‚ivist— st—li—n— di €—leontologi— e ƒtr—tigr—fi— volume IPH noF P ppF IVQEIWH tuly PHIR

e xi‡ qix ƒ exh ƒ€igsiƒ yp „ri pewsv‰ „‚egr‰€egrshei

@gyviy€„i‚eD ehi€reqeA p‚yw „ri €€i‚ vehsxsex

@wshhvi „‚seƒƒsgA yp wyx„i ƒex qsy‚qsy

I PB I

ve ‚e ƒ„‚ehe D we„„iy wyx„eqxe 8 exh‚ie „sx„y‚s

‚e™eivedX pe˜ru—ry TD PHIRY —™™eptedX tune IQD PHIR

uey wordsX possil snse™t—D „r—™hyp—™hid—eD new genus —nd speE its glo˜—l p—leontologi™—l me—ning of its m—rine verteE

™iesD pred—torD widdle „ri—ssi™D p—leoenvironmentF

˜r—te f—un—sF

‚eg—rding the u—lks™hieferzoneD topof weride

e˜str—™tF e new genus —nd spe™ies of fossil „r—™hyp—™hid—e

vimestoneD d—ted v—te v—dini—nD PRH ± PQW w— followE €r—edodromeus s—ngiorgiensis genF nFD nF spF @goleopter—D „r—™hyp—™hiE

d—eD iodromein—eAD with well developed —symmetri™—l m—ndi˜lesD ing ƒto™k—r et —lF @PHIPAD it w—s so f—r renowned for

shorter th—n the he—dD su˜re™t—ngul—r pronotum —nd slender ti˜i—eD

its verte˜r—tesD mostly fishes ˜ut —lso — few reptiles

longer th—n femor—D is des™ri˜ed from the u—lks™hieferzone @uppermost

@vom˜—rdo PHHIY ‚enesto et —lF PHHQY „intori IWWH—Y mem˜er of the weride vimestoneA of the ƒwiss side of wonte ƒ—n

„intori 8 vom˜—rdo PHHUAF roweverD —lso —rthropodsD qiorgioF „r—™hyp—™hid—e —re terrestri—l goleopter— well represented

su™h —s ™on™hostr—™—ns @„intori IWWH˜Y „intori 8 fr—mE in the wesozoi™ fossil re™ord ˜y the extin™t f—mily iodromein—eF p

to nowD more th—n QQ fossil spe™ies ˜elonging to IQ gener— of iodroE ˜ill— IWWIA —nd mysidi—™e—ns @v—rghi et —lFD su˜mittedAD

mein—e h—ve ˜een des™ri˜edF „he new spe™ies is p—rt of — ™olle™tion of

—re import—nt in the u—lks™hieferzone fossil —ssemE fossil inse™ts th—t in™ludes mem˜ers of —t le—st six different ordersF

˜l—gesF f—sed on the e™ology of ™orresponding ext—nt t—x—D this inse™t —ssemE

huring the fieldwork ™—rried out ˜etween IWWU ˜l—ge provides new insights on the p—leoenvironment of wonte ƒ—n

qiorgio —re— during the v—te v—dini—nF „he —ssem˜l—ge in™ludes terresE —nd PHHQ in the vower u—lks™hieferzone —t the †—l w—r—

tri—l t—x— th—t support the presen™e of emerged l—nd —nd t—x— with longE

site h ne—r werideD —˜out IT spe™imens of inse™ts @˜oth

l—sting —qu—ti™ l—rv—l st—ge @—s €le™opter— —nd iphemeropter—A th—t ™onE

—dult —nd l—rv—l formsA h—ve ˜een ™olle™tedF ynly two firm the presen™e of st—˜le fresh w—ter ˜—sinsF

spe™imens h—ve ˜een so f—r des™ri˜ed —s — new genus —nd

spe™ies of iphemeropter— @urzeminski 8 vom˜—rdo

PHHIAF sn the s—me p—perD urzeminski 8 vom˜—rdo

sntrodu™tion

@PHHIA des™ri˜ed —lso — spe™imen of xoto™upes @goleopE

ter—D ymm—tid—eA from the vower weride vimestoneF wonte ƒ—n qiorgio is — fossil v—gerst—tte known È

th

‚e™entlyD — new spe™ies of h—syleptid—e @er™h—eogn—t—A ™entury due to industri—l exploit—tion of sin™e the IW

from the pper u—lks™hieferzone w—s des™ri˜ed ˜y ˜ituminous sh—les —s well —s for the numerous p—leonE

fe™hley 8 ƒto™k—r @PHIIAF „hese findings m—ke the tologi™—l ex™—v—tions th—t took pl—™e m—inly in the l—st

u—lks™hieferzone of wonte ƒ—n qiorgio — unique site ™enturyF sn timeD it h—s yielded sever—l rem—rk—˜le fossils

for widdle „ri—ssi™ inse™tsF of m—rine fishes —nd reptiles —s well —s inverte˜r—tes —nd

‡e herein des™ri˜e — new spe™ies —nd genus of terrestri—l pl—nt rem—insF vying —™ross the st—li—nEƒwiss

iodromein—e @goleopter—D „r—™hyp—™hid—eAF „r—™hyp—E ˜ound—ry to the ƒouth of the geresio v—ke @pigF IAD this

—re— is now in the ‡orld rerit—ge vist of xiƒgy for ™hyd—e in™ludes the single ext—nt su˜f—mily „r—™hyp—E

I hip—rtimento di ƒ™ienze dell— „err— ––erdito hesio99D †i— w—ngi—g—lli QRD sEPHIQQ wil—noD st—lyF iEm—ilX l—ur—Fstr—d—dunimiFitD

