Vecoli Et Al 2011

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Research papers

Cryptospore assemblages from Upper Ordovician (KatianndashHirnantian) strata of Anticosti Island Queacutebec Canada and Estonia Palaeophytogeographic andpalaeoclimatic implications

Marco Vecoli ab Aureacutelien Delabroye c Amalia Spina d Olle Hints e

a Universiteacute Lille 1 Laboratoire laquo Geacuteosystegravemes raquo CNRS FRE 3298 SN5 Citeacute Scienti 1047297que Villeneuve dAscq 59655 Franceb Dipartimento di Scienze della Terra Universitagrave di Cagliari via Trentino 51 09127 Cagliari Italyc Universiteacute Paul Sabatier de Toulouse GET CNRS UMR 5563 14 av Edouard Belin 31400 Toulouse France

d CNR IRPI Perugia Italye Institute of Geology at Tallinn University of Technology Ehitajate 5 19086 Tallinn Estonia

a b s t r a c ta r t i c l e i n f o

Article history

Received 24 January 2011Received in revised form 29 April 2011Accepted 14 May 2011Available online 23 May 2011

Keywords

cryptosporesLate OrdovicianHirnantian

terrestrializationphytogeographypalaeobotany

Rich palynological assemblages have been recovered from deposits of Hirnantian age in Anticosti Island(Queacutebec Canada) and in borehole Valga-10 in southern Estonia The assemblages are well preserved andinclude acritarchs chitinozoans and cryptospores The age of the deposits is well constrained by means of palynomorphs (acritarchs and chitinozoans) as well as sequence stratigraphic and chemostratigraphiccorrelations Cryptospore assemblages from the two localities are similar and are also broadly comparable tothe few known coeval assemblages described elsewhere They include 11 genera and 20 species and testify tothe presence of an extended and diverse 1047298ora during Hirnantian times in Laurentia and for the 1047297rst time alsoin BalticaThe present 1047297ndings contribute to an improved knowledge of origin and early development of vegetative

cover The recovery of diverse and abundant cryptospores in Hirnantian deposits may be related to increasedinput of land-derived sediment during the global sea-level fall linked to the Late Ordovician glaciation but italso demonstrates that the early land plants may have tolerated a wide range of climatic conditions

copy 2011 Elsevier BV All rights reserved

1 Introduction

It is now generally accepted that by mid-Ordovician times(DapingianndashDarriwilian) bryophyte-like land plants had alreadyevolved in parts of Gondwana (Le Heacuterisseacute et al 2007 Rubinstein etal 2010 2011 Strother et al 1996) Cryptospores from the MiddleOrdovician Hanadir Sandstone in Saudi Arabia have been accepted as

the oldestevidenceof land plants because they possess morphologicalanalogs with spores produced by some living hepatics (Gray 1985Richardson 1996b Steemans 2000 Strother et al 1996 Taylor1995 Wellman and Gray 2000 Wellman et al 2003) Slightly olderless diverse and poorly preserved cryptospores with similar mor-phologies to the Hanadir assemblage occur in NW Argentina(Rubinstein et al 2010) Isolated specimens of laevigate permanenttetrads and dyads have been reported from the Klabava Formation(ArenigndashLlanvirn)of Bohemia by Vavrdovaacute (1990) but this report hasyet to be substantiated by further studies Richardson (1996ab)

reviewed all basicmorphological typesof cryptospores which includemonads and permanent dyads and tetrads as well as theirstratigraphic signi1047297cance He described a series of evolutionaryphases of early land vegetation based on the morphological in-novations observed in the fossil record of terrestrial palynomorphs(OrdovicianndashEarly Devonian) Richardson (1996b) proposed a ldquopre-cryptosporerdquo phase to include earliest Ordovician enigmatic spore-

like palynomorphs such as the enigmatic taxa Virgatasporites and Attritasporites which might have been related to freshwater algae orearliest terrestrial vegetation It is important to point out that thesepalynomorphs had been already considered as ldquosporesrdquo by theoriginal report of Combaz (1967) Vecoli (2000 2004) has alsonoted the association of Tremadocian sporomorph assemblageswith palynofacies indicating proximal depositional conditions withpossible in1047298uence of terrestrial sedimentary input The existence of primitive land plants during earliest Ordovician (Tremadocian) timesis also consistent with evidence from recent research suggesting thatterrestrial 1047298oras might have been widespread since the Cambrian(Strother et al 2004 Taylor and Strother 2008 2009 Tomescu et al2009) Cryptospores preserved in sporangia have been described fromsediments as old as Caradoc from the Hasirah Member of the Sa1047297q

Formation Haima Supergroup of Oman (Wellman et al 2003) The

Review of Palaeobotany and Palynology 166 (2011) 76ndash93

Corresponding author at Universiteacute Lille 1 Laboratoire laquo Geacuteosystegravemes raquo CNRS FRE3298 SN5 Citeacute Scienti1047297que Villeneuve dAscq 59655 France Fax +33 320434910

E-mail address marcovecoliuniv-lille1fr (M Vecoli)

0034-6667$ ndash see front matter copy 2011 Elsevier BV All rights reserveddoi101016jrevpalbo201105006

Contents lists available at ScienceDirect

Review of Palaeobotany and Palynology

j o u r n a l h o m e p a g e w w w e l s ev i e r c o m l o c a t e r ev p a l b o






2010) Taking into account the above correlation in western Anticosti

the base of the Hirnantian Stage is recognized at the VaureacutealndashEllis Baytransition It coincides with major faunal and 1047298oral changes andwith the start of a major δ13C excursion (beginning of the HICE sensu

Bergstroumlm et al 2009) occurring in the basal part of the Belonechitina

gamachiana chitinozoan biozone The same stratigraphic horizon islocated in the middle part of the Ellis Bay Fm in eastern Anticosti(Delabroye 2010 Desrochers et al 2010 Fig 2)

WESTERN ANTICOSTI EASTERN ANTICOSTI VALGA-10 DRILL CORE

balls and pillows

calcareous shales

bioherms

Oncolitic grainstones cross beddedgrainstones

dark grey marls

wave-rippled

grainstones

HCS sandstones

Sandstones

lenticular

limestones

nodular limestones

Hardgrounds

tempestites

limestones

argillaceaous limestones

dolomitic limestones

dolostones

argillaceaous dolostones

silty dolostones

marls

dolomitic marls

Fig 2 Lithostratigraphy and chronostratigraphy of investigated sections with location of sampling levels

78 M Vecoli et al Review of Palaeobotany and Palynology 166 (2011) 76ndash93




The OrdovicianndashSilurian boundary is identi1047297ed approximately 2 mabove the base of the Becscie Fm in western Anticosti and about 10 mabove the base of the same formation in eastern Anticosti ( Ancyr-

ochitina ellisbayensis biozone Achab et al 2011 Fig 2 see alsoVerniers and Vandenbroucke 2006)

Withrespecttosedimentarydynamics Desrocherset al (2010)have

identi1047297ed two large Transgressivendash

Regressive cycles in the lower andupper part of the Ellis Bay Fm respectively The regressive phases havebeen interpreted as having been forced by glacio-eustasy and seem tocorrespond to the two well-known Late Ordovician glacial-relatedregressions 1047297rst recorded in North-Gondwana (Delabroye et al 2010Desrochers et al 2010 Loi et al 2010) Each of these glacial cyclescorrelates with a carbon isotope positive excursion (HICE Delabroye2010 Delabroye et al 2010 Desrochers et al 2010)

Correlation of Late Ordovician sequences from Laurentia andBaltica are problematical because no diagnostic Hirnantian graptolitesof the Normalograptus extraordinarius and N persculptus biozones co-occur with palynomorphs in the two areas (see Delabroye and Vecoli2010 Hints et al 2010 for full discussion) In Baltica the base of theHirnantian Stage is thus traced based on chemostratigraphic argu-

ments in correspondence of a major carbon isotope positive excursionin the Porkuni Regional Stage interpreted as the HICE (Hints et al2010 Kaljo et al 2007 2008) The base of this excursion is correlatedto the base of the Spinachitina taugourdeaui biozone in Baltica(Brenchley et al 2003) and to the base of the B gamachiana biozoneon Anticosti (Achab et al 2011 Delabroye 2010 Desrochers et al2010) If we consider that isotopic excursions mark a synchronouschronostratigraphic event then the S taugourdeaui chitinozoanbiozone must be diachronous across the two areas

At present it is not possible to argue in favor of one or the otherhypotheses (ie diachronism of the isotopic signal or of thechitinozoan biozone successions) However in Baltica the latestOrdovician sedimentary successions are thinner and less continuousthan in Anticosti thus preserving a less detailed Late Ordovicianstratigraphicpalaeontologicisotopic record

For the scope of the present paper we consider the position of thebase of the Hirnantian as accepted by workers in the two areas(Fig 2) Accordingly in the Valga borehole the upper Pirgu RegionalStage (upper Katian) is represented by the Jelgava Parojeva and Kuiliformations where Notildelvak (in Potildeldvere 2001) has identi1047297ed theConochitina rugata chitinozoan biozone (Fig 2) B gamachiana hasnot been recovered from the Valga borehole possibly due to theoxidized nature of rocks in the topmost Pirgu Regional Stage (KuiliFm) that are mostly barren of organic-walled microfossils ThePorkuni Regional Stage (Hirnantian) ndash where Kaljo et al (2007) haveidenti1047297ed the HICE ndash is represented by the Kuldiga and Saldusformations (Fig2)The Spinachitina taugourdeaui biozone is identi1047297edin the Bernati Member ndash 1047297rst member of the Kuldiga Fm and theConochitina scabra biozone in the Ēdole Member ndash second and last

memberof the same formation (Fig 2) The base of the Juuru RegionalStage (basal Silurian) lies at the base of the OtildehneFm (Fig 2) All thesestratigraphic units are lithologically characterized by alternatingdolomiticnon-dolomitic marls and limestones (Potildeldvere 2001)

3 Sampling palynological preparations and cryptospore

productive samples

Palynological analyses have been performed on a total of 124samples from Anticosti Island (77 samples from the upper VaureacutealEllis Bay and basal Becscie formations in western Anticosti and 47samples from the upper Ellis Bay and basal Becscie formations ineastern Anticosti) and 14 samples from the Jelgava to the Otildehneformations in the Valga-10 borehole section All samples were treated

by standard palynological acid maceration density separation of theorganic matter and 1047297ltration of the organic-rich residue at 15 μ mSamples produced a rich and extremely well preserved assemblage of

diverse acritarchs and prasinophytes detailed elsewhere (egDelabroye 2010 Delabroye et al 2011)

Light microscope observations were performed on palynologicalslides using an Axioplan 2 Zeiss microscope using Differential Interfer-ence Contrast (DIC) technique in transmitted light Images werecaptured with an Axiocam digital microscope camera and successively

corrected for contrast and brightness using Adobe PhotoshopregSEM observations were done using a FEI Quanta 200 ESEM onpalynological smears on circular glass slides mounted on SEM stubsand gold-coated The circular slides were successively removed andmounted permanently on palynological slides for viewing under lightmicroscope

4 Results cryptospore assemblages

Among the 77 samples analyzed from the western Anticostisection three from the Vaureacuteal Fm and 1047297ve from the Ellis Bay Fmcontained well-preserved cryptospores (Fig 2 and Table 1) Thesection from eastern Anticosti was more productive with 11 samples(two in the Ellis Bay Fm and nine in the basal Becscie Fm) containing

abundant and well-preserved cryptospores (Fig 2 Table 1) The mostdiverse and rich cryptospore assemblages are from samples from theBecscie Fm (eastern Anticosti) This clearly re1047298ects the proximalshallow water mixed siliciclasticndashcarbonate facies of the sediments of the eastern part of Anticosti Island in1047298uenced by terrestrially derivedsedimentation and thus richer in cryptospore content versus thedeeper-water carbonate sedimentary setting of the western sectionsIt also broadly indicates an increase in cryptospore diversity from theKatian up to the uppermost Hirnantian

In the Estonian Valga section cryptospores which are also well-preserved occur abundantly only in 1047297ve samples two from the

Jelgava Fm and one sample from each of the Kuili Kuldiga and Saldusformations (Fig 2 and Table 2) The sample from the latter formation(uppermost Hirnantian) is also the richest and the most diversewhich con1047297rms the trend towards increasing diversity from Katian toHirnantian times noted also in the Anticosti sections The Estonianassemblage comprises 19 cryptospore species 10 of which are sharedwith the assemblages from the two Anticostan sections containing 11species in total The assemblages from Anticosti are thus less diverse

