permian (〰ぁrtinskian to wuchapingian)〰〠conodont ...eprints.whiterose.ac.uk/108745/1/sun et...
TRANSCRIPT
This is a repository copy of Permian (Artinskian to Wuchapingian) conodont biostratigraphy in the Tieqiao section, Laibin area, South China.
White Rose Research Online URL for this paper:http://eprints.whiterose.ac.uk/108745/
Version: Accepted Version
Article:
Sun, YD, Liu, XT, Yan, JX et al. (7 more authors) (2017) Permian (Artinskian to Wuchapingian) conodont biostratigraphy in the Tieqiao section, Laibin area, South China. Palaeogeography, Palaeoclimatology, Palaeoecology, 465 (A). pp. 42-63. ISSN 0031-0182
https://doi.org/10.1016/j.palaeo.2016.10.013
© 2016, Elsevier. Licensed under the Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International http://creativecommons.org/licenses/by-nc-nd/4.0/
[email protected]://eprints.whiterose.ac.uk/
Reuse
Unless indicated otherwise, fulltext items are protected by copyright with all rights reserved. The copyright exception in section 29 of the Copyright, Designs and Patents Act 1988 allows the making of a single copy solely for the purpose of non-commercial research or private study within the limits of fair dealing. The publisher or other rights-holder may allow further reproduction and re-use of this version - refer to the White Rose Research Online record for this item. Where records identify the publisher as the copyright holder, users can verify any specific terms of use on the publisher’s website.
Takedown
If you consider content in White Rose Research Online to be in breach of UK law, please notify us by emailing [email protected] including the URL of the record and the reason for the withdrawal request.
1
Early-Middle Permian conodont biostratigraphy in the Tieqiao1
section,Laibinarea,SouthChina2
Y.D.Sun1,2*,X.T.Liu3,J.X.Yan1,B.Li4,B.Chen5,D.P.G.Bond6,M.M.Joachimski2,P.B.3
Wignall7,X.L.Lai1*4
1. Key Laboratory of Geobiology and Environmental Geology, China University of5
Geosciences,Wuhan430074,P.R.China6
2. GeoZentrum Nordbayern, Universität Erlangen-Nürnberg, Schlossgarten 5, 910547
Erlangen,Germany8
3. Key Laboratory of Marine Geology and Environment, Institute of Oceanology, Chinese9
AcademyofSciences,Qingdao266071,China10
4. Key Laboratory of Marine Mineral Resources, Guangzhou Marine Geological Survey,11
MinistryofLandandResources,Guangzhou510075,China12
5.StateKeyLaboratoryofPalaeobiologyandStratigraphy,NanjingInstituteofGeology13
andPalaeontology,39EastBeijingRoad,Nanjing210008,R.P.China14
6.DepartmentofGeography,EnvironmentandEarthSciences,UniversityofHull,HullHU615
7RX,UK16
7.SchoolofEarthandEnvironment,UniversityofLeeds,LeedsLS29JT,UK 17
*Correspondingauthor:E-mail:[email protected](Y.D.Sun);[email protected](X.L.18
Lai)19
20
2
Abstract21
Permian strata from the Tieqiao section (Jiangnan Basin, South China)22
contain several distinctive conodont assemblages. Early Permian (Cisuralian)23
assemblagesaredominatedbythegeneraSweetognathus,Pseudosweetognathus24
and Hindeodus with rare Neostreptognathodus and Gullodus. Gondolellids are25
absent until the end of the Kungurian stage—in contrast to many parts of the26
worldwheregondolellidsandNeostreptognathodusarethedominantKungurian27
conodonts.AconodontchangeoverisseenatTieqiaoandcoincidedwithariseof28
sealevel inthelatestKungurian:thepreviouslydominantsweetognathidswere29
replacedbymesogondolellids. TheMiddle andLatePermian (Guadalupian and30
Lopingian Series) witnessed a dominance of gondolellids (Jinogondolella and31
Clarkina),thecommonpresenceofHindeodusanddecimationofSweetognathus.32
Twenty one main and six subordinate conodont zones are recognized at33
Tieqiao, spanning the lowerArtinskian to lowermostWuchiapingianStage.The34
main (first occurrence) zones are, in ascending order by stage: the35
Sweetognathus(Sw.)whitei,Neostreptognathodus(N.)pequopensis-Sw.toriyamai,36
andSw.asymmetrican.sp.ZonesfortheArtinskian;theN.prayi,Sw.guizhouensis,37
Sw.iranicus,Sw.adjunctus,Sw.subsymmeticus,Sw.hanzhongensis,Mesogondolella38
(M.) idahoensis andM. lamberti Zones for theKungurian; the Jinogondolella (J.)39
nankingensis Zone for the Roadian; the J. aserrata Zone for the Wordian; the J.40
postserrata,J.shannoni,J.altudaensis,J.prexuanhanensis,J.xuanhanensis,J.granti41
and Clarkina (C.) hongshuiensis Zones for the Capitanian and the C. postbitteri42
3
ZoneforthebaseoftheWuchiapingian.Thesubordinate(interval)zonesarethe43
Pseudosweetognathuscostatus,Pseduosweetognathusmonocornus,Hindeodus(H.)44
gulloides, Pseudohindeodus ramovsi, Gullodus sicilianus, Gullodus duani and H.45
excavatesZones.46
Inaddition, threenewspecies,Gullodus tieqiaoensisn.sp.,Pseudohindeodus47
ellipticae n. sp. and Sweetognathus asymmetrica n. sp. are described. Age48
assignmentsforlesscommonspeciese.g.,Pseudosweetognathusmonocornusare49
reassessedbasedonarichconodontcollection.50
Key words: conodont, Permian, biostratigraphy, Cisuralian, Guadalupian,51
Kungurian,SouthChina52
53
4
1.Introduction54
ConodontsareimportantindexfossilsinthePalaeozoicandTriassic,dueto55
their high speciation rates, geographically widespread distribution and in part56
high abundance in marine sediments. Conodont biostratigraphy provides the57
best method for high-resolution, supra-regional correlations of Permian strata,58
becauseotherkey taxa suchas ammonoidsareoften scarce inmany locations,59
whilst foraminifers and brachiopods are generally long ranging and facies60
controlled and thus less useful for age diagnosis. As a consequence, Permian61
conodonttaxonomyandbiostratigraphyhavebeenthetopicsofextensivestudy62
since the 1950s (e.g., Youngquist et al., 1951; Clark andBehnken, 1971; Ritter,63
1986;WardlawandGrant,1990;Meietal.,1994a;Wardlaw,2000;Nestelletal.,64
2006;Lambert et al., 2010; Shenet al., 2012).The importanceof conodonts in65
stratigraphy is exemplified by their use at Global Boundary Stratotype Section66
and Points: as of 2016, conodonts define of the bases of all but one of the 2967
stagesbetweenthePragian(LowerDevonian)andRhaetian(UpperTriassic),1768
ofwhichhavebeenratifiedbytheInternationalCommissiononStratigraphy.69
Thediversity of Permian conodonts is generally low in comparison to that70
observed for other time periods, with typically less than five genera and two71
dozensofspeciesoccurringinanygivenPermianstage.Conodontzonesarealso72
relativelylongforsomeintervals.Forinstance,thoughsubstantialinvestigations73
have been carried out in West Texas (e.g., Wardlaw, 2000; Nestell et al., 2006;74
Nestell and Wardlaw, 2010a; Wardlaw and Nestell, 2010), only one standard75
5
conodont zone has been established for the Roadian and Wordian stages76
(Henderson et al., 2012). This reflects a true low point in the diversity of77
conodontsduringtheir longevolutionaryhistory.Afurthercomplicationisthat78
minor changes in Permian conodont morphology require a careful taxonomic79
examination of different species.New species are rarely reported from regions80
otherthanWestTexasandSouthChina,perhapsowingtoadecreaseinresearch81
effortandasubstantiallossofexpertiseinrecentyears.82
Establishinga robustbiostratigraphic scheme indifferentareas isessential83
forsupra-regionalcorrelation.Permianconodontshavebeenbeststudiedinthe84
UralsofRussia (EarlyPermian),WestTexas (MiddlePermian)andSouthChina85
(Late Permian and the Permo-Triassic boundary) (e.g., Chuvashov et al., 1990;86
Meietal.,1994b;Zhangetal.,1995;Wardlaw,2000;Lambertetal.,2002;Jianget87
al., 2007; Nestell and Wardlaw, 2010b). The Early to Middle Permian of South88
China has attracted comparatively little research attention and is less89
systematicallystudied.90
This study presents a high-resolution conodont record for the Tieqiao91
section,Guangxi,SouthChina.Newdata,spanningtheArtinskian(EarlyPermian)92
to the early Wuchiapingian (Late Permian), substantially improve existing93
records of the section, first described two decades age in the context of the94
Capitanian-Wuchiapingian(Guadalupian-Lopingian)transition(Meietal.,1994c;95
Wang,2002).96
6
2.GeologicalSetting97
TheYangtzeregionwasalargeisolatedplatformsituatedwithinthePermian98
equatorial Tethys (Fig. 1) with extensive carbonate deposition and diverse99
