthof waterattheSSlocationwas0.19m,withanaverage velocityof0.73m/s(2.4ft/sec).Itwashypothesizedthatthe organicspresentinthesurfacestreamatSScouldbe interferingwiththereddyetracer.Asecondtracerwas injectedusinggreendyeatSSonApril2,2009,awet antecedentcondition.Thistime,thedyearrivedatCSin 15minutes.ThedepthandvelocityatSSwerenot measured,butthedepthwasnoticeablygreaterandflow volumemuchhigheronApril2,2009thanontheprevious tracerdateonAugust28,2008,arelativelydryperiod. AnothertracertestconductedonAugust28,2008resulted inthegreendyetaking18minutestoarrivefromGWto CS,anddyeatthislocationwasnotrepeatedontheApril 2,2009trace. B.G EOCHEMICAL D ATA Tukey’smultiplemeanstestingforgeochemicaldata (Fig.4)showedsignificant( a 5 0.05)differencesinTSS F LOWCHARACTERIZATIONINTHE S ANTEE C AVESYSTEMINTHE C HAPEL B RANCH C REEKWATERSHED UPPERCOASTALPLAINOF S OUTH C AROLINA ,USA 140 N JournalofCaveandKarstStudies, August2013

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andVSSbetweenlocationsSS,CS,andGW.Thehighest valuesweremeasuredatSS,whichvariedbasedonsurface waterconditionsandaveraged5.0mg/LforTSSand 1.5mg/LforVSS.Significantly( a 5 0.05)lowervalues weremeasuredatGWandCS.CShadanaverageTSS valueof0.5mg/LandanaverageVSSvalueof0.2mg/L. GWhadanaverageTSSvalueof0.3mg/Landanaverage VSSvalueof0.1mg/L.ThehighTSSandVSSvaluesat GWmeasuredonthedayofasinkholecollapseinthe suspectedupstreamportionoftheconduitwerenot includedintheaverages.TheextremelyhighTSSand VSSvalueswere18.6and2.5mg/L,respectively,a6100% and2400%increaseintheaveragevalues. Significantdifferences( a 5 0.05)werealsofound betweenthesamplinglocationsfordissolvedcalcium.The measurementsatSPSweresignificantlyhigherthanGW andCS,which,inturn,weresignificantlyhigherthanSS. TheaveragedissolvedcalciumvalueatSPSwas85.1mg/L. Theaveragevaluewas53.3mg/LforGW,46.1mg/Lfor CS,and23.7mg/LforSS.Thelowestmeasuredvalue (15.6mg/L)atSSwas83%lowerthanthehighestmeasured value(91.7mg/L)atSPS. Significantdifferences( a 5 0.05)foralkalinitywere alsoobservedbetweenthesamplinglocations.The measurementsatSPSweresignificantlyhigherthanat GWandCS,which,inturn,weresignificantlyhigherthan attheSS.TheaveragealkalinityvalueatSPSwas 221.3ppm.Theaveragevaluewas146.7ppmforGW, 131.8ppmforCS,and61.2ppmforSS.Thelowest Figure3.DailymaximumelevationsofCSandLake Marion(proxyforCBC)demonstratearelationshipof increasinganddecreasingelevationintandem(elevation scalesonseparateaxesandvarybyafactorof4). Figure4.Meanwaterqualitydataand95%confidenceintervalshownbyverticalbarfor(a)totalsuspendedsolids(TSS)and volatilesuspendedsolids(VSS)(n = 12,nosamplesfromSPS),(b)dissolvedcalciumandmagnesiumions(n = 15exceptat SPSwheren = 5,andSSwheren = 14),(c)specificconductance(n = 24exceptSPSwheren = 5)forthedesignatedlocations (Fig.1),(d)alkalinity,and(e)pH(bothhaven = 19exceptforSPSwheren = 3).SPSwassampledlessoftenduetolowflow volumeatlocation,likelyduetodroughtduringsamplingperiod.GWandSSflowcombinetogetherrightbeforedischargingat CS;SPSisseparatefromtheSanteeCavesystem. A.E.E DWARDS ,D.M.A MATYA ,T.M.W ILLIAMS ,D.R.H ITCHCOCK AND A.L.J AMES JournalofCaveandKarstStudies, August2013 N 141

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measuredvalue(46.0ppm)atSSwas81%lowerthanthe highestmeasuredvalue(236.0ppm)atSPS. Significantdifferences( a 5 0.05)werealsofound betweenthesamplinglocationsforspecificconductance. ThemeasurementsatSPSweresignificantlyhigherthan GWandCS,which,inturn,weresignificantlyhigherthan atSS.TheaveragespecificconductancevalueatSPSwas 397.2 m S.Theaveragevaluewas292.8 m SforGW,274.7 m S forCS,and129.3 m SforSS.Thelowestmeasuredvalue (104.7 m S)atSSwas76%lowerthanthehighestmeasured value(445.0 m S)atSPS. DissolvedmagnesiumandpHvaluesshowlittle variationbetweenthefourlocationsSS,GW,CS,and SPS,asindicatedindifferenceofmeanstestinginFigure4. Valuesfordissolvedmagnesiumweresimilarbetweensites andrangedfrom1.2–1.8mg/Lwithameanconcentration of1.5mg/LatalllocationsexceptSPS,whichhadamean concentrationof1.3mg/L(Figure4).ThepHvalues rangedfrom6.6–7.6SUwithmeansof7.1,7.0,6.9,and7.2 atSS,CS,GW,andSPS,respectively(Fig.4). C.S TABLE I SOTOPE A NALYSES Stableisotopesofwater(Fig.5)plotalongtheCape HatterasLocalMeteoricWaterLine(LMWL),usedhere toapproximatetherangeofsignaturesforprecipitationin theCBCwatershedregion;thisdataisavailablethrough theIAEA’sGlobalNetworkofIsotopesinPrecipitation (GNIP)programandincludestwodifferentversionsofline regressionfits(IAEA/WMO,2006).Thelong-termweightedmeanisotopicsignatureofprecipitationfromtheCape HatterasGNIPstationisalsoincludedinthisfigure. Springwaters(SPS),GWandCSplotsignificantly morenegative(depletedoftheheavierisotopes)andcloser togetherthanSS.Theyalsoplotclosertothelong-term weightedmeanprecipitationsignature,expectedfor groundwaterrechargedbycontemporaryprecipitation. GWplotsmorenegativethanSPS,suggestingthatSPS, withlessupdipaquiferarea,maybeslightlymore influencedbysurfacerecharge.ThisalsoappearsconsistentwithearlierreferencetoStone(pers.comm.,2009) whereGWhasbeenshowntobeacombinationofrecharge andoldwater. On3-18-09,samplesfromallthreelocations(GW,CS andSS)werecollectedduringbaseflowconditions2days aftera25–30mmrainevent.Usingasimplelinearmixing modelapproach(e.g.SklashandFarvolden,1979),with contributingend-membersofGW( d 18 Oof 2 4.38 % )and SS( d 18 Oof 2 3.61 % ),diffusegroundwatercontributionsto CS( d 18 Oof 2 4.17 % )areestimatedtobe73%ofdischarge, with23%derivedfromupstreamsurfacewaterSS. D ISCUSSION ResultsfromthisstudyindicatethattheflowatCSand thespringseepswasrechargedpredominantlyfromdiffuse flowfromthematrix,withaslowresponseofsurfacewater infiltrationtotheconduit.Theterm‘‘slow’’isrelative.In thisstudy,‘‘slow’’impliesthatsurfacewaterdidnot respondwithimmediate(stormevent)movementfromthe surface,intotheaquifermatrix,andthendischargedfrom springseepsintheCBCwatershed. Springsdominatedbydiffuseflowwithslowsurface waterrechargehavefewervariationsinflowthansprings dominatedbyconduitflow.DataanalysisfromWilliamset al.(2012)indicatedthathydrographpeaksatCSwerenot fromGWbaseflowbutmainlySSinputsfromsurface waterrunoffduringrainevents.Independentstableisotope analysisadditionallysuggestedthatdischargeatCSis dominatedbydiffuseflowfromtheaquifermatrix(slow movingwater),contributedheavilyfromtheGWconduit. Alinearmixingmodelusing d 18 Oandassumingcontributingend-membersofGWandSSestimateddiffuse groundwatercontributionstoCStobe73%GWand 23%SS. TheTSSandVSSmeasurementsalsosubstantiatedthe claimofadominantGWcontributionfromdiffuseflowto CS.TSSandVSSvaluesatGWduringthestudyperiod wereconsistentlysmallvaluesandhadasmallstandard deviation.Thiswaslikelyduetosinkholesintheupbasinof thesurfacewatershedroutinelydisplayingnoobvious directopeningstothekarstbelow.Theonlyanomalously largeTSSandVSSvaluesoccurredafterasinkhole collapseinasuspectedupstreamportionoftheconduit servingGW(EdwardsandAmatya,2010).Theotherwise lowTSSandVSSvaluesindicatedatypicallyslowarrival timefromsurfacetogroundwaterinputstoGW.TheTSS andVSSvaluesatCShadonlyslightlylargeraveragethan Figure5.StableisotopesignaturesofSS,CS,SPS,and GWfromMar-09andAug-09samplingevents.LMWLfor theCapeHatterasGNIPstation(includestwoversionsof regressionfit)andthelong-termweightedmeanprecipitation signature(LTM)arealsoincluded. F LOWCHARACTERIZATIONINTHE S ANTEE C AVESYSTEMINTHE C HAPEL B RANCH C REEKWATERSHED UPPERCOASTALPLAINOF S OUTH C AROLINA ,USA 142 N JournalofCaveandKarstStudies, August2013

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GW,butsignificantlysmallerav eragethanSS,indicatingthe dominantflowtoCSduringbaseflowwasclearlyfromGW andnotSS.TheaverageTSSandVSSvaluesweresignificantly largeratSSthaneitherGWorCS,andlikelycontributedto theslightincreaseofTSSandVSSvaluesatCS. Alkalinitymeasurementsalsosubstantiatedthisclaim ofadominantGWcontributionfromdiffuseflowtoCS. ThealkalinitymeasurementatGWduringtheApril2, 2009stormeventwas148.0ppm,veryclosetothe 146.7ppm(5.0ppmstandarddeviation)averagemeasurementatthatlocation.Duringthesamestormevent,the alkalinitymeasurementatCSwas70ppm,almosthalfas muchthantheaveragemeasurementof131.8ppm (20.4ppmstandarddeviation)atthatlocation.The minimalchangeinaveragevalue,withasmallstandard deviation,atGWduringastormeventindicateddiffuse flow.TheflowatCSshowedasignificantdecreasein alkalinity.TheCSflowwaslikelydilutedwiththesurface watersofsignificantlyloweralkalinityfromSS(average alkalinityof61.2ppmwitha6.4standarddeviation). Thealkalinity,dissolvedcalciumandspecificconductancemeasurementsfromthefoursamplinglocationsSS, CS,GW,andSPSallrepresentedvaryingdegreesofdiffuse flowandsurface-groundwaterconnectionsintheSantee LimestonematrixintheCBCwatershed.Thesampling locationSPS,withthehighestofthesemeasurements, representedthedischargewiththelongestresidencetimein theaquifer;andtherefore,longestcontactwiththebedrock andslowestflow.Theseepswerenothydrologically connectedtotheSanteeCavesystem.Morelikely,these seepswereimpactedbythefluctuatingCBCandLake Marionwaterelevation,asdepictedinFigure3(discussed below).TheconduitservingSanteeCaveatGWandCS hadsignificantlyloweralkalinity,specificconductance, anddissolvedcalciumthanSPS,indicatingshorter residencetimeofwaterintheaquifer;andtherefore,faster throughorconduitflowthanSPS.GWhadsimilar,but slightlylarger,alkalinityconcentrationsthanCSduetothe contributiontoCSfromnotonlythepredominantCS baseflowcomponentofGW,butalsoSSbaseflowand stormflowcomponentstoCS.Thelowestvalues,as expected,weremeasuredatthesurfacewatersofa disappearingstreamatSS.Thedifferenceinthechemistry inspringseepsandGWcouldbeduetoseveralfactors. ThenoticeablylargerconduitsizeatGWcouldmovewater fasterthroughtheaquifertoreducecontacttimewiththe limestone.ThegroundwaterconduitfeedingGWalsohad alargersurfacewatershedrechargeareawithsinkholes thatmayallowforquickerinfiltrationofwaterintothe matrixandtotheconduit.Onceinthematrix,theaquifer rechargecouldflowdiffuselyoveralargerareaforGW thanSPS,untilitreachedtheGWconduit.Theseepswere alsohigherinelevationwithintheSanteeLimestoneunit, givingthempotentiallylessvolumeoftheaquifermatrix abovetoyieldadischarge. ThestageelevationatCSandadjacentLakeMarion (equaltotheelevationofthefloodedportionofCBC)also indicatedarelationbetweenfluctuatingbaselevelofCBC anddischargeoftheSanteeLimestonekarstaquiferatCS. IfweignorethepeaksinFigure3,whichweresurface watercontributedfromSL7,thebaseflowstageelevation atthespringbroadlyincreasedanddecreasedwiththe changinglakelevels.Ifthesurface-groundwaterconnectionsinGWwerequick,onewouldnotexpectthebroad severalmonthlongcorrespondences,butratherwould expectrapidchangesintotheconduitthatservesGWand CS.ThecorrespondencesuggestedtheGWsourcereflected arainfallresponseataratemoresimilartothelarge reservoir.GeochemicaldataalsoindicatedaslowsurfacegroundwaterconnectiontypicalforGWandCS(minusthe stormflowcontributionsfromSS),andthehydrologicdata inFigure3werecollectedduringdroughtyears,as indicatedfrommeasuredprecipitationcomparedtolongtermprecipitationrecords(Williamsetal.,2012). Theseresultssuggestedthatduringlowlakelevels,such asatthetimeoffirstdyetrace,thewatertableinthe limestonemightbelowandthesurfacewatergoingintoSS infiltratesdownwardintotheemptyconduits.Thewater maytheneitherbestoreduntilthewatertableincreasesor perhapscontinuesdrainingtoCBCthroughtheaquiferat alowerelevationthanthecavestreampassage,whichwas hypothesizedearlierbyAmatyaetal.(2011).Forthefirst dyetrace,whennodyewasobservedfromSStoCS,the averagelakelevelonAugust28,2008was21.79meters (71.49feet)(USGS,2009).TheaveragelakelevelonApril 2,2009,whenthedyetookonly15minutestoreachCS, was22.96m(75.34feet)(USGS,2009).Thus,a1.17m (3.85foot)fluctuationinlakelevelshowedanimpacton thewatertableinthelimestoneaquiferandcavestream.As lakelevelsdecrease,theaquifermayemptyatlower elevationspringsandmoresurfacewaterinthebasinislost togroundwater,aswasalsoarguedbyAmatyaetal. (2011). ThetraceconductedonAugust28,2008resultedinthe greendyetaking18minutestoarrivefromGWtoCS.The conduitfromGWtoCScanbemostlytraversedandthere wasnoindicationoflosstoalowerconduit.Infact, conduitbaseflowdischargeatGWandCSwaswitnessed onweeklyfieldvisitsduring2009asalwayscontinuousand steady.Because2007–08weredroughtyears,theSantee Limestoneinthisregionappearedtohavesignificant storagecapacity,characteristicofdiffuseflow,tomaintain steadyflowatthespringafteratwo-yeardrought.Even thoughGWmayresultfromflowinmultipleconduits fromanunknownupstreamarea,therechargewaslikely morediffusethandirectrechargefromsurfacewaterand precipitation.Aconceptualmodelforgroundwaterflowin theCBCwatershedisshowninFigure6. Thelackofsignificantdifferencebetweenthelocations forpHparameterindicatedalimitedvalueformeasuring theseindividualparameterstoidentifyflowtypeand A.E.E DWARDS ,D.M.A MATYA ,T.M.W ILLIAMS ,D.R.H ITCHCOCK AND A.L.J AMES JournalofCaveandKarstStudies, August2013 N 143

