Fossil angiosperm wood and its host deposits from the periphery of a dominantly effusive ancient

14 Pages·2010·3.06 MB·English

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AvarietyoffossilwoodsoccuronthenorthernmarginoftheDoupovskéhoryVolcanicComplexinthenorthwestern
partoftheCzechRepublic.Thewoodswereburiedbythreedifferentprocessesreflectingthreedifferentsettings.First,a
largeareabetweenthetownsofKadaňandKláštereciscoveredbyanupto100mthicksequenceoflaharanddebrisava-
lanchedeposits.TheseflowsandavalanchesgatheredwoodofLiriodendron,Lauraceae,Platanus,Cercidiphyllum,
?CraigiaandStyracaceaefromboththevolcaniccomplexslopesandadjacentplains.Second,arichassemblageoffos-
silwoodswiththermophilouselementssuchasLauraceaeandpalmswaspreservedonthenorthernvolcaniccomplex
peripherybyaStrombolianeruptionofamonogeniccone.Third,ashallowlakeformedtothesideofthevolcaniccom-
plex,wherePlatanustrunkswerefossilizedinthetravertine.ThewoodofLiriodendronhasneverbeenpreviouslyre-
cordedinthelocalitiesrepresentingvolcaniccomplexfoothills,butiscommoninthelocallahardeposits.Thisdistribu-
tion leads us to hypothesize that Liriodendron forests dominated higher topographic levels of the Doupovské hory
VolcanicComplex,reaching,butprobablynotexceeding1000ma.s.l.•Keywords:fossilangiospermwood,systematic
palaeobotany, volcanology, taphonomy, geochronology, Tertiary, Czech Republic.
SAKALA,J.,RAPPRICH,V.&PÉCSKAY,Z.2010.Fossilangiospermwoodanditshostdepositsfromtheperipheryofa
dominantlyeffusiveancientvolcano(DoupovskéhoryVolcanicComplex,Oligocene-LowerMiocene,CzechRepub-
lic): systematics, volcanology, geochronology and taphonomy. Bulletin of Geosciences 85(4), 617–629 (5 figures).
CzechGeologicalSurvey,Prague.ISSN1214-1119.ManuscriptreceivedApril21,2010;acceptedinrevisedformSep-
tember 22, 2010; published online November 12, 2010; issued December 20, 2010.
JakubSakala,CharlesUniversity,FacultyofScience,InstituteofGeologyandPalaeontology,Albertov6,12843Praha
2, Czech Republic; [email protected] • Vladislav Rapprich, Czech Geological Survey, Klárov 3, 118 21 Prague,
CzechRepublic;[email protected]•ZoltánPécskay,InstituteofNuclearResearch,HungarianAcademy
of Sciences, Bem tér 18/C, H-4001 Debrecen, Hungary; [email protected]
Volcanicareasoftenprovideexcellentsettingsforpreserva- itsvicinity,relatedtoOligocenetoearlyMioceneactivity
tionoffossilmaterial,especiallyproducingwhenfinepyroc- oftheDoupovskéhoryVolcanicComplex(DHVCinthe
lastic material predominate. Effusive volcanic activity is following text). Prakash et al. (1971) and Sakala &
muchlessconducivetothefossilizationoforganicmaterial. Privé-Gill (2004) described eight different wood types
Lava-builtvolcanoesandvolcaniccomplexesundergoweath- fromthisarea,hencethelocalityisoneoftherichestsites
ering and solid rock decay processes. Weathering products offossilangiospermwoodinCentralEurope(Sakala2006,
maysubstitutetheroleofpyroclasticdepositsindominantly 2007). Temporary exposures excavated in the last fifteen
effusivevolcanoes.Consequentlyincombinationwiththeto- years,includingconstructionofthenewhospitalanddig-
pography,debrisavalanches,debrisflowsandmudflowsmay ging activity for gas pipe and gas-fixtures in Kadaň, of-
beproduced.Suchsecondarysedimentaryprocessesburyre- fered the opportunity for new sampling and reassessment
mainsofbothfaunaandflora(e.g.,Cameron&Pringle1986). ofthetaphonomicmodelfortheareawhichwasbelieved
FossilwoodisrelativelycommonintheTertiaryofthe tohavebeendominatedbydiatremefaciesofmaarvolca-
northwestern part of the Czech Republic. Most are noes (e.g., Kopecký 2010). The aim of our research is to
Cupressaceaes.l.(e.g.,Teodoridis&Sakala2008).Angio- understandhowwoodfragmentswerefossilizedinsettings
spermwoodisfrequentlyfoundinthetownofKadaňand of a dominantly effusive volcanic complex.
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tivity continued until the Early Miocene (ca 29–22 Ma:
Rapprich & Holub 2008). This younger stage formed an
The origin of the Doupovské hory Volcanic Complex extensivecomplexoffoidite/basanite/tephritelavasequen-
(DHVC – Fig. 1) is associated with the formation of the cesupto500mthick(Hradecký1997a,Rapprich&Holub
continentalEger(Ohře)Grabenandascentofmantlederi- 2008).Individuallavaunitsareoftenaccompaniedbyvol-
vedmelts(e.g.,Ulrychetal.2002)alongtheboundarybet- caniclasticdebrites.Thesecoarsegrainedsedimentsarein-
ween the Saxothuringian and Teplá-Barrandian Domains terpreted as lahar or debris avalanche deposits (e.g., Hra-
oftheBohemianmassif(Mlčoch2003).Acomplexmosaic decký 1997b). Hradecký (1997b) described these
of crystalline rocks (including amphibolite, paragneisse, accumulations from the southern and western margins of
orthogneisse,granulite,phylite,variscangranite,etc.)form theDHVC.Inaddition,wehavedocumentedlahardepo-
thebasementoftheDHVC(Mlčoch&Konopásek2010). sitsalsoonthenorthernmarginsoftheDHVC.Generally,
TheentireDHVCrepresentsanerosionalrelicofamulti- the lahar deposits occur in various stratigraphic positions
phasevolcaniccomplex.Activitywasinitiatedintheearli- withintheDHVC,butthedebritesonthenortheasternmar-
est Oligocene (mammal zone MP-21: Fejfar & Kaiser ginspre-datethelavasoftheÚhošťHill(Rapprich2007).
