1485 Haug.vp

12 Pages·2014·3.5 MB·English

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ArthropodcuticlesfromtheMississippian(EarlyCarboniferous,Viséan)HainichenSubgroup,ErzgebirgeBasin,have
beenencounteredinbulk-maceratedsamplesfromaroadcutexposureinChemnitz-Glösa,easternGermany.Thecuti-
clesaredescribedindetailandcomparedwithlargerfaunalcomponentsknownfromthesamehorizons.Partofthespec-
imensrepresentremainsofarthropodappendages,whileothers,thoughexhibitingfinedetailofsurfacetexture,could
notbeassignedtocertainbodyregions.InallcasesitremainsdifficulttoassignthefragmentsfromChemnitz-Glösato
taxaknownfromtheHainichenSubgrouporfromotherlocalitiesyieldingarthropodremainsofsimilarpreservation.
However,itissuggestedthatseveralspecimensmayrepresentscorpionremains,eitherlimbparts(pieceofthepedipalp,
distalclawofwalkinglimb)orfragmentsofthebodysurface.Onespecimen,asingleappendageelement,couldrepre-
sent the first insect fragment from this locality. All fragments appear to be terrestrial faunal components.
Chemnitz-GlösaisoneoftheveryfewLowerCarboniferouslocalitiestoyieldremainsofterrestrialarthropodsandthe
only one outside Scotland. (cid:129) Key words: Mississippian, Lower Carboniferous, arthropod cuticles, Erzgebirge Basin.
HAUG,J.T.,HÜBERS,M.,HAUG,C.,MAAS,A.,WALOSZEK,D.,SCHNEIDER,J.W.&KERP,H.2014.Arthropodcuti-
clesfromtheupperViséan(Mississippian)ofeasternGermany.BulletinofGeosciences89(3),541–552(7figures).
CzechGeologicalSurvey,Prague.ISSN1214-1119.ManuscriptreceivedSeptember20,2013;acceptedinrevisedform
January 13, 2014; published online May 6, 2014; issued June 9, 2014.
JoachimT.Haug(correspondingauthor)&CarolinHaug,LMUMunich,Biocenter–DepartmentofBiologyIIand
GeoBio-Center,GroßhadernerStraße2,82152Planegg-Martinsried,Germany;[email protected]
(cid:129) Maren Hübers & Hans Kerp, Forschungsstelle für Paläobotanik, Institut für Geologie und Paläontologie,
WestfälischeWilhelms-UniversitätMünster,Heisenbergstraße2,48149Münster,Germany(cid:129)AndreasMaas&Dieter
Waloszek,AGBiosystematischeDokumentation,UniversitätUlm,Helmholtzstraße20,89081Ulm,Germany(cid:129)JörgW.
Schneider,TUBergakademieFreiberg,GeologischesInstitut,Bernhard-von-Cotta-Straße2,09596Freiberg,Germany
and Kazan Federal University, Kremlevskaya st. 18, 420008 Kazan, Russian Federation
ThemiddleMississippianHainichenSubgroupcomprises nearChemnitz.Thefaunalrecord,includingadpressionsof
the oldest terrestrial erosional debris (“Early Molasses”) arachnids, myriapods, arthropleurids, eurypterids, shark
oftheVariscanOrogenineasternGermany.Thesemostly eggcapsulesandsingleisolatedfishremains(Nindel1955,
coarseclasticshavebeendepositedintheHainichenSub- Rößler&Schneider1997)hasexclusivelybeendescribed
group of the Hainichen Basin, which is part of the fromtheBerthelsdorfFormationuntil2001(Schneideret
Carboniferous-PermianErzgebirgeBasin(Fig.1;Schnei- al.2005).DuringthereconstructionoftheA4federalmo-
der et al. 2005). The Hainichen Subgroup is divided into torwaynearChemnitz-Glösafrom2001to2003extensive
twoformations(Fig.2).Thelowerone,theOrtelsdorfFor- collectionoffossilplantsandanimalswaspossibleinthe
mation,isdatedasViséan3b(Asbian),theupperone,the Ortelsdorf Formation. The preservation of fossil arthro-
BerthelsdorfFormation,isdatedasViséan3c(Brigantian) podsingeneralis,however,notrestrictedtoadpressions.
(Gaitzschetal.2010).Bothformationshaveyieldedarich SeveralarthropodcuticlesweredescribedfromthePalaeo-
anddiverseflora,whichhasbeendescribed,by,e.g.,Roth- zoic,mainlyfromtheDevonian(e.g.,Størmer1970,1976;
pletz (1880), Sterzel (1884), Máyas (1920) and Hartung Rolfe 1980; Shear et al. 1984) and Pennsylvanian (e.g.,
(1938).TheplantmaterialmainlycomesfromtheOrtels- Winslow 1959, Bartram et al. 1987, Jeram 1994a, Braun
dorf Formation and was sampled in dumps of coal mines 1997,Stankiewiczetal.1998).Cuticlescanprovideadditi-
betweenHainichenandChemnitz.Thesecoalmineswere onalinformationonmorphologicalfeatures,whichcannot
active during the 18th century. Additional material was be gained in such detail from adpression remains. Also
sampledinsandandclaypitsoftheBerthelsdorfformation bulkmacerationofsamplesfromtheOrtelsdorfFormation
(cid:22)(cid:3)(cid:7)(cid:11)(cid:31) (cid:10)!(cid:31)(cid:30) "#$%%(cid:10)&’()*+(cid:10)(cid:31)(cid:30),(cid:29) (cid:29)(cid:30)(cid:31)
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(cid:29)(cid:12)(cid:30)(cid:11)(cid:3)(cid:14)(cid:9)(cid:31) PositionoftheMississippianHainichenBasin(A)andthePennsylvanian-PermianErzgebirgebasins(B:FlöhaBasin,C–D:Zwickau-Oelsnitz
Basin)ineasternGermany(afterSchneider&Romer2010,theirfig.8).