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surface-groundwaterconnectionsinthisparticularwatershed.However,thecomparisonbetweenvariabledissolved calciumandnon-variabledissolvedmagnesiumbetween locationsprovidedinsightintokarstsubsurfaceflow dynamicsgiventhecontacttimewithlimestone. C ONCLUSIONS Theanalysisofhydrologic,geochemicalandstable isotopicdatacollectedwithintheChapelBranchCreek watershedin2009confirmedthefirsthypothesisthatthe dischargeviaCSfromtheSanteeLimestonetoCBCis predominantlyrechargedbyadiffuseflowcomponentfrom thematrixwithslowsurface-groundwaterconnection.The SanteeLimestoneintheupperFloridanaquifer,therefore, behaveslikethekarstinthelowerFloridanaquifer,with thematrix(diffuseflow)contributingheavilytoaquifer dischargeviaconduitstosprings.Analysisandinterpretationofthesedataalsoconfirmedthesecondhypothesisthat thefluctuatingCBCstageelevationsinfluencetheaquifer watertableatSanteeCave.StudiesintheFloridanaquifer havelikewisefoundhydrologicconnectionsbetweenthe surfacewaterinlakesandgroundwaterlevels(Lee,2000; Watsonetal.,2001;Lin,2011). CharacterizingthehydrologyoftheChapelBranch Creekwatershedisessentialforwaterresourceprotection. Thisreservoir-likeembaymenthasbeenontheEPA303(d) listforimpairedwaterbodiesforseveralyears,andboth pointandnon-pointsourcesofwaterpollutionare abundantinthismultipleland-usewatershed.Further studiestoelucidatesurface-groundwaterconnectionsand waterqualityatthespringsalongCBCwouldhelpwith futureBestManagementPracticesforChapelBranch Creekandthegreaterkarstregion. A CKNOWLEGEMENTS Thisstudywasmadepossiblebythesupportfrom SouthCarolinaDepartmentofHealthandEnvironmental ControlÂ’s319GrantAgreement # EQ-7-514(Project # 4OFY06)withUSForestServiceCenterforForest WetlandsResearch(SRS06-CO-11330135-122).Theauthorsacknowledgethefollowingforassistanceinthis particularstudy:IrvinPittsandValerieCarter-Stonewith SCDNRforpermittoaccessSanteeCave;Fieldhelpfrom FStechniciansAndyHarrisonandMattKaswaski;Water sampleanalysisfromLarryMcCord,BrianLynch,Chip Davis,AnitaBrown,LindaWilliamsandDebraGuerry fromSanteeCooperBiological&AnalyticalLaboratory; weyerhaeusercompanyCooperationfromMayorSilas SeabrooksandHermanKellerwiththeTownofSantee andGaryBennettwithSanteeNationalGolfCourse; RangersNathanMaiwaldandAdinFellwithSanteeState Park;SCDHECHydrologistPeteStone;SCDHECAndy Miller,MeredithMurphy,MattCarswell,andNydia Burdick;SCDNRJimScurry,andNPSDenverIngram. ThanksalsotoDr.HerbertSseganeatUniversityof Georgiaforassistancewithstatisticalanalysisofthewater qualitydata.AndspecialthankstotheresidentsofChapel BranchIIfortheirboundlessenergyandlocalsupport. Financialsupportforstableisotopeanalyseswereprovided bytheCanadaFoundationofInnovation,theCanada ResearchChairProgramandNipissingUniversity,ON, Canada.ThankstoDr.KrysChutkofortechnicalsupport onisotopeanalyses.CapeHatterasGNIPstationdata wereaccessedthroughtheInternationalAtomicEnergy AgencyÂ’sGNIPprogramwithanalyticalsupportfromthe UniversityofCopenhagenandtheUSGSRestonStable IsotopeLaboratory. Figure6.Conceptualmodelofgroundwaterflow(diffuseandconduit)betweenSanteeLimestonekarstaquiferandChapel BranchCreekdischargingtoLakeMarion. F LOWCHARACTERIZATIONINTHE S ANTEE C AVESYSTEMINTHE C HAPEL B RANCH C REEKWATERSHED UPPERCOASTALPLAINOF S OUTH C AROLINA ,USA 144 N JournalofCaveandKarstStudies, August2013

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R EFERENCES Amatya,D.M.,Manoj,J.,Edwards,A.E.,Williams,T.M.,andHitchcock,D.R.,2011,SWAT-basedstreamflowandembaymentmodeling ofkarstaffectedChapelBranchwatershed,SC:Transactionsof AmericanSocietyofAgriculturalandBiologicalEngineers,v.54, no.4,p.1311–1323. Amatya,D.M.,Williams,T.M.,Edwards,A.E.,andHitchcock,D.R., 2010,FinalDraftReportandAppendicesA,B,C,andDon‘‘Total MaximumDailyLoad(TMDL)DevelopmentforPhosphorus, ChapelBranchCreek(SC014)’’submittedtoSCDepartmentof Health&EnvironmentalControl,BureauofWater,Divisionof WaterQuality,Columbia,SCApril2010. Ashton,K.,1966,Theanalysisofflowdatafromkarstdrainagesystems: TransactionofCaveResearchGroup,GreatBritain,v.7,p.161–203. Atkinson,T.C.,1977,Diffuseflowandconduitflowinlimestoneterrain inMendipHills,Somerset(GreatBritain):JournalofHydrology, v.35,p.93–100. Clesceri,L.S.,Greenberg,A.E.,andEaton,A.D.,eds.,1998,Standard MethodsfortheExaminationofWaterandWastewater,20 th Edition, Washington,DC,AmericanPublicHealthAssociation. Criss,R.E.,Fernandes,S.A.,andWinston,W.E.,2001,Isotopic, geochemicalandbiologicaltracingofthesourceofanimpactedkarst spring,WeldonSpring,Missouri:EnvironmentalForensics,v.2, p.99–103. Edwards,A.E.,andAmatya,D.M.,2010,SinkholecollapseinSantee, SouthCarolina:PhotoEssay:SouthCarolinaGeology,v.47,p.17–18. Edwards,A.E.,andAmatya,D.M.,2009,Evaluatingforestedkarst topographywithLiDAR:USDAForestServiceBeneaththeForest Fall2009Newsletter;Accessedonwebat:http://fsweb.wo.fs.fed.us/ mgm/geology_notebook.html. ERC,1999,DrainageanalysisofSCDHECdam # D-3746(lowerSantee Shoresdam),Santee,SouthCarolina,Orangeburg,S.C.:Engineering ResourcesCorporation. Fiorillo,F.,2009,Springhydrographsasindicatorsofdroughtsinakarst environment:JournalofHydrology,v.373,p.290–301. Florea,L.J.,andVacher,H.L.,2006,Springflowhydrographs:eogenetic vstelogenetickarst:GroundWater,v.44,no.3,p.352–361. Gunn,J.,1986,Aconceptualmodelforconduitflowdominatedkarst aquifers in Gunay,G.,andJohnson,A.I.,eds.,Proc.Ankara Symposium,‘‘Karstwaterresources’’,July1985,IAHSPublication, v.161,p.587–596. Hockensmith,B.L.,2009,PotentiometricmapoftheFloridanaquiferand tertiarysandaquiferinSouthCarolina-2004:SouthCarolina DepartmentofNaturalResourcesWaterResourcesReport48. Holler,C.Jr.,2000,CavesofSouthCarolina:OldFort,NorthCarolina, HollowHills,91p. IAEA/WMO,2006,GlobalNetworkofIsotopesinPrecipitation.The GNIPDatabase.Accessibleat:http://www.iaea.org/water. Jacobson,D.A.,andLangmuir,D.,1974,Controlsonthequality variationsofsomecarbonatespringwaters:JournalofHydrology, v.23,p.247–265. Jakucs,L.,1959,NeuemethodenderhohlenforschunginUngarnundihre Ergebnisse:DieHohle,v.10,no.4,p.88–98. Katz,B.G.,2004,Sourcesofnitratecontaminationandageofwaterin largekarsticspringsofFlorida:EnvironmentalGeology,v.46, p.689–706. Katz,B.G.,Coplen,T.B.,Bullen,T.D.,andDavis,J.H.,1997,Useof chemicalandisotopictracerstocharacterizetheinteractionsbetween groundwaterandsurfacewaterinmantledkarst:GroundWater, v.35,no.6,p.1014–1028. Lee,T.M.,2000,Effectsofnearshorerechargeongroundwater interactionswithalakeinmantledkarstterrain:WaterResources Research,v.36,no.8,p.2167–2182. Lin,Z.,2011,Estimatingwaterbudgetsandverticalleakagesforkarst lakesinnorth-centralFlorida(UnitedStates)viahydrological modeling:JournaloftheAmericanWaterResourcesAssociation, v.47,no.2,p.287–302. Mahler,B.J.,andLynch,L.,1999,Muddywaters:temporalvariationin sedimentdischargingfromakarstspring:JournalofHydrology, v.214,p.165–178. Martin,J.B.,andGordon,S.L.,2000,Surfaceandgroundwatermixing, flowpaths,andtemporalvariationsinchemicalcompositionsofkarst springs, in Sasowsky,I.D.,andWicks,C.M.,eds.,GroundwaterFlow andContaminantTransportinCarbonateAquifers,A.A.Balkema, Rotterdam,Netherlands,p.5–92. Martin,J.B.,andDean,R.W.,2001,Exchangeofwaterbetweenconduits andmatrixintheFloridanaquifer:ChemicalGeology,v.179, p.145–165. Massei,N.,Lacroix,M.,Wang,H.W.,Mahler,B.J.,andDupont,J.P., 2002,Transportofsuspendedsolidsfromakarstictoanalluvial aquifer:theroleofthekarst/alluviuminterface:JournalofHydrology, v.206,p.88–101. Mathworks,Inc.,2009,MATLABSoftwareversionR2009b,Natick,MA, USA. Moore,P.J.,Martin,J.B.,andScreaton,E.J.,2009,Geochemicaland statisticalevidenceofrecharge,mixing,andcontrolsonspring dischargeinaneogenetickarstaquifer:JournalofHydrology, v.376,p.443–455. Plummer,L.N.,Busenberg,E.,McConnell,J.B.,Drenkard,S.,Schlosser, P.,andMichel,R.L.,1998,Flowofriverwaterintoakarstic limestoneaquifer.1.Tracingtheyoungfractioningroundwater mixturesintheUpperFloridanAquifernearValdosta,Georgia: AppliedGeochemistry,v.13,no.8,p.995–1015. Quinlan,J.F.,andRay,J.A.,1995,Normalizedbase-flowdischargeof groundwaterbasins:Ausefulparameterforestimatingrechargearea ofspringsandforrecognizingdrainagenomaliesinkarstterranes, in Beck,B.F.,ed.,Karstgeohazards,Rotterdam,A.A.Balkema, p.149–164. Ritorto,M.,Screaton,E.J.,Martin,J.B.,andMoore,P.J.,2009,Relative importanceandchemicaleffectsofdiffuseandfocusedrechargeinan eogenetickarstaquifer:anexamplefromtheunconfinedupper Floridanaquifer,USA:HydrogeologyJournal,v.17,p.1687–1698. SCS,1988,SoilSurveyReportofOrangeburgCounty,SC,State ConservationistOffice,Columbia,SC. Shuster,E.T.,andWhite,W.B.,1971,Seasonalfluctuationsinthe chemistryoflimestonesprings:apossiblemeansforcharacterizing carbonateaquifers:JournalofHydrology,v.14,p.93–128. Sklash,M.G.,andFarvolden,R.N.,1979,Theroleofgroundwaterin stormrunoff:JournalofHydrology,v.43,p.45–65. Stone,P.,2009,PersonalCommunication,Hydrologist,SCDepartment ofNaturalResources. USGS,2009,LakeLevelsatElloree,SC.Reston,VA:U.S.Geological Survey. Vesper,D.J.,andWhite,W.B.,2004,Stormpulsechemographsof saturationindexandcarbondioxidepressure:implicationsforshifting rechargesourcesduringstormeventsinthekarstaquiferatFort Campbell,Kentucky/Tennessee,USA:HydrogeologyJournal,v.12, no.2,p.135–143. Watson,B.J.,Motz,L.H.,andAnnable,M.D.,2001,Waterbudgetand verticalconductanceforMagnoliaLake:JournalofHydrologic Engineering,v.6,no.3,p.208–216. White,W.B.,2003,Conceptualmodelsforkarsticaquifers,Karst Modeling,SpecialPublication5,TheKarstWatersInstitute, p.11–16. White,W.B.,1988,GeomorphologyandHydrologyofKarstterrains, NewYork,OxfordUniversityPress,464p. Willoughby,R.H.,2002,GeologicMapoftheSaintPaulQuadrangle, Calhoun,Clarendon,andOrangeburgCounties,SouthCarolina:S.C. GeologicQuadrangleMapGQM-08,scale1:24000,1sheet. Williams,T.M.,Amatya,D.M.,Hitchcock,D.R.,andEdwards,A.E., 2013,Streamflowandnutrientsfromakarstwatershed withadownstreamembayment:ChapelBranchCreek,Journalof HydrologicalEngineering,inpress. Williams,T.M.,2009,Personalcommunication,ProfessorofForest Hydrology,ClemsonUniversity. Worthington,S.R.H.,Davies,G.J.,andFord,D.C.,2000,Matrix, fractureandchannelcomponentsofstorageandflowinapaleozoic limestoneaquifer, in Wicks,C.M.,andSasowsky,I.D.,eds., Groundwaterflowandcontaminanttransportincarbonateaquifers: Rotterdam,Balkema,p.113–128. A.E.E DWARDS ,D.M.A MATYA ,T.M.W ILLIAMS ,D.R.H ITCHCOCK AND A.L.J AMES JournalofCaveandKarstStudies, August2013 N 145

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NEWSPECIESANDNEWRECORDSOFSPRINGTAILS (HEXAPODA:COLLEMBOLA)FROMCAVESINTHE SALEMPLATEAUOFILLINOIS,USA F ELIPE N.S OTO -A DAMESAND S TEVEN J.T AYLOR IllinoisNaturalHistorySurvey,UniversityofIllinoisatUrbana-Champaign,1816S.OakSt.,ChampaignIL61820USA, fsoto@illinois.edu,sjtaylor@illinois.edu Abstract: Thespringtail(Hexapoda:Collembola)faunaofeightcaves(WizardCave, PautlerCave,SpiderCave,Wanda’sWaterfallCave,IllinoisCaverns,StemlerCave, HiddenHandCave,andBatSumpCave)intheSalemPlateauofsouthwesternIllinois (MonroeandSt.Claircounties)wassurveyedin2009usingacombinationofmethods, includingpitfalltraps,Berlese-funnelprocessingoflitter,andhandcollectionsby quadrat,ondrippools,freestandingbait,andrandomlocations.Intotal,forty-nine speciesofspringtailswerefound.Fouraredescribedasnewtoscience( Onychiurus pipistrellae n.sp., Pygmarrhopalitesfransjanssens n.sp, P.incantator n.sp,and P. salemensis n.sp),fourmayrepresentnewspeciesbutthereisinsufficientmaterial availabletopreparefulldescriptions(twospeciesinthegenus Superodontella ,onein Pseudachorutes ,onein Sminthurides ),andthreeothers( Ceratophysella cf. brevis C. cf. lucifuga, and Folsomia cf. bisetosa )areidentifiedtospecies,butdifferencesfromthe nominalspeciessuggestfurtherstudiesmayindicatetheIllinoispopulationsrepresent distinctforms.Inaddition,fiveotherspeciesrepresentnewrecordsforIllinois,and eighteenarenewcaverecordsforthespeciesinNorthAmerica.Thenewrecordsmore thandoublethenumberofspringtailsspeciesknownfromcavesintheSalemPlateau region.Morethanhalf(twenty-nine)ofthespeciesreportedarerankedasrare(S1–S2)at thestatelevel.ThetotalnumberofspringtailspeciesinSalemPlateaucavescouldbe morethantwicewhatisrecordedinthepresentstudy,andmorenewspeciesandstate recordsshouldbefoundwhencavesinotherIllinoiskarstregionsaremorethoroughly examined. I NTRODUCTION Illinois’skarstisdistributedacrossfiveregions(Fig.1) thatcontainnumeroussinkholes,springs,andshallow groundwaterconduits.Fouroftheseregions—Driftless Area,LincolnHills,SalemPlateau,andShawneeHills— containcavesaccessibletohumans.Inadditiontotheir hydrological,recreational,geological,andculturalvalues, thesecavescontainfascinatingassemblagesoflife.The faunamostfamiliartothepublicarebatsandsalamanders, butcavesalsocontainawidevarietyofinvertebrates. AmongthesearetheIllinoiscaveamphipod, Gammarus acherondytes HubrichtandMackin,whichisfederally listedasendangered,theenigmaticcavesnail, Fontigens antrocetes (Hubricht),astate-listedspecies,andasinglesiteendemicIllinoiscavebeetle Pseudanophthalmusillinoisensis BarrandPeck.Onecavespringtail, Pygmarrhopalitesmadonnensis (ZeppeliniandChristiansen),islistedas state-endangeredinIllinois.Numerousotherinvertebrate speciesoccurinIllinoiscaves,includingavarietyofother springtails(Collembola). Springtailsaresmallhexapodscharacterizedbythe presenceoffour-segmentantennae,asix-segmentabdomen,alargevesicle(theventraltube)ontheventralpartof thefirstabdominalsegment,and,inmanyspecies,a jumping-organcomplexformedbythetail-likefurculaand thefurculacatchorretinaculum.Springtailsaremost commonlyfoundinsoilandleaflitter,buttheyhave invadedotherspecializedhabitats,includingcaves.Many soilorleaf-litterspeciesarecommonlyfoundincavesas xenobionts,butsomespeciesarecave-adaptedorcavelimitedanddonotsustainsurfacepopulations(ChristiansenandCulver,1987). InIllinois,themostcommonfamiliesofCollembola reportedfromcavesareHypogastruridae,Onychiuridae, Oncopoduridae,Tomoceridae,Isotomidae,Entomobryidae,andSminthuridae sensulato .Oftheforty-threespecies ofspringtailspreviouslyrecordedfromIllinoiscaves, slightlymorethan25%(elevenspecies)areeithertroglobionts(obligatorilypermanentresidentsofsubterranean habitats)oreutroglophiles(facultativelypermanentresidentsofsubterraneanhabitats).ThegeneraofeutroglophilesortroglobiontsreportedfromIllinoispriorto ourstudyare Typhlogastrura Lethemurus Oncopodura Pseudosinella Sinella,Pygmarrhopalites, and Arrhopalites Pygmarrhopalitessapo (ZeppeliniandChristiansen)and Pygmarrhopalitesmadonnensis (ZeppeliniandChristiansen)aretheonlyspeciesofcavespringtailscurrently knowntobeendemictoIllinois.Thenumberofendemics isprobablyhigherthancurrentinventorieswouldsuggest, F.N.Soto-AdamesandS.J.Taylor–Newspeciesandnewrecordsofspringtails(Hexapoda:Collembola)fromcavesintheSalem PlateauofIllinois,USA. JournalofCaveandKarstStudies, v.75,no.2,p.146–175.DOI:10.4311/2011LSC0257 146 N JournalofCaveandKarstStudies, August2013