2005).TheearlyDHVCfine-grainedvolcaniclasticswere TheoldestlavaatthebaseoftheÚhošťHillwasdatedby
produced by basaltic eruptions of the Strombolian and the K-Ar method as 28.66 ±1.06 Ma (Rapprich & Holub
phreato-magmaticstyle.Pyroclasticmaterialfromthisac- 2008). On the northeastern periphery of the DHVC,
tivity was deposited in subaerial, swampy and lacustrine a group of eroded remnants of monogenetic cones
environments with a total thickness reaching 80 m (Hra- (23–20 Ma: this paper) occur and appear to represent the
decký1997a).Thesedepositscurrentlycropoutontheeas- most recent volcanic activity of the DHVC in the Lower
ternandnortheasternmarginsoftheDHVC.Thevolcanic Miocene(Fig.2).Thesedimentaryinfill(includingcoalse-
activity later became predominantly effusive in character ams)oftheEgerGrabenshortlypostdatesthevolcanicac-
(Rapprich&Holub2008),associatedwithweakStrombo- tivityoftheDHVC.Thissedimentaryperiodisrepresented
lianandpossiblyalsoHawaiianeruptions.Theeffusiveac- tothenortheastoftheDHVCbytheMioceneMostBasin.
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(cid:2)(cid:5)(cid:10)(cid:21)(cid:13)(cid:12)(cid:7)-+ SimplifiedgeologicalmapofthenortheasternperipheryoftheDHVC(adaptedfromtheDigitalGeologicalmapoftheCzechRepublic
1:50,000scaleavailableonhttp://www.geology.cz/extranet/geodata/mapserverandmodifiedwithrespecttonewdataandobservations)andlocationof
samplingsites:1–ZadnívrchHill,2–ProstřednívrchHill,3–Vernéřov,4–Nechranice,5–Zvoníčkov.DigitalElevationModelilluminatedfrom
southwest.
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ATOMKI Laboratories, Debrecen, Hungary. Potassium
concentration was measured using a digitized flame pho-
New specimens of silicified and calcified wood were tometer,CORNING480machinewithaLiinternalstan-
thin-sectionedincompliancewiththestandardtechniques dard. The analyses were controlled by inter-laboratory
andstudiedusingcompoundlightmicroscopy.Anatomical standardsAsia1/65,LP-6,HD-B1andGL-O.Argonwas
descriptionsareinaccordancewiththeIAWAHardwood extracted from the samples by high frequency induction
List(IAWACommittee1989).Thethinsectionsdescribed heating.A38Ar-spikewasintroducedintothesystemviaa
herearepartlyhousedintheNationalMuseuminPrague gaspipettebeforethedegassingbegan.Theisotopicratios
(thosedescribedoriginallybyPrakashetal.1971andepi- were measured on a 15 cm radius magnetic sector-type
type of Cercidiphylloxylon kadanense), and partly in the mass spectrometer under static mode, built in Debrecen,
Chlupáč Museum of Earth History in the Faculty of Sci- Hungary.Balogh(1985)andOdin(1982)describedinde-
ence of Charles University in Prague (new specimens). tailthemethodsappliedhere.Thecalculationofageswas
The approximate age of the lahar deposits was deter- based on atomic constants suggested by Steiger & Jäger
mined by their position beneath a lava dated 28.66 ±1.06 (1977). Analytical errors are quoted for the 68% confi-
Ma (K-Ar: Rapprich & Holub 2008), but the age of the dence level (one standard deviation).
monogenetic volcanism on the northern periphery of the
DHVC was not known. Three samples of lavas from
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monogenic volcanoes (Šumná volcano, Blzeňský vrch
Hill, Jelení vrch Hill) on the northern periphery of the
DHVC were dated using the K-Ar method. All the three AngiospermwoodfromtheDHVC,bothnewlyfoundspe-
samplesweremeasuredasbulk-rocks.Additionally,glass cimens and those described by Prakash et al. (1971) and
and plagioclase fractions were separated from the Sakala&Privé-Gill(2004),canbesubdividedintoseven
groundmass of the Jelení vrch Hill sample to resolve the units called “wood types” sensu Wiemann et al. (1998,
problemoflowpotassiumcontentinthebulk-rocksample. p.85).Thesearelistedusinginformalnamesanddescribed
Geochronological analyses were carried out in the in the following section.
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includesbothtransverseandradialviews(Fig.4A).Other
diagnostic features, such as oval opposite intervessel
pits are visible in the holotype (Prakash et al. 1971,
fig. 33) as well as in the new specimens, e.g., 18/98
(Fig. 4C). Leaves and fruits of Liriodendron are un-
knownfromtheDHVC,butthereareoccurrencesofLi-
riodendronhaueriintheČeskéstředohoříMts(Kvaček
&Teodoridis2007).
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Family Lauraceae
GenusLaurinoxylonFelix emend. Dupéronet al.
(cid:2)(cid:5)(cid:10)(cid:21)(cid:13)(cid:12)(cid:7).+ CalcifiedtrunkofLiriodendroninthelahardepositsexposed
during diggingwork for gas-pipes in Kadaň, southern slope of the
Laurinoxylon czechensePrakash, Březinová & Bůžek
Prostřední vrch Hill.
Figure 4D–G
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Material.–Kadaň-ZadnívrchCNB-2(type);?Nechranice
74/04.
Family Magnoliaceae
Description.–Wooddiffuse-porous,growthringboun-
GenusLiriodendroxylonPrakash, Březinová & Bůžek daries distinct, vessels mostly solitary and in radial
(and oblique) multiples of 2–3, predominantly simple
Liriodendroxylon tulipiferumPrakash, Březinová perforation plates, scalariform perforation plates with
& Bůžek 10–15barsrarelypresentinnarrowvesselelements,he-
Figure 4A–C terocellularrays1-to3-seriatewithsomeenlargedoil/
mucilage cells in the margins, vasicentric axial paren-
Material. – Kadaň-Zadní vrch ZV-23 (holotype), 18/98, chyma.
53/02; Vernéřov 57/02, 61/02.