sedimentaryfacies.Itisanideallocationforconodontstudies.TheLaibinareais100
located in the JiangnanBasin towards the southwesternmargin of theYangtze101
Platform(WangandJin,2000).Aseriesofsuperbsectionsareexposedalongthe102
banks of Hongshui River (Shen et al., 2007) and these have been103
comprehensively studied for the Capitanian-Wuchiapingian transition (e.g.,Mei104
etal.,1994c;Wangetal.,2004;Jinetal.,2006;Wignalletal.,2009b;Chenetal.,105
2011). 106
The Permian strata of the region consist of thick Early Permian platform107
carbonates,subordinateMiddlePermianslopeaswellasbasinalcarbonatesand108
cherts. Late Permian rocks are geographically more variable, including coal109
seams,reefbuild-upsandradiolariancherts(Shaetal.,1990;Shenetal.,2007;110
Qiuetal.,2013).111
ThestudiedsectionatTieqiao(23°42.733'N,109°13.533'E)isexposedon112
thenorthernbankoftheHongshuiRiver,southeastofthetownofLaibin(Figs.1,113
2).ThePermianstratameasure1307mthickandcomprisetheMaping,Chihsia,114
Maokou, Heshan, Wuchiaping and Talung Formations, spanning the earliest115
Permian (Asselian) to the Permian-Triassic boundary (Sha et al., 1990). The116
sectionisveryfossiliferouswithforaminifers,calcareousalgae,crinoids,sponges117
7
andcoralsbeingprolificallyabundant(e.g.,WangandSugiyama,2000;Bucuret118
al., 2009; Zhang et al., 2015), whilst bivalves and ammoinods occur less119
frequently.Well-preservedZoophycostracefossilsarealsoabundant(Gongetal.,120
2010).121
Sha and colleagues (1990) pioneered the study of the Tieqiao section and122
subdividedthesectioninto15Membersand139Beds.Ourstudyfollowsthese123
subdivisions (Figs. 3-5) for consistency and focuses on the stratigraphy and124
conodont zonation of the Chihsia and Maokou formations (Bed 1 to Bed 109).125
TheChihsaFm.generallyrecordsdepositioninacarbonaterampsetting,whilst126
theMaokouFm.comprisesslope tobasin transition facies.The two formations127
are overall 710 m thick and range from the Sakmarian (?) to the128
Capitanian-Wuchiapingianboundary(Figs.3-5).129
3.MaterialsandMethods130
The section was sampled over four field campaigns between 2005-2010.131
During the springof2010, thewaterof theHongshuiheRiver fell to its lowest132
levelofthepasttenyearsduetoaseveredrought,whichallowedustodescribe133
and sample several normally submerged parts of the section (e.g., Bed 17 and134
Bed112).Atotalof374rocksampleswerecollectedwithasamplingresolution135
of~1-2mformostpartsofthesection.Chertsandgrainstonesbearingabundant136
corals and fusulinid foraminifers were avoided during sampling due to137
complications in conodont extraction and low conodont yields. Each sample138
8
weighedbetween2.5and8kg.139
Threehundredandelevensampleswereprocessedinthemicropaleontology140
laboratory at China University of Geosciences (Wuhan) and 63 samples were141
processed at the GeoZentrum Nordbayern, Universität Erlangen-Nürnberg. All142
samplesweredissolvedusing7-10%dilutedaceticacid,wetsievedandair-dried.143
Theinsolubleresidueswereseparatedbyusingheavyliquidfractionation(Jiang144
etal.,2007).Conodontspecimenswerehandpickedusingabinocular.Conodonts145
fromTieqiaoaregenerallywellpreservedwithcolouralternationindexranging146
from1.5to2.5.Atotalof8733specimenswereobtainedfromsampleprocessat147
Wuhanlabandabout~3000specimenswasrecoveredatErlangen.Resultsfrom148
bothlaboratorieswerecrosschecked.149
4.StratigraphyandConodontZonation150
4.1Sakmarian(?)151
The lowermost part of the studied section (Beds 1 to 16, Chihsia Fm.)152
consists mainly of thin-to-medium bedded dark-grey bioclastic micrites, marls153
and black shales (Fig. 2). Brachiopods, gastropods, crinoids, bryozoans and154
spongesarethemostabundantfossils.Ageassignmentforthispartofthesection155
is controversial. Sha et al. (1990) suggested anAsselian age for theunderlying156
MapingFm.andreportedtheoccurrenceofthefusulinaceanEoparafusulinasp.,157
Nankinellasp.,Pamirinasp.,Staffellasp.andPseudofusulinasp.fromBed1toBed158
16,implyingapossibleSakmarianagefortheunit.Meietal.(1998)inferredthis159
9
unittobeof“Longlinian”age—aChineseequivalentoftheArtinskianbyoriginal160
definition (Sheng and Jin, 1994; Jin et al., 1997), now re-defined as Sakmarian161
(Fig. 24.1 in Henderson et al., 2012). Based on consideration of all published162
fossilmaterials,Shenetal.(2007)tentativelyassignthispartofthesectiontothe163
Artinskian.164
Few conodonts were recovered from this part of the section, despite great165
efforts. Many ramiform elements were recovered from Bed 8, but none is166
age-diagnostic.Thepreciseageofthisunitthereforeremainsunresolved.167
4.2Artinskian168
Beds 17 to 26 consist mainly of dark-grey to grey bioclastic pack- and169
grainstones. The conodont assemblage is dominated by Sweetognathus whitei170
andaffinitivespeciesandthusindicatesanArtinskianage.Neostreptognathodus171
andHindeodusarerarewhilstgondolellidsareabsent.ThebaseoftheArtinskian172
stage cannotbepreciselydefinedbecause the first appearancedatum(FAD)of173
Sweetognathus whitei cannot be ascertained due to the inaccessibility of the174
submerged lowerpartofBed17andtheabsenceofdiagnosticconodonts from175
bedsbelowthislevel.Inascendingorder,threeconodontzoneswereestablished176
fortheArtinskian. 177
1).SweetognathuswhiteiZone178
Thelowerlimitofthiszoneisnotdefined.Theupperlimitisdefinedbythe179
FAD of Sweetognathus (Sw.) toriyamai and Neostreptognathodus ex gr.180
10
pequopensis.181
Sw. whitei was one of the most cosmopolitan conodonts during the Early182
Permian (Meietal.,2002). It isknowne.g. fromNorthandSouthChina, Japan,183
U.S.A., Canada and Colombia. (Rhodes, 1963; Igo, 1981; Orchard, 1984; Ritter,184
1986;DingandWan,1990; Jietal.,2004)and isconsideredagoodmarker for185
thebaseofArtinskian. 186
2).Neostreptognathodusexgr.pequopensis-SweetognathustoriyamaiZone187
Lower limit: FAD of Sw. toriyamai and Neostreptognathodus (N.) ex gr.188
pequopensiswithbothtaxafirstoccurringatthesamelevel.Upperlimit:FADof189
Sw. asymmetrica n. sp. The FAD of Sw. bogoslovskajae occurs in this zone. Sw.190
bogoslovskajae is known to co-exist with N. pequopensis in Nevada and has a191
range restricted to the upper “Baigendzhinian” (equivalent to uppermost192
Artinskian to lower Kungurian) (Ritter, 1986). Wang (2002) reported the193
occurrenceofSw. variabilis in this zone (inBed18ofTieqiao).Wehave found194
morphotypes which are similar to Sw. variabilis but the specimens are not195
sufficientlywell-preservedtomakeanidentification. 196
3).Sw.asymmetrican.sp.Zone197
Lower limit: FAD of Sw. asymmetrica n. sp. Upper limit: FAD of198
Neostreptognathodusprayi.HindeoduscatalanoiandSw.cf.windico-occurinthis199
zone. This zone probably straddles the Artinskian-Kungurian boundary due to200
theabsenceoftheN.pnevizoneatTieqiao.201
11
4.3Kungurian202
Kungurianrocks,spanningfromBed27tothelowerpartofBed109,consist203
mainly of medium-to-thick bedded fossiliferous pack- and grainstones with204
commonchertnodulesinthelowerpart.Mediumtothinbeddedlimemudstones205
andwackestonesweregraduallydevelopedhigherintheKungurianstrata,witha206
notable shift in fossil assemblages from a bryozoan- and calcareous207
algae-dominated shallow water facies (Beds 89-99) to a sponge spicule and208
radiolarianrichdeeperwaterfacies(Beds100-111). 209
TheconodontbiostratigraphyofthebasalKungurianstagehasbeenamatter210
of debate (Wang et al., 2011). Kozur et al. (1995) suggested the cline211
Neostreptognathoduspequopensis-N.pnevi tobesuitable foradefinition for the212
Artinskian-Kungurian boundary. Mei et al. (2002) proposed the FAD of N.213
pequopensis or Sw. guizhouensis to define the base of the Kungurian whereas214
Chuvashovetal. (2002) formallyproposed theFADofN.pnevi asdiagnosticof215
the base of the Kungurian, a definition that has been generally accepted216
(Hendersonetal.,2012).However,duetotheabsenceofN.pneviatTieqiao,we217
alternativelysuggestthebaseoftheKungurianbedefinedbytheFADofN.prayi.218
EightconodontzoneswereestablishedfortheKungurian,describedinascending219
orderbelow:220
4).NeostreptognathodusprayiZone221
Lower limit: FAD of Neostreptognathodus prayi. Upper limit: FAD of Sw.222
12
