Radiolaria Siliceous Plankton Through Time

Supplement 1:
Eclogae Geologicae Radiolaria

Helvetiae Siliceous Plankton through
Time
Swiss Journal of
Geosciences
Zeitschrift der I Revue de la

Schweizerischen Geologischen Gesellschaft I Societe Geologique Suisse
Schweizerischen Paliiontologischen Gesellschaft I Societe Paleontologique Suisse
Schweizerischen Fachgruppe ftir Geophysik I Groupement Suisse de Geophysique
Schweizerischen Fachgruppe ftir Sedimentologie (SwissSed) I Groupement Suisse de Sedimentologie (SwissSed)
Schweizerischen Fachgruppe flir Tektonik (Swiss Tectonic Studies Group) I Groupement Suisse des Tectoniciens
(Swiss Tectonic Studies Group)
Contents
BAUMGARTNER, P. O. FENG, Q., MENG,Y., HE, W., Gu, S.:

Preface . . ... SIll A new genus of Entactiniidae (Radiolaria) from the Upper Per-
mian of South China S 67
N., BAUMGARTNER, P.O., CARON, M.:
BANDINI, A.
Turonian Radiolarians from Karnezeika, Argolis Peninsula, HOLLIS, C. J.:
Peloponnesus (Greece) S I Radiolarian faunal turnover through the Paleocene-Eocene tran-
sition, Mead Stream, New Zealand S 79
BECCARO, P.:
Radiolarian correlation of Jurassic siliceous successions of the MARQUEZ, E. J., AITCHISON, J. c. ZAMORAS, L. R.:
Rosso Ammonitico Formation in the Southern Alps and West- Upper Permian to Middle Jurassic radiolarian assemblages of
ern Sicily (Italy) . . . . . . . . . . . .. . S 21 Busuanga and surrounding islands, Palawan, Philippines S 101
DANELIAN, T., LAHSINI, S., DE RAFELIS, M.: WANG, Y.- J., Luo, H., AITCHISON, J. C;
Upper Jurassic Radiolaria from the Vocontian basin of SE Influence of the Frasnian-Famennian event on radiolarian
France . . . . . . . . . .. . . .. S 35 faunas S 127
DE WEVER, P., O'DOGHERTY. L., GORICAN, S.: WONGANAN, N., CARJDROIT, M.:
The plankton turnover at the Permo-Triassic boundary. empha- Middle to Upper Permian radiolarian faunas from chert blocks in
sis on radiolarians. . . . . . . . . . . . S 49 Pai area, northwestern Thailand S 133
DOLVEN, J. K., SKJERPEN, H. A.:

An online micropaleontology database: Radiolaria.org S 63
0012-9402/06/01SIll-V Eclogae geol. Helv. 99 (2006) Supplement 1, SIIl-SV
001 1O.1007/s00015-006-0611-4
Birkhaus er Verlag, Basel, 2006
UNIL I Univers ite de Lausanne
Radiolaria - Siliceous plankton through time

Proceedings of the Tenth Meeting of the International Association of Radiolarian Palaeontologists
INTERRAD X, held at the University of Lausanne, Switzerland, September 2003
Preface
PETER 0. BAUMGARTNER
Radiolaria are a very diverse marine siliceous microplankton obstacle to the elaboration of detailed biochronozones is the
group that have existed at least since the Cambrian to the Re- discontinuous nature of the radiolarian record, due to spotty
cent. Fossil Radiolaria were known since the middle of the pre servation. Biogenous, opaline silica is unstable in the ocean
19th Century and studied by man y scienti sts at the turn of the and in the diagenetic environment. Large amounts of the more
19th120th Century, in part as a consequence of the discoveri es deli cate radiolarian tests become dissolved already during
made in samples collected be the HMS Challenger expediti - their descent in the wat er column and in the bottom sed iments
ons. Intense biostratigraphic radiolarian rese arch was spurted (Takahashi & Honjo 1981). During early diagenesis, opaline
again in the early 1970's by the Deep Sea Drilling Project. silica dissolves and radi olarians becom e preserved either as
Biochronologic zonations based on sample s recovered by casts of the mould that leaves the dissolved test, or as replace-
Ocean Drilling were first proposed for the Tertiary (Riedel & ment by quartz, pyrite or other minerals. In Palaeozo ic and
Sanfilippo 1978) and th e Cre taceous (Fo reman 1973, 1975). Mesozoic samples, usually onl y robust for ms are preserved .
However , the radiolarites, cherty oceanic sedime nts associated Ho wever , exceptionally well preserved samples from special
with ophi olites, had so far only been studied in thin sections. diagene tic enviro nme nts, such as Middle Jurassic man ganese
This chan ged when radiolarian work ers discovered the use of carb on ate nodules (Yao 1997), or euxinic en vironm ents like
hydroflu or ic acid to extract Radiolaria from che rt and othe r the Late Jurassic Solnh ofen Limestone (Z tigeI 1997, Dumitrica
siliceous rocks (Dumitrica 1970, Pessagno & Newport 1972) & Ztigel 2002) show several hundred morphotypes, a diver sity
The new method allowed to work on land samples from moun- that rivals with the diversity of living radiolarians. Th ese sam-
tain ranges that had unde rgone burial diagenesis or even meta- ples teach us, that the average radiol arian assemblage extrac-
morphosis. As a consequence, radiolarian biochronology was ted by harsh chemical treatments from rocks is a poor residue
rapidl y extended into the Jurassic, Triassic and the Palaeozoic of dissolut ion resistant forms. The consequ ence of this una voi-
(Pessagno 1977, Pessagno et al. 1979, Yao et al. 1980, Nak ase- dable fact is the discontinuous record of radiolarian ranges,
ko & Nishimur a 1979, Holdsworth & Jones 1980). A major which greatly hampers biochronologic cor relation. A major
Institut de Geologie et Paleont ologie, Universite de Lausann e, Anth ropole. CH-1015 Lausanne, Switzerland. E-mail: Peter.Baum [email protected]
Preface SIll
step in resolving this problem has been the use of the Unitary essentially based on biochronologic ages and palaeobiogeo-
Associations method (Guex 1977, 1991). Already in early at- graphic affinities of Radiolaria. Much work has dealt with radi-
tempts for a Jurassic-Cretaceous radiolarian zonation (Baum- olarian faunal changes related to major geologic boundaries in
gartner et al. 1980), we realised that this method allows inte- the Palaeozoic (Wang & Luo this volume), at the Permo-
gration of the data on mutual co-occurrence of all considered Triassic boundary (DeWever et al. this volume), and in the
taxa from all studied sections, thus overcoming local gaps of Cenozoic (Hollis, this volume). Mesozoic radiolarian
preservation. Unitary Associations represent maximal sets of biochronology continues to be improved and better calibrated
co-occurring taxa that are put in a stratigraphic sequence by (Danelian this volume, Beccaro this volume, Bandini et al. this
their identification in all studied sections. Starting in 1984 volume).
(Baumgartner 1984a,b) computer programs (Guex and Da- For a number of years, the radiolarian community is con-
vaud 1982) allowed the calculation of Unitary Associations cerned with the development of databases. With the rapid de-
from large data sets. The largest data set so far treated was the velopment of computer technologies, databases have become
MRD database on Middle Jurassic to Early Cretaceous radio- more and more sophisticated and for the last few years only,
larians (Baumgartner et al. 1995). A radiolarian biozonation have become available online. There has been a common
including 22 Unitary Association Zones for the Middle Juras- agreement among radiolarian specialists, that Radiolaria.org is
sic to Early Cretaceous interval was calculated with the Bio- the portal that provides links to most radiolarian databases
graph software (Savary & Guex 1991). Today the majority of and hosts a wealth of information on Radiolaria and radiolari-
Mesozoic radiolarian biozonations are based on Unitary Asso- an workers (Dolven, this volume).
ciations. An excellent example is presented in this volume
(Beccaro, this volume). A new software, UA-Graph (Hammer
et al. 2001) has been developed. REFERENCES
Dating pelagic sediments by radiolarian biochronology has
BAUMGARTNER, P. O. 1984a: Comparison of Unitary Associations and proba-
totally changed our understanding of the geology of Tethyan bilistic ranking and scaling as applied to Mesozoic radiolarians. Computer
and Circumpacific mountain ranges. Suture zones and ophioli- & Geosciences 10(1), 167-184.
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ancient oceans, needed to be dated. A vast campaign of sam- radiolarian zonation based on unitary associations and age of Tethyan ra-
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going on. DANELAIN, T., DUMITRICA-JUD, R., GORICAN, S., GUEX, J., HULL, D.,
Today, Radiolaria are an important microfossil group used KITo, N., MARCUCCI, M., MATSUOKA, A., MURCHEY, 8., O'DOGHERTY,
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Jurassic to Early Cretaceous Radiolarian Biochronology of Tethys Based
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1043.
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BAUMGARTNER, P.O., DE WEVER, P. & KOCHER, R. 1980: Correlation of
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HOLDSWORTH, B. K. & JONES, D. L.1980: Preliminary radiolarian zonation for ontology 27(2),140-190.
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and ice-age Pacific. Marine Micropaleontology 3(3), 229-266. Glacial! interglacial changes in the equatorial part of the Somali Basin
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PESSAGNO, E. A., JR. 1977: Upper Jurassic Radiolaria and radiolarian bios- Osaka Micropaleontologists 10, 155-182.
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PESSAGNO, E. A., JR., FINCH, W. & ABBOTT, P. L. 1979: Upper Triassic Radio- University 23, 135-154.
laria from the San Hipolito Formation. Baja California. Micropaleontol- YAMASHITA, H .. TAKAHASHI, K. & FUJITANI, N. 2002: Zonal and vertical dis-
ogy 25(2),160-197. tribution of radiolarians in the western and central Equatorial Pacific in
PESSAGNO, E. A. & NEWPORT, R. L. 1972: A technique for extracting Radio- January 1999. Deep-Sea Research 11,49,2823-2862.
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Preface SV
0012-9402/06/01S0001-20 Eclogae geol. Helv. 99 (2006) Supplement 1, SI-S20
DOI 1O.1 007/s00015-006-0605-2
Birkh auser Verlag, Basel , 2006
Turonian Radiolarians from Karnezeika, Argolis Peninsula,

Peloponnesus (Greece)
ALEXA NDRE N. BANDINI 1, PETER 0. BAUMGARTNER 1 & MICHELE CARON 2
Key words: Mesozoic, Upper Cretaceo us, Turonian, Radiolaria , Fo raminifera, Teth ys, Pelagonian Zone, Argolis Peninsula, Greece
ABSTRA CT
Near Karnezeika a roughly 140 m thick Upper Cretaceous section consists of species are described and figured in this work. The radiolarian chronos tratig-
interb edded pelagic limestones , cherts and coarse polymict breccias including raphy established by 10 different authors in 11 publications was compared for
ophiolites and shallow water limeston es. At the base, pink pelagic limestones this study and used to establish radiolarian ranges. This exercise shows major
rest on deep ly altered and fractu red Lower Jur assic Pantokrator Limeston e. discrepan cies between authors for the radiolarian ranges of the studied assem-
Th is first pelagic facies is dated as middle Tur onian, based on planktonic blage. Nevertheless, a Turon ian age can be state d based on a synthesis of cited
Fora minifera. Ove r 100 m of coarse ophiolite-carbo nate breccias, interpreted radiolarian ranges. Thi s age is consistent with the age based on planktonic
as a channel or canyon fill in a pelagic environment, document the eros ion of fora minifera. In combining the ages of both Radiolaria and planktonic
the Late Jur assic napp e edifice along the Cre taceo us Pelagonian margin. Foraminifera, the studied samples can be restricted to the late Tu ronian. How-
Above these breccias, we mesured 16 m of principally pink and red pelagic ever , the discrepancies of publ ished radiolar ian ranges call for an urgent,
limestones and radiolarian cherts, in which we recovered well-preserved radi- major revision of the Late Cretaceo us radiolarian biochro nology. The integra-
olarians discussed here. In this interval. the prese nce of planktonic For amin- tion of plankt onic foraminifera with radiolarians may grea tly enhance
fera allows to state a late Turonian to Coniacian age. More than 40 radiolarian biochro nologic resolution in sections where both groups occur.
Introduction
Relatively few studies exist of Upper Cretaceous Radiolaria op hiolite-carbonate breccias yield ed well-preserved radiolari-
from Greece, except for some occurences in the Pindos- ans. From these samples 41 radiolarian species belonging to 18
Ol onos zone (De Wever & Thi eb ault 1981; Thiebault et al. gen era are described and figured in this work. Overall, th e as-
1981; De Wever & Origlia-Devos 1982; Neumann 2003). In the semblage resembles those described by O'Dogherty (1994)
Hellenides , ongoing tectonic activity is reflected in a small- from the lower Turonian. The radiolarian biostratigraphy es-
scaled puzzle of shallow, detrital and pelagic facies in palaeo- tabli shed by the following 11 publications (given with regions)
geographic realms such as the Pelagon ian (Vrielynck 1981). was compared for this study: Dumitrica (1975 , Romani a),
Upper Cretaceous radiolarian occurrences ar e therefore re- For eman (1975, Pacific and 1977, Atlantic), O 'Dogherty (1994,
stricted to times of high silica producticity in pelagic palaeo en - Italy and Spain ), Pessagn o (1976, Californ ia), Riedel & Sanfil-
vironments. One of the objec tives of th is study was to compare ippo (1974, Composite), San filippo & Riedel (1985, Compos-
publ ished radiolarian ran ges give n by different authors for the ite), Schaaf (1985, Composi te) , Taketani (1982, Jap an ),
Upper Cretaceous and to tr y to esta blish an acceptable radi o- Thurow (1988, Atlantic), Vishnevskaya (2001, Russia).
larian age in spite of the differ enc es in radio larian ranges give n Th e presence of abundant planktonic Foraminifera both at
by the various authors. th e ba se of the studied Cr et aceou s section and immediately
For this pr eliminary work we collected 17 samples in a total above the radiolarian samples allow th e comparison between
of 140 m of section for the study of planktonic Foraminifera th e ages determined by radi olari an and foraminiferal bios-
and R adiolari a. 5 samples in a 2.6 m inte rva l above coarse tr ati graph y (Caron 1985).
I Institut de geologie et de paleont ologie, Anth ropole-Dorigny, Un iversite de Lausanne , 1015 Lausann e, Switzerland .
Emai l: Alexandr [email protected]@unil.ch
2 Institut de geologie et de paleotonlogie, Universite de Fribourg, Perolles, 1700 Fribourg, Switzerland . Email: Michele.Caron @unifr.ch
Turonian Radiolarians from Karneze ika SI

Neoautochtonous
~ Peloponnesus I IQuatemary
L:::........=J Conglomerate '-----J alluvial deposits
Tert iary Unit s (Ermi oni • Ad heres)
Adheres Unit Interstratifications of
pelagic limestone
Akros ~ Poros Formation
Nappe 4' • 4 Early Tertiary overthrust
Dhidh lm i·Trapezona Composite Unit

Ermioni Mesoauto- Ligourion Mesoauto-
chtonous Series chtonous Series
Ophiolites Imbricates of
~ ASkliPion Nappe
deep ~ Late Jurassic overthrust
shallow ~ Basal Series
---
deep ~1
shallow ~ Basal Series
I ILower
'-----J Tertiary flysch
Fig. I. Overview of the tectono-stratigraph y of the Argolis Peninsula. Based on: Baumgartner (1985), Vrielynck (1981), Clift (Poros Island) and several unpub-
lished diploma thesis University of Lau sann e (Bandini 2004; Giraud 2005; Glassey 2005).
Geological Setting shallow water limestones that document the ongoing erosion
of the Late Jurassic Pelagonian nappe edifice in a high-relief,
In the Argolis Peninsula, the Juras sic nappe edifice of the east- deeper marine environment.
ern Pelagonian margin is unconformably overlain by several
different Cretaceous series (Vrielynck 1981). The studied sec-
Stratigraphy and sedimentology
tion is located in the Depression of Karnezeika-Stavropodhi, a
complex nappe syncline affected by neotectonic E-W trending Near Karnezeika a roughly 140 m thick section consists of in-
subvertical faults (Fig. 1). The northern edge of this zone is terbedded pelagic limestones, chert s and boulder breccias con-
built by Upper Triassic to Lower Jurassic Pantokrator Lim e- taining abundant ophiolite clasts along with boulders of the
stone (Fig. 2) belonging to the Basal Series of the Dhidhimi- underlying Pantokrator Limestone (Fig. 3). At the base, pink
Trapezona Composite Unit (Baumgartner 1985). These series pelagic limestones rest on deeply altered and fractured Pan-
became overthrust during the late Juras sic by nappes including tokrator Limestone. Over 100 m of coarse ophiolite-carbonate
ophiolites and then the area became deeply eroded during lat- breccias represent a channel or canyon fill in a pelagic environ-
est Jurassic and Early Cretaceous times. ment. The following 16 m are principally pink and red pelagic
Upper Cretaceous pelagic and coarse clastic sediments un- limestones and radiol arian cherts , from which we recovered
conformably overlie the Pant okra tor limestone of the basal se- well-pre served radiolari ans described here.
ries and contain disorganised boulder breccias of basalts and The studied section rests conformably on the breccias de-
S2 A.N . Band ini, P.O. Baumgartner & M. Caron

GEOLOGICAL MAP
EAST OF KARNEZEIKA
1
HHHt=1H
0 2Km
o
QUATE RNARY DEPOSITS
Indifferent
UPPER CRET. ALL OCHTHONOUS SERI ES
~
~~~ Limesto nes, marl s, shallow wa ter &
pelagic breccias
Turonian - Maastrichtian
Tert iary Overthrust s
DHIDHIMI·TRAPEZONA COMPOSITE UNIT
Flysch
Upper Cretaceous · Paleocene
Pelag ic limestones & marls

Turonian - upper Campanian
Ophiolite Breccias
Turonian - upper Campanian
Migdhalitsa Ophiolite Unit

~ Serpentinites
<: Karnez eika
~
- - Jurassic Overthrusts
Basa l Seq uence

p O
~ Potaml Fonmatlon
~ Kimmeridgian
~= _=_=_=~
-=;:-.;:--=:;:--=:;:
Dhlma lna Fonmatlon
Kommendglan
~ Pantokrato r Li mes ton e

~ Norian· Lias
Fig. 2. Geological M ap of the area east of Karnczeik a (modified after V ernez 1990)
scribed above. It is a 16 m thick alternation of marls and red believe that the differences result from locally inclompet e
radiolarian cherts with detrital levels evolving progressively ranges of radiolarian taxa, eithe r due to plaeobiogeographic or
int o a pel agic limestone rich in Gl obotruncanidae. Up section pale oecologic exclusions, or due to preservational (diagenetic )
follow 10 m of disorganised boulder br eccias with a limestone biases. The only way to use th ese ra diolarian zonations is to
inte rbe d, which are in turn un conformabl y covered by quater - maximise the range of each taxon by stacking th e "spartial"
nar y br eccias. ran ges expressed in each publicat ion . In principle, the best way
of doing this is to cre at e U nita ry A ssociations (G ue x
1977-1 991: Baumga rtner et al. 1980; Gu ex & Da vaud 1982,
Radiolarian biochronology
1984; Baumgartner 1984 and Savar y & Gu ex 1991) in using th e
T oday no standard radiolarian bioch ron ology is available for the occurre nce data of well-de fine d tax a only, det ach ed from the
Late Cret aceous. However , several local and regional radiolarian chro nostra tigraphic calibrati on s (Ba umgartner et al. 1995). In
zona tions have been proposed in the past (sec citations above). such a way , we can con struct a range chart for th e Lat e Creta-
In comparing the published ran ge charts it becomes evide nt ceo us that reflects ma ximal ra nges of each taxon with re spect to
th at the chro nostratigraphic ra nge of an y given ta xon proposed the ma ximal ranges of all othe r taxa. Thi s work is in progre ss,
by differ ent authors sho ws major discrep an cies for the studied but not completed (Jack ett et a1. 2002 and Diserens et al. 2003) .
asse mblage (Figs. 4a-b-e--d ) from one pubicati on to the other. For thi s paper the co mpa rison of ran ges is based on th e
Thi s may be du e to uncertainties in the chronostratigraphic ca l- chro nos tratigraphic ran ge of ea ch taxon ex pessed by each au-
ibra tion of radiolarian occur re nces. However, most radiolarian th or. We have simply stacked th ese chronostratigraphic ran ges
zona tions of the Late Cretaceous ar e rather well calibrated by to obta in a minimum and a maximum age for th e existe nce of
mean s of plaktonic foraminifer a and nannofossils. We rather each taxon. We are awar e of the possible errors that may ar ise
Turo nian Radiolarians from Karn ezeik a S3

St ratig raphic log of the Karezeika section , A rg oli s Peninsula (Greece)
140m
130m L1imeston e
Stratigra phic log of the studied part

of the Karezelka section , Argo lls
120m Peninsula (Greece)
Quaternary
Debris now
110m 16m .
Pink pelagic
g!"'"
100m
marls 15m ·n
!iiil~~ ~:
CKAR'3" "0]
Pink pelagic 14m
~8
sorn
marls j AJ76 1350" E il
~ - ~~
13m 2
:3' Progressive passage of
cherts to limestones
6m
80m
5m
..
."
Pink pelagic ~
E
marls ~
c
10m '0.
4m
§ ~
Bloks of
limestone
j
3m J 's""
AI73 320·
60m A17,( 300°
~ .~
2 ~
"0
2m E
A172_190"
SOm "0
E
Ophiolite-carb "0
Al70_090 "
onate breccias 1m .~
""'7 1_0a0
"iii
A169_070
40m Al68_060" !
«
Om
Oebrisflow
Serpent inites
• Radiolarian samplesconsidered for this atudy
30m and basalts •• FOfiIIffiirwfer sa~ les consideftlid for Ihis study
Dolom itic breccias blok s of several meters

Serpen tinites and basalts
20m Dolom itic breccias bloks of several meters
Oph iol ite-carbonate breccias with ol istoliths of

10m
serpentinites
(KA R7"
~~~2" Pink pelagic limeston e
Om
AI13l Deeply altered and fractured Pantokrato r Limestone
Fig. 3. Stratigraphic log of the Karnezeika sec-
Pantokra tor Limestone tion. The inset at right shows the pelagic interval
studied for radiolarians and planktonic foramin-
fera with the samples.
using this procedure. We are, however, in good company, since forniaensis and Afens liriodes (have not been cited from earlier
this procedure was practiced in a series of papers on Mesozoic than early Turonian), found together with Pseudoaulophacus
Radiolaria (e.g. De Wever et al. 1986). putahensis (has not been cited from later than late Turonian).
Despite the major discrepancies between the ranges of Moreover, Dictyomitra urakawensis, Stichomitra communis
each author, a Turonian age can be stated using the procedure and Pseudodictyomitra pseudomacrocephala have not been
described above. This age is mainly based on the presence of cited from later than Coniacian, Crucel/a messinae have not
Patel/ula ecliptica, Patel/ula heroica, Praeconocaryomma cali- been cited from later than early Santonian (Fig. 5).
S4 A.N. Bandini, P.O. Baumgartner & M. Caron

.-
Acaemotyfe rebel/is
O'OOGHERTY, 1994
Acan thocircus hueyi

(PESSAGNO, 1976 )
sensu O'DOG HERTY, 1994
Acanthocircus tympanum
O'OOGHERTY, 1994 ?
Acanthoclrcus venetus
(SQUINABOL, 1914) ?
sensu O'OOGHERTY, 1994
Afens llriodes
RIEOEL & SA NFILIPPO, 1974
Archaeocenosphaera (7) mellifera

O'OOGHERTY,1994
Cruce/fa cectiensis
PESSAGNO, 1971
Fig. ua, Selected T uro nian-Santon ian rad iolari an ranges according to 10 authors as cited in the figure s. Note the major discrepe ncies between authors,
Turonian Radiolarians from Karn ezeika S5

SPECIES PICTURES AUTHORS
Crueel/a messinae
PESSAG NO .1971
1974
1985
Die/yomltra formosa
SQUINABOL , 1904
Dictyomnre mo ntisserei
(SQUINABOL. 1903)
sensu O'OOGHERTY, 1994
1974
1985
Dle/yomllra urakawensls
TAKETANI 1982 Foreman 1975 1977
O' ""rt1994
Pessa no 1976
RIMel and Sanfih 1974
Sanfili and Rldel 1985
Helesium uiocontnum
.....
Durrutnc a 1975
(SQUINABOL, 1903b) Foreman (19751977
' ,~
Pes no 1976
Ried el and Sanhh 197 4
Sanfili and Rrdel 1985
Schaa f 19A5
Tf'lk.e ta ni 1982
0;;
Vishnevska a 12001)
Patel/ufa eefipt/ea
O'OOGHERTY,1994
Rtedet and SanfiJi 1974
SChaaf 1985
Iaketam 11982
Thurow 1988
Vlshnevska a (2001)
Patel/ufa he/ios Durmrnca 1975

(SQUINABOL, 1903 ) Foreman 1197.5 1977
sensu O'OOGHERTY
Riede-l and Sanf,! (197 4)
Thurow 11988
Vishnevska a (2001)
Fig, 4b. Selected Turonian-Santoni an radio larian ra nges according to 10 authors as cited in the figures . Note the major discrepencies between auth or s.
S6 A,N, Bandini , P.O. Baumgartner & M. Caron

Patel/ula neroice
O'OOGHERTY,1994
Praeconocaryomma califomiaensis
PESSAGNO , 1976
Praeconocaryomma lipmanae
PESSAGNO, 1976
Praeconocaryomma universa
PESSAGNO, 1976
Pseudoacanlhosphaera galeata
O'OOGHERTY, 1994
Pseudoacanthosphaera superba
(SQUINABOL, 1904 )
Pseudoa woph a cus p umh en~s

PESSAGNO, 1972
Fig.4c. Selected Turonian-Santonian rad io larian ran ges according to 10 a uthors as cited in the figures. Note the major discrepencie s between authors.
Tu roni an Radiolarians from Karnezeika S7

Pseudoa ulophacus sculptus
(SQUINABOL. 1904)
sensu O'DOGHERTY. 1994
Pseudodictyomitre
pseudomacrocephale
(SQUINABOL , 1903)
Stichomtlra communis
SQUINABO L,1903
sr ichomitra stocki
(CAMPBELL & CLARK . 1944)
sensu O'DOGHERTY. 1994
Tetracanthellipsis euganeu s
SQUINABOL .1903
PeSS8 no 1976
R del and anfili 1974
Santil! and Ridet 1985
ScI1aaf 1985
Tak IRni 19
Thurow 1988
Vishnevska a (2001)
Triactome cellulose Dumitrica 197 5

FOREMAN , 1973 ForemAn 1975 1977
•••• .0
Pes no 1 76
Riedel and Sanfili 1974
Sanfih and Ridel 1985
ScI1aaf 1985
Takelani 1982
Thurow 1988
Voshnevska a (2001)
Triectom e hexeris Dumrtnca 1 75

O'DOGHERTY, 1994 Foreman 1975.1977
P sa no 1976
Riedel and Santili 1974
Sanfili and Rid91 1985
Schaa f 1985
Taketani 1982
Thu 1
Vl$hnevska a 200 1)
Fig. 4d. Selected Turonian-Santonian radiolarian ranges according to 10 authors as cited in the figures. Note the major discrepencies between authors.
S8 A.N. Bandini , P.O. Baumgartner & M, Ca ron

T C S
U
0 A
N N
R
I T
0
N
A 0
C N
I
I I
A
A A 0 0 0 0 0
N <0 Ol
~ 0 N
N N 0,
co
0
1
N
I "', "',
"1"
S
0
~ ~ ~ '"
~
E M L E L <X: <X: <X: <{ <X:
IPseud08ulophaeus seulptus <I> ? i

IPseuooecemnospneers galeata 17 < [> ? I i
ITetracan thellipsis euganeus 16 <I> ? I
IAeaeniotyle rebellis 13 1
IArehaeoeenosphaera (?) mellifera l

Dietyomitra montisserei
IHalesium triaeanth um 31
IPseudoacanthosphaera suoerb« 18
I Triaetoma cettutose I 21 I <t--I> ?I I I I I I I I I I

ITriaetoma hexeris I 22 1 <F"""'l> ?I 1 I I I I I I I I 1
Pseudoaulophaeus putahensis 25 < I I I I
Stiehomitra communis 07 < I I I
Dietyomitra urakawensis 04 < I I I
Pseudodietyomitra pseudomaerocephala 05 < I I
Crueella messinae 34 < I
Aeanthocircus tympanum 11 < > ? I
Aeanthocircus venetus 10 -e > ? I I

Aeanthocircus nuevi 12 -e > I I I
Crueella eaehensis 35 < > I I I I
Dietvomitra formosa I 01 I < > I I r I I I
IPatellula netios I 26 I < 1--- ••••••• > I I I I ~ I
IStiehomitra stocki I 08 I < > I I I I I I
IPraeeonoearyomma lipmanae I 40 I? -c e- .......3-· ··1 I> 1 I r I I 1
IPraeeonoearyomm a universe I 38 I < > I I I I I I
IPatellula eeliptiea I 27 I ~> ? I I I I I I I I I I 1
IPatellula heroiea 28 II> ?I
Praeeonoca omma ealifomiaensis 39
Afens liriodes 09
FORAMINIFERA
CKAR2
IMarginotruncana marianosi CKAR7
Diearinella primitiva I CKAR 14
Margino truneana sigali , CKAR 13 & All6 1350
Marginotruneana sehneegansi CKAR 13 & 14
Marginotruneana corona ta > CKAR 13
Marginotruneana pseudolinneiana > CKAR 14 & AIl6 1350
IMarginotruneana sinuosa 1 I I I I I I I CKAR 14 I