Discussion. – Dupéron-Laudoueneix & Dupéron (2005)
Description. –Wood diffuse-porous, growth ring bounda- inventoried the fossil woods assigned to the Lauraceae.
riesdistinct,vesselsrarelysolitarymostlyinradialmulti- TheoriginalslidesofLaurinoxylondiluviale(Unger)Fe-
plesandclusters,scalariformperforationplateswithabout lix, the type species of Laurinoxylon known since 16th
10 bars, opposite intervessel pits with oval outlines, century as ‘Sündfluthholz’ from the phreatomagmatic
slightlyheterocellularraysmostly3-to4-seriate,andmar- vent breccia at Jáchymov (north of the DHVC, in the
ginal axial parenchyma. Krušné hory Mts/Erzgebirge), were re-discovered in the
FrenchMuseumofNaturalHistory(MNHN)inParisand
Discussion.–Prakashetal.(1971)notedthatthiswood re-described,togetherwithemendedgenericandspecific
type was similar to modern Liriodendron tulipifera L. diagnoses(Dupéronetal.2008).Thefossilwoodbriefly
(seeInsideWood2004–onwards).Marginalparenchyma describedherehascharacteristicsofthemorphogenusLau-
isvisibleinslideG4049oftheholotypespecimenwhich rinoxylon.Prakashetal.(1971)recognizeditasadistinct
(cid:2)(cid:5)(cid:10)(cid:21)(cid:13)(cid:12)(cid:7)/+ A–C–Woodtype1–Liriodendroxylon(A,B:holotypeZV-23ofLiriodendroxylontulipiferum;C:specimen18/98).•A–diffuse-porous
woodwithvesselsrarelysolitary,usuallyinmultiplesandterminalparenchymain3-seriatelines,TS/RLS.•B–slightlyheterocellularraysmostly
3–4-seriateandscalariformperforationplateswithabout10bars,TLS.•C–oppositeintervesselpits,RLS.•D–G–Woodtype2–Laurinoxylon
(holotypeCNB-2ofLaurinoxylonczechense).•D–diffuse-porouswoodwithparatrachealvasicentricparenchyma,TS.•E–heterocellular1–3-seriate
rayswithswollenmarginalcells,TLS.•F–raycellswithoilormucilagecontentinmargin,RLS.•G–detailofasmallvesselelementwithalternate
intervessel pits in lower part and scalariform perforation plate in upper part, TLS. • H–M – Wood type 3 – Platanus (H, K: holotype CNB-6 of
Platanoxylonbohemicum;I,L,M:holotypeZV-24ofSpiroplatanoxyloneuropeanum;J:holotypeCNB-11ofDryoxylonbohemicum).•H–J–dif-
fuse-porouswoodwithdistinctgrowthringsandbroadrays,TS.•K–broadhomocellularraysupto20cellswide,TLS.•L–detailofhomocellularray
andexclusivelyscalariformperforationplateswithabout20bars,TLS.•M–helicalthickeningonthetopofavesselelement,TLS.Scalebars=500μm
in A, B, D, H–K; 200 μm in E, F, L; 50 μm in C, G, M.
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speciesL.czechense.ItdiffersfromL.diluvialeinhaving DryoxylonSchleideninSchmid
generallythinnerrays,mainlyduetonarrowerindividual
raycells.Thisdifferenceinraywidthcanpartlybeexplai- Dryoxylon bohemicumPrakash, Březinová & Bůžek
nedbytheprocessesoffossilizationwhichcouldproduce Figure 4J
moredilatedraycellsinL.diluviale(M.andJ.Dupéron,
pers.comm.).ContrarytoPrakashetal.(1971),weobser- 1971 Dryoxylonbohemicumsp.nov.;Prakash,Březinová
vedoilcellssimilartothoseinL.diluviale(Fig.4E,F)as- & Bůžek, p. 122, pl. 43, figs 64–67.
sociated with ray parenchyma only (but not those
amongst fibres) and very rarely scalariform perforation Material. – Kadaň-Zadní vrch CNB-11 (type).
platesinnarrowvesselelements(Fig.4G).Therefore,itis
not easy to distinguish between these two species. Idio- Description.–Wooddiffuseporous,growthringbounda-
blasts (oil/mucilage cells) associated exclusively with riesdistinct,vesselssolitaryorinshortirregularmultiples
rays are present in several taxa, e.g., Caryodaphnopsis, withscalariformperforationplatesonly(simpleplatesnot
Litsea chinensis or the south-American species of Cin- observed)withabout20bars,sometimeswithspiralthic-
namomum (Richter 1987). As vessel-ray pitting is not kenings,homocellularraysupto20cellswideandapotra-
preserved,itisnotpossibletousecorrectlytheclassifi- cheal diffuse-in-aggregates and scanty paratracheal vasi-
cationsensuRichter(1987).Asaresult,wecannotattri- centric axial parenchyma, rarely crystalliferous.
buteourwoodtoanyparticularlivinggenus.Theperip-
heryoftheDHVChasyieldedthermophilousfloraswith Discussion.–ThethreespeciesrecognizedbyPrakashet
Daphnogene(Kvaček&Teodoridis2007),whichiscon- al.(1971),i.e.,Platanoxylonbohemicum,Plataniniumeu-
sidered to be related to the living Cinnamomum cam- ropaeum(=Spiroplatanoxyloneuropeanum)andthevery
phora (L.) J. Presl (Kvaček et al. 2004, Kvaček pers. poorly preserved Dryoxylon bohemicum, are very similar
comm.). toeachother.Theyallhavelargehomocellularrays,upto
24cellswide,andscalariformperforationplates.Although
there are several differences between them (e.g., smaller
(cid:2)(cid:3)(cid:3)(cid:4)(cid:5)(cid:6)(cid:7)(cid:8)(cid:9)(cid:5)(cid:13)(cid:5)(cid:11) *(cid:4)(cid:26)(cid:6)(cid:26)(cid:8)(cid:3)(cid:13)
vesselelementsinPlatanoxylonbohemicumtypespecies)
theyallseemtobelongtothesametypeofplatanoidwood
Family Platanaceae whoseexactaffinityisuncertain.Wethinkthewoodmust
be related to Platanus neptuni (Ettingsh.) Bůžek, Holý &
PlatanoxylonAndreánszkyexPrakash, Březinová Z.Kvaček(Sakala2006,Kvaček&Manchester2004),the
& Bůžek only member of the Platanaceae present in both volcanic
areasoftheDHVCandtheČeskéstředohoříMts(Kvaček
Platanoxylon bohemicumPrakash, Březinová & Bůžek & Teodoridis 2007). P. neptuni is otherwise known as a
Figure 4H, K nearly complete plant based on foliage branches, isolated
leaves, stipules, staminate inflorescences with pollen in
1971 Platanoxylon bohemicum sp. nov.; Prakash, Březi- situandinfructescences(Kvaček2008).Thespecificsys-
nová & Bůžek, p. 115, pl. 39, figs 44, 46. tematicpositionofP.neptuniwithinPlatanusisexpressed
by a distinct subgenus Glandulosa (see in Kvaček et al.