yieldedseveralarthropodcuticles,whicharedescribedinthis islateViséan(earlyBrigantian)inage(VFZone)(Fig.2).
paper.Wecomparethesecuticleswithotherfossilandextant Although the single-Zircon Pb/Pb evaporation age of
arthropodmaterialandtrytoalignthemwithmacroscopicre- 330 ± 4 Ma of a rhyolithic tuff at the base of the coal-
mainsalreadydescribedfromtheHainichenSubgroup. bearing fossiliferous upper part of the formation (Gehm-
lichetal.1998,Gaitzschetal.2010)isnotveryprecise,it
isconsistentwiththepalynologicaldating.Coalificationin
(cid:27)(cid:6)(cid:10)(cid:20)(cid:13)(cid:12)(cid:4)(cid:26)(cid:28)(cid:9)(cid:17)(cid:20)(cid:4)(cid:14)(cid:3)(cid:12)(cid:20)(cid:13)(cid:9)(cid:20)(cid:21)(cid:8)(cid:9)(cid:17)(cid:14)(cid:4)(cid:5)(cid:6)(cid:8)(cid:15)
the Hainichen Basin is low (sub-bituminous coals;
R ~0.5).Inthefirsthalfofthe20thcenturyoutcropsyiel-
m
Elevensamplesfromdifferentplantdetritus-richhorizons dedoneoftheveryfewViséanmacrofloraswithArchaeo-
andcarbonaceousshalesofdecimeterthicknesswerecol- calamitesradiatus,LepidodendronlosseniiandLyginopte-
lected in the upper Ortelsdorf Formation in Chemnitz- ris bermudensiformis as common elements (Hartung
Glösa(50°52´34.67˝N;12°54´38.73˝E).Thespecimens 1938). The shales exposed near Glösa contain abundant
describedherewerecollectedfromaroadcutoftheA4fe- organicdebris,mainlyplantsincludinglycopsids(Hübers
deral motorway near Chemnitz-Glösa, Free State of Sa- etal.2011)andtheoldestmosses(Hübers&Kerp2012,
xony,Germany,wherea80mthicksuccessionofMissis- Hübersetal.2013).Someofthesamplescontainmacro-
sippian sandstones, shales and occasional coaly layers of scopicplantremainsoftheseedfernLyginopterisbermu-
theupperpartoftheOrtelsdorfFormation,HainichenSub- densiformis and the lycophyte Lepidodendron lossenii.
group,HainichenBasin,isexposed(Schneideretal.2005). Macroscopicfaunalremainscouldnotbeobservedinthe
The Hainichen Basin is a small basin, filled with alluvial samples, but have been discovered during outcrop docu-
plain,alluvialfanandfandeltadeposits.TheupperOrtels- mentation 2001–2002.
dorf Formation at Glösa has been dated palynologically Bulk maceration with 48% hydrofluoric acid yielded
(Jäger & Wierich 2006) and comprises the NM and VF many small plant fragments. Small pieces of sample
zonesofthepalynologicalstandardzonationofClaytonet 2324/2004 were carefully removed from the surface and
al.(1977).Thesamplesarefromtheuppermostpartwhich dissolvedin48%hydrofluoricacid;allothersampleswere
(cid:29)(cid:30)-
(cid:26)(cid:10)(cid:27)(cid:14)(cid:28)(cid:7)(cid:29)(cid:30)(cid:17)(cid:9)(cid:31)(cid:27)(cid:3) (cid:5)(cid:6)(cid:9)(cid:27)(cid:4)(cid:17)(cid:9)(cid:15)(cid:9)!"(cid:6)(cid:28)"(cid:10)#(cid:10)$(cid:9)(cid:14)(cid:3)(cid:6)(cid:7)(cid:14)(cid:4)(cid:5)(cid:13)(cid:9)(cid:11)"(cid:10)(cid:29)(cid:9)(cid:6)(cid:28)(cid:5)(cid:9)(cid:3)##(cid:5)"(cid:9)(cid:16)(cid:7)(cid:13)%(cid:27)(cid:8)(cid:9)(cid:10)(cid:11)(cid:9)(cid:5)(cid:27)(cid:13)(cid:6)(cid:5)"(cid:8)(cid:9)(cid:12)(cid:5)"(cid:29)(cid:27)(cid:8)&
dissolvedcompletely.Cuticlepreparationwascarriedoutas
describedbyKerp(1990)andKerp&Krings(1999).Iso-
lated cuticles were macerated using Schulze’s reagent
(HNO + KClO ), treated with a 4% potassium hydroxide
3 3
solution (KOH ), dehydrated in glycerol and finally
aq
mounted on microscope slides using glycerine jelly. The
slidesarestoredinthecollectionoftheForschungsstellefür
PaläobotanikattheInstitutfürGeologieundPaläontologie,
WestfälischeWilhelms-UniversitätMünster,Germany.