guizhouensis.TheN.prayiZoneisthesecondoldestzoneoftheKungurianinthe223
standardPermianconodontzonation(Hendersonetal.,2012)andsotheN.prayi224
zoneatTieqiaodoesnotindicatethe“true”earliestKungurian(Fig.6).Sw.clarki,225
aspeciesmostcommonlyseeninthelateArtinskian(BeauchampandHenderson,226
1994),alsooccursinthiszone.Thefirstoccurrence(FO)ofSw.clarkiatTieqiao227
isonlyslightlylowerthanthatofSw.guizhouensis.228
5)Sw.guizhouensisZone229
Lowerlimit:FADofSw.guizhouensis.Upperlimit:FADofSw.iranicus.Except230
for in the lower part of this ~180 m thick conodont zone, conodonts are231
relatively rare. The long-ranging species Pseduosw. costatus is the only species232
thatwassparselyrecoveredintheupperpartofthiszone. 233
6)Sw.iranicusZone234
Lowerlimit:FADofSw.iranicus.Upperlimit:FADofSw.subsymmetricus.As235
withtheSw.guizhouensisZone,bothconodontdiversityandabundancearevery236
low.Amajor stratigraphic complication at this level of the section is thatBeds237
76-88areatectonicrepetitionofolderbeds(alsoseeShaetal.,1990). 238
7)Sw.adjunctusZone239
Lowerlimit:FADofSw.adjunctus.Upperlimit:FADofSw.subsymmetrics.Sw.240
cf. paraguizhouensis appears in this zone. Sha et al. (1990) reported the241
occurrenceof“Neogondolella”bisselliinthiszone(Bed91).However,“N”.bisselli242
isanolderspecieswhichoftenco-occurredwiththeArtinskianSw.whiteigroup243
13
(e.g., Behnken, 1975; Clark et al., 1979;Orchard, 1984;Wang, 1994;Mei et al.,244
2002). The occurrence of bisselli obviously contradicts a Kungurian age of the245
hoststrataandalsoisnotconfirmedbyourdataset.246
Sw. adjunctus is also known from the uppermost Victorio Peak Formation247
fromTexasandtheupperPequopFormationfromNevada,USA(Behnken,1975)248
as well as from south-central British Columbia, Canada (Orchard and Forster,249
1988): All of these occurrences are dated to be of late Leonardian age in the250
Permianregionalstratigraphy(=middletolateKungurian).Becauseofthewide251
distribution of Sw. adjunctus, this zone therefore has high potential for252
super-regionalcorrelation.253
8)Sw.subsymmetricusZone254
Lower limit: FAD of Sw. subsymmetricus. Upper limit: FAD of Sw.255
hanzhongensis. This zone correlates to the Kungurian “M. siciliensis-Sw.256
subsymmetricus”zoneinsouthernGuizhou(Meietal.,2002).257
Sw. subsymmetricus is well known from the Kungurian of Guizhou and258
GuangxiinSouthChina,aswellasfromThailandandOman(Meietal.,2002and259
this study; Henderson and Mei, 2003; Metcalfe and Sone, 2008; Burrett et al.,260
2015).TheassertionthatSw.subsymmetricusisrestrictedtotheRoadian(Kozur,261
1993)isincorrect.262
9)Sw.hanzhongensisZone263
Lowerlimit:FADofSw.hanzhongensis.Upperlimit:FADofM.idahoensis.The264
14
FAD of Pseudohindeodus augustus and Pseudohindeodus ramovsi occurs in the265
middle part of this zone. A turnover in the dominant conodont fauna initiated266
during this zone. Hindeodus becomes abundant whilst the abundance and the267
diversityofSweetognathusdecreases.Hindeoduspermicus,H.gulloidesandH.aff.268
wordensisalloccurinthiszone. 269
10)MesogondolellaidahoensisZone270
Lowerlimit:FADofM.idahoensis.Upperlimit:FADofM.lamberti.Conodont271
faunaschange fromHindeodus-Pseduohindeodus-Sweetognathus-dominatedand272
gondolellid-freeassemblagestogondolellid-dominatedassemblagesinthiszone273
(Bed109).Thisshiftinconodontassemblagecoincideswithalithologicalchange274
fromthick-andmedium-beddedwackestonestomorecherty,medium-tothin-275
beddedwacke-andcarbonatemudstones.276
11)M.lambertiZone277
Lower limit: FAD of M. lamberti. Upper limit: FAD of J. nankingensis.278
CarbonateoftheM.lambertiZone(Beds110-111)aremorethinlybeddedwith279
an increasing abundance of sponge spicules and radiolarian tests, indicating280
deepeningandarelativesealevelriseinthelatestKungurian.281
4.4Roadian282
TheRoadianstrataconsistoflessthan5mthickfinelylaminatedandthinly283
bedded carbonatemudstones starting at the topofBed111 (Fig. 2B).Amajor284
sea-levelriseisknownfromtheEarly-MiddlePermiantransitionandismanifest285
15
at Tieqiao by a transition to thinly bedded radiolarian cherts by Bed 112286
(Wordianage).Bythistime,deep,basinalsedimentationwasestablishedinthe287
region. The minor thickness of the Roadian strata may be attributed either to288
condensationduring this sea level rise or tohiatus resulting in a lossof strata289
(duetosuddenlossofcarbonateproductionbelowthecarbonatecompensation290
depth).Onlyoneconodontzoneisrecognized.291
12)JinogondolellanankingensisZone292
Lower limit: FAD of J. nankingensis. Upper limit: FAD of J. aserrata.293
Pseudohindeodusramovsiareabundant.Fine lamination intheupper(Roadian)294
partofBed111indicatesminorbioturbation.295
4.5Wordian296
WordianstrataarepresentedbyBed112tolowermostpartofBed116.The297
sediments consist of thinly bedded radiolarian cherts in the lower part (Beds298
112-113), thickly bedded bioclastic wacke- and packstones in the middle (Bed299
114, also known as “the Great White Bed”) and alternation of cherts and lime300
mudstonesintheupperpart(Bed115-116).Oneconodontzoneisrecognized.301
13)J.aserrataZone302
Lowerlimit:FADofJ.aserrata.Upperlimit:FADofJ.postserrata.TheFADofJ.303
palmata occurs at the same level as the FAD of J. aserrata. This is generally304
consistent with the record in west Texas where the FAD of J. palmata was305
reported very close to the FAD of J. aserrata (Nestell and Wardlaw, 2010a).306
16
Severalspecies,suchasJ.errata,GullodusduaniandthelongrangingspeciesSw.307
hanzhongensis and Pseudohin. ramovsi appear in the middle-upper part of this308
zone. 309
4.6Capitanian310
TheCapitanian(Beds116-119)isthemostintensivelystudiedintervalinthe311
Laibin area (Mei et al., 1994c; Jin et al., 2006; Chen et al., 2009;Wignall et al.,312
2009b).Strataofthisageconsistofmediumbeddedalternatingchertsandlime313
mudstones in the lower part (Beds 116-118) overlain by pack- to grainstones314
(LaibinLimestoneMember,Bed119).Hereweonlygiveabriefdescriptionofthe315
conodontzonesofthisstagesincetheyhavebeenwellstudied.316
14)J.postserrataZone317
Lowerlimit:FADofJ.postserrata.Upperlimit:FADofJ.shannoni. 318
15)J.shannoniZone319
Lowerlimit:FADofJ.shannoni.Upperlimit:FADofJ.altudaensis.320
16)J.altudaensisZone321
Lowerlimit:FADofJ.altudaensis.Upperlimit:FADofJ.prexuanhanensis.This322
interval is characterized by higher extinction rates as well as the onset of323
Emeishanvolcanism(Wignalletal.,2009a;Sunetal.,2010).Lossesincludemany324
foraminifers,calcareousalgae in theequatorialrealmandmanybrachiopods in325
the boreal realm (Bond et al., 2010; Bond et al., 2015). Though there are no326
17
obvious lithological changes in the J. altudaensis Zone at Tieqiao, the last327
appearances of several long-ranging conodonts, such as Gullodus duani, Sw.328
hanzhongensisandPseudohind.ramovsi,areallrecordedinthiszone.329
17)J.prexuanhanensisZone330
Lower limit: FAD of J. prexuanhanensis. Upper limit: the FAD of J.331
xuanhanensis.ThiszonehasnotbeenrecognizedinwesternTexas(Lambertetal.,332
2002).However,itisalsodistinguishableintheTieqiao(Guangxi,thisstudy)and333
atDukou(Sichuan,Meietal.,1994a)sections.IncondensedsectionsinGuizhou,334
the J.prexuanhanensis zone isoftencombinedwith theyounger J. xuanhanensis335
zoneastheJ.prexuanhanensis-J.xuanhanensisassemblagezone(Sunetal.,2010).336
Sw. fengshanensis occurs in this zone. Sw. fengshanensis was established in337
the lateCapitanianstrataatFengshan,northwesternGuangxi(Meietal.,1998).338
At thePenglaitansection,Sw. fengshanensis spans fromtheupper J.postserrata339
zonetothelowerJ.xuanhanensiszone,representingthelastintheevolutionary340
lineageofsweetognathidsinSouthChina(Meietal.,2002).341
18)J.xuanhanensisZone342
Lower limit: FAD of J. xuanhanensis. Upper limit: FAD of J. granti. Many343
maturemorphotypesofJ.shannonioccurinthelowermostpartofthiszoneand344
are very similar to their counterparts from West Texas (Lambert et al., 2002;345
WardlawandNestell,2010).Volcaniclasticmaterialstartstobedepositedduring346
thiszoneandbecomesmorecommonintheoverlyingJ.grantiZone,presumably347
18
corresponding to the large scale of explosive eruptions of the Emeishan Traps348
(e.g.,Sunetal.,2010).349
19)J.grantiZone350