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asmostgroupshavenotbeenstudiedindetail.For example,thegenera Typhlogastrura and Pseudosinella have diversifiedextensivelyincaveselsewhereinNorthAmerica andEurope,andalthoughbotharereportedfromIllinois caves,nospeciesfromeithergenushasbeenidentifiedfor thestate.Inaddition,someeutroglophilesandtroglobionts firstthoughttobewidelydistributedacrosskarstregions arenowknowntorepresentcomplexesofspeciesendemic tojustafewcaves.Themoststrikingexampleofthisisthe Onychiurusreluctus speciescomplex,inwhichfivespecies wererecentlyidentified(Pomorskietal.,2009). Thisreportpresentsfindingsfromaninventoryof springtailscollectedineightcavesinIllinois’sSalemPlateau. M ETHODS F IELD S AMPLING Ateachcave,thedominanthabitattypesinwhich springtailsmightoccurweresampled.Collectionswere madeusingLimburgercheese-baitedpitfalltrapspartially filledwith95%ethanol,Berlese-funnelprocessingoflitter samples,andhandsamplingusinganaspiratoreitherina structuredfashion(timedsearchofdrippoolsandquadrat searchesoncavefloorsandwalls)orbygeneralinspection aroundhaphazardlyplacedcheesesmearsandother potentialsourcesofenergysuchasanimalscatsand rottinglogs.Allspecimenswerepreservedin95%ethanol. EightcaveswereselectedforstudyintheSalemPlateau (Fig.2).Allnecessarypermitsrequiredtoconductthe researchwereobtainedpriortothebeginningoffieldwork. Thecollectiondatesgivenbelowincludetheperiodduring whichpitfalltrapswereexposedinthefield.Littersamples tobeprocessedinBerlesefunnelsinthelaboratorywere usuallycollectedduringthesecondvisittoacave. Collectioninformationforeachcaveisasfollows: USA:IL:St.ClairCo.:WizardCave,Dupo,nearFalling Spring,15–17June2009,SJTaylor,FNSotoAdames,andCAPhillips. USA:IL:MonroeCo.:PautlerCave,3.0miWSWof Waterloo,14–16September2009,SJTaylor,andFN Soto-Adames. USA:IL:MonroeCo:Wanda’sWaterfallCave,7.4miSE ofValmeyer,15–17September2009,SJTaylorand FNSoto-Adames. USA:IL:MonroeCo:SpiderCave,6.5miSofWaterloo, 15–17September2009,SJTaylorandFNSotoAdames. Figure2.DistributionofeightIllinoiscaveswherespringtailswerecollectedduring2009.Grayshadingindicates approximateboundariesofkarstbasedonsinkholemapping (IllinoisStateGeologicalSurvey).Cavelocationsare approximate. Figure1.Illinoiskarstareas,adaptedfromWeibeland Panno(1997)andPannoetal.(1997). F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 147

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USA:IL:MonroeCo:IllinoisCaverns,24–26September 2009,SJTaylor,FNSoto-Adames,AKuhns,E Zaborski,JJacoby,APaprocki,andMPessino. USA:IL:St.ClairCo:StemlerCave,2.7miNEof ColumbiaIL,28–30September2009,SJTaylorand FNSoto-Adames. USA:IL:MonroeCo:HiddenHandCave,3miWof Waterloo,14–16October2009,SJTaylorandFN Soto-Adames. USA:IL:MonroeCo:BatSumpCave,6.6miWSWof RedBud,3–5November2009,SJTaylorandFN Soto-Adames. N OTESTO S PECIES D ESCRIPTIONS ,E COLOGICAL C LASSIFICATIONAND D ISTRIBUTIONAL R ANKING Thenomenclatureofthechaetotaxyin Ceratophysella and Superodontella followsFjellberg(1985)andJordana etal.(1997),respectively.Thechaetotaxyof Pseudosinella followsSzeptycki(1979)andSoto-Adames(2010). Nomenclatureofthechaetotaxyoftheheadandsmall abdomenin Pygmarrhopalites followsChristiansenand Bellinger(1998).TheidentityofheadverticalsetaeM4and M5appearsconfusingintheliterature.WeconsiderM5to formarowwithL2andIL3(rowDinBetschandWaller, 1994),whereasM4formsarowwithL1andIL2(rowC inBetschandWaller,1994).Noneofthespeciesof Pygmarrhopalites reportedherecarriessetaM5,butthe clarificationisnotedbecause P.hirsuta (Christiansen)has beendescribedashavingM5andlackingM4,butan individualidentifiedasrepresentingthisspecies(Zeppelini etal.,2009)examinedbyuscarriesM4(byourconvention) insteadofM5.ThenomenclatureofdistalsetaeinrowsD andEonthelateralvalveofthesmallabdomenis confusing.WehaveidentifiedasE7thesetalabeledE8in ChristiansenandBellinger(1998)becauseinallindividuals examinedbyusthesocketofthissetaalignswithD7(when present)andC7insteadofD8andC8(Figs.15C,F).Seta D9isalwayslongerthansetaeD7–D10andcanbeusedas areferencepointwhensomesetaeintheseriesareabsent. Abbreviationsusedthroughoutthedescriptionsare Ant.,PAO,Th.,andAbd.forantennalsegment,post antennalorgan,thorax,andabdomen,respectively. Typesofthenewspeciesdescribedhereandvouchers forallpreviouslynamedspeciesaredepositedintheIllinois NaturalHistorySurveyInsectCollection. Weassignedeachspeciestoanecologicalclassification followingCulverandPippan(2009),andSket(2008), insteadofthesystemutilizedbyBarr(1963,1968),whichis morefamiliartoNorthAmericanspeleologists.The ecologicalcategoriesconsideredare:troglobiont(TB– obligateandpermanentresidentsofsubterraneanhabitats),eutroglophile(EU–facultativelypermanentresidents ofsubterraneanhabitatsalsofoundinotherhabitats), subtroglophile(SU–obligateorfacultativeresidentsof subterraneanhabitatsthatutilizeotherhabitatsforsome portionoflifecycle)andtrogloxene(TX–sporadic residentsofsubterraneanhabitats[‘‘accidental’’ofBarr 1963,1968]). Rankingsofthespeciesbelowprovideameasureofthe extentofaspeciesdistributionatthestate(S)andglobal (G)level.RankingsarecircumscribedfollowingNatureServeconservationstatusranks(Masteretal.,2009):For stateandglobalimperilment,speciesreportedinfiveor fewerlocalitiesarerankedS/G1;sixtotwentylocalities,S/ G2;twenty-onetoonehundredlocalities,S/G3;uncommonbutnotrare,withsomecauseforlong–termconcern duetodeclinesorotherfactors,S/G4;andwidespreadand common,S/G5.SpeciesintroducedfromotherbiogeographicregionsareindicatedwithIN.Theserankingsare basedonIllinoisandNorthAmericanrecordsobtained fromeithertheCollembolaofNorthAmerica(ChristiansenandBellinger,1998)orthedatabaseofNorth AmericanCollembolarecords(Christiansen,2012).Additionalsourcesofdistributionalinformationarelistedunder individualspeciesaccounts. S PECIES A CCOUNTS H YPOGASTRURIDAE Hypogastrurapannosa (Macnamara),1922—TXS1/G5; NewIllinoisRecord Localities:Wanda’sWaterfallCave,StemlerCave Hypogastrurapannosa ispartofaspeciescomplexthat includes H.essa H.matura, andthePalearctic H.assimilis Hypogastrurapannosa maybeajuniorsynonymof H. assimilis (Babenkoetal.,1994;Thibaudetal.,2004),but wefollowpreviousworkers(Fjellberg,1985;Christiansen andBellinger,1998)inassigningtheepithet pannosa to NorthAmericanpopulationshavingaPAOwithfour papillatearms,mucronallamellaendingabruptlysubapically,andAbd.4withsensillum-likesetap5. ThisspeciesiswidespreadinNorthAmerica,butthisis thefirstreportfromIllinois.Itispossiblethatprevious Illinoisreportsofeither H.matura or,morelikely, H.essa actuallyreferto H.pannosa .Thereisonepreviousreport fromKentuckycaves. Ceratophysellaboletivora (Packard),1873—TXS2/G5 Localities:Wanda’sWaterfallCave,IllinoisCaverns,Bat SumpCave AscircumscribedbyChristiansenandBellinger(1980), basedonthelectotype,thisspeciesischaracterizedby havingAbd.4setap2clearlylongerthanp1andwith7–9 (dependingonthepositionofa5)microsetaeinternaltoa linedrawnacrossa6-p4(Fig.3A),tenenthairscapitate, Yoshii‘‘a’’measure16–18,Ant.4apicalbulbbilobed,and distalinnerdentalsetaeonlyweaklyenlargedbasally.The speciesiseasytoidentifyinalcoholbyhavingrust-redhead andantennae,and,usually,patternlessdarkbluetoblack body. Ceratophysellaboletivora waspreviouslyreportedfrom non-cavesitesinCisco,Markham,andCookcountiesin N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 148 N JournalofCaveandKarstStudies, August2013

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Illinois.Thisspecieshasbeenreportedfromcavesin Missouri,butthisisthefirstrecordforthespeciesin Illinoiscaves. Ceratophysella cf. brevis ChristiansenandBellinger,1980 —EUS1/G5;NewIllinoisRecord Localities:Wanda’sWaterfallCave,BatSumpCave WeidentifyasC.cf. brevis allindividualsshowing charactersofthe C.denticulata speciescomplexasdefined byChristiansenandBellinger(1980),butwithanterior lobesofPAOlessthan1.5 3 thelengthofposteriorlobes andinnerdistaldentalsetaenotsharplybentorbasally enlarged.Ourspecimensarelightbluedorsallyandwhite ventrallyinsteadofdarkbluetoblackaspointedoutby Figure3.Dorsalchaetotaxyoffourthabdominalsegment:(A) Ceratophysellaboletivora ;(B) Ceratophysella cf. brevis ,filledincirclesrepresentmacrosetae,dotsrepresentregularacuminateseta. Superodontella sp.1:(C)labiumandpostlabium; (D)dorsalchaetotaxyofleftsideofhead,arrowpointstosetaabsentin S.substriata ;(E)dorsalchaetotaxyoffirst abdominalsegment. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 149

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ChristiansenandBellinger(1980)andFjellberg(1985)for Alaskanpopulations.Weexaminedelevenspecimens,six fromWanda’sWaterfallandfivefromBatSump,andall haveacuminateserratemacrosetae,setam6presentonTh. 2–3,andlacksetam3onAbd1–4(Fig.3B).Most individualsalsolacksetam4onAbd.1–3,althoughfour specimens(twofromeachcave)carrythissetaonatleast oneoftheabdominalsegments. Fjellberg(1985)reportedtwoformsof C.brevis from Alaska:oneformshouldbeeasytoidentifyas C.brevis by theabsenceofcoarselyserratedsetaeandabsenceofsetae m6onTh.2–3,andm3–4onAbd.1.Thesecondformhas coarsebodyseta,andallthethoracicandabdominalsetae mentionedabovearepresent.Thissecondformwould differfrom C.engadinensis onlyintheproportionof anteriortoposteriorPAOlobesandinnothavingsharply bentsetaeonthedens.Ourspecimensdonotmatcheither ofFjellberg’sformsortheoriginaldescriptionofthe speciesindetailsofthedorsalchaetotaxyofthebodyand mayrepresentadifferentspeciesaltogether.However,the significanceofthemorphologicalvariationobserved amongpopulationsincludedinthe C.denticulata species complexremainsunclear,andsomeothersourceof informationmayberequiredtosortdiscreteunitswithin thegroup. Thisreportrepresentsthefirstrecordof Ceratophysella cf. brevis fromIllinois. C.brevis wasoriginallydescribed fromsurfacehabitatsinWyoming,butnowitisalso knownfromAlaska(Fjellberg,1985)andfromcavesin Kentucky,California,andIndiana.Inadditiontothe NorthAmericanrecords,thespecieshasbeenreported fromSiberia(Babenkoetal.,1994). Ceratophyselladenticulata (Bagnall),1941—EUS5/G5 Localities:IllinoisCaverns,StemlerCave TheindividualssampledinIllinoisCavernshaveawelldevelopedantennalfileandcoarselyserrate,capitate macrosetae. Ceratophyselladenticulata isacommonspecies, widelydistributedthroughouttheNorthernHemisphere (ChristiansenandBellinger,1998;Thibaudetal.,2004).In Illinois C.denticulata hasbeenpreviouslyidentifiedfromat leastsevencavesinCalhoun,Johnson,Union,andMonroe counties. Ceratophysellaengadinensis Gisin,1949—EU/TXS?/G5; NewIllinoisRecord Locality:IllinoisCaverns Thisisanotherspecieswidelydistributedthroughout thetemperatezoneoftheNorthernHemisphere. Ceratophysellaengadinensis differsfrom C.denticulata onlyinthe absenceofsetaa 9 onAbd.5(ChristiansenandBellinger, 1998;Fjellberg,1998).Althoughthespecieshasnotbeen previouslyreportedfromIllinois,itislikelythatinthepast ithasbeenconfusedwithorincludedaspartof C. denticulata; theindividualscollectedinIllinoisCaverns werefoundmixed-inamongmanyspecimensof C. denticulata .However,werecognizebothspeciesbasedon recenthybridizationstudiesthatsuggestthetwoformsare reproductivelyisolatedbypostembryonicisolationmechanisms(Skarz yn ski,2005). Ceratophysella cf. lucifuga (Packard),1888—TBS1/G1; NewIllinoisRecord Locality:Wanda’sWaterfallCave Thespecieslackspigment,butretainsalleyes,andthere islittlechangeintherelativelengthofantennaeandclaw, orinthedevelopmentofsensoryorgans. Onesubadultmale(1.2mmlong)andtwojuveniles (0.89and0.69mm)wereexamined,andtheydifferfrom C. lucifuga, asredescribedbySkarz yn ski(2007),innothaving pigmentontheeyepatch(individualsappearedtobeblind underthedissectingmicroscope),inhavinganundivided apicallobeonAnt.4(trilobedin C.lucifuga ),inhaving4–6 differentiatedsensillaontheventralfileofAnt.4(15in C. lucifuga ),andinthepresenceintheIllinoisspecimensofa supplementarysetabetweenp5andp6onAbd.4.The maledoesnothaveinnerungualteeth,butthetwo juvenileshaveaclearinnerungualtoothonalllegs.Seta m4isabsentonAbd.1–3andpresentonAbd.4ofthe subadult;m4ispresentonallabdominalsegmentsinthe juveniles. Uptothepresentreport, C.lucifuga wasknownonly fromthreecavesinCrawfordandHarrisoncounties, Indiana.Thephysicaldistance( < 330km)andmorphologicaldifferencesbetweentheIllinoisandIndianapopulations suggesttheymayrepresentdistinctspecies,butadditional materialfromIllinoiswillbeneededtodetermineifthe morphologicaldifferencesnotedherearefixed. Xenyllawelchi Folsom,1916—TXS5/G5 Locality:StemlerCave ThiscosmopolitanspeciesoccursthroughoutNorth AmericaandEurasiainsurfaceleaflitter(Thibaudetal., 2004). O DONTELLIDAE Superodontellacornifer (Mills),1934—TXS1/G5 Locality:BatSumpCave ThisspeciesiswidelydistributedthroughoutNorth AmericaandwaspreviouslyreportedfromHardin County,Illinois.However,thisisthefirstrecordofthe speciesfromanIllinoiscave.Thisindividualisasmall juvenilewithonly4dentalsetae. Superodontellasubstriata (Wray),1953—TXS2/G2 Localities:Wanda’sWaterfallCave,IllinoisCaverns Thisspeciesischaracterizedbyhavinganalspines,body tuberclesonheadtoAbd.5circulartosomewhat polygonalincrosssection,butformingridgesandvalleys onAbd.6;Ant.4with7–8bluntsensilla;and2clavate tenenthairsoneachleg.Inaddition,theindividualsfrom Illinoishavedorsalheadsetaec3presentandc2absent (arrowinFig.3D);3labialand7postlabialsetae (Fig.5A);chaetotaxyofTh.2–3asin S.shasta butwith N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 150 N JournalofCaveandKarstStudies, August2013