Material.–Kadaň-ZadnívrchCNB-6(type),73/03;?Ne- 2001,table1)andcorrespondswelltothespecificcharac-
chranice 109/05, 114/06; ?Vernéřov 64/02. ter of our fossil wood (see here below).
ThefossilwoodrelatedtoPlatanusisusuallyclassified
under the mophogenera Platanoxylon or Plataninium. As
SpiroplatanoxylonSüss alreadynoticedbyBrett(1972)andWheeler&Manchester
(2002, p. 97), the classification of fossil Platanus-like
Spiroplatanoxyloneuropeanum(Prakash, Březinová woodsisnotasimplematterbecausetheygenerallydiffer
& Bůžek) Süss from the extant Platanus. Firstly, fossil platanoid wood
Figure 4I, L, M often has scalariform perforation plates only (Wheeler
1995). The only living representative of Platanus which
1971 Plataninium europeanum sp. nov.; Prakash, Březi- bears some resemblence with respect to this feature i.e.
nová & Bůžek, p. 120, pl. 42, figs 59–63. having a higher portion of scalariform to simple perfora-
2007 Spiroplatanoxyloneuropeanum(Prakash,Březinová tion plates, is P. kerrii Gagnep., native to Laos and Viet-
& Bůžek) Süss comb. nov.; Süss, p. 12. nam (Wheeler 1995). Contrary to Prakash et al.’s (1971)
description, and in accordance with Süss & Müller-Stoll
Material. – Kadaň-Zadní vrch ZV-24 (type, CNB-11). (1977,p.50),wehavenotobservedanysimpleperforation
(cid:26)--
(cid:25)(cid:26)(cid:27)(cid:3)(cid:28)(cid:29)(cid:26)(cid:27)(cid:26)(cid:4)(cid:26)(cid:5)(cid:6)(cid:9)(cid:26)(cid:4)(cid:17)(cid:15)(cid:30)(cid:10)(cid:13)(cid:13)(cid:7)(cid:4)(cid:9)(cid:26)(cid:8)(cid:31)(cid:7)(cid:10)(cid:13) (cid:5)!"(cid:9)#(cid:10)(cid:10)$(cid:9)(cid:26)(cid:8)$(cid:9)(cid:7)(cid:6)(cid:13)(cid:9)%(cid:10)(cid:13)(cid:6)(cid:9)$(cid:5) (cid:10)(cid:13)(cid:7)(cid:6)(cid:13)(cid:9)(cid:11)!(cid:10)"(cid:9)(cid:6)%(cid:5)(cid:9) (cid:5)!(cid:7) %(cid:5)!&(cid:9)(cid:10)(cid:11)(cid:9)(cid:26)(cid:9)$(cid:10)"(cid:7)(cid:8)(cid:26)(cid:8)(cid:6)(cid:4)&(cid:9)(cid:5)(cid:11)(cid:11)(cid:3)(cid:13)(cid:7)’(cid:5)(cid:9)(cid:26)(cid:8)(cid:14)(cid:7)(cid:5)(cid:8)(cid:6)(cid:9)’(cid:10)(cid:4)(cid:14)(cid:26)(cid:8)(cid:10)
platesinoursamplesofplatanoidwood.Secondly,asSüss foliage,fruits,seeds,staminateinflorescences,andinsitupo-
& Müller-Stoll (1977) described, some platanoid woods llen(Kvaček2008).
havespiralthickeninginthevesselelements.Thisfeature
is unknown among extant Platanus woods (Wheeler &
(cid:2)(cid:3)(cid:3)(cid:4)(cid:5)(cid:6)(cid:7)(cid:8)(cid:9)(cid:5)(cid:15)(cid:5)(cid:11) ,(cid:4)"(cid:3)(cid:13)
Manchester 2002). Süss (2007) recently created a new
morphogenus,Spiroplatanoxylon,forplatanoidwoodwith
spiral thickening and re-interpreted Platanoxylon and Family Ulmaceae
Plataninium from Kadaň as belonging to this genus. We
can confirm the presence of spiral thickenings (Fig. 4M) GenusUlmusL.
and rare prismatic crystals in axial parenchyma cells in
PlataniniumbutnotinPlatanoxylon.Therefore,weusethe Ulmussp.
denominationSpiroplatanoxylononlyforthePlataninium Figure 5C
typeofwoodinwhichweobservedspiralthickeningand
crystals; as a precaution, we also keep the three Material. – Nechranice 75/04.
morphospeciesasseparatedunits.However,wethinkthey
all represent the same botanical species. Description. – A small branch, 2.5 cm in diameter, with
barkpreserved.Woodring-porous,growthringboundaries
distinct,earlywoodvesselsmainlysolitary,latewoodves-
(cid:2)(cid:3)(cid:3)(cid:4)(cid:5)(cid:6)(cid:7)(cid:8)(cid:9)(cid:5)(cid:14)(cid:5)(cid:11) +(cid:5)!(cid:14)(cid:7)$(cid:7) %&(cid:4)(cid:4)(cid:10))&(cid:4)(cid:10)(cid:8)
selsgrouped,formingtypicalwavytangentialbands,wide
exclusively homocellular rays.