—ndre—FtintoridunimiFit

P hip—rtimento di ƒ™ienze egr—rie e em˜ient—li E niversit—Á degli ƒtudi di wil—noD †i— qiov—nni gelori— P E PHIQQ wil—noD st—lyF

iEm—ilX m—tteoFmont—gn—dunimiFit B gorresponding —uthor

IVR ƒtr—d— vFD wont—gn— wF 8 „intori eF

pigF I E †—l w—r— site h E geogr—phi™

positionF

the deposition—l ˜—sinF eltern—tivelyD it fell dire™tly on ™hin—e @two gener— des™ri˜edX „r—™hyp—™hus —nd ƒystoE

losom—AD for — tot—l of six spe™ies @w—ddison et —lF PHHWA the w—ter surf—™e while flyingD flo—ted for some time

—nd the extin™t su˜f—mily iodromein—e @€onom—renko undergoing — r—pid initi—l de™—y in the w—rm superfi™i—l

IWUUAF iodromein—e is well represented in the fossil reE w—ters ˜efore sinking to the ˜ottomD where ˜uri—l —nd

™ord from the „ri—ssi™ to the gret—™eous —nd it is mu™h fossiliz—tion then took pl—™eF woreoverD sin™e flo—ting

more diverse th—n „r—™hyp—™hin—e @€onom—renko inse™t rem—ins —re su˜je™ted to pred—tionD they were

IWWIAF et presentD —t le—st QQ fossil spe™ies ˜elonging pro˜—˜ly tr—nsported only for short dist—n™esF „he

to IQ gener— of iodromein—e h—ve ˜een des™ri˜ed @€oE

time l—g ˜etween de—th —nd ˜uri—l determined the exE

nom—renko IWUUY ƒori—no et —lF PHHUY rong IWVQY ti— 8

tent of soft tissues de™—yX when the ˜ody re—™hed the

‚en PHIIY ‡—ng et —lF PHIPY €onom—renko 8 †olkov

˜ottom —nd possi˜lyD during initi—l postE˜uri—l ™omp—™E

PHIQAF „he oldest gener— —re from the „ermin—l €ermi—n

tion of the sedimentD the dis—rti™ul—tion ˜e™—me —pp—rE

@€onom—renko 8 †olkov PHIQA —nd only one other

entF sn the spe™imenD the ˜ody l—yout is preserved so

genus @ƒogdodromeus €onom—renkoD IWUUA is from the

we ™—n infer th—t de™—y w—s time limited —nd the deE

„ri—ssi™ @€onom—renko IWUUAF elsoD during our preliE

position—l environment w—s without —ny ™urrent th—t min—ry survey on this inse™t ™olle™tionD the undes™ri˜ed

™ould helpin the dispers—lof the somewh—t loose ˜ody spe™imens were tent—tively —ssigned to six orders

p—rtsF n—mely iphemeropter—D yrthopter—D €le™opter—D imE

„he degree of ™onserv—tion of the other spe™imens

˜iopter—D remipter— —nd goleopter—F f—sed on the ™urE

v—ries gre—tlyD pro˜—˜ly re™ording strong v—ri—tions in rent knowledge —˜out the e™ology of these ordersD ˜—sed

the t—phonomi™—l historyF sn — few inst—n™esD only he—dD on ˜oth ext—nt —nd fossil spe™iesD we provide new inE

thor—x —nd —˜domen outline is preservedD while in other sights on the p—leoenvironment of wonte ƒ—n qiorgio

™—ses —lso —ppend—ges rem—inedF sn —lmost oneEh—lf of —t le—st during the v—te v—dini—nF

the fossils soft tissues preserv—tion o™™ursD pro˜—˜ly due

to — phosph—tiz—tion pro™essD sometimes preserving inE

tern—l stru™tures in det—ilF snse™t t—phonomy

„he spe™imen under study preserves the gener—l

l—yout of the ˜odyF xonethelessD ˜ody segments —nd

w—teri—l —nd methods

legsD —nd to — lesser extentD he—d p—rts —nd —˜domen —re

„he spe™imen h—s ˜een ™olle™ted in lo™—lity h in †—l w—r—D ne—r displ—™ed from their —n—tomi™—l position in the living

weride @g—nton „i™inoD ƒwitzerl—ndA —nd is deposited —t wuseo g—nE —nim—lF „his l—yout suggests th—t the spe™imen went

ton—le di ƒtori— x—tur—le di vug—no @wgƒxAF ƒpe™imen m—nipul—tion through — first st—ge of de™—y in su˜—eri—l —ndGor superE

—nd me—sures were ™ompleted with the —uxili—ry use of the stereo miE

fi™i—l —qu—ti™ environmentD during whi™h soft tissues

™ros™ope vei™— wƒS —nd —n o™ul—r mi™rometerF sm—ges were —™quired

p—rti—lly degr—dedF „hereforeD either the inse™t died

˜y digit—l ™—mer— g—non ios RSH mounted on — m—™hinery in order to

—nd p—rti—lly de™—yed on the groundD its origin—l h—˜iE s™—n the s—mple —t different fo™us l—yers th—t were mounted with erE

t—tD —nd w—s then tr—nsported ˜y — –high9 energy eventD ene ƒt—™ker IFH @ƒtudent iditionAF hr—wings were m—de with ™—mer—

lu™id— —tt—™hed to the mi™ros™opeF su™h —s — flood following he—vy r—ins or just windD to

e new goleopter— from the v—dini—n of wonte ƒ—n qiorgio IVS

pigF P E eA €re—dodromeus s—ngiorE

giensis genF nD spF nF vength

from —pex of m—ndi˜ules to

—pex of elytr— a IHFV mmF fA

dr—wing of the spe™imenF

ƒystem—ti™ p—leontology hes™ription

elmost ™omplete impression in dors—l viewD l—™kE

yrder goleopter— vinn—eusD IUSV

ing —ntenn—eF re—d —nd —˜domen in n—tur—l positionD

ƒu˜order edeph—g— ƒ™hellen˜ergD IVHT single dis—rti™ul—ted elytron preservedF vength from

—pex of m—ndi˜ules to —pex of elytr— of IHFV mmY width p—mily „r—™hyp—™hid—e „homsonD IVSU