Lower limit: FAD of J. granti. Upper limit: FAD of Clarkina hongshuiensis.351
Conodontsareprolificallyabundantinthiszonewithatypicalyieldrateof~100352
specimensperkgrock.353
20)ClarkinahongshuiensisZone354
Lowerlimit:FADofC.hongshuiensis.Upperlimit:FADofC.postbitteri.355
4.7Wuchiapingian356
The early Wuchiapingian (Bed 120) is characterized by deposition of357
extensivebeddedchertswithintercalatedpinkishlimestonelenses.Evidencefor358
a relative sea level fall towards the end of Wuchiapingian is indicated by a359
reductionofchertthicknessup-sectionwithcarbonatesedimentationincreasing.360
Eventually,thisbasinalsettingevolvedintoaspongereeffacies(Beds124-132)361
inthemiddle-lateWuchiapingianlackingconodonts.Clarkinatranscaucasicawas362
found in Bed 134. One conodont zone is established for the earliest363
WuchiapingianatTieqiao:364
21)ClarkinapostibitteriZone365
Lowerlimit:FADofC.postbitteri.Upperlimit:notdetermined.366
5.Subordinatezonesandreassessmentforageassignmentsofrarespecies367
19
Seven subordinate zones are established at Tieqiao, representing interval368
zonesbasedonoccurrencesof longrangingspecies.Thesubordinatezonesare369
less effective for stratigraphic correlation but can provide a valuable reference370
forcaseswhenasingleconodontassemblageisobtainedfromanage-ambiguous371
lithologic unit (e.g., Burrett et al., 2015). In the following section, we first372
describe therangesof thesesubordinatezones in theTieqiaosection, followed373
by comments on the range of the zonal species. A correlation with the main374
conodontzones isshown in figure.6.Note that therangeof thespeciescanbe375
muchlongerthantherespectivezone. 376
1)Pseudosweetognathus(Pseudosw.)costatusIntervalZone 377
Lower limit: FAD of Pseudosw. costatus. Upper limit: FAD of Pseudosw.378
monocornus. ThePseudosw. costatus Zone spans the earlyArtinskian tomiddle379
Kungurian(Bed19toBed94).Elementsof long-rangingspeciesH.minutusare380
abundantinthelowerpartofthiszoneandthereisasingleoccurrenceofH.aff.381
catalanoiinthelowermost.382
Pseudosweetognathuscostatuswasestablished inArtinskianstrataofSouth383
China (Wang et al., 1987) and also reported from Thailand, co-existing with a384
typicalKunguriantaxonSw.subsymmetricus(MetcalfeandSone,2008).Ourdata385
confirm former observations and indicate that the range of Pseudosw. costatus386
extends from the Artinskian Sw. asymmetrica n. sp. Zone to the Kungurian Sw.387
adjunctus Zone. In the middle Kungurian, Pseudosw. costatus was replaced by388
20
Pseudosw.monocornus.389
2)PseudosweetognathusmonocornusIntervalZone390
Lowerlimit:FADofPseudosw.monocornus.Upper limit:FADofH.gulloides.391
ThiszonecomprisesBed94toBed102atTieqiao,andisoflateKungurianage. 392
Li et al. (1989) established the species Pseudosweetognathus monocornus393
(under the genus “Sichuanognathodus”) from the upper part of Maokou Fm. at394
Shangsi.AlateranddetailedstudyofthesamesectionreportedaJinogondolella395
andHindeodusdominatedfaunawhichindicatesanearlyCapitanianageforthe396
upperMaokouFm.(Sunetal.,2008). 397
Pseudosweetognathusmonocornus is found in theupperpartofChihsiaFm.398
and lower part of Maokou Fm. at Tieqiao and here is reassigned a399
middle-Kunguriantoearly-Roadianage.Thisspeciesonlyoccurredwithshallow400
water, high energy assemblage composed of calcareous algae, corals and401
foraminifers found in thickly bedded bioclastic pack- and grainstones.We thus402
speculate that theoccurrenceofPseudosw.monocornusmightbe facies-related,403
anditspresenceinWordiantolowerCapitanianstrataelsewhere(Lietal.,1989)404
cannotbeexcluded.405
3)HindeodusgulloidesIntervalZone406
Lowerlimit:FADofH.gulloides.Upperlimit:FADofPseudohindeodusramovsi.407
This zone occupies Bed 102 and correlates to the middle part of Sw.408
hanzhongensiszone,representingaLateKungurianage.409
21
The species H. gulloides Kozur and Mostler, 1995, ranges from upper410
Kungurian to Roadian. In northeast Thailand, H. gulloides occurs at an411
age-equivalent level as in South China and co-existed with a typical late412
Kungurian assemblage which consists of species Mesogondolella siciliensis,413
Pseudohindeodus oertlii (= angustus) and Sw. subsymmetricus (Burrett et al.,414
2015). In west Texas, the species was recovered from the upper part of Road415
CanyonFm.,representingalateRoadianage(KozurandMostler,1995).416
4)PseudohindeodusramovsiIntervalZone 417
Lower limit: the FAD of Pseudohindeodus ramovsi. Upper limit: the418
occurrence of Gullodus sicilianus. This zone spans from Bed 103 to Bed 115,419
representingalatestKunguriantoWordianage.420
The species Pseudohindeodus ramovsi Gullo and Kozur, 1992 has a much421
longer range than the Interval Zone. Wardlaw (2000) reported sporadic422
occurrences of this species from the Kungurian to Capitanian. Our data are423
consistent with Wardlaw (2000), suggesting that Pseudohindeodus ramovsi424
spanned fromthe lateKungurianSw.hanzhongensis to themiddleCapitanian J.425
altudaensisZone.426
AnotherassociatespeciesinthiszoneisPseudosw.augustus(Igo,1981).This427
specieshasbeenreportedfromcoevalKungurianstratainJapan(Igo,1981;Shen428
et al., 2012), but can also occur inmucholder strata such as in theArtinskian429
(OrchardandForster,1988).430
22
5)GullodussicilianusIntervalZone 431
Lowerlimit:FOofG.sicilianus.Upperlimit:FADofGullodusduani.Thiszone432
coversthemiddlepartofBed115,representingamiddle-lateWordianage.433
Gullodus sicilianus (Bender and Stoppel, 1965) ranges from the Roadian to434
Wordian(Kozur,1993). It isa rare taxon that isknownmostly fromtheTethys435
realmduringtheWordian(Kozur,1995).436
6)GullodusduaniIntervalZone437
Lower limit: FAD of Gullodus duani. Upper limit: prolific occurrence of H.438
excavatus. This zone comprises Bed 115 to Bed 118, covering much of the439
Capitanian. An associated taxon Hindeodus catalanoi ranges through the upper440
partofthiszone.ThoughGulloandKozur(1992)assignedaWordianageforH.441
catalanoi, this form is found in the Capitanian at Tieqiao, suggesting a longer442
rangeofthespeciesthanitsoriginaldefinition.443
Gullodus duani Mei et al. 2002 is a rather rare species in the Guadalupian.444
ThisspecieswasoriginallyrecoveredfromtheMaokouFm.fromGuangxiandis445
onlyknownfromSouthChina.AtTieqiao,thisspeciesisknownfromuppermost446
WordiantomiddleCapitanianstrata.447
7)H.excavatusIntervalZone448
Lower limit: the prolific occurrence of H. excavatus. Upper limit: FAD of C.449
postbitteri (the Capitanian-Wuchiapingian boundary). At Tieqiao, this zone is450
23
representedbytheLaibinLimestoneMember(Bed119)ofalateCapitanianage.451
Hindeodusexcavatus isanother long-rangingspecies in thePermian,but its452
useasazonalfossilderivesfromitsprolificabundanceinthelateCapitanian.453
6.Systematicpalaeontology454
GenusGullodusKozur,1993455
Emended diagnosis: Spathognathodiform elements with a medium to long456
anterior blade and a posteriorly positioned, strongly expanded basal cavity.457
Denticles occur on the blade and above the basal cavity and are inmost cases458
without ornamentations. Denticles are generally 10-15 in number and those459
above the basal cavity can be expanded and form a carina-like structure or460
narrowtransverseridges.Smallcoalesceddenticlesaresometimesdevelopedon461
theanterioredgeformingan“anteriorblade”.Length/heightratioisbetween1.5462
and3.Basal cavities aregreatly expanded,non-ornamentedandoccupy1/3 to463
2/3ofthefullbody.464
Remarks:thediagnosisofthisgenus(Kozur,1993)shouldbeemendedbecauseit465
is often hard to differentiate between Gullodus and Hindeodus. The emended466
diagnosis also includes wider variability of Gullodus species. Key differences467
betweenGullodus andHindeodus are theshapeandpositionof thebasal cavity468
and the length/height ratio: Hindeodus has a more centrally positioned basal469
cavity and lower length/height ratio. A key difference between Gullodus and470
Sweetognathus is that denticles of Gullodus are not ornamented while those of 471
24
Sweetognathus develop nodes. Gullodus can be differentiated from472
Pseudohindeodus because the basal cavity of the latter is more (horizontally)473
expanded and ornamented with a surface apron and occupies!2/3 of the full474