Family Cercidiphyllaceae
Discussion.–Thissinglepieceofwoodwasalreadymenti-
GenusCercidiphylloxylonPrakash,Březinová&Bůžek oned by Sakala (2006). Unfortunately, the very poor pre-
servation only allows its safe attribution to elm wood
Cercidiphylloxylon kadanensePrakash, Březinová thanks to the diagnostic pattern seen in cross-section and
& Bůžek thehomocellularraysseeninradialsection.Acomparison
Figure 5A, B with the fossil elm wood described from Bílina (Sakala
2002) or with other Ulmaceae (Wheeler & Manchester
Material.–Kadaň-ZadnívrchZV-12(holotype),G8113(= 2007)isnotpossible.IntheTertiaryofnorthwesternBohe-
23/98) (epitype). mia, the genus Ulmus is represented by 2 species: U. fis-
cheriasleavesinvolcanicareasofboththeDHVCandthe
Description.–Wooddiffuse-porous,growthringbounda- ČeskéstředohoříMtsandU.pyramidalisintheMostBasin
riesdistinct,vesselsmostlysolitary,withangularoutline, based on leaves and fruits (Kvaček & Teodoridis 2007).
scalariform perforation plates with about 40 bars, hetero-
cellular, mostly 2–3-seriate rays with uniseriate marginal
(cid:2)(cid:3)(cid:3)(cid:4)(cid:5)(cid:6)(cid:7)(cid:8)(cid:9)(cid:5)(cid:16)(cid:5)(cid:11)(cid:5)(cid:17)+!(cid:26)(cid:7)(cid:31)(cid:7)(cid:26)
rows(tails),whichsometimesinterconnectseveralmulti-
seriate ray portions.
Family Malvaceae s.l.
Discussion. – This wood type was originally described by
Prakashetal.(1971)andlaterreviewedindetailbySakala& GenusCraigiaW.W. Smith & W.E. Evans
Privé-Gill(2004)withthedefinitionofanepitypeandcom-
parativestudywiththeextantCercidiphyllaceaeandHama- aff.Craigiasp.
melidaceae.Itwasnoticedthat1)Cercidiphylloxylonkada- Figure 5D–G
nense slightly differed from extant Cercidiphyllum Sieb. &
Zucc.,2)itwastheoldestrecordoftrueCercidiphyllumfossil Material. – Kadaň-Zadní vrch 72/03; Nechranice 70/03,
woodand3)anolderrecordofCercidiphyllum-likewoodof 78/03, 84/04, 89/04, 90/04.
Paleocene/Eoceneagehadtoberelatedtoextinctgenerawith
leavesofTrochodendroidesBerryemend.Craneandfruitsof Description.–Woodsemi-ringporous,growthringboun-
the Nyssidium-type (Sakala & Privé-Gill 2004). Our fossil daries distinct, marked by marginal parenchyma, early-
woodmostprobablyisthewoodofCercidiphyllumcrenatum woodvesselssolitaryorinshortradialmultiplesof2–3,
(Unger)R.W.BrownwhichoccursinallOligocenelocalities latewoodvesselsthick-walled,mainlyinradialmultiples
in the České středohoří Mts (Kvaček & Teodoridis 2007). of2–3,bothwithsimpleperforationplates,stronglyhe-
However,itsoccurrenceintheDHVChasnotbeenprovenso terocellular rays up to 10 cells wide with tile cells
far.C.crenatumisknownasanearlycompleteplantbasedon of the Pterospermum type, apotracheal diffuse to
(cid:26)-(cid:31)
(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:23)
diffuse-in-aggregates and paratracheal vasicentric axial Material. – Kadaň-Zadní vrch CNB-3 (type).
parenchyma.
Description.–Wooddiffuseporous,growthringbounda-
Discussion.–ItwasrecentlysuggestedbySakala(2006)that riesdistinct,vesselssolitary,butmostlyinclustersandra-
thisnewlydiscoveredfossilwoodcouldbeattributedtoMal- dial multiples, scalariform perforation plates with about
vaceaes.l.duetoitsdiagnostictilecellsintherays,andtothe 15–20 bars, heterocellular uniseriate and multiseriate
extant genus Craigia, whose wood was described by Man- (mostly 3- to 4-seriate) rays, axial parenchyma diffuse or
chesteretal.(2006).TilecellsoccurinMalvaceaes.l.andare diffuse-in-aggregates, chambered axial parenchyma with
sometimescitedasarareexampleofsynapomorphybasedon prismatic crystals often present.
woodanatomy(Olson2005,p.514);however,theyarealso
present in several species of Hopea from the Dipterocarpa- Discussion. – The difference in vessel arrangement bet-
ceae(orderMalvales)(e.g.,Manchesteretal.2006,Selmeier weenCoryloxylonnemejciiandC.tertiarumdonotseem
2000,table3).Infact,theCzechwoodtypeissimilartoboth tobesystematicallysignificantandcanbeexplainedbyin-
the extant Craigia and extinct Chattawaya Manchester traspecificorindividualvariability.Weconsiderthatboth
(1980)initsvesseldispositionandcharacteroftilecells.Un- woods represent the same wood type, the affinities of
fortunately,wedidnotobserveanyhelicalthickeningtypical which need re-evaluation. We did not observe the aggre-
of Cragia; the seemingly helically thickened vessels in one gateraysdescribedbyPrakashetal.(1971)sotheconnec-
specimen(Fig.5G)donotrepresenttruespiralsbutonly“co- tion to Corylus becomes questionable. Both woods have
alescent apertures” of intervessel pits (E.A. Wheeler, pers. features seen in the Styracaceae: exclusively scalariform
comm.).Amongfossilrepresentatives,thehelicalthickening perforationplates,poressolitaryorinmultiples,bothuni-
is only present in Reevesia japonoxyla Terada & Suzuki and multiseriate heterocellular rays, diffuse and
(1998).Ontheotherhand,wedidnotobservecrystals,which diffuse-in-aggregates axial parenchyma and prismatic
areverytypicalofChattawaya(Manchester1980,Wheeler& crystalsorsilica(Dickison&Phend1985).Prismaticcrys-
Manchester 2002). Therefore, we call our fossil wood “aff. tals are useful for distinguishing between species within
Craigiasp.”,reflectingthesimilaritytothisgenusexceptfor the extant Styracaceae. These are present only in Bruin-
the typical helical thickening. Although unknown in the smia,HalesiaandStyrax:Bruinsmiadiffersstronglyfrom
DHVC, Craigia represents an important and abundant ele- our wood in lacking growth rings, but a clear distinction
mentintheČeskéstředohoříMtsandalsointheMostandSo- betweenHalesiaandStyraxisnotpossiblebasedonlyon
kolovbasins(Kvaček&Teodoridis2007).Itisgenerallypre- wood (Dickison & Phend 1985). In the David A. Kribs
sent as dispersed fruit valves, but also as complete fruits at wood collection housed in the N.C. State University, we
variousstagesofmaturity,dispersedflowersandflowerbuds observedsimilarstructureintheextantHalesiamacgrego-
withpolleninsitu,oftentogetherwiththeaccompanyingfoli- rii Chun (SJRw 29811) from China, however, it lacked
ageofDombeyopsislobata(Kvaček2008). crystals.Therefore,wearenotabletodistinguishbetween
StyraxandHalesia.IntheTertiaryofnorthwesternBohe-
mia,theonlyrecordofStyracaceaesofarcomesfromthe
(cid:2)(cid:3)(cid:3)(cid:4)(cid:5)(cid:6)(cid:7)(cid:8)(cid:9)(cid:5)(cid:18)(cid:5)(cid:11) +(cid:10)!&(cid:4)(cid:10))&(cid:4)(cid:10)(cid:8)
Early Miocene Cypris formation of the Sokolov Basin –
therearefruitsdesignatedasSinojackiasp.byBůžeketal.