—t ˜—se of elytr— of RFT mm @lengthEwidth r—tio PFQRAF

ƒu˜f—mily iodromein—e €onom—renkoD IWUU

ell three legs on the right side preservedF wid —nd

qenus €r—edodromeus genF novF

hind femor— r—ther l—rge —nd ro˜ustY m—ximum width of

„ype spe™iesX €r—edodromeus s—ngiorgiensis nF spF ˜y monotypyF

right mid femor— is HFUW mmF „i˜i—e thin —nd slenderF

„he preserved t—rsi @sCssA elong—ted —nd slimY only two itymologyX „he n—me derived from the v—tin ver˜ ––pr—edo99D

fore t—rsomers distinguish—˜leD first one enl—rgedF yn whi™h me—ns to preyD ™onsidering its strong —nd sh—rp m—ndi˜les th—t

we supposed to ˜e used to ™—t™h preyD —nd the suffix ––±dromeus99D whi™h the left side preserved only ssCsss femor— —nd sss ti˜i—F

me—ns ––runner99D from its ™ursory legsF

‚ight fore ti˜i—e @length IFWP mmA h—r˜ouring — simple

vo™us typi™usX ƒite hD †—l w—r— ne—r werideD wendrisioD g—nE

sul™—te —ntenn— ™le—ner org—n with enl—rged dist—l —re— ton „i™inoD ƒwitzerl—ndF

—nd —n —pi™—l spur @HFPR mmAF pive t—rsi @tot—l length hi—gnosisX „he gener—l h—˜itus of the new t—xon @genus or

spe™iesAD — simple sul™—te —ntenn— ™le—ner in the dist—l p—rt of the ti˜i—eD IFTP mmA on s right leg ™le—rly visi˜leF ƒlender ti˜i—eF

the met—™ox— sep—r—ting the thor—x —nd —˜domenD the met—episternum

re—d ˜igD longer th—n widerD length of QFH mm

re—™hing the meso™ox—l ™—vities prompts its —s™ription to the to the

m—ndi˜les ex™luded —nd of QFRP m—ndi˜les in™ludedD f—mily „r—™hyp—™hid—eD su˜f—mily iodromein—eF „he new t—xon is deE

width —t the eyes PFUT mmF w—ndi˜les —symmetri™—l s™ri˜ed —s new genus —nd spe™ies due to the presen™e of — unique set of

—nd tri—ngul—rD ro˜ust —nd sh—rpwith — sm—ll tooth in ™h—r—™tersX well developed —symmetri™—l m—ndi˜lesY re™t—ngul—r pronoE

tumD longitudin—l groove —˜sentY short met—ventriteY —nd ˜ig he—d with the —nterior se™ondEthird of the inner m—rginF „ot—l

l—rge eyes on the upper surf—™eF

length of IFQV mmD width HFWT mmD lengthEwidth r—tio

of IFRRF €—rti—l impression of l—˜i—l p—lp visi˜le ˜etween

€r—edodromeus s—ngiorgiensis spF nF m—ndi˜lesF

pigF PeD f €ronotum wide @QFI mmAD —nterior m—rgin slightly

™on™—ve with pointed —pi™es @left sideAD posterior m—rgin

rolotypeX wgƒx VRSIF

str—ightF

itymologyX „he n—me derived from the ™olle™ting —re— of

e˜domen p—rti—lly visi˜le with light impressions wonte ƒ—n qiorgio @ƒwitzerl—ndGst—lyAF

of segment—tion preserved on the left sideF vength of hi—gnosisX —s for the genusD ˜eing the only spe™iesF

IVT ƒtr—d— vFD wont—gn— wF 8 „intori eF

—˜domen IFS times the length of mesoE —nd met—thor—™es ˜—se —nd on —n often —noxi™ ˜ottomD —s indi™—ted ˜y

™ommon l—min—ted limestone or m—rlyElimestone l—yers ™om˜inedF

—nd the —lmost gener—l —˜sen™e of ˜iotur˜—tion @„intori

his™ussion IWWH˜Y „intori 8 ‚enesto IWWHY „intori 8 vom˜—rdo

IWWWY vom˜—rdo et —lF PHIPAF gommon —re —lso ™l—yE „he spe™imen h—s ˜een des™ri˜ed ˜—sed on the

™hips ˜edsD often ri™h —lso in d—rk —lg—lEfilm fr—gmentsD preserved morphologi™—l ™h—r—™ters even if some key

pro˜—˜ly rel—ted to storms —ffe™ting the sh—llower p—rt ™h—r—™ters —re l—™kingD pro˜—˜ly due to su˜—ere—l degr—E

of the ˜—sin or the threshold tow—rd the open w—tersF d—tion ˜efore ˜uri—lF fetterEpreserved fe—tures in™lude

„he u—lks™hieferzone ˜—sin w—s ™le—rly —lso sometimes the gener—l h—˜itusD the m—ndi˜les —nd the very develE

—ffe™ted ˜y sudden fresh w—ter supply following stormy oped he—d with dors—l eyesF „he new genus differs from

he—vy r—insD —s there is re™ord of sever—l levels ri™h in €etrodromeus @€onom—renko 8 †olkovD PHIQAD €erE

™on™hostr—™—ns ™rust—™e—ns @„intori IWWH—Y „intori 8 mund— @€onom—renko 8 †olkovD PHIQAD u—r—tom— @€oE

fr—m˜ill— IWWIAF purthermoreD freshw—ter ™ould indu™e nom—renkoD IWUUAD ƒinodromeus @‡—ng et —lFD PHIPAD

strong sudden ™h—nges in the l—goon—l environment oriE €s—™odromeus @€onom—renkoD IWUUAD ˆin˜ini— @rongD

gin—ting m—ss mort—lity events —mong m—rine fishes IWVQA —nd portiseode @ti— 8 ‚enD PHIIA for the —˜sen™e