elementlength.475
Based on the revised diagnosis, Gnathodus sicilianus Bender and Stoppel,476
1965 should remain as Gullodus sicilianus as suggested by Kozur (1993).477
However, Pseudohindeodus catalanoi Gullo and Kozur (1992) and Gullodus478
hemicircularis Kozur, 1993 should belong to Hindeodus, rather than479
PseudohindeodusorGullodus.480
Occurrence:KunguriantoCapitanian481
482
Gullodustieqiaoensisn.sp.SunandLai483
Plate4,figs.6,7.484
Noreportedspecimensaresimilartothisspecies.485
Etymology:Fromthenameofthesectionfromwherethespeciesisdescribed.486
Holotype:SpecimenS1_060 (Pl.4,fig.6)fromsample41-1ofBed41,ChihsiaFm.,487
TieqiaoSection,SouthChina.488
Paratype:SpecimenS1_062 (Pl.4,fig.7)fromsample41-2ofBed41,ChihsiaFm.,489
TieqiaoSection,SouthChina.490
Diagnosis:AGullodusspecieswithahighlength/heightratioandarobustcusp.491
25
Description:Bodyslimandelongated.Length/heightratiois~2.Thecuspistall,492
wideandrobust,normallytwiceashighasthedenticlesandthreetimeswider493
thanthedenticles.13-17denselyarrayeddenticlesdeceaseinheightposteriorly.494
Posteriordenticleson thebasalcavityaremoreexpandedandthuswider than495
therest.Theycanbelowerandmorefused.Thebasalcavityisexpanded,leafor496
irregularshapedandoccupiestheposterior2/3oftheelement.Thewidestpoint497
isintheposterior1/4to1/3.498
Remarks: This species has a very high length/height ratio and a posteriorly499
positioned, expanded but non-ornamented basal cavity that extends to the500
posterior end. It thus belongs to Gullodus rather than Hindeodus or501
Pseudohindeodus.502
Occurrence:lowerChihsiaFm.(earlyKungurian),Tieqiao,SouthChina503
504
GenusHindeodusRexroad&Furnish,1964505
HindeoduscatalanoiGulloandKozur,1992506
Plate7,Figs.6-8.507
PseudohindeoduscatalanoiGulloandKozur,1992p.225,pl.5,fig.A.508
HindeodusgulloidesBurrettetal.2015,p.111-113,Fig.6,figs.J-I.509
Diagnosis: A Hindeodus species that is triangular shaped with 2 to 3 anterior510
coalesceddenticlesand12-15denselyarrayeddenticles.511
26
Remarks: The species resembles its Artinskian-Kungurian and “Roadian”512
predecessors H. hemicircularis Kozur 1993 and H. gulloides Kozur andMostler,513
1995.Theyallhavetwotothreeanteriordenticles.However,H.hemicircularisis514
sub-semicircularshapedandhasfewerbutwiderdenticleswhileH.gulloides is515
moreelongatedandhasamuchbroadercuspthanthecurrentspecies. 516
Occurrence: upper Maokou Fm. (middle-late Capitanian), Tieqiao, South China;517
WordianofSicily.518
Hindeodussp.ASunandLai519
Plate4,Figs.23,26.520
Diagnosis: A Hindeodus species whose outline is close to that of an isosceles521
triangle.522
Description:Bodytriangularshapedwitha longanterioredge.Anteriorangle is523
around 45炣-60炣. Two or three small coalesced denticles may develop on the524
anterior edge. Medium sized cusp followed by three low denticles. Posterior525
denticlesaretallerandwideranddecreaseinheighttowardstheposteriorend.526
Thebasalcavityismediallyexpandedandcentralpositioned.527
Remarks:ThespeciesresemblesH.permicusbutdiffersbyitsoutlineandshapes528
ofdenticles.529
Occurrence:upperKungurian,basalMaokouFm.ofSouthChina530
GenusPseudohindeodusGulloandKozur,1992531
27
Pseudohindeodusellipticaen.sp.SunandLai532
Plate4,fig.13;Plate7.fig.14.533
Pseudohindeodussp.Wang,1995,pl.1,figs.1a,1b.534
Etymology:Fromtheovalshapeofthebasalcavityofthespecies.535
Holotype:SpecimenS7_001 (Pl.7,fig.14)fromsample104-2ofBed104,Maokou536
Fm.,TieqiaoSection,SouthChina.537
Paratype:SpecimenS2_075 (Pl.4,fig.13)fromsample104-2ofBed104,Maokou538
Fm.,TieqiaoSection,SouthChina.539
Diagnosis:AGullodusspecieswithanasymmetricalbasalcavitythatisnearoval540
inshape.541
Description:Elementissmallandrounded.Cuspis large,robustandhigherand542
broaderthananyfollowingdenticles.The5-8denticles immediatelybehindthe543
cusparethinandmorefusedwitheachotherandthuscanappearasaridge.The544
last 4-6 denticles are the largest amongst all denticles. They are lower, more545
roundedinshapeandrelativelyevenlyspacedwitheachotherwithasmallgap546
inbetween.Thebasalcavityisdecoratedwithanapron,horizontallyexpanded,547
asymmetricalandveryrounded.Theoutlineofthebasalcavityisclosetoanoval.548
Remarks:thespeciesresemblesPseudohindeodusramovsi.However,ramovsihas549
aneartriangularbasalcavitywhilstPseudohindeodusellipticaen.sp.hasamore550
roundedbasalcavity.551
28
Occurrence:upperKunguriantolowermostRoadian,basalKufengFm.andlower552
MaokouFm.ofSouthChina.553
554
GenusSweetognathusClark,1972555
Sweetognathusasymmetrican.sp.SunandLai556
Plate1,Figs.1,7,14,17.557
Etymology:Thespeciesnamereferstoitsasymmetricanteriortransverseridges.558
Holotype:SpecimenS1_018(Pl.1,Fig.1)fromsample18-1ofBed18,ChihsiaFm.,559
TieqiaoSection,SouthChina.560
Paratypes:Specimen S1_037 (Pl. 1, Fig.7) from sample 22-2 of Bed22, Chihsia561
Fm.,TieqiaoSection,SouthChina.562
Diagnosis:AType III sweetognathid (definition followsRitter,1986)withshort563
bladeandasymmetricanteriortransverseridges.564
Description:Shortblade,oftenbearing4-6denticles;thecuspismoderatelybig,565
the firstdenticle is thebiggestandveryoften fusedwith thecuspand formsa566
high robust cusp; the other denticles are much smaller, lower and more fused567
toward to the carina. The first two denticles are occasionally both very high,568
robustand triangular inshape.Transverseridgesareclearly incised.Thereare569
commonly 6 to 8 transverse ridges. The first one or two ridges are always570
asymmetricallydeveloped—inmostcasestheleftnodesaremissing.Thewidest571
partof thecarina is inthemiddle.Thebasalcavity is leaf- toheart-shapedand572
29
moderatelyexpanded,occupyingtheposteriorhalfofthefullelementlength. 573
Remarks:This species is similar to Sw. subsymmetricus.Both speciesdeveloped574
asymmetricanteriortransverseridges.However,thecurrentspeciesdiffersfrom575
Sw. subsymmetricus by: 1) the length ratio of free blade/carina < 1 (most576
commonly1/3),whereasthatofSw.subsymmetricusisgenerally≥1;2)thecuspis577
large tall and robust, whereas that of Sw. subsymmetricus is moderately large,578
comparedwithotherdenticlesontheblade;3)anapparentgapbetweenblade579
and carina; Sw. subsymmetricus has low small denticles connecting blade and580
carina;4)Sw. subsymmetricushasa lessexpandedbasal cavityandanarrower581
carina, therefore appears more “slim”; 5) gaps between transverse ridges that582
becomelagertowardtheposteriorend. 583
ThoughSw.subsymmetricusandSweetognathusasymmetrican.sp.mayhave584
close affinities,Sw.asymmetrica n. sp. is restricted to theArtinskian to earliest585
Kungurian whereas Sw. subsymmetricus is found in younger rocks of late586
Kungurian to Roadian age (Kozur, 1995). Many reported occurrences of Sw.587
subsymmetricus inpre-middle-Kungurianstrata (thathavenotbeen illustrated)588
shouldbereassessed.589
The paratype shares a few common featureswithSw. variabilis. They both590
havetwobigtriangular-shapeddenticles.Thekeydifferenceisthepositionofthe591
basalcavity.Sw.variabilishasabasalcavityneartheposteriorend.Inaddition,592
Sw.variabilishasalongblade(blade/carinaratio!1)andfivetransverseridges593
30
with the widest being near the posterior end. Sw. asymmetrica n. sp. has a594
blade/carinaratioalways<1,andusuallysevenormoretransverseridgeswhile595
the widest occurs near the middle of the body. In addition, neither of Sw.596
subsymmetricus and Sw. variabilis should be considered as synonyms of Sw.597
paraguizhouensisandSw.guizhouensis(Shenetal.,2012).598
Occurrence:Artinskian-basalKungurian,basalChihsiaFm.ofSouthChina599
Note:Thespecimenshowninfig.3inPl.4seemlyhasagapbetweenbladeand600
carina.ThisisonlybecauseinthelateralviewtheSEMtookthepartwithmissing601
nodesintheasymmetricanteriortransverseridges.602
Sweetognathussp.A603
Pl.1fig.9604
Diagnosis: A Type III sweetognathid with tall and slim denticles and narrow605
ridges.606
Description:Bodyelongatedwithaheight/lengthratio≈ 1/2.Cusptallandslim,607
at least twice as high as any following denticles. The cusp is immediately608
followedby five to six very slimdenticles.The first and thirddenticles are the609
lowest. A gap is developed between the fourth and fifth denticles. Nodes are610
short,forming5-7lowandgenerallyevenlyspacedridges.611
Sweetognathustoriyamai(Igo,1981)612