Table 1

Cryptospore occurrences in the western (A) and eastern (B) Anticosti sections

Sample T e t r a h e d r a l e t e s g r a y a e

T e t r a h e d r a l e t e s m e d i n e n s i s

P s e u d o d

y a d o s p o r a l a e v i g a t a

R i m o s o t

e t r a s p r o b l e m a t i c a

G n e u d n a s p o r a d i v e l l o m e d i a

V e l a t i t e t r a s r u g o s a

S e g e s t r e

s p o r a r u g o s a

C h e i l o t e

t r a s s p 2

D y a d o s p o r a m u r u s a t t e n u a t a

Sample R i m o s o t


C h e i l o t e

t r a s s p 1

C h e i l o t e

t r a s s p 2


V e l a t i t e t r a s l a e v i g a t a


P s e u d o d


T e t r a h e d r a l e t e s g r a y a e



EB42 EB324

EB132 EB320

EB18 EB319

EB111 EB318

EB109 EB316

EB94 EB314

EB4 EB313

EB2 EB312

EB311

EB310

EB309

EB308EB346

EB366

No cryptospores No cryptospores

No cryptospores

79M Vecoli et al Review of Palaeobotany and Palynology 166 (2011) 76ndash93




but generally the taxonomic composition of the assemblages fromEstonia and Anticosti are similar The similarity is even more evident

at the generic level with eight out of eleven genera shared betweenthe assemblages from the two areasThe assemblages include tetrads and dyads either naked or

enveloped as well as alete monads Tetrads (either naked or enveloped)are the dominant morphotype throughout the study sections They areespecially diverse in the palynologically richest samples from theuppermost Hirnantian levels

5 Systematic paleontology

Cryptospores are treated herein as an informal taxonomic groupWe follow this de1047297nition of cryptospores ldquoNon-marine sporomorphs(non-pollen grains) without those typical visible haptotypic featuressuch as trilete marks or furrows which characterize tracheophytespores and pollen grains Single grains or monads permanent dyadsand tetrads are included as are sporomorphs separated from polyadswhich may or may not preserve contact areasrdquo (from Strother 1991)Within the cryptospores subgroups can be distinguished based onpresenceabsence of morphological features following the subdivi-sion and terminology proposed by Wellman and Richardson (1993)Richardson (1996ab) and Wellman and Richardson (1996) The termldquomiosporerdquo is used herein as a general term indicating all fossil plantspores smaller than 200 μ m (Steemans 2000) thus including bothcryptospores and trilete spores

The synonymy listings include the basionym any emendationspublished records which are considered important for a correctinterpretation of a particular taxon concept as well as any publishedincorrect identi1047297cation of the particular taxon

51 Naked fused cryptospore tetrads

Group characterized by permanent tetrads without outer enve-lopes and with spores fused together No lines of attachment on thetetrad surface marking the position of planes of attachment betweenthe spores

Genus Cheilotetras Wellman and Richardson 1993Type species Cheilotetras caledonica Wellman and Richardson 1993

Cheilotetras caledonica Wellman and Richardson 1993Plate I 25 Plate III 4710

1993 Cheilotetras caledonica Wellman and Richardson pp 162ndash

163 pl 1 1047297gs 1ndash7Dimensions 30 36 40 45 μ m (four specimens measured)Occurrence Valga-10 drillcore Jelgava Fm sample VA46 (3457)

Kuili Fm sample VA42 (3290) Kuldiga Fm Edole Mb sample VA40(3200) Saldus Fm Brocēni Mb sample VA38 (3132)

Previous records This species has been previously reported onlyfrom post-Ordovician strata Llandovery Deerhope Fm PentlandHills Inlier Scotland (Molyneux et al 2008) Wenlock LesmahagowHagshaw Hills and North Esk Inliers of the Midland Valley of Scotland(Wellman and Richardson 1993) Henshaw Fm Pentland Hills InlierScotland (Molyneux et al 2008) WenlockndashLudlow StonehavenGroup Scotland (Wellman 1993) WenlockndashPridoli Rumney InlierSouth Wales (Burgess and Richardson 1995) Ludfordian Burgsvik

Fm Gotland Sweden (Hagstrom 1997) Ludlow (Gorstian)ndash

PridoliArisaig Group Nova Scotia Canada (Beck and Strother 2001)Lochkovian Clam Bank Fm Western Newfoundland Canada (Burdenet al 2002) Tawil Fm Saudi Arabia (Steemans et al 2007)

Cheilotetras sp 1Plate II 16ndash18

Description Permanent fused cryptospore tetrad (no line of attachment visible) characterized by sub-rounded sur1047297cially laevi-gate spores

Dimensions 32 40 44 μ m (three specimens measured)Occurrence Eastern Anticosti Becscie Fm Fox Point Mb samples

EB314 and EB324Comparison In contrast to typical Cheilotetras caledonica the

present specimens do not show the distinct ldquo1047298angerdquo extendingbeyond junctions between adjacent spores Also the individual sporeswhich comprise the tetrad are smaller

Cheilotetras sp 2Plate I 10

Description Permanently fused cryptospore tetrad (no line of attachment visible) characterized by sub-triangular spores These are

joined at thickened equatorial crassitudes about 2ndash3 μ m Sporescompletely laevigate

Dimensions 17 (19) 22 μ m (ten specimens measured)Occurrence Western Anticosti Ellis Bay Fm member 1 sample

EB109 Eastern Anticosti Ellis Bay Fm Lousy Cove Mb sample EB

346 Valga-10 drillcore Jelgava Fm sample VA46 (3457) Kuili Fmsample VA42 (3290) Kuldiga Fm Edole Mb sample VA40 (3200)Saldus Fm Brocēni Mb sample VA38 (3132)

Comparison The present specimens are distinguished by otherCheilotetras species essentially for their diminutive size they arebroadly similar to the previously reported Cheilotetras sp in Wellman

Table 2

Cryptospore occurrences in the Valga-10 borehole section



P s e u d o d y a d o s

p o r a l a e v i g a t a


S e g e s t r e s p o r a

r u g o s a

C h e i l o t e t r a s c a l e d o n i c a

C h e i l o t e t r a s s p 2

D y a d o s p o r a m

u r u s a t t e n u a t a

R i m o s o t e t r a s p r o b l e m a t i c a

H i s p a n a e d i s c u

s s p

A b d i t u s d y a d u

s l a e v i g a t u s

V e l a t i t e t r a s s p 1



l a e v i g a t a


m e m b r a n i f e r a

S p h a e r a s a c c u s g l a b e l l u s

V e l a t i t e t r a s l a

e v i g a t a

V e l a t i t e t r a s r u

g o s a

A l e t e c r y p t o s

p o r e

VA38 3132

VA40 3200

VA42 3290

VA44 3355

VA46 3457

Plate I Cryptospores from Anticosti Island and Valga-10 borehole (scale bar 10 μ m)

1 Tetrahedraletes medinensis Strother and Traverse emend Wellman and Richardson 1993 Sample VA38 (3132)2 5 Cheilotetras caledonica Wellman and Richardson 1993 Sample VA38 (3132)3 6 8 Velatitetras sp 1 Samples 3 8 VA38 (3132) 6 VA 46 (3456)4 7 Velatitetras sp 2 Sample VA38 (3132)

9 Rimosotetras problematica Burgess 1991 Sample EB 36610 Cheilotetras sp 2 Sample EB 34611 Velatitetras rugosa (Strother and Traverse) Steemans Le Heacuterisseacute and Bozdogan 1996 Sample EB 34612 Velatitetras laevigata Burgess 1991 Sample VA38 (3132)





Plate I





Plate II Cryptospores from Anticosti Island (scale bar 20 μ m EF England Finder)

1 3 4 10ndash12 Tetrahedraletes grayae Strother 1991 (1slideEB308 EF B29 3 slide EB314EF M1924 slide EB309EF Q18 10slideEB311 EFG4211 slide EB318EF J39 12slide EB309 EF U381)

2 5 6ndash9 13ndash15 Tetrahedraletes medinensis (Strother and Traverse) Wellman and Richardson 1993 (2 slide EB4[1] EF T434 5 slide EB2[1] EF L38 6 slide EB308 EF M38 7slide EB309 EF J314 8 slide EB309 EF R44 9 slide EB 314 EF F193 13 slide EB314 EF B221 14 slide EB2[1] EF N314 15 slide EB111[2] EF L261)

16ndash18 Cheilotetras sp 1 Wellman and Richardson 1993 (16 slide EB314 EF R24 17 slide EB314 EF G324 18 slide EB314 EF J321)19ndash20 Rimosotetras problematica Burgess 1991 (19 slide EB4[1] EF P24 20 slide EB111 EF L26)





(1996) Cheilotetras sp in Wang et al (1997) and Cheilotetras sp A inRubinstein and Vaccari (2004)

52 Naked unfused cryptospore tetrads

Permanent tetrads constituted of discrete spores with line of

attachment clearly visible on the tetrad surface and without enclosingenvelope The tetrads are never recorded completely dissociatedsuggesting that they are dispersed intact and remain permanentlyattached

Genus Tetrahedraletes Strother and Traverse emend Wellman andRichardson 1993

Type species Tetrahedraletes medinensis Strother and Traverseemend Wellman and Richardson 1993

Tetrahedraletes medinensis Strother and Traverse emend Wellmanand Richardson 1993

Plate I 1 Plate II 2 5 6ndash9 13ndash15 Plate III 1ndash3 5 Plate IV 41214

1971 lsquo

Spore tetrads in tetrahedral con1047297

gurationrsquo

Gray and Boucotp 919 1047297g 1h1971 lsquoTetrad of rather thick walled spore-like alete palynomorphsrsquo

Cramer pl 4 1047297g 11972 lsquoNon-miospore tetradic palynomorphrsquo Cramer and Diacuteez

p 116 pl 36 1047297gs 79 and 841979 Tetrahedraletes medinensis Strother and Traverse p 8 pl 1

1047297gs 5 14ndash171979 Nodospora burnhamensis Strother and Traverse p 10 pl 1

1047297g 11 pl 2 1047297g 1sic 1993 Tetrahedraletes medinensis (Strother and Traverse)

emend mdash Wellman and Richardson p 165 pl 2 1047297gs 8 10ndash12For an exhaustive synonymy see Strother (1991 p 122) and

Wellman and Richardson (1993 p 165)

Dimensions 22 (24) 30 μ m (28 specimens measured)Occurrence Western Anticosti Vaureal Fm samples EB2ndashEB94

Eastern Anticosti Becscie Fm Fox Point Member samples EB308ndash

EB311 EB314ndashEB318 and EB320 Valga-10 drillcore Jelgava Fmsample VA46 (3457) Kuili Fm sample VA42 (3290) Kuldiga FmEdole Mb sample VA40 (3200) Saldus Fm Brocēni Mb sampleVA38 (3132)

Previous records This species has been recorded worldwide andranges from Ordovician (Caradoc) to Devonian Jacobson and Achab(1985) already reported the occurrence of T medinensis in theVaurealFm of Anticosti Other selected previous Ordovician occurrences areas follows Caradoc type section Shropshire UK (Wellman 1996)Late Ordovician (CaradocndashAshgill late KatianndashHirnantian) BedinanFm southeastern Turkey (Steemans et al 1996) Naumlr borehole

Gotland Sweden (Gray 1988 Le Heacuterisseacute 1989) E1ndash81 boreholenortheastern Libya (Richardson 1988) Kalpintag Fm South XinjiangChina (Wang et al 1997) Mallowa Salt Carribuddy Basin Australia(Foster and Williams 1991) Hirnantian type Llandovery area south-west Wales ( persculptus graptolite Biozone in this area the speciesranges up to the turriculatus graptolite Biozone upper LlandoveryBurgess 1991)upper memberof the Salar del RincoacutenFm (Rubinsteinand Vaccari 2004) Latest Ashgill or earliest Llandovery CedabergFm South Africa (Gray et al 1986)

Tetrahedraletes grayae Strother 1991Plate II 1 3 4 10ndash12

1981Tetrahedraletes sp Duf 1047297eld and Legault Pl 2 1047297g 6

1982 aff Tetraletes CramerndashVavrdovaacute pl 1 1047297g 81984 spore tetrad type A Vavrdovaacute pl 2 1047297g 11988 Tetrahedraletes sp Richardson pl 19 1047297g 1

1991 Tetrahedraletes grayii Strother pp 221ndash222 pl 1 1047297gs 1ndash32001 Tetrahedraletes grayaeStrother 1991mdash Steemans p 9 1047297g 515