@„intori IWWH—Y vom˜—rdo et —lF PHIPA —ggr—v—ting the of pun™tu—tions —nsGor stri—e on elytr—F „he gener— €etE

gener—l w—ter ™onditions thus ™—using —˜ortion in the rodromeus @€onom—renko 8 †olkovD PHIQAD €ermund—

m—rine reptile v—rios—urus pregn—nt fem—les @‚enesto @€onom—renko 8 †olkovD PHIQA —nd u—r—tom— @€onoE

et —lF PHHQAF „he freshEw—ters flowing —˜ove the s—lted m—renkoD IWUUA show — lower he—d widthGlength r—tio

ones ™ould —lso indu™e — tempor—ry w—ter str—tifi™—tionD ™omp—red to €r—edodromeus genF nF „he new genus

preventing — gener—l mixingF „his m—de the lower w—ter m—rkedly differs from portiseode —nd from ˆin˜ini—

l—yer less —nd less oxi™ due to the org—ni™ m—tter de™—y —lso for the sh—pe of ˜odyD pronotum —nd he—dF st —lso

—t the ˜ottomF es terrestri—l dwellersD inse™ts were pro˜E differs from ƒogdodromeus @€onom—renkoD IWUUA in the

—˜ly less —ffe™ted ˜y su™h eventsX howeverD storms ™ould longer —˜domen —nd in the sh—pe of the pronotumF

h—ve thrown the inse™ts over the ˜—sin itselfF gomp—red to €l—ty™ox— @€onom—renkoD IWUUA it differs

‡e h—ve identified IQ still undes™ri˜ed spe™imens in the sh—pe of pronotum —nd m—ndi˜lesD in the shorter

—s ˜elonging to six different ordersX iphemeropter—D he—d ™—psule —nd sm—ller eyesF €r—edodromeus genF nF

yrthopter—D €le™opter—D im˜iopter—D remipter— —nd differs from nd— @€onom—renkoD IWUUA —nd from €erE

goleopter—F „hey in™lude ˜oth terrestri—l —nd —qu—ti™ mund— @€onom—renko 8 †olkovD PHIQA in the width

formsD the l—tter —s —dult —nd l—rv—l st—gesF rere we —nd sh—pe of the pronotumD whi™h l—™ks the pointed

™onsider —s —qu—ti™ the inse™t groups th—t require —n —nterior m—rgins —nd is less m—rkedly n—rrower th—n

—qu—ti™ environment during —t le—st one development—l the ˜—se of the elytr—F gomp—red to u—r—dromeus @€oE

st—geF nom—renkoD IWUUA the new t—xon h—s legs with longer

„—˜le I shows the list of the identified t—x—D the femor—D protruding ˜eyond l—ter—l m—rgins of the ˜ody

num˜er of spe™imens for e—™h t—xon —nd their living —nd ti˜i—e longer th—n femor—F „he ™omp—rison with

environmentD ˜—sed on the —ssumption th—t the living feipi—o™—r—˜us @rongD IWVQA is diffi™ultD sin™e the holoE

environment of fossil groups ™—n ˜e inferred from th—t type found ˜y rong is preserved in ventr—l viewD whi™h

of ext—nt represent—tivesF „wo more spe™imens reE we —re missingF enyw—yD —pp—rently in feipi—o™—r—˜us

m—ined unidentifiedF the —˜domen is shorterD the met—thor—x is quite longer

ƒix spe™imensD ˜elonging to four different ordersD —nd femor— —re less strong —nd thi™k th—n in €r—edodroE

—re terrestri—lX they in™lude yrthopter—D im˜iopter— meusF

—ndD —mong goleopter—D „r—™hyp—™hid—e —nd gupediE

d—ec sn —dditionD the morphologi™—l fe—tures of the

two remipter— spe™imens —re ™loser to those of terresE

„he u—lks™hieferzone inse™t —ssem˜l—ge

tri—l f—milies th—n to those of —qu—ti™ onesF emong terE

—nd its p—leoenvironment—l impli™—tions

restri—l groupsD „r—™hyp—™hid—e —re pred—tors —nd modE

„he deposition—l environment for the u—lks™hieE ern represent—tives of the group @„r—™hyp—™hin—eA dwell

ferzone is th—t of — sh—llow l—goonD —dj—™ent to — ™—r˜oE in loose soils in definitely terrestri—l h—˜it—ts @ƒ™hull et

n—ti™ pl—tform @ƒF ƒ—lv—tore holomiteAF „ow—rd ‡estE —lF PHHIAF „he yrthopter— —nd the two remipter— ™olE

xorthG‡est it f—™es — deeper ˜—sin @€erledoE†—renn— le™ted —re terrestri—l —nd the ext—nt represent—tives of

porm—tionA —nd the ™omplex system of ™—r˜on—ti™ pl—tE the groups —re typi™—lly phytoph—gousF

forms of the isino porm—tion further to the i—stD with ‡ithin the fossil inse™ts of wonte ƒ—n qiorgioD

somewh—t limited ™onne™tion to the open —nd deeper —qu—ti™ forms —re well representedF ‡e h—ve —ssigned

se— @„intori IWWH—Y „intori 8 vom˜—rdo IWWWY vom˜—rE seven spe™imens @—dults —nd l—rv—eA to —qu—ti™ groupsD

n—mely iphemeropter— @—mong theseD „intorin— meriE do et —lF PHIPAF ƒediment—tion took pl—™e ˜elow w—ve

e new goleopter— from the v—dini—n of wonte ƒ—n qiorgio IVU

Order Family Species N° of specimens Living environment Strictly freshwater for reproduction and life through all the Tintorina Ephemeroptera Tintorinidae 2 (adults) immature stages; in clean still or running waters. meridensis Larvae feed on plant or, more rarely, animal debris Strictly freshwater for reproduction and life through all the Ephemeroptera nd* nd* 1 (nymphal stage) immature stages; in clean still or running waters. Larvae feed on plant or, more rarely, animal debris Preferentially terrestrial habitats although some species live Orthoptera nd* nd*1 on the shores of ponds, streams, lakes, and rivers phytophagous, otherwise carnivorous/predatory Freshwater, clean, cold and running waters. Plecoptera nd* nd* 2 (1 adult, 1 nymph) Phytophagous or predatorial habits Strictly terrestrial and phytophagous; feeding on leaf litter, moss, bark and lichen. Subsocial behavior, can build tunnels Embioptera nd* nd*1 and chambers woven from the silk they produce on rocks and the bark trees, or in leaf litter Hemiptera nd* nd* 2 Terrestrial, phytophagous or parasitic Strictly terrestrial habitat, both in the larval and adult stage, Praedodromeus Coleoptera Trachypachidae 1 with predatory habits, extant species (Trachypahinae) dwell sangiorgiensis in loose soils. Strictly terrestrial. Larvae: wood borers, typically live in fungus-infested wood, feeding on dead, fungi infested wood. Coleoptera Cupedidae? nd* 1 (elytron, fragment) Adults live in dead subterranean wood such as tree roots, buried structural timber and the rotten cores of living trees or stumps Coleoptera Coptoclavidae? nd* 1 (adult) Freshwater. Predatory habits