Forcomparison,differentfossilandextantarthropods
were studied. This includes isolated legs of an undeter-
mined pseudoscorpion, Brachistosternus sp. (Scorpiones,
Chelicerata), Scutigera coleoptrata (Chilopoda, Myria-
(cid:29)(cid:12)(cid:30)(cid:11)(cid:3)(cid:14)(cid:9)" Age, subdivision, facies pattern and fossil content of the
poda), and Oniscus asellus (Isopoda, Crustacea), which
Hainichen Subgroup (after Gaitzschet al. 2010, their fig. 3).
weremaceratedin10%KOHfortwoweeks.(Thecuticle
ofazygentomaninsecttreatedinthesamewayprovedto
betoomuchdamagedforfurtherprocessing.)The(remain- (propertyoftheAxelrodInstitute,UniversityofGuelph,on
ing) cuticles were rinsed in water, placed between two longtermloantotheROM)weredocumentedasthesepro-
glass slides and left there for drying. This procedure was videfinedetailsofdifferentappendages.Specimensfrom
usedtoobtaincomparabledatafromextantspecimens,asit Crato were documented in the same way as those from
is supposed to partly resemble the taphonomic and Mazon Creek. The specimen from the Rhynie Chert was
preparational history of Glösa arthropod remains. Subse- documentedwithtransmittedlightonaZeissAxioskop2
quently the dried specimens were documented using a equippedwithagrey-scaleAxioCam.Animagestackwas
Zeiss Axioskop microscope equipped with a SkopeTek recordedandprojectedasaminimumintensityprojection
DCM510ocularcameraundertransmittedpolarizedlight. using the software ImageJ.
Images were recorded in the form of stacks, several for
eachspecimen.ThesestackswerefusedwithCombineZM
andCombineZP.Thefusedimageswerethenstitchedwith !(cid:14)(cid:15)(cid:10)(cid:3)(cid:12)(cid:7)(cid:4)(cid:12)(cid:6)(cid:21)(cid:9)(cid:6)(cid:16)(cid:9)(cid:4)(cid:5)(cid:14)(cid:9)(cid:20)(cid:3)(cid:4)(cid:5)(cid:3)(cid:6)(cid:7)(cid:6)(cid:8)(cid:9)(cid:10)(cid:11)(cid:4)(cid:12)(cid:10)(cid:13)(cid:14)(cid:15)
AdobePhotoshopCS3orMicrosoftImageCompositeEdi-
tor (details on procedure in Hauget al.2009). (cid:2)(cid:3)(cid:4)(cid:5)(cid:3)(cid:6)(cid:7)(cid:6)(cid:8)(cid:9)(cid:10)(cid:7)(cid:7)(cid:11)(cid:12)(cid:8)(cid:10)(cid:13)(cid:11)(cid:9)(cid:11)(cid:14)(cid:11)(cid:15)(cid:11)(cid:12)(cid:4)(cid:16)
Foranoverviewoftaxapossiblyrepresentedbythecu-
ticle remains from Glösa, different arthropods from the ThesamplesfromGlösacontainfourcuticularremainsof
Upper Carboniferous Mazon Creek Lagerstätte, Illinois, different arthropod appendage elements:
USA(ROM43581trigonotarbid;ROM45545Geralinura (cid:129) Specimen“Glösa-2324/2004-018”(Fig.3A):Appen-
carbonaria,thelyphonidwhipscorpion;ROM45559scor- dage element, ca 1100 µm long, width increasing
pion; ROM 45571 Latzelia primordialis, scutigeromorph fromproximaltodistalend,upto400µm.Surfaceex-
centipede; ROM 47504 Hesslerella shermani, aquatic hibiting pore-like structures, ca 10 µm in diameter,
isopod;ROM61543pterygoteinsect;ROM61608diplo- whichmayrepresentinsertionsofsetae.Threedistal
pod;ROM61541Dasyleptussp.,archaeognath)andEarly movableelements,446µm,390µmand227µmlong.
Permian Carrizo Arroyo deposits, New Mexico, USA Triangular movable setae (60–70 µm long) arranged
(ROM 57075 Dasyleptus sp.) from the collection of intworows.
the Royal Ontario Museum (ROM) Toronto were docu- (cid:129) Specimen“Glösa-ProfilKS11-099”(Fig.3B):Presum-
mentedunderpolarizedlightwithaCanonRebelT3icam- ably terminal sickle-shaped claw, about 820 µm long.
era and an MP-E 65 mm macro lens. Also several stacks Two smaller spines arise medio-proximally from the
were obtained from each specimen and processed as de- claw.
scribed above. Additionally, an isolated appendage frag- (cid:129) Specimen “Glösa-Profil KS27-101” (Fig. 3C): Appen-
ment of a trigonotarbid specimen from the Devonian dageelement,ca690µmlongand400µmwide.Surface
Rhynie Chert, Scotland (RRC 3702, Remy Rhynie Chert exhibitspore-likestructures,probablyhairbases;some
Collection at the Forschungsstelle für Paläobotanik, hairs still attached.
Münster) and leg details of different insect specimens (cid:129) Specimen “Glösa-Profil KSgr.Pr.-056” (Fig. 3D): Pre-
[ROM63044(AI190)grylloid;ROM63045(AI459)blat- sumablyterminalappendageelement,ca1.65mmlong,
toid; ROM 63046 (AI 751) tettigoniid; ROM 63047 upto170µmwide,arcuate.Smallserrationsonconcave
(AI2649)orthopteran;ROM63048(AI3211)notonectid lateralsideofthelimb.Serrationsupto25µmlong.Sur-
heteropteran]fromtheCretaceousCratoFormation,Brazil face rough. No special structures present.
(cid:29)(cid:30)!
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% &
!
(cid:2)
(cid:29)(cid:12)(cid:30)(cid:11)(cid:3)(cid:14)(cid:9)# Arthropodappendageelements.(cid:129)A–Glösa-2324/2004-018.Appendageelementwithdistalmovableelementsandtriangularmovable
setae,scalebar=100µm.(cid:129)B–Glösa-ProfilKS11-099.Clawwithdenticles,scalebar=100µm.(cid:129)C–Glösa-ProfilKS27-101.Arthropodappend-
ageelementshowinghairs(indicatedbyarrows),scalebar=100µm.(cid:129)D–Glösa-ProfilKSgr.Pr.-056.Sickle-shapedelementwithserrations,scale
bar=100µm.
