Scientific Papers Natural History Museum The University of Kansas
Transcript of Scientific Papers Natural History Museum The University of Kansas
Upper Carboniferous Insects from the Pottsville Formation of Northern Alabama
(Insecta: Ephemeropterida, Palaeodictyopterida, Odonatoptera)
By
Roy J. BeckemeyeR 1,2 and michael S. engel1,3
1 Division of Entomology (Paleoentomology), Natural History Museum, Biodiversity Institute, and Department of Ecology and Evolutionary Biology, 1501 Crestline Drive – Suite 140, University of Kansas, Lawrence, Kansas 66045;
E-mail: (RJB) [email protected]; (MSE) [email protected] 957 Perry Ave., Wichita, Kansas 67203-3141
3 Corresponding author
CONTENTSABSTRACT..............................................................................................................2inTRodUcTion ...................................................................................................2 acknowledgemenTS ...........................................................................................3MATERIALSANDMETHODS............................................................................3 geological SeTTing ...........................................................................................3 meThodS .............................................................................................................3SYSTEMATICPALEONTOLOGY........................................................................4 SUpeRoRdeR ephemeRopTeRida peaRSe, 1947.....................................................4 Anniedarwinianewgenus........................................................................4 Anniedarwinia alabamensisnewspecies...........................................6 SUpeRoRdeR palaeodicTyopTeRida peaRSe, 1936..............................................6 Pharciphyzelusnewgenus........................................................................6 Pharciphyzelus lacefieldinewspecies................................................9 Camptodiaphanewgenus.......................................................................11 Camptodiapha atkinsoni newspecies..............................................11 Agaeoleptopteranewgenus.....................................................................13 Agaeoleptoptera uniotemplanewspecies........................................14 SUpeRoRdeR odonaTopTeRa lameeRe, 1900....................................................14 Oligotypus tuscaloosaenewspecies................................................14DISCUSSION.........................................................................................................17LITERATURECITED............................................................................................17
Scientific PapersNatural History MuseumThe University of Kansas
©NaturalHistoryMuseum,TheUniversityofKansas ISSNNo.1094-0782
21October2011 Number44:1–19
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INTRODUCTION
TheCarboniferousinsectfaunaofNorthAmericahasreceivedconsiderableattentionsincetheinitialstudiesbyScudderandhis contemporariesondeposits in Illinois,Pennsylvania,RhodeIsland,Kansas,Massachusetts,andtheMaritimeProvincesofCanada(e.g.,Scudder,1868a,1868b,1868c,1878,1879,1885a,1885b,1893,1895;Dana,1864;Sellards,1904;Handlirsch,1906a,1911,1919).Sub-sequently,CarpenterandKukalová-Peck,amongothers,provided a series of contributions extensively revisingandfurtherdocumentingtheCarboniferousinsectfauna(e.g.,Carpenter,1933,1938,1940,1960,1963b,1964,1965,1967,1980,1983,1987,1992a,1992b,1997;CarpenterandRichardson, 1968, 1971, 1976; Richardson, 1956; Cope-land, 1957; Kukalová-Peck, 1987, 1997a; Kukalová-Peckand Richardson, 1983; Burnham, 1983; Béthoux, 2006,2007,2008,2009;BéthouxandBriggs,2008:videetiamBé-thoux et al., 2008, for considerationofKukalová-Peck’shypotheses).Supplementingthesemorecomprehensiveworkshavebeenvariousisolatedrecordsfromdepositsin Utah, Missouri, Tennessee, New Mexico, Ohio, andKentucky(e.g.,Carpenter,1967,1970;Lewis,1979;NelsonandTidwell, 1987;McComasandMapes, 1988;Brauck-mannetal.,1993;Rasnitsynetal.,2004).Outsideofthese,workontheCarboniferousofNorthAmericahasslowedinrecentdecadesandnewdepositsbearinginsectshavenotbeenforthcoming.However,recentcollectingintheUpperCarboniferousPottsville Formation, primarily atthe Union Chapel Mine in north-central Alabama, hasuncoveredinsectmaterial,albeituncommonly(Lacefield,2000;Atkinson,2005).Althoughrepresentedentirelyasfragmentsofwings,thesecompressionsareexceptionallywellpreservedanddocumentasimilarbreadthofordinal
diversityalreadywellknownfromsimilar-ageddepositselsewhereinNorthAmericaandEurope.
Hereinweprovideapreliminaryaccountoftheinsectdiversitybasedonfivewellpreservedinsectwingsrecov-eredoverthecourseofseveralyearsfromthePottsvilleFormation. The McWane Science Center has an addi-tionalinsectcomprisedoftheapexofasmallwing(veryroughlyestimatedtobethedistal15–20%).Nocatalogueor accession numberwas assigned to the specimen, al-thoughitwasaccompaniedbyapaperlabelwiththean-notations: “Object: dragonflywing /UCMNo. [blank]/Horizon: Pottsville Fm. / Locality:UCM/Collectedby:PrescottAtkinson/Date:2008.”Despitethelabelat-tributionof“dragonfly”,thepreservedportion(approxi-mately8by14mm) is too incomplete toallowdefinitetaxonomicassignmentof this fossil at either familialorordinallevel.Thisspecimenandthosedescribedhereinmakeup theonly insect fossils,other than ichnofossils,currently known from the Pottsville Formation ofAla-bama.TheUniversityofKansasismountinganexcava-tion in thehopes ofmore fullydocumenting the insectfaunaofthePottsvilleFormation.
acknowledgmenTS
WeextendourthankstoDr.PrescottAtkinson,who,in his role as Vice President of theAlabama Paleonto-logicalSociety(APS)andProjectManagerfortheUnionChapel Mine (UCM), invited our review and study ofthesefossils. Healsogenerouslyprovidedlodgingandlocaltransportationfortheauthors’respectivevisits,andmade arrangementswith themuseums formaking the
ABSTRACT NewUpperCarboniferous(Pennsylvanian,WestphalianA)insectsaredescribedfromlo-calitiesinthePottsvilleFormationofnorthernAlabama(includingtheUnionChapelMine).Fivespeciesarerecordedinfivepalaeopterousordersandcomprisingfivegenera(fournewtoScience).Newtaxaproposedare:Anniedarwinia alabamensisnewgenusandspecies(Ephemeropterida:Syntonopterodea:Syntonopteridae);Pharciphyzelus lacefieldi new genus and species (Palaeodictyopterida: Palaeodictyoptera:Homoiopteridae),Camptodiapha atkinsoninewgenusandspecies(Palaeodictyopterida:Diaphanopterodea:Namurodiaphidae);Agaeoleptoptera uniotemplanewgenusandspecies(Palaeodictyopterida:Megasecoptera:Ancopteridae);andOligotypus tuscaloosaenewspecies(Odonatoptera:Protodonata:Paralogidae). Eachtaxonisdescribed,fig-ured,andcomparedwithcloserelativesinCarboniferousandPermiandeposits. Camptodiaphanewgenusextendsthegeographicalrangeof the familyNamurodiaphidae into theCarboniferousofNorthAmerica.ThediagnosisofthefamilyAncopteridaeisexpandedtoaccommodateAgaeoleptopteranewgenusratherthanproposeanothermonogenericfamily.ThedistributionofAncopteridaeisextendedgeographicallyintoNorthAmericaandtemporallyintotheUpperCarboniferous.Akeytothegeneraofancopteridsisprovided.
Key Words: Alabama; taxonomy;Pennsylvanian; Syntonopterodea; Syntonopteridae; Ephemeroptera;Palaeodictyoptera;Lycocercidae;Diaphanopterodea;Namurodiaphidae;Megsecoptera;Ancopteridae;Pro-todonata;Paralogidae.
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specimens and working space available. Dr.Atkinsonand theAPSwere instrumental in campaigning for thepreservationoftheUCMlocality,nowknownformallyastheStevenC.MinkinPaleozoicFootprintSite,aholdingoftheAlabamaDepartmentofConservationandNaturalResources(ADCNR).TheStateLandsDivisionofADC-NRisacknowledgedfortakingtheUnionChapelMinesiteundertheirprotectionandtherebymakingpossiblethefuturerecoveryofsuchspecimensasthosedescribedin thispaper. PrescottAtkinsonandMiloWashington,APSFieldTripCoordinator, alsohosted theauthorsonfieldtripstotheUCMlocality. Dr.RandyMecredy,Di-rector,andMichaelDressler,CollectionsAssistant,oftheAlabamaMuseumofNaturalHistory,andMr.JunEber-sole,CollectionsManagerattheMcWaneScienceCenter,wereallverysupportiveandhelpful.Theyprovidedfirst
classworking spaceandequipment, and therebymadethetimespentintheirfacilitiesefficient,productive,andpleasant. Thehospitalityandprofessionalismofall theaboveare sincerelyappreciated. In the initial stagesofidentifying several of these fossil wings, the charac-ter list and charactermatrix of Prokop andRen (2007),werequitehelpful.Lastly,wearegratefultotheeffortsofDr.CarstenBrauckmann,Dr. JakubProkop,Dr. JörgSchneider,andDr.Kirsten Jensen, the lastaseditor, forconstructiveinputandimportantcontributionsthatsig-nificantly improved themanuscript. Financial supportwasprovidedbyU.S.NationalScienceFoundationgrantDEB-0542909(toMSE).ThisisacontributionoftheUni-versityofKansasNaturalHistoryMuseumEntomologyDivision(Paleoentomology).