Fig. 5. Summary of T uronian to Santoni an ra diolaria n and foraminfe r ages obt ain ed for the studied samples. Th e radi olarian ran ges represent maximum ran ges
acco rding to the 10 authors cited in figure 4.
Tu roni an Radiolarians from Karnezeik a S9

Systematic Paleontology of Radiolaria Archaeocenosphaera (?) sp,
(Plate 2, Figs. 20-21)
T he suprage ne ric sys te matics pre sented her e foll ow De
Wever et al. (2001) . The syno nymies give n incl ude th e origi -
Remarks. - Cortical shell very lar ge, spherical with symme tri-
nal descr iption of ea ch taxon a nd additiona l synonymies
cal meshwork. Cortical she ll with very large pol ygon al por es.
fro m th e fo llow ing publicati on s: Dumi trica (1975), Forem an
A medullary spongy cortical she ll can be seen th rough th e
(1975, 1977), Q 'Dogherty (1994) , Pe ssagno (1976), R ied el &
large por es of th e first one, Very lon g spines occur at pore
Sanfilippo (1974), Sanfilippo & R iedel (1985), Schaaf (1985),
junctions, Spines with circular cross -sect ion.
T ak etan i (1982), Thurow (1988), Vishnevs kaya (2001). For
furthe r synonymies th e reade r is refer ed to th ese publica-
Genus TRIACTOMA ROST 1885
tions.
Triactoma cellulosa FOREMAN 1973
Class ACTINOPODA
Sub class RADIOLARIA MOLLER 1858
Sup erorder POLYCYSTINA EHRENBERG 1838 1973 Triaetoma eel/ulo sa new species F OREMAN, p. 259, pI. 2, figs 9-10: pI.
Order SPUMELLARIA EHRENBERG 1875 16, fig. 9.
Superfam ily ACTINOMMACEA (A) HAECKEL 1862, emend. 1994 Triaetorna eel/ulosa FO REMAN. - O' DOG HERTY, p. 300-301, pI. 54, figs
DE WEVER et al. 2001 19-23.
Family P ARVIVACCIDAE PESSAGNO & YANG in PESSAGNO

et al. 1989 Tria ctoma hexeris Q'DOGHERTY 1994
Subfamily ACAENIOTYLINAE YANG 1993 (Plate 1, Figs, 32- 33)
Genus ACAENIOTYLE FOREMAN 1973 1994 Triaetoma hexeris n. sp. O ' D OGII ERTY. p. 303, pI. 55, figs. 14-21.
A caeniotyle rebellis Q 'DOGHERTY 1994 Triactoma sp, aff. T. hexeris Q 'DOGHERTY 1994
(Plate 1, Figs. 21-22) (Plate 1, Fig. 34»
1994 Acaenio tyle rebel/is n. sp. O' DOGHERTY, p. 287-288 , pI. 51, figs. 5-10. Remarks. - Te st with cor tica l shell less rouded th en previous
species, but more he xagon al thiner. Por es ar e not clearl y
Acaeniotyle sp, A hexagon al.
(Plate 1, Fig. 23)
Superfamily ACTINOMMACEA (B) HAECKEL 1862, emend.
Remarks. - Test with two primar y spines well preser ved an d at DE WEVER et al. 2001
the bottom right part of th e she ll a th ird spine. The an gle be - Family ACTINOMMIDAE HAECKEL 1862
tween the three primary spines is 90 degrees (perhaps it had
origina lly four primary spin es"), Genus PSEUDOACANTHOSPHAERA Q'DoGHERTY 1994
Acaeniotyle sp, B Pseudoacanthosphaera galeata Q 'DOGHERTY 1994

(Plate 1, Fig. 24) (Plate 1, Fig. 26)
Remarks. - Specimen not ver y well pr eserved with onl y one 1994 Pseudoacanthosp haera galeata n. sp. O ' DOGHERTY, p. 297, pI. 53, figs.
16-19.
spine . Test with tub ercl es.
Family XIPHOSTYLIDAE HAECKEL 1881 Pseudoacanthosphaera superba (SQUINABOL 1904)

(Plate 1, Fig. 27)
Genus ARCHAEOCENOSPHAERA PESSAGNO & YANG in
PESSAGNO et al. 1989 1904 Trisphaera supe rba n. sp. S QUINABOL, p. 190, pI. 2, fig. 13.
1994 Pseudoacanthosp haera sup erba ( S QUINABOL). - O 'DOGHERTY, p.
298-299, pI. 54, figs. 5-10.
Archaeocenosphaera (?) mellifera Q 'DOGHERTY 1994
(Plate 2, Fig. 18-1 9)
Pseudoacanthosphaera sp, aff. P. spinosissima (SQUINABOL
1988 Hem icryptoeapsa po lyhedra D UMITRICA. - THUROW" p. 40 1, pI. 1, fig.
1904)
\. (Plate 1, Fig. 28)
1988 Hemicryptoea psa sp. cr. H. po lyhedra D UMITRICA. - TH UROW, p. 401.
pI. 5, fig. 2. Remarks. - Te st with a small spinose cortical she ll and two
1994 Arehaeoee no sp haera? m ellife ra n. sp. O ' DOGH ERTY, p. 375-376, pI.
long three-bladed primary spines .
74, figs. 1- 5.
S10 A.N. Bandini , P.O. Baumgart ner & M. Caron

Pseudoacanthosphaera (?) sp, size. Cortical shell with about ten elliptica l pore frames on
(Plate 1, Fig. 29) ea ch mamma,
Remark s. - Test with ellipsoidal cortical she ll and three, maybe Superfamily PYLONIACEA H AE CKEL 1881
four primary spines (the fourth one is not visible ). Primar y Subsup erfamily DACTYLIOSPHAERILAE SOUI NASOl 1904
spines three-bladed. Meshwork developing small secondary Family HAGIASTRIDAE RI ED EL 1971
spines at pore vertices.
Genus CRUCELLA PESSAGNO 1971
Genus TETRACANTHELLIPSIS S O UINABOL 1903
Crucella messinae P ESSAGNO 1971
Tetracanthellipsis euganeus S O UINABOL 1903 (Plate 2, Figs. 14-15)
(Plate 1, Fig. 25)
1971 Cr ucella m essina e n. sp. PESSAGNO. p. 56, pI. 6, figs. 1- 3.
1903 Tetracanthellipsis eugan eus n. sp. SQUIN ABOL, p. 117. pI. 8. fig. 9. 1975 Crucella sp. FOREMAN, p. 6 12. pI. 10. fig. 7: pI. 20. fig. 9.
1994 Tetracanth ellipsis eugan eus SQUI NABOL. - O 'OOGHERTY. p. 295-296. 1976 Crucella messinae PESSAGNO. - PESSAGNO. p. 32, pI. I , figs. 4.
pI. 53. figs. 8-10. non 1982 Crucella messinae PESSAGNO. - TAKETANI, p. 50. pI. 9. fig. 17.
1988 Crucella messinae PESSAGNO.- THUROW" p. 399, pI. 5, fig. 22.
? 1988 Crucella sp. B THUROW" p. 399. pI. 2. fig. 15.
Sup erfamily ACTINOMMACEA (C) HA ECKEL 1862, emend. 1994 Crucella messinae PESSAGNO. - O 'DOGHERTY. p. 368. pI. 70, figs.
D E WEVER et al. 2001 21-2 4, pI. 71. figs. 1-6.
Family CONOCARYOMMIDAE LI PMAN 1969 2001 Crucella cf. m essinae PESSAGNO. - VISHNEVSKAYA . p. 158, pI. 114,
fig. 10.
Genus PRAECONOCARYOMMA P ESSAGNO 1976

Crucella cachensis PESSAGN O 1971
Praeconocaryomma universa P ESSAG NO 1976 (Plate 2, Figs. 16-17)
1971 Crucella cachensis n. sp. PESSAGNO, p. 53, pI. 9. figs. 1- 3.
1976 Praecon ocaryomma un iversa n. sp. PESSAGNO. p. 42. pI. 6. fig. 14- 16. 1976 Crucella cachensis PESSAGNO. - PESSAGNO, p. 31. pI. 3, figs. 14- 15.
1982 Praecon ocaryomma un iversa PESSAGNO. - TAKETANI. p. 47. pI. 1. figs. 1982 Crucella cachensis PESSAGNO. - TAKETANJ. p. 50, pI. 9, fig. 16.
Ja - b, 4: pI. 9, fig. 4. 1988 Cruce lla cachensis PESSAGNO. - THUROW" p. 399. pI. 2. fig. 13.
2001 Praecon ocaryomma uni versa PESSAGNO. - v lSHNEVSKAYA. p. 179. pI. 1994 Crucella cachensis PESSAGNO. - O 'O OGHERTY, p. 370 , pI. 71. figs.
21, fig. 3: pI. 24. fig. I : pI. 97. fig. I : pI. 113, fig. 5: pI. 125. fig. 1- 2: pI. 15- 22.
126 , fig. I. 2001 Cruce lla cachensis PESSAGNO. - VISHNEVSKAYA, p. 158, pI. 95, fig. 5:
200 1 Praecon ocaryomma ? universa PESSAGNO. - vISHNEVSKAYA, p. 179, pI. 125, fig. 12; pI. 129, fig. 3.
pI. 2 1, figs. II.
2001 Praecon ocaryomma ex gr. un i versa PESSAGNO. - vI SHNEVSKAYA. p. Sub superfamily PATULIBRACCHIILAE PESSAGNO 1971
179. pI. 80, figs. 4; pI. 8 1. fig. I.
Family PATULIBRACCHIIDAE P ESSAGNO 1971
Praeconocaryomma californiaensis PE SSAGNO 1976 Genus HALESIUM PESSAGN O 1971

Halesium triacanthum (SO UINABO l 1903) sensu O'DOGH ER-
1976 Praecon ocaryomma californiaensis n. sp. PESSAGNO. p. 41. pI. 7. fig. 1--8. TY, 1994
1982 Praecon ocaryomma calif orniaensis PESSAGNO - TAKETANI, p. 47. pI.
(Plate 2, Fig. 10)
I, figs. Za-c: pI. 9. figs. 1- 2.
1903 Dictyastrum triacanthos n. sp. SQUI NABOL, p. 121, pI. 9, fig. 28.
Praeconocaryomma Iipmanae P ESSAG NO 1976 ? 1988 Halesium quadratum PESSAGNO. - THUROW" p. 401, pI. 2, fig. 10.
(Plate 2, Figs. 26-27) 1994 Halesium triacanthum (SQUINABOL). - O'OOGHERTY, p. 350--351, pI.
65, figs. 9-14.
1976 Praecon ocaryomma lipman ae n. sp. PESSAGNO, p. 41-42, pI. 4, fig.
12- 13.
Halesium sp.
1982 Praecon ocaryomma lipma nae PESSAGNO. - TAKETANI. p. 47. pI. 9. fig. 3.
Praeconocaryomma sp,
Remarks. - Test with 3 relatively thick and small ra ys,
Genus PESSAGNOBRACCHIA K O WR & MosTLER 1978

Rema rks. - Elliptical cortical shell in outline with numerous
equally spaced, cone-like mammae. Cortical shell with or with-
Pessagnobracchia sp,
out rad ial spines circular in cros s-section projecting from cen-
(Plate 2, Fig. 13)
tcr of each mamma. Cortical she ll with pore frames of uniform
Tu ron ian Radiolarians from Karnezeika S11

Remarks. - Spongy three -ra y ed t e st with irregular arrang e - Patellula ecliptica O 'D OGH ERTY 19 94
ment o f p ores on rays. ( P late 2, Figs. 3-4)
Family PSEUDOAULOPHACIDAE RI EDEL 1967 1994 Patellula ec/iptica n. sp. O ' OOOHERTY, p. 329, pI. 61, figs. 1-5 .
Subfa mily PSEUDOAULOPHACINAE RIED EL 1967 Patellula heroica O 'DOGH ERTY 19 94
( P late 2, Figs . 5-6)
Genus DACTILYODISCUS S OUINABOL 1903 1994 Patellula heroica n. sp. O'OOOHERTY, p. 330, pI. 61, figs. 6-11.
Patellula sp,
Dactyliodiscus sp, ( P la te 2, Fig. 7 )
( P late 2 , Figs. 8-9 )
Remarks. - T est flattened e lli pso idal w it h fifteen spine s r adiat-
Remarks. - T est is disc-shaped a n d circu la r in outline w ith a ing in the same equatorial plane .
v ariable number of equatorial s pines. M e shwork spongy w ith
irregul arly pore frames. Poorly d efined central raised area. Order ENTACTINARIA K OZUR & MOSTLER 1982
Upper a n d lower surfaces of the test with small tubercles. Family SATURNALIDAE D EFLANDRE 1953
Subfamily SATURNALINAE D EFLANDRE 1953
Genus PSEUDOAULOPHACUS P ESSAGNO 1963
Genus ACANTHOCIRCUS S OU1NABOL 1903
Pseudoaulophacus sculptus (SOU INA BOL 1904) sensu 0'-
D OGHERTY 1994 Acanthocircus venetus (S OU1NABOL 1 9 14) s e n s u O'D OGH ERTY
(P la t e 1 , Figs. 35-36) 1994
( P late 1, Figs. 15-16)
1904 Theodiscus sculptus n. sp. SQUII'ABOL, p. 200, pI. 4, fig. 9.
1988 Alievium superbum « Ce no manian » for m (SQUINABOL). - THUROW.. 1914 Saturnalis venetus n. f. SQUINABOL, p. 269, 299. pI. 20[1]. fig. 2; pI.
p. 397, pI. 5, fig. 11. 24[5], fig. 1.
1988 A lievium sp. A. T HUROW, p. 397, pI. 5, fig. 12. ? 1975 Spo ngosaturnalis horridus (SQUINABOL). - FOREMAN, p. 610, pI. 4,
1994 Pseudoauloph acus sculptus (SQUINABOL). - O'OOOHERTY, p. fig. 3.
319-320 , pI. 59, figs. 1--4. 1975 Spongosaturnalis hueyi gro up (PESSAONO) . - FOREMAN, p. 611, pI.
1B, figs. 1-3; non pI. 1A . figs. 7- 8 (=A. tym panum ?).
1975 Spo ngosaturnalis (?) prec/arus new species FOREMA N, p. 611, pI. l A ,
Pseudoaulophacus putahensis P ESSAGNO 1972
figs. 4--5; pI. 4, fig. 8.
( P late 1 , Fig s . 3 7-38) 1994 Acanthoci rcus venetus (SQ U INA BOL). - O 'OOOHERTY, p. 256, pI.
45, figs. 1--8.
1972 Pseudoaulophacus putahensis n. sp. PESSAONO, p. 301, pI. 27, fig. 1.
1976 Pseudoaulophacus putahensis PESSAONO. - PESSAONO, p. 28, pI. 3, fig.
13. Acanthocircus tympanum O'DOG HERT Y 1994
1988 Pseudoaulophacus put ahensls PESSAONO. - T HUROW" p. 404, pI. 2, fig. ( P lat e 1, Figs. 17-18)
4.
1994 Pseudoaulophacus putah ensis PESSAONO. - O'OOOHERTY, p. 320-321,
pa rs 1975Spongosaturnalis hueyi gro up (PESSAONO). - FOREMAN, p. 611, pI.
pI. 59, figs. 5-13.
l A, figs. 7-8; non pI. IB, figs. 1-3 (=A. venetus?)
2001 Pseudoaulophacus putah ensis PESSAONO. - VISHNEVSKAYA, p. 181, pI.
1994 A canthocircus tympanum n. sp . O'OOOHERTY, p. 259-260, pI. 45,
130, fig. 7.
figs. 17-24.
Sup erfamily SPONGURACEA H AECKEL 1862

Acanthocircus hueyi ( PESS AGNO 1976 ) s e n s u O 'DOGHERTY 19 94
Family SPONGURIDAE H AECKEL 1862
( P late 1 , Fig s. 19-2 0 )
Genus PATELLULA K OZLOVA 19 72 1975 Spongosaturnalis hueyi (PESSAONO). - FOREMAN, p. 611, pI. 1A , fig. 6;
pI. 4. fig. 10.
Patellula helios ( S OU INA BOL 19 03 ) s e n s u O 'DOGHERTY 19 94 1976 Spo ngosaturnalis hueyi n. sp. PESSAONO, p. 39, pI. 12, fig. 1.
( P late 2, F ig s . 1-2) 1994 Acanthocircus hu eyi (PESSAONO). - O' DOOHERTY, p. 260-261, pI. 46,
figs. 1- 5.
200! Spo ngosaturnalis hueyi (PESSAONO). - VISHNEVSKAYA. p. 186, pI. 122,
1903 Stylotrochus helios n. sp. SQUINABOL, p. 124, pI. 10, figs. 23-23a.
fig. 2.
1976 Pseudoau lophacus lenticulatus (WH ITE). - PESSAONO, p. 28, pI. 9,
200! Spongosa turnalis ex. gr. hu eyi (PESSAONO). - VISHNEVSKAYA, p. !86,
figs. 11-1 2.
pI. 92, figs. 7--8, pI. 95, figs. 1-3.
1982 Pseudoa ulophacus lenticulatus (WH ITE) . - TAKETANI, p. 51, pI. 10,
fig. 11.
1985 Pseudoaulophacus lenticulatus (W H IT E) . - SANFILIPPO & RIEDEL, p. Order NASSELARIA EH RENBERG 1875
596, te xt-figs. 6.4a- b. Superfam ily ARCHAEDICTYOMITRACEA PESSAGNO
1994 Patellula helios (SQUINABOL). - O'OOOHERTY, p. 327-328, pI. 60, figs.
1976
19-24.
S1 2 A .N. Bandini, P.O. Baumgartner & M . Caron

Family ARCHAEDICTYOMITRIDAE PESSAGNO 1976 1988 Theocon us sp. A cf. T. co ronatus SOUINABOL. - TH UROW.. p. 407, pI.
4, figs. 3-4.
2001 Spo ngo caps ula aff. zam o raensis (PESSAGNO). - V ISHNEVSKAYA, p.
Genus DICTYOMITRA ZITTEL 1876 186. pI. 77, fi g. 5; pI. 90. f igs. 5-6.
Dietyom itra formosa SQ UIN ABOL 1904

Sup erfamily EUCYRTIDIACEA EHRENBERG 1847
family PSEUDODICTYOMITRIDAE PESSAGN O 1 977
1904 Dictyomi tra fo rmosa n. sp. SOUINABOL. p. 232. pI. 10. fi g. 4.
1974 Dictyomitra to rq uata FOREMAN. - RI EDEL & SANFILIPPO . p. 778. pI. 5. Genus PSEUDODICTYOMITRA P ESSA GNO 1977
figs. 1.2 and 4.
1975 Dictyomitra du odecim costata (SQUINABOL). - FOREMAN. p. 6 14. pI. 7. Pseudodietyomitra pseudomacroeephala ( SQU INABOL 1903)
fig. 8; pI. IG. fi g. 5.
1976 Dictyom itra fo rmosa SOUINABOL. - PESSAGNO. p. 5 1. pI. 8. fi gs. 10-12.
(Plate 1, Figs. 7-8 )
1982 D ictyomitra formosa SOUINABOL. - T AKETANI. p. 58. pl. 4. f igs. 6a-b:
pI. I I, f ig. 13. 1903 D ictyomitra pseud omacrocepha la n. sp. SQUINABOL,p. 139, pI. 10, fig. 2.
1985 D ictyomitra formosa SOUINABOL. - SCIIAAF" te xt-fig.I 1. p. 250. 1974 Dictyomitra ma crocephala SOUINABOL. - RIEDEL & SANFILIPPO,
1988 D ictyo m itra formosa SOUINABOL - T HUROW.. p. 400, pI. I, fi gs. 23 p. 778. pI. 4. figs. 10--11; pI. 14, fig . 11.
and 25. 1975 Dictyomitra pseudomacrocephala SOUINABOL. - DUMITRI CA.. tex t-
1994 D ictyom itra formosa SQUINABOL. - O' DOGHERTY. p. 80. pI. 4, fi gs. fig.2 ,19.
8- 12. 1975 Dictyomitra pseudomacroceph ala SOUINABOL. - FOREMAN. p. 6 14,
2001 Di ctyomitra formosa SOUINABOL. - VI SHNEVS KAYA, p. 160, pI. 25. pI. 7, fig. 10.
fig . 10. 1976 Dictyomitra (?) pseudoma crocephala SQUINABOL. - PESSAGNO. p. 53.
pI. 3, figs. 2-3.
1982 Pseudodictyomitr a pseudom acroceph ala (SOUINABOL) . - T AKETANI,
Dictyomitra sp, cr. Di formosa S Q UIN A BO L 1904 p. 61, pI. 5. fi gs. 4a-b; pI. 12, figs. 7-8.
(Plate 1, Fig. 3) 1985 Pseu dod ictyomitra pseudomacrocephala (SOUINABOL). - SANFILIPPO
& RI EDEL. p. 608, text -figs. 10. l a- b.
(species number 2)
1985 Pseudodictyo m itra pseudoma croceph ala (SOUINABOL). - SCHAAF.
Dictyomitra montisserei (SQUINABO L 1903) sensu O 'D OGH E R- text-fig.11. p. 250
TY 19 94 1988 Pseudod ictyom itra pseudoma crocephala (SOUINABOL). - T HUROW"
(Plate I, Figs. 4-5 ) p. 405. pI. 1. fi g. 13: pI. 3, figs. 11-1 6.
1994 Pseudodictyomitra pse udomacrocephala (SOUINABOL).- O 'O OGHERTY.
p. 108- 109. pI. 8. figs . 5-8.
1903 Stichopho rmis Mon tis Serei n. sp. SOUINABOL. p. 137. pI. 8, f ig. 38.
2001 Pseudodictyom itra pseudomacrocephala (SOUINABOL) . - VISIl-
1975 Dictyomitra sp. A. FOREMAN. p. 615. pI. I G . fig. 7; pI. 2G . f igs. 18 and
NEVSKAYA. p. 183- 184, pI. 20. fig. 6; pI. 24, fi g. 10; pI. 100, fig . 3;
20.
pI. 129. figs. 5, 9 and 10.
1982 Archa edictyomitra sp. A . T AKETANI, p. 58. pI. 4. fi gs. Sa- b.
? 1988 A rchae dic tyo rnitra lacrim ula (FOREMAN). - THUROW.. p. 397. pI. 3.
fi g. 8. Family EUCYRTIDIIAE EHREN BER G 1847
1988 A rchae dictyo m itra simplex PESSAGNO. - TH UROW.. p. 398, pI. 3, fig . 9.
1994 Dictyomi tra m ontisserei (SQUINABOL 1903b). - O 'DOGHERTY, p. 77.
pI. 3. fi gs. 1-29.
Genus STICHOMITRA CA Y EUX 1897
Stichomitra communis SQ UINABOL 1903

Dictyomltra urakawensis (TAKETANI1982)
(Plate 1, Fig. 6)
1903 Sticho m itra communis n. sp. SOUINABOL, p. 141, pI. 8, fi g. 40.
? 1975 Dictyomitra sp. - D UMITRICA.. text-fig.Z; 8.
1975 Sticho m itra sp. O UMITRICA" text -fig.2, 21.
1982 Dictyom itra uraka wen sis n. sp. T AKETANI. p. 59. pI. 4. fi gs. 8a-b: pI.
1975 Sticho m itra spp. cf. D. tek schaensis A LIEV. - FOREMAN, p. 6 15. pI. 2H,
11. f ig. 16.
fi gl.
1982 Stic ho m itra co m m unis SOUINABOL. - TAKETA NI. p. 54. pI. 3. fig. 9;
Sup erfamily AMPHIPYNDACEA RI ED EL 1967 pI. I I . fig. 5.
Family SPONGOCAPSULIDAE P ESSA GNOI977 1988 Stic ho mitra co mmunis SOUINABOL. - T HUROW" p. 406. pI. 4. fi g. 10.
1988 Sticho mitra sp. cf. S. co m m un is SOUINABOL. - TH UROW" p. 406, pI. 4.
fi g. 9.
Genus TORCULUM O'D OGHERT Y 19 9 5 1994 Sti cho mitra co m m unis SOUINABOL. - O 'OOGHERTY, p. 144-1 45,
pI. 17. fi gs. 6-16.
Toreulum eoronatum ( SQUINABOL I904) 200 1 Sticho mi tra com m unis SOUINABOL. - V ISHNEVSKAYA. p. 188. pI. 23.
fig. 8; pI. 79. fig. 3; pI. 129. fig. 8.
(Plate 1, Fig. 9)
1904 Th eo con us co ronatus n. sp. SOUINABOL. p. 220. pI. 8. fig . 3. Stichomitra stocki ( C A MP BELL & CLA RK 1944) sensu 0 '-
1976 Sticho m itra (?) zam o raensis n. sp. PESSAGNO. p. 54. pI. 3. fi gs. 7-9. D OGHERTY 1994
1982 Spo ngocaps ula (?) zam o raensis (PESSAGNO). - TA KETANI. p. 62. pI. 5. (Plate 1, Figs. 12-13 )
figs. 6a-b; pI. 12, fi gs. 12-13.
1988 Theocon us coronatus Gro up SOUINABOL.- TH UROW" p. 407, pI. 4, fi gs. 2.
1944 Stichocapsa (?) stock i n. sp, CAMPBELL & CLARK, p. 44, pI. 8, fi gs. 31-33.
Turoni an Radiolarians from Karnezeika S 13

1974 Amphipyndax stocki (CAMPBELL & CLARK). - RIEDEL & SANFILIPPO, Discussion and Conclusions
p.775, pI. 15, fig. 11; pI. 11, figs. 1-3.
1975 Amphipyndax stocki (CAMPBELL & CLARK). - DUMITRICA, text- The radiolarian chronostratigraphic ranges used for this paper
fig.2.23. are based on the comparison of taxon ranges established by 10
1982 Amphipyndax stocki (CAMPBELL & CLARK). - TAKETANI, p.52, pI. 2,
figs. 9a-b; pI. 10, figs. 13-14. different authors (see list in introduction). By stacking
1982 Amphipyndax sp. TAKETANI, p.52, pI. 10, fig. 16. chronostratigraphic ranges, we obtain a maximum range for
1988 Stichomitra (?) sp. A. THUROW" p.406, pI. 1,fig. 17. the existence of each taxon. Despite possible inaccuracies of
1994 Stichomitra stocki (CAMPBELL & CLARK). O'DOGHERTY, p. 147-148 calibration, the radiolarian age given by this procedure is con-
and 150, pI. 18, figs. 9-15.
sistent with the age based on planktonic foraminifera (late
2001 Amphipyndax stocki (CAMPBELL & CLARK). - VISHNEVSKAYA, p.146,
pI. 1,fig. 13; pI. 4,fig. 11-13; pI. 6,fig. 12; pI. 16, fig. 1;pI. 93, fig. 4-5; Turonian to Coniacian). In combining the radiolarian and the
pI. 94, fig. 8 and 10; pI. 99, fig. 1-3and 8;pI. 114, fig. 12. planktonic foraminifer ages, the samples would be restricted to
2001 Amphipyndax stocki (CAMPBELL & CLARK) var. A VISHNEVSKAYA.- the late Turonian. However, the major discrepancies of pub-
VISHNEVSKAYA, p.146-147, pI. 16, fig. 2-6; pI. 26, fig. 6;pI. 100, fig. 4; lished radiolarian ranges call for an urgent, major revision of
pI. 123, fig. 16-21 and23.
2001 Amphipyndax stocki (CAMPBELL & CLARK) var. B VISHNEVSKAYA.- the Late Cretaceous radiolarian biochronology, a project that
VISHNEVSKAYA, p147, pI. 3,fig. 6;pI. 12, fig. 3and 5;pI. 15, fig.I-5. is underway (Jackett et al. 2002 and Diserens et al. 2003). The
2001 Amphipyndax stocki (CAMPBELL & CLARK) var. C VISHNEVSKAYA.- integration of planktonic foraminifera with radiolarians may
VISHNEVSKAYA, p.147, pI. 3,fig. 2 and 4;pI. 14, fig. 1-3.V greatly enhance biochronologic resolution in sections where
both groups occur.
This is the first time that Late Cretaceous radiolarians are
described from the Argolis Peninsula.
NASSELLARIA INCERTAE SEDIS
Genus AFENS RIEDEL & SANFILIPPO 1974 Acknowledgements