Dimensions 32 (45) 50 μ m (12 specimens measured)Occurrence Western Anticosti Vaureal Fm sample EB2 Ellis Bay

Fm member 1 sample EB111 Eastern Anticosti Becscie Fm Fox

Point Mb samples EB309 EB311 EB314 and EB318 Valga-10drillcore Jelgava Fm sample VA46 (3457) Kuili Fm sample VA42(3290) Kuldiga Fm Edole Mb sample VA40 (3200) Saldus FmBrocēni Mb sample VA38 (3132)

Remarks and comparisons The diameter of typical T grayae from thetype locality in North America is around 35 μ m (Strother 1991) Thepresent specimens on average have larger size more similarly tothe population of the sporomorphs of the morphotype Nodosporandash

Tetrahedraletes sp A of Richardson (1988) showing a dimensionalranges from 29 to 52 μ m

Previous records Upper CaradocndashLower Telychian subsurface of northeastern Libya (Richardson 1988) Ashgill Couches Fort AtkinsonDolomite McQuokata Group Illinois (Strother 1991) OostduinkerkeboreholeBrabant Massif Belgium (Steemans2001)KosovFmCentral

Bohemia Czech Republic (Vavrdovaacute 1982) Llandovery Jupiter FmAnticosti Island Queacutebec Canada (Duf 1047297eld 1985) Medina GroupNiagara Gorge Lewiston New York (Llandovery Miller and Eames1982) Early Silurian Centerville Fm central Ohio USA (Taylor 2002)

Genus Rimosotetras Burgess 1991Type species Rimosotetras problematica Burgess 1991

Rimosotetras problematica Burgess 1991Plate I 9 Plate II 1920 Plate III 1112 Plate IV 1320

1971 lsquoSpore tetradrsquo Gray and Boucot 1047297g 1(g)1985 lsquoLoose tetradsrsquo Richardson p 29 pl 15 1047297gs 5 and 61988 Nodospora burnhamensis lsquoloosetetradrsquo Richardson pl 191047297gs

11 and 121991 Rimosotetras problematica Burgess pp 586ndash587 pl 1 1047297gs 12

14 and 15


Fm member 1 sample EB111 Eastern Anticosti Ellis Bay Fm LousyCove Mb sample EB366 Becscie Fm Fox Point Mb samples EB316ndash

EB318 Valga-10 drillcore Jelgava Fm sample VA46 (3457) Kuili Fmsample VA42 (3290) Saldus Fm Brocēni Mb sample VA38 (3132)

Previous records This species has a long stratigraphic range havingbeen frequently recorded in strata across the Late Ordovician throughSilurian such as in the type Llandovery area (HirnantianndashearlyLlandovery from persculptus to cyphus graptolite biozones Burgess1991) southeastern Turkey (Caradoc to Ludlow Bedina and Dada

formations Steemans et al 1996) North Africa (Caradoc to lateLlandovery subsurface of northeastern Libya Richardson 1988) andnorthwestern Argentina (Salar del Rincoacuten Formation Puna regionRubinsteinand Vaccari2004)FurtherSilurian occurrencesare as followsLlandovery Medina Group Niagara Gorge New York USA (Miller andEames 1982) Tuscarora Fm Pennsylvania USA (Strotherand Traverse1979) Qusaiba Member of Qalibah Fm Saudi Arabia (Steemans et al2000 Wellmanet al 2000) Tiangua Fm SerraGrande Group Brazil (LeHeacuterisseacute et al 2001) Villa Maria Fm ParanaacuteBasin Brazil (Mizusaki et al2002) Itacurubi Group Paranaacute Basin Paraguay (Mendlowicz Maulleret al 2004) Wenlock type Wenlock area Shropshire UK (Burgess andRichardson1991)MidlandValleyofScotland( Wellman and Richardson1993) Ballytoohy Fm Upper Clew Bay complex Clare Island Ireland(Higgs and Williams 2001) Ludlow Tanezzuft and Acacus formations

MG-1 borehole Tunisia (Spina and Vecoli 2009)Thespecieshas alsobeen recordedfrom Early Devonian(Lochkovian)

strata of the Tawil Fm in Saudi Arabia (Steemans et al 2007)





53 Envelope-enclosed unfused cryptospore tetrads

Permanent tetrads constituted of discrete spores with line of attachment clearly visible on the tetrad surface Unfused tetrads havebeen reported enclosed within laevigate or variously ornamentedenvelopes (eg Burgess 1991 Burgess and Richardson 1991 Gray

1985 Richardson 1988)

Genus Velatitetras Burgess 1991Type species Velatitetras laevigata Burgess 1991

Velatitetras laevigata Burgess 1991Plate I 12 Plate III 13ndash19 Plate IV 8

1985 lsquoObligate tetrad with smooth perisporersquo Gray p 177 pl 11047297gs 2 and 3

1988 lsquoSpore tetrad with smooth or possibly degraded reticulateenvelopersquo Gray p 355 pl 1 1047297g 3

1988 Nodospora sp B mdash Richardson p 941991 Velatitetras laevigata Burgess p 583 pl 1 1047297gs 5 and 6

Dimensions 34 (40) 50 μ m (10 specimens measured)Occurrence Eastern Anticosti Becscie Fm Fox Point Mb sample

EB308 Valga-10drillcore Saldus FmBrocēni MbsampleVA38(3132)Remarks and comparisons The average diameter of V laevigata in

Burgess (1991) description is 27 μ m The present specimens arelarger as are those described by Wellman (1996) from the type areaof the Caradoc Series and the North African specimens attributed toNodospora sp B by Richardson (1988)

Previous records Velatitetras laevigata has been mainly documen-ted from Ashgill (late KatianndashHirnantian) to Llandovery worldwideOnly one record from Lochkovian of Saudi Arabia has been recentlypublished (Steemans et al 2007) Some relevant occurrences arereported here as follows

CaradocndashAshgill Bedinan Fm Turkey (Steemans et al 1996)Ashgill Oostduinkerke borehole Brabant Massif Belgium (Steemans2001) Kalpintag Fm Southern Xinjiang China (Wang et al 1997)Elkhorn Fm and Preachville Mb Drakes Fm Kentucky USA(Gray 1985 1988) Upper CaradocndashLower Telychian subsurfaceof northeastern Libya (Richardson 1988) Latest Ordovicianndashlate

Rhuddanian from Scrach to Crychan Fm SW Wales (Burgess 1991)Llandovery Salar del Rincoacuten Fm Puna Region Argentina (Rubinsteinand Vaccari 2004) Tiangua Fm Serra Grande Group (Le Heacuterisseacuteet al 2001) and Villa Maria Fm Paranaacute Basin (Mizusaki et al2002) Brazi l Itacurubi Group 269-R1 borehole ParanaacuteBasin Paraguay (Mendlowicz Mauller et al 2004) Qusaiba Mb

Nuayyim-2 and Hawiyah-152 boreholes Saudi Arabia (Steemans etal 2000 Wellman et al 2000) Wenlock Fish Bed Fm Haghaw Hillsinlier and Logan Fm Lesmahagow Inlier Scotland (Wellman andRichardson 1993) Ballytoohy Fm Clare Island Ireland (Higgsand Williams 2001) LudlowndashPridoli Dadas Fm Turkey (Steemanset al 1996) Tanezzuft and Acacus formations Tunisia (Spina andVecoli 2009)

Velatitetras rugosa (Strother and Traverse) Steemans Le Heacuterisseacuteand Bozdogan 1996

Plate I 11 Plate IV 1915ndash17

1979 Nodospora rugosa Strother and Traverse 1979 p 10 pl 21047297g 21985 lsquoobligate tetrads with1047297rmly adherent rugose perisporersquo Gray

p 176 pl 1 1047297

gs 10ndash

131985 lsquoobligate tetrads with rugose perisporersquo Gray p 177 pl 21047297g191988 Nodospora rugosa Strother and Traverse Richardson p 94

pl 18 1047297gs 3ndash6 pl 19 1047297gs 2ndash41991 Velatitetras rugulata Burgess 1991 pp 585ndash586 pl 1 1047297g 101996 Velatitetrasrugosa (Strother andTraverse)emend mdash Steemans

Le Heacuterisseacute and Bozdogan pp 61 63 pl 5 1047297gs 11 and 12 text-1047297g 5

Dimensions 36 (40) 45 μ m (10 specimens measured)Occurrence Western Anticosti Ellis Bay Fm sample EB42 Eastern

Anticosti Ellis Bay Fm Lousy Cove Mb sample EB346 Becscie FmFox Point Member sample EB314 Valga-10 drillcore Saldus FmBrocēni Mb sample VA38 (3132)

Previous records This species has been already recordedin Anticosti Island by Richardson and Ausich (2007 Velleda Mbof Ellis Bay Fm Hirnantian) Other occurrences are as followsCaradocndashHirnantian Bedinan Fm Turkey (Steemans et al 1996)Ashgill Oostduinkerke borehole Brabant Massif Belgium (Steemans2001) AshgillndashEarly Llandovery Hlasna Treban Czechoslovakia(Vaacutevrdovaacute 1988 1989) Early Llandovery (Rhuddanian) Bronydd

Plate III Cryptospores from Valga-10 borehole (scale bar 20 μ m EF England Finder)

1ndash3 5 Tetrahedraletes medinensis (Strother and Traverse) Wellman and Richardson 1993 (1 slide VA38 3132[2] EF N293 2 slide VA40 3200[3] EF F274 3 slide VA383132[2] EF G223 5 slide VA38 3132[2] EF L18)

4 7 10 Cheilotetras caledonica Wellman and Richardson 1993 (4 slide VA38 3132[2] EF N40-1 7 slide VA38 3132[1] EF O28-2 10 slide VA38 3132[2] EF M262)6 Gneudnaspora divellomedia (Tchibrikova) Balme 1988 var minor Breuer Al-Ghazi Al-Ruwaili Higgs Steemans and Wellman 2007 (slide VA40 3200[1] EF N37-3)8 Abditusdyadus laevigatus Wellman and Richardson 1996 (slide VA38 3132[2] EF K38-2)

9 Sphaerasaccus glabellus Steemans Higgs and Wellman 2000 (slide VA38 3132 [2] EF N25-3)11 12 Rimosotetras problematica Burgess 1991 (12 slide VA38 3132[2] EF L182)13ndash19 Velatitetras laevigata Burgess 1991 (13 slide VA38 3132[1] EF L51 14 slide VA38 3132[2] EF O473 15 slide VA38 3132[2] EF K433 16 slide VA38 3132[2] EF

K473 17 slide VA38 3132[2] EF K302 18 slide VA38 3132[2] EF E352 19 slide VA38 3132[2] EF R22)20 Envelope-enclosed alete cryptospore (slide VA38 3132[2] EF O243)

Plate IV Cryptospores from Valga-10 borehole (VA) and from Anticosti Island (EB) (scale bar 20 μ m EF England Finder) (see on page 86)

1 9 15 16 17 Velatitetras rugosa (Strother and Traverse) Steemans Le Heacuterisseacute and Bozdogan 1996 (1 slide VA38 3132[2] EF M29 9 slide VA 38 3132[2] L44 15 slideEB314[1] EF P224 16 slide EB42[1] EF Q24 17 slide VA38 3132[2] H333)

2 3 Segestrespora rugosa (Johnson) Burgess 1991 (2 slide VA38 3132[2] EF P22 3 slide VA38 3132[2] EF J392)4 12 14 Tetrahedraletes medinensis (Strother and Traverse) Wellman and Richardson 1993 (4 slide VA38 3132[2] EF J25 12 slide VA38 3132[2] EF J184 14 slide VA38

3132[2] EF Q344)5 7 Dyadospora murusattenuata (Strother and Traverse) Burgess and Richardson 1991 (5 slide VA40 3200[1] EF Q45 12 slide VA38 3132[2] EF O401)6 Hispanaediscus sp (Cramer) Burgess and Richardson 1991 (slide VA42 3290[1] EF O132)8 Velatitetras laevigata Burgess 1991 (slide VA38 3132[2] P38)

10 19 Pseudodyadospora laevigata Johnson 1985 (10 slide VA38 3132[2] EF O473 19 slide VA38 3132[2] EF N25)11 Segestrespora laevigata Burgess 1991 (slide VA38 3132[2] EF K422)13 20 Rimosotetras problematica Burgess 1991 (13 slide EB318[1] EF P292 20 slide VA38 3132[2] EF M37)18 Segestrespora membranifera (Johnson) Burgess 1991 [slide VA38 3132[2] EF J333)