geological SeTTing
The insect fossils described in this paperwere col-lected from the Early Pennsylvanian (Westphalian A)Pottsville Formation, Mary Lee coal zone, in northernAlabamafromlocalitiesassociatedwithstripmines.AllthesitesareintheBlackWarriorcoalbasin(Murrieetal.,1976); twoare inWalkerCountyandone inTuscaloosaCounty(Fig.1). MostofthefossilsarefromtheformerUnionChapelMineinWalkerCounty,areclaimedminesite now preserved as the Steven C. Minkin PaleozoicFootprint Site, a Carbonifereous plant and ichnofossil(invertebrateandvertebratetrackway)lagerstätteknownforproducinglargenumbersofhighqualityfossils(Butaet al., 2005). Themine covers parts of the eastern halfofsec.21andthewesternhalfofsec.22,T.14S,R.6W,intheCordova7.5-minutetopographicquadrangle(Pa-shin,2005).Asthefossilsarefoundinoverburdenspoils,the exact stratigraphicpositionof theirorigin isnot al-waysdeterminable.ThegeneralizedstratigraphicsectionshowninFig.1(afterPashin,2005)showsthattheshalesin thealternatingshale-sandstonesequencesbelowandabovetheMaryLeecoalbedsarethesourceoftheterres-trialplantandtrackwayfossils.ThethreeUnionChapelMinespecimenswereallfoundbetween2000and2008;twoareassociatedwithplantmaterials,onewith tetra-podtrackways.
Oneofthefossilinsectspecimenswasfoundin1988inaspoilpileassociatedwithanunidentifiedstripmineatWindhamSprings,TuscaloosaCounty. Morespecificlocation information for this site is not available. An-otherwascollectedin1993atanopenpitcoalmine“ap-
proximately100yardseastofAlabamaHighway13,ap-proximately2milesnorthofEldridgeinWalkerCounty”(Lacefield,pers.comm.,2010)(Sec.5,T.13S,R.10W).Ac-cordingtoLacefield,theminewasoperatedbytheHaleyBrothersandBarbourCoalCompany,withheadofficesinHaleyville,Alabama.Ithassincebeenreclaimedandthe site is now part of the right-of-way for a four-lanehighway.Thisspecimencamefrom“ashalesequenceofthePottsvilleFormation,possiblyanintervalbetweentheJaggercoalseamandtheMaryLeecoal”(Lacefield,pers.comm.,2010).
meThodS
All specimens were photographed with a digitalmacro camera (Casio Exilim EX-FH20) and all exceptUCM2368and2369werescannedat1200–2400pixel-per-inch resolution using a Hewlett-Packard flat bed scan-ner. In addition, photomicrographs using aNikon 990digital camerawith external strobe flash (Nikon SB-26)weremadeofmostofthespecimenstoprovidevenationdetails.Flashorientationwasoptimizedtoshowdetailsofinterest.Imageprocessingsoftware(AdobePhotoshop6.0,AdobeIllustrator10,andXARAExtreme4.0.4966DL)wasusedtooverlayimagesasrequiredtoproducethefi-nalreconstructiondrawingsofwingshapeandvenation.Finalphotographicimagesforthefigurespresentedhereareforlightsourcesfromthetopleft.Theclassificationfollowed isgenerally thatofGrimaldiandEngel (2005)exceptasnoted.
MATERIALSANDMETHODS
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SYSTEMATICPALEONTOLOGY
SUpeRoRdeR ephemeRopTeRida peaRSe, 1947
HereinweuseEphemeropterida in the sense of in-cluding Syntonopterodea (Syntonopteridae), Permo-plectoptera (Protereismatidae), andEphemeroptera, butnotTriplosoboptera (Triplosobidae)as itsaffinitieswithPalaeodictyopterida,orasanodebetweenEphemerop-terida and Palaeodictyopterida, are becoming increas-inglyclear(e.g.,ProkopandNel,2009).
OrderSYNTONOPTERODEALaurentiaux,1953FamilySYNTONOPTERIDAEHandlirsch,1911
Anniedarwinianewgenus
Type species.—Anniedarwinia alabamensisBeckemey-erandEngelnewspecies.
Diagnosis.—Wings [First interpreted as a fore andhindwingpair,thefossilactuallycomprisesrightandleft
forewingswithonewingfoldedovertheother] lackingarchedictyon;originsoflongitudinalveinsunknown(ap-proximatelybasal30%ofrightforewingandbasal20%ofleft forewingmissing); ScP terminateson costalmarginapproximatelyone-quarterwing lengthbasadapex;RAunbranched;RPforkingbeyondmidwing;hindwingRAandMP briefly contiguous (but not fusing) basad one-quarter wing length; MA forking basad midwing, MPforkingdistad toMAfork;CuAforkingone-thirdwinglength;CuPunbranched(Fig.2); left forewingCuPandAA1 in contact at one-fifth wing length; anteriormostbranchAA terminating on posterior margin near mid-wing;intercalaryveinsbetweenterminalbranchesoflon-gitudinalveins,especiallynumerousnearwingapex.
Etymology.—The new genus-group name honorsCharles Darwin’s humanity by remembering his sec-ondchildandyoungestdaughter,AnneElizabeth“An-nie”Darwin (1841–1851). Darwin nursed his ten-year-olddaughterthroughthefinalstagesofherillness.Her
Fig.1.OccurrenceofnorthernAlabamaPottsvilleFormationfossilinsectsandageneralizedgeologicsection(afterPashin,2005)oftheEarlyPennsylvanianPottsvilleFormationMaryLeecoalzone.OutlinesofWalkerandTuscaloosacountiesindicated;circularsymbolsmarkthelocationsofthethreesitesfromwhichfossilswerecollectedwhiletheshadedareaindicatestheextentofthePottsvilleFormationinAlabama.
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deathbrokeher father’sheart:“Wehave lost the joyofourhousehold,and thesolaceofouroldage: shemusthaveknownhowwelovedher;ohthatshecouldknownow how deeply, how tenderlywe do still...” (CharlesDarwin,30April1851:Keynes,2001).
Discussion.—Anniedarwinia is much more slenderthan Lithoneura Carpenter, 1938, the only other genusfor which a nearly complete wing is available (lengthtowidthratiosof2.4forL. mirifica Carpenter,1944hind
wing,3.0 forA. alabamensisnewspecies right forewing,2.9forA. alabamensisnewspeciesleftforewing).Theonlysyntonopteridforwhichportionsofbothforeandhindwings areknown isLithoneura lameereiCarpenter, 1938,for which the basal third to half of thewings are pre-served.Inthatspecies,thebaseofthehindwingisquitewideandthewingsharplytapereddistally;moreover,thecostalmarginismuchmorenarrowsothatthehindwingshapedepartssignificantlyfromtheoverallovalshapeof
Fig.2. Anniedarwinia alabamensisnewgenusandspecies(Syntonopterodea:Syntonopteridae). Reconstructionofvenationofleftandrightforewingsofholotype(UCM1076a);veinsidentifiedandconvexity(+)orconcavity(–)(wingindorsalaspect)noted.
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theforewing.InL. mirifica,however,thehindwingisnotsignificantlywidenedatthebaseandisellipticalinshape,approachingtheshapeoftheknownforewings.Thusitisdifficulttoallocatesignificancetothedifferenceinslen-dernessbetweentheforewingofA. alabamensis newspe-ciesandthehindwingofL. mirifica.Moreover,MAisnotfusedwithRPintheleftforewingofA. alabamensisnewspecies,insteadclearlyabuttingitforabriefdistancebe-forediverging,whiletheseveinsfuseforashortdistanceinLithoneura.Finally,CuPiscurvedposteriorlyinLitho-neura,while thisvein is straight inAnniedarwinia. ThislattertraitismoresimilartoAnglolithoneuraProkop,Nel,andTenny,2010,fromwhichAnniedarwiniadiffersintheanal veins apparently not zigzagged (zigzagged inAn-glolithoneura)andScPandRAmorewidelyspaced(morecloselypositionedinAnglolithoneura)(Prokopetal.,2010)(Figs.2–4).Likeallofthesetaxa,Anniedarwiniahasacor-rugatewinglackinganarchedictyon,hasCuPsimple,hasMAwithananteriorcurvesuchthatitrunsalongorfuseswithRPforashortdistance,andhasAA1touchingCuPatonepoint(Fig.2).FromGallolithoneuraGarrouste,Nel,andGand,2009,Anniedarwiniadiffersby the forkingofMApriortotheforkingofRPinbothwings,theforkofMPmuchmorestronglydistad the forkofCuA(distadbutrelativelycloseinGallolithoneura),thelatterveinfork-ingmuchmorebasadthaninGallolithoneura (Garrousteetal.,2009).
Anniedarwinia alabamensisnewspeciesFigs.2–4
Ephemeropterasp.;Atkinson,2005:171–173,fig.1A,1B,2.