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Figure4.Chaetotaxyof Superodontella sp.1,shortarrowspointatsetaeabsentin S.substriata :(A)mesothorax;(B) abdomen4–5;(C)sternaofabdomen3–4,longarrowpointsanteriorly;(D)cuticlesculpturingandstructureofsetaeon abdomen5. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 151

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setam4absent(arrowinFig.4A);Abd.1–3chaetotaxy reducedto3anteriorand4posteriorsetae(Fig.5B);Abd. 4(Fig.5C)withp5clearlylongerthanp6;Abd.5with2 anteriorsetae(Fig.5C);sternaofAbd.3–4with3anterior and2posteriorsetae(arrowinFig.4Cpointsatseta absent). Thisisaforestleaflitterspeciesoriginallydescribed fromIllinois,andithasbeenreportedfromChampaign, Clark,Jackson,Logan,andPopecounties.Additional recordsforthespeciesincludeNorthCarolina,Connecticut,Quebec(Therrienetal.,1999),andBritishColumbia. ThisisthefirstrecordofthisspeciesfromcavesinIllinois. Superodontella sp.1—TX?S1?/G1? Locality:Wanda’sWaterfallCave Thesingleindividualcollectedissimilarto S.shasta in lackinganalspines,inhavingbodytuberclesangulateor polygonalincrosssectiondorsallyonheadandbody, althoughontheposteriorpartofAbd.5thetubercles becomeroundedorcircular(Fig.4D),inhavingrelatively shortungues,andinthepresenceof6bluntsensillaonAnt. 4.However,itdiffersfrom S.shasta inhavingarelatively bluntmouthcone(Fig.3C),inhaving1,2,2clavatetenent hairsonthepro-,meso-,andmetathoraciclegs,respectively,andinthatthelongestposteriorsetaeonAbd.4 Figure5.Chaetotaxyof Superodontellasubstriata :(A)labiumandpostlabium;(B)abdomen1;(C)abdomen4–5. Onychiurus pipistrellae n.sp.:(D)sensillaofthirdantennalsegmentsenseorgan. N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 152 N JournalofCaveandKarstStudies, August2013

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areapicallyexpandedtoweaklybluntandserrate.Other potentiallyinformativecharactersobservedintheindividualfromIllinois,butunknownin S.shasta are6labialand 7postlabialsetae(Fig.3C);dorsalheadsetaec2andc3 present(Fig.3D);Th.2–3setaea2anda6absent,andm5 present(Fig.4A);Abd.1–3with4anteriorand5posterior setae(Fig.3E);Abd.4withsetaa3displacedposteriorly, andp6aslongasp5,butclearlythicker(Fig.4B);Abd.5 with3anteriorsetae(Fig.4B);andsternaofAbd.2–4with 3,4and2setae,respectively(Fig.4C,onlyAbd.3–4 shown). Superodontella sp.2—TX?S1/G1? Locality:Wanda’sWaterfallCave ThesingleindividualcollectedinWanda’sWaterfall Caveisajuvenileandappearstorepresentanewspecies characterizedbyaveryshortfurculawithonlytwodental setae,PAOwiththreearms,tenenthairsonalllegs acuminate,analspineswelldeveloped,bodytubercles roundorovalincrosssection,andAnt.4with7welldifferentiatedbluntsensillaanddorsalheadrowconly withsetac3present.Thespeciesisdarkblueanddoesnot showcharacterstypicallypresentincave-adaptedspecies. N EANURIDAE Pseudachorutesaureofasciatus (Harvey),1898—TXS5/G5 Locality:StemlerCave Thiswidespreadspeciesisfoundinforestleaflitter fromBritishColumbiaandCaliforniatoPennsylvaniaand Florida.InIllinoisthespecieshasbeenreportedfrom Calhoun,Johnson,andUnion(GuthrieCave)counties. Pseudachorutesaureofasciatus isalsoknownfromHunter’s Cave,JacksonCounty,Iowa. Pseudachorutes sp.—TX?S1/G1? Locality:Wanda’sWaterfallCave ThesingleindividualcollectedinWanda’sWaterfall Caveappearstorepresentanundescribedspecies.Thisis ablackspecies,withoutmorphologicaladaptationsto subterraneanlife.Thisislikelyasurfaceleaflitterform accidentallyfoundinsidethecave.Afterthisindividualwas identifiedasapotentialnewspecies,severalcollectingtrips weremadetoWanda’sWaterfall,butnoadditional specimenswerefound. Vitronuragiselae Gisin,1950—TXIN? Locality:Wanda’sWaterfallCave Thisisprobablyanintroducedspeciesnowcommonin moistwoodchipbedsincentralIllinois. Vitronuragiselae wasfirstreportedfromNorthAmericabyChristiansen andBellinger(1998),althoughthespecieswasknowntobe establishedonthecontinentbeforethat(RichardSnider, MichiganStateUniversity,personalcommunication).In IllinoisthespecieswaspreviouslyobservedinChampaign Countybytheseniorauthor.Thisisthesecondrecordof V.giselae fromNorthAmericancaves;thespecieswas alreadyreportedfromSwampRiverCave,Tennessee. Althoughthisspecieshasareducednumberofeyesand iswhitewhenpreservedinalcohol,itisnotasub-or eutroglophile.Livingspecimensarebrightorangeandthe reducednumberofeyesischaracteristicofthelineage. O NYCHIURIDAE Onychiuruspipistrellae n.sp.—TB?S1?/G1? MaterialExamined:Holotype,INHSCollectionNumber:551,651;USA,IL,MonroeCo,BatSumpCave,6.6mi WSWofRedBud,handcollectedoncavefloor,swarming aroundcheesebait,3–5November2009,SJTaylorandFN Soto-Adames;Female,slidemounted.Theslidewiththe holotypealsoincludesoneindividualof Heteraphorura subtenua (Folsom)andoneindividualof Folsomiastella ChristiansenandTucker.Paratypes,INHSCollection Number551,652-56;9malesandfemalesonslidesand 69adultsandjuvenilesofbothsexesinalcoholwithsame collectioninformationasholotype.Additionalparatypes, INHSCollectionNumber551,657-60;USA,IL,St.Clair Co,StemlerCave,2.7miNEofColumbiaIL,hand collectedoncaveflooranddrippool,28–30September 2009,SJTaylorandFNSoto-Adames;3mountedonslides and5inalcohol. Etymology:Theepithetreferstothetypelocality,Bat SumpCave. Description:Largestmaleandfemale1.7mmand2.2 mmlong,respectively.SubapicalsensillaofAnt.4 smooth,roundedandrestinginashallowpit;subapical invaginationasis O.reluctus ;basalsensillalongerthan subapical.Ant.3senseorganwithfiveguardsetae,five papilla,tworods,andtwosmoothsensilladistally expandedandbent,withmiddlegroove(Fig.5D);basal sensillashorterthanbasalsensillaonAnt.4.Postantennal organwith12–15compoundvesicles.Apicalsetaofouter maxillarylobeunilaterallycoarselyserrateonapicalthird, sublobularplatewithtwoappendages.Labialpalpwith6 proximalsetae;thickened-bluntterminalsetapresentonly onpapillaAandB;papillaEwith3guardsetae.Dorsal pseudocelli32/033/33353;ventral2/000/1112;subcoxae1– 3with222pseudocelli.Ventralparapseudocelli00/000/ 11100?(viewofthedorsalanalvalveisobstructedinall individualsexaminedandthepresenceorabsenceofa parapseudocelluscannotbeestablished).Ventralthoracic setaeabsent.Prothoraxwitheightsetae.DorsalchaetotaxyofTh.2andAbd.1asinFigures6Aand6B;tipof macro-andmesosetaeacuminateandwide(Fig.6C); medialsetaabsentonAbd.1(Fig.6B)andpresenton Abd.4(Fig.6E);Abd.VIsetaa0absentandp0present, spinesstronglyconical.Subcoxae1–3with555setae. Ventraltubewith6–7 + 6–7setae.Furculascarswith4 shortposteriorsetaeinasinglerow(Fig.6D),homology ofotherfurcula-associatedsetaedifficulttoassess,astheir arrangementvariesasymmetricallywithinindividuals. Dorsalanalvalvewithsetaea0andc0present,setab0 absent(Fig.6F);lateralanalvalveswith4hr,6inner,4 medialand5outersetae(Fig.7A).Pro-andmetathoracic F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 153

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tibiotarsiwith9,8,1setaeinwhorlsA–C,respectively; mesothoracictibiotarsuswith9,8,2setaeinwhorlsA–C (Figs.7B,7C);setaMpresent;tenenthairacuminate. Unguiswithouterteeth,innerteethabsent.Unguiculus withoutbasallamellaandaslongas,tomarginallylonger than,inneredgeofunguis. Remarks:Table1summarizesdifferencesbetweenthe newspeciesandotherNorthAmericanmembersofthe O. Figure6. Onychiuruspipistrellae n.sp.,dotsrepresentsetaeandcirclespseudocelli,longarrowspointanteriorly:(A)dorsal chaetotaxyofmesothorax;(B)dorsalchaetotaxyofabdomen1;(C)typicalmacroseta,arrowpointsatdetailofapicalend;(D) furculascarandassociatedsetae,arrowpointstoanteriorend;(E)chaetotaxyofmedialsectionofabdomen4,arrowpointsat detailofpseudocelliornamentation;(F)upperanalvalve. N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 154 N JournalofCaveandKarstStudies, August2013

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Figure7.Chaetotaxyof Onychiurus spp.,dotsrepresentregularacuminatesetae,dotdiameterisroughlyproportionaltoseta length:(A) O.pipistrellae n.sp.,analvalves;(B) O.pipistrellae n.sp.,mesothoracictibiotarsus,anteriorview,inneristoward topoffigure,arrowindicatessetaabsentfrompro-andmetathoracictibiotarsi;(C) O.pipistrellae n.sp.,mesothoracic tibiotarsus,posteriorview,inneristowardtopoffigure;(D) O.wilchi ,abdomen1,syntypedepositedattheIllinoisNatural HistorySurvey;(E) O.wilchi asabove,butprothorax,circlerepresentspseudocellus;(F) Pseudosinellaaera ,dorsalhead chaetotaxy,dotsandopencirclesrepresentmicro-andmacrosetae,respectively. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 155

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reluctus speciescomplex.Mostindividualsof O.pipistrellae n.sp.willkeyoutto O.steinmanni Pomorski,Furgol,and Christiansen,2009inPomorskietal.(2009),butthenew speciesiseasilydistinguishedfrom O.steinmanni bythe numberofpseudocellionTh.2–3,thenumberofsetaeon ventraltube,thenumberofPAOvesicles,and,apparently, thenumberofproximalsetaeonthelabialpalp.Thenew speciesismostsimilarto O.reluctus Christiansen,1961, fromwhichitdiffersinlackingpseudocellionTh.1,the numberofsetaeontheventraltube,theshapeofthe macrosetaltip,andpossiblythenumberofPAOvesicles. AnadultfemalefromBatSumpcavehas1 + 1pseudocelli onTh.1,butitretainsthelownumberofsetaeonthe ventraltube,theshapeofthemacrosetae,andaPAOwith only13vesicles. Onychiuruspipestrellae n.sp.wascollectedabout50to 60kmsouthwestofthetypelocalityof O.wilchi Wray, 1950.FollowingWray’s(1950)originaldescription,the twospeciesshouldbeeasytodistinguishbythecombined absenceofdorsalpseudocellionTh.1andAbd.1in O. wilchi andthenumberofPAOvesicles.However,the syntypesof O.wilchi depositedattheIllinoisNatural HistorySurveyexaminedcarry1and3pseudocellionTh. 1(Fig.7E)andAbd.1(Fig.7D),respectively.This suggeststhat O.wilchi isaseniorsynonymofeither O. reluctus or O.pipistrellae n.sp.Unfortunately,the charactersneededtodecidebetweenthealternatives(i.e., numberofsetaontheventraltube,numberofvesicleson thepostantennalorgan,andshapeofthetipofthe macrosetae)arenotvisibleinthetypesstudied,andthe statusof O.wilchi remainsunresolveduntilfreshtopotypicalmaterialisobtained. Thedistributionof O.pipestrellae isunclear.The populationsfromBatSumpandStemlercaveswere previouslysampledandidentifyas O.reluctus (Lewis etal.,2003).However,theredescriptionof O.reluctus by Pomorskietal.(2009)makesitclearthatthepopulation inBatSumpandStemlerCaverepresentanewspecies.It ispossiblethatotherrecordsof O.reluctus incavesfrom southernIllinois(e.g.,FogelpoleCave,HiddenHand Cave,FultsSaltpeterCave),Missouri,andKentucky maybereferabletothenewspecies.Thefiveother describedspeciesinthe O.reluctus speciescomplexare foundincavesinVirginia,Indiana,Wisconsin,Iowa,and Colorado. Thalassaphoruraencarpata (Denis),1931—EUS2/G5 Localities:BatSumpCave,StemlerCave ThiscommonspeciesiswidespreadacrossNorth America. Thalassaphoruraencarpata isknownfrom Cumberland,Jackson,Johnson(FirestoneCreekCave), Piatt,andWaynecountiesinIllinois.Thisspeciesis relativelycommoninsubterraneanhabitats,asithasbeen reportedfromcavesinMissouri,Indiana,Texas,and Hawaii. Table1.Diagnosticcharactersforspeciesinthe Onychiurusreluctus speciesgroup.Charactersfor O.wilchi takenfromWray(1950)andfromthreeindividualsof thesyntypicseriesdepositedattheIllinoisNaturalHistorySurvey(INHS).CharactersforallotherpreviouslynamedspeciesfollowPomorskietal .(2009). Species Numberof PAOVesicles Numberof DorsalPseudocelli Numberof VentralPseudocelli Shapeof Macrosetaetip NumberofSetae VentralTube NumberofSetae SubcoxaeLegs1–3 Unguicular Lamella O.pipistrellae n.sp.12–1532/033/333532/000/1112broadlyacuminate6–7555absent O.reluctus Christiansen andBellinger 15–1632/133/333532/000/1112truncate10555absent O.furcisetosus Pomorski, FurgolandChristiansen 16–1732/133/333532/000/1212bifurcate8555absent O.nathanieli Pomorski, FurgolandChristiansen 1532/033/333542/000/1112acuminate6455present O.reluctoides Pomorski, FurgolandChristiansen 14–1532/133/333542/000/1212acuminate9555absent O.steinmani Pomorski, FurgolandChristiansen 16–1732/022/333532/000/1112acuminate9555absent O.wilchi Wray(Fig.18in originaldescription) 18–2032/022/02233…………absent O.wilchi INHScotypes…32/133/3????…acuminate?……absent N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 156 N JournalofCaveandKarstStudies, August2013