Fm type Llandovery area South-West Wales (Burgess 1991)Llandovery Tuscarora Fm Pennsylvania ( Johnson 1985 Strotherand Traverse 1979) Medina Group Niagara Gorge New York(Miller and Eames 1982) Jupiter Fm Quebec Canada and PoorValley Ridge Mb Virginia USA (Gray 1985) Salar del Rincoacuten

Fm Puna Region Argentina (Rubinstein and Vaccari 2004) TianguaFm Serra Grande Group Brazil (Le Heacuterisseacute et al 2001) VillaMaria Fm Paranaacute Basin Brazil (Mizusaki et al 2002) ItacurubiGroup 269-R1 borehole Paranaacute Basin Paraguay (MendlowiczMauller et al 2004) Qusaiba Mb of Qalibah Fm Saudi Arabia

Plate III





Plate IV (caption on page 84)





(Steemans et al 2000 Wellman et al 2000) LlandoveryndashLudlowAllenport Pennsylvania USA (Beck and Strother 2008) Ludlowuppermost Pitinga Fm Urubu River area western Amazon BasinBrazil (Steemans et al 2008) LudlowndashLochkovian Tanezzuft AcacusTadrart formations Ghadamis basin MG-1 borehole Tunisia (Spinaand Vecoli 2009)

Velatitetras sp 1Pl I 3 6 8

Description Probable cryptospore tetrad composed by tightly-adherent sub-circular spores Line of attachment is not clearly evidenton the tetrad surface the junction seems to be entirely fusedEnvelope laevigate and folded

Dimensions 50 52 54 μ m (three specimens measured)Occurrence Valga-10 drillcore Saldus Fm Brocēni Mb sample

VA38 (3132)

Velatitetras sp 2Pl I 4 7

Description Probable cryptospore tetrad characterized by tightly-adherent sub-circular spores Line of attachment at the junctionsbetween spores is not clearly distinguishable Envelope ornamentedby an irregular reticulum formed by low muri about 3ndash4 μ m apart

Dimensions 39 40 μ m (two specimen measured)Occurrence Valga-10 drillcore Saldus Fm Brocēni Mb sample

VA38 (3132)

54 Envelope-enclosed fused cryptospore dyads

Group characterized by pseudodyads enclosed within a variouslyornamented envelope

Genus Segestrespora Burgess 1991Type species Segestrespora membranifera (Johnson) Burgess 1991

Segestrespora membranifera (Johnson) Burgess 1991Plate IV 18

pp 1985 Dyadospora membranifera Johnson p 336 pl7 1047297gs 1ndash35 and 6 non pl 7 1047297g 7

1988 Pseudodyadospora cf Dyadospora membranifera Richardsonp 94 pl 15 1047297gs 1ndash2

1991 Segestrespora (Dyadospora) membranifera (Johnson 1985)comb nov mdash Burgess p 588 pl 2 1047297gs 2ndash5

Dimensions 30 36 μ m (two specimens measured)Occurrence Valga-10 drillcore Saldus Fm Brocēni Mb sample

VA38 (3132)Previous records CaradocndashHirnantian Bedinan Fm Turkey (Stee-

mans et al 1996) Ashgill Oostduinkerke borehole Brabant MassifBelgium (Steemans2001) HirnantianndashLlandovery(Rhuddanian) fromScach to Bronydd Fm type Llandovery area South-West Wales(Burgess 1991) Early Llandovery Hlasna Treban Czechoslovakia(Vaacutevrdovaacute 1989) LlandoveryQusaiba Mbof QalibahFm Saudi Arabia(Steemans et al 2000 Wellman et al 2000) Villa Maria Fm ParanaacuteBasin Brazil (Mizusaki et al 2002) Salar del Rincoacuten Fm Puna RegionArgentina (Rubinstein and Vaccari 2004)

Segestrespora laevigata Burgess 1991Plate IV 11

pp 1985 ldquoDyadospora membraniferardquo Johnson p 336 pl 7 1047297g 7non pl 7 1047297gs 1ndash3 5 6

1991 Segestrespora laevigata Burgess 1991 p 589 pl 2 1047297g 1

Dimensions 33ndash38 μ m (two specimens measured)Occurrence Valga-10 drillcore Saldus Fm Brocēni Mb sample

VA38 (3132)Previous records CaradocndashHirnantian Bedinan Fm Turkey

(Steemans et al 1996) Ashgill Oostduinkerke boreholeBrabant Massif Belgium (Steemans 2001) HirnantianndashLlandovery

(Rhuddanian) from Scach to Bronydd Fm type Llandovery areaSouth-West Wales (Burgess 1991) Llandovery Qusaiba Mb of Qalibah Fm Saudi Arabia (Steemans et al 2000 Wellman et al2000) Tiangua Fm Serra Grande Group Brazil (Le Heacuterisseacute et al2001) Villa Maria Fm Paranaacute Basin Brazil (Mizusaki et al2002) Itacurubi Group 269-R1 borehole Paranaacute Basin Paraguay(Mendlowicz Mauller et al 2004)

Segestrespora rugosa (Johnson) Burgess 1991Plate IV 23

1985 Pseudodyadospora rugosa Johnson p 337 pl 8 1047297gs 2ndash61988 Pseudodyadospora sp C Richardson p 951991 Segestrespora (Pseudodyadospora) rugosa (Johnson 1985) mdash

Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9

Dimensions 35 (40) 42 μ m (seven specimens measured)Occurrence Western Anticosti Vaureal Fm sample EB94Valga-10

drillcore Jelgava Fm sample VA46 (3457) Kuili Fm sample VA42(3290) Kuldiga Fm Edole Mb sample VA40 (3200) Saldus FmBrocēni Mb sample VA38 (3132)

Previous records This species has a wide geographic distributionstratigraphically ranging from the Upper Ordovician to the LlandoveryCaradocCaradoc typearea southern Britain (Wellman1996) CaradocndashAshgill (SandbianndashHirnantian) borehole E1-1 northeastern Libya(Richardson 1988) Ashgill (late KatianndashHirnantian) Oostduinkerkeborehole Brabant Massif Belgium (Steemans 2001) Latest Katianndash

Hirnantian Bedinan Fm southeastern Turkey (Steemans et al 1996)Hirnantian Kalpintag Fm southern Xinjiang China HirnantianndashearlyLlandovery (Rhuddanian) Llandovery type area SW Wales (Burgess1991) Early Llandovery (Rhuddanian) Qusaiba Member Qalibah FmSaudi Arabia (Steemans et al 2000 Wellman et al 2000) MedinaGroup Niagara Gorge Lewiston New York USA (Miller and Eames1982) Llandovery Tiangua Fm Serra Grande Group (Le Heacuterisseacute et al2001) and Villa Maria Fm Paranaacute Basin Brazil (Mizusaki et al 2002)Itacurubi Group 269-R1 borehole Paranaacute Basin Paraguay (MendlowiczMauller et al 2004)

55 Envelope-enclosed unfused cryptospore dyads

Group characterized by true dyads enclosed within a variouslyornamented envelope

Genus Abditusdyadus Wellman and Richardson 1996Type species Abditusdyadus histosus Wellman and Richardson

1996

Abditusdyadus laevigatus Wellman and Richardson 1996Plate III 8

1996 Abditusdyadus laevigatus Wellman and Richardson p 68 pl4 1047297gs 7ndash8

Dimensions 56 μ m (one specimen measured)Occurrence Valga-10 drillcore Saldus Fm Brocēni Mb sample

VA38 (3132)Previous records Ashgill (late KatianndashHirnantian) Oostduinkerke

borehole Brabant Massif Belgium (Steemans 2001) Early Llandovery(Rhuddanian) Qusaiba Mb Qalibah Fm Saudi Arabia (Steemans et al2000 Wellman et al 2000) Llandovery Villa Maria Fm Paranaacute





Basin Brazil (Mizusaki et al 2002) Llandovery (Telychian)ndashLudlow(Ludfordian) from Keefer Fmto Wills Creek FmPennsylavania USA(Beck and Strother 2008) Lochkovian Lower Old Red SandstoneLorne Scotland (Wellman and Richardson 1996)

56 Naked fused cryptospore dyads (pseudodyads)

Group comprising dyads which are not enclosed within anenvelope and which does not show a recognizable line of attachment

Genus Pseudodyadospora Johnson 1985Type species Pseudodyadospora laevigata Johnson 1985Pseudodyadospora laevigata Johnson 1985Plate IV 1019

1985 Pseudodyadospora laevigata Johnson p 337 pl 7 1047297g11pl81047297g 7

Dimensions 31 (34) 54 μ m (18 specimens measured)Occurrence Western Anticosti Vaureal Fm samples EB2 EB94

Ellis Bay Fm member 2 sample EB132 member 6 sample EB42Eastern Anticosti Becscie Fm Fox Point Mb samples EB309 EB314EB316 Valga-10 drillcore Jelgava Fm samples VA46 (3457) VA44(3355) Kuili Fm sample VA42 (3290) Kuldiga Fm EdoleMb sample VA40 (3200) Saldus Fm Brocēni Mb sample VA38(3132)

Previous records P laevigata ranges from Late Ordovician to EarlyDevonian although it is particularly abundant in AshgillndashLlandoverystrata Richardson and Ausich (2007) reported the record of P laevigata

from the Velleda Mb of the Ellis Bay Fm (Hirnantian)Other records are as follows Ashgill Kalpintag Fm Southern

Xinjiang China (Wang et al 1997) Oostduinkerke borehole BrabantMassif Belgium (Steemans 2001) Bedinan Fm Turkey (Steemanset al 1996) AshgillndashEarly Llandovery Hlasna Treban Czech Republic(Vaacutevrdovaacute 1988 1989) Early Llandovery (Rhuddanian) Qalibah FmQusaiba Mb Saudi Arabia (Steemans et al 2000 Wellman et al2000) Llandovery Medina Group Niagara Gorge New York (Millerand Eames 1982) Tuscarora Fm USA ( Johnson 1985) Villa MariaFm Paranaacute Basin Brazil (Mizusaki et al 2002) Itacurubi Group269-R1 borehole Paranaacute Basin Paraguay (Mendlowicz Mauller et al2004) WenlockndashLudlow Stonehaven Group Scotland (Marshall1991) LudlowndashPridoli Dadas Fm Turkey (Steemans et al 1996)LudlowndashLochkovian Tanezzuft Acacus Tadrart formations Ghada-mes Basin MG-1 borehole southern Tunisia (Spina and Vecoli 2009)Lochkovian Clam Bank Fm Western Newfoundland Canada (Burdenet al 2002)

57 Unfused naked cryptospore dyads (true dyads)

This group comprises dyads which readily separate into two aletespores (Burgess 1991)

Genus Dyadospora Strother andTraverse1979 emendBurgess andRichardson 1991

Type species Dyadospora murusattenuata Strother and Traverseemend Burgess and Richardson 1991

Dyadospora murusattenuata Strother and Traverse emend Burgessand Richardson 1991

Plate IV 57

1979 Dyadospora murusattenuata Strother and Traverse 1979p 15 pl 3 1047297gs 9 10

1989 lsquoDyadrsquo Turnau and Jakubowska 1047297g 261991 Dyadospora murusattenuata Strother and Traverse emend mdash

Burgess and Richardson p 614 pl 2 1047297gs 7 9

Dimensions 30 (35) 44 μ m (16 specimens measured)Occurrence Western Anticosti Ellis Bay Fm member 1 samples

EB109 EB111 Eastern Anticosti Becscie Fm Fox Point Mb samplesEB316 EB318 Valga-10drillcore JelgavaFm sample VA46(3457) KuiliFmsampleVA42(3290) Saldus FmBrocēni Mb sampleVA38(3132)

Previous records This species has a long stratigraphic range

(Ordovicianndash

Devonian) and a common worldwide distribution Someselected Ordovician occurrences are as followsCaradoc Capillas Fm Sierras subandinas NW Argentina (Rubin-

stein 2005) Upper CaradocndashUpper Ashgill E1-1 borehole Libya(Richardson 1988) Ashgill Kosov Fm HlasnaTreban Czech Republic(Vaacutevrdovaacute 1988 1989) Preachersville Mb Drakes Fm Ohio USA(Taylor 1995) Kalpintag Fm Southern Xinjiang China (Wang et al1997) Hirnantian Oostduinkerke borehole Brabant Massif Belgium(Steemans 2001) Bedinan Fm Turkey (Steemans et al 1996)Hirnantian Salar del Rincoacuten Fm Puna Region Argentina (Rubinsteinand Vaccari 2004) Llandovery Qusaiba Mb Qalibah Fm SaudiArabia (Steemans et al 2000)