Diagnosis.—Asforthegenus(vide supra).Description.—Both wings. ScP, RA, RP, and ante-
riorbranchofMAstraight,parallelandequally spacedroughly3mmapartatmidwing;trueveinsandinterca-lary veins fairlyuniformly spaced alongposterio-distalmargins of wing; no crossveins preserved (Figs. 2–4).Right forewing. Distal 70% of wing, preserved length79.9mm,preservedwidth37.8mm,estimatedtotalwinglength114mm;ScPterminatingjustbasalofthree-quar-tersofwinglength;RAunbranched,terminatingat95%ofwinglength;RPbranchesat60%winglength,RPwithfivebranches,numerousintercalaryveins,branchester-minating indistal15%ofwing length;MAforking justbasalto50%winglength,MAwiththreebranches,anteri-orbranchrunningparalleltoposteriorbranchofRPuntiljustdistalof65%winglength,thenturningrathersharp-lytowardposteriormargin,terminatingjustbasalto85%wing length;MP forkingbetween45–50%wing length;CuAforkingnear35%winglength;CuPterminatingjustbasalto60%winglength;analveinsterminatinginbasalhalfofwing;intercalaryveinsnumerousalongposterior
marginandapexofwing. Left forewing.Distal80%ofwing, preserved length 95.4mm, preservedwidth 39.9mm,estimatedtotalwinglength115mm;ScPterminat-ingbasaltothree-quarterswinglength;RAunbranched,terminatingjustdistalto95%ofwinglength;RPbranch-ingat two-thirdswinglength,6-branched,withnumer-ousintercalaryveins,branchingterminatingindistal10%ofwing;basal3mmofpreservedportionofRPandMAcontiguous,separatingat25%ofwinglength;MPforkingbasalto60%winglength,3-branched,branchesterminat-ingbasalto75%ofwinglength;CuAforkingatone-thirdwing length, portion of CuA beyond 50%wing lengthnot preserved; CuP unbranched, terminating on poste-riormarginat55%winglength,mostanterioranalveinincontactat20%winglength;posteriorportionsofwingmissingfromone-totwo-thirdswinglength;numerousanalveins,terminatingbasalto50%winglength;numer-ousintercalaryveinsatposteriormarginofwingindis-tal one-quarter ofwing; left forewingmaximumwidthbetween30and50%wing length, right forewingabout6%wider than left forewing(althoughthedifference inwidthcouldconceivablybe representativeof theactualinsect,itcouldalsobetheresultoftaphonomiceffects).
Holotype.—PV2005.7.2.252.1 (UCM 1076a-part)/2005.7.2.252.2 (UCM 1076b-counterpart) (Figs. 3,4),University ofAlabamaMuseumofNaturalHistory,Tuscaloosa,Alabama;nearlycompleteforewingpairwithone foldedover theother;basal20–30%ofwingsmiss-ing;PottsvilleFormation(Pennsylvanian),UnionChapelMine,WalkerCounty,Alabama,July2000,T.P.Atkinsoncollector.
Etymology.—The specific epithet is taken from theState of Alabama, from which the type material origi-nates.
SUpeRoRdeR palaeodicTyopTeRida peaRSe, 1936(=RoSTRapalaeopTeRa wooTTon and kUkalová-peck, 2000)
OrderPALAEODICTYOPTERAGoldenberg,1877FamilyHOMOIOPTERIDAEHandlirsch,1906b
Pharciphyzelusnewgenus
Type species.—Pharciphyzelus lacefieldi BeckemeyerandEngelnewspecies.
Diagnosis.—Forewing(basedontheexpandedareabetween ScP and the costalmargin in the basal half ofthewing)withR,M,andCuallbranchinginbasalthirdof wing and ScP extending beyond midwing (incom-pletelypreservedbutbasedoncoarseandpositionScPlikelyextendsnearlytowingapex);pronouncedbendinstems ofmainveins near base;R branching just distad
carboniferouS inSectS froM alabaMa 7
branching of M (Fig. 5A); RP branching beyond wingmidlength(Fig.5A);MAunbranched,originofMAnearand basad first branch ofMP;MP area triangular andmultibranched; CuA unbranched; CuP branched, CuAandanteriorbranchofCuPconvergentatposteriormar-ginofwing(Fig.5A);convexridgeinwingmembraneatbaseofanalveinsformingananalbrace;crossveinsverynumerousandmostlyreticulate(Fig.5)(somecrossveinsanastomosing but not reticulate as in true BreyeriidaeHandlirsch, 1906b, contra tentative identification inAt-kinson,2005).
Etymology.—Thenewgenus-groupnameisacombi-nationoftheGreekwordspharci(meaning,“awrinkle”),for the convex ridge in themembraneof thewing thatforms the anal brace, andphyzelos (meaning, “shy”), inrecognitionoftherarityofPottsvilleFormationinsectfos-silsfromAlabama.Thenameismasculine.
Discussion.—ThefamilyHomoiopteridaewasestab-lished byHandlirsch (1906b) to contain several speciesofinsectsfromtheUpperCarboniferousofCommentry,France, andwas later revisedbyKukalová (1969),whonoted that “...the families Homoiopteridae, Breyeriidae
Fig.3.Photomicrographofholotype(UCM1076a,part)ofAnniedarwinia alabamensisnewgenusandspecies(Syntonopterodea:Syntonopteri-dae)(PV2005.7.2.252.1).
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[Handlirsch,1906b],Graphiptilidae[Handlirsch,1906b],and Lycocercidae [Handlirsch, 1906a] form a phylo-genetic unit...” She also stated that, unlikeBreyeriidaeand Graphiptilidae, Homoiopteridae and Lycocercidaesharedapronouncedbendinthestemsofthemainveins
inthebasalthirdofthewings.ShenotedthatCuAwassimpleandcurvedinLycocercidae,butbranchedinHo-moiopteridae,andthatthelatterfamilywasalsocharac-terizedbya“scleroterizedstripposteriortothecosta.”Inaddition,Kukalováindicatedthatshehadseenthestrip
Fig.4.Photomicrographofholotype(UCM1076b,counterpart)ofAnniedarwinia alabamensisnewgenusandspecies(Syntonopterodea:Synto-nopteridae)(PV2005.7.2.252.2).
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inmostoftheHomoiopteridaespecimensshereviewed,butthestriphasseldombeencitedinsubsequentdescrip-tionsofhomoiopteridtaxa.In1983,Kukalová-PeckandRichardson described Mazonopterum wolfforum and as-signedittoHomoiopteridae;thespecieshadCuAsimple,buttheauthorsdidnotcommentonhavingthereincom-promised one of the distinguishing characters betweenLycocercidae andHomoiopteridae. In that samepaperthey synonymizedRoechlingia hitleri Guthörl, 1933 [cor-rected from “Röchlingia” toRoechlingia by BrauckmannandBecker, 1992]with Scepasma gigasHandlirsch,1911;theholotypeofR. hitleri hadasimpleCuAandbyimpli-cationScepasmawouldhaveshared thischaracter; later,Brauckmann and Becker (1992) reinterpreted the fossilofS. gigasandverifiedthesimpleCuAanddescribedanadditionalspecies,S. mediomatricorum.Carpenter(1992c)continuedtolist“CuAsimple”asacharacterofLycocer-cidae,“CuAbranched”ascharacteristicoftheHomoiop-teridae,althoughhelistedMazonopterumasahomoiop-terid. HeplacedScepasma infamily incertae sedis,basedon theholotype’s fragmentarynature, apparentlymiss-ing the additional material represented by Roechlingia.Mostrecently,Prokopetal.(2006)describedAnglopterum magnificum asanewspeciesofHomoiopteridae; itpos-sessedCuAsimple.
Kukalová-PeckandRichardson(1983)listedanothercharacterofthenewspeciesofHomoiopteridaetheyde-scribed (Mazonopterum wolfforum, Larryia osterbergi, andTurneropterum turneri):ananalbracecomprisedofacon-vex ridge in thewingmembrane in the formof an arcthattiedthebasesoftheanalveinstogether.TheywentbackandreviewedotherpreviouslydescribedHomoiop-teridae[Boltopruvostia robusta(Pruvost,1919),Homoioptera woodwardiBrongniart,1893,Homoioptera gigantea Agnus,1902,Ostrava nigraKukalová,1960,Scepasma gigasHan-dlirsch,1911,Thesoneura americanaCarpenter,1944],andfoundthatallsharedthisconvexanalbrace.Prokopetal.(2006)alsofoundtheanalbracetobepresentonA. mag-nificum.Kukalová-Peck(1997b),inadiscussionofbasalarthropodstructures,identifiedthe“analbrace…formedby[a]stiffenedmembraneinconjunctionwiththebasalportionofAA”asanautapomorphyofthePalaeoptera.However,thebracehasneverbeendescribedaspresentonanylycocercidspecies,andweareunawareofithav-ingbeenidentifiedinotherfamiliesofPalaeodictyoptera(althoughitshouldtobenotedthatthisstructuremighteasilybeinterpretedasataphonomicartifactandmissedunlessonewerespecificallylookingforit).
Pharciphyzelus lacefieldi newspecies,possessesbothasimpleCuAandananalbraceformedbyaconvexridgeinthewingmembrane.WethusplaceP. lacefieldiinthefamilyHomoiopteridaeratherthanLycocercidae,notinginpassing that the familyLycocercidae shouldat some
pointinthefutureberevisitedandreinterpretedtoclarifyitsrelationshiptoanddistinctionfromHomoiopteridae.