(cid:17)(cid:4)(cid:5)(cid:11)(cid:3)(cid:9)(cid:10)(cid:3)(cid:4)(cid:5)(cid:3)(cid:6)(cid:7)(cid:6)(cid:8)(cid:9)(cid:18)(cid:19)(cid:4)(cid:20)(cid:18)(cid:14)(cid:11)(cid:9)(cid:21)(cid:3)(cid:10)(cid:13)(cid:15)(cid:11)(cid:12)(cid:4)(cid:16)
andmoreorlessregularlyarrangedpores.Poresgenerally
15–20 µm in diameter, sometimes containing hair bases
ThesamplesfromGlösaalsocontainseveralfragmentsof (Fig.4B),largestpore161µmindiameterwithremnantsof
arthropodcuticle,whichcannotbeassignedtocertainbody thin cuticle at the margin (Fig. 4D). Some specimens ex-
parts.However,theirwell-preservedsurfacesexhibitmany hibitingsmallpapillae(Fig.4C).Papillaeca3µmindiam-
structural details. eter.
–ArthropodCuticleType1:Glösa-ProfilKS62/2-037 –ArthropodCuticleType2:Glösa-ProfilKS62/2-001,
to -041, -043 to -053, -056, -057, -061; Glösa-Profil -003,-004,-030,-031;Glösa-2324/2004-095(Fig.4E-G):
KSgr.Pr.-014, -043, -060; Glösa-Profil KS 11-016, -038, Thickcuticlewithstrongsclerotizationandroughtubercu-
-092; Glösa-Profil KS27-105; Glösa-Profil KS34/35-024 lar surface, occasionally with pore-like structures
(Fig.4A–D):Cuticlefragmentswithisodiametricstructure (Fig. 4F).
(cid:29)(cid:12)(cid:30)(cid:11)(cid:3)(cid:14)(cid:9)$ ArthropodCuticleType1andArthropodCuticleType2.(cid:129)A–Glösa-ProfilKS62/2-040.ArthropodCuticleType1,overviewshowing
isodiametricstructureandpores,scalebar=100µm.(cid:129)B–detailofA,showingporewithhairbase,scalebar=10µm.(cid:129)C–Glösa-ProfilKS62/2-041.Ar-
thropodCuticleType1showingsmallpapillae(indicatedbyarrows),scalebar=10µm.(cid:129)D–Glösa-ProfilKS62/2-061.ArthropodCuticleType1show-
ingexceptionallylargeporewithremnantsofthincuticleatthemargin,scalebar=100µm.(cid:129)E–Glösa-ProfilKS62/2-001.ArthropodCuticleType2,
overviewshowingroughtubercularsurface,scalebar=100µm.(cid:129)F–detailofD,showingpore-likestructures,scalebar=100µm.(cid:129)G–Glösa-Profil
KS62/2-003. Arthropod Cuticle Type 2, specimen showing differences in strength of sclerotization, scale bar = 100 µm.
(cid:29)(cid:30)(cid:30)
(cid:26)(cid:10)(cid:27)(cid:14)(cid:28)(cid:7)(cid:29)(cid:30)(cid:17)(cid:9)(cid:31)(cid:27)(cid:3) (cid:5)(cid:6)(cid:9)(cid:27)(cid:4)(cid:17)(cid:9)(cid:15)(cid:9)!"(cid:6)(cid:28)"(cid:10)#(cid:10)$(cid:9)(cid:14)(cid:3)(cid:6)(cid:7)(cid:14)(cid:4)(cid:5)(cid:13)(cid:9)(cid:11)"(cid:10)(cid:29)(cid:9)(cid:6)(cid:28)(cid:5)(cid:9)(cid:3)##(cid:5)"(cid:9)(cid:16)(cid:7)(cid:13)%(cid:27)(cid:8)(cid:9)(cid:10)(cid:11)(cid:9)(cid:5)(cid:27)(cid:13)(cid:6)(cid:5)"(cid:8)(cid:9)(cid:12)(cid:5)"(cid:29)(cid:27)(cid:8)&
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!(cid:12)(cid:15)(cid:10)(cid:11)(cid:15)(cid:15)(cid:12)(cid:6)(cid:21)(cid:9)(cid:6)(cid:16)(cid:9)(cid:4)(cid:5)(cid:14)(cid:9)(cid:20)(cid:3)(cid:4)(cid:5)(cid:3)(cid:6)(cid:7)(cid:6)(cid:8)(cid:9)(cid:10)(cid:11)(cid:4)(cid:12)(cid:10)(cid:13)(cid:14)
rates(Fig.6B)andchilopods(Figs5Finlet,6C).Yet,simi-
(cid:16)(cid:3)(cid:20)(cid:30)(cid:17)(cid:14)(cid:21)(cid:4)(cid:15)
larstrongedgescanalsobefoundamongpterygoteinsects,
for example orthopterans (Fig. 7A). Pterygote insects are
While cuticle fragments can provide quite different data alsothebestexamplespossessingprominentandmovable
compared to adpression fossils of arthropods, identifying setae, for example blattoids (Fig. 7B) and grasshoppers
the taxonomic affinities of the cuticle fragments can be among the orthopterans (Fig. 7C). In blattoids, however,
challenging.Inthiscasetheinvestigationofthefewcuticle thestrongedgesarelessdefinitethanthatseenonthecuti-
fragmentsisanimportanttask,duetothestratigraphicori- cle fragment. Additionally the spines are not arranged
ginofthespecimens.TheCarboniferousisacrucialperiod in strictly proximo-distally oriented rows in blattoids
forthediversificationofterrestrialarthropodsinparticular, (Fig.7B).Thisisthecaseingrasshoppers(Fig.7C)andin
including chelicerates, insects and myriapods (e.g., Rolfe general in orthopterans (Fig. 7A, D). Orthopterans also
1980,Rasnitsyn&Quicke2002,Grimaldi&Engel2005). have well defined edges, along which the spines arise
Cheliceratesarerepresentedbyvariousalreadyspecialized (Fig.7A).Inconclusion,theelementcouldrepresentanap-