Family Styracaceae 1996(recordedasHalesiacrassainKvaček&Teodoridis
2007).
GenusCoryloxylonPrakash, Březinová & Bůžek
(cid:28)(cid:3)(cid:6)(cid:23)(cid:8)(cid:9)(cid:3)(cid:6)(cid:3)(cid:10)(cid:20)(cid:7)(cid:8)(cid:9)(cid:16)(cid:7)(cid:4)(cid:12)(cid:16)(cid:5)(cid:14)(cid:12)(cid:9)(cid:17)(cid:3)(cid:6)(cid:3)(cid:10)(cid:20)
Coryloxylon nemejciiPrakash, Březinová & Bůžek
Figure 5H–J
Materialforstudywassampledfromfivelocalities(Fig.2)
1971 Coryloxylonnemejciisp.nov.;Prakash,Březinová& representingthreedifferentvolcanogenicsedimentaryde-
Bůžek, pp. 116–118, pl. 40, figs 48–52. posits:
Material. – Kadaň-Zadní vrch CNB-4 (type).
(cid:19)(cid:20)(cid:4)(cid:21)(cid:22)(cid:5)(cid:23)(cid:24)(cid:25)(cid:26)(cid:27)(cid:5)(cid:28)(cid:24)(cid:3)(cid:29)(cid:6)(cid:30)(cid:9)(cid:4)(cid:21)(cid:22)(cid:5)(cid:23)(cid:24)(cid:25)(cid:26)(cid:27)(cid:5)(cid:31)(cid:9)(cid:24)(cid:21) (cid:30)(cid:3)(cid:23)
!"(cid:3)(cid:25)(cid:20)"#(cid:6)#(cid:9)(cid:29)(cid:5)(cid:10)(cid:27)(cid:5)(cid:12)(cid:5)(cid:20)(cid:21)(cid:4)(cid:5)(cid:13)(cid:5)#(cid:21)(cid:5)$#%&(cid:5)(cid:12)’
Coryloxylon tertiarumPrakash, Březinová & Bůžek
Figure 5K–M
Localities1,2and3aresituatedwithinthevolcaniclastic
1971 Coryloxylon tertiarum sp. nov.; Prakash, Březinová debritesoftheDHVC(Fig.2).Thelocalitieswereartifi-
& Bůžek, pp. 118–120, pl. 41, figs 53–58. cial trenches (length × width × depth = 10 × 1 × 1 m at
(cid:26)-
(cid:25)(cid:26)(cid:27)(cid:3)(cid:28)(cid:29)(cid:26)(cid:27)(cid:26)(cid:4)(cid:26)(cid:5)(cid:6)(cid:9)(cid:26)(cid:4)(cid:17)(cid:15)(cid:30)(cid:10)(cid:13)(cid:13)(cid:7)(cid:4)(cid:9)(cid:26)(cid:8)(cid:31)(cid:7)(cid:10)(cid:13) (cid:5)!"(cid:9)#(cid:10)(cid:10)$(cid:9)(cid:26)(cid:8)$(cid:9)(cid:7)(cid:6)(cid:13)(cid:9)%(cid:10)(cid:13)(cid:6)(cid:9)$(cid:5) (cid:10)(cid:13)(cid:7)(cid:6)(cid:13)(cid:9)(cid:11)!(cid:10)"(cid:9)(cid:6)%(cid:5)(cid:9) (cid:5)!(cid:7) %(cid:5)!&(cid:9)(cid:10)(cid:11)(cid:9)(cid:26)(cid:9)$(cid:10)"(cid:7)(cid:8)(cid:26)(cid:8)(cid:6)(cid:4)&(cid:9)(cid:5)(cid:11)(cid:11)(cid:3)(cid:13)(cid:7)’(cid:5)(cid:9)(cid:26)(cid:8)(cid:14)(cid:7)(cid:5)(cid:8)(cid:6)(cid:9)’(cid:10)(cid:4)(cid:14)(cid:26)(cid:8)(cid:10)
Prostřednívrch)andearthworks(length×width×depth= thevalleyamidsthillsrepresentingremnantsoftheDHVC
300×200×2matVernéřov,300×100×1-3and300× lava sequences, but the contact is not exposed at present.