Pl.1,fig.12,15.613
31
NeostreptognathodustoriyamaiIgo,1981,p.42-43,pl.6,figs.1-16614
SweetognathuswhiteiIgo,1981,Pl.7,fig.7?615
Remarks: The denticles of this species point forwards. The carina is616
lens-shaped—thus the widest is near the middle. There is a short and narrow617
ridgeconnectingthebladeandcarina.Theridgeisrelativelyhighanteriorlyand618
decreasesinheighttowardsthecarina,thusgivingatriangularshapeiflaterally619
viewed. 620
Comparisons:Theshortnarrowridgebetweenthefreebladeandcarinaisoneof621
the most distinguishable features of this species. This species has a broad,622
lens-likecarina,whichissimilartoSw.behnkeni.However,thelatterspecieshas623
“ledge-like” decorations on the carina, whereas Sw. toriyamai is decorated by624
lowertransverseridges.625
Occurrence: Artinskian, basal Chihsia Fm. of South China andKuchibora Fm. of626
Japan.627
628
7.Conclusions:629
Adetailed conodontbiostratigraphic and taxonomic studyof theEarly and630
Middle Permian strata at Tieqiao, South China has enabled recognition of 21631
mainand7subordinateconodontzones.Threenewspeciesareestablished.The632
followingconclusionscanbedrawn:633
32
1) The Tieqiao strata record a change in conodont faunas from Early Permian634
Sweetognathus dominated assemblages to Middle Permian gondolellids635
dominatedassemblagesfromthelatestKungurianonwards.Thisshiftcoincided636
witharelativesea-levelriseandchangetodeeperwaterfacies.637
2) The Early Permian Sweetognathus fauna represents an important evolution638
lineageandashallower(surface?)watergroup,whichevolvedinparalleltothe639
contemporarybutpossiblydeeper-dwellingMesogondolellafauna.640
3) The Chihsia Fm., which had been in many cases erroneously regarded as a641
MiddlePermianunit, isofEarlyPermianage.ItspanstheArtinskiantothelate642
Kungurian whilst the overlying Maokou Fm. straddles the Early and Middle643
Permian from the late Kungurian to latest Capitanian. The Chihsia/Maokou644
lithological boundary is thus locally not suitable for defining the Early-Middle645
Permianboundary(Kungurian-Roadianstageboundary).646
4)SpeciessuchasJ.palmataandJ.errataoccurattime-equivalentstratigraphic647
levels at Tieqiao as in west Texas, suggesting that they can be used for648
intercontinentalcorrelations.649
650
Acknowledgements651
Manycolleagueshelpedwithfieldandlabwork,includingH.S.Jiang(Wuhan),652
F. Li (now Guilin), Q. Li (now Qingdao), D. Lutz (Erlangen), F. Nenning (now653
Münster), W. Pan (now Guangzhou), L.N. Wang (Wuhan), Y.J. Yan (now654
33
Lianyungang)andW.Q.Xue(Wuhan).H. IgoandB.R.Wardlaware thanked for655
providing conodont literature. Y.D. Sun acknowledges the Alexander von656
Humboldt Foundation for a fellowship. This study is supported by the Chinese657
Fundamental Research Funds for the Central Universities (CUG130615), 973658
Program(2011CB808800),theNaturalScienceFoundationofChina(41272044;659
41472087;41572002)andthe111Project(B08030).DBacknowledgesfunding660
fromtheUKNaturalEnvironmentResearchCouncil(grantNE/J01799X/1).661
662
References663
Beauchamp,B.,Henderson, C.M., 1994.TheLowerPermianRaanes,GreatBear664
Cape and Trappers Cove Formations, Sverdrup Basin, Canadian Arctic -665
Stratigraphy and Conodont Zonations. Bulletin of Canadian Petroleum666
Geology,42(4):562-597.667
Behnken, F.H., 1975. Leonardian and Guadalupian (Permian) Conodont668
Biostratigraphy in Western and Southwestern United States. Journal of669
Paleontology,49(2):284-315.670
Bender, H., Stoppel, D., 1965. Perm-Conodonten. Geologisches Jahrbuch, 82:671
331-364.672
Bond,D.P.G.,Hilton,J.,Wignall,P.B.,Ali,J.R.,Stevens,L.G.,Sun,Y.,Lai,X.,2010.The673
MiddlePermian (Capitanian)massextinctionon landand in theoceans.674
Earth-ScienceReviews,102(1-2):100-116.675
Bond, D.P.G., Wignall, P.B., Joachimski, M.M., Sun, Y., Savov, I., Grasby, S.E.,676
Beauchamp,B.,Blomeier,D.P.G.,2015.AnabruptextinctionintheMiddle677
Permian (Capitanian) of the Boreal Realm (Spitsbergen) and its link to678
anoxiaandacidification.GeologicalSocietyofAmericaBulletin,127(9-10):679
1411-1421.680
34
Bucur, I.I., Munnecke, A., Granier, B., Yan, J., 2009. Remarks on the Permian681
dasycladaleanalgaSinoporellaleeiYabe,1949.Geobios,42(2):221-231.682
Burrett, C., Udchachon, M., Thassanapak, H., Chitnarin, A., 2015. Conodonts,683
radiolarians and ostracodes in the Permian E-Lert Formation, Loei Fold684
Belt,IndochinaTerrane,Thailand.GeologicalMagazine,152(01):106-142.685
Chen, B., Joachimski, M.M., Sun, Y.D., Shen, S.Z., Lai, X.L., 2011. Carbon and686
conodont apatite oxygen isotope records of Guadalupian-Lopingian687
boundary sections: Climatic or sea-level signal? Palaeogeography688
PalaeoclimatologyPalaeoecology,311(3-4):145-153.689
Chen,Z.Q.,George,A.,Yang,W.R.,2009.EffectsofMiddle-LatePermiansea-level690
changes and mass extinction on the formation of the Tieqiao skeletal691
mound in the Laibin area, South China. Australian Journal of Earth692
Sciences,56(6):745-763.693
Chuvashov,B.I.,Dyupina,G.V.,Mizens,G.A.,Chernykh,V.V.,1990.BasicSectionsof694
CarboniferousandLowerPermianofWesternSlopeofUrals.UralBranch695
AcademicScienceofRussia,Sverdlovsk,369pp.696
Clark,D.L.,Behnken,F.H.,1971.ConodontsandBiostratigraphyof thePermian.697
GeologicalSocietyofAmericaMemoir,127:415-439.698
Clark,D.L.,Carr,T.R.,Behnken,F.H.,Wardlaw,B.R.,Collinson,J.W.,1979.Permian699
ConodontBiostratigraphyintheGreatBasin.In:Sandberg,C.A.,Clark,D.L.700
(Eds.),ConodontbiostratigraphyoftheGreatBasinandRockyMountains.701
BrighamYoungUniversity,GeologyStudies,pp.143-150.702
Ding,H.,Wan,S.,1990.TheCarboniferous-Permianconodontevent-stratigraphy703
in the South of the North China Platform. Courier Forschungsinstitut704
Senckenberg,118:131-155.705
Gong, Y.-M., Shi, G.R., Zhang, L.-J., Weldon, E.A., 2010. Zoophycos composite706
ichnofabricsandtiersfromthePermianneriticfaciesinSouthChinaand707
south-easternAustralia.Lethaia,43(2):182-196.708
Gullo, M., Kozur, H., 1992. Conodonts from the pelagic deep-water Permian of709
central Western-Sicily (Italy). Neues Jahrbuch für Geologie und710
35
PaläontologieAbhandlungen,184(2):203-234.711
Henderson,C.M.,Mei,S.,2003.Stratigraphicversusenvironmentalsignificanceof712
PermianserratedconodontsaroundtheCisuralian-Guadalupianboundary:713
new evidence from Oman. Palaeogeography, Palaeoclimatology,714
Palaeoecology,191(3-4):301-328.715
Henderson, C.M., Davydov, D.I., Wardlaw, B.R., 2012. The Permian Period. In:716
Gradstein,F.M.,Ogg,J.G.,Schmitz,M.D.,Ogg,G.M.(Eds.),TheGeologicTime717
Scale2012.Elsevier,pp.653-679.718
Igo, H., 1981. Permian conodont biostratigraphy of Japan. Palaeontological719
Society of Japan special papers, 24. Palaeontological Society of Japan,720
Tokyo,51pp.721
Ji, Z.S., Yao, J.X., Jin, X.C., Yang, X.N.,Wang, Y.Z., Yang,H.L.,Wu, G.C., 2004. Early722
PermianConodontsfromtheBaoshanBlock,WesternYunnan,China.Acta723
GeologicaSinica,78(6):1179-1184.724
Jiang, H.S., Lai, X.L., Luo, G.M., Aldridge, R., Zhang, K.X., Wignall, P.B., 2007.725
Restudyofconodontzonationandevolutionacross theP/Tboundaryat726
Meishansection,Changxing,Zhejiang,China.GlobalandPlanetaryChange,727
55:39-55.728
Jin, Y.G., Wardlaw, B.R., Glenister, B.F., Kotlyar, G.V., 1997. Permian729
chronostratigraphicsubdivisions.Episodes,20(1):10-15.730
Jin, Y.G., Shen, S.Z., Henderson, C.M., Wang, X.D., Wang, W., Wang, Y., Cao, C.Q.,731
Shang,Q.H.,2006.TheGlobalStratotypeSectionandPoint(GSSP)forthe732
boundary between the Capitanian and Wuchiapingian Stage (Permian).733
Episodes,29(4):253-262.734
Kozur, H., 1993. Gullodus n. gen.–a new conodont genus and remarks to the735
pelagic Permian and Triassic of Western Sicily. Jb. Geol. B. -A., 136(1):736
77-87.737
Kozur,H.,1995.PermianConodontZonationanditsImportanceforthePermian738
StratigraphicStandardScale.Geol.Palaont.Mitt.Innsbruck,20(165-205).739
Kozur, H., Mostler, H., 1995. Guadalupian (Middle Permian) conodonts of740
36
sponge-bearinglimestonesfromthemarginsoftheDelawareBasin,West741
Texas.GeologiaCroatica,48(2):107-128.742
Lambert,L.L.,Wardlaw,B.R.,Nestell,M.K.,Nestell,G.P.,2002.LatestGuadalupian743
(Middle Permian) conodonts and foraminifers from West Texas.744
Micropaleontology,48(4):343-364.745