The field work in the Argolis Peninsula was partly financed by the Societe
A/ens liriodes RIEDEL & SANFILIPPO 1974 Acadernique Vaudoise and the University of Lausanne, Switzerland. The
(Plate 1, Fig. 14) salaries are paid by the Swiss National Science Foundation (grant n° 2000-
063762 and n° 200021-105845). We are thankful for the loan of the Vernez'
thin sections bythe Musee Cantonal de Geologie, Lausanne, Switzerland. We
1974 Afens liriodes new genus and new species RIEDEL & SANFILIPPO, p.
aregrateful forhelp in thefield byFrance Girault and Thierry Glassey; assis-
775, pI. 11, fig.11; pI. 13,figs. 14-16.
tance in the laboratory by Sebastien Bruchez, Marc-Olivier Diserens and
1985 Afens liriodes RIEDEL & SANFILIPPO. - SANFILIPPO & RIEDEL, p.624,
Sarah-Jane Jackett; and for discussions with Paulian Dumitrica. The manu-
text-figs. 13.3a--c. script has benefited from careful reviews by Atsushi Matsuoka and Luis 0'-
1985 Afens liriodes RIEDEL & SANFILIPPO. - SCHAAF, text-fig.! 1,p.250.
Dogherty.
1988 Afens liriodes RIEDEL & SANFILIPPO. - THUROW" pI. 2,fig.1.
1994 Afens liriodes RIEDEL & SANFILIPPO. - O'DOGHERTY, p. 246, pI. 42,
figs. 23-26.
SPECIES LIST
Planktonic Foraminifera
Acaeniotyle rebellis O'DOGHERTY 1994 (PI. 1, Figs. 1-2)
In the following, we use the ranges proposed by Caron (1985).
Acaeniotyle sp. A (PI. 1, Fig. 3)
The first pelagic facies at the top of the Pantokrator Limestone
Acaeniotyle sp, B (PI. 1, Fig. 4)
is dated as middle Turonian by the presence of Helvetoglo-
Acanthocircus hueyi ( PESSAGNO 1976) sensu O'DOGHERTY
botruncana helvetica (PI. 3, Fig. 1, early Turonian - middle
1994 (PI. 2, Figs.18-19)
Turonian) and Marginoglobotruncana marianosi (PI. 3, Fig. 2,
Acanthocircus tympanum O'DOGHERTY 1994 (PI. 2, Figs.
middle Turonian -late Turonian).
16-17)
In the pelagic limestone rich in Globotruncanidae at the
Acanthocircus venetus (SQUINABOL 1914) sensu O'DOGHERTY
top of the radiolarian cherts, the presence of Dicarinella prim-
1994 (PI. 2, Figs. 14-15)
itiva (PI. 3, Fig. 9, middle Turonian - Coniacian), Margin-
A/ens liriodes RIEDEL & SANFILIPPO 1974 (PI. 2, Fig. 33)
otruncana sigali (PI. 3, Figs. 5 and 12, middle Turonian - early
Archaeocenosphaera (?) mellifera O'DOGHERTY 1994 (PI. 1,
Santonian), M. schneegansi (PI. 3, Figs. 3 and 7, middle Tur-
Figs. 5-6)
onian - early Santonian), M. coronata (PI. 3, Fig. 4, middle
Archaeocenosphaera (?) sp. (PI. 1, Figs. 7-8)
Turonian - early Campanian), M. pseudolinneiana (PI. 3, Figs.
Crucella cachensis PESSAGNO 1971 (PI. 1, Figs. 29-30)
6,10 and 11, middle Turonian - early Campanian) and M. sin-
Crucella messinae PESSAGNO 1971 (PI. 1, Figs. 27-28)
uosa (PI. 3, Fig. 8, late Turonian - early Santonian) allow to
Dactyliodiscus sp, (PI. 2, Figs. 1-2)
state a late Turonian to early Santonian age without further
Dictyomitra formosa SQUINABOL 1904 (PI. 2, Figs. 20-21)
precision.
Dictyomitra montisserei (SQUINABOL 1903) sensu O'DOGHERTY
1994 (PI. 2, Figs. 23-24)

Dictyomitra sp, cf. Di formosa SQUINABOL 1904 (PI. 2, Fig. 22) BAUMGARTNER, P.O., GUEX, J. & DUMlTRICA, P. 1995: Concepts of the sys-
tematic and biostratigraphic work. In: BAUMGARTNER, P.O et al. (Eds.):
Dictyomitra urakawensis T AKETANI 1982 (PI. 2, Fig. 25) Middle Jurassic to Lower Cretaceous Radiolaria of Tethys: Occurrences,
Halesium sp, (PI. 1, Figs. 32-33) Systematics, Biochronology, 11-15. Memoires de Geologie (Lausanne),
Halesium triacanthum (SQUINABOL 1903) sensu O'OOGHERTY 23. Institut de Geologie et de Paleontologic, Universite de Lausanne.
1994 (PI. 1, Fig. 31) CAMPBELL, A.S. & CLARK, B.L. 1944: Radiolaria from Upper Cretaceous of
Middle California. Geological Society of America Special Paper, 57,1-61.
Patellula ecliptica O'OOGHERTY 1994 (PI. 2, Figs. 9-10)
CARON. M. 1985: Cretaceous planktic foraminifera. In: BOLLI, H.M.,
Patellula helios (SQUINABOL 1903) sensu O'OOGHERTY 1994
SAliDERS, J.B. & PERCH-NIELSEN, K. (Eds.): Plankton stratigraphy, 17-86.
(PI. 2, Figs. 7-8) Cambridge University Press, Cambridge! New York.
Patellula heroica O'OOGHERTY 1994 (PI. 2, Figs. 11-12) DE WEVER, P. & ORIGLIA-DEVOS, I. 1982: Datations nouvelles par les Radio-
Patellula sp, (PI. 2, Fig. 13) laires de la serie des Radiolarites s. I. du Pinde-Olonos, (Grece). C. R.
Acad. Sc. Paris, 294, 399-404.
Pessagnobrachia sp, (PI. 1, Fig. 34) DE WEVER, P. & THIEBAULT, F. 1981: Les Radiolaires d'age jurassique superieur
Praeconocaryomma californiaensis PESSAGNO 1976 (PI. 1, acretace superieur dans les radiolarites du Pinde-Olonos (Presqu'ile de Ko-
Figs. 21-22) roni; Peloponnese meridional, Grece), Geobios, 14(5), 577-609.
Praeconocaryomma lipmanae PESSAGNO 1976 (PI. 1, Figs. DE WEVER, p .. GEYSSANT, J.R., AZEMA, J .. DEVOS, I., DUEE, G., MANIVIT, H.
& VRIELYNCK. B. 1986: La coupe de Santa Anna (zone de Sciacca, Sicile):
23-24)
une synthcse biostratigraphique des apports des macro-, micro- et nan no-
Praeconocaryomma sp, (PI. 1, Figs. 25-26) fosiles du Jurassique superieur et Cretace inferieur. Revue de Micropa-
Praeconocaryomma universa PESSAGNO 1976 (PI. 1, Figs. leontologie, 29 (3),141-186.
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Fig. 14) nation of late Cretaceous radiolarites in orogenic environments: an exam-
Pseudoacanthosphaera sp. aff. P. spinosissima (SQU[NABOL ple from accreted teeranes of southern Costa Rica. Interrad X 2003, Ab-
1904) (PI. 1, Fig. 16) stracts & Programme, University of Lausanne, Switzerland, 49-50.
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Bahama Basin (Site 5A, JOIDES Leg 1) and the Great Valley Sequence, THUROW, J. 1988: Cretaceous radiol arians of the North Atlantic Ocean; ODP
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5-83. (Site 398). In: BOILLOT, G. , WINTERER, E .L., (Eds.): Proceedings of the
PESSAGNO, E.A . 1976: Radiolarian zonati on and stratigraphy of the Upper Ocean Drilling Program, Scientifi c results (G alicia margin; coverin g Leg
Creta ceous portion of the Great Valley Sequ ence , California Coast 103 of the cruises of the drilling vessel JOIDES Resolution, Ponta Delga-
Range s. Micropaleontology spec . Publ ., 2, 1-95. da, Azores, to Bremerhaven, Germany , 25 April 1985-19 June 1985), 103,
RIEDEL, W.R . & SANFILIPPO, A. 1974: Radiolari a from the southern Indian 379-4 18, Texas A. & M. Uni versity, Ocean Dr illing Program, Colle ge Sta-
Ocean , DSDP Leg 26. In: DAVIES, T.A. , et al. (Eds.): Initial Rep ort s of tion , TX , United Stat es.
the Deep Sea Drilling Proj ect , 26 (D urba n, South Afric a to Frernantle, VERNEZ, G. 1990: Etude geo logique et miner alogique de la Vallee de Paliorni-
Au stralia; Sept-Oct, 1972),771-781. U.S. Gove rnment Printing Office, nos entre Karnezeika et Stavrop odh i. Unpublished diplom a the sis, Uni-
Washington, D.C. versity of Lau sanne .
SANFILI PPO, A. & RIEDEL, W.R. 1985: Creta ceous Radi olaria. In: BOLLI , H.M., VISHNEVSKAYA, V. S. 2001: Jur assic to Cretace ous radiolarian biostratigraph y
SAUDERS, J.B. & PERCH-NIELSEN, K. (Eds.): Plankton stratigraph y, of Ru ssia. GEOS, Moscow , 376 p.
631-7 12. Cambridge Un iversit y Pres s, Ca mbridge/ New York . VRIELYNCK, B. 1981: Evolution paleogeographique et structurale de la
SAVARY, J . & G UEX, J . 1991: BioGraph : un nouveau pro gramme de con strue- presqu'tle d'Argolide (G rece) , Revue de geologie dynamique et de geo-
tion des correlations biochronologiqu es basees sur les associati ons uni- graphie physique , 23(4), 277-288.
tair es. Bulletin de la Societe Vaudoise des Scienc es Naturelles, 80 (3) ,
317- 340. Manu script received Januar y 2004
Revision accepted February 2005
Plate 1
SEM·lllustrations of Upper Cretaceous radiolarians from Karnezeika, Argolis Peninsula (Greece)

Figures 1-2 Acaeniotyle rebellis O'DOGIIERTY 1994 A168_06 (Figs. 1 and 2)
Figure 3 Acaeniotyle sp, A A174_300
Figure 4 Acaeniotyle sp, B A170_090
Figures 5-6 Archaeocenosphaera (?) melllfera O 'DOGHERTY 1994 AI72_190 (Figs. 5 and 6)
Figure s 7-8 Archaeocenosphaera (?) sp, A173_320 (Figs. 7 and 8)
Figures 9- 10 Triactoma cellulosa FOREMAN 1973 AI74_300 (Figs. 9 and 10)
Figur es 11-12 Triaetoma hexeris O 'D OGHERTY 1994 A173_320 (Figs. 11 and 12)
Figure 13 Triactoma sp. aff. T. hexeris O 'D OGHERTY 1994 AI72_19O
Figure 14 Pseudoacanthosphaera galeata O'D OGHERTY 1994 A173_320
Figure 15 Pseudoacanthosphaera superba (SOUINABOL 1904) A174_300
Figure 16 Pseudoacanthosphaera sp, aff, P. spinosissima (SOUINABOL 1904) A173_320
Figure 17 Pseudoacanthosphaera (?) sp. A173_320
Figure 18 Tetracanthellipsis euganeus SOUINABOL1903 A173_320
Figures 19-20 Praeconocaryomma universa PESSAGNO 1976 AI72_190 (Figs. 19 and 20)
Figures 21-2 2 Praeconocaryomma califomiaensis PESSAGNO 1976 AI72_19O(Figs. 21 and 22)
Figure s 23-2 4 Praeconocaryomma lipmanae PESSAGNO 1976 AI74_300 (Figs. 23 and 24)
Figure s 25-26 Praeconocaryomma sp, A174_300 (Fig.25 and 26)
Figure s 27-28 Crucella messinae PESSAGNO 1971 AI68_060 (Fig. 27), AI72_19O(Fig. 28)
Figure s 29- 30 Crucella cachensis PESSAGNO 1971 AI74_300 (Fig. 29), A173_320 (Fig. 30)
Figur e 31 Halesium triacanthum (SOUINABOL 1903) sensu O 'D OGH ERTY 1994 A168_060
Figure s 32-33 Halesium sp, A168_060 (F ig. 32), A173_320 (Fig. 33)
Figures 34 Pessagnobrachia sp. A174_300
S16 A.N . Bandini, P.O. Baumgartner & M. Caron

Turonian Radiolarians from Karnezeika S17
Plate 2
SEM·iIIustrations of Upper Cretaceous radiolarians from Kamezeika, Argolis Peninsula (Greece)

Figures 1-2 Dactyliodiscus sp, AI72_190 (Figs. I and 2)
Figures 3-4 Pseudoaulophacus sculptus (SQUINABOL 1904) sensu O'D OGHERTY1994 A I72_190 (Figs. 3 and 4)
Figures 5-6 Pseudoaulophacus putahensis PESSAGNO 1972 A173_320 (Figs. 5 and 6)
Figures 7-8 Patellula helios (SQUINABOL 1903) sensu O'D OGH ERTY 1994 AI74_300 (Figs. 7 and 8)
Figures 9- 10 Patel/ula ecliptica O'D OGHERTY1994 A173_320 (Figs. 9 and 10)
Figures 11- 12 Patel/ula heroica O'D OGHERTY 1994 AI74_300 (Figs. I I and 12)
Figure 13 Patel/ula sp. AI72_190
Figures 14-15 Acanthocircus venetus (SQUINABOL 1914) sensu O'DOGHERTY1994 AI72_19O (Fig. 14), A173_320 (Fig. 15)
Figures 16-17 Acanthocircus tympanum O 'DOGHERTY 1994 AI74_300 (Fig. 16), A173_320 (Fig. 17)
Figures 18-19 Acanthocircus hueyi (PESSAGNO 1976) sensu O'D OGHERTY 1994 A174_300 (Fig. 18), A173_320 (Fig. 19)
Figures 20-2 1 Dictyomitraformosa SQUINABOL 1904 Al72_19O (Fig. 20), A173_320 (Fig. 21)
Figure 22 Dietyomitra sp. cr. Di formosa SQUINABOL 1904 AI72_19O
Figures 23-2 4 Dictyomitra montisserei (SQUINABOL1903) sensu O'D OGHERTY 1994 AI68_060 (Fig. 23), AI72_19O (Fig. 24)
Figure 25 Dictyomitra urakawensis TAKETANI 1982 AI72_190
Figure 26 Torculum coronatum (SQUINABOL 1904) AI72_190
Figures 27-28 Pseudodictyomitra pseudomacrocephala (SOUINABOL 1903) A173_320 (Figs. 27 and 28)
Figures 29- 30 Stichomitra communis SQUINABOL 1903 A173_320 (Figs. 29 and 30)
Figures 31- 32 Stichomitra stocki (CAMPBELL & CLARK 1944) sensu O 'D OGHERTY1994 A168_060 (Fig. 31), A1 73_320 (Fig. 32)
Figure 33 Afens liriodes RIEDEL& SANFILIPPO 1974 A174_300
SI8 A N. Bandini , P.O. Baumgartner & M. Caron

Turonian Radiolarians from Karnezeika S19
Plate 3
Optical microscope illustrations of Upper Cretaceous planktonic foraminfers from Kamezeika, Argolis Peninsula (Greece)
Figure 1 Helvetoglobotruncana helvetica (BOLLI 1945) CKAR2
Figure 2 Marginotruncana marianosi (DOUGLAS 1969) CKAR7
Figure 3 Marginotruncana schneegansi SIGAL1952 CKAR13
Figure 4 Marginotruncana coronata (BOLLI 1945) CKAR13
Figure 5 Marginotruncana sigali (REICHEL1950) CKAR13
Figure 6 Marginotruncano pseudolinneiana PESSAGNO 1967 CKAR14
Figure 7 Margtnotruncana schneegansi SIGAL1952 CKAR14
Figure 8 Marginotruncana sinuosa PORTHAULT 1970 CKAR14
Figure 9 Dicarinella primitiva (DALBIEZ1955) CKAR14
Figure 10 Morginotruncana pseudolinneiana PESSAGNO 1967 AI76_1350
Figure 11 Marginotruncana pseudolinneiana PESSAGNO 1967 AI76_1350
Figure 12 Marginotruncana sigali (REICHEL 1950) AI76_1350

0012-9402/06/01S021-13 Eclogae geol. Helv. 99 (2006) Supplement 1, S21-S33
DOI10.1007/s00015-006-0607-0
Birkhauser Verlag, Basel, 2006
Radiolarian correlation of Jurassic siliceous successions of the Rosso

Ammonitico Formation in the Southern Alps and Western Sicily (Italy)
PAOLA BECCARO
Key words: Radiolarians, Amm onit es. Biostrat igraph y, Unit ary Associat ions Zones, Rosso Ammonitico Medio, Middle-Late Jurassic,Italy
Parole chiave: Radiolar i, Amm oniti , Biostr at igrafia, Zone ad Associazioni Unitarie , Rosso Amm on itico Medio, Gi urassico medio-super iore, Italia
A BST RACT RIASSUNTO
This paper deals with the radiolarian biostratigraphy of Middle-Upper Jurassic La ricerca riguarda la biostratigrafia a radiolari di successioni pelagiche silicee
pelagic siliceous successions of the Southern Alps and Western Sicily (Italy ). riferibili al Giurassico Medio-Superiore delle Alpi Meridionali e della Sicilia
Th e crucial complement to this research is the occurrence of ammonites in the occide ntale (Italia). L'asp etto piu impo rtante e la presenza di ammoniti sia
studied successions (Rosso Amm on itico Medi o: the intermediate siliceou s nelle successioni studiate (Rosso Amm onitico Medio: porzione interm edia si-
memb er of the Rosso Ammonitico Formation ), as well as in the under- and licea della Formazione del Rosso Amm on itico) sia in quelle sotto- e sovra-
overlying sediments (Rosso Ammonitico Inferiore and Rosso Ammonitico Su- stanti (rispell ivame nte Rosso Amm onitico Inferiore e Rosso Amm on itico Su-
periore, respectively). The abund ance of rad iolarians in all successions allow per iore). L'abb ond anza di radiolari in tull e Ie sezioni studiate ha perm esso di
to analyse them for a twofold purp ose: to date directly most of (he successions, analizza rli sia per datar e dirett arnent e la maggior parte delle successio ni sia
and to impro ve the calibra tio n of radiolar ian biozones thanks to the occur- per miglior are la calibrazione delle biozone a rad iolari grazie alia presen za
rence of ammonites. The biostr atigraph ic analysis has been carried out using delle ammoniti. L'a nalisi biostr at igrafica e stata effettuata con il met odo delle
the Unita ry Associations meth od, and six new radiolarian biozone s have been Associazioni Unitarie e sono state definite sei nuove biozone a radiolari: la co-
defined: the combined occur rence of radiolarians and ammonites provided a pre senza di radiolari ed ammoniti ha cosl fornit o una nuo va zonazione a ra-
new Bath onian to late Kimmerid gian radiolarian zonat ion for the South ern diolari per l'int ervallo di temp o Batonian o-Kimm erid giano delle A lpi Meri-
Alps and Western Sicily (Ita ly). The new Unitary Association Zone s show a dionali e della Sicilia occident ale (Italia). Le Zone ad Associazioni Unitarie
good reprodu cibility throughout the investigate d successions, and mak e possi- ide ntificate mostr ano una buo na riproducibilita tra Ie sezioni stra tigrafiche
ble a first direct da ting and cor relatio n by rad iolarians of the Rosso Am- analizzate ed hanno perm esso una prim a datazione e correlazione dirett a a ra-
monitico Medi o. Fur therm ore, the radiolar ian biozones revea l a significant di- diolar i del Rosso Ammonitico Medio. Le biozone a radiolari hanno inoltre ri-
achro nism for both the lower a nd the uppe r limit of the Jurassic pelagic velato un significativo diacron ismo sia per il limite inferiore sia per il limite su-
siliceo us facies in the Alpine and Sicilian sections. In the light of new radio lar- per iore de lle facies pelagiche silicee nelle sezioni alpine e siciliane. Alia luce
ian biozo nes, the age assignme nts of the Ceniga (Southern Alps) and the San- dei nuovi dati biostrat igrafici forniti dai radiolar i sono infine discusse Ie eta
t ' Ann a (Sicily) sections, and the ranges of some taxa are discussed . della facies silicea nelle sezioni di Ceniga (Alpi Merid ionali ) e di Sant ' Ann a
(Sicilia) e la distribuzi one stratigrafica di alcuni taxa.
Introduction
The Sicilian chain and the Southern Alps represent seg-
Th e study of siliceous successions in the Southern Alps and in ments of the Alpine collision al belt formed during the "T er-
Western Sicily (Italy) benefits from the fact that such successi- tiary" time along th e boundary between the European and
ons are approximately coeval and may be refe rred to anal o- African plates. The Jurassic paleog eographic settings of Sicily
gous paleogeographic settings. These similarities enable to and the Alps result from the bre akup of the Triassic carbon ate
comp are radiolarian assemblages of different geographic areas platform and the subseque nt extensional mo vements. The
and to correlate different Teth yan paleog eographic region s. plat form was divided in several areas with different subside nce
Th e paleogeographic dom ains are two pelagic plateaux (Tra - rates, and different paleog eographic domains were formed
panese Domain in Western Sicily and Tr ento Plateau in the (pe lagic plateaux, basins and limited areas of platforms) (Figs.
Southern Alps) and one basin (Sicanian Domain in Western 1 and 2). During the Middl e Jurassic a general deepening took
Sicily) (Figs. 1 and 2). The age of the studied successions ran- place and the deposition of sediments in the Rosso Arnmoniti-
ges fro m the Bathonian to the Kimmeridgian. co facies occurred on the pelagic plateaux up to Late Jurassic
I Dipartimento di Scienze della Te rra. Via Valperga Caluso 35, 10125 Tori no. Italy. Ema il: [email protected]
Jura ssic radiolarians, Southern Alps , Sicily S21

.
Sicanian
SA
Ioni an
fill Fig. 1. An attempt of paleogeograph ic recon-
struction of the Sicilian area at the Late Tri assic
Ej time (Catalano et al. 1996). Favignana , Balala di
o Reeh Baida and 1nici MI. sections belon g to the pelagic
plateau of the Trap ane se Domain ; Sant'Anna sec-
• Slopes
tion belon gs to the Sicanian Basin. The paleogeo-
D Basinal areas graphi c location of the stratigraphic sections is
only indicative.
(Winterer & Bosellini 1981; Catalano et al, 1996). The Rosso under- and overlying sediments and, in some instances, also
Ammonitico sediments consist of condensed red nodular lime- within the siliceous successions: This fact motivates the analy-
stone rich in ammonites, and are subdivided in three members sis of radiolarian assemblages in the selected sections. The
whose ages differ between the Southern Alps and Sicily. The abundance of radiolarians enables to study them for a twofold
lower part (RAI: Rosso Ammonitico Inferiore) spans the late purpose: to date directly most of the siliceous facies (whose
Bajocian-early Callovian in the Alps and the Bathonian-mid age was generally based on the ages of the bracketing forma-
Oxfordian in Sicily. The upper part (RAS: Rosso Ammonitico tions) and to improve the calibration of radiolarian zones by
Superiore) starts in the mid Oxfordian in the Alps and in the ammonite zones.
early Kimmeridgian in Sicily. RAI and RAS are separated by This paper deals with the main results of the author's PhD
the intermediate siliceous member (RAM: Rosso Ammonitico research (Beccaro 2002) where radiolarian assemblages were
Medio), assigned to upper Callovian-mid Oxfordian in the studied for the first time in the following sections: Fornazzo
Alps and mid Oxfordian-Kimmeridgian in Sicily. The interme- Strada, Fornazzo Cava, Castello Inici, Balata di Baida, Favig-
diate siliceous member represents the studied sediments in all nana (North-western Sicily) (Beccaro 2002, 2004a) and Cava
the stratigraphic sections except for Sant 'Anna (Sicily) . Vianini (Southern Alps) (Beccaro 2002). Concerning the San-
Very few authors have studied the Jurassic radiolarian as- t'Anna (South-western Sicily) and the Ceniga (Southern Alps)
semblages of the Southern Alps and Western Sicily. Kocher sections, new radiolarian data improved the former biochrono-
(1981) was the first author who studied radiolarians from the logic assignments. New species of Nassellaria (Fultacapsa
Southern Alps but mainly in the Lombardian basin. The Ceni- ozvoldo vae, Podobursa andreai, Podobursa vannae, Loopus
ga section was first described by Fogelgesang (1975) , and then doliolum martae) and Spumellaria (Emiluvia peteri, Triactoma
studied for radiolarians by Baumgartner (1984) and Baumgart- enzoi) from the cited sections have been described in Beccaro
ner et al. (1995b). The Coston delle Vette section was first de- 2004b.
scribed by Dal Piaz (1907) , then analysed by Bosellini & Dal
Cin (1968) and Della Bruna & Martire (1985), and finally stud-
Lithologic description of the stratigraphic sections
ied for radiolarians by Beccaro (1998) and Beccaro et al.
(2002). The Cava Vianini was only described by Papa (1994) In Western Sicily five stratigraphic sections of the Rosso Am-
for sedimentologic purposes. Concerning Western Sicily, sev- monitico Medio (RAM) have been studied in the Trapanese
eral authors studied the geology for different aims (Giunta & Domain (pelagic plateau) and one section in the Sicanian
Liguori 1972, 1973; Wendt 1964, 1971; Catalano et a1. 1989; Basin (Fig . 1). The geographic location of these sections is
Cecca et al. 2001; Catalano et al. 2002 among others). Never- shown in Figure 3.
theless, the radiolarian papers regarding the Sicanian Basin TRAPANESE DOMAIN (North-western Sicily) - Fornazzo
have been mainly focused on the Sant'Anna section (Riedel & sections are located at the Inici Mt. near Castellammare del
Sanfilippo 1974; Mascle 1973, 1979; Baumgartner et al. 1980; Golfo. The Fornazzo Strada section crops out along the road
Origlia-Devos 1983; Baumgartner 1984; De Wever et al. 1986; to Fornazzo quarry. The RAM is 26 m thick and consists of
Aita 1987; De Wever 1995) because of its rich fossil content well-stratified red siliceous limestone alternating with calcare-
(ammonites, belemnites, brachiopods, echinoderms, nannofos- ous marlstone. Nodules and beds of dark red chert are abun-
sils). Other Sicanian sections were studied by Kito et al. (1990) dant (Fig . 4). The Fornazzo Cava section crops out in the old
and Kito & De Wever (1992,1994). Fornazzo quarry, and the Castello Inici section is located in
Generally, the close association of ammonite-bearing beds the southwestern side of the Inici Mt. In both sections the
with levels containing well-preserved radiolarians is quite rare. RAM is incomplete and it is 8 m and I3 m thick, respectively.
In the investigated sections, the ammonites occur both in the The lithology is the same as at Fornazzo Strada and the main
S22 P. Beccaro
r:~~
'-, AUSTROALP INE DOI'vlAIl\'
TRENTO
lnsubric Fault l. ine

PLATEAU
.~
.~
/~
cv • 0 rRl ULI
( Maggiore.
.Cumo Lake
J ;! ..:::- .L
....- CE
Bc llu no
.:s> /. I'LATFOR:Yl
'-oj
O~al dO ~1t.
I" Lake ,";;/ Fig. 2. Geographic location of the stratigraphic
~ sections and present day distribution of Mesozoic
"
LOl'vlBARDIAN BASIN
.
I..:
• Gar da I .akc.,~ \~ .'~ .

paleogeographic domains in the Southern Alps
(Italy) (Bosellini et al. 1981; Martire 1992). CE:
Milan 50 kin
• Veron a ..' Veniee Ceniga, CV: Coston delle Vette, VN: Cava
Vianini.
Fig. 3. Geographic location of the stratigraphic

sections in Western Sicily (Italy). BB: Balata di
50 km Baida, FV: Favignana, 1M refers to the three sec-
N SICILY ---- - - _ tions of the Inici Mt. (Fornazzo Strada, Fornazzo
--- Cava and Castello Inici), SA: Sant'Anna,
difference is the occurrence of macrofossils (ammonites and dark siliceous limestone and beds chert (light brown, red and
belemnites; Fig. 4). The Balata di Baida section crops out black in colour). Some levels of bentonites occur at the top
along the orographic left of the Sarcona River at Balata di of the section (Fig. 4). The radiolarian preservation varies
Baida village (near Inici Mt.). The RAM is 21 m thick, and it from moderate to very good. The Ceniga section crops out
consists of an alternation of nodular red limestone and vari- near the Sarca River, south of Ceniga village (North of the
ously coloured chert beds; thin levels of marls tone are wide- Garda Lake). In this section it is also possible to observe the
spread along the section (Fig. 4). The radiolarian preservation Early Jurassic platform deposits of San Vigilio Oolite
in the Inici sections is moderate. The Favignana succession is (oolitic-bioclastic grainstone). The top of San Vigilio Oolite
located in the southern coast of the Favignana island (Egadi is overlain by a very thin RAI (20 em) formed by pink pelag-
Archipelago). The section is 2 m thick, and it consists of thin ic limestone and bearing a hardground at its top. The RAM
limestone and marlstone alternating with thin chert beds black is 9 m thick, and it consists of an alternation of red siliceous
and red in colour (Fig. 4). Neither the base nor the top of the limestone thinly stratified and thin whitish marlstone. Six
siliceous succession crop out. The radiolarian preservation is levels of bentonites occur in the middle part of the section
very good. (Fig. 4). The radiolarian preservation is poor. The Coston
SICANIAN DOMAIN (South-western Sicily) - The basinal delle Vette section is located in the Feltrine Alps (Dolomiti
succession of Sant'Anna crops out at about 1 km NE of San- Bellunesi). Here, the time-equivalent facies of the RAM is
t'Anna village (near Sciacca). The stratigraphic section is 9 m the Fonzaso Formation, which reaches a thickness of 100 m.
thick, and consists of a regular alternation of whitish limestone The Fonzaso Fm. consists of a quite regular alternation of
and marls tone; the chert is rare. The base is not exposed packstone and wackestone with subordinate oolitic grain-
(Fig. 4). The radiolarian preservation is good. stone and mudstone, and chert ribbons and nodules (Fig. 4)
In the Southern Alps three stratigraphic sections have (Della Bruna & Martire 1985; Beccaro 1998; Beccaro et al.
been studied in the Trento Plateau (Fig. 2). The Cava Viani- 2002). The central part of the formation is characterized by
ni section crops out in an active quarry close to Madonna the oolitic grainstone resedimented from the Friuli Platform.
della Corona Sanctuary (eastern side of the Garda Lake). The radiolarian preservation varies from moderate to very
The RAM is 10 m thick, and it consists of an alternation of good.
Jurassic radiolarians, Southern Alps, Sicily S23

[JJ
N
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-e
tl:l
, :nJ"'Jil'tt~
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Ammo nite Zones