58 Hilate cryptospores

These forms consist of alete monad spores hemispherical inlongitudinal view with a hilum and an equatorial thickening Theexine may be laevigate or variously ornamented Closely similarmonads have been observed partly united at the contact area as loosedyads

Genus Gneudnaspora Balme emend Breuer Al-Ghazi Al-RuwailiHiggs Steemans and Wellman 2007

1988 Gneudnaspora Balme p 1241991 Laevolancis Burgess and Richardson pp 606ndash6072007 Gneudnaspora Balme emend mdash Breuer Al-Ghazi Al-Ruwaili

Higgs Stemans Wellman pp 47ndash48Type species Gneudnaspora kernickii Balme 1988Remark Breuer et al (2007) considered that Gneudnaspora

Balme 1988 needed a revision and emendation because Balme(1988) had misinterpreted the presence of mostly irregular ldquoscarsrdquoon the proximal face of the spore as trilete or monlete markingsFollowing this emendation Laevolancis Burgess and Richardson1991 would become undistinguishable from hence junior synonymof Gneudnaspora

Gneudnaspora divellomedia (Chibrikova) Balme 1988 var minor

Breuer Al-Ghazi Al-Ruwaili Higgs Steemans and Wellman 2007Plate III 6

1959 Archaeozonotriletes divellomedium Chibrickova p 65 pl 91047297g 4

pp 1988 Gneudnaspora divellomedium (Chibrickova) Balme

p 125 pl 3 1047297gs 1ndash71991 Laevolancis divellomedia (Chibrikova) Burgess and Richard-

son p 607 pl 2 1047297gs 4 and 6 cum syn

2007 Gneudnaspora divellomedia (Chibrikova) Balme 1988 varminor Breuer et al p 48 pl 3 1047297gs 10ndash16

Dimensions 23 (37) 42 μ m (seven specimens measured)Occurrence Western Anticosti Ellis Bay Fm member 1 sample

EB111 member 6 sample EB42 Eastern Anticosti Becscie Fm FoxPoint Mb samples EB314 EB318 and EB320 Valga-10 drillcore

Jelgava Fm sample VA46 (3457) Kuili Fm sample VA42 (3290)Kuldiga Fm Edole Mb sampleVA40 (3200) Saldus Fm Brocēni Mbsample VA38 (3132)

Remarks Breuer et al (2007) introduced the variety ldquominor rdquo to

accommodate small (28ndash34 μ m in diameter) specimens of Gneudnas- pora divellomedia which always bear visible ldquoscarsrdquo on the proximalsurface (hilum) The present specimens appear to be attributable to





this variety even if they are slightly larger on average Severalprevious records of G divellomedia also show slightly larger dimen-sional ranges mostly falling in between the average diameters of thedivellomedia (54 μ m) and minor (31 μ m) morphotypes Accordinglywe include in the present variety the portion of the populations of G divellomedia recorded in previous works which is consistent with

the size ranges provided in the original diagnosis of Breuer et al(2007)Previous records G divellomedia var minor has a worldwide

distribution from Ordovician to Devonian Richardson and Ausich(2007) reported the occurrence of this form in the Velleda Mb of theEllis Bay Fm in eastern Anticosti

Ordovician Lower Ordovician Dapingian()ndashDarriwillianZanjoacuten and Labrado formations Sierras Subandinas northwesternArgentina (Rubinstein et al 2010 2011) Ashgill KalpintagFm Southern Xinjiang China (Wang et al 1997) HirnantianOostduinkerke borehole Brabant Massif Belgium (Steemans 2001)

Silurian Llandovery of subsurface central Saudi Arabia (Steemanset al 2000 Wellman et al 2000) type Wenlock area ShropshireEngland (Burgess and Richardson 1991) Ludlow (Ludfordian) Argiles

Principales Formation northern Ghadames Basin southern Tunisia(Vecoli et al 2009)Earliest Devonian (Lochkovian) Lorne Scotland (Wellman and

Richardson 1996) EarlyndashMiddle Devonian (PragianndashGivetian) Jauf and Jubah formations northern Saudi Arabia (Breuer et al 2007)

Genus Hispanaediscus (Cramer) Burgess and Richardson 1991Type species Hispanaediscus verrucatus Cramer 1966

Hispanaediscus spPlate IV 6

Description Amb subrounded distal exine ornamented withclosely spaced verrucae in part coalescing to form muri 1ndash2 μ mwide and about 1 μ m high Hilum very thin and transparent

Dimensions 34 μ m (one specimen measured)Occurrence Valga-10 drillcore Kuili Fm sample VA42 (3290)

59 Envelope-enclosed monads This group consists of laevigate orvariously ornamented monad enclosed within a laevigateor variouslyornamented envelope

Genus Sphaerasaccus Steemans Higgs and Wellman 2000Type species Sphaerasaccus glabellus SteemansHiggs and Wellman

2000 p 102ndash103 Pl 4 1047297gs andashb

Sphaerasaccus glabellus Steemans Higgs and Wellman 2000Plate III 9

1996 lsquoLaevigate monads enclosed within a laevigate envelopersquo

Wellman p 118 Pl 3 1047297gs 18ndash202000 Sphaerasaccus glabellus Steemans Higgs and Wellman 2000

p 102ndash103 Pl4 1047297gs andashb


VA38 (3132)Previous records Caradoc Acton Scott Beds Caradoc type area

Southern Britain UK Hirnantian upper member of the Salar delRincoacutenFm (Rubinstein and Vaccari 2004) Llandovery (Rhuddanian)Qusaiba Mb of Qalibah Fm Saudi Arabia (Steemans et al 2000) LateLudlow uppermost Pitinga Fm Urubu River Area Northern Brazil(Steemans et al 2008)

Alete cryptosporePlate III 20

Description Sub-circularaletemonadornamented by closely spacedanastomosing rugulae The spore appears as completely enclosed by anunsculptured laevigate envelope


VA38 (3132)

6 Discussion and conclusions

Wellman (1996) noted that cryptospore assemblages display littleevolutionary changes during the Darriwilian (Middle Ordovician)ndashmid-Telychian (late Llandovery early Silurian) interval and subse-quent work (Steemans 2000 Wellman and Gray 2000) hassupported this observation Strother (2000) considered that taxo-nomic overlap between Caradoc and Ashgill occurrences might be duemore to lumping than to true similarity In any case it appears thatmorphological and evolutionary innovation in spore morphology wasrather limited during this interval For example no signi1047297cant plantextinction events are known from the Hirnantian (Gray 1984 1988Steemans 2000) If early embryophytic vegetation was not signi1047297-

cantly affected by the Late Ordovician glaciation it could indicate thatit possessed a large tolerance to climatic variation In her proposedsubdivision of land plant evolution into a series of major evolutionaryevents of planetary signi1047297cance Gray (1993) attributed the earlyDarriwilian (Middle Ordovician)ndashto late early Silurian interval to theldquoEoembryophytic Evolutionary Levelrdquo of plant evolution Stratigra-phically this corresponded to her previously established (Gray 19841985) Microfossil Assemblage (MA) Zone I extended downward toinclude the basal Llanvirn (basal Darriwilian) The concept of Eoembryophytic was established by Gray (1993) on the basis of thedominance in the spore record of nondissociating spore tetrads intetrahedral con1047297guration The observed homogeneity of cryptosporeassemblages throughout the mid-Ordovicianndashearly Silurian timeinterval prevents the establishment of cryptospore-based biostrati-graphic schemes of high stratigraphic resolution The presentlyavailable cryptospore zonations include this time interval in onesingle zone (eg the already cited MA1 Zone of Gray 1985 theDyadospora murusattenuatandashD murusdensa Assemblage Zone of Richardson and Edwards 1989 or the time-equivalent Velatitetras

rugulatandashDyadospora murusdensa Assemblage Biozone of Richardson1988 1996b) while subdivision at the subzone level are based onminornot well constrained andoftenlocal or not imprecisely de1047297nedchanges in cryptospore assemblages (eg the subdivisions proposedin Richardson 1996b) Gray (1988) suggested that increase incryptospore size might be a useful trend for determining the strat-igraphical position within Microfossil Assemblage Zone I noting thatthe average diameter of Ordovician tetrads approached 30 μ m whilemid-late Silurian cryptospores displayed average diameters close to50 μ m However the validity of this method has never been tested in

subsequent works In our Late Ordovician assemblages tetrads showconsiderable variation in size range frequently reaching up to 50 μ min diameter (eg specimens of Velatitetras) and there is no apparentcorrelation between size and stratigraphic position The assemblagesdescribed herein are precisely dated by means of independent bio-stratigraphic data (chitinozoans) and are also stratigraphically con-strained by chemostratigraphic (HICE)arguments Even if the numberof cryptospore-producing samples is not large a trend highlightingincrease in diversity from upper Katian to upper Hirnantian strata isclearly delineated by our dataset of crytpospore stratigraphic dis-tribution (Tables 2 and 3) Both in Anticosti and in Estonia the mostdiverse and also the richest samples come from upper Hirnantianstrata However whether this trend re1047298ects a real diversi1047297cation of the palaeo1047298ora or is a sampling effect of increased input from land-

derived clastic sedimentation is dif 1047297cult to ascertain In bothhypotheses clearly the Late Ordovician glaciation seems not to affectthe biodiversity of cryptospores as it has been noted before (eg





Steemans et al 2010 Wellman 1996) The diversity trend observedis thus in complete agreement with the cryptospore biodiversitycurves compiled by Steemans (2000) and Strother (2000) Interest-ingly recent investigations have shown that the Hirnantian glaciation

did not cause signi1047297cant extinctions also in microphytoplankton(acritarchs) assemblages worldwide (Delabroye et al 2011 Vecoli2008)

Comparison of our Laurentian and Baltican material with previ-ously described cryptospore suites from worldwide localities (Fig 3Table 3) also shows that no signi1047297cant provincialism can be detectedon the basis of plant microremains for the Late Ordovician to earlySilurian time interval It should be noted that up to now the mostrecent available syntheses of Ordovician cryptospore palaeogeogra-phical distribution (Edwards and Wellman 2001 Steemans et al2010) did not report the presence of cryptospores in Baltica

We have attempted to quantify the taxonomic similarity betweenour assemblages and previously published assemblages of similar agefrom selected localities (Tables 3 and 4) The similarity has beencalculated by the ldquoshared species between two given assemblagesrdquoldquototal species in the two assemblagesrdquo ratio Only assemblagescontaining at least ten taxa have been considered for this comparisonThe results are discussed here on the basis of currently acceptedpalaeogeographic reconstructions (Fig 3 Table 4) The Anticostiand Valga assemblages described in this paper have about 50 of species in common Relatively high similarities (about 30 of sharedspecies) are evidenced between the AnticostindashValga assemblages andassemblages from Avalonian localities (southern Britain and south-west Wales) similarity with the Belgian assemblage described bySteemans (2001) is slightly lower (ca 25) Based on these resultsone could observe that taxonomic similarities are generally higherbetween palaeogeographical domains that are geographically closeseparated by narrow oceans Generally speaking similarity withGondwanan localities (separated by wide oceanic basins) is lowalthough our Laurentian and Estonian assemblages do share a

2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3

Fig 3 Location of reported Late Ordovician (KatianndashHirnantian) cryptosporeassemblages plotted on palaeogeographical reconstruction obtained using BugPlatesoftware (Torsvik 2009) at 445 My 1 Valga-10 drillcore Estonia (this study)2 Anticosti Island (this study) 3 Southwest Wales(Burgess 1991) 4 Southern Britain(Wellman 1996) 5 Belgium (Steemans 2001) 6 Southeastern Turkey (Steemanset al 1996) 7 Saudi Arabia (Steemans et al 2009) 8 Northeast Libya (Richardson1988) 9 Northwest Argentina (Rubinstein 2005 Rubinstein and Vaccari 2004)10 Czech Republic (Vavrdovaacute 1988 1989) 11 South China (Wang et al 1997)12 USA (Gray 1988) 13 Canada (Gray 1988) 14 Sweden (Gray 1988) 15 South

Africa (Gray et al 1986) 16 North China (Yin and He 2000) 17 Australia (Foster andWilliams 1991) 18 USA (Strother 1991) 19 Western Libya (Gray et al 1982)

Table 4

Similarity index among cryptospore assemblages (percentage of shared species ontotal species) from the study localities versus other selected localities Numbers referto localities as in Fig 3 and Table 3