groups such as true scorpions (Fig. 5A) and the once di- pendage element of a pterygote insect related to
verse,butnowextinctTrigonotarbida(Fig.5B),butalsoby orthopterans, possibly the tibia of the metathoracic ap-
lesswell-knowntaxasuchasThelyphonida,thewhipscor- pendage. These are usually quite elongate, the specimen
pions(Fig.5C).WhiletheHainichenSubgroupalsoyiel- couldinthiscaserepresentonlythedistalpartofthisele-
dedremainsofaquaticarthropods,i.e.Eurypterida,thecu- ment,whiletheproximalpartwouldbemissing.Theproxi-
ticle fragments described here do not appear to represent malrimofthespecimencouldwellbeinterpretedasrepre-
remains of eurypterids. Insects are well represented by senting an artificial rim caused by fragmentation. This
non-flyingforms,suchasArchaeognatha(Fig.5E),butes- specimen, if indeed representing an orthopteran, would
pecially by pterygote insects (Fig. 5D), which are an im- represent the first insect fossil from this locality.
portantpartoftheCarboniferousfaunas.Myriapodsarere- Specimen“Glösa-ProfilKS11-099”(Fig.3B)isaclaw
presented by in-groups of Chilopoda, such as with denticles. Similar tarsal or pre-tarsal claws occur in
Scutigeromorpha(Fig.5F),butalsoDiplopodaarewellre- variousterrestrialarthropods,suchasdifferentchelicerate
presented (Fig. 5G). Another terrestrial group of arthro- taxa (Fig. 6A, B), myriapods (Fig. 6C), and also insects
pods,thewoodlice(Isopoda),anin-groupofEucrustacea, (Fig. 7B, D). Even the dactylus of a wood louse appears
have not been found in the Carboniferous yet, although veryclawlike(Fig.6D).Yet,mostoftheseclawsappearto
aquatic isopods make their appearance in the Carbonife- besimpleclawswithoutfurtherarmature.Theonlyexam-
rous (Fig. 5H). plebearingadditionalsmallerspine-likestructureslikethe
Accordingly, the Glösa cuticles could be remains of fossilcanbeseenonthetarsalclawoftheheredocumented
any of the representatives of the terrestrial groups, al- scorpion (Brachistosternus sp.; Fig. 6B). This makes it
though until now mainly chelicerates and myriapods are likelythatthefossilrepresentsthetarsalclawofascorpion,
known from the same locality. Possible affinities are dis- yet other affinities cannot be excluded completely.
cussed for each specimen in the following. Specimen“Glösa-ProfilKS27-101”(Fig.3C)isacuti-
Specimen “Glösa-2324/2004-018” (Fig. 3A) could be cle fragment that provides the fewest prominent details.
interpretedasrepresentingthesub-terminalelementofan Stillquitemanydetailsarepreservedsuchaspores,which
appendagewithaputativeterminalclaw.Yet,itdoesnot originallyaccommodatedhairstructures.Theopeningson
bearasingleterminalelement,buttherearethreemovable the surface are reminiscent of those developed in
ones. This morphology would be unusual for a terminal trigonotarbidcuticles(Fig.5Binlet),andsimilarlylooking
structure.Additionally,thedistalopeningofthelargeele- slitsensillaarecommononthecuticleofarachnidsingen-
mentissignificantlylargerthanthethreesmallelements, eral.However,hairysurfaces–andthereforesurfaceswith
and much more likely accommodated a further terminal holesafterthesetaearelost–areinfactcommonamong
limbelement.Itisalsodifficulttojudgeifthethreesmall terrestrialarthropods(e.g.,Fig.5Einlet)andmakeanas-
elements lie “inside” the large element, i.e. between two signment even more challenging than for the other speci-
cuticlelayers,whichwouldpointtoatrueterminalclaw,or mens.
whethertheylieoutside,representingequivalentsofstrong Specimen “Glösa-Profil KSgr.Pr.-056” (Fig. 3D) is a
movable setal elements. possiblydistalsickle-shapedelementwithserrationsalong
Besidesthisuncertaintyaboutthetruenatureofthese itsinneredge.Suchanelementcouldbeadactylus,e.g.,of
supposed terminal structures, the specimen possesses an(terrestrial)isopod(Fig.6D),butitappearsratherlong
strongproximo-distallyrunningedgesequippedwithlarge for such a structure. Elongate tarsal elements of insects
triangular setae, which appear to be movable (Fig. 3A). (Fig. 7D, E) have also structural similarities to the fossil
Strongedgesequippedwithprominentspinesorsetaeare, specimen.Suchstructurescontinueusuallyintoapre-tar-
to some degree, also found on representatives of chelice- sus(Fig.7E,F),butnosuchstructurecanbediscernedon
(cid:29)(cid:30).