Cava Vianini Radiolarian UAZ-5A Hybonotum Zone: ea rly Tith.
Legend
UAZ F: early Kimm. pars to late Kimm. Becken Z one: lat e Kimm .
~ Siliceous limestone ~ RAt and RAS Cavour;Zone: late Kimm.
UAZ E: late? Oxf. to early Kimm. pars Divisum Zone: earty Kimm.
~ Strabfied limestone ~ Chert beds and nodules
UAZ 0 : mid?~ate? Oxf. Strombecki Zone : ea ~y Kirnrn.
m Marly limestone B Bentonites Transvorsan'um Zona: topmostmid Oxt.
M a rtstone (!, Ammonite s
UAZ C: mid Ox!. Plicatilis Zo ne: mid Oxf .
UAZ B: earty Call. pars to ea~y Ox!. ~3 : mid?-Iate? Oxt.
:ili:i San Vigilio Oolite < t Belemnites
&2: earty Call.
~ Nod ula r limestone UAZ A: early? mid Bath. to ea~y Call. pars , 0 1: ea~y Bath,
Fig, 4, Radiolarian biostratigraphic correlation of pelagic siliceous facies between Western Sicily and the Southern Alps by means of the six radiolarians biozones UAZ A-F. Three sections from the
Southern Alps (Ceniga, Cava Vianini, Coston delle Vette), five sections from North-western Sicily (Fornazzo Strada, Fornazzo Cava, Castello Inici, Balata di Baida, Favignana), and one section from
South-western Sicily (Sant'Anna) have been dated and correlated, The investigated successions represent the intermediate pelagic siliceous member (RAM) of the Rosso Ammonitico Frn. in all the sec-
tions, except at Coston delle Vette (where the Fonzaso Fm. is the time-equivalent siliceous facies of the RAM) and at Sant'Anna (where a basinal section crops out). The age of the biostratigraphic units
UAZ A-F is provided by calibration with ammonite assemblages found both in the studied successions and in the under- and overlying sediments.
A B c o E F Unitary Associations Zones for Sicily and Southern Alps
1 1 1 1 1 1 1 1 1 1 2 Unitary Association. (UAs)
123 4567 890123 4567890
• . . .. ,. .. . ' . . . . - . . . • .. Eucyrtidiellum unuma~n~e dentalum B"'UlKoARTNIEA
• . , ,. . . ., C . . . 0' c .c Styiocaps. oblon gulB KocH~JI
• . . .. .., ~ , . .g ~ .' ~ Vnume emin ll/us lCHl1Vo'M& y U)
: : : . : :: : : q ..:: ~~ ~~/:~:I~:c'::t~R~~HEA) v. :
- . . . . . . . 3 . , . . 6 §r . , . . ~ . . Mlnfv sus fNiJt;;ili~ pnJet; uad alupe n! l-S BAlNCAA THER & BARTOLINI
. . ii . . . . 3 ~ . ... ~ - Mlrifvs U$ fra gili.s s I B~ TNER
-. ~ ~ '.".'.;Q.l 9
~
..
..... . , . . 3' · ,
A " P.seuaoeucyrtrs fm"" HULL
A cafmiot~psis varliltus Vf1riatu s (OlVO~DO'''''')
• - . • • • • • ·oJ • , •• ~ • • • • ? . Tethys etta dhJmena ~n .s/.s s:I. e..
UMGA.Rr",E.1P
. " . .. : : S'_ . . , "tJ • . .... r. tradltryrn a co ,.,.lito$~m$is 5 I (p[$$A(i. No)
• . . . . . . . . . . ..~ . . A C8M IOtylopSlS variatu .s s I (OZ\l'Ol.OO"t,oA)
• . ...•. • . • . . • " . , •• ~ •. Telh~etta dhimen~M$J! up. A SM SU Baumgartner e t al. 1995c.
• • • • ' • • ' 0 • •• ~ . , Eucyrt idieJlu m l,mUmaen$e $ I (YAO)
• • • • • • • • • • • • • • • , • , • • ~ 0 , Podobuf$a polyac~nth~ (F ISCHU)
• • • • • • • • • • • • , • • • • • • • 41 • , Pa hniindromeda spp .
• • • • • • • • • • 2:; , •• , 0 •• 0 , Em iIuvi. premyogii ~ItTN~R
• • • • • • • • • • • ' •• , •. 3 . . Perispyridium ordll1arium gr. ( PESSA.GNO)
• • • • • • • • • • • • • . . . ;3 . , Irensnsuum bre vicostafum Qr, (~ DCNA)
• • • • • • • • • • • • • • • . . .. Risto !, alrissim. s.l. (Rusr)
• ••••••••••••••• . , • . , Parahswm Carpatlt iCum Wtoz. &. D E WE \lE.R
• • • • • • • • • • • • • • • • . ' , • , Paron~~ na bandyi P U SAGIoI O
A..•·••...•••...•.••
• •• • ••••••••••••• • . ,
•
.:...:..- • •
C •••
•••••••••••••••• • . .
. .. .. . . . . . • . . . . . . . . . . . . ,.
0 • , o. ••• • ••• , ••
Pou/~$ sp. at!. P ocu!~tus Of. W£vu sensu Baumgartner et at. 1 99~
Bemoullius deera (B AUUQUTNER)
Angulobf8cchia puris imae nsis (PI!.SSAGIofO)
Tnacloma ~rab /ake; YAnG & WANG
KiJinora carenarum (M...r$t)()r.A )
~ : : : . . , .. :: : : : : : ::: . : : : : : . : : : ;:~;~::::e:~%::e~uY:(Y~)
~ ::: ,~ : :::::: ~ : : ~ : : ~ : : : : : ~ : : ~~~=::d=~:e~~;:S=naensiS (BAU~TNfR)
~ •••• , • . ,....... . . , . , •• , . . . WlllinfJdellum (?) ma..euec~. CoRTESE.
~ :::::::: :: :: ~ : ~ ~ : : ~ : ~~~~~Iu;";'::~~~:::TA
:?,0.:rr ... • • • ••
•• ••
• • •• ••
•••••
•• •
•
. ...
•• .,
• • .
0 Telretrabs lea lis (ONOlOO'VAl
Eucyrtidie//um ptyc1um (RIEDEL &. S ,Io.Nf"llIPPO)
Acanthocin;;us suboblorJ9Us e.! (YAal
0: :••:••:: :• :: ~ : ~~x":~~~u;,~~ss~~7~(:~NMGA~rN[~)
0
g: . • • • •• 0 • Loopu~ do/ iolum marta e BECCN!O

~ . • • • • • • . Syrir)gocapsa 'poA
(') . • • • • • •• Trrrabs ewmgi s.l. (PfSS...c;NO)
~ : :. :. :. :. :. :: J:~~:S~~;~:;k~s{r:c:~"IO)
Qlo . .. Podoburu Ina CiJnthll (FI5C H~ t) gr.
~ • 0 • • 0 • 0 • Tntrab s naY'( P(U,I,(j,NO)
• • o.
• •• • • • • • • , PanfaneJ /rum nedeli P f SSACNO

• •• • • • • • ' • O'- nda Sp B sensu Hull 1997
• • , . 0. ' •• ' • • , 0 Homoeoparoan~"a argoJidens ;s B AuWaAA TNER
. • .• • • • • •. , •. , . , WriSiri~lIum nrpitfhicum DlIM. r ~ 1CA
B::: : : :::: '.0 ~ ~ ~ ~:;;:~~::J~a:~~: ~~~T:~~AR TNER )

~ • • • • • • • , • • , RiJto/. ltltissim~ a/tis~m. (RVST)
§; •• • •••••• ' . Zhamoioellum venfncosum O UlIIlTAICA
N .. • • • • • • • • , . Tntr,bs exotica (P f SSAGIoI O)
• ' . • • • • • • • X ltus magnu s e ~TN E . 0 ,
• •• • • • • • • • • Emifuvia orea B ~TN E .ll

, .• • • • • • • • • • • Podobursa vlltnae BE-ceAR O
" , ·, • , •• .• . , • • MJnfu~us gulJdlt/upens ls PESS.r.c;,..O
C: •• •
•
•
•
•
•
•
• •
• • • , PodotHJfSlt SfMosa (OzVO~DOVA)
• • Tnac10ma blake! ( P rsaAGl NO)
'--- , ,
, •• , • •• •• • 0 Z!Jamoide flum (') exquiSitum Hvu
C
i!::
•• , • . , •. , • . Zanola comuta
• ••• • 0 • Loopusdo/;o lum
(~TN f R )
OllU'T ~ 1CA
0 •
••
• • • ' • Saitoum ievium DE WE VErt
• • • • HexIJS.tum.lis suboblongus (y '(0)
3 Cingulotums urpata DUUlTRICA
o a: • • 0 ,
• . ' • TmClOffl. fOfll'm.n~lt M UZJl'oOt

.!l. 0 • • . . Em i/wia pe terl B ECCARO
~ • •• Tetratr.bs bulbou BA UMQAIl'T..-EJI
• •• Minf usus dian ae s.l. ( K.ut Af A)
• ••
AfChaeodictyomitra apiarium (R OST)
:• •
• Prot unuma j JJ po nicus MATs uOKA & Y"O
• •
• • • •
Eml /uvia hopson; P U 9 01roO..c)
• Trltrabs ewin~i waneS
•• i (PtM"C.NO)
• N.pora deoweWcn 8AlJ htOARTN[ 1II
• • 0
• • . . Napora Iospe nsi s P[ S$AGkO

• • • • Angu/oblBCChia biard ina/rs Or.rot.OOWl
''....•
'
•
•
• ,
•
•
,
• • • ZhamoidltHum o vum DUM m~ 1CA
0 Paron."IJa pygmae. B A~JIf TNER
• , •
• • . . Emifuvie peSs.gnoi ' I. F OIt [ ..... N

• • • . Futtac.p sa spnaeorlu (OlYOl.OO\f") 5.1.
~
o '•• .
••
•
·..•••
. . . , Emiluvia ordmaria Ol'YolDOVA
• • ' . . , . MlrJfusus dian8lt mifJor 8~TN E R
•••• ' Tritrabs rtHxJodaetylu$ B AlIM t)A It'W[ 1t
..
PodocaPS' amphJtreptera F OR EN""
• • Emi/uvia ulJimi B ~TN( rt & O ll...,TRIC..
• .'
Minfu sus dianae dia n8ft (K....,UI UI:)
••
.'.'..
TriactomiJ IJfhonianum R UST
Wrang6' Jlium Okamura ; {MauTANI}
LoopcJs pruYlltivu$ ( MAT8\K)O(,Jl &. y~)
• PSltudOeuCyrlls sp. B sl!Insu IMdz ' 991
:
.'••
Fultacap!Ja !Jphaeriu (OZ'YOl.O(NJo,)
Teo~rtus ca/Marrus (OZ\rOLDO\'A)
E ..• ••
RJsto/~ nodou Hofr l
..••'.
Saltoum defCourli WI 02 & D E W EVER Fig. 5. Range chart: occurrences of radiolarian taxa used for
A caeonjot~ umbJJicaf. (RUST)
Synrl90Capsa spi ne /Hera BA Litr.tCAItT"'ER the biozonation. The software BioGraph grouped 100 select-
Napora bonelJ" PessoiIgno . WH"llN & YOI
ed species in 20 Unitary Associations which have been manu-
F· PStwOoeucyrffS reticul~ri$ M " T!?tUQll.A & Y NJ
ally assembled in six Unitary Associations Zones: UAZ A-F.

Radiolarian biozones UAZ A-F bly included in the very condensed RAJ (Fig. 4). UAZ C
is assigned to mid Oxfordian by the occurrence of ammonite
The radiolarian biostratigraphy illustrated in this paper has assemblages belonging to Plicatilis and Transversarium
been carried out using the Unitary Associations method (Guex Zones at the Castello Inici section (Savary 2000), and
1977,1991) and the software BioGraph (Savary & Guex 1991, Transversarium Zone at the Fornazzo Cava section (Bovero
1999), and represents a new regional zonation for the Southern 2000).
Alps and Western Sicily (Italy). About 130 radiolarian species
have been identified from 8 stratigraphic sections, and 100 taxa
UAZ 0 (mid?-late? Oxfordian)
have been retained to construct the zonation (Fig. 5). Also
added to this database are the radiolarian occurrences of the UAZ D is the most widespread and best recorded biozone in
Coston delle Vette section (Trento Plateau, Southern Alps) all the sections except Sant'Anna (Sicanian Domain, South-
(Beccaro et al. 2002), and the raw data of two Baumgartner's western Sicily; Fig. 4). Species belonging to UAZ D are shown
samples (POB 1703 and POB 1704) of the Ceniga section in Plate 1, Figures 14 and 19-24. UAZ D is questionably as-
(Trento Plateau, Southern Alps) (Baumgartner 1984; Baum- signed to mid-late Oxfordian due to the age assignment of the
gartner et al. 1995b). The adding of the Baumgartner's samples well calibrated UAZ C (mid Oxf.) and the occurrence of am-
was necessary in order to get a radiolarian record also from monites referred to the boundary mid-late Oxfordian (Papa
the upper part of the Ceniga section that lacked in well pre- 1994) at the base of RAS in the Cava Vianini section (South-
served samples from PhD field work . Six Unitary Associations ern Alps; Fig. 4).
Zones (UAZ A-F) have been defined (Fig. 5), and the
chronostratigraphic value of each UAZone is provided by cal- UAZ E (late? Oxfordian - early Kimmeridgian pars)
ibration with the ammonites found in the RAM of the studied
UAZ E is well recorded in all sections except Cava Vianini
sections and in the under- and overlying sediments (RAI and
(Southern Alps), where the time-equivalent facies is the RAS
RAS) .
(Fig. 4). Species belonging to UAZ E are shown in Plate 1,
Figures 25-31. Ammonites belonging to Strombecki Zone
UAZ A (early?-mid Bathonian - early Callovian pars)
(early Kimmeridgian) occur at the very top of UAZ E at the
UAZ A occurs only at the Coston delle Vette section (South- Fornazzo Strada section (North-western Sicily) (Bovero 2000)
ern Alps; Fig. 4). Some species characteristic of UAZ A are il- so that it is likely that UAZ E also comprises the late Oxfor-
lustrated in Plate 1, Figures 1-3. UAZ A is assigned to dian.
early?-mid Bathonian - early Callovian pars thanks to the Bentonite levels occur at the top of UAZ D at the Cava
presence of early Bathonian ammonites at the top of RAI at Vianini and Ceniga sections , and at the very base of UAZ E at
the Coston delle Vette section (Della Bruna & Martire 1985), the Ceniga section. The time intervals expressed by UAZ 0
and the early Callovian pars age assignment of the following (mid?-late? Oxfordian) and UAZ E (late? Oxfordian-early
UAZB. Kimmeridgian pars) suggest that the volcanic activity took
place also in the late Oxfordian-early Kimmeridgian time, and
UAZ B (early Callovian pars - early Oxfordian) not only in the mid Oxfordian as stated in the literature (Mar-
tire 1989; Baumgartner et al. 1995b).
UAZ B is present at the Cava Vianini and Coston delle Vette
sections (Southern Alps; Fig. 4). Species belonging to UAZ B
UAZ F (early Kimmeridgian pars -late Kimmeridgian)
are shown in Plate 1, Figures 4-11. The base of UAZ B is
dated as early Callovian pars due to the presence of early UAZ F has been recognized at the Coston delle Vette
Callovian ammonites at the top of RAJ (Papa 1994) at the section (Southern Alps), and at the Fornazzo Strada and
Cava Vianini section. The upper part of UAZ B is constrained Sant'Anna sections (Western Sicily; Fig. 4). Species occur-
to early Oxfordian by the well-dated UAZ C. ring into UAZ F are illustrated in Plate 1, Figures 32-35 .
UAZ F spans all the Kimmeridgian due to the presence of
Divisum Zone (early Kimmeridgian) at the base of the RAS
UAZ C (mid Oxfordian)
in the Coston delle Vette section (Southern Alps) (Dal Piaz
UAZ C has been recognized in the Fornazzo Strada, For- 1907), and of Cavouri Zone (late Kimmeridgian) at the For-
nazzo Cava and Castello Jnici sections (North-western Sicily; nazzo Strada section (North-western Sicily) (Bovero 2000).
Fig. 4). Species belonging to UAZ C are illustrated in Plate The extremely base of the RAS at the Fornazzo Strada sec-
1, Figures 12-13 and 15-18. In the Alpine sections UAZ C tion is dated as late Kimmeridgian (Beckeri Zone) (Bovero
seems missing but no evidence of stratigraphic gaps has been 2000). In the Sant'Anna section (South-western Sicily) the
found. Most likely, UAZ C was not recognized at the Coston top of the pelagic siliceous succession is assigned to UAZ F,
delle Vette section due to the cover, and in the Cava Vianini and is overlain by ammonite-bearing limestone belonging to
section due to the low frequency sampling. At the Ceniga the Hybonotum Zone (early Tithonian) (De Wever et al.
section the time interval corresponding to UAZ C is proba- 1986).
S26 P. Beccaro
Diachronism of the Jurassic siliceous facies between the Concerning the Ceniga section, the time interval stated in
Southern Alps and Western Sicily the previous papers (Baumgartner 1984; Baumgartner et a~.
1995b) spans the mid Callovian to early Tithonian. The radi-
The stratigraphic correlation through UAZ A-F reveals a sig-
olarian assemblages of the new samples CE 0.80 and CE 5.20
nificant diachronism for the lower as well as for the upper limit
belong to U AZ D (Fig. 4) and assign most of the section to
of the pelagic siliceous facies in the Alpine and Sicilian sec-
mid?-late? Oxfordian. The radiolarian content of the new
tions.
samples CE 5.77, CE 5.90 and CE 6.90 belong to UAZ E
In the Southern Alps (Trento Plateau) the siliceous deposi-
(Fig. 4), and refer the upper part of the section to late? Ox-
tion began in the early?-mid Bathonian-early Callovian pars
fordian-early Kimmeridgian pars. The last two samples (POB
(UAZ A) at the Coston delle Vette section, in the ~arl'y
1703 and POB 1704) are from Baumgartner's sampling and
Callovian pars - early Oxfordian (UAZ B) at the Cava Viani-
they were assigned to late Oxfordian-early Tithonian (Baum-
ni and in the mid?-late? Oxfordian (UAZ D) at the Ceniga
se~tion (Fig. 4). In North-western Sicily (Trapanese Domain) gartner 1984). The radiolarian assemblages of these samples
belong now to UAZ E (late? Oxf.-early Kimm. pars). I~ ~c
the siliceous deposition started in the mid-late Oxfordian:
count of this research, the age of the Rosso Ammonitico
UAZ C (mid Oxf.) at the Fornazzo Strada, Fornazzo Cava and
Medio at the Ceniga section is restricted to mid? Oxfordian-
Castello Inici sections; UAZ D (mid?-late? Oxf.) at the Balata
early Kimmeridgian pars (UAZ D-E) (Fig. 4). This ~ge a.s-
di Baida and Favignana sections. In the basinal section of San-
signment highly differs from those stated for other sections m
t' Anna the onset of the siliceous facies is the youngest: U AZ E
the Trento Plateau, where the RAM is referred to late
(late? Oxf.-early Kimm. pars) (Fig. 4). On the other hand,
Callovian - mid Oxfordian (e.g., Cava Vianini, this paper;
UAZ D (mid?-late? Oxf.) corresponds to the final phase of
Martire 1996).
the siliceous deposition at the Cava Vianini section (Southern
Concerning the Sant' Anna section, several authors as-
Alps), and UAZ E (late? Oxf.-early Kimm. pars) corresponds
signed it to a variety of ages: mid Oxfordian to mi~ Tith~n
to the final phase of the siliceous deposition in the Ceniga sec-
ian (Origlia-Devos 1983), mid Oxfordian to early Kimrnerid-
tion (Southern Alps). UAZ F (early Kimm. pars-late Kimm.)
gian (De Wever et al. 1986), mid-late Oxfordian to l~te
indicates the end of the siliceous facies at the Coston delle
Oxfordian-early Kimmeridgian (De Wever 1995). The radio-
Vette (Southern Alps), Fornazzo Strada and Sant'Anna
larian content of the new sample SA 0.35 belongs to UAZ E
(Western Sicily). The diachronism of the ending of the
(Fig. 4) and constrains the first layers of the su~cess~on to
siliceous facies in the Fornazzo Cava, Castello Inici and Favig-
late? Oxfordian-early Kimmeridgian pars. The radiolarian as-
nana sections is only due to the incompleteness of the succes-
semblages of the new samples SA 5.10 and SA 9.10 belong to
sions (Fig. 4).
UAZ F (Fig. 4) and date the middle and upper parts of the
As can be inferred by the above discussion, the diachro-
section to early Kimmeridgian pars-late Kimmeridgian. This
nism of the siliceous facies occurs within the same paleogeo-
age assignment is consistent with the early Tithonian age. of
graphic domain as well (Fig. 4). The three Alpine sections
the ammonite assemblages found at the base of the overlying
were located in different sectors of the Trento Plateau, and the
nodular limestone (De Wever et al. 1986). The present study
diachronism suggests that local topography and/or tectonic
now restricts the age of the pelagic siliceous succession of the
movements primarily controlled the onset of the siliceous de-
Sant'Anna section to the late? Oxfordian-late Kimmeridgian
position. Different bottom morphology and t~ct~nic~ may
(UAZ E-F).
have influenced different spatial and temporal distribution of
the siliceous facies: the oldest onset and the youngest end took
place at Coston delle Vette (located in a transitional area be- Age assignment discussion for some taxa
tween Trento Plateau and Belluno Basin) while the youngest
After the calibration of UAZ A-F through ammonite zones,
onset was at Ceniga (located near the strongly tectonic active
the stratigraphic distribution of some taxa can be compared
Garda Escarpment). On the other hand, the Sicilian sections
with those of Baumgartner et al. (1995a). In the following dis-
belonging to the Trapanese Domain were paleogeographically
cussion the lettered UAZones (e.g., UAZ A) refer to the
very close to each other, and the diachronism of the onset
new biozones illustrated in the present paper; the numbered
amongst them is minor (only Favignana section shows a rather
UAZones (e.g., UAZ 8-13) refer to the biozones of Baum-
younger onset but its base does not crop out).
gartner et al. (I 995a). Some age considerations were illustrat-
ed also in Beccaro (2004a) concerning Eucyrtidiellum
unumaense s.l. (Y AO) and Williriedellum (?) marcucciae
Age assignment discussion on the siliceous facies at Ceniga
CORTESE. The ranges of these species remained unchanged
(Southern Alps) and Sant'Anna (Sicily)
after the new zonation by UAZ A-F: Eucyrtidiellum
In the light of new radiolarian biozones UAZ A-F, the age as- unumaense s.l. (Y AO) and Williriedellum (?) marcucciae
signments of the Ceniga (Trento Plateau, Southern Alps) and CORTESE extended to mid Oxfordian (and not only to early
the Sant'Anna (Sicanian Domain, South-western Sicily) sec- Oxfordian as the previous assignment of Baumgartner et al.
tions have been discussed. 1995a).
Jurassic radiolarians, Southern Alps. Sicily S27

Podobursa polyacantha (FISCHL!) (PI. 1, Fig. 14) Nine stratigraphic sections of the Rosso Ammonitico
Medio or time-equivalent siliceous facies have been analysed
Its range is assigned to UAZ 5-8 (latest Baj.-early Bath. to mid
in the Southern Alps and Western Sicily. Six new radiolarians
Call.-early Oxf.) but in the studied sections P. polyacantha
biozones (UAZ A-F) have been defined (Figs. 4 and 5): UAZ
reaches the very base of UAZ D (mid?-late? Oxf.) (Fig. 5).
A (early?-mid Bath.-early Call. pars), UAZ B (early Call.
pars-early Oxf.), UAZ C (mid Oxf.), UAZ D (mid?-late?
Ristola altissima altissima (ROST) and Ristola altissima nodosa Oxf.), UAZ E (late? Oxf.-early Kimm. pars), UAZ F (early
HORI (PI. 1, Figs. 13 and 30, respectively) Kimm. pars-late Kimm.). The chronostratigraphic value of
each UA Zone is provided by calibration with the ammonites
The age of R. altissima altissima is stated as UAZ 7-12 (late
found in the same stratigraphic sections. The new UAZones
Bath.-early Call. to early-early late Tith.). In the studied sec-
show a good reproducibility throughout the investigated suc-
tions R. altissima altissima first appears in UAZ C (mid Oxf.),
cessions, and make possible the first correlation by radiolari-
and its ancestral form R. altissima major BAUMGARTNER & DE
ans of the intermediate siliceous member of the Rosso Am-
WEVER occurs in samples assigned to U AZ B (early Call. pars-
monitico Fm. between the Southern Alps and Western Sicily
early Oxf.) at the Coston delle Vette section (Southern Alps)
(Fig. 4).
(Beccaro et al. 2002). Furthermore, R. altissima altissima dis-
The newly defined radiolarian biozones also confirm the
appears at the top of UAZ E (late? Oxf.-early Kimm. pars)
diachronism of the Middle Jurassic siliceous deposition in the
when R. altissima nodosa first appears. R. altissima nodosa was
studied area. On the Trento Plateau (Southern Alps) the
comprised in the synonymy of R. altissima altissima in Baum-
siliceous deposition began in the early?-mid Bathonian - early
gartner et al. 1995c and this fact explains the long range (UAZ
Callovian pars (UAZ A) at the Coston delle Vette section, in
7-12: late Bath.-early Call. to early-early late Tith.) of the lat-
the early Callovian pars - early Oxfordian (UAZ B) at the
ter species. R. altissima nodosa is very probable the transition-
Cava Vianini, and in the mid?-late? Oxfordian (UAZ D) at
al form between R. altissima altissima and R. cretacea (BAUM-
the Ceniga section (Fig. 4). In the Trapanese Domain (North-
GARTNER), whose range is UAZ 12-17 (early-early late Tith. to
western Sicily) the siliceous deposition started in the mid-late
late Val.).1t is therefore reasonable that the range of R. altissi-
Oxfordian: UAZ C (mid Oxf.) at the Fornazzo Strada, For-
ma altissima (UAZ 7-12: late Bath.-early Call. to early-early
nazzo Cava and Castello Inici sections, and UAZ D (mid?-
late Tith.) is restricted to UAZ C-E (mid Oxf. - early Kimm.
late? Oxf.) at the Balata di Baida and Favignana sections. In
pars).
the Sicanian Basin (South-western Sicily) the onset of the
siliceous facies at the Sant'Anna section was in UAZ E (late?
Syringocapsa spinellifera BAUMGARTNER (PI. 1, Fig. 34)
Oxf.-early Kimm. pars) (Fig. 4).
Its range is referred to UAZ 9-12 (mid-late Oxf. to early-early In the light of the new radiolarian biozones UAZ A-F, the
late Tith.) but in the studied sections S. spinellifera does not age assignments of the Ceniga and Sant'Anna sections have
appear before UAZ F (early Kimm. pars-late Kimm.) (Fig. 5). been compared with the literature data. The age of the Ceniga
The age assignment of this taxa should be reconsidered in the section has been restricted to mid? Oxfordian - early Kim-
light of the present data. meridgianpars (UAZ D-E), and the age of the Sant'Anna sec-
tion is now referred to the late? Oxfordian - late Kimmerid-
gian (UAZ E-F) (Fig. 4).
Tetratrabs bulbosa BAUMGARTNER and Tetratrabs zealis
Thanks to the calibration with ammonites, some range con-
(OZVOLDOVA) (PI. 1, Figs. 21 and 22, respectively).
siderations have been stated for certain taxa. The age of
T. zealis (UAZ 4-13: late Baj. to latest Tith.-earliest Berr.) dis- Podobursa polyacantha (FISCHL!) is extended to mid-late Ox-
appears towards the top of UAZ E (late? Oxf.-early Kimm. fordian. At least for the studied area, it could be possible that:
pars) whereas is stated to reach the earliest Berriasian (Baum- Syringocapsa spinellifera BAUMGARTNER does not appear be-
gartner et al. 1995c). T. bulbosa is assigned to UAZ 7-11 (late fore early Kimmeridgian, Tetratrabs bulbosa BAUMGARTNER
Bath.-early Call. to late Kimm.-early Tith.) but it first appears does not appear before the mid Oxfordian, the range of Risto-
in UAZ D (mid?-late? Oxf.). la altissima altissima (ROST) is restricted to mid Oxfordian -
early Kimmeridgian pars.
The radiolarian biostratigraphy by means of new UAZ
Conclusions
A-F is a contribution for a better-defined radiolarian zonation
The main conclusions of this research concern the definition of of the Jurassic Mediterranean Tethys. All radiolarian data pro-
six new radiolarian biozones for the Southern Alps and West- vided by this research will be used both to improve the zona-
ern Sicily (Italy), the dating of the intermediate pelagic tion of the INTERRAD Jurassic-Cretaceous Working Group
siliceous member (Rosso Ammonitico Medio) of the Rosso (Baumgartner et al. 1995d), and to create a database for the
Ammonitico Fm. in the studied area, and the correlation of the definition of new radiolarian biozones for the Jurassic Medi-
investigated successions by means of the new radiolarian bio- terranean Tethys.
zones.
S28 P. Beccaro
Acknowledgements BOSELLINI, A., MASETTI D. & SARTI M. 1981: A Jurassic "Tongue of the
ocean" infilled with oolitic sands: the Belluno Trough, Venetian Alps,
italy. Marine Geology, 44, 59-95.
I would like to thank the supervisors of my PhD thesis: Prof. Peter O. Baum-
BOVERO, A. 2000: Analisi paleontologica e stratigrafica del Rosso Ammoniti-
gartner (Universite de Lausanne. Switzerland), Dr. Spela Gorican (Paleonto-
co (Giurassico medio-superiore) nella sezione Fornazzo di Monte Inici,
loski Institut ZRC-SAZU, Ljubljana, Slovenia), Prof. Marta Marcucci (Uni-
Sicilia Occidentale. Graduation Thesis, University of Turin, Italy, (un-
versity of Florence, Italy) and Dr. Luca Martire (University of Turin, Italy). I
publ.), 1-173.
am also grateful to the reviewer Dr. Luis O'Dogherty (Universidad de Cadiz,
CATALANO, R, DI STEFANO, P. & KOZUR, H. 1989: Lower Permian Albaillel-
Spain) for having improved this paper by helpful comments.
lace a (Radiolaria) from Sicily and their stratigraphic and paleogeographic
The scanning electron micrographs have been taken at the University of Flo-
significance. Rend. Accad. Sc. Fis. Mat. Napoli, IV, LVI, CXXVIII, 80--113.
rence and University of Turin (Italy), at the University of Bristol (Great
CATALANO, R, DI STEFANO, P., SULLI, A. & VITALE, F.P. 1996: Paleogeogra-
Britain) and at the Paleontoloski Institut ZRC-SAZU (Ljubljana, Slovenia).
phy and structure of the Central Mediterranean: Sicily and its offshore
area. Tectonophysics 260, 291-323.
CATALANO, R, Lo CICERO, G. & SULLI, A. 2002: Geology of Sicily: an intro-
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Ocean, DSDP Leg 26. In: DAVIES, T.A., LUYENDYK, B.P. et a!. (Eds.): Ini- Jurassic Passive Continental Margin, Southern Alps, Italy. Amer. Ass.
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taires. Bull. Geo!. Lausanne, 313, 317-340.
S30 P. Beccaro
Plate 1
Scanning electron micrographs of the most important radiolarians used for the definition of the UAZ A-F are illustrated. For each photo are given the code of
the section, the number of the sample, the UAZone of the sample, and the magnification. The codes of the stratigraphic sections are: BB-Balata di Baida, CI-
Castello Inici, CV-Coston delle Vette, FZC-Fornazzo Cava, FV-Favignana, IN-Fornazzo Strada, SA-Sant'Anna, VN-Cava Vianini.
1 - Eucyrtidiellum unumaense dentatum BAUMGARTNER, CV 60, UAZ A, x250