1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3

Comparison of cryptospore assemblages fromthe studylocalities(1 Valga2 Anticosti)and other published localities (3 SW Wales 4 Southern Britain 5 Belgium 6 SETurkey 7 Saudi Arabia 8 NE Libya 9 NW Argentina 10 Czech Republic 11 China)Numbers refer to map of Fig 3 Only assemblages containing a signi1047297cant number of cryptospores have been considered

Miospores 1 2 3 4 5 6 7 8 9 10 11

Abditusdyadus laevigatus x x Abditusdyadus laevigatus x Abditusdyadus histosus x Ambitisporites avitus x Ambitisporites sp x Aneurospora sp x Archaeozonotriletes chulus xCheilotetras caledonica xCheilotetras sp x xCheilotetras sp A xCheilotetras sp 1 xCheilotetras sp 2 x xChelinospora sp xDyadospora murusattenuata x x x x x x xDyadospora murusdensa x x xcf Dyadospora sp B xEnvelope-enclosed alete cryptospore xGneudnaspora chibrikovae xGneudnaspora divellomedia x x x x x xHispanaediscus sp xHispanaediscus sp A xImperfectotriletes patinatus x xImperfectotriletes vavrdovae x x x x x xLeiotriletes sp xNaked apiculate monad xNaked apiculate tetrad xNaked laevigate monad xNodospora sp xPseudodyadospora laevigata x x x x x x xPseudodyadospora cf laevigata xPseudodyadospora petasus x x xPseudodyadospora sp A xPseudodyadospora sp xReticulate monad x

Rimosotetras problematica x x x x x xSegestrespora burgessii xSegestrespora laevigata x x x x xSegestrespora membranifera x x x x x x xSegestrespora rugosa x x x x x x xSegestrespora rugosa x xSegestrespora sp A x x xSegestrespora sp C xSphaerasaccus glabellus x x xStegambiquadrella contenta xSynorisporites sp xTetrad type cf Nodospora sp D xTetrahedraletes grayae x x x xTetrahedraletes medinensis x x x x x x x x x x xVelatitetras anatoliensis x xVelatitetras laevigata x x x x x x x xVelatitetras reticulata x

Velatitetras retimembrana x x xVelatitetras rugosa x x x x x x xVelatitetras sp A x xVelatitetras sp 1 xVelatitetras sp 2 xVestidusdyadus qalibahinus x





signi1047297cant number of taxa with Turkey (peri-Gondwana assemblagedescribed by Steemans et al 1996) and northern China (showed as aseparate terrane at near-equatorial palaeolatitudes close to the marginof Gondwana in the reconstruction by Torsvik 2009 assemblagedescribed by Wang et al 1997)

It is evident that these results should be considered only as

preliminary and much more data will be needed before a morecomplete picture of Ordovician phytogeography as well as of earlyplant communities response to climatic variation is 1047297nally achievedIn this respect the present work by adding some more details on thetaxonomic diversity of cryptospores from two well-dated localitieswhich were previously under-investigated should be considered as acontribution towards such a goal

Acknowledgements

We would like to thank Philippe Recourt Sylvie Reacutegnier andEveline Rogie (Geacuteosystegravemes Lille) for technical assistance and helpwith SEM work Thisworkhasbeen conducted aspartof the PhD workof AD at the Laboratoire Geacuteosystegraveme of Universiteacute Lille 1 under the

supervision of MV Research costs including travel expenses weresupported by an ECLIPSE II (ldquoTerrestrializationrdquo) grant awarded byINSUCNRS to MV Additionally MV Acknowledges a ldquoVisitingProfessorrdquo grant (2010) from the University of Cagliari Italy wherethis work was completed AS is grateful to Dr F Guzzetti (IRPI CNRItaly) Prof R Bonaurio (Faculty of Agriculture Perugia UniversityItaly) and Dr A Boscherini (Geological Survey Umbria Region Italy)OH acknowledges support from the Estonian Science Foundationgrant 7640 The constructive remarks of two anonymous referees areacknowledged and contributed to improve the paper

This study is a contributionto theANR project ldquoGlobal perspectiveson the Terrestrialization processrdquo No 2010 BLAN 607 01 (coordinatorM Vecoli)

References

Achab A Paris F 2007 The Ordovician chitinozoan biodiversi1047297cation and its leadingfactors Palaeogeography Palaeoclimatology Palaeoecology 245 5ndash19

AchabA Asselin EDesrochersA Riva JFarley C2011 Chitinozoan contributiontothe development of a new Upper Ordovician stratigraphic framework for AnticostiIsland Geological Society of America Bulletin 123 186ndash205

Balme BE 1988 Miospores from Late Devonian (Early Frasnian) strata CarnarvonBasin Western Australia Palaumleontographica Abt B 209 (4) 109ndash166

Barnes CR 1988 Stratigraphy and Palaeontology of the OrdovicianndashSilurian boundaryinterval Anticosti Island Quebec Canada In Cocks LRM Rickards RB (Eds) AGlobal Analysis of the OrdovicianndashSilurian Boundary Bulletin of the BritishMuseum Natural History Geology 43 pp 195ndash219

Beck JH Strother PK 2001 Silurian spores and cryptospores from the Arisaig GroupNova Scotia Canada Palynology 25 127ndash177

Beck JH Strother PK 2008 Miospores and cryptospores from the Silurian section atAllenport Pennsylvania USA Journal of Palaeontology 82 857ndash883

Bergstroumlm SM Chen X Gutierrez-Marco JC Dronov A 2009 The new chronostrati-graphic classi1047297cation of the Ordovician System and its relations to major regional

series and stages and to δ13

C chemostratigraphy Lethaia 42 97ndash

107Brenchley PJ Carden GAF Hints L Kaljo D Marshall JD Martma T Meidla TNotildelvak J 2003 High-resolution stable isotope stratigraphy of Upper Ordoviciansequences constraints on timing of bioevents and environmental changesassociated with mass extinction and glaciation Geological Society of AmericaBulletin 115 89ndash104

Breuer P Al-Ghazi A Al-Ruwaili M Higgs KT Steemans P Wellman CH 2007Early to Middle Devonian miospores from northern Saudi Arabia Miospores Revuede micropaleacuteontologie 50 27ndash57

Burden ET Quinn L Nowlan GS Bailey-Nill LA 2002 Palynology andmicropaleontology of the Clam Bank Formation (Lower Devonian) of WesternNewfoundland Canada Palynology 26 185ndash215

Burgess ND 1991 Silurian cryptospores and miospores from the type Llandoveryarea southwest Wales Palaeontology 34 575ndash599

Burgess ND Richardson JB 1991 Silurian cryptospores and miospores from the typeWenlock area Shropshire England Palaeontology 34 601ndash628

BurgessNDRichardson JB 1995 Late Wenlockto early Pridolicryptospores andmiosporesfrom south and southwest Wales Great Britain Palaeontographica Abt B 236 1ndash44

Chibrickova EV 1959 Spory iz Devonskih Boleye Drevinkh otlozheni Bashkirii

Bashkirskii Filnal Institut Geologii Akademiya Nauk SSR pp 1ndash116Combaz A 1967 Un microbios du Treacutemadocien dans un sondage dHassi-Messaoud

Actes de la Socieacuteteacute Linneacuteenne de Bordeaux (B) 104 1ndash26

Copper P 2001 Reefs during the multiple crises towards the OrdovicianndashSilurianboundary Anticosti Island eastern Canada and worldwide Canadian Journal of Earth Sciences 38 153ndash171

Cramer FH 1971 Implications from mid-Palaeozoic palynofacies transgressions forrates of crustal movements especially in the Wenlockian Anals da AcademiaBrasileira de Ciencias 43 51ndash66

Cramer FH Diez MDCR 1972 North American Silurian palynofacies abd theirspatial arrangement acritarchs Palaeontographica Abt B 138 107ndash180

Delabroye A 2010 Dynamiques du phytoplancton agrave travers la limite Ordovicienndash

Silurien aux faibles paleacuteolatitudes (Laurentia Icircle dAnticosti Queacutebec Canada Baltica Valga Estonie) Compleacutements aux donneacutees des hautes latitudes glaciairesdu Gondwana Ph D Thesis Universiteacute de Lille 1 France 1ndash464

Delabroye A Vecoli M 2010The end-Ordovician glaciation and the HirnantianStagea global review and questions about late Ordovician event stratigraphy Earth-Science Reviews 98 269ndash282

Delabroye A Achab A Asselin E Copper P Desrochers A Farley C Ghienne J-F Joachimski M Loi A Munnecke A Paris F Vecoli M 2010 Toward a re1047297nedldquoHirnantian Composite Stratigraphical Modelrdquo for Laurentia Baltica and North-Gondwana IPC3 Programme amp Abstracts Book Symposium 8 The Great OrdovicianBiodiversi1047297cation Event causes andconsequences London June28ndash July3 2010 p 142

Delabroye A Vecoli M Hints O Servais T 2011 Acritarchs from the Ordovician ndash

Silurian boundary beds of the Valga-10 drill core southern Estonia (Baltica) andtheir stratigraphic and palaeobiogeographic implications Palynology 35 4ndash45

Desrochers A Farley C Achab A Asselin E Riva JR 2010 A far-1047297eld record of theend Ordovicianglaciation the Ellis Bay Formation Anticosti Island Eastern CanadaPalaeogeography Palaeoclimatology Palaeoecology 296 248ndash263

Duf 1047297eld SL 1985 Land-derived microfossils from Jupiter Formation (Upper

Llandoverian) Anticosti Island Quebec Journal of Palaeontology 59 1005ndash1010Duf 1047297eld SL Legault JA 1981 Acritarch biostratigraphy of Upper OrdovicianndashLower

Silurian rocks Anticosti Island Queacutebec preliminary results In Lespeacuterance PJ(Ed) Subcommission on Silurian Stratigraphy OrdovicianndashSilurian BoundaryWorking Group Field meeting Anticosti-Gaspeacute Queacutebec Stratigraphy andPaleontology 2 pp 91ndash99

Edwards D Wellman CH2001 Embryophyteson landthe Ordovicianto Lochkovian(Lower Devonian) record In Gensel PG Edwards D (Eds) Plants Invade theLand Columbia University Press New York pp 3ndash28

Foster CBWilliamsGE 1991Late OrdovicianndashEarly Silurian agefor theMallowaSaltof the Carribuddy Group Canning Basin Western Australia based on occurrencesof Tetrahedraletes medinensis Strother and Traverse 1979 AustralianJournal ofEarhSciences 38 223ndash228

Gray J 1984 OrdovicianndashSilurian land plants the interdependence of ecology andevolution Special Papers in Palaeontology 32 281ndash295

Gray J 1985 The microfossil record of early land plants advances in understanding of early terrestrialization 1970ndash1984 In Chaloner WG Lawson JD (Eds)Evolution and Environment in the Late Silurian and Early Devonian PhilosophicalTransactions of the Royal Society London pp 167ndash195

Gray J 1988 Land plant spores and the OrdovicianndashSilurian boundary Bulletin of theBritish Museum of Natural History (Geology) 43 351ndash358

Gray J 1993 Major Paleozoic land plant evolutionary bio-events PalaeogeographyPalaeoclimatology Palaeoecology 104 153ndash169

Gray J Boucot AJ 1971 Early Silurian spore tetrads from New York Earliest newworld evidence for vascular plants Science 173 918ndash921

Gray J Massa D Boucot AJ 1982 Caradocian land plant microfossils from LibyaGeology 10 197ndash201

Gray J Theron JH Boucot AJ 1986 Age of the Cedarberg Formation South Africaand early land plant evolution Geological Magazine 123 (4) 445ndash454

Hagstrom J 1997 Land-derived palynomorphs from the Silurian of Gotland SwedenGFF 119 301ndash316

Harris MT Sheehan PM Ainsaar L Hints L Maumlnnik P Notildelvak J Rubel M 2004Upper Ordovician sequences of western Estonia Palaeogeography Palaeoclimatol-ogy Palaeoecology 210 135ndash148

Higgs KT Williams DM 2001 A Silurian micro1047298ora from the Ballytoohy Formation(Upper Clew Bay complex) of Clare Island New Survey of Clare Island 2

Hints L Hints O Kaljo D Kiipli T Maumlnnick P Notildelvak J Paumlrnaste H 2010

Hirnantian (latest Ordovician) bio- and chemostratigraphy of the Stirnas-18 corewestern Latvia Estonian Journal of Earth Sciences 59 1ndash24 Jacobson SR Achab A 1985 Acritarch biostratigraphy of the Dicellograptus

complanatus graptolite zone from the Vaureal formation (Ashgillian) AnticostiIsland Quebec Canada Palynology 9 165ndash198