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Heteraphorurasubtenua (Folsom),1917—EUS2/G5 Locality:Wanda’sWaterfallCave Thisispartofacomplexthatincludesthreeother species( H.bima H.casa ,and H.tala ).Theactual distributionof H.subtenua isunclear,asoldidentifications mighthaveconflatedtheidentityofallfourforms. ConfirmedrecordsindicatethespeciesrangesfromAlaska andBritishColumbiatoMaineandNorthCarolina, althoughitappearstobeabsentintheregionbetween Iowa-Missouri-ArkansasandBritishColumbia.InIllinois thespecieshasbeenreportedfromAlexander,Champaign, Coles,Jackson,LaSalle,Union,Vermillion,andWashingtoncounties. Heteraphorurasubtenua hasbeenpreviouslyreportedfromcavesinAlaska,Illinois,Indiana, NorthCarolina,Texas,andWestVirginia. T ULLBERGIDAE Mesaphorurasilvicola (Folsom),1932—TXS1/G5 Localities:Wanda’sWaterfallCave,BatSumpCave Mesaphorurasilvicola iswidespreadinNorthAmerica andcommoninsurfaceleaflitter.InIllinois, M.silvicola is knownfromJackson,Monroe,andVermillioncounties. ThespecieswaspreviouslyreportedfromacaveinIndiana. I SOTOMIDAE Hemisotomathermophila (Axelson),1900—TXS1?/G5 Localities:Wanda’sWaterfallCave,StemlerCave Thisspeciesiscommonlyfoundonsurfaceleaflitter acrosstemperateandtropicalregionsoftheworld(Potapov, 2001;MariMuttandBellinger,1990),andinIllinoisitwas previouslyknownfromChampaignCounty.Thisappearsto bethefirstrecordforthisspeciesfromIllinoiscaves. Desoriatrispinata (MacGillivray),1896—EUS5/G5 Localities:Wanda’sWaterfallCave,IllinoisCaverns Thiscommonsurfacespeciesiswidelydistributedacross temperateregionsoftheNorthHemisphere(Potapov,2001) andrelativelycommonincaveleaflitterinNorthAmerica. TheindividualsfromtheSalemPlateauhavethreeinsteadof fourlabralpapillae,acharactersharedwithsomepopulationsfromMissouricaves(ChristiansenandBellinger, 1998).ThelabralcharactermaydefineanisolatedpopulationrestrictedtotheSalemPlateauregiononbothsidesof theMississippiriver. Folsomia cf. bisetosa Gisin,1953—TXS1/G5;NewIllinois Record Locality:StemlerCave ThesingleindividualcollecteddeepinStemlerCave agreeswiththedescriptionof F.bisetosa providedby Fjellberg(2007)inallcharactersexceptinhavinglong unilaterallyserratemacrosetaeonallsegments(macrosetae aresmoothin F.bisetosa )andinhavingdorsalmanubrial setaml1(ml1absentin F.bisetosa ).Thismaybethesame formidentifiedas F. cf. bisetosa fromIndianabyWaltzand Hart(1996). Folsomiabisetosa isdistributedacrosstheOld Worldarcticandsubarcticregions,anditspresenceinan Illinoiscaveseemsunlikely.Additionalmaterialisneeded toconfirmtheidentityofthespecies. Folsomiacandida Willem,1902—EUS5/G5 Localities:Wanda’sWaterfallCave,IllinoisCaverns,Bat SumpCave,StemlerCave Thisisacommonlitterandsoilspecieswidespreadin NorthAmericancaves. Folsomiaprima Mills,1931—EUS2/G5 Localities:IllinoisCaverns,BatSumpCave Thiscommonsurfaceleaflitterspeciesisoftenfoundin caves.InIllinoisthespecieshasbeenreportedfromcaves inHardinCountyandfromsurfaceleaflitterin Champaign,Vermillion,andWilliamsoncounties. Folsomiastella ChristiansenandTucker,1977—EUS5/G5 Localities:BatSumpCave,StemlerCave ThisspeciesisdistributedacrosstemperateNorth AmericaandEurasia(Potapov,2001).InNorthAmerica, F.stella isoftenfoundincaves.InIllinois,thespeciesis previouslyknownfromAlexander,Champaign,Monroe, andVermillioncounties. Folsomiavariabilis Fjellberg,1984—TXS1/G5;New IllinoisRecord Localities:Wanda’sWaterfallCave,StemlerCave Thisisthefirstrecordfor F.variabilis fromIllinoisand fromcaves.Thespecieswasoriginallydescribedfrom populationsfoundinmossesat11,000ft(3,400m)of elevationinColorado(Fjellberg,1984). Isotomiellaminor Scha ¨ffer,1896—EUS5/G5 Locality:WizardCave Thisspeciesiscommoninsurfaceandcavehabitats throughoutNorthAmerica. Parisotomanotabilis (Scha ¨ffer),1896—EUS5/G5 Localities:Wanda’sWaterfallCave,IllinoisCaverns, HiddenHandCave Parisotomanotabilis isacommonsurfacespecies, widelydistributedacrosstemperateregionsoftheNorthernHemisphereandprobablythemostcommonspeciesof springtailinsurfaceleaflitterinNorthAmerica.The speciesisoftenfoundincaves. Proisotomasepulcralis Folsom,1902—TXS1/G1;New IllinoisRecord Locality:StemlerCave ThisisthefirstrecordfromIllinoisandfromcavesfor thisuncommonspecies. Proisotomasepulcralis waspreviouslyknownonlyfromWashingtonD.C.,Michigan,and Pennsylvania. T OMOCERIDAE Lethemurusmissus Mills,1949—TBS2/G2 Locality:PautlerCave ThistroglobiontiswidespreadincavesintheSalem Plateau. Lethemurusmissus wasoriginallydescribedfrom F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 157

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JerseyCountyIllinois,butisnowknowntooccurin Kentucky,Missouri,Alabama,Indiana,Virginia,and Colorado. Pogonognathellusflavescens (Tullberg),1871speciescomplex—EUS5/G5 Localities:WizardCave,PautlerCave,Wanda’sWaterfall Cave,SpiderCave,IllinoisCaverns,BatSumpCave, HiddenHandCave,StemlerCave Speciesinthe P.flavescens complexarethemost commonmembersofthefamilyTomoceridaeinsurface leaflitterandincavesinNorthAmerica.Membersofthe P. flavescens complexappeartobewidespreadinNorth AmericaandEurope.Untilrecently,allformsinthe complexinNorthAmericawereassignedtothenominal species,althoughthelargeamountofmorphological variationreportedfromthroughoutitsrangesuggestedthat itrepresentedaspeciescomplex(Christiansen,1964). Recently,Felderhoffetal.(2010)andParketal.(2011) concludedthat,basedonmolecularevidence, P.flavescens doesnotoccurinNorthAmericaandallormostforms previouslyidentifiedassuchareendemictoNorthAmerica. Mostspeciesinthecomplexhavebeenidentifiedonlyby usingmoleculardata,andjustahandfulofthemhavebeen diagnosedusingmorphologicalcharacters.Thus,the identityoftheformscollectedintheSalemPlateaucaves willremainunclearuntiltheirDNAcanbestudied. Pogonognathellusnigritus (Maynard),1951—TXS5/G5?; NewIllinoisRecord Localities:SpiderCave,IllinoisCaverns,BatSumpCave Thisisacommonsurfaceleaflitterspeciesinwooded areasoftheSalemPlateau.Theactualdistributionof P. nigirtus inNorthAmericaisnotknown,becauseinthepast ithasbeenincludedaspartofthevariationof P.elongatus Maynard(Felderhoffetal.,2010),hencethequestionmark forglobalranking.Thisisthefirstrecordforthespecies fromcaves. O NCOPODURIDAE Oncopoduraiowae Christiansen,1961—TB/EUS2/G2 Localities:WizardCave,Wanda’sWaterfallCave,Illinois Caverns,HiddenHandCave,StemlerCave Thisisthemostcommonspeciesof Oncopodura found inIllinoiscaves.Recentcollectionsofthisspecieson surfaceleaflitter(IL:MonroeCo.,Valmeyer,SaltLick Point,N38.30644W90.30475,leaflitterbetweenrock outcropsonblufftop,13May2011,F.N.Soto-Adames) indicatethespeciescanmovefreelyabovegroundunder appropriateconditions.Thespecieshasbeenpreviously reportedfromotherlocalitiesinMonroeCounty,Illinois, andincavesinIowa,Missouri,andSouthDakota. E NTOMOBRYIDAE Pseudosinellaaera ChristiansenandBellinger,1980—EU S2/G5 Locality:WizardCave Complementtothedescriptionof P.aera basedon individualscollectedinWizardCave:Freshlykilled individualsuniformlylightblue,pigmentmoreintenseon eyepatch.Scalespresentonlyonhead,trunk,andcoxae. ApicalbulbofAnt.4absent;subapicalsenseorgan‘‘Y’’ shaped(Fig.8A),appearingcapitateinsideviewandin smallindividuals.SenseorganofAnt.3withmainsensilla (i.e.,numbers2and3inChenandChristiansen,1993) slightlybent,laterallyexpandedwithdensecentralrachis (Fig.8B);sensilla1and4longer,andsensillum5shorter, thansensilla2–3;atleastonelenticularorganpresent.Eyes 2 + 2.Dorsalheadchaetotaxyasin(Fig.7F):macrosetae A0,A2andA3present;A1coarselyciliate,butnota macroseta;otherdorsalheadsetaenarrowandsmooth; M0present,insertedalmostbetweenA3;rowSwith4 + 1 setae,S0inlinewithS3;rowPawith3setaeand bothriotrixPa6,Pa5amicroseta.Prelabralsetaeciliate, allothersetaesmooth,labralintrusionandpapillaenot seen.Maxillarcapitulumwithfourteeth.Sublobularplate ofoutermaxillarylobewiththreeseta-likeappendages; subapicalandapicalsetaesmoothandsubequal.Labial papilaEwith4guardsetae,lateralappendagestraight, bluntandreachingtipofpapilla.Labialpalpproximal setaeYandZsubequal.Labialtrianglechaetotaxywith anteriorsetaesmooth,posteriorsetaecoarselyciliate, exceptforr,whichisacuminate,shortandsmooth; formulaas M1M2r EL1L2A1-5(Fig.8C).Allpostlabial setae,exceptL2andO1,coarselyciliate;columnsI,C,E,L andOwith4,1,2,3,2setae,respectively;setaeL2andO1, smoothandacuminateasr;oneadditionalsetasimilartor externaltocolumnO.Allpleuralandperistomalsetae coarselyciliate,pss1-2bothriotrix-like.Trunkmacrosetae formula00/0100 + 2.Mesothoracichoodnotdeveloped, collarformedby2–3rowsofacuminatetoweaklyapically bentmacrosetae.Mesothoraxnotpolychaetotic.Metathoraxwithoutp2p,butotherwisewithafullcomplementof setae;setaep2lessthan2 3 thelengthofa3.Abd.1with10 posteriorsetae(Fig.8D)arrangedinasinglerow,butnot evenlydistributedalongrow,setaa3insertedverycloseto m3anda5tom4;a6present,notpairedtosensillaas.Abd. 2(Fig.8E)withmi,ml,Lm,andLlfan-shaped;a2,a3, a2p,m4,andp5pciliate,allothersetaeeitherdenticulate orsmooth;a3insertednearandreachingas;m3andm5 macrosetae,a6present.Th.3(Fig.8F)witha2,a6,am6, andallsupplementarysetaefan-shaped;a3notreachingas; ashalfthelengthofm3;d2present,a7insertedanteriorto andreachingam6;m7smoothmicrosetainsertedinrow withp5;p7aciliatemicroseta;m7enlarged.Bothriotrichal complexofAbd.4(Fig.9A)withsetaspresentandfanshaped;a,m,D1,Pi,andPefan-shaped;C1p,T3,andD1p ciliate.OtherchaetotaxyofAbd.4asinFigure9B:B5and B6macrosetae,B5insertedjustanteriororontheline betweenA5-C2;C1asmoothmicrosetae;D3,E2,E3,F1, andF3largemacrosetae;T6,T7,D2,andE1small macrosetae;F2amicrosetae;microsetaeposteriortoE3 present;posteriorsetae3 + 3.Trochanteralorganwith23 N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 158 N JournalofCaveandKarstStudies, August2013

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Figure8. Pseudosinellaaera :(A)supapicalorganoffourthantennalsegment;(B)senseorganofthirdantennalsegmentand associatedsensilla;(C)labialtriangle.Tergalchaetotaxy,dots,opencirclesandcrossedcirclesrepresentmicro-and macrosetaeandpseudopores,respectively;(D)abdomen1;(E)abdomen2;(F)abdomen3. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 159

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setaeinfemales,15inthesingleadultmalestudied,and4 inonesmalljuvenile.Internalfaceofallfemorawith5 conicsetae.Alltibiotarsiwithoneposteriorbluntor acuminatemacrosetainsertednearbasalthirdofsegment. Tenenthairweaklycapitate,appearingacuminateorblunt atlowmagnification(Figs.9C,D).Proportionofunguiculus:tenenthair:posteriorsmoothsetaonhindlegsas 1:1.3:1.3.Unguiculuslanceolate,posteriorlamellaserrate onforelegs,apparentlysmoothonhindlegs.Unguis (Figs.9C,D)with4internalteeth;basalteethnotaligned, Figure9. Pseudosinellaaera ,dots,opencircles,trianglesandcrossedcirclesrepresentmicro-andmacrosetae,fan-shaped setae,andpseudopores,respectively:(A)abdomen4bothriotrichalcomplex;(B)completechaetotaxyofabdomen4;(C) prothoracicclawcomplex;(D)metathoracicclawcomplex. N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 160 N JournalofCaveandKarstStudies, August2013