Coleoptera nd* nd* 1 (abdomen fragment) nd*

„—˜F I E possil —ssem˜l—ge —nd environment—l fe—turesF BxoteX nd a not determined

densisAD €le™opter— —nd —qu—ti™ goleopter— @gopto™l—viE fe™hley 8 ƒto™k—r @PHIIAF €resen™e of iphemeropter—

d—ecAF ell these t—x— need perm—nent fresh w—ter pools @urzeminski 8 vom˜—rdo PHHIA —nd h—syleptid—e

or sm—ll stre—ms to spend —t le—st one of their developE @fe™hley 8 ƒto™k—r PHIIA spe™imens points to — quite

ment—l st—ge whi™hD in ext—nt w—yflies @iphemeropter—A ™lose sour™e —re— espe™i—lly if we ™onsider th—t the three

—nd ƒtoneflies @€le™opter—A ™—n l—st for up to three ye—rs spe™imens of h—syleptus tri—ssi™us h—ve ˜een found on —

@qrim—ldi 8 ingels PHIRAF single m—ss mort—lity surf—™e @vom˜—rdo et —lF PHIPAF

snterestingly fl—ttode—D whi™h —re the most —˜unE „his me—ns th—t the tr—nsport h—d not to ˜e long otherE

d—nt —nd diverse groupin the m—in „ri—ssi™ inse™t —sE wise — mu™h l—rger s™—ttering of the spe™imens would

sem˜l—ges @enderson et —l IWWVY w—n™uso et —l PHHUY h—ve t—ken pl—™eF

ƒh™her˜—kov PHHVAD —re —˜sent from our fossil —ssemE „he only other re™ord of l—nd dwellers for the

˜l—geF ƒu™h — p—ttern w—s re™orded only from ––e™ozone weride vimestone reg—rded the pl—nts @ƒto™k—r 8 uusE

T99 of the wolteno porm—tion @g—rni—nAD where horseE t—ts™her PHIHAD even if the —uthors did not give — p—E

t—ils were the domin—nt veget—tion @enderson et —lF leoenvironment—l interpret—tion of the l—nd from where

IWWVAF f—shkuev et —l @PHIPA h—ve des™ri˜ed the fossil the pl—nt rem—ins ™—me fromF ƒto™k—r 8 uust—ts™her

inse™t —ssem˜l—ge from the pper funts—ndstein of @PHIHA only wrote —˜out — possi˜le –—rid9 environment

vower pr—n™oni— —nd „huringi—D d—ted enisi—nD in™ludE quite f—r from the ˜—sinF roweverD we h—ve —lso to keep

ing more th—n QHH spe™imensD where ™oleopter— repreE in mind th—t the flor— des™ri˜ed ˜y ƒto™k—r 8 uusE

sent from RV to UV7 of ™olle™ted m—teri—l ˜ut fl—ttode— t—ts™her @PHIHA refers to the g—ssin— fedsD thus —t the

—re —lso presentF topof the vower weride vimestoneD when the gener—l

‚eg—rding the life environment of the u—lks™hieE p—leoenvironment—l ™onditions were pro˜—˜ly different

ferzone inse™tsD the emerged l—nd during the v—te v—diE from those of the u—lks™hieferzoneF ƒo f—rD only m—rine

ni—n in the wonte ƒ—n qiorgio —re— h—s never ˜een dwellersD other th—n the terrestri—l pl—nts —nd — single

pre™isely re™ordedD though some fresh w—ters h—d to inse™t fr—gment @urzeminski 8 vom˜—rdo PHHIAD h—ve

˜e supposed owing to the presen™e of ™on™hostr—™—ns ˜een found in the lower weride vimestone ™omp—red to

@„intori IWWH˜AF „he ƒ—n ƒ—lv—tore holomite surE the sometimes ™ommon esterids @„intori IWWH˜Y vomE

rounded the weride vimestone ˜—sin@sA —nd possi˜ly ˜—rdo et —lF PHIPA in sever—l horizons of the vower —nd

h—d sm—ll tempor—ry isl—nds —s —lre—dy supposed ˜y widdle u—lks™hieferzoneF „he only simil—rity ˜etween

„intori @IWWH˜AD urzeminski 8 vom˜—rdo @PHHIA —nd the u—lks™hieferzone —nd the vower weride vimestone

IVV ƒtr—d— vFD wont—gn— wF 8 „intori eF

†erte˜r—te levels is the presen™e of more m—rly limeE iv—ns @IWVPA suggests th—t „r—™hyp—™hid—e were

stonesD usu—lly l—min—ted —nd ri™her in org—ni™ m—tterF pred—tors or xylomy™etoph—gousF qround ™over from

„husD the preserv—tion windows seem to ˜e rel—ted to — veget—tion w—s s™—r™eD —nd ˜eetles were f—st runners in

higher ™l—y ™ontent th—n in the well ˜edded limestone order to run —™ross the ˜—re —re—s without in™urring in

th—t m—kes upmost of the lower —nd middle weride pred—tionF „his hypothesis fits well with €r—edodroE

vimestoneF ‡hi™h is the origin of the ™l—y —nd in whi™h meus genF nF gener—l h—˜itusD whi™h —ppe—rs slender

w—y it is rel—ted to ˜ottom ™onditions f—vor—˜le to ni™e —nd —gileD with longD thin ™ursory legsF w—ndi˜les —re

preserv—tion is still to ˜e —n—lyzedF well developed —nd —ppe—r —pt to pred—tionD eyes —re of