Johnson NG 1985 Early Silurian palynomorphs from the Tuscarora Formation inCentral Pennsylvanianand their paleobotanical and geological signi1047297cance Reviewof Palaeobotany and Palynology 45 307ndash360

Kaljo D Martma T Saadre T 2007 Post-Hunnebergian Ordovician carbon isotopetrend in Baltoscandia its environmental implications and some similaritieswith that of Nevada Palaeogeography Palaeoclimatology Palaeoecology 245138ndash155

Kaljo D Hints L Maumlnnik P Notildelvak J 2008 The succession of Hirnantian eventsbased on data from Baltica brachiopods chitinozoans conodonts and carbonisotopes Estonian Journal of Earth Sciences 57 197ndash218

Le Heacuterisseacute A 1989 Acritarches et kystes dalgues prasinophyceacutees du Silurien degotland Sueacutede Palaeontographia Italica 76 57ndash302

Le Heacuterisseacute A Melo JHG Quadros LP Grahn Y Steemans P 2001 Palynologicalcharacterization and dating of the Tiangua Formation Serra Grande Group

northern Brazil In Melo JHG Terra GJS (Eds) Correlaccedilacirco de SequumlecircnciasPaleozoacuteicas Sul-Americanas Ciecircncia-Teacutecnica-Petroacuteleo Seccedilatildeo Exploraccedilatildeo dePetroacuteleo Rio de Janeiro 20 pp 25ndash42





Le Heacuterisseacute A Al-Ruwaili M Miller M Vecoli M 2007 Environmental changesre1047298ected by palynomorphs in the early Middle Ordovician Hanadir Member of theQasim Formation Saudi Arabia Revue de Micropaleacuteontologie 50 3ndash16

Loi A Ghienne J-F Dabard M-PParis F Botquelen A Christ N Elaouad-DebbajZGorini A Vidal M Videt B Destombes J 2010 The Late Ordovician glacio-eustatic record from a high-latitude storm-dominated shelf succession the BouIngarf section (Anti-Atlas Southern Morocco) Palaeogeography Palaeoclimatol-ogy Palaeoecology 296 332ndash358

Long DGF Copper P 1987 Stratigraphy of the Upper Ordovician upper Vaureal andEllis Bay formations eastern Anticosti Island Quebec Canadian Journal of EarthSciences 24 1807ndash1820

Marshall JEA 1991 Palynology of the Stonehaven Group Scotland evidence for aMid-Silurian age andits geologicalimplications GeologicalMagazine128 283ndash286

Mendlowicz Mauller P Pereira E Grahn Y Steemans P 2004 Anaacutelise bioestrati-graacute1047297ca do intervalo Llandoveriano da Bacia do Paranaacute no Paraguai Oriental RevistaBrasiliera de Paleontologia 7 199ndash212

Miller MA Eames LE 1982 Palynomorphs from the Silurian Medina Group (LowerLlandovery) of the Niagara Gorge Lewiston New York USA Palynology 6 221ndash254

Mizusaki AM Melo JHG Lelarge ML Steemans P 2002 Vila Maria FormationParanagrave BasinBrazilmdash an example of integrated geochronological and palynologicaldatings Geological Magazine 139 453ndash463

Molyneux SG Barron HF Smith RA 2008 Upper LlandoveryndashWenlock (Silurian)palynologyof thePentland Hills inliers Midland Valleyof ScotlandScottishJournalof Geology 44 151ndash168

Petryk AA 1981 Stratigraphy sedimentology and paleogeography of the upperOrdovicianndashlower Silurian of Anticosti Island Quebec In Lespeacuterance PJ (Ed)Subcomission on Silurian Stratigraphy Boundary Working Group Field Meeting

Anticosti-Gaspeacute Queacutebec 1981 2 Stratigraphy amp Paleontology pp 11ndash39Potildeldvere A (Ed) 2011 Valga (10) drill core Geological Survey of Estonia Tallinn

Estonian Geological Sections Bulletin 3 pp 1ndash50Richardson JB1985 LowerPalaeozoic sporomorphstheir stratigraphical distribution

and possible af 1047297nities In Chaloner WG Lawson JD (Eds) Evolution andEnvironment in the Late Silurian and Early Devonian Philosophical Transactions of the Royal Society of London Series B Biological Sciences 309 pp 201ndash205

RichardsonJB1988Late Ordovicianand Early Siluriancryptosporesand miospores fromnortheast Libya In El-Arnauti A Owens B Thusu B (Eds) Subsurface Palynos-tratigraphy of Northeast Libya Garyounis University Press Benghazi pp 89ndash109

Richardson JB 1996a Taxonomy and classi1047297cation of some new early Devoniancryptospores from England In Cleal CJ (Ed) Studies on Early Land Plant Sporesfrom Britain Special Papers in Palaeontology 55 pp 7ndash40

Richardson JB1996b Chapter 18A Lowerand Middle Palaeozoic records of terrestrialpalynomorphs In Jansonius J McGregor DC (Eds) Palynology Principles andApplications American Association of Stratigraphic Palynologists FoundationHouston pp 555ndash574

Richardson JG Ausich WI 2007 Late OrdovicianndashEarly Silurian cryptosporeoccurrences on Anticosti Island (Icircle dAnticosti) Quebec Canada Canadian Journalof Earth Science 44 1ndash7

Richardson JB Edwards D 1989 Sporomorphs and plant megafossils In HollandCH Bassett MG (Eds) A Global Standard for the Silurian System GeologicalSeries No 9 Cardiff National Museum of Wales Cardiff pp 216ndash226

Rubinstein CV 2005 Ordovician to Lower Silurian palynomorphs from the Sierrassubandinas (Subandean ranges) northwestern Argentina a preliminary report InSteemans P Javaux E (Eds) Pre-Cambrian to Palaeozoic Palaeopalynologyand Palaeobotany mdash Carnets de Geacuteologie Notebooks on Geology Brest Memoir200502 Abstract 09 (CG2005_M0209)

Rubinstein CV Vaccari NE 2004 Cryptospores assemblages from the OrdovicianSilurian boundary in the Puna Region NW Argentina Palaeontology 471037ndash1061

Rubinstein CV Gerrienne P De la Puente GS Astini RA Steemans P 2010 EarlyMiddle Ordovician evidence for land plants in Argentina (eastern Gondwana) TheNew Phytologist 188 365ndash369

Rubinstein CV Vecoli M Astini RA 2011 Biostratigraphy and palaeoenvironmentalcharacterization of the Middle Ordovician from the Sierras Subandinas (NWArgentina) based on organic-walled microfossils and sequence stratigraphy

Journal of South American Earth Sciences 31 124ndash

138Spina A Vecoli M 2009 Palynostratigraphy and vegetational changes in the Siluro-Devonian of the Ghadamis Basin North Africa Palaeogeography Palaeoclimatol-ogy Palaeoecology 282 1ndash18

Steemans P 2000 Miospore evolution from the Ordovician to Silurian Review of Palaeobotany and Palynology 113 189ndash196

Steemans P 2001 Ordovician cryptospores from the Oostduinkerke borehole BrabantMassif Belgium Geobios 34 3ndash12

Steemans P Le Hegraverisse A Bozdogan N 1996 Ordovician and Silurian cryptosporesand miospores from southeatern Turkey Review of Palaeobotany and Palynology 9335ndash76

Steemans P Higgs KT Wellman CH 2000 Cryptospores and trilete spores from theLlandovery Nuayyim-2 Borehole Saudi Arabia In Al-Hajri S Owens B (Eds)Stratigraphic Palynology of the Palaeozoic of Saudi Arabia GeoArabia BahrainSpecial Volume 1 pp 92ndash115

Steemans P Wellman CH Filatoff J 2007 Palaeophytogeographical and palaeoe-cological implications of a miospore assemblage of earliest Devonian (Lochkovian)age from Saudi Arabia Palaeogeography Palaeoclimatology Palaeoecology 250237ndash254

Steemans P Rubinstein C Melo JHG 2008 Siluro-Devonian miospore biostratig-raphy of the Urubu River area western Amazon Basin northern Brazil Geobios 41(2) 263ndash282

Steemans P Le Heacuterisseacute A Melvin J Miller MA Paris F Verniers J WellmanCH 2009 Origin and radiation of the earliest vascular land plants Science324 353

Steemans P Wellman CH Gerrienne P 2010 Palaeogeographic and palaeoclimaticconsiderations based on Ordovician to Lochkovian vegetation In Vecoli MCleacutement G Meyer-Berthaud B (Eds) The Terrestrialization Process ModellingComplex Interactions at the BiospherendashGeosphere Interface Geological Society of London Special Publication 339 pp 49ndash58

Strother PK 1991 A classi1047297cation scheme for the cryptospores Palynology 15219ndash236Strother PK 2000 Cryptospores the origin and early evolution of the terrestrial 1047298ora

In Gastaldo RA DiMichele WA (Eds) Phanerozoic Terrestrial Ecosystem ThePaleontological Society Papers 6 pp 3ndash19

Strother PK Traverse A 1979 Plant microfossils from the Llandoverian andWenlockian rocks of Pennsylvania Palynology 3 1ndash21

Strother PK Al-Hajri S Traverse A 1996 New evidence for land plants from thelower Middle Ordovician of Saudi Arabia Geology 24 55ndash58

Strother PK Wood GD Taylor WA Beck JH 2004 Middle Cambrian cryptosporesand the origin of land plants Memoirs of the Association of AustralasianPalaeontologists 29 99ndash113

Strother PK Vecoli M Beck J 2009 Paleopalynology of the Kanosh Shale at FossilMountain Utah Abstracts 9th North American Paleontological ConventionUniversity of Cincinnati Ohio USA June 21ndash26 2009 Cincinnati Museum CenterScienti1047297c Contributions N 3 p 203

Taylor WA 1995 Ultrastructure of Tetrahedraletes medinensis (Strother and Traverse)Wellman and Richardson from the Upper Ordovician of Southern Ohio Review of Palaeobotany and Palynology 85 183ndash187

Taylor WA 2002 Studies in cryptospore ultrastructure variability in the tetrad genusTetrahedraletes and type material of the dyad Dyadospora murusattenuata Reviewof Palaeobotany and Palynology 119 325ndash334

Taylor WA Strother PK 2008 Ultrastructure of some Cambrian palynomorphs fromthe Bright Angel Shale Arizona USA Review of Palaeobotany and Palynology 15141ndash50

Taylor WA Strother PK 2009 Ultrastructure morphology and topology of Camrbian palynomorphs from the Lone Rock Formation Wisconsin USA Reviewof Palaeobotany and Palynology 153 296ndash309

Tomescu AMF Pratt LM Rothwell GW Strother PK Nadon GC 2009Carbon isotopes support the presence of extensive land 1047298oras pre-dating theorigin of vascular plants Palaeogeography Palaeoclimatology Palaeoecology 28346ndash59

Torsvik TH 2009 BugPlates Linking Biogeography and Palaeogeography wwwgeodynamicsno

Turnau E Jakubowska L 1989 Early Devonian miospores and age of the ZwolenFormation (Old Red Sandstone Facies) from Ciepielow IG-1 Borehole AnnalesSocietatis Geologorum Poloniae (Rocznik Polskiego Towarzystwa Geologicznego)59 391ndash416

Vavrdovaacute M 1982 Recycled acritarchs in the uppermost Ordovician of BohemiaCasopis pro mineralogii a geologii 27 337ndash345

Vavrdovaacute M 1984 Some plant microfossils of the possible terrestrial origin fromthe Ordovician of central Bohemia Vestnik Ceskeho geologickeho ustavu 59165ndash170

Vavrdovaacute M 1988 Further acritarchs and terrestrial plant remains from the LateOrdovician at Hlasna Treban (Czechoslovakia) Casopis pro mineralogii a geologii33 1ndash10

Vavrdovaacute M 1989 New acritarchs and miospores from the Late Ordovician of HlasnaTreban Czechoslovakia Casopis pro mineralogii a geologii 34 403ndash420

Vavrdovaacute M 1990 Coenobial acritarchs and other palynomorphs from the ArenigLlanvirn boundary Prague Basin Vestnik Ustredniho ustavu geologickeacuteho 65237ndash242

Vecoli M 2000 Palaeoenvironmental interpretation of microphytoplankton diversitytrends in the CambrianndashOrdovician of the northern Sahara Platform Palaeogeo-graphy Palaeoclimatology Palaeoecology 160 329ndash346

Vecoli M 2004 Stratigraphic and palaeoenvironmental distribution of organic-walledmicrofossils in CambrianndashOrdovician transitional strata of borehole Bir Ben Tartar-