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onetoothslightlylargerthanother,basaltoothendingat < 46%(range43–49%;n 5 3)ofinnerclawlength;distal unpairedtoothaslargeasbasalteethandendingat < 74% (range70–77%;n 5 3)ofinnerclawlength;externalteeth conspicuous.Anteriorfaceofventraltubewith9to14 setae,4oftheminsertedalongventralgroove,with1distal marginmacroseta;lateralvesicleswith4smoothand5to 10stronglyciliatesetae;distalmarginofposteriorfacewith 3medialand3or4lateralsetae,basalventralchaetotaxy notclearlyseenonanyspecimen,butapparentlywith13 setae(5 + 5and3unpaired).Allmanubrialsetaeciliate; dorsalmanubrialplatewith2internaland5external, coarselyciliatesetae.Denstubercleabsent.Mucronalteeth subequal;mucronalspinewithminutebasaltooth. Remarks:TheindividualsfromWizardCavediffer fromtheoriginaldescriptionofthespeciesonlyinhaving dorsalheadmacrosetaeA3present. Pseudosinellaaera is verysimilarto P.argentea Folsom, P.flatua Christiansen andBellinger,and P.granda ChristiansenandBellinger, butitiseasilydistinguishedfromallbythepresenceof2 + 2 eyes.Inaddition,itcanbedistinguishedfrom P.flatua by thepresenceofciliateprelabralsetae,from P.argentea by theabsenceofsetam4ionAbd.2,andfrom P.granda by thepresenceofonlyoneheadmacrosetaA0,bythe presenceofshortacuminatelabialsetar,andbyhavinga ciliatea3onAbd.2.Additionaldifferencesbetweenthe fourspeciesarelistedinTable2. InIllinois Pseudosinellaaera hasbeenpreviously reportedfromcavesinGallatinandJohnson(Firestone CreekCave)counties. P.aera appearstohaveamostly southerndistribution;theloc alitiesinIllinoisareator nearthenorthernmostlimitofthespecies.Thisspecies doesnotshowstrongadaptationstocavehabitat,but almostallrecords,fromMexicotoIllinois,arefrom caves. Pseudosinellaargentea Folsom,1902—EUS2/G5? Localities:SpiderCave,IllinoisCaverns Ascurrentlycircumscribed,thisspeciesshowsconsiderablemorphologicalvariationthroughoutitsdistribution,suggestingitlikelyrepresentsaspeciescomplex (ChristiansenandBellinger,1998).Thespecimensfrom theSalemPlateauhave4 + 4posteriorsetaeonAbd.4 (Fig.10A),andthetenenthaironalllegsisacuminate. Thisisconsistentwiththepopulationof P.argentea reportedfromMaidenCave,WestVirginia(Soto-Adames, 2010). ThisisacommoncavespeciesinIllinois.Ithasbeen previouslyreportedfromCaroll,Hardin(Brown’sHole Cave),andMonroe(FultsSaltpeter,Fogelpole,Spiderand Wanda’sWaterfallcaves)counties.Thespeciesalso appearstobecommonincavesinKentucky,Missouri, Arkansas,Indiana,Alabama,Tennessee,Washington, Virginia,WestVirginia,Pennsylvania,andConnecticut. SurfacepopulationsinIllinoishavebeenreportedfrom Champaign,Clark,Edgar,Johnson,andUnioncounties. Coecobryatenebricosa (Folsom),1902—EUS5/G5(IN?) Localities:WizardCave,PautlerCave,Wanda’sWaterfall Cave,IllinoisCaverns ThisisaspeciescommonlyfoundincavesinNorth America.PreviousreportsfromIllinoiscavesincludeRose HoleandPautlerCave,bothinMonroeCounty. ThisspecieswasoriginallydescribedfromtheWashington,D.C.,area,butcurrentlyitisknowntooccur aroundtheworldinprotectedhabitatssuchasgreenhouses andcaves(ChenandChristiansen,1997).Thegenus Coecobrya isalmostexclusivelyEastAsianindistribution (ChenandChristiansen,1997),anditislikelythat C. tenebricosa representsanearlyintroduction,duringhistoric times,intoNorthAmerica. Entomobrya sp. Locality:WizardCave Thesingleindividualcollectedisinanearlyinstarand notidentifiabletospecies. Homidiasocia Bo ¨rner,1909—TXIN Locality:SpiderCave Thisisanintroducedspecies.Almostallspeciesof Homidia arerestrictedtoEastAsiaandOceania;onlytwo ( H.socia and H.sauteri )areknownfromNorthAmerica. Theoldestrecordof H.socia inNorthAmericaappearsto befrom1970,fromGeorgia(K.Christiansen,Collembola recordsdatabase).Thehistoricalcollectionofspringtailsat theIllinoisNaturalHistorySurveythatgoesbacktothe secondhalfofthe1800s,doesnotincluderepresentatives ofthisspecies,despiteitnowbeingthemostcommonform foundingrassesgrowingalongcountryroadsinChampaignCounty. Homidiasocia wasfirstnoticedinacavein JohnsonCounty,southernIllinois,in1973(Christiansen andBellinger,1980,1998;K.Christiansen,Collembola recordsdatabase)andinChampaignCountybythesenior authorin1988.Thespeciesisalsoknownfromcavesin HarrisonandCrawfordcounties,Indiana. N EELIDAE Megalothoraxminimus (Willem),1900—EUS5/G5 Locality:Wanda’sWaterfallCave Thisisacommonsurfaceleaflitterspeciesfrequently foundincaves.ThespecieshasbeenrecordedfromGrundy, Lawrence,andWashingtoncountiesinIllinois.Thisappears tobethefirstrecordforthespeciesfromIllinoiscaves. Megalothoraxtristani (Denis),1933—TXS1/G1 Locality:WizardCave Thisiseitherararespeciesorithasbeengenerally confusedwiththemorewidelydistributed M.incertus Megalothoraxtristani waspreviouslyreportedfromIllinois byBonet(1948). Neelidesminutus (Folsom),1901—EUS5/G5 Locality:Wanda’sWaterfallCave Thisisacommonsurfaceleaflitterspeciesoftenseenin cavesamples.InIllinoisthespecieshasbeenpreviously F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 161

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Table2.Comparisonof Pseudosinellaaera andP. argentea fromSalemPlateauto P.flatua and P.granda. Character Species(Caves) P.argentea (SpiderCave andIllinoisCaverns) P.aera (WizardCave) P.grandaP.flatua Eyenumber0200 HeadmacrosetaA01121 Prelabralsetaeciliateciliateciliatesmooth Sizeoflabialsetam1&m2 a M1 < M2 M1 < M2 M1 < M2 m1 M2 Labialsetarsmooth,thinwalledblunt microseta smoothacuminatemicrosetaabsentorsmoothconicreducedsmoothacuminate microseta Headventralgroovesetae4ciliate4ciliate4ciliate3smooth1ciliate Abd.2setaa3ciliateciliatesmoothsmooth Abd.2setaa2pciliate& < a2weaklyciliate& a2ciliate& a2ciliate& a2 Abd.2seta4mipresentabsentabsentabsent Abd.4posteriorsetae43…… Tenenthairacuminateweaklytruncateacuminateacuminate Ventraltubeanteriorsetae99–149 b 11–13 Ventraltubeposteriorsetae (total/distalmargin) 19/922–24/9–1113/714/? Ventraltubelaterodistalsetae7–8;3–4ciliate10–14;5–10ciliate8–9;4–5ciliate11–13 a Underlineandwithoutunderlineindicatethesetaeisciliateorsmooth,respectively.Upper-andlower-case‘m’correspondstomacro-ormicroseta e,respectively. b Theoriginaldescriptionof P.granda statesthereare6–7setaeontheanteriorfaceoftheventraltube(ChristiansenandBellinger,1996),buttheirfigure46shows9setae. N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 162 N JournalofCaveandKarstStudies, August2013

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reportedfromCalhoun,Cook,Gallatin,Jackson,Lake,La Salle,Lawrence,andWaynecounties. S MINTHURIDIDAE Sphaeridiaserrata (FolsomandMills),1938—TXS1/G3 Locality:Wanda’sWaterfallCave Thisisanuncommonspeciespreviouslyreportedfrom HerodandPopecountiesinIllinois.Thisisthefirstrecord ofthisspeciesfromIllinoiscaves. Sminthurides sp.—TXS1/G1 Locality:BatSumpCave Figure10. Pseudosinellaargentea ,dots,andopencirclesrepresentmicro-andmacrosetae,respectively:(A)posteriorchaetotaxyof abdomen4. Pygmarrhopalitesfransjanssens n.sp.,dots,andopencirclesrepresentmicrosetaeandspine-likesetae,respectively: (B)detailofverticalchaetotaxyofthehead,leftside;(C)generalch aetotaxyofface;(D)subapicalsen sillaoffourthantennalsegment. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 163

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Thisspecieshasauniquecombinationofcharactersnot seeninotherNearcticspecies.Thisisnotacave-adapted form.Onemaleandonefemalewereseen,andformal descriptionofthespeciesmustawaitthestudyoffurther material. A RRHOPALITIDAE Pygmarrhopalitesfransjanssens n.sp.—TB/EU?S1/G1? MaterialExamined:Holotype,INHSCollection Number551,634;USA,IL,St.ClairCo.,WizardCave, Dupo,nearFallingSpring,adultfemale,slide-mounted, collectedinpitfalltrapintwilightzone,15–17June2009, SJTaylor,FNSoto-AdamesandCAPhillips.Paratypes, INHSCollectionNumber551,635-36;2slide-mounted adultfemalesonindividualslidesand8otherindividualsof undeterminedsexinalcoholwithsamecollectionlocality asholotype;2paratypesofundeterminedsexcollectedwith anaspiratorunderrottinglogintwilightzone;8paratypes collectedinpitfalltrapsinbothdarkandtwilightzones. Etymology:ThisspeciesisdedicatedtoFransJanssens, DepartmentofBiology,UniversityofAntwerp,Antwerp, Belgium,inrecognitionofhiscontributionstospringtail taxonomythroughthedevelopmentofcollembola.org. Description:Largestindividual0.65mm.Background colorwhite,withdarkbrownscatteredoverhead,body,and allappendages.Ant.4with5–6subsegmentsinproportions as26–32:10–11:10–11:10–11:10–11:27–29.Ant.4subapical sensillacapitate(Fig.10D),eachshortpreapicalsubsegmentwith11setae.Ant.3withaweakbasalbulge (Fig.11F);senseorgan(Fig.11G)withtworodsensillain ashallowdepression,setaeApiandApeappearingthin walledandwithdrawnoutapices,butotherwisenormally acuminate,Aaiarodsensilla.Eyes1 + 1inadarkpatch. DorsalheadchaetotaxyasinFigures10BandC,with verticalsetaeM4,IL1-3,L1-2,andA3shortspine-like; M5absent.Apicalsetaeofoutermaxillarylobebifurcate, sublobularplatewiththreeappendages(Fig.11A).Labial palppapillaEwith3guardsetae.Latero-posteriorsetae (lp1)2.5–3.5 3 setac2(Fig.11B).Smallabdomenwithout denticlesorspines,chaetotaxyasinFigure10C:C1 bifurcate,C2-6andC8smooth,enlargedatbasebut withouttunica,C7andC9longbutnotbasallyenlarged; setaC2 < 1.3 3 B2andC3 < 2.2 3 D3;setaeD7–8absent, D9typicallylong;femaleappendage(Fig.11D,E) palmate,withsmoothstemandshallowbranches,sitting onacircularpapilla.Metatrochanterlongerthanitis wide(Fig.12A),with5setae.Allclawswith1innertooth (Figs.12B,C);dorsaltunicasmooth,coveringdistalhalf ofL1–2,and4/5ofL3.Unguiculusofalllegswithinner tooth,outstandingonL1butsominuteonL3astobe visibleonlyinsomeperspectives;apicalunguicular filamentsurpassinglengthofunguisonalllegs.Tenaculumwith2setae.Manubriumwith4 + 4dorsalsetae. Densdorsally(Fig.12D)with3inner(L),6dorsal(D1–2, ID1–4)and7external(E)setae;setaeL1–3,E1andE3 spine-like.Densventrallywith2unpairedsetae.Mucro withspatulateapex. Remarks:Thisspeciesischaracterizedbythecombined presenceof3sublobalsetaeonthemaxillarypalp,4 + 4 dorsalmanubrialsetae,apalmatefemaleappendage,and smallabdomenwithsetaeC1bifurcateandsetaeD7–8 absent.Table3listsvaryingcharactersforthegroupof specieswithpalmatefemaleappendage,smallabdomen setaC1bifurcate,andtwounpairedventralsetaondens. Pygmarrhopalitesfransjanssens n.sp.ismostsimilarto P. incantator n.sp.,fromwhichitdiffersinthenumberof sublobalsetaeonthemaxillarypalp,thenumberofguard setaeonlabialpapillaE,thenumberofdorsalmanubrial setae,theabsenceofD7–8onthesmallabdomen,andthe shapeofmetathoracicunguiculus. Thenewspecieskeysoutto P.principalis (Stach)in Bretfeld(1999),butthedifferencesbetweenthetwoforms arenotclearbecausethestateofsomecharactersinthe Europeanformremainindispute.Stach’s(1945)original descriptionof A.principalis doesnotmentionthe conditionofsmallabdomensetaC1.Vargovitsh(2009) pointsoutthatGisin’s(1947)Figure2showsC1as bifurcate,themucroaspointedinsteadofspoonshaped, andthesmallabdomensetaeinseriesCasbasally expandedinsteadofenlargedbutsimple.Inaddition, Gisin(1947)depictsheadverticalsetaeA3,IL1–2,andL1 asstronglyspine-likeanddistinctfromthoseinseriesM, andsetaM4isabsent.Fjellberg(1984)firstreported P. principalis fromNorthAmerica,buttheconditionofC1is notmentioned,andhisFigure9Cshowswhatappearsto beverticalheadsetaM5presentandM4absent.Fjellberg (2007)reportedthatFennoscandianpopulationsof A. principalis haveverticalcephalicM1-3,butnotM4–5, maxillarypalpwithtwosublobalhairs,andlabialpalp papillaEwith3guardsetae.Fjellberg(2007)doesnot mentionthenumberofmanubrialsetaeorthecondition ofC1.Vargovitsh(2009)describedanewsubspecies, P. principalisskelicus ,whichhediagnosedbasedonthe relativelengthofantennaetocephalicdiagonalandthe presenceofannulationsonAnt.4ofmales.Vargovitsh (2009)presentedthemostcompletedescriptionfor P. principalis sofarpublished,buthedoesnotmentionthe numberofsetaeonthemaxillarypalporlabialpapillaE. Inviewofthedifferencesbetween P.fransjanssens n.sp., P.principalisskelicus ,and P.principalis inthesenseof Gisin(1947)andFjellberg(2007),weoptedtodescribethe formcollectedatWizardCaveasanewspecies. Pygmarrhopalitesincantator n.sp.—EU/TB?S1/G1 MaterialExamined:Holotype:INHSCollectionNumber:551,638;USA,IL,St.ClairCo.,WizardCave,Dupo, nearFallingSpring,adultfemale,slide-mounted,collected inpitfalltrapintwilightzone,15–17June2009,SJTaylor, FNSoto-AdamesandCAPhillips.Paratypes,INHS CollectionNumber551,639-40;1adultfemaleand1adult N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 164 N JournalofCaveandKarstStudies, August2013

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malemountedonindividualslideswithsamecollection informationasholotype. Etymology:Theepithetofthenewspeciesrefersto WizardCave,thetypelocality. Description:Largestindividual1.0mm.Background colorwhite,withorangespotsscatteredoverhead,body, andallappendages.Ant.4with6subsegments(Fig.12E) inproportionsas36–37:11:10:10:10:25.Ant.4subapical sensillacapitateasin P.fransjanssens n.sp.,eachshortsub terminalsubsegmentwith11setae.Ant.3withoutbasal bulge(Fig.12F);senseorganasin P.fransjanssens n.sp., with2rodsensillainindependentshallowdepressions, Figure11. Pygmarrhopalitesfransjanssens n.sp.:(A)dorsalviewofoutermaxillarypalpandlobe;(B)bigabdomensetaec2 andlp1;(C)lateralviewofcompletechaetotaxyoffemaleanalvalves,insetdorsalviewofsetaC1fromanotherindividual; (D)dorsalviewoffemaleanalappendage;(E)lateralviewoffemaleanalappendage;(F)thirdantennalsegment;(G)detail ofapicalsenseorganofthirdantennalsegment. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 165

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setaeApiandApeappearingthinwalledandwithdrawn outapices,butotherwisenormallyacuminate,Aaiarod sensilla.Eyes1 + 1inadarkpatch.Dorsalheadchaetotaxy (Figs.13A,B)withsetaeA3,IL1,M4,andL1–3weakly spine-like;M5absent.Apicalsetaeofoutermaxillarylobe bifurcate(Fig.12G),butbasalspinecloselyappressed againstapicalsetaeandvisibleonlyinsomeperspectives; sublobularplatewithoneappendage.LabialpalppapillaE with4guardsetae(Fig.12H).Smallabdomenwithout denticlesorspines;chaetotaxyasinFigure13C:C1 bifurcate,C2–6smooth,enlargedatbasebutwithout extensions,baseofC7-8notenlarged;setaC2 < 1.2 3 B2 andC3 < 1.7 3 D3;setaeD7–D8present;femaleappendage (Fig.13D)palmate,withsmoothstemanddeepbranches, someofwhichoriginateclosetothemiddleofthestem, appendagesittingonacircularpapilla.Metatrochanter rectangular,with4anteriorand1posteriorsetae.Allclaws with1innertooth(Figs.13E,F),toothstrongestonL3; Figure12. Pygmarrhopalitesfransjanssens n.sp.:(A)metatrochanter;(B,C)pro-andmetathoracicclawcomplexes, respectively;(D)completechaetotaxyoffurcula,L = innercolumn,D = dorsal,ld = laterodorsal,E = outercolumn. Pygmarrhopalitesincantator n.sp.:(E)fourthantennalsegment;(F)thirdantennalsegment;(G)lateralviewofouter maxillarypalpandlobe;(H)ventralviewoflabialpalppapillaE,terminalsetaomitted. N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 166 N JournalofCaveandKarstStudies, August2013