„he u—lks™hieferzone is often ™h—r—™terized ˜y gre—t dimensionsF „he new genus does not resem˜le the

finely l—min—ted ˜eds re™ording se—son—l —ltern—tion —t more —n™ient gener— €etrodromeus @€onom—renko et

the —noxi™ ˜ottom of the m—rine ˜—sinD influen™ed ˜y †olkovD PHIQAD €ermund— @€onom—renko et †olkovD

™ontinent—l —re—sF vight l—yers —re ˜elieved to h—ve ˜een PHIQA —nd ƒogdodromeus @€onom—renkoD IWUUAD —nd —pE

deposited during the dry ––summer99 —nd d—rk l—yers pe—rs more simil—r to the tur—ssi™ gener— u—r—tom— @€oE

during the ––winter99 r—ining se—son @„intori IWWH—AF e nom—renkoD IWUUA —nd u—r—dormeus @€onom—renkoD

™omp—rison of the surf—™e ™olor on whi™h the inse™ts —re IWUUA —nd in gener—l to the g—r—˜ids h—˜itusF

preserved prompted us to —ssign —dult st—ges of —qu—ti™ „—king into —™™ount —ll the t—phonomi™ —nd enE

t—x— @gopto™l—vid—ecD €le™opter—D iphemeropter—A vironment—l elementsD — possi˜le s™en—rio —t — ™lose disE

m—inly to the ––summer99 dry se—son @light l—min—eAF t—n™e from the deposition—l ˜—sin is th—t of — l—nd with

„he l—rv—l st—ges @iphemeropter—D €le™opter—A —re —sE ponds or sm—ll l—kesD some of them perm—nent th—t were

signed m—inly to the ––winter99 r—iny se—son @d—rk l—miE ™onne™ted to —nd through — riverine systemF et le—st in

n—eAD indi™—timg they were present through the whole the r—iny se—son floodings ™ould ˜ring fresh w—ter

ye—rD —s were the perm—nent fresh w—ter ˜—sinsF dwellers su™h —s estherids —nd inse™t l—rv—e to the ˜—sinF

„he emerged l—nds supporting those fresh w—ters were

pro˜—˜ly not too f—rD —llowing for the terrestri—l inse™t

gon™lusions

spe™imens f—lling into the l—goon to ˜e preserved —lmost

int—™tF „he l—nd pro˜—˜ly —lso supported woods formE „he new t—xon is one of the oldest gener— of „r—E

ing —t le—st — loose org—ni™ m—tter ri™h soil to form the ™hyp—™hid—e des™ri˜ed so f—rF ‚e™entlyD €onom—renko

living environment for terrestri—l inse™ts with pred—tory 8 †olkov @PHIQA des™ri˜ed two new gener— from the

—nd phytoph—gous h—˜itsF „he ™omplex p—leoenvironE termin—l €ermi—n of f—˜ii u—men9D uuznetsk f—sinD ƒiE

ment of the wonte ƒ—n qiorgio —re— th—t is ™oming in ˜eri—F „he other „ri—ssi™ genusD ƒogdodromeus @€onoE

the light —t le—st for the v—te v—dini—n timeD m—y pro˜E mor—nkoD IWUUAD ™omes from the w—dygen porm—tion

—˜ly expl—in the high ˜iodiversity for the fish —ssemE @uyrgyzst—nAD d—ted i—rly g—rni—n @ƒ™her˜—kov PHHVAD

˜l—ge of the u—lks™hieferzoneF e™tu—llyD fish spe™ies —l˜eit ƒ™her˜—kov suggests — v—te v—dini—n —geD ˜—sed

@more th—n PH —™ross the u—lks™hieferzoneA outnumE on the fossil inse™t f—un— primitive ™h—r—™teristi™sF €—E

˜ered other —qu—ti™ m—™roorg—nisms su™h —s m—rine pier —nd ™olle—gues @PHHSA des™ri˜ed the ˜eetles ™olle™E

reptiles @I spe™iesA —nd ™rust—™e—ns @PEQ spe™iesAD so f—r tion from qreÁs — †oltzi—D the fossil l—gerst—tten from the È

no mollusks or e™hinoderms h—ving ˜een re™orded from enisi—n lo™—lity in the †osgesD pr—n™eF yne of the spe™iE

the u—lks™hieferzone itselfF „his must ˜e t—ken into —™E mens @€—pier 8 —lF PHHSF pFIVUD spF QID fig UeA ™—n ˜e

™ount when ™omp—ring the verte˜r—tes f—un—s —™ross the re™ognized —s — „r—™hyp—™hid—eD —lthough its system—ti™

„ethys —fter the re™ent dis™overy of sever—l widdle position w—s not st—tedF €r—edodromeus genF nFD ™olE

„ri—ssi™ verte˜r—te sites in ƒouthern ghin— @ti—ng et —l le™ted in ƒouthern iuropeD therefore exp—nds the r—nge

PHHWY „intori et —lF PHIQ—D PHIQ˜AD espe™i—lly when ™omE of distri˜ution of „ri—ssi™ „r—™hyp—™hid—e to the west

p—ring sites of simil—r —ge su™h —s the ˆingyi @quizhou —nd ™ontri˜utes to support the ide— th—t the high diverE

€rovin™eD ghin—A —nd the u—lks™hieferzone f—un—sD ˜oth sity o˜served in the „r—™hyp—™hid—e fossil re™ords from

d—ted to the v—te v—dini—n @ti et —lF PHIPY „intori et —lF tur—ssi™ to gret—™eous ™ould h—ve st—rted during the

