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

Scientific PaPerS, natural HiStory MuSeuM, tHe univerSity of KanSaS2

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.

carboniferouS inSectS froM alabaMa 3

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


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.


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.


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


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).


[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).


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.


Fig.5.Pharciphyzelus lacefieldi newgenusandspecies(Palaeodictyoptera:Homoiopteridae):A.Reconstructionofwingvenation;BRdenotesanalbridgeformedbyconvexridgeofstiffenedmembranesensuKukalová-PeckandRichardson(1993).B.Photomicrographofholotype(noknowncounterpart),comprisingbasal70%offorewing(PI1993.0001.0001).


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.


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.


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).


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


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.


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).


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.

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