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dorsaltunicaonalllegssmooth,coveringapicalthirdof clawonL1-2and4/5ofL3claw.UnguiculusofL1witha minuteinnertooth,unguiculusofL2–3toothless,apical unguicularfilamentsurpassinglengthofunguisonalllegs. Tenaculumwith2setae.Manubriumwith5 + 5dorsalsetae. Densdorsally(Fig.14A)with3inner(L)and6dorsal(D12,ld1–4)setae;seriesEwithamaximumof7setaeinfemales and6inmale;setaeL1–3,E1,andE3spine-like.Dens ventrallywith2unpairedsetae.Mucrowithspatulateapex. Remarks:Bothfemaleshaveonedenswith7Esetae andonewith6setae.InbothcasesthemissingsetaisE6. Inonefemale,thedenswithoutE6alsohasonly5dorsal setae. Thisspeciesisdistinguishedfrom P.hirtus asdescribed byChristiansenandBellinger(1998)andZeppeliniand Christiansen(2003)bytheabsenceofheadverticalsetaM5, bytheshapeofthefemaleappendageand,perhaps,bythe presenceofasmoothtunicaonallclaws.Oneindividual fromWisconsindepositedattheINHSandidentifiedas P. hirtus (Zeppelinietal.,2009)isidenticaltothespecimens fromWizardcaveinhavingonlyfourverticalheadsetaein seriesM(M5absent),thenumberofdorsalmanubrialsetae, generalshapeoffemaleappendage,andthenumberof maxillaryandlabialpalpssetae. Pygmarrhopalitessapo (ZeppeliniandChristiansen),2003 —TBS1/G1 Locality:PautlerCave Theindividualsstudiedhave6 + 6dorsalmanubrialsetae, maxillarypalpwithbifurcateapicalsetaand3sublobal appendages,andlabialpapillaEwith3guardsetae. ThisspeciesisendemictoMonroeCounty,andit previouslywasreportedfromFrog,Pautler,Jacobs,and RoseHolecaves(ZeppeliniandChristiansen,2003). Pygmarrhopalitessalemensis n.sp.—TBS1/G1 MaterialExamined:Holotype:INHSCollection Number551,641;Illinois,St.ClairCo,StemlerCave, 2.7miNEofColumbiaIL,adultfemale,slide-mounted, handcollectedoncavefloorindarkzone,28–30September 2009,SJTaylorandFNSoto-Adames.Paratypesonslides withINHSCollectionNumbers551,642-50;samelocality asholotype,1slide-mountedadultfemaleand3adultsof undeterminedsexinalcohol,handcollectedondrippool, darkzone;Illinois,MonroeCo.,Wanda’sWaterfallCave, 7.4miSEofValmeyer,2femalesand1maleadultson individualslidesand1adultofundeterminedsexinalcohol collectedinpitfalltrapsandbyhandoncavefloor15–17 September2009,SJTaylorandFNSoto-Adames;Illinois, MonroeCo.,IllinoisCaverns,1adultfemalemountedona slide,pitfalltrapindarkzone,2additionaladultsand1 juvenileofundeterminedsexinalcohol,pitfallandleaf litter,24–26September2009,SJTaylor,FNSoto-Adames, AKuhns,EZaborski,JJacoby,APaprocki,andM Pessino;Illinois,MonroeCo.,SpiderCave,6.5miSof Waterloo,1slide-mountedadultmale,pitfalltrap,15–17 September2009,SJTaylorandFNSoto-Adames;Illinois, Table3.Diagnosticcharactersforspeciesof Pygmarrhopalites withbifurcateC1,palmatefemaleappendageandtwounpairedventralsetaeondens. Species Ant.4 Subsegments Head Vertical SetaM5 OuterPlate Maxillary PalpSetae LabialPalp EGuard Setae Small Abdomen C1 SmallAbdomen DSeries PosteriortoD6 Female Appendages Branches/Stem Dorsal Manubrial Setae Number ofSetae onDens P.fransjanssens n.sp.6 2 33bifurcate2shallow/smooth47 P.incantator n.sp.6 2 14bifurcate4deep/smooth56–7 P.hirtus (Christiansen) 6 + ……bifurcate4shallow/smooth…6 P.arcus (Zeppelini andChristiansen) 7 2 ……bifurcate3shallow/smooth…7 P.hubbardi (Zeppelini andChristiansen) 7 + ……bifurcate4shallow/denticulate…7 P.principalisskelicus Vargovitsh a 6 2 …2?normal bifurcate? 3–4shallow/smooth77 a Although P.principalisskelicus hasunbranchedC1isitincludedinthetablebecauseitseemsmostsimilarto P.fransjanssens n.sp.fromamongPalearcticforms,andbecause P.principalis issometimesdescribedashaving bifurcateC1. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 167

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Figure13. Pygmarrhopalitesincantator n.sp.,dots,andopencirclesrepresentmicrosetae,andspine-likesetae,respectively: (A)detailofverticalsetaeofthehead;(B)generalchaetotaxyofface(C)completechaetotaxyofsmallabdomen,lateralview; (D)femaleanalappendage;(E,F)pro-andmetathoraxicclawcomplex,respectively. N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 168 N JournalofCaveandKarstStudies, August2013

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MonroeCo.,HiddenHandCave,3miWofWaterloo,2 adultfemalesand1juvenile,slide-mounted,7additional adultsandjuvenilesinalcohol,pitfalltrapsindarkzone, 14–16October2009,SJTaylorandFNSoto-Adames. Etymology:ThenewspeciesisnamedaftertheSalem Plateauregion,whereitseemstobewidespreadincaves. Description:Largestindividual0.89mm.Background colorwhite,withdarkorangespotsonheadcovering Figure14. Pygmarrhopalites spp.,dotsrepresentregularacuminatesetae: Pygmarrhopalitesincantator n.sp.:(A)complete dorsalchaetotaxyoffurcula,L = innercolumn,D = dorsal,ld = laterodorsal,E = outercolumn. Pygmarrhopalitessalemensis n.sp.:(B)completechaetotaxyofface;(C)fourthantennalsegment;(D,E)pro-andmetathoraxicclawcomplex,respectively. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 169

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clypeus,areabetweenantennalbasesandvertex;genaland labralareaswhite;largeabdomenwithscatteredorange spotsdorsolaterally,sometimesformingilldefinedstripes; smallabdomenandallappendageswhite.Ant.4with5 subsegments(Fig.14C)withproportionsas40:10:10:10:30 ofthetotalAnt.4length.Ant.4with4welldefinedwhorls ofsetaecorrespondingtotheapicalwhorlonsubsegmentI andeachoneofsubsegmentsII–IV;thenumberofsetae/ sensillaoneachwhorlas10/10/12/12;basalwhorlsmissing 2sensillapresentondistalwhorls.Ant.4subapicalsensilla capitate.Ant.3withoutbasalbulge;senseorganwithtwo rodsensillainindependentshallowdepressions,setaeApi Figure15. Pygmarrhopalitessalemensis n.sp.,chaetotaxyoffemalesmallabdomen,dotsandopencirclesrepresentmicroandmacrosetae,respectively:(A)seriesCsetaeofholotype,StemlerCave;(B)setaeC 4 andC 5 ,femaleparatypefromStemler Cave;(C)completechaetotaxyoflowervalve,paratypefromSpiderCave;(D,E)detailoffemaleanalappendage,paratypes fromWanda’sWaterFallandStemlerCave,respectively;(F)organizationofcompletechaetotaxyoflowervalveinholotype. N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 170 N JournalofCaveandKarstStudies, August2013

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andApeappearingthinwalled,basallyswollenandwith drawnoutapices,butotherwisenormallyacuminate,Aaia rodsensilla.Eyes1 + 1indarkorangepatch.Allvertical headsetaenormal(Fig.14B),M5absent.Apicalsetaeof outermaxillarylobebifurcate;sublobularplatewiththree hairs.LabialpapillaEwith4guardsetae.Smallabdomen withoutdenticlesorspines,chaetotaxyofseriesCasin Fig.15A:C1simpleandsmoothorveryfinelyserrate;C2– 6andC8withbasalserrations,denticlesorteeth,and lateralextensionsvariouslydevelopedaccordingtoindividualsandlocalities(Figs.15B,C);setaeC7andC9 longbutnotbasallyenlarged;setaD2/F3 5 1.2to1.8 (mode 5 1.6;3/7)C2/B2 5 0.9to1.1(mode 5 1.1;5/7)and C3/D3 5 1.8–3.1(mode 5 2.0;2/7);setaeD7–10present (Figs.15C,F);femaleappendageapicallysquareor rounded(Figs.15D,E),withshortserrationscovering apical1/3–2/3andsittingonaheart-shapedpapilla. Metatrochanterrectangular,with4anteriorand1posteriorsetae.Allclawswith1innertooth.Unguiculuswith innertoothlarge,singleandbasalonL1andL2 (Fig.14D),andabsentorsmall,duplicated,anddistalon L3(Fig.14E);apicalunguicularfilamentacuminateand surpassinglengthofunguisonalllegs.Tenaculumwith2 setae.Manubriumwith6 + 6dorsalsetae.Densdorsally (Fig.16A)with3inner(L),6dorsal(D1-2,ld1-4)and7 external(E)setae;setaeL1-3,E1andE3spine-like.Dens ventrally(Fig.16B)with2unpairedsetae.Mucroapically acuminateand0.6–0.7 3 (mode 5 0.7;4/7)aslongasdens. Remarks:OneindividualfromWanda’sWaterfallCave has10/11/11/11setae/sensillaonwhorlsI–IV.Thetwo individualsfromStemlerCavearemissingdentalsetaE6 ononedensbutnottheother.OneindividualfromStemler Cavehasoneproximalandonedistalinnerunguicular toothonL1-2,butonlythedistaltoothonL3.One individualfromHiddenHandCavehas3tenacularsetae. Pygmarrhopalitessalemensis n.sp.,belongstoagroup ofMidwesternspeciescharacterizedbyhavingfive subsegmentsonAnt.4andsmallabdomenseriesCsetae sculpturedorwithlateralextensions.Thefivespeciesin thegroupdifferindetailsofthesculpturingofsetaein seriesC,shapeofthefemaleappendage,numberofdental spines,numberofventralunpairedsetaeondens,and numberofheadverticalsetaeinseriesM(Table4).The newspeciesseemsintermediatebetweentherecently described P.sapo and P.leonardwoodensis Zeppelini, TaylorandSlay,2009.Thethreespeciescanbe distinguishedbydenschaetotaxy,patternofsculpturing ofsmallabdomensetaeC3-6,andfemaleappendage accordingtoTable4.Inaddition,in P.leonardwoodensis theinnertoothontheprothoracicclawsisbasal,whilein P.salemensis thetoothisinsertnearthemiddleofthe claw; P.sapo carries3guardsetaeinlabialpapillaE, whereas P.salemensis has4guardsetae.Differenceswith otherspeciesaredetailedinTable4. K ATIANNIDAE Sminthurinushenshawi (Folsom),1896—EUS5/G5 Locality:BatSumpCave Thisisacommonsurfacespecieswidespreadacross NorthAmerica.InIllinois,thespecieshasbeenpreviously reportedfromJackson,Champaign,Coles,Cook,DuPage, Kane,Lake,Randolph,Richland,andWoodfordcounties. Figure16. Pygmarrhopalitessalemensis n.sp.,completedorsalchaetotaxyofdens:(A)dorsal,L = innercolumn,D = dorsal, ld = laterodorsal,E = outercolumn;(B)ventral. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 171

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Table4.Diagnosticcharactersforspeciesof Pygmarrhopalites fromthemidwesternUSAstateshavingfivesubsegmentsonthefourthantennalsegmentandsmall abdomensetaeinseriesCeithersculpturedorwithlateralextensions. SpeciesColor Numberof HeadVertical MSeta SmallAbdomen SeriesCSetae Ornamentation Small Abdomen SeriesCSetae Extensions Female Appendage Female Appendage Papilla Numberof UnpairedVentral SetaeonDens Spines on Dens a P.salemensis n.sp.white,with extensive orangepattern 4basallyserratepresentnarrowpaddle, shortteeth symmetrical onapical 1/3–2/3 heart-shaped25 P.leonardwoodensis Zeppelini, TaylorandSlay white4singlebasal tooth orsmooth present orabsent narrowpaddle, shortteeth asymmetrical onapical1/3 heart-shaped25 P.sapo (Zeppelini andChristiansen) white4denticulateabsentnarrowpaddle, shortteeth symmetrical onapical1/3 heart-shaped21 P.madonnensis (Zeppelini andChristiansen) …4smoothpresentpalmate,long apicalteeth circular25 P.lewisi (Christiansen andBellinger) white5basallyserrateabsentnarrowpaddle, shortteeth symmetrical onapical1/3 heart-shaped12 P.ater (Christiansen andBellinger) white4basallyserratepresentserrate,seta-like acuminate elongate,pointed21 P.pygmaeus (Wankel) reddishbrown4smoothabsentnarrowpaddle, shortteeth symmetrical onapical1/3 circular25 a WhenfivedentalspinesarepresenttheyarealwaysL1-3,E1andE3;whentwospinesarepresenttheyareL1andE1;whenonespineispresentitisalwaysE1 N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 172 N JournalofCaveandKarstStudies, August2013

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Sminthurinushenshawi iscommonincaves,butthisisthe firstrecordfromthishabitatinIllinois. B OURLETIELLIDAE Bourletiella sp. Locality:SpiderCave Thisisanearlyinstarindividualnotidentifiableto species.Membersofthisgenusarenotcommonlyfoundin caves.Thisislikelyanaccidental. D ICYRTOMIDAE Ptenothrix sp. Locality:SpiderCave Thesinglejuvenilecollectedisnotidentifiableto species.Severalspeciesof Ptenothrix frequentcaves, including P.atra P.marmorata ,and P.maculosa .In Illinois, P.atra isthespeciesmostcommonlyreported fromcaves. D ISCUSSION SamplingofeightcavesintheSalemPlateauregion usingacombinationofmethodsyieldedforty-ninespecies, sixteenofwhichrepresentnewrecordsforIllinois.Thenew recordsincludefourspeciesdescribedasnew,fourspecies thatarelikelynew,butwithinsufficientmaterialforproper descriptions,andthreeotherformsthatwereassigned namesbutaresufficientlydistinctthatastudyofadditional materialmayshowthemtoalsorepresentnewspecies. EighteenspeciesarereportedforthefirsttimefromIllinois caves.Seventeenspeciesarerankedasrareforthestate (S1),buteightofthesearewidelydistributedacrossNorth Americaandthestaterankingiseitheranartifactofthe relativelypoorknowledgeofthefaunaofthestateorthe resultofunresolvedtaxonomicissues(e.g., Onychiurus pipistrellae n.sp.).Someoftheotherspeciesrankedasrare areprobablytrulyrareinthestate,eveniftheyare widespreadelsewhereonthecontinent,becausethey representthelimitofthedistributionalrangeforthe species(e.g., Pseudosinellaaera Folsomiabisetosa,Proisotomasepulcralis ,and Megalothoraxtristani ).Others,such as Pygmarrhopalites spp.,areprobablytrulyrareand endemictotheregion. Twelvespecieshavemorphologicalcharactersthat suggestsomelevelofadaptationtocavesandareclassified aseithertroglobionts,oreu-orsubtroglophiles.Seven species( Ceratophysellalucifuga,Onychiuruspipistrellae n. sp., Lethemurusmissus,Pygmarrhopalitesfransjanssnes n. sp., P.incantator n.sp., P.sapo ,and P.salemensis n.sp.) arecurrentlyknownonlyfromcaves,although L.missus andperhaps O.pipistrellae n.sp.arewidelydistributed acrossunconnectedcavesystems,suggestingtheyareable tomigratethroughprotectedsurfacehabitats.Most collectionsof Oncopoduraiowae havebeenmadeincaves, butrecentsurfacecollectionsincavebearingareassuggest amechanismtoexplainitswidespreaddistribution.Itis possiblethatsimilarmechanismsmaysupportthemovementanddispersalof L.missus and O.pipistrellae n.sp. throughsurfaceleaflitter. Folsomiacandida,F.stella, Pseudosinellaargentea,P.aera, and Coecobryatenebricosa showweakmorphologicaladaptationstocaves,allare widelydistributedacrossNorthAmericaandsurface populationsarenotrare. Sevenofthecavessurveyedwerepreviouslysampled byLewisetal.(2003).Theypublishedfindingsrelating primarilytotroglobionts,whereasthepresentstudy reportsonfindingsforspringtailsofallecological classificationsfoundincaves.Inaddition,ourstudy focusedonlyonspringtails,whereasLewisetal.(2003) surveyedallcaveinvertebrates.Inallinstanceseachcave sampledinthepresentsurveyyieldedmorespringtail speciesthanreportedbyLewisetal.(2003)(Table5).Most speciesreportedbyLewisetal.(2003)werecollectedagain duringthepresentsurvey.FiverecordslistedbyLewis etal.(2003)werenotconfirmedbythepresentstudy (Table5).Theabsencesofcollectionsof Pseudosinella fromWanda’sWaterfallCave,of Pygmarrhopalitescarolynae (ChristiansenandBellinger),of Onychiurus ‘‘ reluctus ’’fromHiddenHandCave,andof Sensillanuraillina (ChristiansenandBellinger)fromBatSumpCaveduring thepresentsurveyiscurious,giventhatthesewerethe Table5.ComparisonofspringtailspeciesrecordedfromcavesintheSalemPlateau(MonroeandSt.Claircounties,Illinois) byLewisetal.(2003)relativetothenumberrecordedinthepresentstudy. Cave Lewis etal.(2003) Present Study Speciesin Common Lewisetal.(2003)Species notFoundinPresentStudy PautlerCave2420 Wanda’sWaterfallCave1220 Pseudosinella sp.nr. argentea SpiderCave1710 StemlerCave11510 IllinoisCaverns3132 Lethemurusmissus HiddenHandCave341 Pygmarrhopalitescarolinae Onychiurus ‘‘ reluctus ’’ BatSumpCave2121 Sensilanuraillina Total137785 F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 173