Pharciphyzelusnewgenusdiffers frommostHomoi-opteridaegenerainhavingCuAsimple.AsopposedtoMazonopterum, Pharciphyzelus new genus has a shorterandmore sharply curved stem ofMP, the stems ofM,CuA,andAA1aremoresharplybent(22°/11°,36°/24°,38°/33°,respectively);stemofCunearlybisectsanglebe-tweenCuAandCuPinPharciphyzelus,stemofCuparalleltoCuPwithCuAarching sharply forward inMazonop-terum.PharciphyzelusdiffersfromScepasmaandMazonop-teruminhavingmorenumerousandmorecloselyspacedbranchesofMPandinCuPsinuousratherthanstraighttosmoothlyposteriorlycurved.Anglopterumhasaverywide field between RP andMA and between the stemofMandCuAwhilethesefieldsaremoderatelywideinPharciphyzelus.
Pharciphyzelus lacefieldi newspeciesFig.5
Palaeodictyopterasp.;Lacefield,1993:11,unnumberedfig.Palaeodictyoperasp.;Lacefield,2000:68,unnumberedfig.Breyeriasp.;Atkinson,2005:173–174,fig.4.
Diagnosis.—Asforthegenus(vide supra).Description.—Basalportionofwing,76.1mmlong,
38.4mmwide(estimatedtocomprise70%oftotalwinglength, estimated total length ca. 105 mm). PreservedportionofScPcloselyparallelingR;maximumdistancebetweencostalmarginandScP5mm,distancebetweenScPandR2mm;stemsofMandCuparallelandcontigu-ousbutnotcoalesced;stemsofR,M,andCustraightandparallel untilCudiverges sharplyposteriorly, atwhichpointRandMareprominentlyarchedandreflexed;be-tweenseparationofRPfromRAandfirstbranchingofRP,thecostalmargin,ScP,RA,RP,andMAnearlystraight,parallel and closely spaced; RP distally branched (Fig.5A);stemofRP longer thanstemofR;stemofMfromdivergenceofCuAlongerthanstemofMP;stemofCuPnearlytwiceaslongasstemofCuafterdivergencefromM; anterior branch of CuP following a sinuous courseinitially, diverging from CuA, then converging towardCuAatposteriorwingmargin;analveinsnumerousanddensely branched, anal veins branch basally; course ofanal, cubital,andmedialveins initiallysmooth,butbe-comereticulateasposteriormarginisapproached;cross-veinsfineanddenselyreticulate;convexridgeintheformofanarcthroughmembraneinanalareaformingananalbracethatpassesthroughtheanalveinsatanearlyper-pendicularangle,locatedataradiusofabout1cmfrompreservedbase.
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Fig.5.Pharciphyzelus lacefieldi newgenusandspecies(Palaeodictyoptera:Homoiopteridae):A.Reconstructionofwingvenation;BRdenotesanalbridgeformedbyconvexridgeofstiffenedmembranesensuKukalová-PeckandRichardson(1993).B.Photomicrographofholotype(noknowncounterpart),comprisingbasal70%offorewing(PI1993.0001.0001).
carboniferouS inSectS froM alabaMa 11
Holotype.—PI993.0001.0001 (Fig. 5B), University ofAlabamaMuseum ofNaturalHistory, Tuscaloosa,Ala-bama;partialforewingindorsalaspect;nocounterpart;PottsvilleFormation(Pennsylvanian),coalmine2miN.ofEldridge,WalkerCounty,Alabama,16January1993,J.Lacefieldcollector.
Etymology.—Thespecificepithetisapatronymhon-oring JimLacefield,Alabamageologist andpaleontolo-gistandcollectoroftheholotype.
OrderDIAPHANOPTERODEAHandlirsch,1919FamilyNAMURODIAPHIDAE
Kukalová-Peck&Brauckmann,1990
Camptodiaphanewgenus
Type species.—Camptodiapha atkinsoni BeckemeyerandEngelnewspecies.
Diagnosis.—Stems of ScP and R straight at wingbase,withoutdeflection(Fig.6A);MAwithsharpbendatconvergencewithRP,veinsdonotcoalescebutarejoinedbyshortcrossvein;CuAwithsharpbendatconvergencewithMbut veins do not coalesce (presence or absenceofcrossveinunknownbecauseofdefectinrocksurface:Figs. 6A, 7); ScP terminating on costalmargin distal towingmidlengthbutwellshortofapex;MAandCuAun-branched(Fig.6A);RAandanteriorbranchofRPparallelandwidelyseparatedoverdistalhalfofwing;RPwithsixbranches;MPwithsevenbranches;CuPwithatleastthreebranchespreserved; all branchingof longitudinal veinsdichotomous. Differentiated from Diaphanopteridae:DiaphanopterabyMAandRPandCuAandMconverg-ingbutnotcoalescingandwithshortcrossveinsversuscoalescence inDiaphanopteridae,andbyCuAandCuPwellseparatedandnotconvergentwithashortcrossveinasinDiaphanoptera(BéthouxandNel,2003).
Etymology.—Thenewgenus-groupnameisacombi-nationoftheGreekwordkamptos(meaning,“bending”or“flexible”), inreferenceto thesharplybent longitudinalveinsMAandCuA,andthesuffixdiapha,fromthefamilyname.Thenameisfeminine.
Discussion.—The family Namurodiaphidae is rareand heretofore known only from a single specimen,Namurodiapha sippelorum Kukalová-Peck and Brauck-mann, 1990. The holotype is a nearly complete insectfrom Hagen-Vorhalle of Germany and was deemed tobeofparticularinterestsinceitpossessedwhatwaspre-sumedtobeaprimitiveformofwingbracingthroughco-alescenceofMAandRPandofCuAandM,afeaturethatispresentincertainpalaeopterousinsectsandsomeNe-optera.Inthisfamily,coalescencedoesnotoccur,buttheveins convergeandare connectedbya short crossvein.
Theoccurrenceof Camptodiapha atkinsoniinthePennsyl-vanianPottsvilleFormationextendstherangeofthefam-ilytoNorthAmerica.ThenewspeciesisroughlytwicethesizeofN. sippelorum,whichhadawinglengthof37mm.
Sinitshenkova(2002)andGrimaldiandEngel(2005)bothplacedthefamilyNamurodiaphidaeinMegasecop-tera,althoughKukalová-PeckandBrauckmanndescribedit inDiaphonopterodea. However, the trimerous tarsi,absenceofanarchedictyon,andsimpleMAsupport,asdoes the putative presence of neopterouswing flexion,inclusionofthefamilywithinDiaphanopterodeaandweaccordinglyreturnthefamilytothisorder.
Camptodiapha atkinsoni newspeciesFigs.6–7
Diagnosis.—Asforthegenus(vide supra).Description.—Hind wing (determined by straight
anteriormarginofwing),71.7mmpreservedlength,74mm estimated total length,maximumpreservedwidth19.3mm,estimatedmaximumwidth21mm;basalthirdofwingmissinganteriortoR;measurementsinmmdis-taltooriginCuA:stemofMdivergingfromRat5.8mm;CuAdivergingfromM(closestconvergenceofCuAandM) at 5.6mm; first branch CuP at 8.5mm; originMA10.7mm;originRPat12.9mm;MPforking8.9mmdis-taltooriginMA;MAbendingsharplyatpointofclosestconvergencewithRPat8.9mm,shortcrossvein joiningMAandRP;ScPintersectingcostalmarginattwo-thirdswinglength;firstbranchofRP24.7mmdistaltooriginRP,RPwithsixbranches; indistalhalfofwingRAandanteriorbranchRProughlyparallelandseparatedby2mm;branchesofRPintersectingposteriorwingmarginin distal quarter ofwing;MAunbranched, terminatingbeyond three-quarterswing length;MP forkingat levelofMAconvergencewithRP,MPwithsevenbranches,allterminatingonposteriormarginbeyondwingmidlength,intersectingmarginatanglesof60±5º;CuAunbranched,bending sharplyat convergencewithM, althoughonlyshortstemofMAafteroriginpreserved(Fig.7);missingportionofCuAbetweenoriginandconvergencewithM(soitisunknownwhetherornottherewerem-cuacross-veins:Fig.7);CuAterminationnotpreserved(posteriorthirdofbasalhalfofwingdistal toAA2missing);CuPwithatleastthreebranches,butportionsofbranchesdis-tal to forkingmissing;preservedportionsof analveinsbranched; crossveins faint and poorly preserved, butstraightandnotreticulate.Counterpartcomprisingdis-tal two-thirds of wing less anterior margin, distal halfshowingonlycourseoflongitudinalveinsthroughanob-scuringthinlayerofmatrix;portionofwingmembrane(whererevealed)withanirregularbandedcolorpattern.
Scientific PaPerS, natural HiStory MuSeuM, tHe univerSity of KanSaS12
Holotype.—WSC#MSC9334 (UCM3045), McWaneScienceCenter; part a nearly completehindwing lack-
ingposteriorportionofbasalhalfandcostalregionante-riortoRinbasalthird(Fig.6B),counterpartrepresenting
Fig.6.HindwingofCamptodiapha atkinsoni newgenusandspecies(Diaphanopterodea:Namurodiaphidae):A.Reconstructionofwingvena-tion.B.Photomicrographofpart(WSC#MSC9334–UCM3045)comprisingnearlycompletewingindorsalaspectabsentonlytheverybase;tosamescaleasinfigure6A.C.Photomicrographofcounterpart,posterio-distalhalfofwinginventralaspect,portionofmembranecoveredbythinlayerofmatrix,visibleportionofwingmembranerevealingextensivecolorpattern.
carboniferouS inSectS froM alabaMa 13
thedistal60%ofwinglackinganteriormarginandwithathinlayerofmatrixcoveringthedetailsofwingvena-tionandmembraneoverallbut thecentral thirdof thewing (Fig. 6C); specimens on thin slab of rock roughly2mmthickthatwasfoundinassociationwithtetrapodtrackways;PottsvilleFormation(Pennsylvanian),UnionChapelMine,Walker County,Alabama;March 2005, P.Atkinsoncollector.