2 - Stylocapsa oblongula KOCHER, CV 60, UAZ A, x200
3 - Unuma echinatus ICHIKAWA & YAO,CV 60, UAZ A, x150
4 - Stylocapsa catenarum MATSUOKA, VN 2.85, UAZ B, x350
5 - Praewilliriedellum convexum (YAO),VN 2.85, UAZ B, x200
6 - Stichocapsa robusta MATSUOKA, VN 2.85, UAZ B, x200
7 - Triactoma parablakei YANG & WANG, VN 0.40, UAZ B, x100
8 - Tricolocapsa plicarum s.l. YAO, VN 2.85, UAZ B, x250
9 - Tethysetta dhimenaensis dhimenaensis (BAUMGARTNER), VN 2.85, UAZ B, x250
10 - Eucyrtidiellum ptyctum (RIEDEL & SANFILIPPO), VN 0.40, UAZ B, x250
11 - Williriedellum carpathicum DUMITRICA, VN 2.85, UAZ B, x200
12 - Podobursa vannae BECCARO, CI 4, UAZ C, x100
13 - Ristola altissima altissima (ROST), IN 3, UAZ C, x100
14 - Podobursa polyacantha (FISCHLI), CE 5.20, UAZ D, x100
15 - Xitus magnus BAUMGARTNER, CI 5, UAZ C, x120
16 - Eucyrtidiellum unumaense s.l. (YAO), FZC 3, UAZ C, x250
17 - Tritrabs hayi (PESSAGNO), CI 5, UAZ C, x75
18 - Williriedellum (?) marcucciae CORTESE, IN 1, UAZ C, x300
19 - Zhamoidellum (?) exquisitum HULL, IN 3, UAZ D, x200
20 - Angulobracchia biordinalis OZVOLDOVA, BB 11.70, UAZ D, x200
21 - Tetratrabs bulbosa BAUMGARTNER, CI 8, UAZ D, x60
22 - Tetratrabs zealis (OZVOLDOVA), IN 3, UAZ D, x60
23 - Napora lospensis PESSAGNO, IN 8, UAZ D, xl00
24 - Emiluvia orea BAUMGARTNER, CI 8, UAZ D, x80
25 - Mirifusus dianae minor BAUMGARTNER, CI 15, UAZ E, x100
26 - Mirifusus dianae dianae (KARRER), SA 0.35, UAZ E, xl00
27 - Pseudoeucyrtis sp. B sensu WIDZ 1991, SA 0.35, UAZ E, xl00
28 - Podocapsa amphitreptera FOREMAN, CV 122, UAZ E, x130
29 - Eucyrtidiellum nodosum WAKITA, IN 19, UAZ E, x300
30 - Ristola altissima nodosa HORI,BB 15.10, UAZ E, x150
31- Emiluvia ultima BAUMGARTNER & DUMITRICA, SA 0.35, UAZ E, xl00
32 - Pseudoeucyrtis reticularis MATSUOKA & YAO, IN 32, UAZ F, x120
33 - Acaeniotyle umbilicata (ROST), IN 32, UAZ F, x100
34 - Syringocapsa spinellifera BAUMGARTNER, CV 125.6, UAZ F, x100
35 - Napora boneti PESSAGNO, WHALEN & YEH, IN 30, UAZ F, x150
S32 P. Beccaro
00l2-9402/06/OlS035-13 Eclogae geol. Helv. 99 (2006) Supplement 1, S35-S47
DOl 10.1007/sOO0l5-006-0603-4
Upper Jurassic Radiolaria from the Vocontian basin of SE France

TANIEL DANELIAN 1 , SALIM LAHSINl l & MARC DE RAFELIS 2
Key words: Radiolaria. Vocontian basin. France, Upper Jurassic

Mots cles: Radiolaires, Bassin Vocontien, France, Jurassique superieur
ABSTRACT RESUME
Two sections of the Vocontian basin (southeast France) were explored for Deux coupes du bassin Vocontien (Sud - Est de la France) ont ete examinees
their radiolarian content. Preservation in calcite impeded their extraction from pour leur contenu en radiolaires. Leur conservation en calcite dans la coupe de
the Chateauneuf d'Oze section, which is well dated by ammonites. Fortunate- Chateauneuf d'Oze (bien datee par des Ammonites) n'a pas permis leur extra-
ly, oligospecific assemblages of pyritized radiolaria were yielded by twelve ction. Par contre, des assemblages oligospecifiques de radiolaires pyritises ont
limestone beds of the Meouge section. In general, preservation worsens to- pu etre extraits de douze niveaux calcaires de la coupe de Meouge. En general,
wards the top of the section. The extracted fauna is dominated by Nassellaria la conservation des Radiolaires est moins bonne vers Ie haut de la coupe. La
and more particularly by Archaeodictyomitridae, Pseudodictyomitridae and faune extraite est dominee par des Nassellaires et plus particulieremcnt des
Williriedellidae. It is the first time that some species are reported outside their Archaeodictyomitridae, Pseudodictyomitridae et Williriedellidae. Quelques
type area or from Western Tethys. Amongst the published zonations the especes sont signalees pour la premiere fois en dehors de leur region-type ou
scheme of Baumgartner et al. (1995a) is found to he the most useful and allows dans la Tethys occidentale. Parmi les biozonations publiees, celie de Baum-
correlation of the studied part of the section with the mid-late Oxfordian to gartner et al. (1995a) s'avere la plus utile et permet de dater la coupe etudiee
late Kimmeridgian-early Tithonian time interval. avec la fourchette d'age Oxfordien moyen-superieur a Kirnmeridgien supe-
rieur-Tithonien inferieur.
Introduction
In spite of the enormous progress made over the last thirty nary stratigraphic studies because its pelagic sedimentary se-
years on our knowledge of Mesozoic Radiolaria (see De quences are particularly thick and relatively complete (e.g. Jan
Wever et al. 2001 and references therein) we are still far from du Chene et al. 1993,2000; Groupe Francais d'etude du Juras-
unravelling the full biochronological potential of this plankton sique 1997).
group. This is due to the large number of species that are still Radiolaria were known to be present in the Vocontian
poorly known from a taxonomic and stratigraphic point of Basin (Beaudoin 1977), but observations were based on thin
view and also to the small number of radiolarian-bearing section. Recent reports by Lambert (1999) and Gardin et al.
samples, for which the age is calibrated independently of (2000) on Lower Cretaceous (Valanginian-Hauterivian) pyri-
Radiolaria themselves. The latter point is also important when tized Radiolaria constitute the only modern study for the Vo-
describing the dynamic of radiolarian macroevolutionary contian basin. We present here radiolarian assemblages ex-
changes through time (Danelian & Johnson 2001). tracted from Upper Jurassic strata of the Vocontian Basin and
The Vocontian Basin, situated in the south-eastern part of discuss their biostratigraphic implications.
France (Fig. 1), represents a privileged area for multidiscipli-
1 Micropaleontologie, Universite Pierre-et-Marie-Curie. - C.N.R.S-UMR 5143 Tour 46-56, 5° etage, Case 104,4 Place Jussieu, 75252 Paris Cedex 05.
Email: [email protected]
2 Biomineralisations et Paleoenvironnernents, Universite Pierre-et-Marie-Curie - FR32. CEPAGE Tour 56--55, 5° etagc, Case 116,4 Place Jussieu,
75252 Paris Cede x 05
Upper Jurassic Radiolaria from SE France S35

(1995) established a composite type mega-sequence for its
Upper Jurassic sedimentary sequences and highlighted the
presence of a number of different lithostratigraphic patterns.
These are easily identifiable and correiatable throughout the
basin with the bio-chronostratigraphic framework established
by Atrops (1982) on the basis of ammonites found in various
sections (Fig. 2A). Following an interval dominated by Oxfor-
dian marls, Kimmeridgian sedimentary facies are essentially
N o
- -- 50km
made of alternating limestones and marls, that become clearly
t calcareous in the Tithonian (Atrops 1982; Pederneiras 1995;

De Rafelis 20(0).
Studied material
Two sections of the Vocontian basin are investigated for Radi-

olaria (Fig. 1). Chateauneuf d'Oze represents one of the refer-
ence sections for the Oxfordian - Kimmeridgian boundary in-
terval of the subalpine regions because of its richness in am-
monites and quality of outcrops (Atrops 1982; De Rafelis et al.
2001). It is situated a few kilometres to the South-East of
Veynes on the left bank of Drouzet, along the Departmental
road D20 between Veynes and Chiiteauneuf d'Oze (44°31 'N-
5°53'E). Situated on the northern side of the Serres Jurassic
high, it represents an area of average sedimentation rates
(Moussine-Pouchkine et al. 1998). The section is about 100
• Marseille metres thick and consists of a regular alternation of marls and
limestones interrupted by a 3 metre thick slump. Limestones
are fairly homogeneous and consist of slightly bioturbated and
organic matter-rich pelagic facies, often rich in ammonites.
Fig. 1. Outcrops of the Vocontian basin in SE France and location of the two
studied sections: Chateauneuf d'Oze and Meouge. The Meouge section is situated along the departmental
road 0 942 (44°12'N-5 °33'E), about twenty kilometers to the
south-southwest of Chateauneuf d'Oze. Situated at the south-
ern flank of the Serres Jurassic high, it also represents a depo-
sitional environment of average sedimentation rates. The out-
crop is about one hundred metres thick and starts close to the
Palaeogeographic and stratigraphic framework
point where the road dips downhill beside the river Meouge.
The tectonic evolution of the Vocontian Basin is directly Although still poorly studied, the Meouge section has the ad-
linked to the opening of the Ligurian branch of Tethys and the vantage of being almost continuous and unaffected by the
formation of its European margin (Dercourt et al. 1993). Sedi- gravitational event present at Chateunauf d'Oze (De Rafelis
mentary variations reflect the different stages of the opening 2000). It is entirely composed of alternating marls and lime-
of Tethys in this area. Break up of the hercynian basement be- stones organised in pluridecimetric beds with an easily work-
ginning in the Triassic led to the formation of a number of able stratonomy (Fig. 2B). As at Chateauneuf d'Oze, pelagic
basins (Languedocian, Dauphinois, Provencal and Vocontian), limestones at Meouge are very homogeneous (micritic). It is
well defined by the presence of submarine ridges (Dubois & onl y towards the top of the sequence (bed 181 onwards,
Delfaud 1989). The basins were more clearly separated during Fig. 2B) that they display some different facies (nodular lime-
the Early and Middle Jurassic tectonic subsidence, which was stones, calcarenites with bioclasts or finely laminated calcaren-
followed by a thermal subsidence during the Late Jurassic and ites) . Macro- and microfossils are rare. The presence of Glo-
Early Cretaceous. From the middle Oxfordian a generalized bochaetes, Saccocoma, Tubiph ytes and a few foraminifera are
subsidence affected the Vocontian passive margin and allowed reported (De Rafelis 2000; Langoisseux 2001). The few am-
the development of highly homogeneous sedimentary facies monites discovered do not provide any precise biostratigraphic
both within the basin and on its edges (Dubois & Delfaud information (Atrops, pers. comm.). However, the presence of
1989). At this time, marine sedimentary environments reached lithostratigraphic units displaying distinct stratonomic pat-
their maximal extension. terns, recognizable throughout the Vocontian basin, are pre-
The interest of the Upper Jurassic sequences of the Vocon- sent in the Meouge section and provide some clues as to the
tian Basin lies with the high quality of its outcrops. Pederneiras age of this section (Pederneiras 1995; Moussine-Pouchkine et
S36 T. Danielan, S. Lahsini & M. De Rafelis

Composite column
r- :2~ ~f),--,,·o
-= Meouge
z 5"§
:! ~~ Section
z ":(5
N 165
0
J:
f-
i= ~-----'::::.:'../J
178
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:J
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II:
UJ
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'"o
UJ
CD
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:J
164
Z mass ive
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C> _ . :J limestones
a:: . .:>
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)(
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0
~ ::l
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a.
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~
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o
I
....z
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o
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------
Sal
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'"
s
c ~::::-
z
«
C> 0:
o:: ::! -c
w 3§
:::;: !;
" dJ -
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0::
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0
0:: 1-- D
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~
:J
a. :>
a. ....
=> -c
~ .:J
~ D
~
~
CD Mg-21 Fig. 2. A) Compo site lithostratigraphic column of
27
the Uppe r Juras sic megasequence of the Vocon-
tian basin, between the "Terres noires" formation
E23 Limestone at the bolt om and the massive Tithonian lime-
_ Marl stones at the top (modified after Pederneiras
1995). B) Detailed lithostratigraph ic column of
10
the Meouge section, including bed numb ers and
tentative lithostratigraphic corr elations with the
Upper Jur assic megasequ ence (after De Rafelis
2000). Arrow s indicate limestone beds that yield-
ed radiolaria.
Upper Jurass ic Radiolaria from SE Fra nce S37

Tab le 1. Occur rence of rad iolar ian taxa iden tified in samples o f the Me ou ge section.
Species/Samples Mg Mg Mg Mg Mg Mg Mg Mg Mg Mg Mg Mg
2 3 21 29 37 58 77 98/99 100 102 132 133
Acastea sp.cf. A remusa HULL
•
Ar chaeo dictyomitra apiarium (ROsT)
•
Archaeodictyom itra etrusca CHIARI et al.
• cf.
Archaeodictyomitra pa tricki KOCH ER

• •
Arc haeo dictyomitra sp.aff. A. pa tricki KOCII ER
•
Archaeo dictyomitra shengi YANG
• •
Archaeo dictyomitra spelae CHIARI e t al.
•
Archaeodictyomitra sp.A
•
Cinguloturris fu siforma HORI cf. •
Emiluv ia p entapo rata STEIGER & STEIGER
•
Eucyrtidiellum pty ctum (RIEDEL & SANFILIPPO)
• •
Gongylothorax f avosus DUMITRICA
• • • • • • •
Loopus doliolum DUMITRICA
• •
Loop us venus/us (CHIARI et al. )
• • • cf. • •
Pantane llium oligop orum (VINASSA )
•
Praeconocaryomma scaterba HULL
•
Protunuma j aponicus MATSOUKA & YAD
• • • •
Ristola altissima ssp.cf. R. a. altissima (ROST)
•
Sait oum pa gei PESSAGNO
•
Spongocapsula palmerae (PESSAGNO)
•
Stichocapsa tuscan ica CHIARI et al.
•
Sun a sp.
•
Tethy setta (?) sp.
•
Triactomaforemonae MUZAVOR
•
Tripocy clia sp.cf. Tluciae JUD
•
Tritrabs sp.cf. T exotica (PESSAGNO)
•
Williri edellu m carpathic um DUMITRICA
• •
Williriedellum crys tallinum DUMITRICA
• • •
Zhamoidellum ovu m DUMITRICA
• • • • • • • • • • •
al. 1998; De Rafelis et al. 1997). A bove the "Terres Noires" Balderum horizon (Divisum A mmon ite zone) , and finally (4)
Formation one can thu s recognize from bottom to top (1) an a bundl e of massive limestones (beds 153 to 162) which could
interval of massive limeston es (beds 38 to 61) commonly pre- possibly represent the ones occurring at the base of the Kim-
sent within the Planula Ammon ite zone, (2) a bundl e of lime- mer idgian (Acanthicum Ammon ite zone) . In summary, al-
stone beds known as «the gro up of 14 (G-14»> (beds 113 to though no direct biostrat igraphic age assignment existe d for
128), commonly pres ent within the Loth ari Ammonite sub- the Meouge section prior to our study, it could be tent atively
zone, (3) a limestone proj ection (be ds 136 to 140), which is consider ed as coverin g the upp er Oxfordian to upp er Kim-
tentat ively correlated with the one occurring within the meridgian interval.
S38 T. Danielan, S. Lahsini & M. D e Rafelis

Radiolarian fauna and age evidence for the Meouge section. Some species are found for
the first time outside their type area (i.e. Loopus venustu s) or
Samples were first processed with hydrochloric acid , which was
outside the Palaeopacific tCinguloturris fusiforma) . Much
followed by a brief (30 min.) bath in diluted hydrofluoric acid
work is still left to be done in order to refine the biochronolog-
(HF 5%) . Sample size was in the order of several tens of crn' .
ical potential of Upper Jurassic radiolaria.
Although present in abundance in the limestones of
Chateauneuf d'Oze section , Radiolaria are entirely calcified
and no identifiable forms were extracted. However, twelve Systematic Palaeontology
lime stone and marly limestone sam ples from Meouge section
Species and genera are discussed in alphabetical order. The
yielded pyritized identifiable Radiolaria, which display differ-
suprageneric classification followed in this work is after De
ent degrees of preservation and abundance . The occurrence of
Wever et al. 2001 and is not repeated here. The synonymy lists
all radiolarian taxa identified in samples of Meouge section ar e
contains, only a few representative studies.
given on Table 1. The identified fauna is oligospecific, with a
dominance of Nassellaria over Spumellaria. Samples Mg-2,
Mg-21, Mg-77 and Mg-102 yielded the best preserved assem-
Genus Acastea YANG 1993
blages, displaying a diversity of about a dozen species.
Type species: Acaeniotyle diaphorogona FOREMAN 1973
Amongst Nassellaria, representatives of families Archaeodic-
tyomitridae, Pseudodictyomitridae and Williriedellidae are the
Acastea sp.cf. A. remusa HULL
most abundant. In general, preservation worsens towards the
(PI. I , Fig. 1)
top of the section, only four taxa being identified in samples cf. A castea remu sa n.sp. HULL 1997, p. 35, pI. 11, figs. 2, 6, 17.
Mg-132 and Mg-133.
Remarks. - The illustrated morphotype recalls A. remusa by
Amongst the published radiolarian zonations the scheme
its blunt spinal tips and presence of deep distally tapering
of Baumgartner et al. (1995a) was found to be the most useful.
longitudinal grooves running on the ridges of its massive
The co-occurrence of species Zhamoid ellum ovum and Gongy-
spin es.
lothorax [avo sus in samples Mg-2 and Mg-100 (table 1) allows
their assignment to Unitary Associ ation Zones (UAZ) 9-10,
Genus Archaeodictyomitra PESSAGNO 1976
which are correlated with the middle Oxfordian to early Kim-
Type species: Archaeodictyomitra squinaboli PESSAGNO 1976
meridgian. This is inferred from th e first occurrence (FO) of
Zhamoidellum ovum and the last occurrence (LO) of Gon gy-
Archaedictyomitra etrusca CHIARI , CORTESE& MARCUCCI
lothorax fa vosus. However, following the disco very of an ex-
(PI. I, fig. 4)
ceptionally well -preserved radiolarian fauna in the Mariana Archaeodictyomitra etrusca n.sp. CHIARI, CORTESE & MARCUCCI in Chiari et
trench (Matsuoka 1998), we now know that G. [avosus existed al. 1997, p. 63, pI. I , fig. 9-10 .
until Berriasian time, at least in the Palaeopacific. Conse-
Remarks. - This species can be distinguished from A. suzukii
quently, the age of the Meouge section can be only correlated
AlTA by the presence of more numerous costae (13-14 costae
with the mid-late Oxfordian to late Kimmeridgian-early
visible per half perimeter).
Tithonian , based on the single occurrence of Z. ovum (UAZ 9-
11) throughout the quasi-totality of the radiolarian-bearing
Archaeodictyomitra patricki KOCHER
samples. On the other hand, Emiluvia pentaporata (= E. bisel-
(PI. 1, Fig. 5-7)
lea) is considered in the biozonation of Baumgartner et al. Archaeodictyomitra patricki n.sp. KOCH ER 1981, p. 57, pI. 12, fig. 14-17 .
(1995a) as constrained to UAZ 11. However, it is found to Archaeodictyomitra sp.cf. A .patricki KOCH ER; Chiari et al. 1997, pI. 1 fig. 16.
occur in middle Oxfordian to lower Kimmeridgian (U AZ
Remarks. - This small four-segment species is characterized by
9-10) samples of the Ionian zone (Danelian 1995) and in Ox-
its conical proximal part becoming an inverted trapezoid dis-
fordian strata of Roumania (Durnitrica, pers. commun.).
tall y. Faint traces of segmental divisions are often visible on
the outline. 10-12 longitudinal costae visible per half perimeter
run up to the chephalis.
Conclusion
Radiolaria occur abundantly in the ammonite-rich Chateau-
Archaeodictyomitra sp.aff. A. patricki Kocher
neuf d 'Oze section, but their preservation in calcite did not
(PI. 1, Fig. 8)
allow extraction of any identifiable microfauna. On the con - aff. A rchaeo dictyomitra patricki KOCH ER 1981, p. 57, pI. 12. fig. 14-17 .
trary, an oligospecific and moderately diverse pyritized Radio- cf. Ar chaeodictyomitra sp.aff. Asquinaboli PESSAGNO, CHIARI et al. 1997, pI. I ,
larian fauna was extracted from the Meouge section. Although fig. 17.
the biochronologic resolution of Radiolaria does not allow a Remarks. - This morphotype differs from A. patricki by its
precise age assignment (mid-late Oxfordian to late Kimmerid- more elongated outline and lesser number of costae (about 10
gian-early Tithonian), it is currently the only biostratigraphic visible per half perimeter).
Upper Jur assic Radiolaria from SE France S39

Archaedictyomitra shengi YANG Remarks. - Two distinct morphotypes were observed in our
(PI. 1, Fig. 9-10) material. Morphotype A (fig. 16) displays a trapezoidal ab-
Archaeodictyomitra shengi n.sp, YANG 1993, p.64, pI. 1, domen, bearing 7 prominent costae visible per half-circumfer-
Archaeodictyomitra labronica n.sp. CHIARI, CORTESE & MARCUCCI in Chiari et
ence. Morphotype B (fig. 17) displays a much more rounded
al. 1997, p. 64, pI. 1, fig. 11-12.
Not Archaeodictyomitra shengi YANG,Hori 1999, fig. 7-13 (= A. spelae Chiari abdomen, bearing c. 12 faintly developped costae visible per
et al. 1997). half-circumference. In that respect it resembles E. bortolotii
which nevertheless displays a more trapezoidal abdomen.
Remarks. - We do agree with Hori (1999) in considering A.
labronica as a junior synonym of A. shengi YANG 1993 because
it is practically impossible to differentiate these elongated, sub-
Genus Loopus YANG 1993 sensu Hull (1997) and Dumitrica
conical to subcylindrical morphotypes with smooth outline.
et al, (1997)
However, the specimen illustrated by Hori seems to us to be
Type-species: Pseudodictyomitra primitiva MATSUOKA & YAO
related to A. spelae because the last three segments are well in-
1985
dividualised on the outline. The known age range of this
species is Oxfordian to Tithonian.
Loopus doliolum DUMITRICA
(PI. 1, Fig. 19-20)
Archaeodictyomitra sp. A Dictyomitra sp. C, YAO 1984, pI. 3, fig. 4.
(PI. 1, Fig. 12) Pseudodictyomitra sp. Csensu Yao, Gorican 1994, p. 84, pI. 22, fig. 15, ? fig. 14.
Remarks. - Small (total length : 110 urn) conical form bearing « Dictyomitra sp. C» in Yao, Danelian et al. 1996, fig. 4d-e.
11 costae visible per half perimeter. It differs from A. patricki Loopus doliolum n.sp.; DUMITRICA in Dumitrica et al. 1997, p. 30, pI. 5, fig. 3,
5, 14
by its smaller size and absence of an inverted trapezoidal distal
part. Remarks. - Morphotypes that are not clearly constricted at
their distal part are included under this species. They are all
Genus Cinguloturris DUMITRICA in Dumitrica & Mello 1982 characterized by a single row of pores and faint costae inter-
Type-species: Cinguloturris carpatica DUMITRICA rupted by well-marked broad stictures. Its known age range is
in Dumitrica & Mello 1982 Oxfordian (Gorican 1994) to Berriasian (Dumitrica et al.
1997).
Cinguloturris fusiforma HORI
(PI. 1, Fig. 13) Loopus venustus (CHIARI, CORTESE & MARCUCCI)
Cinguloturris fusiforma n.sp. HORI 1999, p. 93, fig. 9.3-9.6, 11.6. (PI. 1, Fig. 21-23)
Remarks. - Only two specimens of Cinguloturris were Cinguloturris (?) venusta n.sp. CHIARI,CORTESE & MARCUCCI in Chiari et al.
1997, p. 66, pI. 2, fig. 4-5.
found in our material. One of them is incomplete and is ques-
tionably assigned to C. fusiforma. (pI. 1, fig. 14). This species Remarks. - This species is characterized by segments display-
was previously known only from the Palaeopacific (Japan and ing a single row of open pores situated in their middle part.
Far East Russia). C. fusiforma is likely to represent an offshoot Thin costae are placed above and below each pore in such a
lineage of C. carpatica from which it differs by its spindle- way that an upper and a lower row of depressions or relict
shaped contour. pores are formed on each postabdominal segment. Costae do
not loop around the pores. L. venustus is closely related to
Genus Emiluvia FOREMAN 1973 L. primitivus and might be its ancestor.
Type-species: Emiluvia chica FOREMAN 1973
Genus Pantanellium PESSAGNO 1977
Emiluvia pentaporata STEIGER & STEIGER Type-species: Pantanellium riedeli PESSAGNO 1977
(PI. 1, Fig. 15)
Emiluvia pentaporata n.sp. STEIGER & STEIGER 1994, p. 458, pI. 1, fig. 9-10 Pantanellium oligoporum (VINASSA)
Emiluvia bisellea n.sp. DANELIAN in Baumgartner et al. 1995b, p. 196, pI. 4018,
(PI. 2, Fig. 1)
fig. 1-4.
Ellipsoxiphus oligoporus n.sp. VINASSA 1899, p. 228, pI. 17, fig. 44.
Sphaerostylus oligoporus (VINASSA); Sanfilippo & Riedel 1985, p. 590, fig. 4.5;
Genus Eucyrtidiellum BAUMGARTNER 1984 Matsuoka 1992, pI. 3, fig. 4.
Type-species: Eucyrtidium (?) unumaensis YAO 1979 Pantanellium oligoporum (VINASSA); Chiari et al. 1997, pI. 3, fig. 8.
Eucyrtidiellum ptyctum (RIEDEL et SANFILIPPO) Genus Tethysetta DUMITRICA in Dumitrica et al. 1997
(PI. 1, Fig. 16-17) Type-species: Tethysetta pygmaea DUMITRICA et al. 1997
Eucynidium ptyctum n.sp.; Riedel & Sanfilippo 1974, p. 778, pI. 5, fig. 7, pI. 12,
fig. 14; not fig. 15.
Tethysetta (?) sp.
Eucyrtidiellum ptyctum (RIEDEL & SANFILIPPO); Baumgartner et al. 1995b,
p. 214, pI. 3017, fig. 1-8; Hull 1997, p. 82, pI. 48, fig. 10. (PI. 2, Fig. 10)
Remarks. - A single specimen found in our material recalls