(cid:26)(cid:10)(cid:27)(cid:14)(cid:28)(cid:7)(cid:29)(cid:30)(cid:17)(cid:9)(cid:31)(cid:27)(cid:3) (cid:5)(cid:6)(cid:9)(cid:27)(cid:4)(cid:17)(cid:9)(cid:15)(cid:9)!"(cid:6)(cid:28)"(cid:10)#(cid:10)$(cid:9)(cid:14)(cid:3)(cid:6)(cid:7)(cid:14)(cid:4)(cid:5)(cid:13)(cid:9)(cid:11)"(cid:10)(cid:29)(cid:9)(cid:6)(cid:28)(cid:5)(cid:9)(cid:3)##(cid:5)"(cid:9)(cid:16)(cid:7)(cid:13)%(cid:27)(cid:8)(cid:9)(cid:10)(cid:11)(cid:9)(cid:5)(cid:27)(cid:13)(cid:6)(cid:5)"(cid:8)(cid:9)(cid:12)(cid:5)"(cid:29)(cid:27)(cid:8)&
&
%
(cid:2)
!
’ (cid:29)
)
(cid:25)
(cid:29)(cid:12)(cid:30)(cid:11)(cid:3)(cid:14)(cid:9)( Differentarthropodsforcomparison.MainlyterrestrialformsfromtheCarboniferous,MazonCreek,USA.(cid:129)A–ROM45559.Scorpion,
scalebar=2mm.(cid:129)B–ROM43581.Trigonotarbid,scalebar=3mm;Binlet.–RRC3702.Cuticleremainsofanappendageofatrigonotarbidfromthe
DevonianRhynieChert,Scotland.(cid:129)C–ROM45545.Geralinuracarbonaria,thelyphonid(whipscorpion),scalebar=3mm.(cid:129)D–ROM61543.
Pterygoteinsect,scalebar=3mm.(cid:129)E–ROM61541.Dasyleptussp.,archaeognath(winglessinsect),scalebar=3mm;Einlet.–ROM57075.Closeup
ofalegofaDasyleptussp.specimenfromtheEarlyPermianofCarrizoArroyo,NewMexico,USA.(cid:129)F–ROM45571.Scutigeromorphcentipede
Latzeliaprimordialis,scalebar=3mm;Finlet–closeupofalegofthecounterpartofF.(cid:129)G–ROM61608.Diplopod,scalebar=4mm.(cid:129)H–ROM
47504.Hesslerella shermani, aquatic isopod, scale bar = 2 mm.
(cid:29)(cid:30)/
(cid:2)(cid:3)(cid:4)(cid:4)(cid:5)(cid:6)(cid:7)(cid:8)(cid:9)(cid:10)(cid:11)(cid:9)(cid:12)(cid:5)(cid:10)(cid:13)(cid:14)(cid:7)(cid:5)(cid:8)(cid:14)(cid:5)(cid:13)(cid:9)(cid:15)(cid:9)(cid:16)(cid:10)(cid:4)(cid:17)(cid:9)(cid:18)(cid:19)(cid:20)(cid:9)(cid:21)(cid:20)(cid:9)(cid:22)(cid:23)(cid:24)(cid:25)
thefossil(Fig.3D).Thefossilcould,duetoitslength,either Interestingly,mostarachnidsfromtheBerthelsdorfForma-
representthedactylusofachelipedofacrustacean,orthe tionhavebeenfoundinasandstonebedtogetherwithcom-
“digitusmobilis”ofacheliceratancheliped.Asothercuti- monremainsoftheotherwiseraremesophilousplantfossil
clefragmentsmightrepresentscorpionremains,itishighly Sphenopteris nindeliana (Rößler & Schneider 1997). It
likely that this specimen represents the digitus mobilis of couldbeassumed,thatthosearachnidshavepreferreddrier
the pedipalp (claw-bearing second appendage) of a scor- areas in this river landscape.
pion. ThefossilrecordofEarlyCarboniferousterrestrialar-
ArthropodCuticleType1(Fig.4A–D)bearsinitssur- thropods is very poor with only few localities, because
face structure similarities to the scorpion cuticle depicted Lower Carboniferous strata are mostly marine (Shear &
in Stankiewicz et al. (1998, fig. 2g) and also to that of Kukalová-Peck1990).Alsoplantlocalitiesareratherfew,
Attercopus fimbriunguis (see Selden et al. 2008, their and their total number in central and northwest Europe
fig. 1), an arachnid probably in the evolutionary lineage barelyexceedsafewdozens;mostareinthelateViséan.
leadingtowardsmodernaraneans.AlsoArthropodCuticle Recently,twoTournaisian(Courceyan)localitieswithar-
Type2(Fig.4E–G)canbecomparedwiththatofascor- thropods have been reported from southern Scotland
pion, as depicted in Jeram (1994a, his pl. 7, figs 7, 8). (TweedBasin,ScottishBorders).Bothfossilassociations
include myriapods and scorpions (Smithson et al. 2012),
which,however,havenotbeendescribedindetailyet.The
(cid:25)(cid:14)(cid:21)(cid:14)(cid:3)(cid:20)(cid:13)(cid:9)(cid:8)(cid:12)(cid:15)(cid:10)(cid:11)(cid:15)(cid:15)(cid:12)(cid:6)(cid:21)(cid:9)(cid:20)(cid:21)(cid:8)(cid:9)(cid:10)(cid:6)(cid:21)(cid:10)(cid:13)(cid:11)(cid:8)(cid:12)(cid:21)(cid:30)(cid:9)(cid:3)(cid:14)(cid:17)(cid:20)(cid:3)*(cid:15)
other locality is East Kirkton also in southern Scotland
(West Lothian) (Clarkson et al. 1994). The East Kirkton
Theentirefining-upwardsequenceoftheupto800mthick depositsareBrigantianinageliketheGlösabedswhichare
Ortelsdorf Formation is dominated by compositionally described here.