1 (Tt-1 Ghadamis Basin southern Tunisia) Memoirs of the Association of Australasian Palaeontologists 29 13ndash30Vecoli M 2008 Fossil microphytoplankton dynamics across the OrdovicianndashSilurian

boundary Review of Palaeobotany and Palynology 148 91ndash107Vecoli M RiboulleauA Versteegh GJM 2009Palynologyorganic geochemistry and

carbon isotope analysis of a latest Ordovician through Silurian clastic successionfrom borehole Tt1 Ghadamis Basin southern Tunisia North Africa palaeoenvir-onmental interpretation Palaeogeography Palaeoclimatology Palaeoecology 273378ndash394

Verniers J Vandenbroucke TRA 2006 Chitinozoan biostratigraphy in the Dobs LinnOrdovicianndashSilurian GSSP Southern Uplands Scotland GFF 128 195ndash202

Wang Y Li J Wang R 1997 Latest Ordovician cryptospores from southern XinjiangChina Review of Palaeobotany and Palynology 99 61ndash74

Wellman CH 1993 A land plant microfossil assemblage of Mid Silurian age from theStonehaven Group Scotland Journal of Micropalaeontology 12 47ndash66

Wellman CH 1996 Cryptospores from the type area of the Caradoc Series in southernBritain In Cleal CJ (Ed) Studies on Early Land Plant Spores from Britain SpecialPapers in Palaeontology 55 pp 103ndash136

Wellman CH Gray J 2000 The microfossil record of early land plants Philosophical

Transactions of the Royal Society of London Series B 355 717ndash732Wellman CH Richardson JB 1993 Terrestrial plant microfossils from Silurian inliers

of the Midland Valley of Scotland Palaeontology 36 155ndash193





Wellman CH Richardson JB 1996 Sporomorph assemblages from the lsquoLower Oldred Sandstonersquo of Lorne Scotland In Cleal CJ (Ed) Studies on Early Land PlantSpores from Britain Special Papers in Palaeontology 55 pp 41ndash101

Wellman CH Higgs KT Steemans P 2000 Spore assemblages from a Siluriansequence in Borehole Hawiyah-151 from Saudi Arabia In Al-Hajri S Owens B(Eds) Stratigraphic Palynology of the Palaeozoic of Saudi Arabia GeoArabiaBahrain Special Volume 1 pp 116ndash133

Wellman CHOsterloff PLMohiuddinU 2003Fragmentsof the earliest land plantsNature 425 282ndash285

Yin L He S 2000 Palynomorphs from the transitional sequences between Ordovicianand Silurian of northwestern Zhejiang South China Palyno1047298oras and Palynomophsof China 186ndash202





Research papers

Cryptospore assemblages from Upper Ordovician (KatianndashHirnantian) strata of Anticosti Island Queacutebec Canada and Estonia Palaeophytogeographic andpalaeoclimatic implications

Marco Vecoli ab Aureacutelien Delabroye c Amalia Spina d Olle Hints e

a Universiteacute Lille 1 Laboratoire laquo Geacuteosystegravemes raquo CNRS FRE 3298 SN5 Citeacute Scienti 1047297que Villeneuve dAscq 59655 Franceb Dipartimento di Scienze della Terra Universitagrave di Cagliari via Trentino 51 09127 Cagliari Italyc Universiteacute Paul Sabatier de Toulouse GET CNRS UMR 5563 14 av Edouard Belin 31400 Toulouse France

d CNR IRPI Perugia Italye Institute of Geology at Tallinn University of Technology Ehitajate 5 19086 Tallinn Estonia

a b s t r a c ta r t i c l e i n f o

Article history

Received 24 January 2011Received in revised form 29 April 2011Accepted 14 May 2011Available online 23 May 2011

Keywords

cryptosporesLate OrdovicianHirnantian

terrestrializationphytogeographypalaeobotany

Rich palynological assemblages have been recovered from deposits of Hirnantian age in Anticosti Island(Queacutebec Canada) and in borehole Valga-10 in southern Estonia The assemblages are well preserved andinclude acritarchs chitinozoans and cryptospores The age of the deposits is well constrained by means of palynomorphs (acritarchs and chitinozoans) as well as sequence stratigraphic and chemostratigraphiccorrelations Cryptospore assemblages from the two localities are similar and are also broadly comparable tothe few known coeval assemblages described elsewhere They include 11 genera and 20 species and testify tothe presence of an extended and diverse 1047298ora during Hirnantian times in Laurentia and for the 1047297rst time alsoin BalticaThe present 1047297ndings contribute to an improved knowledge of origin and early development of vegetative

cover The recovery of diverse and abundant cryptospores in Hirnantian deposits may be related to increasedinput of land-derived sediment during the global sea-level fall linked to the Late Ordovician glaciation but italso demonstrates that the early land plants may have tolerated a wide range of climatic conditions

copy 2011 Elsevier BV All rights reserved

1 Introduction

It is now generally accepted that by mid-Ordovician times(DapingianndashDarriwilian) bryophyte-like land plants had alreadyevolved in parts of Gondwana (Le Heacuterisseacute et al 2007 Rubinstein etal 2010 2011 Strother et al 1996) Cryptospores from the MiddleOrdovician Hanadir Sandstone in Saudi Arabia have been accepted as

the oldestevidenceof land plants because they possess morphologicalanalogs with spores produced by some living hepatics (Gray 1985Richardson 1996b Steemans 2000 Strother et al 1996 Taylor1995 Wellman and Gray 2000 Wellman et al 2003) Slightly olderless diverse and poorly preserved cryptospores with similar mor-phologies to the Hanadir assemblage occur in NW Argentina(Rubinstein et al 2010) Isolated specimens of laevigate permanenttetrads and dyads have been reported from the Klabava Formation(ArenigndashLlanvirn)of Bohemia by Vavrdovaacute (1990) but this report hasyet to be substantiated by further studies Richardson (1996ab)

reviewed all basicmorphological typesof cryptospores which includemonads and permanent dyads and tetrads as well as theirstratigraphic signi1047297cance He described a series of evolutionaryphases of early land vegetation based on the morphological in-novations observed in the fossil record of terrestrial palynomorphs(OrdovicianndashEarly Devonian) Richardson (1996b) proposed a ldquopre-cryptosporerdquo phase to include earliest Ordovician enigmatic spore-

like palynomorphs such as the enigmatic taxa Virgatasporites and Attritasporites which might have been related to freshwater algae orearliest terrestrial vegetation It is important to point out that thesepalynomorphs had been already considered as ldquosporesrdquo by theoriginal report of Combaz (1967) Vecoli (2000 2004) has alsonoted the association of Tremadocian sporomorph assemblageswith palynofacies indicating proximal depositional conditions withpossible in1047298uence of terrestrial sedimentary input The existence of primitive land plants during earliest Ordovician (Tremadocian) timesis also consistent with evidence from recent research suggesting thatterrestrial 1047298oras might have been widespread since the Cambrian(Strother et al 2004 Taylor and Strother 2008 2009 Tomescu et al2009) Cryptospores preserved in sporangia have been described fromsediments as old as Caradoc from the Hasirah Member of the Sa1047297q

Formation Haima Supergroup of Oman (Wellman et al 2003) The

Review of Palaeobotany and Palynology 166 (2011) 76ndash93

Corresponding author at Universiteacute Lille 1 Laboratoire laquo Geacuteosystegravemes raquo CNRS FRE3298 SN5 Citeacute Scienti1047297que Villeneuve dAscq 59655 France Fax +33 320434910

E-mail address marcovecoliuniv-lille1fr (M Vecoli)

0034-6667$ ndash see front matter copy 2011 Elsevier BV All rights reserveddoi101016jrevpalbo201105006

Contents lists available at ScienceDirect

Review of Palaeobotany and Palynology

j o u r n a l h o m e p a g e w w w e l s ev i e r c o m l o c a t e r ev p a l b o











balls and pillows

calcareous shales

bioherms


dark grey marls

wave-rippled

grainstones

HCS sandstones

Sandstones

lenticular

limestones

nodular limestones

Hardgrounds

tempestites

limestones



dolostones


silty dolostones

marls

dolomitic marls

















productive samples











Table 1




P s e u d o d


R i m o s o t




S e g e s t r e


C h e i l o t e

t r a s s p 2




C h e i l o t e

t r a s s p 1

C h e i l o t e

t r a s s p 2




P s e u d o d





EB42 EB324

EB132 EB320

EB18 EB319

EB111 EB318

EB109 EB316

EB94 EB314

EB4 EB313

EB2 EB312

EB311

EB310

EB309

EB308EB346

EB366


No cryptospores

































Table 2








r u g o s a







s s p






l a e v i g a t a





e v i g a t a


g o s a


p o r e

VA38 3132

VA40 3200

VA42 3290

VA44 3355

VA46 3457








Plate I





















1971 lsquo


gurationrsquo




























14 and 15






























p 176 pl 1 1047297

gs 10ndash























Plate III














VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References









































































































































balls and pillows

calcareous shales

bioherms


dark grey marls

wave-rippled

grainstones

HCS sandstones

Sandstones

lenticular

limestones

nodular limestones

Hardgrounds

tempestites

limestones



dolostones


silty dolostones

marls

dolomitic marls

















productive samples











Table 1




P s e u d o d


R i m o s o t




S e g e s t r e


C h e i l o t e

t r a s s p 2




C h e i l o t e

t r a s s p 1

C h e i l o t e

t r a s s p 2




P s e u d o d





EB42 EB324

EB132 EB320

EB18 EB319

EB111 EB318

EB109 EB316

EB94 EB314

EB4 EB313

EB2 EB312

EB311

EB310

EB309

EB308EB346

EB366


No cryptospores

































Table 2








r u g o s a







s s p






l a e v i g a t a





e v i g a t a


g o s a


p o r e

VA38 3132

VA40 3200

VA42 3290

VA44 3355

VA46 3457








Plate I





















1971 lsquo


gurationrsquo




























14 and 15






























p 176 pl 1 1047297

gs 10ndash























Plate III














VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References













































































































































productive samples











Table 1




P s e u d o d


R i m o s o t




S e g e s t r e


C h e i l o t e

t r a s s p 2




C h e i l o t e

t r a s s p 1

C h e i l o t e

t r a s s p 2




P s e u d o d





EB42 EB324

EB132 EB320

EB18 EB319

EB111 EB318

EB109 EB316

EB94 EB314

EB4 EB313

EB2 EB312

EB311

EB310

EB309

EB308EB346

EB366


No cryptospores

































Table 2








r u g o s a







s s p






l a e v i g a t a





e v i g a t a


g o s a


p o r e

VA38 3132

VA40 3200

VA42 3290

VA44 3355

VA46 3457








Plate I





















1971 lsquo


gurationrsquo




























14 and 15






























p 176 pl 1 1047297

gs 10ndash























Plate III














VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References






























































































































































Table 2








r u g o s a







s s p






l a e v i g a t a





e v i g a t a


g o s a


p o r e

VA38 3132

VA40 3200

VA42 3290

VA44 3355

VA46 3457








Plate I





















1971 lsquo


gurationrsquo




























14 and 15






























p 176 pl 1 1047297

gs 10ndash























Plate III














VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References


































































































































Plate I





















1971 lsquo


gurationrsquo




























14 and 15






























p 176 pl 1 1047297

gs 10ndash























Plate III














VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References


















































































































































1971 lsquo


gurationrsquo




























14 and 15






























p 176 pl 1 1047297

gs 10ndash























Plate III














VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References




















































































































































p 176 pl 1 1047297

gs 10ndash























Plate III














VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References




































































































































Plate III














VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References











































































































































VA38 (3132)




VA38 (3132)




















Burgess pp 589ndash

590 pl 2 1047297

gs 7ndash

9








1996

























Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References

















































































































































Plate IV 57







(Ordovicianndash

















































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References


























































































































































VA38 (3132)















2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References






































































































































2

13

14

19

101

154

5 7

8

612

17

16

9

1811

3



Table 4


1 Estonia

2 Anticosti

1 E s t o n i a

2 A n t i c o s t i

3 W a l e

s

4 B r i t a i n

5 B e l g i u m

6 T u r k e y

7 S a u d

i A r a b i a

8 L

i b y a

1 0 C z e c h i a

9 A r g e n t i n a

1 1 C

h i n a

500

500

421

400

368

360

250

273 200

222 294

200

240

258 125

118

217

267

259

182

Table 3


Miospores 1 2 3 4 5 6 7 8 9 10 11











Acknowledgements




References





































































































































Acknowledgements




References































































































































Vecoli Et Al 2011

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Transcript of Vecoli Et Al 2011