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shortestcaveincludedinthepresentstudyand,withthe exceptionofWanda’sWaterfallCave,wesamplednearly thefulllengthoftheaccessiblepassagesatthesesites. Pygmarrhopalitescarolynae and P.salemsis n.sp.,theonly Pygmarrhopalites collectedinHiddenHandCave,differ sharplyincolorpattern,smallabdomensetaeornamentationandfemaleanalappendage,andhindclawmorphology,anditisnotlikelythetwospeciescouldbeconfused. Sensillanuraillina isprobablyatrogloxeneorsubtroglophile,andalthoughthespecieswasnotcollectedinsideBat SumpCaveduringourvisits,itwastakeninsurfaceleaf littersamplednearthecaveentranceonthesamedaythe cavewasvisited. Theabsenceof Lethemurusmissus insamplesfrom IllinoisCavernsisnotsurprising.IllinoisCavernsisalarge, complexsystem,andmostareasofthesystemwerenot sampledduringthepresentstudyduetotimeconstraints. Samplingeffortfocusedonthosesectionsofthecavemost impactedbythelargenumberofvisitorsthattourthissystem everyyear;thecavehassincebeenclosedtothepublicinan attempttohelpmanagewhitenosesyndromeofbats. Itistellingaboutthegeneralstateofourknowledgeof thespringtailfaunaofIllinoisthatleaflittersamples collectedinsidecavesattheentrance,haveyieldedthree newspeciesandsixnewstaterecordsofwhatareclearly surfaceleaflitterspecies. Itisofsomeconcernthat Coecobryatenebricosa, the onlymemberofthe Sinella Coecobrya generacomplex, acomplexoftypicallyeu/subtroglophileortroglobiont species,isaninvasivespecies.Itisnotclearwhattheroleof thisintroducedspringtailspeciesmightbeinfragilecave ecosystems.CavesintheShawneeHills,southoftheSalem Plateauregion,harborthreenativespeciesofeutroglophiles/troglobiontsinthegenus Sinella (Christiansenand Bellinger,1998).Itispossiblethattheintroducedform couldmovesouth,invadecaves,andextirpatethenative springtailspecies. Therelativelylargenumberofnewspeciesandrecords forthestateandthepotentialthreattonativeeutroglophilesandtroglobiontsbyintroducedspeciespointtothe needforcontinueddetailed,taxon-focusedsamplingof cavesystemsinIllinois.Onlythroughintensesamplingwill webeabletoidentifycommunitiesunderthreatandhave theinformationneededtomakemoreeffectivemanagementandconservationdecisions. A CKNOWLEDGEMENTS Wethankallvolunteersandscientistswhohelpedwith fieldwork,includingJoAnnJacoby,AndyKuhns,Annie Paprocki,MassimoPessino,ChrisPhillips,BobWeck,and EdZaborski.WethankLesleyDeemforhervaluable assistanceinsortingmaterialinthelaboratory.Weare gratefultoallthelandownerswhoprovidedaccesstotheir caves,includingDonCoonsoftheKarstConservancyof Illinois,JohnA.Nelze nandShaneSellerswiththeBoy ScoutsofAmerica,Joseph‘‘Mic’’Middletonwiththe IllinoisDepartmentofNaturalResources,StolleQuarry (especiallyJohnCramer),ThomasA.LeChienandKatie FeigenbutzoftheAmberWavesLandTrust,Boband NancyWeck,RaymondandKathleenMarchwinski,and PaulFree.KellyNeal(IllinoisNaturePreservesCommission)workedcloselywithusonpermitsnecessarytocarry outthiswork.JohnLovaas,MonaColburn,BobWeck, FrankWilhelm,andPhilipMossoftheIllinoisSpeleologicalSurveyprovidedaccesstodetailsofcavelocationsand maps.TheIllinoisEndangeredSpeciesProtectionBoard providedfundingtocovertravelcosts,laboratorysupplies, andlaboratoryassistance. R EFERENCES Babenko,A.B.,Chernova,N.M.,Potapov,M.B.,andStebaeva,S.K., 1994,CollembolaofRussiaandAdjacentCountries,Family Hypogastruridae,Moscow,Nauka,336p.InRussian. Barr,T.C.,Jr.,1963,Studiesonthecavernicole Ptomaphagus oftheUnited States(Coleoptera:Catopidae):Psyche,v.70,p.50–58. Barr,T.C.,Jr.,1968,EcologicalstudiesintheMammothCaveSystemof Kentucky:I.Thebiota:InternationalJournalofSpeleology,v.3, p.147–204. Betsch,J.-M.,andWaller,A.,1994,Chaetotaxicnomenclatureofthe head,thoraxandabdomeninSymphypleona(Insecta:Collembola): ActaZoologicaFennica,v.195,p.5–12. Bonet,F.,1947,(1948),Monograf adelafamiliaNeelidae(Collembola): RevistadelaSociedadMexicanadeHistoriaNatural,v.8, p.131–192. Bretfeld,G.,1999,SynopsesonPalearcticCollembola.Volume2. Symphypleona:StaatlichesMuseumfu ¨rNaturkundeGo ¨rlitz,AbhandlungenundBerichtedesNaturkundemuseumsGo ¨rlitz,v.71,318p. Chen,Jian-Xiu,andChristiansen,K.A.,1993,Thegenus Sinella with specialreferenceto Sinellas.s. (Collembola:Entomobryidae)of China:OrientalInsects,v.27,p.1–54.doi:10.1080/00305316.1993. 10432236. Chen,Jian-Xiu,andChristiansen,K.A.,1997,Subgenus Coecobrya ofthe Genus Sinella (Collembola:Entomobryidae)withspecialreferenceto thespeciesofChina:AnnalsoftheEntomologicalSocietyofAmerica, v.90,p.1–19. Christiansen,K.A.,1964,ArevisionoftheNearcticmembersofthegenus Tomocerus (CollembolaEntomobryidae):Revued’E cologieetBiologieduSol,v.1,p.639–677. Christiansen,K.A.,2012,KenChristiansenCollembolaCollection: GrinnellCollege,Grinnell,Iowa.http://web.grinnell.edu/courses/bio/ collembola/maintable_menu.asp[accessedJanuary31,2012]. Christiansen,K.A.,andBellinger,P.F.,1980,TheCollembolaofNorth AmericanorthoftheRioGrande,ataxonomicanalysis,firstedition, Grinnell,Iowa,GrinnellCollege,1322p. Christiansen,K.A.,andBellinger,P.F.,1998,TheCollembolaofNorth AmericanorthoftheRioGrande,ataxonomicanalysis,second edition,Grinnell,Iowa,GrinnellCollege,1518p. Christiansen,K.A.,andCulver,D.C.,1987,BiogeographyanddistributionofcaveCollembola:JournalofBiogeography,v.14,p.459–477. Culver,D.C.,andPipan,T.,2009,TheBiologyofCavesandOther SubterraneanHabitats,Oxford,OxfordUniversityPress,254p. Felderhoff,K.L.,Bernard,E.C.,andMoulton,J.K.,2010,Surveyof Pogonognathellus Bo ¨rner(Collembola:Tomoceridae)inthesouthern Appalachiansbasedonmorphologicalandmoleculardata:Annalsof theEntomologicalSocietyofAmerica,v.103,p.472–491.doi:10. 1603/AN09105. Fjellberg,A.,1984,CollembolafromtheColoradoFrontRange,U.S.A.: ArcticandAlpineResearch,v.16,p.193–208. Fjellberg,A.,1985,ArcticCollembolaI—AlaskanCollembolaofthe FamiliesPoduridae,Hypogastruridae,Odontellidae,BrachystomellidaeandNeanuridae,EntomologicaScandinavicaSupplement,no.21, 126p. N EWSPECIESANDNEWRECORDSOFSPRINGTAILS (H EXAPODA :C OLLEMBOLA)FROMCAVESINTHE S ALEM P LATEAUOF I LLINOIS ,USA 174 N JournalofCaveandKarstStudies, August2013

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Fjellberg,A.,1998,TheCollembolaofFennoscandiaandDenmark.Part I:Poduromorpha,Leiden,BrillAcademicPublisher,FaunaEntomologicaScandinavicaseries35,184p. Fjellberg,A.,2007,TheCollembolaofFennoscandiaandDenmark.Part II.EntomobryomorphaandSymphypleona,Leiden,BrillAcademic Publisher,FaunaEntomologicaScandinavicaseries42,266p. Gisin,H.,1947,SurlesinsectesApte rygotesduParcNacionalSuisse. Espe `cesetgroupementseue daphiques:Re sultatsdesrecherches scientifiquesentreprisesauParcNationalsuisse,v.2,p.75–91. Jordana,R.,Arbea,J.I.,Simo n,C.,andLucia n ez,M.J.,1997, Collembola,Poduromorpha:Madrid,MuseoNacionaldeCiencias Naturales,ConsejoSuperiordeInvestigacionesCient ficas,series FaunaIbe rica,v.8,807p. Lewis,J.J.,Moss,P.,Tecic,D.,andNelson,M.E.,2003,Aconservation focusedinventoryofsubterraneaninvertebratesofthesouthwest Illinoiskarst:JournalofCaveandKarstStudies,v.65,p.9–21. MariMutt,J.A.,andBellinger,P.F.,1990,ACatalogoftheNeotropical Collembola,IncludingNearcticAreasofMexico,Gainsville,Florida, SandhillCranePress,seriesFloraandFaunaHandbookno.5,237p. Master,L.,Faber-Langendoen,D.,Bittman,R.,Hammerson,G.A., Heidel,B.,Nichols,J.,Ramsay,L.,andTomaino,A.,2009, NatureServeConservationStatusAssessments:FactorsforAssessing ExtinctionRisk,Arlington,Virginia,NatureServe,57p. Panno,S.V.,Weibel,C.P.,andLi,W .,1997,KarstregionsofIllinois, Champaign,IllinoisStateGeologica lSurvey,openfileseries1997-2,42p. Park,Kyung-Hwa,Bernard,E.C.,andMoulton,J.K.,2011,Threenew speciesof Pogonognathellus (Collembola:Tomoceridae)fromNorth America:Zootaxa,v.3070,p.1–14. Pomorski,R.J.,Furgo,M.,andChristiansen,K.A.,2009,Reviewof NorthAmericanspeciesofthegenus Onychiurus (Collembola: Onychiuridae),withadescriptionoffournewspeciesfromcaves: AnnalsoftheEntomologicalSocietyofAmerica,v.102, p.1037–1049.doi:10.1603/008.102.0612. Potapow,M.,2001,SynopsesonPalearcticCollembola.Volume3. Isotomidae:StaatlichesMuseumfu ¨rNaturkundeGo ¨rlitz,AbhandlungenundBerichtedesNaturkundemuseumsGo ¨rlitz,v.73,603p. Skarz yn ski,D.,2005,Taxonomyoftheceratophysellanlineage(Collembola:Hypogastruridae)inthelightoflaboratoryhybridization studies:Zootaxa,v.1043,p.47–59. Skarz yn ski,D.,2007,Aredescriptionof Ceratophysellalucifuga (Packard) (Collembola,Hypogastruridae)fromNorthAmericancaves:Journal ofCaveandKarstStudies,v.69,p.275–278. Sket,B.,2008,Canweagreeonanecologicalclassificationof subterraneananimals?:JournalofNaturalHistory,v.42,p.1549– 1563.doi:10.1080/00222930801995762. Soto-Adames,F.N.,2010,Twonewspeciesanddescriptivenotesforfive Pseudosinella species(Hexapoda:Collembola:Entomobryidae)from WestVirginian(USA)caves:Zootaxa,v.2331,p.1–34. Stach,J.,1945,Thespeciesofgenus Arrhopalites Bo ¨rnoccurringin Europeancaves,AcademiaPolonaLitterarumetScientarium,Acta MuseiHistoriaNaturalisno.1,47p. + plates. Szeptycki,A.,1979,ChaetotaxyoftheEntomobryidaeanditsphylogeneticalsignificance.Morpho-systematicstudiesofCollembola,IV, Krako w,PolskaAkademiaNauk,ZakadZoologiiSystematyczneji Dos wiadczalnej,219p. Therrien,F.,Changnon,M.,andHe bert,C.,1999,Listedesespe `cesde CollembolesduQue bec:Bulletindel’entomofaune,v.21,p.8–11. Thibaud,J.-M.,Schulz,H.-J.,anddaGamaAssalino,M.M.,2004, SynopsesonPalearcticCollembola.Volume4.Hypogastruridae: StaatlichesMuseumfu ¨rNaturkundeGo ¨rlitz,Abhandlungenund BerichtedesNaturkundemuseumsGo ¨rlitz,v.75,287p. Vargovitsh,R.,2009,NewcaveArrhopalitidae(Collembola:Symphypleona)fromtheCrimea(Ukraine):Zootaxa,v.2047,p.1–47. Waltz,R.D.,andHart,J.W.,1996,AchecklistoftheIndianaCollembola (Hexapoda):ProceedingsoftheIndiananAcademyofScience,v.105, p.29–41. Weibel,C.P.,andPanno,S.V.,1997,KarstTerrainsandCarbonate BedrockinIllinois,Champaign,IllinoisStateGeologicalSurvey, Illinoismap8,1:500000. Wray,D.L.,1950,SomenewNearcticCollembola:Psyche,v.57, p.95–100 + 1plate.doi:10.1155/1950/65134. Zeppelini,D.,andChristiansen,K.A.,2003, Arrhopalites (Collembola: Arrhopalitidae)inU.S.caves,withthedescriptionofsevennew species:JournalofCaveandKarstStudies,v.65,p.36–42. Zeppelini,D.,Taylor,S.J.,andSlay,M.E.,2009,Cave Pygmarrhopalites Vargovitsh,2009(Collembola,Symphypleona,Arrhopalitidae)in UnitedStates:Zootaxa,v.2204,p.1–18. F.N.S OTO -A DAMESAND S.J.T AYLOR JournalofCaveandKarstStudies, August2013 N 175


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