Etymology.—Thespecificepithet isapatronymrec-ognizingDr. T. PrescottAtkinson, collector of this andseveral other insect specimens from the Union ChapelMine locality;he isoneof theAlabamaPaleontologicalSocietymembersresponsiblefortheprotectionandpres-ervationofthisimportantfossillocality.
OrderMEGASECOPTERAHandlirsch,1906aSuborderEUBLEPTOPTERACrampton,1930
(=EubleptidodeaLaurentiaux,1953)FamilyANCOPTERIDAEKukalová-Peck,1975
AncopteridaeKukalová-Peck,1975:3.Typegenus:An-copteraKukalová-Peck,1975.
Diagnosis.—Wingsslender,butapparentlynotpeti-olate; almost equally broad to slightly tapered beyondwing midlength; apex broadly rounded to slightly ta-pered; ScP extendingbeyondwingmidlength; ScP andRAclosetocostalmargindistally;basesofRandMnot
coalescedbutstemofRPandMmaybecontiguousnearbase;RParisingbasally;Mbranchingproximally;MAandCuA simple; RP,MP, andCuP branched; costalmarginnearlystraightbutbrieflyconvexbasaltooratwingmid-length; crossveinsnumerous, forming intercalaryveins;posteriormarginofwingsinuoustomildlyundulate.
Comments.—The family Ancopteridae was estab-lishedbyKukalová-Peck (1975) for theLowerPermiangenus,Ancoptera Kukalová-Peck, 1975, from theMora-vian Obora deposits. Ancoptera is much broader, lessslender, and has a straighter and less concave costalmarginandamuchmoreundulatoryposteriormarginthanAgaeoleptoptera. Themajorvenational characteris-tics listed in the familial diagnosis are consistentwiththose identified for Ancopteridae by Kukalová-Peck(1975)andCarpenter(1992c).Thedifferences(especiallylackofdefinitivepreservationoftheposterio-basalpor-tionof thewing, theattenuatedformofAgaeoleptopteraandthealmostsinuousratherthanundulatoryposteriormargin)mightbesufficienttowarrantestablishmentofanewfamily.However,weprefertotakeaconservativeapproachandplacethenewgenuswithinAncopteridae,extendingthedefinitionandexpandingthegeographicrange and temporal extent of the family. If additionalmaterialisfoundinthefuture,thisassessmentmayhavetoberevisited.
key To geneRa of ancopTeRidae
1. Wing long but only slightly narrow,with broadlyroundedapex (wing lengthabout4–5 timeswidth),wingnearlysamewidthdistadandbasadmidwing;posteriormarginwithmildundulations;costalmarginstraightwithslightconvexityinbasalthird;RPbranchesinbasalthirdofwing(LowerPermian:Obora,Moravia,CzechRepublic).....................AncopteraKukalová-Peck
—. Wingverylongandnarrowwithslightlytaperedapex(winglengthabout7timeswidth),wingtaperedandnarrowedslightlydistadmidwing;posteriormarginslightly sinuous; costalmarginnearly straightwithdefinite convexity atmidwing;RPbranchesnearermidwing(UpperCarboniferous:UnionChapelMine,Alabama,USA)....................Agaeoleptopteranewgenus
Agaeoleptopteranewgenus
Type species.—Agaeoleptoptera uniotemplaBeckemey-erandEngelnewspecies.
Diagnosis.—Wingextremelyslender,apparentlynotpetiolate, slightly narrower distally than basal to mid-
Fig.7.Detailsofbaseofradial-cubitalspaceandportionsofCuAinhindwingofCamptodiapha atkinsoninewgenusandspecies(Diapha-nopterodea:Namurodiaphidae).ThepreservedportionofthebranchofCuAatitsorigin,andthepreservedcourseofCuAafteritsconvergencewithMallowreconstructionofthecourseofCuAasillustratedinfigure6A.ThemissingportionoftherocksurfacelikelycontainedacrossveinbetweenCuAandMsimilartotheonebetweenMAandRP(Fig.6).
Scientific PaPerS, natural HiStory MuSeuM, tHe univerSity of KanSaS14
wing,posteriormarginslightlysinuous(Fig.8A);costalmargin,ScP,andRAallparallelandveryclosetogetherdistally; RP apparently separating from RA at extremebaseofwing,RPwith three terminalbranches;RPandMcontiguousproximally(Fig.8A);stemofMveryshort;MAandCuA simple (Fig. 8A);MPandCuPbranched;intercalaryveinsinradialandmedialfields.
Etymology.—The genus-group name is a combina-tionoftheGreekwordsagaios(meaning,“elegant”),leptos(meaning,“slender”),andpteron(meaning,“wing”).Thenameisfeminine.
Agaeoleptoptera uniotempla newspeciesFig.8
Megasecopterasp.;Atkinson,2005:175,fig.6.
Diagnosis.—Asforthegenus(vide supra).Description.—Preserved length 80 mm, estimated
total length 83mm; preservedwidth 11.8mm; ratio oflength towidth 7.0, placing it in associationwithmostslenderoffossilwings(Kukalová-Peck,1975);maximumwingwidth justdistal to one-quarterwing length, cos-talmarginwithnoticeable convexity just basal towingmidlength; posterio-basal portion of wing absent, butimpressions of shape visible onmatrix of part indicatebaseofwingtobebroadratherthanpetiolate(Fig.8.1);only basal and terminal portions of ScPpreserved, ScPterminatingatthree-quarterswinglength;costalmargin,ScP,RA,andRPeachseparatedbyabout1mmat10%ofwinglength;RAnearlystraight,withslightcurvature,terminatingnearapex;RA-RPbranchingnotpreserved,RP contiguouswithM in basal 10% ofwing (Fig. 8A);portionofRPfrom40%winglengthtoabout70%winglengthnotpreserved,branchingofRPnotpreserved,butwith three terminal branches, with numerous interca-laryveins,RPfieldinterminal15%ofwinglength;MAsimple,branchingfromMat15%winglength,terminat-ingat83%winglength;firstMPforkjustbasaltowingmidlength,MPwiththreeterminalbranchesandseveralintercalaryveins(Fig.8A);CuAandMPparallelandwellseparated(about1mm)basally,CuAandanteriorbranchofCuPnotpreservedbeyondwingmidlength,estimatedthatCuAwouldterminateat60%winglength;CuPandAA1notpreservedinbasal20%ofwing;AA1terminat-ingatabout40%winglength;fewcrossveinspreserved.
Holotype.—PV2005.0007.0260.001 (UCM2368) (part)(Fig. 8B) / PV2005.0007.0260.002 (UCM2369) (counter-part) (Fig. 8C), University ofAlabamaNaturalHistoryMuseum,Tuscaloosa,Alabama;nearlycompletewingonlargeslabs(approximately30by40cm)thatalsocontainfossil plant material; Pottsville Formation (Pennsylva-
nian),UnionChapelMine,WalkerCounty,Alabama,P.Atkinsoncollector.
Etymology.—ThespecificepithetisacombinationoftheLatintermsunio(meaning,“union”)andtemplum(meaning,“temple”),inrecognitionofUnionChapel,thenamesakeoftheminethatisthetypelocalityforthespecies.
SUpeRoRdeR odonaTopTeRa lameeRe, 1900
OrderPROTODONATABrongniart,1893FamilyPARALOGIDAEHandlirsch,1906b
GenusOligotypusCarpenter,1931Oligotypus tuscaloosaenewspecies
Fig.9
Diagnosis.—Largeandslenderspecies(130–160mmintotallength)withcostalmarginconcavelycurved,ScPterminating on costal margin near midwing, posteriorbranchesof longitudinalveins terminatingmorenearlyperpendiculartoposteriorwingmargin(Fig.9),termina-tionsoflongitudinalveinsmorebasalthaninotherOli-gotypusspecies.
Description.—Distal half of forewing (basal half offorewing,hindwing,bodyunknown):preservedlength77 mm, estimated to comprise distal 50–60% of winglength,estimatedlengthofcompletewing130–160mm;wingslender,maximumwidth23mm;ScPinterceptingcostalmargin46mmbasalofwingapex,estimatedtoberoughlythree-quartersofwinglengthfrombase;distancebetweencostalmarginandScPequaltodistancebetweenScPandRAat levelof initialbranchingofRP;RAverycloselyparallelingcostalmargindistaltoterminationofScP (Fig.9A);anteriorbranchofRPdiverging fromRAdistal to termination of ScP; branches of RPwidely di-vergent;MAunbranched,paralleltoposteriorbranchofRP; anteriorbranchofMPparallel toMA,MAbranch-ingslightlydistaltoRPbranching;MProughlyparalleltoposteriorbranchofMA;crossveinsnotwellpreserved.