Lambert, L.L., Gorden L. Bell, J., Fronimos, J.A., Wardlaw, B.R., O.Yisa, M., 2010.746
ConodontbiostratigraphyofamorecompleteReefTrailMembersection747
near the type section, latest Guadalupian Series type region.748
Micropaleontology,56(1-2):233-253.749
Li,Z.S.,Zhan,L.P.,Dai,J.Y.,Jin,R.G.,Zhu,X.F.,Zhang,J.H.,Huang,H.Q.,Xu,D.Y.,Yan,750
Z.,Li,H.M.,1989.StudyonthePermian-Triassicbiostratigraphyandevent751
stratigraphy of northern Sichuan and southern Shaanxi. Geological752
Memoirs,9.GeologicalPublishingHouse,Beijing,435pp.753
Mei,S.L., Jin,Y.G.,Wardlaw,B.R.,1994a.Successionof conodontzones fromthe754
Permian ‘Kuhfeng’ Formation, Xuanhan, Sichuan and its implication in755
globalcorrelation.ActaPalaeontologicaSinica,33(1):1-23.756
Mei,S.L.,Jin,Y.G.,Wardlaw,B.R.,1994b.SuccessionofWuchiapingianconodonts757
from northeast Sichuan Province and its worldwide correlation. Acta758
MicropalaeontologicaSinica,11:121-139.759
Mei, S.L., Jin, Y.G., Wardlaw, B.R., 1994c. Zonation of conodonts from the760
Maokouan-Wuchiapingian boundary strata, SouthChina. Palaeoworld, 4:761
225-233.762
Mei, S.L., Jin, Y.G., Wardlaw, B.R., 1998. Conodont Succession of the763
Guadalupian-LopingianBoundary Strata in LaibinofGuangxi, China and764
WestTexas,USA.Palaeoworld,9:53-76.765
Mei,S.L.,Henderson,C.M.,Wardlaw,B.R.,2002.Evolutionanddistributionofthe766
conodontsSweetognathusandIranognathusandrelatedgeneraduringthe767
Permian, and their implications for climate change. Palaeogeography,768
Palaeoclimatology,Palaeoecology,180(1-3):57-91.769
Metcalfe, I., Sone, M., 2008. Biostratigraphy and palaeobiogeography of Lower770
37
Permian (lower Kungurian) conodonts from the Tak Fa Formation771
(Saraburi Limestone), Thailand. Palaeogeography, Palaeoclimatology,772
Palaeoecology,257(1-2):139-151.773
Nestell, M.K., Nestell, G.P., Wardlaw, B.R., Sweatt, M.J., 2006. Integrated774
biostratigraphy of foraminifers, radiolarians and conodonts in shallow775
anddeepwaterMiddlePermian(Capitanian)depositsofthe“Raderslide”,776
GuadalupeMountain,WestTexas.Stratigraphy,3(3):161-194.777
Nestell, M.K., Wardlaw, B.R., 2010a. Jinogondolella palmata, a new Gondolellid778
conodontspeciesfromtheBellCanyonFormation,MiddlePermian,West779
Texas.Micropaleontology,56(1-2):185-194.780
Nestell, M.K., Wardlaw, B.R., 2010b. Radiolarians and conodonts of the781
Guadalupian(MiddlePermian)ofWestTexas:advancesintaxonomyand782
biostratigraphy.Micropaleontology,56(1-2):1-6.783
Orchard, M.J., 1984. Early Permian conodonts from the Harper Ranch beds,784
Kamloops area, southern British Columbia. Paper - Geological Survey of785
Canada,84-1B:207-215.786
Orchard, M.J., Forster, P.J.L., 1988. Permian conodont biostratigraphy of the787
Harper Ranch Beds, near Kamloops, south-central British Columbia.788
Energy,MinesandResourcesCanada,pp.1-27.789
Qiu, Z., Wang, Q., Zou, C., Yan, D., Wei, H., 2013. Transgressive–regressive790
sequences on the slope of an isolated carbonate platform (Middle–Late791
Permian,Laibin,SouthChina).Facies,10.1007/s10347-012-0359-4:1-19.792
Rhodes, F.H., 1963. Conodont from the topmost Tensleep Sandstone of the793
eastern Big Norn Mountains, Wyoming. Journal of Paleontology, 37:794
401-408.795
Ritter,S.M.,1986.Taxonomicrevisionandphylogenyofpost-EarlyPermiancrisis796
bisselli-whiteiZoneconodontswithcommentsonlatePaleozoicdiversity.797
GeologicaetPalaeontologica,20:139-165.798
Sha, Q.A., Wu, S.W., Fu, J.M., 1990. An integrated investigation on the Permian799
System of Qin-Gui areas, with discussion on hydrocarbon potential.800
38
SciencePress,Beijing.801
Shen, S.Z.,Wang,Y.,Henderson,C.M.,Cao,C.Q.,Wang,W.,2007.Biostratigraphy802
and lithofacies of the Permian System in the Laibin-Heshan area of803
Guangxi,SouthChina.Palaeoworld,16(1-3):120-139.804
Shen,S.Z.,Yuan,D.X.,Henderson,C.M.,Tazawa,J.,Zhang,Y.C.,2012.Implications805
of Kungurian (Early Permian) conodonts from Hatahoko, Japan, for806
correlation between the Tethyan and international timescales.807
Micropaleontology,58(6):505-522.808
Sheng,J.Z.,Jin,Y.G.,1994.CorrelationofPermiandepositsofChina.Palaeoworld,809
4:14-113.810
Sun, Y.D., Lai, X.L., Jiang, H.S., Luo, G.M., Sun, S., Yan, C.B., Wignall, P.B., 2008.811
Guadalupian (Middle Permian) Conodont Faunas at Shangsi Section,812
Northeast Sichuan Province. Journal of China University of Geosciences,813
19(5):451-460.814
Sun,Y.D.,Lai,X.L.,Wignall,P.B.,Widdowson,M.,Ali,J.R.,Jiang,H.S.,Wang,W.,Yan,815
C.B., Bond, D.P.G., Védrine, S., 2010. Dating the onset and nature of the816
MiddlePermianEmeishanlargeigneousprovinceeruptionsinSWChina817
using conodont biostratigraphy and its bearing on mantle plume uplift818
models.Lithos,119(1-2):20-33.819
Wang, C.Y., Ritter, S.M., Clark, D.L., 1987. The Sweetognathus complex in the820
Permianof China: implications for evolution andhomeomorphy. Journal821
ofPaleontology,61:1047-1057.822
Wang,C.Y.,2002.PermianconodontsfromLaibinandHeshan,Guangxi.Bulletin823
of Nanjing Institute of Geology and Palaeontology, Acdemia Sinica, 15:824
180-190.825
Wang, W., Cao, C.Q., Wang, Y., 2004. The carbon isotope excursion on GSSP826
candidate section of Lopingian-Guadalupian boundary. Earth and827
PlanetaryScienceLetters,220(1-2):57-67.828
Wang, X.-D., Sugiyama, T., 2000. Diversity and extinction patterns of Permian829
coralfaunasofChina.Lethaia,33(4):285-294.830
39
Wang, Y., Jin, Y.G., 2000. Permian palaeogeographic evolution of the Jiangnan831
Basin, South China. Palaeogeography, Palaeoclimatology, Palaeoecology,832
160(1-2):35-44.833
Wang,Y.,Wang,J.,Chen,J.,Wang,W.J.,Shen,S.Z.,Henderson,C.M.,2011.Progress,834
Problems and Prospects on the Stratigraphy and Correlation of the835
KungurianStage,EarlyPermian(Cisuralian)Series.ActaGeologicaSinica,836
85(2):387-398.837
Wang,Z.H.,1994.EarlyPermianConodontsfromtheNashuiSection,Luodianof838
Guizhou.Palaeoworld,4:203-224.839
Wardlaw,B.R.,Grant,R.E.,1990.Conodontbiostratigraphyof thePermianRoad840
CanyonFormation,GlassMountains,Texas.U.S.GeologicalSurveyBulletin,841
1895:63-66.842
Wardlaw,B.R.,2000.GuadalupianconodontbiostratigraphyoftheGlassandDel843
Norte Mountains. In: Wardlaw, B.R., Grant, R.E., Rohr, D.M. (Eds.), The844
Guadalupiansymposium-SmithsonianContributionstotheEarthSciences,845
pp.37-87.846
Wardlaw, B.R., Nestell, M.K., 2010. Latest Middle Permian conodonts from the847
ApacheMountains,WestTexas.Micropaleontology,56(1-2):149-183.848
Wignall,P.B.,Sun,Y.,Bond,D.P.G., Izon,G.,Newton,R.J.,Védrine,S.,Widdowson,849
M.,Ali,J.R.,Lai,X.,Jiang,H.,Cope,H.,Bottrell,S.H.,2009a.Volcanism,Mass850
Extinction, and Carbon Isotope Fluctuations in the Middle Permian of851
China.Science,324:1179-1182.852
Wignall, P.B., Védrine, S., Bond, D.P.G., Wang, W., Lai, X.-L., Ali, J.R., Jiang, H.-S.,853
2009b.Faciesanalysisandsea-levelchangeattheGuadalupian–Lopingian854
Global Stratotype (Laibin, South China), and its bearing on the855
end-Guadalupian mass extinction. Journal of the Geological Society,856
166(4):655-666.857
Youngquist, W.L., Hawley, R.W., Miller, A.K., 1951. Phosphoria conodonts from858
southeasternIdaho.JournalofPaleontology,25(3):356-364.859
Zhang,K.X.,Lai,X.L.,Ding,M.H.,Liu, J.,1995.ConodontSequenceand itsGlobal860
40
CorrelationofPermian-TriassicBoundaryinMeishanSection,Changxing,861
Zhejiang Province. Earth Science-Journal of China University of862
Geosciences,20(6):669-676.863
Zhang,Z.,Wang,Y.,Zheng,Q.F.,2015.MiddlePermiansmallerforaminifersfrom864
the Maokou Formation at the Tieqiao section, Guangxi, South China.865
Palaeoworld,24(3):263-276.866
867
Figurecaptions868
Fig. 1 Middle Permian palaeogeographic reconstructions of South China and869
Laibinarea(afterWangandJin,2000;Wignalletal.,2009b).870
871
Fig.2Fieldphotographsofthestudiedsection.A,anoverviewoftheChihsiaFm.872
inthelowerpartofthesection.Adiggerinthefarside(bluesquare)asscale.B,a873
close reviewof fine laminatedBed111-112 transition (Roadian-Wordian). The874
pen(~15cmlong)asscale.875
876
Fig. 3 Log of the lower part of Tieqiao section (Asselian to Kungurian) with877
conodontrangesandzonation.878
879
Fig. 4 Log of the middle part of Tieqiao section (Kungurian to Wordian) with880
conodont ranges and zonation. Keys are the same as in Fig. 3. Note that the881