T dhimenaensis with its fusiform outline and the presence of MARCUCCI, M. & STEIGER. T . 1995b: Rad iolar ian ca ta log ue a nd syste ma t-
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spinose ridges on intersegmental divisions. Un certainties are
BAUMGARTNER, P.O ., O 'DOGHERTY, L., G ORICAN, S., U RQUHART, E.,
due to the presence of 4 rows of por es per segment dipla yed PILLEVUIT, A . & D E W EVER, P. (E ds .): Middl e Ju ras sic to Lower Cre ta-
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Genus Tripocyclia HAECKEL 1881
basin : Ie Jurassique terrnin al-B erriasien des Chain es Sub alpines rnerid-
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RAYNAUD, J .-F. , R ENARD, M ., STEFFEN, D ., STEINHAUSER, N., STRASSER, PESSAGNO, E . A Jr. , SIX, W. M. & YANG, Q . 1989: The X iph ost ylidae H aeckel
A ., STROHMENGER, C & VAIL, P. 1993: Seq ue nce-stra tigraphic int erpret a- and Pa rvivaccidae , n. fa rn., (Radiolaria) fro m the North A me rican J uras -
tion o f Upper Tithonian-Berriasian refe re nce sections in South- East sic. Micropaleontology. 35 (3), 193-255.
Fr an ce: a multidisciplinary appro ach . - Bull . ce ntr es Rech. Explor.-Pro d. R IEDEL, W. R & SANFILIPPO, A 1974: R ad iolaria from the southern Indian
Elf Aq uit ain e 17, 151-181. Ocean, DSDP leg 26. In: D AVIES, T. Aet al. (Eds.): Initial reports of the
JAN DU CHENE, R, ATROPS, F. , D E R AFELlS, M. , E MMANUEL, L. & R ENARD, Deep Sea Drilling Proj ect. U . S. Governme nt Pinting Office , Wa shin gt on,
M. 2000 : Pal ynology and sequence stratigra phy in the tethyan uppe r ox- D . C, 26, 771- 814.
ford ian- lowe r kimmeridgian, S-E France , Bull. ce ntre R ech . Explor. Prod . R OST, D . 1885: Beitrage zu r Kenntnis der fossi len Radiolarien aus Geste ine n
E lf-Aquitaine 22, 273- 321. der Kreide . Pale onto gr aph ica, 34, 181-213.
JUD, R 1994: Bioch ron ol ogy a nd Systematics of Ea rly Cre taceous R ad iola ria SANFILIPPO, A . & RIEDEL, W .R 1985: Cre taceous R adi olar ia. In: BOLLI, H .M..
of the Western Tethys . Me rnoi res de Geologie (La usa nne) 19, 147 p., 24 SAUNDERS, J .B. & PERCH-NI ELSEN, K. (Ed s.): Plan k ton Stra tigraphy.
pis. Ca mb ridge U niv. Press, 573-630.
KOCIlER, R N. 1981: Bioch ronostra tigr ap hisch e Untersuc hunge n oberju rassis- STEIGER E . & STEIGER 1. 1994: New Rad iolar ia from th e "R uhpoldiger Ma r-
cher radiolarie n fiihrender Gesteine , insb esonde re de r Siida lpen. Mit- mor " of U rsc h lau ( La te J urassic , C hie mga u A lps, Ba varia ). Abha ndl.
teilu ngen a us dem Geologischen Inst itu t der Eidgenossische n Technis- Geol. Bundesan stalt -A . 50, 453-466.
che n Hochschule und de r Universitlit Z ur ich , Ne ue Fo lge 234 ,1-184. Vt NASSA DE R EGNY, P.E . 1899 : I radiolari de lle fta niti tito niane di Carpena
LAMBERT, E . 1999 : Le s Radiol aires cre taces d u Bassin Vocontie n : Stratigra - (Spezia) . Palaeontograp hia ita lica 4, 217-238 .
phie, Paleoen vironnem en t. T he se MN HN , 209 p., 5 pis. YAO, A . 1979: Radiol ar ian fa una from the Mino Belt in th e n ort hern part
LANGOISSEUX, O . 200 1: Strati gr aphi e seq uentie lle basee sur une ap proche of the Inu yam a A rea, Ce nt ra l J apan , Pa rt II: Nassella ria I. J ou rn al of
m ult idisciplinaire: Ie Kirnmer idgie n inferie ur de la fosse vocontienne (S - Geosciences , O sak a City Un iver sity 22, 21-72.
E de la Fra nce). Me rno ire. D iplom e d 'e tud es superieures d 'inge nieu r geo- YAO, A . 1984: Su bdi vision of the Me so zoic complex in Kii-Yura area , so ut h-
logu e. Univ. de G en eve. 89 p. west Ja pan an d its bearing on the Mesozo ic basin de vel opme nt in th e
MATSUOKA, A . 1992: Jurassic a nd Ea rly Cretaceous radio laria ns fro m Leg 129, so uthe rn Ch ichibu te rra ne. Jo urnal of Geosciences , Osak a City University
Sites 800 a nd 801, wes te rn Pacific Ocean. In: LARSON, R.L. e t al. (Eds.): 27, 41-103.
Proceed ings of the O cea n Drilling Progra m, Scientific R esults, Co llege YANG, Q . 1993: Taxon om ic studies of U pper Jurassic (Ti thonia n) R ad io lar ia
Sta tio n, T X (Oc ea n Drilling Pro gram ) 129, 203- 220. from the Ta ma n Format ion , eas t-ce ntral Me xico . Pal aeoworl d , Specia l
MATSUOKA, A 1998: Faunal composi tio n of earliest Cr e ta ceous (Berriasian) Issue 3, Nanjing U nive rsity Press, 164 p., 26 pis.
radi ola ria from the Mariana Trenc h in the wes te rn Pacific . News of O sa ka
Microp aleontologists, Spec. Vol. 11: 165- 187.
MATSUOKA , A. & YAO, A 1985: Lat est J urassic Radiolarians from th e Tori- Manuscr ipt received January 2004
nosu Group in Southwest Japan. Osaka Ci ty U niv., J. Geosci. 28,125-145 . Revision acce pted February 2005
S42 T . D ani elan , S. Lahsini & M . D e R afelis

Plate 1
Scanning Electron Micrographs of Radiolaria extracted from samples of the Meouge section. Bar scale (upper right) is equal to 100 J.Im for all figures.
1) Acastea sp.cf. A. remusa HULL, Mg-100; 2-3) Archaeodictyomitra apiarium (RUST), Mg-2; 4) Archaeodictyomitra etrusca CHIARI et al. Mg-29; 5-6) Archaeodic-
tyomitra patricki KOCHER, Mg-29; 7) A. patricki, Mg-2; 8) Archaeodictyomitra sp.aff. A. patricki KOCHER, Mg-2 ; 9) Archaeodictyomitra shengi YANG,Mg-29; 10)
A. shengi, Mg-77; 11) Archaeodictyomitra spelae CHIARI et aI., Mg-77; 12) Archaeodictyomitra sp. A, Mg-2; 13) Cinguloturris fusiforma HORI, Mg-100; 14) Cin-
guloturris sp.cf. C. fusiforma HORI, Mg-77; 15) Emiluvia pentaporata STEIGER & STEIGER, Mg-100; 16) Eucyrtidiellum ptyctum (SANFILIPPO & RIEDEL), Morpho-
type A, Mg-77 ; 17) E. ptyctum, Morphotype B, Mg-2; 18) Gongylothorax favosus DUMITRICA, Mg-37; 19) Loopus doliolum DUMITRICA, Mg-37; 20) L. doliomum,
Mg-29; 21) Loopus venustus (CHIARIet al.), Mg-2; 22) L. venustus, Mg-2; 23) L. venustus, Mg-133.

PLATE 1

Plale2
Scanning Electron Micrographs of Rad iolaria extracted from samples of the Meouge section. Bar scale (upper right) is equal to 100 urn for all figures except of
fig. 7 (=200 urn),
I) Pantane/lium oligopo rum (VINASSA), Mg-2; 2) Praecon ocaryomma scatebra HULL, Mg-21; 3) Protunuma japo nicus MATSUOKA & YAO, Mg-37; 4) P. japonicus,
Mg-37; 5) Ristola altissima cf.ssp. R. a. altissima (ROST) sensu Baumgartn er et al. 1995b, Mg-37; 6) Saitoum pagei PESSAGNO, Mg-21; 7) Spongocapsula palm erae
(PESSAGNO), Mg-21; 8) Stichocapsa tuscanica CIIIARI, CORTESE & MARCUCCI, Mg-77; 9) Suna sp., Mg-77; 10) Tethysetta (1) sp., Mg-77; 11) Triactom a fo remanae
MUZAVOR; Mg-77; 12) Tripocyclia sp.cf. Tiluciae J UD, Mg-100; 13) Tritrabs sp.cf. T. exo tica (PESSAGNO), Mg-37; 14) Wil/iriedel/um carpathicurn DUMITRICA, Mg-
77 ; IS) W. carpathicum , Mg-21; 16) Wi/liriede//um crystallinum DUMITRICA, Mg-37; 17) Zham oidellum ov um DUMITRICA, Mg-37 ; 18) Z. ovum , Mg-21; 19) Z.
ov um , Mg-2.
S46 T. Danie lan, S. Lahsini & M. De Rafelis

PLATE 2

DOl 10.1007/s00015-006-0609-y
The plankton turnover at the Permo-Triassic boundary,

emphasis on radiolarians
PATRICK DE WEVER 1, LUIS O'DOGHERTy2 & SPELA GORICAN3
Key words: Permian, Triassic, extinction, radiation, protists, Radiolaria

Mots cles: Permien, Trias, extinction, radiation, protoctistes, radiolaires
ABSTRACT RESUME
The examination of plankton biodiversity through Permian-Triassic period L'examen de la biodiversite du plancton au passage Permien-Trias semble
seems to display different patterns of evolution depending of the scale of study montrer differents types de manifestation selon l'echelle d'observation aussi
(taxonomy stratigraphy or biogeography). In this paper we present the state of bien en terme de stratigraphie que de taxonomie ou de geographic. A cette
the art of the plankton turnover at the Permo-Triassic and we review more epoque, Ie plancton etait surtout represente par les radiolaires. A une echelle
precisely the pattern of extinction and recovery of radiolarians during such pe- globale cette periode est certes marquee par des extinctions mais elles sont
riod, because at that time, plankton was essentially represented by radiolari- progressives comme Ie prouvent les deux ordres de radiolaires: les Albaillela-
ans. At a global scale the end-Permian to early Triassic period is marked by ria et des Latentifistularia. Neanmoins cette periode est surtout marquee par
several strong extinctions in the marine realm, and in the radiolarians they une enorme diversification post-crise, plus encore que par l'extinction. A une
occur progressively as exemplified by two orders of radiolarians: Albaillelaria echelle moyenne (ex. niveau generique et celui d'un pays) les renouvellements
and Latentifistularia. Nevertheless, this period is marked by a tremendous de faune semblent impressionnants, alors qu'a l'echelle des especes et de bas-
post-crisis diversification in both at family and generic level, more than extinc- sins particuiiers, on est conduit 11 se demander si des modifications resultent de
tion; an actualized revision of the diversity at the family level is also offered in reelles crises biologiques ou de problernes de conservation. II est amusant de
our review. At a moderate scale (i.e. genera and in a region) the modifications constater que plus Ie nombre d'etudes est eleve, plus la diversite apparait gran-
appear impressive while at specific and regional domain the message is not so de et 11 I'oppose, plus elles sont rares, plus Ie provinciaiisme est evoque.
clear, one can wonder if some crisis manifestation results from a taxonomic ac-
cident or from preservative conditions. In fact, strangely enough more the
number of studies is, more the diversity is high, and oppositely less they are,
more the provincialism is evoked.
sic crisis within the sedimentary successions containing well-

1. Introduction
preserved fossils indicate that during the Early Mesozoic a
Mesozoic deep-sea sedimentary rocks are widespread, the suc- genuine radiation of microplankton took place, one that in-
cessions are relatively complete and little altered, and they cluded heterotrophic as well as photoautotrophic protists. Un-
have been sampled extensively, Undoubtedly, the appearance doubtedly, the radiolarians are the most important and domi-
of evolutionary innovation among the plankton is enhanced by nant group among the microplankton after the Permo-Triassic
these circumstances, furthermore, many of the best preserved crisis. This is the reason why we analyse in this paper the
Paleozoic sedimentary rocks were deposited in shallow, epi- changes that occurred in this group.
continental seaways where open-ocean plankton might not be
present or abundant. The minute calcite plates of calcareous
1.1 Bioevents concept and faunal turnover
nannoplankton, delicate opaline silica tests (radiolarians) and
frustules (diatoms) might well be destroyed in more highly al- The idea of global bioevents is under discussion since the very
tered sediments, and if early dinoflagellates lacked archeopy- beginning of the science of Earth history. Georges Cuvier
les, their affinities would not be recognizable. Nonetheless, the (1769-1832) already recognized that most intervals occurred
spectacular diversity changes recorded after the Permo-Trias- in Earth history from time to time, each of them with a strong
Correspondance: Patrick De Wever

1 Museum National d'Histoire Naturelle, 43 Rue Buffon, F-75005 Paris, France. Email: pdewever@mnhnJr
2 Facultad de Ciencias del Mar, Universidad de Cadiz, 11510 Puerto Real, Spain. Email: [email protected]
3 Paleontoloski Institut Ivana Rakovca ZRC SAZU, Novi trg 2, SI-1Ooo Ljubljana, Slovenia. Email: [email protected]
Radiolarian plankton turnover, Permo-Triassic boundary S49

Table I. Effects of the main crisis of biod iversity
Families Genera
on familie s and genera with indicati on of calcu -
lated species loss (simplified from Jabl onski 19% ) Observed Calculated Observed Calculated
extinction ( %) species loss ( %) extinction ( %) species loss ( %)
End Ordovician 26 84 60 85
Late De vonian 22 79 57 83
End Permian 51 95 82 95
End Tria ssic 22 79 53 80
End Cretaceous 16 70 47 76
faunal turnover, later called catastrophe, extinction event, fau- those of species , because gen era are erected accord ing to
nal change , bio-event, and so on. The concept of catastrophe major features of morphology.
seems somewhat exaggerate as described and emphasized later The problem of estimating the importance of extinctions
by d'Orbigny, but one should set this idea in its proper context: and recovery depends upon the involved taxonomic level and
at that time the Earth was supposed to be approximately 6000 the way the counting is conducted. Sepkoski's family and
years old only. genus level (1989) summaries have allowed the species-level
Bioevents, often in connection with a significant change in extinction intensities of the five major extinctions to be calcu-
lithology, were used by d'Orbigny and most scientists con- lated (Table 1).
cerned in the first three quarters of the 19th century in order to The fossil record of planktonic life has the advantage of
subdivide the Phanerozoic. The result of these works is the abundant specimens, good stra tigra phic control, closely spaced
stratigraphic scale. Nevertheless, thr ee decades ago the fact of samples, and broad geographic distribution. The qual ity, how-
bio-event was disputed and even if these have now been ac- ever, varies substantially from group to group on the one hand ,
cepted, the global synchronicity of the events has often been and is largely dependent on the desired stratigraphic level of
doubted , and the question as to causation has been discussed precision on the other hand .
very controversially. If we still use the term catastrophism, we At a global scale, the Permi an-Triassic boundary was char-
are not using it in its original meaning (Cuvier's definition), acteri zed by the extinct ion of 50% of invertebrate families and
but accepting his valid observation of the existence of bio- 90% of invert ebrate species; extinction of fusulinid s, rugose
events . According to the main pha ses these bioe vents are and tabulate corals (orders), 2 orders of bryozoans , productids
named extinction-radiation or faun al turnover sequence. (order) and several ord ers of articulate brachiopods, trilobites
(class), euryperids (orders), blastoid s (sub-phylum) and sever-
al subclasses of atta ched echinoderms (mostly crino ids).
1.2 Scale of time
Some supposed catastrophes were in fact only the result of a
1.4 Estimation and counting: how variations can be illustrat ed
lack of record and they are no mor e considered as catastrophic
events. One should also be aware of global vs. regional cata- For several years, numerous researchers have been greatly
strophes. If the distinction is quite easy in theory, practically it interested in the study of biodiversity and its evolution thr ough
is not so obvious, as it will be discussed later herein. time. These studies pointed out several crisis periods. Never-
Biotic crises and events are sometimes distinguished. Dif- theless, it must be mentioned that the results depend on the
ferentiating from an event, a crisis spans a relatively long time methodology used. Some researchers simply counted the num-
interv al. This is the case where over a relatively long period ber of taxa presented in a publication without any taxon omic
the extinction rate exceeds the originat ion rate . Crises do not analysis. That is to say that the number of taxa in a particul ar
only occur in times of long-term environmental changes or work often depends on the taxonomic point of view of the
fluctuations, but also as a termin al phase of a taxon , when the author(s) of the used publicat ions. Hence simple counting
diversity of a group of organisms decreases, sometimes to- might be a source of mistak es. It is, therefore, advantag eous if
wards to zero. the counting of taxa is done by one or several specialists with
practice in the respecti ve gro up and after a thorough taxo-
nomic review.
1.3 Crisis: at which taxonomic level?
Biodiversity changes in the paleont ological record show that
1.5 Recovery
mass extinctions and recoveries occurred during some relative-
ly short time intervals. The se events are named and analysed A study by David Jablonski (1996) shows that recover y from
as "e volutionary crises". When considering crisis one might mass extinctions differs from one geographical region to an-
discuss the taxonomic level involved, the message can vary ac- other. Regions differ greatly from each other not only in terms
cording to the category. Plots of generic turnover should pro- of which and how quickly species diversified, but also in the
vide a better indication of morphological innovation than ratio of surviving local species to foreign invaders. In general,
S50 P. De Wever, L. O'Dogherty & S. Gorican

Phanerozoic patterns of phytoplankton radiation and extinc- In this palaeogeographic reconstruction, the Late Permian
tion parallel those documented for skeletonized marine inver- was characterized by falling sea levels (Ross & Ross 1988) ex-
tebrates, both augmenting and constraining thought about posing large areas in the northern high latitudes. But superim-
evolution in the oceans. Plants and animals may be the most posed on this longer trend a large-scale transgressions is record
conspicuous fossils in Phanerozoic rocks, but planktonic pro- in China, Iran, western Tethys, and in central Europe. This
tists are the most abundant. transgression became even more pronounced in the latest Per-
Much of our ability to discriminate among various hypothe- mian (Wignall et al. 1998), just before the Permo-Triassic tran-
ses explaining the evolutionary changes within geologic times sition, and reached its maximum in the Early Triassic, with
not only depends on careful and detailed comparisons of evolu- flooding of many areas of the Tethys and North America. In
tionary patterns, recognised in trophically, ecologically, repro- fact the Permo-Triassic boundary interval was a time of second
ductively, and developmentally different groups of organisms, order highstand of sea-level. This reinforces the fact that the
but also is essential to know how the earth system climate dri- end-Permian mass extinction cannot, in any regard, be consid-
ves the evolutionary process during critic periods of the past. ered as the product of sea-level fall (Hallam & Wignall 1999).
The time for recovery in the earliest Triassic was also high- Increasing values of 87Sr/86S r since the latest Permian-Early
ly variable depending on marine oxygen levels, like it has been Triassic time (Korte et al. 2003) are related to enhanced conti-
recently shown (Twitchett et al. 2004). These authors ob- nental weathering under humid climatic conditions in the latest
served a faster recovery in well-oxygenated environments Permian and the lack of a dense land vegetation in the Early
(seamounts) after the P-T extinction in the middle Gries- Triassic, prior to the Spathian, which are in agreement with the
bachian. An apparent delay in recovery after the end-Permian observed sea-level trend. A consequence of this flooding was
extinction event was observed in deeper-marine settings due to the spread of anoxic bottom waters into the shallow marine
widespread and prolonged benthic oxygen restriction. habitat during the early stages of transgression, which in turn
may have played an important role during the aftermath of the
P-Tr extinction event (Hallam 1989).
2. The Permo-Triassic Setting: Scenario for a crisis
In order to establish the duration of the Permo-Triassic ex-
tinction several attempts were made and estimates for the du-
2.1 Environmental constraints
ration of the extinction vary according to different inclinations
According to the classic Permian paleogeography, only one in fixing relevance to particular fossil groups or sampling loca-
continent (Pangea) was present at that time and surrounded by tions. Holser & Magaritz (1992) suggest a duration of 5-10 Ma
only one super-ocean (Panthalassa). In this land-sea distribu- for the crisis, whereas Hallam & Wignall (1997) prefer a rapid
tion, the concomitant drastic decrease of shelf surface added but not instantaneous mass extinction, although unfavourable
by the Carboniferous-Permian glaciation (Scheffler et al. 2003) climatic and environmental conditions prevailed during most
resulted in a drastic dynamic change of the global ocean circu- of Early Triassic (Wignall & Hallam 1992; Bottjer et al. 1996).
lation (Fiuteau et al. 2001). Radiometric ages point to a duration of 0.7-0.3 Ma for the ex-
An important event, occurring over a very brief span of ge- tinction interval, or at least for the interval between the 813C
ological time (initiated by 251.7 ± 0.4 Ma and terminated by decline and the P-Tr boundary (Bowring et al. 1998). Other
250.2 ± 0.3 Ma; Kamo et al. 2003) took place in this scenario: studies using estimates of sedimentation rates suggest more
the Siberian flood volcanism. A comparatively extensive flood rapid rates of ecosystem collapse, about 10-30 kyr for marine
volcanism occurred somewhat earlier in China (Emeishan extinctions (Twitchett et al. 2001; Rampino & Alder 1998).
traps, about 259 ± 3 Ma old; Courtillot & Renne 2003). The However, analysis of terrestrial flora (Looy et al. 2001) gives a
large amount of volcanic gases played a primary role in the different picture. The time lag between terrestrial ecosystem
Permo-Triassic extinction (Wignall 2001) but the rapid and collapse and plant extinction had a duration of 0.5-0.6 million
strong negative 813C excursion (c.a. 4-6%0), (Benton & Twitch- years, showing also a temporary diversity increase after the
ett 2003) is too great to be explained solely by the catastrophic negative 813C excursion.
collapse of ocean primary productivity in the late Changxingian
(Wang et a11994; Hallam & Wignall 1997). In this scenario and
2.2 For various groups: real extinctions or incompleteness of
an additional source of 0 2 to the ocean-atmosphere system is
the fossil record?
required, but the causes of the isotopic excursion seem to be
complex and multiple (Korte et al. 2004). In this sense, the Early Mesozoic radiating protists include taxa characterized by
Siberian volcanism triggered an initial global warming at the P- both siliceous and calcareous skeletons, as well as unmineral-
Tr boundary, which in turn melted clathrate bodies, releasing a ized groups. Possibly the break up of continents and the parti-
large input of methane into the atmosphere causing more tioning of the world's oceans, including strengthened circula-
warming (Heydary & Hassanzadeh 2003). The process contin- tion and upwelling of nutrients, contributed to the ecologic op-
ued in a positive feedback spiral that has been termed the 'run- portunities for diversification of planktonic organisms. After
away greenhouse' phenomenon (Benton & Twitchett 2003) the Permian crisis during the Mesozoic, planktonic protists un-
leading to the most dramatic extinction known. derwent remarkable evolutionary innovations and radiations.

s § ~ § !::
ECHIDNlNlDAE - - - ~
SECUICOLLACTIDAE - - -
PSEUDOROTASPHAER/DAE
----- --- ---
CERATO/KJSC/DAE -
CORYTHOECIDAE
ALBA/LLELLIDAE ALBAILLELLARIA
FOLLICUCULLIDAE
--+-
PALACANTHOLITHIDAE
RUZHENCEVISPONG/DAE
I. 1- -----1- ----
PSEUDOLITHELIIDAE
LATENTIFISTULIDAE LATENTIFISTULARIA
ORMISTONELLIDAE II
CAULETELLIDAE
INANIGUmDAE
PROVENTOCITIDAE - - - -
ENTACTINIIDAE - - - - - - - - - - - - - - - - 1 1 - - - - +.;..+...;.+:-#---
PALAEOSCENIDIIDAE
SPONGENTACTINIIDAE
m PALAEOLITHOCYCLIIDAE
z PYLENTONEM/DAE
~
POLYENTACTINIIDAE
HEXASTYLIDAE
o EPTINGIiDAE 1o;-..... 4'-U----~
:j HINDEOSPHAERIDAE Io;-'~#-.:.j
z MULTIARCUSELLIDAE H-:I-+#- '~ I
»:;0
PENTACTINOCARPtDAE
CENTROCUBIDAE
i> HEPTACLADIDAE
THALASSOTHAMNIDAE
I•
• • • •• • •• • ••• •••••• • •• • • • •••• •• ••• ~
CAPNUCHOSPHAERIDAE
SPONGOSATURNAL OID/DAE
SATURNALIDAE
KUNGALARIIDAE
ARCHOCYRTIDAE ••• •••••••••••• ••• • •

ARCHAEOSEMANTIDAE _ ............ ............. I-
POPOFSKYELLIDAE • • • •• • • •
POULPIDAE I ;"!-+..j.l.~-----4--
ACROPYRAMIDIDAE
TR/PEDURNULIDAE
FOREMANELLINIDAE
z BULBOCYRTIIDAE
»en PLANISPINOCYRTIIDAE
SPONGOSILICARMIGERIDAE
en PLAGIACANTHIDAE ----4- ~
m ULTRANAPORIDAE
r CUNlCULIFORMIDAE
r
»
:;0
NABOLELLIDAE
RUESTICYRTIIDAE I·
s SPONGOLOPHOPHAEN/DAE
MONICASTER/CIDAE
PSEUDOSATURNIFORMIDAE
DEFLANDRECYRTIIDAE
PSEUDODICTYOMITRIDAE
EUCYRTIDIIDAE
LIVARELLIDAE
CANOPTIDAE
BAGOTIDAE
I
PARVICINGULIDAE
? ANAKRUSIDAE - - - -
ARCHAEOSPONGOPRUNIDAE -
RELINDELLIIDAE - ~
OERTLISPONGIDAE • ·1r .'
en GOMBERELLIDAE • ' 11 .... ----~
"'C PYRAMISPONGIIDAE j-::..j.:~-----4----
C ACTINOMMIDAE •• • • • • • • • • • • • • ••• ••• • ••••• ••••• •••• ~
:i: LlTHELIDAE
m SPONGURIDAE ~
r XIPHOSTYLIDAE
~ ANGULOBRACCHIIDAE
PATRULIIDAE •
:;0 I "".;..;f-- - - - +-
s VEGHICYCLIIDAE
HEXAPOROBRACHIIDAE
PANTANELLIIDAE
HAGIASTRIDAE
lJ
Fig. 1. Stratigraphic ranges of radiolarian families through time (Redraw n fro m De Wever et al. 200]) .
S52 P. De Wever, L. O'Doghert y & S. Gorican

Radiolarians, dinoflagellates, calcareous nannoplankton, di- fossilized. In this sense the picture displayed by the fossil
atoms, silicoflagellates, and planktonic foraminifers either ap- record is biased. We will show later how this mechanism is il-
peared and radiated for the first time, or diversified from pre- lustrated in the radiolarian record at the Permo-Triassic crisis.
existing forms into a greater variety of new types. The causes Among Paleozoic plankton, remains of radiolarians (to-
of the early Mesozoic plankton revolution remain uncertain; gether with those of acritarchs) are particularly abundant, al-
probably they are as complex as the factors that promoted the though representatives of other planktonic groups such as
radiation of skeleton metazoans and protists at the beginning tintinnids and chrysophytes are found sporadically. We there-
of the Paleozoic. The fluctuations of diversity in planktonic fore focus hereafter on radiolarians, undoubtedly the domi-
protists have been suggested many times to be a function of nant plankton fossils in the Paleozoic and Triassic oceans.
variations in primary productivity and heterogeneity of water They first occurred in the Cambrian (Won & Below 1999) and
masses. During intervals of strong heterogeneity of water became morphologically complex through time (like
masses and high productivity, the oceans can support many Foraminifera) with the number of species increasing signifi-
specialized species that live effectively, at least for a part of cantly in the Ordovician and Devonian.
their life-cycles, within a narrow range of environments. When
productivity drops and oceanic water masses become less
3. The radiolarian record
structured and more homogeneous, narrowly adapted species
disappear. Obviously, as it has been shown by paleontology,
3.] Diversity of radiolarian families at a global scale
petrography and geochemistry, the oceans' physical structure
provides opportunities for plankton speciation and the biolog- Some evaluation of biodiversity for the Phanerozoic was at-
ic inter-actions of constituent populations. Unfortunately, such tempted several years ago for all fossil groups at the family
biologic interactions are not understood, even for living plank- level and these included radiolarians (Benton 1993). This ten-
ton, and so explanations concentrate on types of oceanograph- tative analysis, however, does not present a correct image of
ic changes that are known to correlate with stratigraphic the radiolarian world: only 35 families are referred to the
changes in diversity. whole geologic period whereas more than 130 are known (De
The fossil record indicates that recovery of ecosystems af- Wever et al. 2001, 2003). Some analyses were also done at the
fected by the crisis needs about 5 Ma (Bottjer et al. 1996; Schu- Permo-Triassic (Erwin 1993; Yao & Kuwahara 1997; Kozur
bert & Bottjer 1995), which seems to be achieved during 1998) but these curves also show contradictory results. In order
Anisian time (Gall et al. 1998). The apparently low recovery to give a realistic picture we present herein curves of radiolari-
together with the poor quality of the fossil record characteriz- an diversity following the recent work of De Wever et al.
ing the interval aftermath the P-Tr extinction modifies the per- (2001) who have homogenised the taxonomic criteria used for
ception of the magnitude of the end-Permian extinction, as the Paleozoic, Mesozoic and Cenozoic, and updated the strati-
well as, it masks the paths undergone by the recovery. These graphie ranges for all families known. At a global scale (Figs. 1,
particular patterns have led to explain the recovery in terms of 2), rapid changes in the biodiversity of radiolarian assemblages
recolonization of damaged Early Triassic environments by the appear to be fairly rare throughout the geological record ex-
so-called "Lazarus taxa", that is, super-specialized taxa that cept during the Early to Middle Triassic, after the major mass
migrate to "unknown" refuges during extinction periods and extinction event which occurred at the Permian/Triassic
that reappear "unchanged" after millions of years when ade- boundary. During this time interval of nearly 20 Ma a large
quate environmental conditions are re-established. One may number of families occurred for the first time (Fig. 2).
wonder if it is just the simple variation in the quality of fossil Among the six radiolarian orders occurring in Paleozoic-
record which masks our perception of the extinction event Mesozoic deposits (Fig. 1), three were more or less extinct at
magnitude and hides devlopments during the recovery phase. the Late Paleozoic crisis (Albaillellaria, Archaeospicularia and
According to Twittchett (2001) the incompleteness of the Latentefistularia). Only one order clearly crossed the event
fossil record in the Early Triassic was caused by a sharp de- (Entactinaria), and two (Nassellaria, Spumellaria) sporadicaly
crease in productivity at the end of the Permian. The drop of occured before, but clearly diversified after the crisis.
productivity during the crisis in turn lead to a reduction of the Representatives of the Archaeospicularia are the oldest ra-
biomass, which must be readjusted by decrease in abundance diolarian order and the only one that clearly became extinct
(but retaining body size) or decrease in body size (but with before the Permo-Triassic crisis. The Entactinaria had a con-
similar population size). Those forms that reduce their popula- tinuous and regular diversification during the Paleozoic, and
tion sizes playa "Lazarus role" giving an apparent extinction, crossed the Permian/Triassic boundary to be rather common
because they reduce in number so much that they are below and diversified in the Mesozoic.
paleontological detection levels (Wignall & Benton 1999). Evolution within the Albaillellaria seems to have been con-
Only when the primary productivity returns to pre-crisis nor- tinuous and regular during the Paleozoic. During the Car-
mal levels, Lazarus taxa seem to "reappear". Obviously, those boniferous and the Permian, the Albaillellidae underwent a
taxa that reduce their body size, but retain or even increase great and biostratigraphically important diversification. In the
their population size would be the best candidates to become Late Carboniferous, simplifications of the internal spicule gave
Radiolarian plankton turnover. Permo-Triassic boundary S53