200×3matZadnívrch).Similardepositswereobservedin WecategorizethissiteasEarlyDHVCvolcaniclasticson
allthreelocalities.Thesedimentsformsubhorizontalbeds thebasisofdataobtainedduringthelate18thandearly19th
upto2mthick,representedbyapoorlysortedpolymictic century,whenceladonitewasexploitedintheareaofZvo-
(fragments of various types of basaltic rocks), níčkov and eastern foothills of the Úhošť Hill. Großkopf
coarse-grained(basalticbouldersupto40cmindiameter) (1932)andMüller(1936)describedthesedimentshosting
matrix-supporteddebrite.Thevolcanigenicmatrixconsists the travertine as a sequence of alternating layers of tuffs,
offinebasalticdetritusandclinopyroxenecrystals.Theto- claysandlimestone.Thelithologicalschemeissupported
talthicknessofthelahardepositsobservedontheJezerní by a 114 m deep borehole (Müller 1936).
hora Hill reaches 100 m. Xenolithic clasts occur rarely,
they were found in some depositional units of debrites
)(cid:12)(cid:3)(cid:23)(cid:18)(cid:13)(cid:3)(cid:9)(cid:3)(cid:6)(cid:3)(cid:10)(cid:20)
aroundKlášterecandtheyconsistofhorizontallystretched
violetclayswithquartzcrystals,mostprobablygneissar-
gillized by weathering already prior to the volcanic acti- Threemonogeneticvolcanoesfromthelatestphaseofvol-
vity.Thedepositscontainabundanttreetrunks,uptoseve- canicactivityonthenorthernperipheryoftheDHVCwere
ralmeterslongand1mindiameter,andfragmentsoftree dated using the K-Ar method: Šumná Volcano, Blzeňský
branches.Thetrunksaredistributedrandomlyinthehost vrchHillandJelenívrchHill.Despitethesimilarsettingof
rock and subhorizontally orientated (Fig. 3). The number thethreevolcanoes,theircompositiondifferssignificantly.
of trunks exposed during occasional excavations was not ThelavaoftheŠumnávolcanohasatrachybasalticcompo-
highenoughtoevaluatethepossiblepreferredorientation sition,apicrobasalteruptedontheJelenívrchscoriacone,
statistically. theČachovickývrchHilliscomposedofanalteredbasa-
nite(notsuitableforK-Aranalysis)andalimburgiteforms
the Blzeňský vrch lava. The trachybasalt of the Šumná
((cid:9)(cid:25)(cid:26)(cid:24)(cid:20)(cid:21)#(cid:25)(cid:9) !"(cid:3)(cid:25)(cid:20)"#(cid:6)(cid:7)(cid:5)(cid:14)(cid:5)#(cid:21)(cid:5)$#%&(cid:5)(cid:12)’
Volcanowassampledinthecastle-trenchonthetopofthe
ŠumnáHill(541ma.s.l.).Theremnantofalavalakeinside
Opalizedandcalcifiedwoodfragmentsoccuratthebaseof thescoriaconecraterisexposedthere.Apicrobasaltfeeder
fine- to medium-grained scoriae on the banks of the Ne- dykeofascoriaconeremnantcropsoutonthepeakofthe
chranice dam on the southern slopes of the Čachovický JelenívrchHill(363ma.s.l.).Basanitelavawassampled
vrchHill.Thegeologicalclassificationwasreconstructed intheactivequarrycurrentlyoccupyingtheentireBlzeň-
onthebasisofseveralsmalloutcrops.Thelowerpartofthe skývrchHill.Suitablepotassiumconcentrationsproduced
sectionisaccessibleonlywhentheNechraniceLakeisem- reasonabledatafortheŠumnáVolcano(20.58±0.66Ma)
ptiedduringdryseasons.Thebasementoftheprofilecon- andthelavaofBlzeňskývrchHill(22.51±1.06Ma).The
sistsofthefossil-weatheredgneisseoverlainbyapyroclas- K OcontentinthepicrobasaltoftheJelenívrchHilldoes
2
tic deposit. Locally, relics of pre-volcanic sediments – notexceed0.5%.Hence,acceptableresultscouldbeob-
sandstonesderivedfromcrystallinerocks,arealsopresent. tainedneitherbyprocessingabulk-rocksamplenorbypro-
The non-welded, loose pyroclastic rocks do not create a cessing the plagioclase/glass fraction separated from the
realoutcropthereandthethicknesscanonlybeestimated groundmass.
as not exceeding 1 m. The pyroclastic deposit consists of The data obtained from Šumná volcano and Blzeňský
basaltic scoria fragments, 1–2 cm in diameter. The frag- vrch Hill yield an Aquitanian age and can be correlated
mentsoffossilizedwoodscanonlybefoundwherepyroc- with the youngest lava of the Úhošť Hill profile (22.09 ±
lasticsoccur.Thefossilizedwoodwascollectedinanarea 0.73 Ma, Rapprich & Holub 2008).
200×20m,onthebanksoftheNechraniceLake.Thepy-
roclasticdepositsareoverlainbyafewmetersthickbasani-
(cid:25)(cid:5)(cid:4)(cid:23)(cid:21)(cid:4)(cid:4)(cid:5)(cid:3)(cid:9)(cid:7)(cid:3)(cid:9)(cid:7)(cid:17)(cid:8)(cid:11)(cid:18)(cid:3)(cid:9)(cid:3)(cid:14)(cid:20)(cid:7)(cid:8)(cid:9)(cid:16)(cid:7)(cid:4)(cid:17)(cid:13)(cid:8)(cid:17)(cid:5)(cid:10)(cid:13)(cid:8)(cid:11)(cid:18)(cid:20)
ticlavaflow,whichextendstothesitefromthetopofČa-
chovický vrch Hill.
Thispaperfocusesonfossilizedwoodfromfivesitesfrom
thenorth-easternmarginsoftheDHVC,representingthree
(cid:19)(cid:23)(cid:3)(cid:21)(cid:22))*(cid:3)(cid:23) !"(cid:3)(cid:25)(cid:20)"#(cid:6)(cid:7)(cid:5)(cid:15)(cid:5)#(cid:21)(cid:5)$#%&(cid:5)(cid:12)’
different types of volcaniclastic deposits (Fig. 2).
TheareabetweenthetownsofKadaňandKlášterec–
CalcifiedtreetrunkswerefoundintheabandonedZvoníč- where the localities Zadní vrch Hill, Prostřední vrch Hill
kovvillageamongfragmentsoflacustrinetravertineconta- and Vernéřov are situated (localities 1–3 in Fig. 2) – is
ining numerous cavities that originated from herbaceous mainly composed of polymictic volcanogenic debrites.
stems. The travertine containing the tree trunks occurs in Thesepolymicticmatrix-supportedvolcanogenicdeposits
(cid:26)-.