immature terrigenous clastics, with subordinated thin Someinterestingcomparisonscanindeedbemadewith
coalseams.Thelowerpartoftheformationformsthever- the East Kirkton lake deposits, even though this locality
tical and lateral transition from a subaqueous near-shore hasyieldedmuchmoreandmuchlargerspecimensthanthe
fandeltaenvironmentintoterrestrialdistalalluvialfande- heredescribedChemnitz-Glösalocality.Thelakewassur-
posits.Theupperpart,yieldingtheheredescribedarthro- roundedbyagymnosperm-pteridospermforest(Galtier&
pod cuticles, consists of floodplain and channel deposits Scott1994,Scottetal.1994),inhabitedbyavarietyofter-
anddepositsoflocalswampsformingdecimetre-thickcoal restrial invertebrates and vertebrates (Clarkson et al.
seamsandcarbonaceousshale.Theveryshallowbutwide 1994).Findsofterrestrialarthropodsaredominatedbydif-
channelgeometryandbed-loadtransportpointtobraided ferentontogeneticstages(13mmto700mm)ofthegiant
to anastomosing river systems. Vertically buried centi- Pulmonoscorpius kirktonensis, the top predator on land
metre thick stems of lycopsids and Archaeocalamites, as (Jeram1994b).Severalmyriapods(Shear1994)areknown
well as finely rooted horizons, are not rare. Layers with as well as one harvestman (Dunlop & Anderson 2005).
current-oriented,inplaceschaotictrunksandtwigsofAr- Eurypteridsarerepresentedbytheupto1.5mlongsuppos-
chaeocalamites, lycopsids and other plants may indicate edly amphibious-terrestrial Hibbertopterus scouleri
flood events. Temporary flood plain ponds and lakes, re- (Jeram & Selden, 1994) (Whyte 2005). The fish fauna of
mainingafterflash-floodevents,mayhavebeenspawning the lake consisted of different actinopterygians, acantho-
placesoffemalexeanacanthidandhybodontsharksasindi- diansandarhizodontidsarcopterygianaswellashybodont
catedbymass-occurrencesoftheireggcapsules.Thecom- andxenacanthsharks(Coates1994,Paton1994)–thelat-
monnessofterrestrialadaptedarthropods,especiallyscor- ter fishes are the producers of the shark egg capsules
pions,isindicatedbythecuticles;scorpioncuticlesappear known from the Hainichen Subgroup (Schneider et al.
to be generally better preservable than those of other 2010).
non-mineralisedarthropods,probablyduetometalsenclo- The Brigantian (late Viséan) marks the start of an in-
sedinthecuticle(Jeram2001).Thisabundanceofremains creased radiation of terrestrial arthropods (Ward et al.
ofterrestrialarthropodsfitswellinthepictureofanopen 2006)andtetrapods(Smithsonetal.2012)afteratimein-
riverlandscapewithinstablevegetationofmainlyhygro- tervalreachingfromthelatestDevoniantotheHolkerian
philous to mesophilous plants. Similarly, the adpression (~ middle Viséan) that is characterized by a general ab-
fossilsofarthropodsfromtheBerthelsdorfFormationare senceofterrestrialarthropodsandtetrapods.Thisperiodis
dominatedbyterrestrialforms.Mostcommonareremains generallyknownasRomer’sGap.Ithasbeenhypothesized
of the gigantic arthropod Arthropleura (the so far oldest that the absence of terrestrial arthropods and tetrapods is
known–Schneideretal.2010)followedbytrigonotarbid relatedtolowoxygenlevels,whichinhibitedtheseanimals
andphalangiotarbidarachnids(Rößler&Schneider1997). withpoorlydevelopedrespiratorystructuresfromleaving
The preferred habitat of Arthropleura were, most likely, thewater(Wardetal.2006).Whetherthishypothesiscan
looselyvegetatedriverlandscapes(Schneideretal.2010). bemaintainedorwhethertheabsenceofterrestrialfaunais
(cid:29)(cid:30),
(cid:26)(cid:10)(cid:27)(cid:14)(cid:28)(cid:7)(cid:29)(cid:30)(cid:17)(cid:9)(cid:31)(cid:27)(cid:3) (cid:5)(cid:6)(cid:9)(cid:27)(cid:4)(cid:17)(cid:9)(cid:15)(cid:9)!"(cid:6)(cid:28)"(cid:10)#(cid:10)$(cid:9)(cid:14)(cid:3)(cid:6)(cid:7)(cid:14)(cid:4)(cid:5)(cid:13)(cid:9)(cid:11)"(cid:10)(cid:29)(cid:9)(cid:6)(cid:28)(cid:5)(cid:9)(cid:3)##(cid:5)"(cid:9)(cid:16)(cid:7)(cid:13)%(cid:27)(cid:8)(cid:9)(cid:10)(cid:11)(cid:9)(cid:5)(cid:27)(cid:13)(cid:6)(cid:5)"(cid:8)(cid:9)(cid:12)(cid:5)"(cid:29)(cid:27)(cid:8)&
(cid:2) %
& !
(cid:29)(cid:12)(cid:30)(cid:11)(cid:3)(cid:14)(cid:9)+ Macerated cuticles of extant arthropod appendages and details of their distal claws. (cid:129) A – pseudoscorpion, scale bar = 0.1 mm.
(cid:129)B–Brachistosternussp.(Scorpiones,Chelicerata),scalebar=0.3mm.(cid:129)C–Scutigeracoleoptrata(Chilopoda,Myriapoda),scalebar=0.5mm.