Late
Rhaetian
Norian
Indosinian/
Yenshanian
orogenies
1 Appearance
of family
,
U
(J)
(J)
~
0::::
I-
Middle
Carnian
Ladinian
Chert
~
~
•
Anisian
Anoxia gap
Spathian
I
Smithian
Early Scythian Dienerian
Evaporites ~
~ I
Griesbachian
Changhsingian I
Late
Lopingian
Guadalupian
Wuchiapingian
Caoilanian
Wnrrli. n
II
Ir t
Siberian
Euxinia • I
Roadian traps
Leonardian Family
Artinskian
diversity:
I
Z Emeishan
« c
('IJ
2 'C
0::::
ui
0...
Early
('IJ
~
E
Pangean
u .-
j
('IJ
Cisura lian (J)
assembly
Glaciation
Asselian
wt I- I I I
o 10 20 30 40 50
Fig. 2. Radiolarian diversity at the family level facing the main events of the Permo-Triassic boundary. The number of appearance of new families clearly corre-
sponds to the periods of main stresses; modified after Kidder & Worsley (2004).
rise to the last Albaillellaria (Follicucullidae and Palacantholi- (Ishigaum and Cauletella, cf Takemura et al. 2003) seem to
tidae). During the Late Permian the evolution of some albail- cross the PIT crisis, and then the order became extinct in the
lellarians underwent two different directions: one was to build Early Triassic. The presence of Spumellaria in the Paleozoic is
a complex shell (Neoalbaillella), the other was to simplify this still uncertain; in fact, many of the Paleozoic spherical radio-
shell (Follicucullus); only the latter crossed the boundary (Bra- larians, previously considered to be Spumellaria contain the
gin 1991; Sugiyama 1992, 1997; Yao & Kuwahara 1997). It is remains of an inner spicule, indicative of Entactinaria (De
known that under environmental stresses, some radiolarian Wever et al. 2001). Since their identification depends on the
species seem to be able to adopt a regressive/primitive charac- study of the innermost skeleton, and in most cases this is ab-
ter. Often this process happens along with a reduction or even sent or not well preserved, they could be confused with some
loss of skeleton (Guex 1981, 2001; De Wever et al. 2003). entactinarians (in which the initial spicule is dissolved), with
The evolution of Nassellaria was very slow during the Late which they share their general morphology. One could consid-
Paleozoic and they practically disappeared during the Permian er, for example, that they are represented by some many-lay-
so that one can say that the main nassellarian groups appeared ered spongy skeletons occurring in the Permian assigned to the
during the Triassic. The oldest Triassic nassellarians are mor- Relindellidae and Archaeospongoprunidae. Nevertheless,
phologically similar and possibly related to Paleozoic Ar- studies in Oregon and South China (Blome & Reed 1992;
chaeosemantidae. Shang et al. 2001) show that true spumellarians (Pantanelli-
The Permian is not marked by the appearance of new or- idae-like forms) seem to be already present before the PIT
ders of radiolarians, although some representatives of proba- boundary. More recently some representative of several fami-
ble spumellarians occur together with the last Latentifistularia lies were reported from uppermost Permian from China (Feng
(Cauletellidae). Some representatives of Latentifistularia et al. in progress); they are respectively Pyramispongidae

Norther High
ICS Faunal Southern High Latitude Tropica l
EPOCHS
stages
Latitude 813 C PDB ('11.0)
stages
Chert Color Chert Color
-2 o 4 -40 -30 -20 -10
C t:
ctl Spathian Ql
J::
oxic anoxic
:i: U
Ql
C "0
Ql Ql
6 Smithian 0:::
Bio-Geosphere Perturbation
Induan Dienerian
----
~.~
C Ol
ctlc
c J:: .-
f)
.~ '2
E .!l!
t:
'-
Q) ,S
0> C
ctl
Thuringian Ql
'0, (propa rlj J::
o.. E- C U
"0
Ql III
Q) 3 '0. "0 o:::, t:
Ql
Ql
-
co ~
ctl
-l
:c
o
0::: >- J::
~u
::J (9
3:
260 .4 . 0 7 ..L..._ _..l..- ---lL..-- I . L..- ....L..._- L_ _- - ' - - - l _....L ..l..- .L..-_....L.._'--_ _- - '
Fig. 3. Stratigraphy of events according to their geogra phic (latitud inal) position. It points out the lack of inform at ion for the critical levels; Carb on isot opes from
Bent on & Twitchell (2003) a nd Atudorei (1998). Sulfur isotope from Kato el al. (2002).
(Tetrapaurinella, Paurinella), Gomberellida e ( Tam onella) and In the aftermath of the mass extinction, when th e last rep-
Oertli spon gidae iParaoertlisp on guss , Since the se genera ex- resentatives of the orders Alba illellaria and Latentifistul aria
tent upward, but have not been yet mentioned in the Lower compl etely disappeared, with the exception of some gen era:
Tri assic, they are considered as Lazaru s taxa by these authors. Follicucullus, Ishigaum, Cauletella which are mentioned till the
According to the dat a reviewed above , no new orders of Early Tri assic, (Sugiyama 1997; Takemura et al. 2002), the
radi olari ans appeared after the Pe rmian -Triassic crisis. Th e Earl y Triassic fauna s wer e characterized by extremely low di-
only exception might be Nassellaria, if we consider that Meso- versity and poor preser vation (Hori et al. 2003; Sashid a 1983,
zoic nassellarians are not direct descendants of Paleozoic rep- 199] ; Sugiyama 1992, 1997; Suzuk i et al. 2002). Th ese faun as
resent atives, all of which disapp ear ed by the end of the Car - mostly contain entactinarian s (Cryptostephanidium, Entactinia,
boniferous. The existing linea ges of radiolarians, nevertheless. Hegleria, Parentactinia, Pseudostylosphaera, Tibor ella), some
produced an impressive number of new morphotypes grouped sparse mono- and dicyrtid nassellarians (Archaeosemantis,
into new families. Of the 14 families known in the Permian, 9 Ho zmadia, Poulpus, Spongosilicarmiger?), and rare spumel-
continued to exist in the Mesozoic and among the 55 families larians with inner spicule (Gl om eropyle, Plafkerium?). In addi-
known in the Triassic, 25 appeared during the Early and Mid- tion to this extremely low divers ity, the poor preservation and
dle Tri assic. The Middle Triassic was undoubtedly the main th e scarcity of good outcrops, limit the information that is now
epoch for radiolarian radi ation; mor e than a third of the tot al available about Early Tri assic faun as (Fig. 3). Furthermore, we
number of radiolarian families, recognized from the Cambrian stress the importance pla yed by the apparent abrupt disap-
to the Pre sent originated during th is impo rtant peri od of pearan ce of radiolarian chert s that were common in the deep-
plankt on radiation (Figs. 1, 2). marin e sections of South China, Japan, and Western Canada in
What could have caused these tremendous radiations and the Late Permian (Fig. 4). T hey reapp eared only in the Spath i-
evolutionary inno vation s? For the most part the causes rem ain an afte r a "chert gap" of 7 to 8 Ma (Isozaki 1994; Kakuwa
uncertain , but one of them could be the bre ak-up of cont inent s 1996). This is, in part, the rea son why the abrupt chan ge in ra-
and the partitioning of the world 's oceans that produced diolar ians is amplified after this long period of instability. For
strengthe ned circulation, as suggested by ocean modelling, and the reasons mentioned abo ve the curre nt information available
induced up-welling of nutrient s (De Wever et al. 1994). It is for the radiolarians durin g the Early Tria ssic should be consid-
known th at during the Earl y Mesozoic, modern oceanic basins ered quite incomplete and the evolutionary pattern undergone
began to form (Neotethys spreading) and new oceanic current by the gro up at this interval remains poorl y known .
systems appeared. Numerous microplates began their separa - Follo wing the "che rt gap ", the rapid diversification of Tri-
tion in the Tethys Ocean , crea ting new seaways in the western assic radiolarian assemblages during the Early to Late Triassic
Tethys and mod ifying oceanic circulation. (De Wever & is the most prolific that has ever been recorded in the Phanero-
Baudin 1996). zoic. For example, the extincti on of radiolarians between the
Radiolarian plankt on turn over , Permo-Triassic boundary S55

60- ,- - ----r ,
30 .--- - ----,-----,
PERMIAN TRIASSIC PERMIAN TRIASSIC
50
40 20
IV 1Il Fig. 4. Numb ers of quot ed families and genera
Q; 30 oS!
c: E near the Permian-Triassic Bound ary according to
CI> IV
C) U. several authors. and from raw dat a (Ume da 2002 +
20 10 Sugiyama 1997 and Kuwahara & Yao 2001 + Yao
& Kuwahara 1997) and from synthetic data (Yao
10 & Kuwahara 1997 and De Wever et al. 2(03) .
For genera all curves follow the same trend, they
0+-- - - -+-- - - -+-- - - --\ 0-'-- - - +-- - - +-- - --1 decrease from Wuchiapingian to Scythian. For
Wuch . Chang. Scyth. Wuch. Chang . Scy th .
families appears a discrepancy between curves. In
fact information from Yao & Kuwahara (1997)
--tr- Yao & Kuwah ara ; 1997 (synth.) -+- Kuwa . & Yao; 200 1 + Yao & Kuwa.; 1997 (raw data)
have bee n complied but not pre viously homo-
-+- Umeda; 2002 + Sugiyama; 1998 (raw data) --0- De Wever et al.: 200 1 (synth.) genised. Data treatment influences the conclusions.
Permian and Triassic (Fig. 1) seem s to have been a slow pro- (5- 6%0) is recorded (Hallam & Wignall 1997; Benton &
gressive and important phenomenon. But, the most distinctive Twit chett 2003; Korte et al. 2004). The available data for the
featu re for this event is not th e diminishing number of taxa at diversity at generic or family level are insufficient to correlate
both family and generic level s near th e PfT boundary, but to th e carbon isotope cur ve. We can only observe th at ext inc-
rather an important diversification that followed it, during th e tion (and th e drop on diver sity) at famil y or generic level dur-
Triassic: nine families cros s the PfT boundary (six of these Pa- ing the end-Permian (cf. Fig. 3) seem s to be link ed with de-
leozoic families became ex tinct by th e end of the Triassic) and cre asing values of ol3e. But, it see ms ob vious that any inter-
45 new families originated during the Triassic; in other words pretati on of the ol3C signal with out taking into account othe r
the fauna is almost complet ely ren ewed . T he Figure 4 displ ays geo indicators and a det ailed correlati on has actually not much
a similar tendency between family and gen eric curves, but a pale obiological significanc e.
rever se tendenc y is observed in the dat a from Yao & Kuwa -
hara (1997). The differences can be explained because th e
3.2 P- T radiolarian div ersity at a detailed scale
supra -generic assignments into familie s follow ed by these au-
thors is different from that we ha ve used for our curves (De
A vailable data
Wever et al. 2001). Like it was mentioned in the foreword of
th is art icle, Figure 4 well-displays how the biodiversity (espe- Only 21 years of invest igation covers all our informati on o n
cially at family level) is strongly biased by the taxonomy em- E arly Triassic radiolarians; a rath er moderate period of re-
ployed in the analysis of faun al turnovers. Irrespective to this search, and with still few dat a. Table 2 reports the available
correlation, the Middle Triassic is a period of unprecedented taxonomic data for the Early Triassic: 19 papers from the cir-
blooming and accelerated evolution of Entactinaria, Nassellar- cum-Pacific region and only 12 pap ers from the Tethyan area
ia and Spumellaria (Fig. 1). This figur e clearly shows that En- (So uth China, Thailand and Turkey) (Fig. 5). It is also interest-
tactinaria represent the most num er ou s radiolarian group that ing to note that the majority of publications deal with Spath ian
survived both, the PfT boundary and the Early Triassic crisis, radi olari ans. The earliest Tri assic (Griesbachian) radiolarians,
however, the nassellarians represent the gro up with the most on the other hand , are known fro m New Zealand onl y and
rapid evolution and diversification dur ing th e Triassic and wer e first described in 2002. Th e review clearly shows the
later. Thi s fact well illustrates how the nassellarians displ ay a weakness of our knowled ge abo ut the radiolarian stratigraph y
powerful reco ver y during po st-e xtinction per iods as has be en during this phase of reco ver y, and the number of taxa ide nti-
signalled for the Mesozoic time (O 'Dogherty & Guex 2002) fied at genera or species level is extremely low. It is now
Since th e Anis ian the entactinarian s, nassellarians and spumel- kn own how poor preservation can influence the apparent com-
larians diversifi ed simultaneously resulting in new familie s th at po sition of the fauna (O' Dogherty et al. 2003). The small num-
will become th e base for th e spe ctac ular radiation at gen eric ber of species illustrated or describ ed by publication (Ta ble 2)
and species level. also stro ngly contributes to an unr eali stic vision about th e tax-
Usually under anoxic environments radiolarian diversity onomic re-organization and th e radi ation of radiolarians du r-
minima occur simultaneously with the Ol3C positi ve excursion ing th e Early Triassic. Th is insuffic ienc y of data, evidently, can
(O 'Dogherty & Guex 2002) . During th e Changxingian a minor be , in some cases, provision ally translated into a wrong inter-
positive peak is concomitant with th e end Permian extinction pretation about the paleob iogeogr aphy and phylogeny at both
(cf. Holser et al. 1991), but just before the Permo-Triassic or der and family levels.
boundary a conspicuous neg at ive carbon isotopes excursion All these papers report th e pr esence of highly abundant
S56 P. De Wever, L. O'Dogherty & S. Go rican

Table 2. Synthesis of published papers concern- Faunal stage Author GenJSp. n.sp
ing Early Triassic radiolarians with respectively:
Age, author(s), number of recorded genera/num- Griesbachian Takemura et al. 2002, 2003 (Waipapa, New Zealand) 51?
ber of species and nov. sp. This table underlines
Kozur et al. 1996 (Karakaya, Turkey) 2/2 2
how new study implies new taxa, as a result of
Sashida et al. 1998 (Phatthalung, Thailand) 5/5
the very poor knowledge of these fauna of that Dienerian
Feng et al. 2001 (Hunan, China) 2/2
precise period.
Kamata et al. 2003 (Waipapa, New Zealand) 7/?
Blome & Reed 1992 (Oregon, USA) 4/4

Smithian Yao & Kuwahara 1997 (Inuyama, Japan) 4/7
Yao & Kuwahara 1999 (Ziyun,China) 2/2
Sashida 1983. 1991 (Kanto, Japan) 7/13 8

Zhang et al1999 (Nadanhada, China)
Bragin 1991 (Shikote Alin, North Eastern Russia) 4/4
Sashida & Igo 1992, Sashida et al. 1993 (Phatthalung, Thailand) 6/7 1/4
Feng 1992 (Southwest China)
Feng & Liu 1993 (Yunnan, China)
Sugiyama 1992, 1997 (Gifu, Japan) 22/26 19
Nagai & Mizutani 1993 (Gifu, Japan) 11/14
Spathian Kusunoki & Imoto 1996 (Kyoto, Japan)
Yao & Kuwahara 1997 (Inuyama, Japan) 7117
Isogawa et al. 1998 (Ashio, Japan) 12(17) 60
Kamata 1995, 1999 (Tochigi, Japan) 16/30 5
Feng et al. 2000 (Yunnan, China) 12/27
Sashida et al. 2000 (Northern Thailand) 38/50 4
Yao & Kuwahara 2000 (Guizhou,China) 11/14
Suzuki et al. 2002 (Ashio, Japan) 15/30
Hori et al. 2003 (Kaka Point, New Zealand) 7/15 3
Entactinaria along with rare Nassellaria and undeterminable assic boundary. As was mentioned earlier, the preservation of
Spumellaria. The Late Permian extinction undoubtedly played radiolarian faunas were so strongly affected by the end Permi-
an important role in the evolutionary history of Radiolaria. A an environmental collapse that a conspicuous "radiolarite gap"
quasi-complete renovation took place during the aftermath of has been described (Isozaki 1994, 1997; Kakuwa 1996) from
the Permian crisis at generic level (only a few genera belonging the uppermost Permian (Clarkina iranica Zone of conodont)
to 10 families), although at order level all the groups seems to until the end of the lower Olenekian (top of the Smithian) last-
cross the critical boundary (Fig. 1). It is interesting to note the ing about 10 Ma. But recent investigations show that this gap,
presence of some radiolarians with Triassic affinities (i.e.: pan- spread over the world, does not exist in high southern latitudes
tanelliids) in the uppermost Paleozoic strata (Shang et al. (New Zealand) where radiolarites have been recently de-
2001), as well as some Paleozoic forms (Fig. 3) which have scribed (Kamata et al. 2003; Takemura et al. 2002, 2003).
been recorded in the Early Triassic (Sugiyama 1992, 1997; Yao At the family level and global scale, we have seen that the
& Kuwahara 1997, 1999). number of radiolarian families was not affected (Figs. 1, 4).
But, oppositely, radiolarian species (and genera) were strongly
affected at low latitudes but not in high southern latitudes
Discussion
(Kozur 1998). This discrepancy has been interpreted by Kozur
Evoking a crisis requires precise definition of the involved (op. cit) as a differential effect of extinction in southern high
level (taxonomic, geographic or stratigraphic) because a phe- latitude with respect to low or northern latitudes due to a huge
nomenon may appear sharp at one scale but not at a different volcanic activity in northern hemisphere (Siberia and China).
one. That is why at the family level and global scale radiolari- The long phase of recovery would be due to immigration from
ans did not seem to be strongly affected by the Permo-Triassic the southern high latitude cold water (less affected than those
crisis, but seems very sharp at specific level (i.e. 96% of species of high northern and low latitudes) rather than from tropical
disappeared, cf Kozur & Mostler 1982). Other authors (Suzu- waters, which would be a faster process (Kozur 1998, Kozur et
ki et al. 2002) from a database of published data propose at al. 1996). For this author, this process would be slow (c.a. 5
least three distinct turnovers during the Early Triassic time, Ma; late Scythian to early Anisian) because it would require at
the most conspicuous turnover being that of the Late first the adaptation of some elements of the cold water fauna
Olenekian-Anisian boundary (143 species disappearing and to warm tropical conditions. In addition, the recovery of the
118 new species appearing). radiolarians, for this author, started at the base of the Spathian
The small quantity of data is undoubtedly related to the where, however, not yet a distinct radiation of Triassic ele-
scarcity of localities with a complete record for the Permo-Tri- ments occurred but mainly Paleozoic (Late Devonian-Missis-

Fig. 5. Paleogeographic map for the Early Trias-
sic (adapted from Hallam & Wignall (1997), dis-
playing the het erogeneous distribution of studied
sites for radiolarian s. The number of publication
for each region is as follow: Oregon: I; Turk ey: 1.
3D· Thailand: 4: S.China : 7; Japan: 13: NE Russia:l ;
New Ze aland : 4.
It clearly reveal that besides tropical western Pan-
thalassa almost nothing is known (only 1 locality
for Oregon , Turkey, New Zealand and NE Rus-
sia).lt is therefor e difficult to have a representativ e
scope of events which happens at that time. Note
the type of projection (with a sigmoid equat or)
which seems to locate New Zealand close to south
pole while it is at 300S , as mentioned on figure.
sippian) "Lazarus taxa" reappeared from "unknown refuges", served, being more fragile, so some may have appeared and
which are the origin for some phylogenetic connection at fam- evolved during this time of environmental crisis without any
ily or even at order level. The proposed " Lazaru s recovery" is fossil record. The strong reduction of biomass at time of the
very big, about 50% of the genera that disappeared at the PfTr crisis is undoubtedly related to the collapse of the primary pro-
boundary re-appeared only in the Olenekian-Middle Triassic ductivity. In order to equilibrate the global biomass reduction,
interval. As shown here above, real data neither support nor some strategies can be drawn (Fig. 6) . Like in several groups
contradict this interpretation, they are just insufficient at pre - (e .g. Foraminifera, Leven & Korgachin 2001) radiolarians may
sent time . strongly decrease their population but maintain their body
From a paleobiogeographic point of view, some paleolati- size, as it is the case of some Paleozoic relict forms (some al-
tudinal differences were mentioned for Anisian radiolarians, baillellids belonging to the genus Follicucullus) that could be
where stratigraphic and taxonomic data are quite abundant considered to be "Lazarus forms" (in the sense of scarce
and consistent (Bragin 1991; Sugiyama 1997; Kozur & Mostler record due to a poor preservation rate). Also some forms that
1994). In this sense, Aita & Bragin (1999) established the reduce the complexity of their skeleton by partial loss of the
genus Glomeropyle for a large, thick-walled form occurring ex- outer components (=proteromorphosis sensu Guex 2001) , but
clusively in high (south and north) latitudes, where other non- moreover having a more delicate skeleton, are more suscepti-
Tethyan species co-occur with well known Tethyan represen- ble to be dissolved, and therefore reinforce the Lazarus effect
tatives (Hori et al. 2003). The same type of reconstruction (i.e. Latentifistulidae). On the other hand, other forms reduce
using a compilation of the published data for the late Early their body size under environmental stress by, i) reducing the
Triassic (Olenekian) has been presented (Suzuki et al. 2002) . size of the entire skeleton (as is the case of large spherical
These authors find a strong provinciality between the radio- spumellarian-type forms of the late Permian (Shimizu et al.
larian faunas of the western Panthalassa and the eastern 2004), or ii) by losing the external skeleton (proteromorphosis,
Tethys. Nevertheless, the non-homogeneous data used in the i.e. Entactinaria forms); the population size being not neces-
comparison (4 studies for the eastern Tethys against 14 papers sarily reduced. It seems likely that many spicular forms be-
for the western Panthalassa) and different stratigraphic levels longing to entactinids in the earliest Triassic in the aftermath
led them to consider that radiolarian provinces were signifi- of the crisis are the result of a severe loss of the outer skeleton.
cantly distinct during late Olenekian because there is only a This simple and economic architecture is a potential source for
5% of coincidence at specific level. Comparing the same data new phylogenetic developments and would explain also the
at a generic level rises up the coincidence to 50%. Such differ- sudden appearance of so many new groups (families, genera
ences, in our opinion, probably derived from an inhomoge- and species) since the beginning of the Middle Triassic when
neous taxonomy (mostly open nomenclature assignations) in the preservation of radiolarian skeletons improved dramatical-
the utilised synthesis, partly at least due to the bad preserva- ly. The strong diversification also coincides with the rising val-
tion recorded in this interval worldwide. ues of strontium isotope ratios (Korte et al 2003), interpreted
as the consequence of Early Triassic sea level rise and the re-
turn to normal geo-biosphere conditions.
Alternative explanation
It is interesting to note that in spite of the low diversity as-
It is clear that the plankton response to the adverse environ- semblages displayed in the earliest Triassic, all the information
ment of the Late Permian is complex and unfortunately badly available shows a clear domination by Entactinaria, with many
documented. Forms with thin skeletons have not been pre- spicular forms, some Spumellaria and mono- or dicyrtid Nas-

>-
II~daPlaH"
~
w weak Real Extinctions
> radiation
o
~
o
w
~
radiation
reducing body volume Lazarus effect
* C> h.
~
-c
0<) H >tt-- CI)
CI) Z C>
¢- <> ~
RICH >r(.. >n-- -c :U)
~
I-
~
0 0 w
FOSSILIZATION as w I
~ U
,
retained population reduced population reduced population
Appar ent Extinctions
AA
Z outer skeleton
0 skeleton
i= reduction loss
U
Z
i=
Q-<)¢¢ @GJ)
AA
X
w
uJ
~
o, -c> Q- ¢
Spumellaria
QJ)fg)
Entactinaria Latentifistularia Albaillelaria
REDUC ING BODY VOLUME - SIMPLIF ICATION CONSTANT BODY VOLUME
Fig. 6. Stress effects on radiolarian popul ations (see explanation in text )
sellaria. Multicycrtid Nassellaria do not occur befor e the Tri assic, and can be due to the extremely low abu ndances of
Anisian. These sha pes fit perfectly with the ability of some planktonic fauna (basically radiolarians) but it could also be a
specie s to adopt a regressive/primitive character such as spicu- result of exceptionally poor preserv ation. Both phenomena are
lar forms and monocyrtids (De Wever 1982, De Wever et al. undoubtedly connected. For this reason, the apparent faun al
2003) . dissimilarity displa yed betwe en some local studies, actually can
All the data available toda y show that the picture we have not be considered as a solid argument for discussing an early
for Early Triassic radiolarian faun a is very far from being com- and high provinciali sm because it would lead to an unrealistic
plet e. We agree with Wignall & Bent on (1999) who emphasize paleobiogeography mod el for the group at Early Triassic time .
that the abundance of Lazarus taxa (largel y favoured by Kozur A supplementary problem is that larger the interval lacking
1998, in the aftermath of the Late Permi an extinctions is most data is, smaller the probability is to illustrate the origin of the
probably the reflection of the extreme rar ity of these organ- phylogenetic lineages . Indeed, the main period of diversifica-
isms at this time, as well as, the incompleteness of the strati- tion (Spathian) corresponds to the period for which the num-
graphic record (radiolaritic gap) and its poor preservation. ber of publications is the highest (2 for Griensbachian, 4 for
the Dienerian, 2 for Smith ian and 17 for Spathian). Thi s coin-
cidence reminds that of Sheeh an (1977) who showed that the
Final consideration
highest specific diversity of Phanerozoic marine invertebrate
The low diversity displayed by the currently available data re - per Ma corresponds to the largest areal exposure of rocks of a
flects the poor qua ntity of the pelagic fossil record at the Early given age .
Radiolarian plankton turn over , Permo-Triassic boundary S59

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S62 P. De Wever, L. O'Dogherty & S. G ori can

DOl 10.1007/s00015-006-0602-5
An online micropaleontology database: Radiolaria.org

JANE K. DOLVEN 1 & HANS A. SKJERPEN 2
Key words: Online databases, paleoinforrnatics, www.radiolaria.org
ABSTRACT
Growing amounts of available radiolarian data require better information- is reviewed by appointed web-editors to ensure quality and correctness.
management systems. Managing, preserving and exchanging information can Species names, descriptions, synonyms and references are based on published
be done by using online databases with a web interface. Radiolaria.org material, and this is linked to a picture-archive showing representatives of the
(www.radiolaria.org), an example of such a web-database. contains species in- species from different geographical areas and/or periods. In addition, Radio-
formation and other resources, and has been designed by and for the radiolar- laria.org also serves as a communal website with e.g. news, events and discus-
ian community. In Radiolaria.org researchers can contribute with information sion forums.
and this body of data is then made freely available. All submitted information
1. Introduction the system is of a high quality; 3) to transform the data into a

suitable format for storage; 4) to enter the data into the sys-
Paleoinformatics, the area of paleontology concerned with
tem; and 5) to have the data quality controlled (by web-edi-
management of information, including preservation of system-
tors). However, once such a system is operational, it has the
atic information and expertise, was first introduced in the pale-
potential to make micropaleontological work more efficient by
ontological literature by MacLeod & Guralnick (2000) and
making better use of the information and resources that al-
MacLeod et al. (2000). These authors pointed out that, due to
ready exist. Eventually, similar initiatives may be interconnect-
the increasing pace of micropaleontological data gathering,
ed creating networks of knowledge bases.
there is a need for more efficient ways of storing, accessing,
In the following sections, we will present the website Radi-
exchanging, analyzing and presenting data. These demands are
olaria.org (www.radiolaria.org). The idea of making Radiolar-
met by the adoption of more sophisticated information-man-
ia.org came up during late spring 2000, due to our need of hav-
agement technology, paired with training in the development
ing a database with information (images, descriptions, refer-
and use of such technologies. There is also a need to coordi-
ences, synonyms, geographic and stratigraphic distribution
nate initiatives in these areas to ensure maximum benefits for
etc.) about radiolarian species. Previously we had used books,
the research community as a whole.
reprints, library-copies, and various computer documents to
Online databases (defined herein as databases with a web
keep information at hand, but a database would make the in-
interface) are one effective way of managing micropaleonto-
formation more easily available. Instead of making a local sys-
logical data. The information is entered or accessed easily, at
tem, we decided to put the database online so other radiolari-
any time, from anywhere with the use of a web browser. These
an researchers also could access and add data. The first version
web-databases enable easy searching and comparison of data,
of Radiolaria.org was presented at the Interrad-meeting in
and make it easier to extract needed data in a suitable format,
2000 in California, where it received positive feedback from
which is important for data exchange.
fellow researchers. Since then, there has been both functional-
Building online databases requires large amounts of time
ity and content added to the system. Radiolaria.org is now an
and work: 1) to develop a robust and user-friendly paleoinfor-
established source for radiolarian information online, with sev-
matic system; 2) to gather and ensure that any data added into
eral thousand requests every week from all over the world.
1 Natural History Museum, University of Oslo, Pb. 1172 Blindern, 0318 Oslo, Norway. Email: [email protected]
2 Virtual One as, Pb. 229, 2021 Skedsrnokorset, Norway. Email: [email protected]
An online micropaleontology database: Radiolaria.org S63