Holotype.—PI1988.001.001 (Fig. 9B); Alabama Mu-seumofNaturalHistory,UniversityofAlabama,Tuscalo-osa,Alabama;singlewing(nocounterpart)representingforewing (on basis ofmaximum convexity of posteriormarginnearwingmidlength) inventralaspect,extend-ing from just distal of branching ofRP to near apex ofwing; Pottsville Formation (Pennsylvanian), “In spoilpile”,stripmineatWyndam[sic:correctspellingWind-ham]Sprs.,TuscaloosaCounty,Alabama,1October1988,J.Hall,K.Gaddy,collectors.
Etymology.—Thespecificepithetrecognizestheleg-endaryMississippianChiefTuscaloosa,famousforlead-ingabattleagainst theconquistadorHernandodeSoto
carboniferouS inSectS froM alabaMa 15
and for whom the city and county of Tuscaloosa,Ala-bamawerenamed.
Discussion.—ParalogidaeoccuroverawideareaofthecentralandeasternUnitedStatesinNorthAmerica,with specimens recorded from the Permian of Kansas,and the Carboniferous of Rhode Island, Illinois, andnowAlabama. Recently,Nel et al. (2009) reported thefirstparalogidfromEurope,describingthenewspeciesParalogopsis hispanicus, fromCordoba, Spain. The fam-ily Paralogidae was established by Handlirsch (1906b)forthegenusParalogusScudder,1893,towhichheaddedhisgenusParalogopsisHandlirsch,1911,afewyearslater.Carpenter (1931) subsequentlydescribed thegenusOli-gotypus, andmany years later newly circumscribed thefamily,relegatingParalogopsis longipesHandlirsch,1911tofamilyIncertae sedis (Carpenter,1960,1992c). Carpenter(1960)alsoprovidedcorrectionstoportionsofScudder’s(1893) description and Fraser’s (1957) illustration anddiscussionofParalogus.Carpenter(1960)notedthatthemajordifferencesbetweenthetwoparalogidgeneraarethemuchgreaterconvexityoftheposteriormarginofthewinginParalogus,andthetruebranchingoftheveinsin
theMAfieldofParalogusasopposedtotheirregularve-nationpatternoftheoriginsoftheveinsintheMAfieldinOligotypus. Nonetheless,allparalogidshaveScPter-minatingnearer thewingmidlength than theapex, thebranchesofRPwidelydivergent,andarelativelyreducednumberofcrossveinsbycomparisonwithMeganeuridae.
Oligotypus tuscaloosaediffersfromO. tillyardiCarpen-ter,1931inoverallsize(thewingofthelatterspeciesis50mmlongand11mmwide,abouta third the sizeofthe new species), and in beingmore slender (width tolength5.6–7mminO. tuscaloosae,4.5mminO. tillyardi).ThelongitudinalveinsandbranchesterminateatamorenearlyperpendicularangletotheposteriorwingmargininO. tuscaloosae and inParalogus versusatmorenearlyacute angles inO. tillyardi andO. makowski CarpenterandRichardson, 1971; thus the terminationsof the lon-gitudinal veins inO. tuscaloosae aremore basal than inO. tillyardi andO. makowski.ItshouldbenotedthatNelet al. (2009) recently revised the Permo-Carboniferousgriffenflies,andquestionedtheplacementofO. makowskiinParalogidae;theydidnot,however,definitivelyassignthespecies toanother family. WethereforecompareO.
Fig.8.Agaeoleptoptera uniotempla newgenusandspecies(Megasecoptera:Ancopteridae):A.Reconstructionofwingvenationwithconvexity(+)orconcavity(-)ofwingindorsalaspectnoted.B.Photomicrographofpart(UCM2368)(PV2005.007.0260.001).C.Photomicrographofcounter-part(UCM2369)(PV2005.007.0260.002).Photographsandillustrationtosamescale.
Scientific PaPerS, natural HiStory MuSeuM, tHe univerSity of KanSaS16
tuscaloosaewith both O. tillyardi andO. makowski. Car-penterandRichardson(1971)estimatedO. makowski tobe90mminlengthand18mmwideforaratioof5.0,nearlyasslenderasO. tuscaloosae,andbothspecieshaveamoretaperedapexthanthatofO. tillyardi. TheforewingScPterminatesmoredistallyinO. tuscaloosaethanintheotherOligotypusspecies.Thatandthesharpbendofthelongi-tudinalveinstowardtheposteriormarginmeansthatScPterminatesatthelevelofterminationofMPinO. tusca-
loosae,butatthelevelofterminationofCuPintheotherspeciesofOligotypus.Theargumentmightbemadethatthesumofthesedifferenceswouldbesufficientforustohave established anewgenus forO. tuscaloosae. How-ever, since the description is based solely on the distalhalfofthewing,weprefertotakethemoreconservativestanceofplacingthespeciesasbasalwithinOligotypusforthemomentanduntilmorecompletely-preservedmate-rialisdiscovered.
Fig.9. Oligotypus tuscaloosaenewspecies(Protodonata:Paralogidae):A.Reconstructionofwingvenation,withvenationalnomenclaturefol-lowingthatofRiekandKukalová-Peck(1984)asamendedbyNeletal.(1993),Bechly(1996),Flecketal.(2003),andRehn(2003);theuseof“+”or“–”afteraveinidentityindicatesconvexityorconcavity,respectively.B.Photomicrographofholotype;distalhalfofforewingindorsalaspect(PI1988.0001.0001).
carboniferouS inSectS froM alabaMa 17
DISCUSSION
TodatetheinsectsrecoveredfromthePottsvilleFor-mationareallofpalaeopterousorders,lineagesthatwerediverseandabundantduringtheLatePaleozoic.Whileindicative of a unique fauna, all of the taxa recoveredthusfarexhibitconnectionswithothermajordepositsinNorthAmericaandEurope.Giventhediversityofprimi-tiveneopteraninsects,includingearlyDictyoptera,pres-entinotherCarboniferousdepositsinNorthAmerica,itissurprisingthatsuchtaxahavenotbeenrecoveredyetfromthedeposit inAlabama. Although insectsarenotabundantinthedeposit,continuedexcavationlikelywillreveal additional diversity, particularly primitive Ne-optera, andperhaps includingPaoliidae,putatively themost basal lineage of neopteran insects. ThePottsvilleinsectsareofconsiderableinterestgiventhattheyarethesoutheasternmostrecordsofUpperCarboniferousinsects
intheUnitedStatesandtherebyrepresentaninterestingbiogeographic comparison with the well documentedandmorenortherlyorwestern faunasof Illinois,Penn-sylvania,andKansas.Thecurrentsampling,whilesmall,highlightsthatnumerouslineageswerepresentandthatinsects should be intensely sought from the PottsvilleFormation.Inaddition,allofthespecimensrecoveredtodateareoflargerinsectsanditispossiblethatpastcol-lectingeffortsoverlookedmorefaintlypreservedormin-utewings,representingthevastmajorityofinsectdiver-sity(Carpenter,1992c;BeckemeyerandHall,2007),evenduringthehyperoxicatmospheresoftheLatePaleozoic.Thus,thepotentialvalueofmaterialfromthesedepositsis considerable for their broader implication to under-standLatePaleozoicinsectdiversityanddistributions.
LITERATURECITED
Agnus,A.N. 1902. Description d’un néuroptère fossile nouveau,Homoioptera gigantea.BulletindelaSociétéEntomologiquedeFrance7:259–261,+1pl.
Atkinson,T.P.2005.ArthropodbodyfossilsfromtheUnionChapelMine.AlabamaPaleontologicalSocietyMonograph1:169–176.
Bechly,G. 1996.MorphologischeUntersuchungen amFlügelgeäderderrezentenLibellenundderenStammgruppenvertreter(Insecta;Pterygota;Odonata) unter besonderer Berücksichtigung derPhylogenetischenSystematikunddesGrundplanesder*Odonata.Petalura,SpecialVolume2:1–402+[i].
Beckemeyer,R.J., and J.D.Hall. 2007.Theentomofaunaof theLowerPermianfossilinsectbedsofKansasandOklahoma,USA.AfricanInvertebrates48(1):23–39.
Béthoux,O. 2006. Revision ofCacurgusHandlirsch, 1911, a basalPennsylvanianArchaeorthoptera (Insecta:Neoptera).BulletinofthePeabodyMuseumofNaturalHistory47(1–2):29–35.
Béthoux,O. 2007. Emptying the Paleozoicwastebasket for insects:MembersofaCarboniferous‘protorthopterousfamily’assignedtonaturalgroups.Alavesia1:41–48.
Béthoux,O.2008.RevisionandphylogeneticaffinitiesofthelobeattidspeciesbronsoniDana,1864andsilvaticaLaurentiaux&Laurentiaux-Vieira, 1980 (Pennsylvanian;Archaeorthoptera).ArthropodSystematicsandPhylogeny66(2):145–163.
Béthoux,O.2009.Theearliestbeetleidentified.JournalofPaleontology83(6):931–937.
Béthoux,O., andD.E.G. Briggs. 2008.HowGerarus lost its head:Stem-groupOrthoptera andParaneoptera revisited. SystematicEntomology33(3):529–547.
Béthoux,O.,andA.Nel.2003.RevisionofDiaphanopteraspeciesandnewdiagnosisofDiaphanopteridae(Palaeoptera:Diaphanopterodea).JournalofPaleontology77(5):1016–1020.