PHIQAF widdle „ri—ssi™F

yf the ext—nt gener— of „r—™hip—™hyd—eD „r—™hyE

p—™hus inh—˜its the rolo—r™ti™ region —nd ƒystolosom— e™knowledgementsF ‡e th—nk gristin— vom˜—rdo for the h—rd

work during long ye—rs of field work —nd for her ™ontri˜ution to the the xeotropi™—l region @qrim—ldi et —lF PHHRAF „r—™hyE

knowledge of the fossils from the u—lks™hieferzoneF w—rkus pel˜erD p—™hid—e fossil re™ord from the „ri—ssi™ to the gret—™E

then ™ur—tor —t the wuseo g—nton—le di stori— x—tur—le in vug—noD

eous is more —˜und—nt in v—ur—si—D ˜ut spe™imens h—ve

supported our ex™—v—tions providing the logisti™ —nd fin—n™i—l supportF

˜een des™ri˜ed from the eustr—li— „r—l˜—g—r fossil fish

„he —uthors would like to th—nk the two —nonymous reviewers for

˜ed @fe—ttie 8 every PHIPA —nd from the gret—™eous of their ™onstru™tive ™omments whi™h h—ve resulted in —n improved

m—nus™riptF the ƒ—nt—n— porm—tionD fr—zil @w—isey IWWIAF

e new goleopter— from the v—dini—n of wonte ƒ—n qiorgio IVW

‚ipi‚ixgiƒ

enderson tFwFD enderson rFwF 8 grui™ksh—nk eF‚FsF nu™le—r genes @goleopter—X g—r—˜id—e —nd „r—™hyp—E

@IWWVA E v—te „ri—ssi™ i™osystem of the woltenoG ™hid—eAF oolF ƒ™rFD QVX RQETPF

vower illiot fiome of ƒouthen efri™—F €—l—eontologyD w—n™uso eFgFD q—llego yFpF 8 w—rtinsExeto ‚FqF @PHHUA E

RIX QVUERPIF „he „ri—ssi™ inse™t f—un— from the vos ‚—stros porE

f—shkuev eFD ƒell tFD eristov hFD €onom—renko eFD ƒinitE m—tion @fermejo f—sinAD v— ‚ioj— €rovin™e @ergentiE

shenkov— xF 8 w—hler rF @PHIPA E snse™ts from the n—AX its ™ontextD t—phonomy —nd p—leo˜iologyF emeE

funts—ndstein of vower pr—n™oni— —nd „huringi—F €—E ghini—n—D RR@PAX QQUEQRVF

l—entolF tFD VTXIUSEIVSF w—isey tF qF @IWWIA E ƒ—nt—n— fossilsX —n illustr—ted —tl—sF

fe—ttie wFqF 8 every ƒF @PHIPA E €—l—eoe™ology —nd p—E „FpFrFD xeptune gityD RSW ppF

l—eoenvironment of the tur—ssi™ „—l˜r—g—r fossil fish €—pier pFD xell eFD qr—uvogelEƒt—mm vF8 q—ll tF gF @PHHSA E

˜edD qulgongD xew ƒouth ‡—lesD eustr—li—F el™herinE v— diversite des goleopter— @snse™t—A du „ri—s d—ns le

g—D QTX RSIERTSF xordEist de l— pr—n™eF qeodiversit—sD PU@PAX IVIEIWWF

fe™hley qF 8 ƒto™k—r ‚F @PHIIA E „he first wesozoi™ re™ord €onom—renko eFqF @IWWIA E edeph—g—F snX er9noldi vF†FD

of the extin™t —pterygote inse™t genus h—syleptus @snE herikin †F†FD €onom—renko eFq 8 xikritin vF

se™t—X er™h—eogn—th—X wonur—X h—syleptid—eA from @idsA E wesozoi™ goleopter—F f—™khuys €u˜lisherD

the „ri—ssi™ of wonte ƒ—n qiorgio @ƒwitzerl—ndAF €—E PVS ppF

l—eodiversityD RX PQEQUF €onom—renko eFqF 8 †olkov eFxF @PHIQA E edemosynoides

iv—ns wFiFqF @IWVPA E i—rly evolution of the edeph—g— ± —si—ti™us w—rtynovD IWQTD the i—rliest unown wemE

some lo™omotor spe™ul—tionsF „he goleopterists fulE ˜er of —n ixt—nt feetle p—mily @snse™t—D goleopter—D

lettinD QT@RAX SWUETHUF „r—™hyp—™hid—eAF €—leontolF tFD RU@TAX THIETHTF

qrim—ldi hF 8 ingel wFƒF @PHHRA E ivolution of the snse™tsF ‚enesto ƒFD vom˜—rdo gFD „intori eF 8 h—nini qF @PHHQA E

g—m˜ridge niversity €ressD USS ppF xothos—urid em˜ryos from the widdle „ri—ssi™ of

rong ‰FgF @IWVQA E widdle tur—ssi™ possil snse™ts in xorth xorthern st—lyX —n insight into the vivip—rity of

ghin—F qeologi™—l €u˜lishing rouseD feijingD volF of xothos—ursc tF †ertF €—leontolFD PQ@RAX WSVEWTIF

tte PPQ ppF ‘in ghinese“F ƒh™her˜—kov hF iF @PHHVA E w—dygenD „ri—ssi™ v—gerst— È

num˜er oneD ˜efore —nd —fter ƒh—rovF el—vesi—D PX IIQE ti gFD ƒh—ng ‡FD hi—o qFD wot—ni ‚F 8 „intori eF @PHIPA E

IPRF fiodiversity —nd ƒtr—tigr—phi™ distri˜ution of the first

ƒori—no gFD hel™los ƒFˆF 8 €onom—renko eFqF @PHHUA E l—te v—dini—n @widdle „ri—ssi™A m—rine verte˜r—te p—uE

feetle —sso™i—tions @snse™t—X goleopter—A from the n— ± ˆingyi p—un— from ƒouth ghin—F tF †ertF €—leonE

f—rremi—n @vower gret—™eousA of ƒp—inF el—vesi—DID tolFD QP @supplF QAX IIUF

VIEVVF ti— „FD vi—ng rFD gh—ng rF 8 ‚en hF @PHIIA E e new genus

ƒto™k—r ‚FD f—umg—rtner €FyF 8 gondon hF @PHIPA E snteE —nd spe™ies of fossil iodromein—e from the ‰ixi—n

gr—ted v—dini—n ˜ioE™hronostr—tigr—phy —nd geo™hroE porm—tion of ‡estern vi—oningD ghin— @goleopter—X

nonology of wonte ƒ—n qiorgio @ƒouthern elpsD edeph—g—X „r—™hyp—™hid—eAF oot—x—D PUQTX TQETVF