(cid:129) D –Oniscus asellus(Isopoda, Crustacea), scale bar = 0.4 mm.
(cid:29)(cid:30)0
(cid:2)(cid:3)(cid:4)(cid:4)(cid:5)(cid:6)(cid:7)(cid:8)(cid:9)(cid:10)(cid:11)(cid:9)(cid:12)(cid:5)(cid:10)(cid:13)(cid:14)(cid:7)(cid:5)(cid:8)(cid:14)(cid:5)(cid:13)(cid:9)(cid:15)(cid:9)(cid:16)(cid:10)(cid:4)(cid:17)(cid:9)(cid:18)(cid:19)(cid:20)(cid:9)(cid:21)(cid:20)(cid:9)(cid:22)(cid:23)(cid:24)(cid:25)
(cid:2) % & ! ’
(cid:29)(cid:12)(cid:30)(cid:11)(cid:3)(cid:14)(cid:9). Legsofdifferentinsectsforcomparison.AllspecimensfromtheCretaceousCratoFormation,Brazil.(cid:129)A–ROM63044.Tibiaofagrylloid.
Smallspinesalongtheedgeappeartobemovable,scalebar=0.8mm.(cid:129)B–ROM63045.Tarsusanddistalpartoftibiaofablattoid.Tibiabearinglarge
movablespines.Tarsuswithpairedpre-tarsalclaw,scalebar=0.4mm.(cid:129)C–ROM63046.Tibiaofatettigoniid(grasshopper)bearingverylargemovable
spines,scalebar=0.5mm.(cid:129)D–ROM63047.Tarsusandpre-tarsusofanorthopteran.Tarsuselongateinwardcurvedwithaninnerserratededge.Pre-tar-
sal claw well-developed, scale bar = 0.5 mm. (cid:129) E – ROM 63048. Tarsus and pre-tarsus of a possible notonectid bug, scale bar = 0.8 mm.
ratheralackofappropriatedepositsandlocalitiesresulting fordiscussionsofstratigraphyandsedimentologyaswellastech-
inabiasofthefossilrecordneedstobetested.Thefirstre- nicalsupport.JTHwasfundedbytheAlexandervonHumboldt
portbySmithsonetal.(2012)suggeststhatmanytetrapod FoundationasaFeodor-LynenPostdocfellowandbyYaleUni-
versity,andJTHandCHwereadditionallysupportedbyD.E.G.
lineages have their origin much earlier than previously
Briggs(NewHaven).J.-B.Caron,P.FentonandJ.Waddington
thought.
(all Toronto) spent a significant amount of time in making the
Careful analyses of bulk-macerated samples from
specimensfromtheRoyalOntarioMuseumavailabletoJTHand
Chemnitz-Glösa resulted in the recognition of herba-
CH,forwhichtheyareheartilythanked.Wearealsogratefulfor
ceous lycopsids (Hübers et al. 2011) and the discovery
thelong-lastingsupportofS.Liebau(Tübingen)providingaccess
oftheoldestmosses(Hübers&Kerp2012,2013).This toaZeissAxioskop2attheUniversityofUlm.Wethankallpeo-
contributionstressesthatthismethodmayalsorevealan ple involved in providing freely available software such as
unexpected variety of terrestrial arthropods, and thus CombineZM/ZP,MicrosoftImageCompositeEditor,Gimpand
clearlydemonstratesitsgreatpotentialofsedimentsthat OpenOffice.
at first sight only seem to contain organic debris.
Even though the preservation is fragmentary, the
-(cid:14)(cid:16)(cid:14)(cid:3)(cid:14)(cid:21)(cid:10)(cid:14)(cid:15)
taphocoenosisfromthelateViséanHainichenSubgroup
isoneoftheveryfewoccurrencesofEarlyCarbonifer-
ousterrestrialarthropodsandtoourknowledgetheonly BARTRAM,K.M.,JERAM,A.J.&SELDEN,P.A.1987.Arthropodcu-
one from outside Scotland. Together with previously ticlesincoal.JournaloftheGeologicalSociety144,513–517.
DOI 10.1144/gsjgs.144.3.0513
published reports on the biota of the Hainichen Sub-
groupthesefindsprovideimportantinformationforthe BRAUN, A. 1997. Vorkommen, Untersuchungsmethoden und
Bedeutung tierischer Cuticulae in kohligen Sediment-
reconstruction of an Early Carboniferous terrestrial en-
gesteinen des Devons und Karbons. Palaeontographica,
vironment.
Abteilung A 245, 83–156.
CLARKSON,E.N.K.,MILNER,A.R.&COATES,M.I.1994.Palaeoec-
ologyoftheViséanofEastKirkton,WestLothian,Scotland.
(cid:2)(cid:10)*(cid:21)(cid:6),(cid:13)(cid:14)(cid:8)(cid:30)(cid:14)(cid:17)(cid:14)(cid:21)(cid:4)(cid:15)
Transactions of the Royal Society of Edinburgh, Earth Sci-
ences84, 417–425. DOI 10.1017/S0263593300006210
Financial support was provided by the Deutsche For- CLAYTON,G.,COQUEL,R.,DOUBINGER,J.,GUEINN,K.J.,LOBOZIAK,
schungsgemeinschaft (DFG grant KE 584/17-1). We thank S.,OWENS,B.&STREEL,M.1977.Carboniferousmiosporesof
S.Schultka(Berlin)andB.Gaitzsch(Freiberg)formakingpartof westernEurope:illustrationandzonation.MededelingenRijks
theGlösamaterialavailable;additionallyB.Gaitzschisthanked Geologische Dienst 29, 1–71.
(cid:29)(cid:29)