Béthoux, O., N.P. Kristensen, andM.S. Engel. 2008. Hennigianphylogeneticsystematicsandthe‘groundplan’vs.‘post-groundplan’approaches:A reply toKukalová-Peck. Evolutionary Biology35(4):317–323.
Brauckmann,C.,andR.Becker.1992.EinneuesRiessen-InsektausdemOber-KarbondesSaarlandes(Palaeodictyoptera:Homoiopteridae).GeologicaetPalaeontologica26:135–141.
Brauckmann,C.,D.R.Chestnut,Jr.,andJ.R.Jennings.1993.Newspilapterid
insect from the Breathitt Formation (middle Pennsylvanian,WestphalianB) of easternKentucky,USA.Neues Jahrbuch fürGeologieundPaläontologie,Monatshefte1993(11):641–647.
Brongniart,C.1893.Recherchespourserviràl’histoiredesinsectesfossilesdestempsprimaires,précédéesd’uneétudesurlanervationdesailesdes insectes [2volumes]. SaintÉtienne,France:Théolier etCie.495pp.,+37pls.
Burnham, L. 1983. Studies onUpperCarboniferous insects. 1. TheGeraridae(OrderProtorthoptera).Psyche90(1–2):1–57.
Buta, R.J.,A.K. Rindsberg, andD.C. Kopaska-Merkel, eds. 2005.PennsylvanianfootprintsintheBlackWarriorBasinofAlabama.AlabamaPaleontologicalSocietyMonograph1:1–387.
Carpenter,F.M.1931.TheLowerPermianinsectsofKansas.Part2.TheordersPalaeodictyoptera,Protodonata, andOdonata.AmericanJournalofScience,Series521:97–139.
Carpenter,F.M.1933.AnewmegasecopteronfromtheCarboniferousofKansas.UniversityofKansasScienceBulletin21(9):365–367.
Carpenter, F.M. 1938. TwoCarboniferous insects from the vicinityofMazonCreek, Illinois.American Journal of Science, Series 536:445–452.
Carpenter,F.M.1940.CarboniferousinsectsfromtheStantonFormation,Kansas.AmericanJournalofScience,Series5238:636–642.
Carpenter,F.M.1944.CarboniferousinsectsfromthevicinityofMazonCreek,Illinois.IllinoisStateMuseum,ScientificPapers3(1):9–20,+4pls.
Carpenter,F.M.1960.StudiesonNorthAmericanCarboniferousinsects.1.TheProtodonata.Psyche67(4):98–110.
Carpenter, F.M. 1963a.Amegasecopteron fromUpperCarboniferousstratainSpain.Psyche70(1):44–49.
Carpenter,F.M.1963b.StudiesonNorthAmericanCarboniferousinsects.2.ThegenusBrodioptera, from theMaritimeProvinces,Canada.Psyche70(1):59–63.
Carpenter, F.M. 1964. Studies onNorthAmericanCarboniferousinsects.3.AspilapteridfromthevicinityofMazonCreek,Illinois(Palaeodictyoptera).Psyche71(3):117–124.
Carpenter,F.M.1965.StudiesonNorthAmericanCarboniferousinsects.4.ThegeneraMetropator,Eubleptus,HapalopteraandHadentomum.Psyche72(2):175–190.
Carpenter,F.M.1967.StudiesonNorthAmericanCarboniferousinsects.
Scientific PaPerS, natural HiStory MuSeuM, tHe univerSity of KanSaS18
5.PalaeodictyopteraandMegasecopterafromIllinoisandTennessee,withadiscussionoftheorderSypharopteroidea.Psyche74(1):58–84.
Carpenter,F.M.1970.FossilinsectsofNewMexico.Psyche77(4):400–412.Carpenter, F.M. 1980. Studies onNorthAmericanCarboniferous
insects.6.UpperCarboniferousinsectsfromPennsylvania.Psyche87(1–2):107–119.
Carpenter, F.M. 1983. Studies onNorthAmericanCarboniferousinsects. 7. The structure and relationships ofEubleptus danielsi (Palaeodictyoptera).Psyche90(1–2):81–95.
Carpenter,F.M.1987.ReviewoftheextinctfamilySyntonopteridae(orderuncertain).Psyche94(3–4):373–388.
Carpenter,F.M.1992a.StudiesofNorthAmericanCarboniferousinsects.8.NewPalaeodictyopterafromKansas,U.S.A.Psyche99(2–3):141–146.
Carpenter, F.M. 1992b. Studies onNorthAmericanCarboniferousinsects. 9.A new species of Eubleptidae fromMazonCreek(Palaeodictyoptera).Psyche99(2–3):147–152.
Carpenter,F.M.1992c.SuperclassHexapoda,pp.1–655,inKaesler,R.L.(ed.),TreatiseonInvertebratePaleontology,PartR,Arthropoda4[volumes3and4].Boulder:GeologicalSocietyofAmerica.xxii+655pp.
Carpenter,F.M.1997. Insecta,pp.184–193, inShabica,C.W.,andA.A.Hay(eds.),Richardon’sGuidetotheFossilFaunaofMazonCreek.Chicago:NortheasternIllinoisUniversity.xvii+[i]+308+[1]pp.
Carpenter,F.M.,andE.S.Richardson,Jr.1968.MegasecopterousnymphsinPennsylvanianconcretionsfromIllinois.Psyche75(4):295–309.
Carpenter, F.M., andE.S.Richardson, Jr. 1971.Additional insects inPennsylvanianconcretionsfromIllinois.Psyche78(4):267–295.
Carpenter,F.M.,andE.S.Richardson,Jr.1976.StructureandrelationshipsoftheUpperCarboniferousinsect,Eucaenus ovalis(Protorthoptera:Eucaenidae).Psyche83(3–4):223–242.
Copeland,M.J.1957.ThearthropodfaunaoftheUpperCarboniferousrocksoftheMaritimeProvinces.MemoirsoftheGeologicalSurveyofCanada286:1–110.
Crampton,G.C.1930.ThewingsoftheremarkablearchaicmecopteronNotiothauma reediMcLachlanwithremarksontheirprotoblattoidaffinities.Psyche37(1):83–103.
Dana,J.D.1864.OnfossilinsectsfromtheCarboniferousformationinIllinois.AmericanJournalofScience,Series237:34–35.
Fleck, G., G. Bechly, X.Martínez-Delclòs, E.A. Jarzembowski, R.Coram, andA.Nel. 2003. Phylogeny and classification of theStenophlebioptera(Odonata:Epiproctophora).AnnalesdelaSociétéEntomologiquedeFrance39(1):55–93.
Fraser,F.C.1957.AreclassificationoftheorderOdonata.RoyalZoologicalSocietyofNewSouthWales,Handbook12:1–133.
Garrouste, R.,A.Nel, andG.Gand. 2009.New fossil arthropods(NotostracaandInsecta:Syntonopterida)intheContinentalmiddlePermianofProvence(Bas-ArgensBasin,France).ComptesRendusPalevol8(1):49–57.
Goldenberg,F.1877.FaunaSaraepontanafossilis.DiefossilenThiereausderSteinkohlenformationvonSaarbrücken,2Heft.Saarbrücken,Germany:Möllinger.54pp.,+2pls.
Grimaldi,D.,andM.S.Engel.2005.EvolutionoftheInsects.Cambridge,U.K.:CambridgeUniversityPress.xv+755pp.
Guthörl,P.1934.DieArthropodenausdemKarbonundPermdesSaar-Nahe-Pflaz-Gebietes.AbhandlungenderKöniglich-PreussischenGeologischenLandesanstalt164:1–219.
Handlirsch,A.1906a.RevisionofAmericanPaleozoicinsects.ProceedingsoftheUnitedStatesNationalMuseum29:663–820.
Handlirsch,A. 1906b.DieFossilen InsektenunddiePhylogeniederrezentenFormen:EinHandbuchfürPaläontologenundZoologen.Leipzig,Germany:Engelmann.ix+1–640pp.,+pls.1–36.
Handlirsch,A.1911.NewPaleozoicinsectsfromthevicinityofMazonCreek,Illinois.AmericanJournalofScience,Series431(194):297–326,353–377.
Handlirsch,A.1919.RevisionderpalaeozoischenInsekten.DenkschriftenderKaiserlichenAkademiederWissenschaften,Wien,mathematisch-naturwissenschaftlicheKlasse96:511–592.
Keynes,R.2001.Annie’sBox:CharlesDarwin,HisDaughterandHumanEvolution.London,U.K.:FourthEstate.xiv+331pp.
Kukalová, J. 1960. New Palaeodictyoptera of the Carboniferousand Permian of Czechoslovakia. SborníkÚstředního ústavuGeologického,oddílGeologický25:239–251,+2pls.
Kukalová,J.1969.RevisionalstudyoftheorderPalaeodictyopteraintheUpperCarboniferousshalesofCommentry,France.PartII.Psyche76(4):439–486.
Kukalová-Peck, J. 1975.Megasecoptera from theLowerPermian ofMoravia.Psyche82(1):1–19.
Kukalová-Peck,J.1985.EphemeroidwingvenationbasedonnewgiganticCarboniferousmayflies andbasicmorphology,phylogeny, andmetamorphosisofpterygoteinsects(Insecta,Ephemerida).CanadianJournalofZoology63(4):933–955.