ƒwitzerl—ndAF ƒwiss tF qeos™iFD TH@IAX PQWEPTWF ti—ng hFD wot—ni ‚FD r—o ‡FD ‚ieppel yFD ƒun ‰FD „intori eFD

ƒto™k—r ‚F 8 uust—ts™her iF @PHIHA E „he v—dini—n flor— ƒun F 8 ƒ™hmitz vF @PHHWA E fiodiversity —nd ƒeE

from the g—ssin— ˜eds @weride vimestoneD wonte quen™e of the widdle „ri—ssi™ €—nxi—n m—rine ‚eptile

ƒ—n qiorgioD ƒwitzerl—ndAX prelimin—ry resultsF ‚ivF p—un—D quizhou €rovin™eD ghin—F e™t— qeolF ƒinFD VQX

stF €—leontolF ƒtr—tFD IIT@PAX IUQEIVVF RSIERSWF

„intori eF @IWWH—A E „he e™tinopterygi—n pish €roh—le™ites urzeminski ‡F 8 vom˜—rdo gF @PHHIA E xew fossil ipheE

from the „ri—ssi™ of xorthern st—lyF €—l—eontologyD QQX meropter— —nd goleopter— from the v—dini—n @widdle

ISSEIURF „ri—ssi™A of g—nton „i™ino @ƒwitzerl—ndAF ‚ivF stF €—E

„intori eF @IWWH˜A E istherids from the u—lks™hieferzone leontolF ƒtr—tFD IHU@IAX TWEUVF

@„ri—ssi™A of vom˜—rdy @xF st—lyAF snX ‚o˜˜— iF vom˜—rdo gF @PHHIA E e™tinopterygi—ns from the widdle

@idFA E etti Rë ƒimposio di i™ologi— e €—leoe™ologi— „ri—ssi™ of xorthern st—ly —nd g—nton „i™ino @ƒwitE

dell— gomunit—Á fentoni™heD wuseo ‚egion—le di zerl—ndAX —n—tomi™—l des™riptions —nd nomen™l—tur—l

ƒ™ienze x—tur—li @IWVVAX WSEIHSD „orinoF pro˜lemsF ‚ivF stF €—leontolF ƒtr—tFD IHU@QAX QRSEQTWF

„intori eF 8 fr—m˜ill— iF @IWWIA E ƒexu—l dimorphism in — vom˜—rdo gFD „intori eF 8 „on— hF @PHIPA E e new

gon™hostr—™—n popul—tion from the v—te v—dini—n of spe™ies of ƒ—ngiorgioi™hthys @e™tinopterygiiD ƒemioE

ƒouthern g—l™—reous elps @xF st—lyAF snX uiel—nEt—rE notiformesA from the u—lks™hieferzone of wonte

owosk— FD reintz xF 8 x—krem rFeF @idsA E pifth ƒ—n qiorgio @widdle „ri—ssi™Y werideD g—nton „i™iE

ƒymposium on wesozoi™ „errestri—l i™osystems —nd noD ƒwitzerl—ndAF follF ƒo™F €—leontolF stF SI@QAX PHQE

fiot—F gontri˜utions €—leontologi™—l wuseumD niE PIPF

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„intori eF 8 vom˜—rdo gF @IWWWA E v—te v—dini—n fish f—uE uFeFD g—nnone tFtF 8 qutel ‚F‚F @PHHWA E wonophyly

n—s from vom˜—rdy @xEst—lyAX str—tigr—phy —nd p—E of terrestri—l —deph—g—n ˜eetles —s indi™—ted ˜y three

IWH ƒtr—d— vFD wont—gn— wF 8 „intori eF

dle „ri—ssi™ ˆingyi p—un— E eviden™e from the ex™—E leo˜iologyF snX err—ti— qF 8 ƒ™hultze rFE€F @idsA E

€ro™eedings of the ƒymposium ––wesozoi™ fishesX sysE v—tion —t xim—igu of ‡ush— histri™tD ˆingyi gityD

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tem—ti™s —nd fossil re™ord99X RWSESHRF quizhou €rovin™eF e˜str—™t ˜ook of the P snterE

„intori eF 8 ‚enesto ƒF @IWWHA E e new v—rios—urus from the

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u—lks™hieferzone @ ppermost v—dini—nA of †—l™eresio

†erte˜r—te p—un—sD ƒeptem˜er IHEISD PHIQD ˆingyiD

@†—reseD xF st—lyAF follF ƒo™F €—leontolF stFD PW@QAX QHWE

ghin—X TQETSF

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‡—ng fFD h—ng rF 8 €onom—renko eFqF @PHIPA E wesozoi™

„intori eFD ritij „FD ti—ng hFD vom˜—rdo gF 8 ƒun F @PHIQ—A

„r—™hyp—™hid—e @snse™t—X goleopter—A from ghin—F

E „ri—ssi™ —™tinopterygi—n fishesX the re™overy —fter the

€—l—eontologyD SSD PX QRIEQSQF

endE€ermi—n ™risisF sntegrF oolF ynEline IHGPHIQY

h—ng rF @IWWUA E i—rly gret—™eous inse™ts from the h—l—zi hysX IHFIIIIGIURWERVUUFIPHUUF

porm—tion of the hixin f—sinD tilin €rovin™eD ghin—F „intori eFD ti—ng hF‰FD wot—ni ‚FD ƒun F‰FD ti gFD ou

ˆFh 8 w— vF„F @PHIQ˜A E p—un—l sequen™e of widE €—leoworldD UX USEIHQF