reportedoccurrenceofMesogondolellabisselliinBed91(Shaetal.,1990)cannot882
41
beconfirmedbyourdataset(fordetailsseediscussionoftheSw.adjunctuszone).883
Keysisthesameasinfigure3.884
885
Fig.5LogoftheupperpartofTieqiaosection(WordiantoWuchiapingian)with886
conodontrangesandzonation.Keysisthesameasinfigure3.887
888
Fig. 6 Correlation chart of the Early-Middle Permian with standard conodont889
zonation(Hendersonetal.2012)andTieqiao(thisstudy)andNashui(Meietal.,890
2002) sections. 1.,Pseudosw.monocornus; 2.,H. gulloides; 3.,G. sicilianus; 4.,H.891
excavates.892
893
Plate1.SEMimagesofTieqiaoconodonts-genusSweetognathus.Barscalefor100894
μm, ‘a’ for oral view, ‘b’ for lateral view. Default is oral view. 1, 7, 14, 17.895
Sweetognathus asymmetrica n. sp., 1, holotype, S1_018 (18-1); 7, paratype,896
S1_037 (22-2); 14, S_001 (18-1); 17, S_006 (24A); 2, 16. Sweetognathus whitei897
(Rhodes, 1963), 2, S1_019 (18-1); 16, S_005 (23A); 3, 8. Pseudosweetognathus898
costatus Wang, Ritter and Clark, 1987, 3, S1_021 (19-2), 8, S1_025 (21-2). 4.899
Sweetognathussp.4,S1_023(19-2);5.TransitionalformfromSw.inornatustoSw.900
asymmetrica n. sp., S1_038 (22-2); 6. Sweetognathus sp., S1_031 (22-1). 9.901
Sweetognathus sp. A., S1_026 (21-2); 10. Sweetognathus inornatus Ritter, 1986,902
S1_030(22-1);11.Sweetognathuscf.bogoslovskajaeKozurinKozurandMostler,903
42
1976, S1_020 (18-1); 12, 15. Sweetognathus toriyamai (Igo, 1981), 12, S_002904
(17c),15,juvenileform,S_003(17c);13.Neostreptognathodusexgr.pequopensis,905
S_004(17c).906
907
Plate 2. SEM images of Tieqiao conodonts—genera Sweetognathus,908
Pseudosweetognathus,NeostreptognathodusandHindeodus.Scalebaris100μm,909
‘a’fororalview,‘b’forlateralview.Defaultisoralview.1,2,4,9.Sweetognathus910
bogoslovskajae Kozur in Kozur and Mostler, 1976, 1, S1_039 (22-2), 2, S1_043911
(24-3), 4, juvenile, S1_057 (39-1), 9, S1_051 (27-1); 3, 12. Sweetognathus912
guizhouensis Bando et al., 1980, 3. S1_048 (26-3), 12. S1_055 (39-1); 5-7.913
PseudosweetognathuscostatusWang,RitterandClark,1987,5,S1_047(26-2),6.914
S1_049 (26-4), 7. S1_045 (25-2); 8. Neostreptognathodus prayi Behnken, 1975,915
gerontic form, S1_046 (25-2); 10. Sweetognathus clarki Morphotype I, (Kozur,916
1976),S1_050(27-1);11.Sweetognathus inornatusRitter,1986,S1_053(29-1);917
13. Hindeodus aff. catalanoi, S1_022 (19-2); 14, 16, 17. Hindeodus minutus918
(Ellison, 1941), 14, S1_044 (25-1), 16, S1_056 (39-1), 17, S1_052 (28-1); 15.919
Hindeodussp.S1_042(23-3).920
921
Plate 3. SEM images of Tieqiao conodonts—genera Sweetognathus and922
Pseudosweetognathus. Scalebar is 100μm, ‘a’ for oral view, ‘b’ for lateral view.923
Default is oral view. 1-3.Pseudosweetognathus costatusWang,Ritter andClark,924
43
1987,1.S1_065(52-2);2.S1_066(58-1),3.S1_068(65-2);4,5.Sweetognathus925
iranicusKozur,1975,4,S1_069(66-3),5,S1_071(71-1);6.Pseudosweetognathus926
monocornus(DaiandZhang,1989),S2_001(94-2);7,9,19.Sweetognathussp.7.927
S1_072 (90-7), 9. S2_002 (97-2), 19. S2_020 (100-1); 8, 10, 11. Sweetognathus928
adjunctus (Behnken, 1975), 8. S2_004 (97-2), 10. S2_005 (97-2), 11. S1_076929
(91-1); 12, Sweetognathus cf. paraguizhouensis S1_078 (91-3); 13. Transitional930
formbetweenSweetognathusiranicusandSweetognathushanzhongensis,S2_007931
(99-4); 14.Sweetognathus subsymmetricsWang,Ritter andClark, 1987, S2_039932
(100-3);15-18.Sweetognathushanzhongensis(Wang,1978),15,S2_038(100-5),933
16,S2_018(100-1),17,S2_010(100-1),18,S2_028(100-3).934
935
Plate 4. SEM images of Tieqiao conodonts—genera Sweetognathus, Gullodus,936
Hindeodus and Pseudohindeodus. Scale bar is 100 μm, ‘a’ for oral view, ‘b’ for937
lateral view. Default is oral view. 1. Sweetognathus sp. S2_049 (102-2); 2-4.938
SweetognathussubsymmetricsWang,RitterandClark,1987,2.S2_051(102-3);3.939
S2_056(102-4);4.S2_058(102-4);5,15.SweetognathusiranicusKozur,1975,5,940
S2_082(105-3),15,S2_072(103-2);6-7.Gullodustieqiaoensisnsp.,6.holotype,941
S1_060 (41-1), 7. paratype, S1_062 (41-2); 8. Gullodus sicilianus (Bender and942
Stoppel, 1956), S3_020 (115-4); 9, 10, 16, 18. Transitional forms between943
HindeodusandPseudohindeodus.Notethattheseelementsdevelopedweakapron944
structures on basal cavities. 9. S1_074 (91-1), 10. S1_075 (91-1), 16. S2_057945
(102-4),18.S2_053(102-3);11,12.GullodusduaniMeietal.,2002,11.S1_013946
44
(TQ-28), 12. S3_022 (115-7); 13. Pseudohindeodus ellipticae n. sp. Sun and Lai,947
paratype, S2_075 (104-2); 14.Pseudohindeodus ramovsi Gullo andKozur, 1992,948
S2_073(103-4);17.Hindeoduscf.wordensisWardlaw,2000;17,S2_060(102-5);949
19, 20. Hindeodus cf. julfensis 19, S2_014 (100-1), 20. S2_011 (100-1); 21.950
Hindeoduscf.permicus,S2_050(102-2);22,36.Hindeodussp.22,S2_061(102-5),951
36, S2_022 (100-3); 23, 26. Hindeodus sp. A. 23, S2_068 (103-2); 26, S2_084952
(106-1). 24, 25, 27, 30-32, 34, 35. Hindeodus permicus (Igo, 1981) 24. S2_081953
(105-3),25.S2_034(100-4),27.S2_026(100-3),30.S2_083(105-3),31.S2_016954
(100-1), 32. S2_062 (103-1), 34. S2_071 (103-2); 35, S2_067 (103-2). 28, 29.955
Hindeodusminutus (Ellison,1941),28.S2_027(100-3);29.S2_021(100-2);33.956
HindeodusgolloidesKozurandMostler,1995,S2_066(103-2).957
958
Plate 5. SEM images of Tieqiao conodonts—genera Mesogondolella and959
Jinogondolella.Scalebaris100μm,‘a’fororalview,‘b’forlateralview.Defaultis960
oralview.1,7.transitionaltypebetweenM.lambertitoJ.nankingensis,1,S3_001961
(109-2),7,S3_012(113-2);2.MesogondolellalambertiMeiandHenderson,2002,962
S3_002(109-2);3,8.Mesogondolellasp.,3,S3_004(109-3),8.S3_013(113-7);4.963
Mesogondolellaidahoensis(Youngquist,Hawley,Miller,1951),S3_005(110-2);5,964
6.Mesogondolellasiciliensis(Kozur,1975),5.S3_007(111-1);6.S3_006(111-1);965
9.Jinogondolellanankingensis(Ching,1960),S3_011(111-5);10-11.Jinogodolella966
errataWardlawandNestell, 2000,10. S3_019 (115-3),11. S3_017 (115-2);12.967
Jinogondolella aserrata (Clark and Behnken, 1979), S3_021 (115-4); 13, 16.968
45
Jinogondolellasp.,13.SP_051(115-3),16,S3_037(116-7);14,15.Jinogondolella969
postserrata (Behnken, 1975), S3_028 (116-1); 15. S3_036 (116-7); 17, 18.970
Jinogondolellashannoni(Wardlaw,1994),17.S3_040(116-8);18,S3_030(116-2);971
19.Jinogondolellaaltudaensis(Kozur,1992),S3_033(116-3).972
973
Plate6. SEM imagesofTieqiao conodonts—genera JinogondolellaandClarkina.974
Scale bar is 100 μm, ‘a’ for oral view, ‘b’ for lateral view and ‘c’ for back view.975
Defaultisoralview.1.Jinogondolellaprexuanhanensis(MeiandWardlaw,1994),976
S4_004 (TQ-11); 2. Jinogondolella cf. prexuanhanensis SP_010 (118-2); 3-5.977
Jinogondolellashannoni(Wardlaw,1994),3.S3_062(118-2);4.S4_006(TQ-17+);978
5,SP_014(118-2);6,17.Jinogondolellasp.,6,SP_013(118-2),17,S_035(119A);979
7. Jinogondolella xuanhanensis (Mei and Wardlaw, 1994), 06-70_023 (TQ-6f);980
8-11. Jinogondolella granti (Mei and Wardlaw, 1994), 8. 06-70_024 (TQ-6f), 9.981
06-70_022 (TQ-6f), 10. TQ6f_010 (TQ-6f), 11. 06-70_027b (TQ-6f); 12-13.982
Clarkina postbitteri Mei and Warldlaw, 1994, 12. S6_054 (TQ-1), 13. C6_040a983
(TQ-1). 14. Clarkina sp., S6_055 (TQ-1); 15. Clarkina transcaucasica Gullo and984
Kozur, 1992, S4_003 (134-9); 16. Clarkina hongshuiensis Henderson, Mei and985
Wardlaw,2002,S_029(TQ-1).986
987
Plate 7. SEM images of Tieqiao conodonts—genera Hindeodus, Jinogondolella,988
Mesogondolellaand Sweetognathus.Scalebar is100μm, ‘a’ fororalview, ‘b’ for989
46
lateral view. Default is oral view. 1, transitional form between Sweetognathus990
bogoslovskajae and Sweetognathus inornatus, S_007 (24A); 2. Sweetognathus991
inornatus Ritter, 1986, S_008 (26C); 3. Sweetognathus fengshanensis Mei and992
Wardlaw,1998,S_016(117-3);4.Mesogondolellaidahoensis(Youngquist,Hawley993
andMiller,1951),S_009(109-2);5.Jinogondolellapalmata(NestellandWardlaw,994
2010), S_025 (111-1-2); 6-8. Hindeodus catalanoi (Gullo and Kozur, 1992), 6,995
S3_052 (117-2), 7, S3_052 (117-2), 8, S3_043 (116-12); 9-10. Sweetognathus996
hanzhongensis (Wang, 1978), 9, S4_012 (TQ-25), 10, S_025 (115-8). 11-12.997
Pseudohindeodusaugustus (Igo,1981),11, S7_007 (102-4),12, S7_005 (102-4).998
13.Pseudohindeodussp.S7_003(104-2).14.Pseudohindeodusellipticaen.sp.Sun999
andLai,holotype,S7_001(104-2).1000