(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:23)
withhorizontallyorientatedtreetrunksreflectsedimenta- volcaniclastics.Thesedepositsaretraditionallydescribed
tion from debris flow – lahar. The older interpretation by as“tuffs”,butnomodernvolcanologicalstudyhasyetbeen
Kopecký (e.g., 2010) considering maar volcanoes is re- focussedonthem,mainlyduetotheabsenceofgoodexpo-
jected here because the debrites display no signs of sures. The fine-grained volcaniclastics that underlie the
phreatomagmatic fragmentation, xenolith content is low Úhošť lavas can most probably be correlated with the
andtheirpresenceislimitedsolelytosomelayersandthe fine-grained volcaniclastics with limestone intercalations
debrites comprise subhorizontal beds of thickness up to knownfromthesoutheasternmarginsoftheDHVC(mam-
2m.Ontheotherhand,diatremebrecciaspenetratingcrys- mal zone MP-21: Fejfar & Kaiser 2005, i.e. 34 Ma). The
talline units should contain one type of basaltic rock (not fine-grainedvolcaniclasticsatZvoníčkovthereforerepre-
various types of basaltic rocks as from the DHVC) and sent products of early DHVC activity in the Early
highcontentofupper-crustalxenoliths.Numeroustypesof Ruppelian.
volcanic rocks would be present only if diatreme pene- The coarse-grained and strongly vesiculated character
tratedthepre-existingvolcanicsequencewhichisnotthe ofthepyroclastsatNechranicereflectsalocallow-energy
case in the northern DHVC periphery. magmatic-gas driven volcanic eruption (Strombolian).
Thelaharsmusthaveoriginatedfromaterrainwithhigh The wood assemblage at Nechranice is relatively rich. It
relative elevation – most likely in the central part of the containsnotonlythetypesdescribedabove,butalsonew
DHVC. The volcanic complex very probably reached alti- unidentified angiosperm wood (with at least three new
tudesof1000ma.s.l.initscentralpartatthattime(thehigh- types,probablyrelatedtosubtropicalLauraceaeandother
est part of the early DHVC edifice is preserved in an ero- families)andonesilicifiedpalmstem(No.121/07),which
sionalremnantonthePustýzámekHill,927ma.s.l.inthe aresubjectofongoingstudies.Theplantswereburiedrap-
centreoftheDHVC).Thetreesweredraggedbythemass idlybythepyroclasticfallout.Thesubsequentlavaeffu-
flowsduringtheirdescentdowntheslopesofthevolcano. sion protected the accumulation from being eroded. The
Theprocessmayexplainthehighspeciesdiversityfromthe monogenetic volcanism on the northern periphery can be
Zadní vrch Hill. The lower species diversity of the Pro- assignedtotheAquitanianaccordingtothenewdatapre-
střednívrchandVernéřovlocalitiesisveryprobablydueto sentedinthispaper.However,wewerenotabletoanalyze
the limited extent of occasional exposures. We consider exactly the Běchovický vrch Hill, the surrounding
Liriodendron-dominatedforeststobetypicalofhigheralti- monogenic volcanoes yielding data within the range of
tudesonthevolcaniccomplexflanksasthiswoodtypehas 20.58±0.66to22.51±1.06Ma.Themonogenicactivity
notbeenobservedinthelowlanddepositssurroundingthe onthenorthernmarginoftheDHVCwascontemporane-
volcaniccomplex.Liriodendrontodayisastraight,talltree, ouswiththeyoungestlavaoftheÚhošťprofile(Rapprich
whichisfortwo-thirdsofthetrunkfreeoflateralbranches & Holub 2008).
and famous for its considerable height; nowadays it is the
tallestofallEasternUnitedStatesangiospermtrees,andcan
(cid:29)(cid:3)(cid:9)(cid:23)(cid:6)(cid:21)(cid:4)(cid:5)(cid:3)(cid:9)(cid:4)
attain60minheight(Beck1990).Wefoundapieceatrunk
40cmindiameter(Fig.3)whichwouldroughlycorrespond
toanatleast30mtalltree(Beck1990,table1).Theclastic We have distinguished three types of deposits reflecting
material of the sediment is derived from the decay of an threedifferentvolcanicenvironments:1)lahardepositson
EarlyRuppelianvolcanicedifice.Thelaharsareoverlainby theZadnívrchHill,ProstřednívrchHillandatVernéřov,
alavaremnant(Rapprich2007)correlatedwithlavadated 2) pyroclastic deposits of a local monogenic cone at Ne-
28.66±1.06Ma(Rapprich&Holub2008).Hence,wemay chraniceand3)lacustrinesedimentsonaplainatthefoot
supposetheageofthelaharsasLateRuppelian. ofthevolcanonearZvoníčkov.TheobtainedK-Aragesfor
Thelacustrinelimestone(travertine)atZvoníčkovcon- the monogenic cones on the DHVC northern periphery
tainingplatanoidwoodformsintercalationsinfine-grained are the youngest among the volcanic rocks of the entire
(cid:2)(cid:5)(cid:10)(cid:21)(cid:13)(cid:12)(cid:7)7+ A,B–Woodtype4–Cercidiphylloxylon(epitypeG8113ofCercidiphylloxylonkadanense).•A–diffuse-porouswoodwithmostlysolitary
vessels,angularinoutline.TS.•B–detailofalongscalariformperforationplatewithabout50bars,TLS.•C–Woodtype5–Ulmus(specimen75/04):
Outerpartofaring-porouswoodwithmainlysolitaryearlywoodvesselsandlatewoodvesselsinwavytangentialbands,TS.•D–G–Woodtype
6–?Craigia(D–F:specimen72/03;G:specimen89/04).•D–Woodsemi-ringporouswithfoldedearlywoodvessels,TS.•E–stronglyheterocellular
rayswithtilecells,TLS.•F–detailoftilecellsofPterospermumtype,RLS.•G–coalescentaperturesofalternateintervesselpitsmimickinghelical
thickening,TLS.•H–M–Woodtype7–Coryloxylon(H–J:holotypeCNB-4ofCoryloxylonnemejcii;K–M:holotypeCNB-3ofCoryloxylontertiarum).
•H,K–diffuse-porouswoodwithnumerousvesselsinradialmultiples,TS.•I,L–mostly3–4-seriatestronglyheterocellularrays,TLS.•J–two
scalariformperforationplateswithlessthan20barsinthemiddle,RLS.•M–scalariformperforationplatewithlessthan20barsfromasideview,TLS.
Scale bars = 500μm in A, C, D, H, K; 200μm in E, F, I, L; 100μm in B, G, J, M.
(cid:26)-(cid:26)