Kukalová-Peck, J. 1987.NewCarboniferousDiplura,Monura, andThysanura, the hexapodgroundplan, and the role of thoraciclobesintheoriginofwings(Insecta).CanadianJournalofZoology65(10):2327–2345.
Kukalová-Peck,J.1997a.MazonCreekinsectfossils:Theoriginofinsectwings and clues about the originof insectmetamorphosis, pp.194–207,inShabica,C.W.,andA.A.Hay(eds.),Richardon’sGuidetotheFossilFaunaofMazonCreek.Chicago:NortheasternIllinoisUniversity.xvii+[i]+308+[1]pp.
Kukalová-Peck,J.1997b.Arthropodphylogenyand‘basal’morphologicalstructures,pp. 247–268, inFortey,R.A., andR.H.Thomas (eds.),ArthropodRelationships.London:ChapmanandHall.xii+383pp.
Kukalová-Peck,J.,andC.Brauckmann.1990.Wingfoldinginpterygoteinsects, and the oldestDiaphanopterodea from the early LateCarboniferous ofWestGermany.Canadian Journal ofZoology68(6):1104–1111.
Kukalová-Peck,J.,andE.S.Richardson,Jr.1983.NewHomoiopteridae(Insecta:Palaeodictyoptera)withwing articulation fromUpperCarboniferousstrataofMazonCreek,Illinois.CanadianJournalofZoology61(7):1670–1687.
Lacefield,J.1993.Arecentdiscovery.NatureSouth:TheMagazineoftheAlabamaNaturalHistorySociety3(4):11.
Lacefield,J.2000.LostWorldsinAlabamaRocks:AGuidetotheState’sAncient Life andLandscapes. Tuscaloosa:AlabamaGeologicalSociety.123pp.
Lameere,A. 1900.Manuelde la faunadeBelgique:Tome II, Insectesinférieurs.Bruxelles,Belgium:Lambertin.858pp.
Laurentiaux,D.1953.ClassedesInsectes(InsectaLinné1758),pp.397–527,inPiveteau,J.(ed.),TraitédePaléontologie,TomeIII,LesFormesUltimesd’Invertébrés,MorphologieetÉvolution:Onychophores,Arthropodes,Échinodermes,Stomocordés.Paris,France:MassonetCie.1063pp.
Lewis,S.E.1979.Anewspeciesofinsect(Protorthoptera:Narkemidae)from theVerdigris Formation (Pennsylvanian) ofwest centralMissouri.JournalofPaleontology53(3):754–756.
McComas,G.A., andR.H.Mapes. 1988. Fauna associatedwith thePennsylvanianfloralzonesofthe7-11Mine,ColumbianaCounty,northeasternOhio.OhioJournalofScience88(1):53–55.
Murrie,G.W.,W.P.Diamond,andS.W.Lambert.1976.Geologyof theMaryLeeGroupofCoalbeds,BlackWarriorCoalBasin,Alabama.Washington,D.C.:BureauofMines,UnitedStatesDepartmentoftheInterior.ii+49pp.
Nel,A.,X.Martínez-Delclòs,J.-C.Paicheler,andM.Henrotay.1993.Les‘Anisozygoptera’ fossiles.Phylogénie et classification (Odonata).Martinia,NuméroHors-Série3:1–311.
Nel,A.,G.Fleck,R.Garrouste,G.Gand,J.Lapeyrie,S.M.Bybee,andJ. Prokop. 2009. Revision of Permo-Carboniferous griffenflies(Insecta:Odonatoptera:Meganisoptera)baseduponnewspeciesand redescriptionof selectedpoorlyknown taxa fromEurasia.Palaeontographica,AbteilungA, Paläozoologie-Stratigraphie289(4–5):89–121.
Nelson, C.R., andW.D. Tidwell. 1987.Brodioptera stricklani n. sp.
carboniferouS inSectS froM alabaMa 19
(Megasecoptera:Brodiopteridae),anewfossilinsectfromtheUpperManningCanyonShaleFormation,Utah(lowermostNamurianB).Psyche94(3–4):309–316.
Pashin,J.C.2005.PottsvillestratigraphyandtheUnionChapelLagerstätte.AlabamaPaleontologicalSocietyMonograph1:39–58.
Pearse,A.S.1936.ZoologicalNames:AListofPhyla,Classes,andOrders.Durham:DukeUniversityPress.24pp.
Pearse,A.S.1947.ZoologicalNames:AListofPhyla,Classes,andOrders.Durham:DukeUniversityPress.26pp.
Peng,D.-C.,Y.-C.Hong, andZ.-J. Zhang. 2005.Namurian insects(Diaphanopterodea)fromQilianshanMountains,China.GeologicalBulletinofChina24(3):219–234.[InChinese]
Prokop,J.,andA.Nel.2009.SystematicpositionofTriplosoba,hithertotheoldestmayfly,fromUpperCarboniferousofCommentryincentralFrance (Insecta: Palaeodictyopterida). Systematic Entomology34(4):610–615.
Prokop,J.,andD.Ren.2007.NewsignificantfossilinsectsfromtheUpperCarboniferousofNingxia innorthernChina (Palaeodictyoptera,Archaeorthoptera).EuropeanJournalofEntomology104(2):267–275.
Prokop, J., R. Smith, E.A. Jarzembowski, andA.Nel. 2006.Newhomoiopterids from theLateCarboniferousofEngland (Insecta:Palaeodictyoptera).ComptesRendusPalevol5(7):867–873.
Prokop,J.,A.Nel,andA.Tenny.2010.OnthephylogeneticpositionofthepalaeopteranSyntonopteroidea(Insecta:Ephemeroptera),withanewspeciesfromtheUpperCarboniferousofEngland.Organisms,DiversityandEvolution10(4):331–340.
Pruvost,P.1919. Introductiona l’étudede terrainhouillerduPas-de-Calais:La faune continentaledu terrainhouillerduNordde laFrance,pp. 97–321, inMemoirepour servir a l’explicationde lacartegeologiquedelaFrance.Paris:ImprimereNationale.xxxii+584pp.,+29pls.
Rasnitsyn,A.P.,D.S.Aristov,A.V.Gorochov, J.M.Rowland, andN.D.Sinitshenkova. 2004. Importantnew insect fossils fromCarrizoArroyoand thePermo-Carboniferous faunalboundary.Bulletinof theNewMexicoMuseumofNaturalHistoryandScience25:215–246.
Rehn,A.C.2003.Phylogeneticanalysisofhigher-levelrelationshipsofOdonata.SystematicEntomology28(2):181–239.
Richardson,Jr.,E.S.1956.PennsylvanianinvertebratesoftheMazonCreekarea,Illinois.Fieldiana:Geology12(1):3–76.
Riek,E.F.,andJ.Kukalová-Peck.1984.Anewinterpretationofdragonflywing venation based upon Early Carboniferous fossils fromArgentina (Insecta:Odonatoidea) andbasic character states inpterygotewings.CanadianJournalofZoology62(6):1150–1166.
Scudder,S.H.1868a.RemarksontwofossilinsectsfromtheCarboniferousformationinAmerica.ProceedingsoftheBostonSocietyofNaturalHistory11:150–151.
Scudder, S.H. 1868b.Descriptions of fossil insects, foundonMazonCreek,anearMorris,GrundyCo.,Ill.GeologicalSurveyofIllinois3:566–572.
Scudder, S.H. 1868c. [Descriptions of Palaeozoic insects fromNovaScotiaandNewBrunswick],pp.202–206,inDawson,J.W.,OnsomeremainsofPalaeozoic insectsrecentlydiscovered inNovaScotiaandNewBrunswick.CanadianNaturalistandGeologist3:202–206.
Scudder,S.H.1878.ACarboniferoustermesfromIllinois.ProceedingsoftheBostonSocietyofNaturalHistory19:300–301.
Scudder,S.H.1879.Palaeozoiccockroaches:Acompleterevisionofthespeciesofbothworlds,withanessaytowardtheirclassification.MemoirsoftheBostonSocietyofNaturalHistory3:23–134.
Scudder,S.H.1885a.Palaeodictyoptera:OrtheaffinitiesandclassificationofPaleozoicHexapoda.MemoirsoftheBostonSocietyofNaturalHistory3:319–351.
Scudder,S.H.1885b.Newgeneraandspeciesoffossilcockroaches,fromtheolderAmericanrocks.ProceedingsoftheAcademyofNaturalSciences1885:34–39.
Scudder,S.H.1893.InsectfaunaoftheRhodeIslandcoalfield.BulletinoftheUnitedStatesGeologicalSurvey101:1–21.
Scudder,S.H.1895.Revisionof theAmericanfossilcockroaches,withdescriptionsofnewforms.BulletinoftheUnitedStatesGeologicalSurvey124:5–176.
Sellards,E.H.1904.AstudyofthestructureofPaleozoiccockroaches,withdescriptionsofnewformsfromtheCoalMeasures.AmericanJournalofScience,Series418:113–134,213–227.
Sinitshenkova,N.D.2002.SuperorderDictyoneurideaHandlirsch,1906(=Palaeodictyopteroidea),pp.115–124,inRasnitsyn,A.P.,andD.L.J.Quicke (eds.),History of Insects.Dordrecht, TheNetherlands:KluwerAcademicPublishers.xii+517pp.
Wootton,R.J.,andJ.Kukalová-Peck.2000.FlightadaptationsinPalaeozoicPalaeoptera(Insecta).BiologicalReviews75:129–167.
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