2. Radiolaria.org the people that are (or have been) contributing to Radiolar-
ia.org. It also gives the names and contact information for the
Radiolaria.org includes general pages explaining what radio-
web-editors responsible for the different geographical areas/
larians are, as well as more detailed pages with taxonomic data
time periods. "Radpeople" is a username- and password-pro-
on different species. It can be accessed free of charge, through
tected section providing addresses and e-mails of radiolarian
a standard web browser, and uses hot-links for navigation.
researchers. (If interested, contact the webmaster on Radio-
Most of the information is contained in a relational database,
laria.org). To locate specific information it is possible to type
and the webpages are generated from this content.
in one or more words on the "Search"-page. The user selects
The most important feature of Radiolaria.org is probably
whether it is a search on species, synonym, reference, descrip-
the "species information", and the link between what re-
tion or archive. The result is displayed with hot-links to the rel-
searchers name different radiolarians (or used to name them,
evant pages. Introduction pages like "What are Radiolarians",
i.e. synonyms) and pictures of how the species look like in dif-
"Quaternary and Tertiary radiolarians" written by various au-
ferent areas/periods. Having this information online, where re-
thors, contain general information about radiolarians for
searchers can contribute and discuss species concepts, may be
non/novice researchers.
one step in the right direction in the process of bringing taxon-
omy into conformity, which is very important for both local
and regional biostratigraphic and paleoclimatic work. 2.2. Species information
Information about a radiolarian species can be found in sever-
al ways:
2.1. Home page
The web-interface of Radiolaria.org consists of several sections 1) By scrolling the species list and looking for names (listed
that divide content and functionality into more manageable re- alphabetically) or pictures;
ports and tools. On the home page (Fig. 1) you will find links 2) By navigating via the stratigraphic time scale (adopted
to most pages and sections included in the site. A detailed de- from the International Commission on Stratigraphy) and
scription and screenshot of every page is beyond the scope of geographic links listed on the right-hand side of the home
this paper, we will here rather give an overview and some ex- page (e.g. by clicking on "Holocene (Recent)" and select-
amples of the database features and functionality. ing "Nordic Seas"). When selecting the area of interest, the
Under "About radiolaria.org" you will find general infor- different species are listed under either Nassellaria,
mation about the website, the idea behind it, how the system Spumellaria, or Phaeodaria. The number of species found
works, and a user manual (.pdf) on how to contribute informa- in each category is also indicated;
tion to the system. "Species list" contains names (listed alpha- 3) By using the search-engine.
betically), pictures, and links to all radiolarian species in the
database. The "Archive" section includes (scanned) plates and Under each species it is possible to find the original and/or
captions of important taxonomic publications, which we have other descriptions, images, synonyms, references and web links
got the necessary permissions to reproduce. As many books (if any) to other resources on the internet. There is also a dis-
and journals with original plates and species descriptions are cussion forum where radiolarian researchers can exchange
getting old, out of print, or somehow unavailable, it is of great ideas and opinions, add comments and suggestions, and post
importance to digitize and make these publications available related non-published material. Each species is linked to a ge-
online. "Web links" contains links to other online radiolarian ographic/stratigraphic distribution. The user can navigate be-
resources available on the Internet. "InterRad" informs about tween different pages by clicking on the links in the submenu:
the International Association of Radiolarian Paleontologist The "Images"-link displays a list of photographs for the se-
(InterRad) meetings, membership, key people and the "Rad- lected species. Information about location, sample, latitude,
folks" mailing list. The official "Newsletter" can be down- longitude and photographer is shown next to each picture.
loaded from a password protected page. "News" keeps re- Availability of various specimen pictures from different ocean-
searchers updated on what is happening within the website and ic regions and time intervals may help point out regional/strati-
the radiolarian community in general. Under "Mystery Rad" it graphic variations within the species. This is of importance, e.g.
is possible to post a picture and information about an unknown in comparative morphological analysis.
species and let the community share their opinions and sug- The "Synonyms" displays a list of registered synonyms for
gestions as to which species it may be. "Radiolarian art" is a the species, sorted in ascending chronological order. The full
page that links science and art. At present (October 2005) it reference to each synonym is found when selecting "Refer-
contains pictures of eleven radiolarian models made in white ences". This online facility may give the user a more updated
clay by the Swedish artist Eva Bjerke, several pictures of a and complete overview of available information relating to a
large quilt made by the Canadian artist Barbara J. West, and species compared to the consultation of single publications.
pictures of three plexiglass models of living radiolarians made All areas/periods where the species has been registered are
by the German artist Dagmar Borgwart. "Contributors" lists found under "Distribution". (As this is work in progress, not
S64 J. Dolven & H.A. Skjerpen

r ,
Rad io laria .or g
I I Radio laria .org is an online database containing infonnat ion about
radio larians - fossil and recent. \\ IIh image ~. descnpnons, references. T r '
" syno nyms and links 10 other online resour ces . more ...
",'
101 K
Radiolaria 31'"C' holop lankt omc protozoa I·s.a ponodI- •
widely distributed in the oceans. The )' occur
througho ut the water column from near 1< 110
surface to hundred s ofmeters depth . As with
many planktonic organi sms . their abun dance 1<
in a geographical region i~ related to q uality
R l'<''l'l of the water mass, including such ..-ariablcs
"h as temperature, salinity. productivity. ami
available nut rie nts. more ...
&til.obrian Art
Radiolarians have been an inspiration fo r many artists . These pag'-""S
show examples ofthcir works . more ..,
Artist: Art ist :

lIarbara W"", Eva Bje rke
(Canada) (Sweden)
Norge<fonkningsrld
Fig. 1. The home page of Radiolaria.org (October 2(05) .
all areas/periods may have yet been included.) The presence of the page. This will tak e the contributor through severa l
a species should be documented by a picture and preferably a steps (pages), wher e different forms are filled out. In form
referenc e to a book or paper. Th e name of the person who one (Species and origin al description) and two (Other de-
contributed the information is indicated in parenthesis. scription), the genus and species name along with at least
The display of species names, descriptions, synonyms and one description (including author and year) is entered.
pictures together has proven very impo rtant in making the Synonym(s) and Reference(s) are inserted in form thr ee,
species concept more homogenous betw een users, as scientists and Image(s) with related information is added in form
use differ ent taxonomies and will therefore give different four. All fields marked with an asterisk (*) are required
name s to the same specimen. Un fortunately, Radiotaria.org field s that need to be filled in order to submit the data into
does not prevent the possibility of the same species being reg- the system. Once the data entry is completed, the user can
istered und er two different nam es in two geographical areas/ preview the contributed information. After approval, the
peri ods. data is submitte d along with the contributor's nam e and e-
mail address. Th e contributor is also suggested to notify the
web-editors bye-mail about the new additions.
2.3. Contributing information
2) A user may also add single pieces of information to a
One important aspect of Radiolaria.org is that any radiolarian species that is already registered in the system. This is done
researcher can add information (provide d it meets certain cri- by selecting the species of interest, locating the "Add "-link
teria, see section 2.4.) into the dat abase. This can be done in on top of each page in the submenu (e.g. Add description ,
severa l ways: Add image , Add synonym/refere nce) , filling out form es),
and submitt ing the information .
1) A contributor can add a species by selecting the stra ti-
graphic /geographic location wher e the species is known to When data are submitte d into the system (either way de-
be pre sent and click on the "Add species" link on top of scribed above), it is not instantly made visible on th e site, but
An online micropal eont ology database: Radiolaria.org S65

placed in an administration area and flagged for review. The • Support for a wider variety of data, particularly quantita-
review process is described in more detail under the following tive radiolarian data (including sample information like lo-
section. cation, geographic coordinates, age etc.). This data could
then be used further, e.g. in a Geographical Information
System, to plot species distribution maps.
2.4. Administration and Quality control
• A Software Archive with links to useful applications for ra-
The goal of the Radiolaria.org-project is to gather high-quality diolarian researchers (e.g. data analysis packages including
information about radiolarians from all time periods and make common statistical, plotting and modeling functions).
this available online. One way to acquire data of good quality • A Taxonomic Identification Tool that displays species ac-
is to use information published mainly in peer-reviewed books, cording to selected characteristics (e.g. number of spheres/
journals and electronic publications. In Radiolaria.org, the segments, number of spines, pore size/shape). Each species
content of species names, descriptions, synonyms and refer- is registered with a certain number of characteristics, allow-
ences is based solely on published data. ing searches and drill-downs on taxonomic keys.
Copyright laws affects online information, as this is a form
of publishing. Ownership and different use-rights for data can Radiolaria.org belongs to the radiolarian community and the
be a source of conflicts, especially regarding already published people that have invested their time and expertise building it.
material. We are aware of copyright issues and are working on While the system is currently hosted on a private server, it will
solving them. be transferred to a research institution (doing radiolarian re-
To further ensure the quality of the submitted data, web- search) for further development and hosting - preferably long
editors are allowed to log on to the system and check submit- term. This transfer will take place after Radiolaria.org has
ted material flagged for review. A web-editor is assigned a cer- reached a defined goal of content (e.g. when the Recent radio-
tain geographic or stratigraphic area, according to his/her ex- larians catalogue is completed). The new hosting research in-
pertise, and reviews all information related to this area. The stitution must be committed to run Radiolaria.org on the fol-
web-editors ensure that: 1) the information submitted is (with lowing conditions:
regards to species descriptions, synonyms and references) • Keep Radiolaria.org freely accessible for all.
based on published material with correct citations; and 2) the • Maintain the system, and preferably develop it further.
images, distribution and web-links are correctly assigned. If a • Agree to hand the system over to a new institution, should
web-editor finds errors, judges the data entry to be insufficient the institution not be able to comply with the above condi-
or wrong, or needs to contact and discuss with the contributor, tions.
he/she can use the name and email that was submitted with the
content. The review process (checking of submitted material Radiolaria.org was built for the radiolarian community, and
before it is made available online on the website) also prevents will exist as long as this community shares the work and the
intentional sabotage by online pranksters. benefits in our quest for better science.
Acknowledgement
3. Future and goals
Many thanks to Dr. Giuseppe Cortese for reviewing the paper and providing
A complete online catalogue with high quality information re- useful comments and suggestions. We also like to acknowledge Dr. Norman
quires different people with different expertise to donate many MacLeod and Prof. Kjell Bjorklund for reading and correcting an earlier ver-
hours of work. While some work in Radiolaria.org is spon- sion of this manuscript. Radiolaria.org is from 2003-2005 funded by the Nor-
wegian Research Council (Project number: 157834/432).
sored by grants, most is based on voluntary work without mon-
etary compensation. An important part of reaching the goals
of Radiolaria.org is to recruit area/period-experts willing to REFERENCES
contribute with their time and expertise. Although most radio-
MACLEOD, N. & GURALNICK, R, 2000: Paleoinformatics. In: LANE, RH.,
larian researcher already have a fully booked schedule, we
STEININGER, F., KAESLER, RL., ZIEGLER, W., LIPPS, J.H. (Eds.): Fossils
hope that many will see the importance of this project and will and the future; paleontology in the 21st century. Senckenberg-Buch, 74:
be able to provide help to build up the database in the future. 31-36.
Information systems evolve over time, with new functional- MACLEOD, N., DIVER, P., GURALNICK, R, LAZARUS, D. & MALMGREN, B.,
2000: Computers, quantification, and databases. In: LANE, RH.,
ity ever adapting to new users and needs. What new features
STEININGER, F., KAESLER, RL., ZIEGLER, W., LIPPS, J.H. (Eds.): Fossils
will be requested by the radiolarian community is hard to pre- and the future; paleontology in the 21st century. Senckenberg-Buch, 74:
dict, but some of the currently planned features for Radiolar- 191-201.
ia.org include:
Manuscript received April 2004
Revision accepted February 2005
S66 J. Dolven & H.A. Skjerpen

0012-9402/06/OlS067-12 Eclogae geol. Helv. 99 (2006) Supplement 1, S67-S78
DOl 10.1007/s00015-006-0608-z
A new genus of Entactiniidae (Radiolaria) from the Upper Permian

of South China
QINGLAU FENGl, YOUYAN MENG, WEIHONG HE & SONGZHU Gu
Key words: Taxonomy, Radiolarian, Late Permian, South China
ABSTRACT
Four new species and one indeterminate species of radiolarians from the nal pore frames composed of thick but narrow bars, which is very similar in
Changxiangian (Upper Permian) of southern Guangxi, South China are intro- morphology to some genera within the Mesozoic subfamily Pantanelliinae
duced. They all belong to one new genus: Megaporus n. gen. This new genus is PESSAGNO (1977), but differs from the latter in possessing initial spicules with-
characterized by two concentric spherical shells with pentagonal and hexago- in medullary shell.
divided in ascending order into three formations: the Heshan,

Introduction
Changxing and Dalong formations (Fig. 1). All specimens de-
The studies on the mass extinction of Permian radiolarians in- scribed in the present paper were collected from the Dongpan
dicated that the Changxingian was a main extinction interval Section near Dongpan Village (22°16.196'N, 107°41.505'E). In
for the Permian radiolarian fauna and the main extinction this section, the Dalong Formation overlies Changxingian
event was a gradual process (Kakuwa 1996; Vishnevskaya limestone of the Changxing Formation and is overlain by mud-
1997; Kuwahara & Yao 1998). Our investigations in South stones of the Lower Triassic Luolou Formation containing
China, however, prove that some taxa, such as Entactinaria Ophiceras tingi TrEN and Claraia dieneri NAKAZAWA. There-
(Kozur & Mostler 1982), had high diversity during the late fore, the Dalong Formation is upper Changxingian. The lower
Changxingian, and then experienced rapid extinction at the part of the Dalong Formation is mainly composed of mud-
end of the Changxingian. In the high diversity stage, some new stones and siliceous rocks; the middle part, of bedded siliceous
taxonomic units developed. One of these units, described here- rocks with shales; and the upper part, of siliceous mudstones,
in as Megaporus n. gen., is characterized by having two con- bentonites and muddy siliceous rocks with Permian ammo-
centric spherical shells with large pores, which is very similar to nites, bivalves and brachiopods, for example: Huananoceras ct.
some genera of the Mesozoic subfamily Pantanelliinae PES- perornatum CHAO & LIANG, Laibinoceras ct. compressum
SAGNO (1977) in morphology, but differs from the subfamily in YANG, Qiangjiangoceras sp., Euchondria jiangxianensis Gu &
possessing initial spicules within the medullary shell. LIU, Euchondria dalongensis YIN, Leptochondria intermedia
YIN, Paracrurithyris pigmae (LIAO), Martinia sp., Spinomar-
ginifera and so on (Fig. 2). Abundant well-preserved radiolari-
Stratigraphy
ans were obtained from the bedded siliceous rocks, including
The fauna described herein was collected from an area located Neoalbaillella optima ISHIGA, KITO & IMoTo, Albaillella trian-
between Dongmen and Liuqiao Towns, Fusui County, gularis ISHIGA, KITO & IMOTO, Albaillella yaoi KUWAHARA,
Guangxi Zhuang Autonomous Region (Fig. 1) and the strata and so on together with the radiolarians described in this
in this region mainly consist of Upper Permian and Lower Tri- paper, which belong to the upper Changxingian Neoalbaillella
assic successions (BGMRGZAR 2001). The Upper Permian is optima Zone (Kuwahara et al. 1998; Yao et al. 2001) (Fig. 2).
I Faculty of Earth Sciences, China University of Geosciences, Wuhan 430074, China. E-mail: [email protected]
A new genus of Late Permian Entactiniidae 67

B
Chongzuo
o
Fig. 1. A and B: Locality map of the studied area;

C: Geologic map of the studied area and locality
of the studied section.
Systematic Paleontology
Discussion. - The structure of this new genus resembles that
All specimens described in this paper are deposited in Muse- of Entactinosphaera FOREMAN (1963) from which it differs by
um of China University of Geosciences, Wuhan, People's the presence of very large pores on the cortical shell, and by ir-
Republic China. regular arrangement of the primary pines on the cortical shell.
It is also similar to some genera of Pantanelliinae PESSAGNO
Class ACTINOPODA (1977), such as Cana MEKIK (2000), Gorgansium PESSAGNO &
Subclass RADIOLARIA BLOME (1980), and Pantanellium PESSAGNO (1977), but differs
Superorder POLYCYSTIDA EHRENBERG 1838, emend. from the latter by having initial spicule in the medullary shell.
RIEDEL1967 The new genus is assigned to the Entactiniidae RIEDEL (1967)
Order ENTACTINARIA KOZUR & MOSTLER 1982 because it has two latticed shells and an initial spicule.
Family Entactiniidae RIEDEL1967 Etymology. - mega (Latin, adj.) = large; porus (Latin, adj.)
= pore.
Type genus. - Entactinia RIEDEL1967 Occurrence. - Upper Permian, Dalong Formation in south-
ern Guangxi, China.
Diagnosis. - Spherical to subspherical Entactinaria with com-
monly six to eght-spined initial spicule. Shell latticed, with one Megaporusjini FENGnew species
or more shells (De Wever et al. 2001). Plate 1, Figures 1-19; Plate 2, Figures 1-3, 8, 9, 12
Occurrence. - Ordovician to Lower Jurassic. ?Cryptostephanidium sp., SHANG, CARIDROIT & WANG 2001,
pI. 4, fig. 20.
Genus Megaporus FENGnew genus
Type species. - Megaporus jini FENGn. sp. Diagnosis. - A species of Megaporus with five to six three-blad-
ed primary spines irregularly arranged on the cortical shell.
Description. - Test small, with two concentric shells: cortical Description. - Test small, spherical, and with five to six ap-
and medullary shells. Cortical shell spherical, and composed proximately equal three-bladed primary spines. Spines gradually
of pentagonal and hexagonal pore frames with two to six decrease in width distally. They are irregularly distributed on
three-bladed primary spines; pore frames thick and narrow; the cortical shell, the angle between two close spines being vari-
pores very large. Medullary shell small, consisting of pentago- able and not always at right-angle. Cortical shell is composed of
nal, hexagonal and irregular pore frames, and connected with pentagonal and hexagonal pore frames with massive but short
cortical shell by some three-bladed and cylindrical beams. secondary spines at the pore frame vertices besides the primary
There is an initial spicule inside the medullary shell, and the spines; pore frame thick and narrow; pores very large, about
spicule is relatively strong compared to the small medullary seven to ten pores present on outer surface of hemisphere.
shell. Medullary shell small, consisting of pentagonal, hexagonal and
S68 Q. Feng et al.

48-87 (average 72, holotype 70), diameter of pore of cortical
'J;
:;
...
...:
1)
.s: Columnar sec t io n and shell 19-32 (average 26, holotype 29); diameter of medullary
';: L.:..
:; E shell 25-27 (average 26, holotype 28), width of pores on
tr: '-' sa mp le numbers
:E medullary shell 5-9; width of primary spines near base 12-25
u (average 20, holotype 21); length of primary spines 48-106 (av-
'" ... erage 80, holotype 56) .
'" L.:..:::
t'::
';:
0 ..................... , Occurrence. - This species is only known from the middle
f-
.... part of the Dalong Formation (upper Changxingian) in the
I1J
~
0
-
-l
0
C la rai o dieneri
Dongpan Section, southern Guangxi, China.
-l Discussion. - This species is similar to Cana elegans MEKIK
_._. ._. Ophiceras lingi (2000) from the Lower Cretaceous of Northwest Turkey in the
..... _-------- Hua nanoce ra s
-- - characteristics of the cortical shell, but differs from the latter in
------- - -
1:; --..... - - - - - 1-<11 binocer as
---_ .- ._.__ - Nuculopsis
_---------_.
Euchondrta
.. having approximately equal primary spines irregularly distrib-
c,
c,
:J
_-_._._....-- Martinia
---_._._
..
------------
-------
uted on the cortical shell and an initial spicule.
- - - Spi nomargi ntfera
---- - - - - - Paracrurtthyris
-...........
,;:----- Megaporus yini FENG new species
- - - ......
.•••••.•....
----
---
_----------
--- ..
Plate 2, Figures 4-7, 10, 11; Plate 3, Figures 16-20; Plate 4,
(, - )
----- Figures 1-9
---
-------
6· ·1
0-5
e -- Diagnosis. - A species of Megaporus with three unequal,

-
..0...
-- coplanar primary spines and well-developed nodes.
""
E
:;
- Description. - Test is small, with two spherical concentric
C :;: --
t::
-0 ----
:-ct
~
r- L.:.. - -
-- shells: cortical and medullary shells. Cortical shell composed of
;:: en ~
I1J
i::
-- --
-------
pentagonal and hexagonal pore frames with well-developed
0-
....
:;
a.
- 0
o"" --- --
nodes at the pore frame vertices. Thickness of bars on the pore
frames about 3 times their width. Pore very large, about six to
a. nine pores present on out surface of hemisphere. Medullary
~ ----- --
--- - shell with polygonal pore frames, and connected with the
-- - [1 nodes on the cortical shell by several beams. Three coplanar
primary spines are similar in configuration: three-bladed and
------::..:"'_--
------- gradually decreasing in width toward distal end, but unequal in
-------_
-------... § S hale
length and asymmetrical in arrangement: two spines (basal
--------
--------.
- - - - - - - t:."':.1 M udst o
-----::-::-:::-::-=~
TIL' spines) nearly equal in length and closer together, often short-
....:; - --
- - - - - - § Bent onite er than third spine (polar spine). There are initial spicules in-
--, -- ----------,.;
-
-------- B ~i~I~$et~~l:
:::0 -------- side the medullary shell.
...: --------
--------
L.:.. --------
-l -------- Etymology. - This species is named for Prof. Hongfu Yin of
en -------- ~ : : : :I :i~~~~:I1~
i:: --
---- -----
---- China University of Geosciences (Wuhan) in honor of his
X
~ rf:f:f:f:f=t:Ei ~ ~:l~~::u~
S ili ceous
rock many contributions to the study of the Permian-Triassic
~ boundary.
~
:<:
..=:
------
----_ . rock
U (:g Lim estone Type. - Holotype, Plate 4, Figure 1, sample C6-5, catalog
number X0301-15.
Fig, 2. Columnar section showing the stratigraphic position of the radiolarian Measurementsiutiu. - Based on 16 specimens. Length of
fauna described in this paper. polar spine 37-63 (average 52, holotype 38), width of polar
spine near base 11-24 (average 17, holotype 11); length of basal
spine 35-53 (average 42, holotype 35), width of basal spine
irregular pore frames, and connected with cortical shell by some near base 11-23 (average 16, holotype 11); diameter of cortical
three-bladed and cylindrical beams. There are initial spicules in- shell 57-88 (average 71, holotype 57); diameter of pore of cor-
side the medullary shell, but the number of spicules is unclear. tical shell 19-35 (average 29, holotype 19).
Etymology. - This species is named for Prof. Yugan Jin of Occurrence. - This species is only collected from the mid-
the Nanjing Institute of Geology and Palaeontology in honor of dle part of the Dalong Formation (upper Changxingian) in the
his many contributions to the study of the Permian stratigraphy. Dongpan Section, southern Guangxi, China.
Type. - Holotype, Plate 1 Figure 14, sample C6-3, catalog Discussion. - This species resembles some species of the
number X0301-14. Mesozoic Gorgansium PESSAGNO & BLOME (1980) in outline,
Measurements (urn). - Based on 24 specimens, of which but differs from the latter in having a smaller test, larger pores
only a few possess a medullary shell. Diameter of cortical shell on cortical shell and an initial spicule.
A new genus of Late Permian Entactiniidae S69

Megaporus yangi FENG new species rounded longitudinal ridges alternating with three wide longitu-
Plat e 3, Figures 1--6, 12-14 dinal grooves, and gradually decreasing in width at distal direc-
tion, but they are unequal in length and asymmetrical in arrange-
Diagnosis. - Megaporus with two unequal polar spines and ment: long primary spine is nearly two times the length of the
well-developed nodes on cortical shell. short spine, and minimum angle between them about 125°.
Description. - Test small, with two concentric shells: corti- Etymology. - Unicus-a-um (Latin, adj.) = unique.
cal and medullary shells. Cortical shell spherical, composed of Type. - Holotype, Plate 3, Figure 7, sample C6-5, catalog
pentagonal and hexagonal pore frames (predominantly hexag- number X0301-17.
onal) with well-developed nodes at the pore frame vertices; Measurements (urn) . - Based on three specimens. Length
nodes long, massive , and triradiate. Thickness of bars on the of long polar spine 57-83 (average 70, holotype 83), width of
pore frames approximately four times their width. Pore very long polar spine near base 23-30 (average 28, holotype 30);
large , about seven to ten pores pr esent on outer surface of length of short polar spine 43-52 (average 48, holotype 43),
hemisphere. Two polar spines are similar in configuration, but width of short polar spine near base 18-27 (average 22, holo-
unequal in length and obliquely opposed to each other. Both type 20); diameter of cortical shell 80-90 (average 85, holotype
spines are made up of three narrow, rounded longitudinal 83); minimum angle between both polar spines 105°-140° (av-
ridges alternating with three wide longitudinal grooves, and erage 125°, holotype 103°).
gradually decrease in width at distal direction. Medullary shell Occurrence. - This species is only collected from the mid-
with polygonal pore frames, and connected with the nodes on dle part of the Dalong Formation (upper Changxingian) in the
the cortical shell by several triradiate and cylindrical beams. Dongpan Section, southern Guangxi, China.
There is an initial spicule inside medullary shell. Discussion. - This species is similar to Denize magnifica
Etymology. - This species is named for Prof. Qun Yang of MEKIK (2000) from the Lower Cretaceous of northwestern
Nanjing Institute of Geology and Palaeontology in honor of Turkey in morphology, but different from the latter in having
his many contributions to the study of the Jurassic Radiolaria. an obtuse angle between both polar spines and well-developed
Typ e. - Holotype, Plate 3, Figure 1, sample C6-3, catalog nodes. It is distinguished with Megaporus yangi n. sp . by
number X0301-16. strongly asymmetrical arrangement of both polar spines.
Measurements (urn). - Based on 9 specimens. Length of
long polar spine 53-83 (average 75, holotype 77), width of long Megaporus sp.
polar spine near base 19-37 (average 28, holotype 30); length Plate 2, Figure 13; Plate 3, Figures 10, 11
of short polar spine 46-57 (average 51, holotype 50), width of
short polar spine near base 28-40 (average 32, holotype 30); di- Description. - Test small, with two concentric shells: cortical
ameter of cortical shell 64-79 (average 71, holotype 71). and medullary shells. Cortical shell spherical, composed of pen-
Occurrence. - This species is only collected from the mid- tagonal and hexagonal pore frames (predominantly pentago-
dle part of the Dalong Formation (upper Changxingian) in the nal) with poorly developed nodes at the pore frame vertices .
Dongpan Section, southern Guangxi, China. Thickness of bars on the pore frames approximately three times
Discussion. - The morphology of this species is similar to their width. Pore very large and about seven to nine pores pre-
some species of the genus Pantanellium PESSAGNO (1977) from sent on outer surface of hemisphere. Two polar spines are simi-
Upper Triassic-Lower Cretaceous in morphology, but differs lar in configuration, but unequal in length and obliquely op-
from the latter in having an initial spicule inside medullary posed to each other. They are with three narrow, rounded lon-
shell and longer nodes on the cortical shell. gitudinal ridges alternating with three wide longitudinal
grooves, and gradually decreasing in width at distal direction.
Megaporus unicum FENG new species Occurrence. - Changxingian, the Dalong Formation in
Plate 3, Figures 7-9 southern Guangxi, China.
Discussion. - It differs from Megaporus yangi n. sp. and
Diagnosis. - Megaporus with two unequal polar spines be- Megaporus unicum n. sp. in having poorly developed nodes at
tween which the minimum angle is 103°-140 °. the pore frame vertices.
Description. - Test small with two concentric shells: cortical
and medullary shells. Cortical shell spherical, composed of pen-
tagonal and hexagonal pore frames (predominantly pentagonal)
with well-developed nodes at the pore frame vertices; nodes Acknowledgments
long, massive, and triradiate. Thickness of bars on the pore This study has been supported by NSFC (Project No. 49972002. 40232025).
frames about three times their width. Pore very large, about eight We wish to express oursincere thanks to Prof. Qun Yang and Dr.Hui Luo of
pores present on outer surface of hemisphere. Medullary shell Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sci-
ences, Prof. Haoruo Wu of Institute of Geology and Geophysics, Chinese
small, with polygonal pore frames. and connected with the nodes Academy ofSciences, and Prof. Jonathan A ichison oftheUniversity ofHong
on the cortical shell by several triradiate and cylindrical beams. Kong for their valuable suggestions. We aregrateful to Prof. Dr. Helmcke and
Two polar spines are similar in configuration: consisting of three Dr. R. Ingavat-Helmcke for their help during thecourse ofthestudy.
S70 Q. Feng et al.

REFEREN CES Pessagno, E.A . Jr. 1977: Lo wer Cre ta ceo us radi olarian biostra tigra phy of the
G rea t Va lley Sequen ce and Fran ciscan Co mplex, Cali forn ia Coast
BUREAU OF G EOLOGY AND MINERAL R ESOURCES OF G UANGX I ZHUANG Ran ges. Co ntributions to th e Cushma n Fo unda tio n of For amin ifer al Re-
A UTONOMOUS REGION, 2001: 1:50,000 Liuq iao geologic map and explana- sea rch, Special Pu blicat ion 15, 1-8 7.
tor y no tes (in Chinese). PESSAGNO, E.A . JR.. & BLOME, C D . 1980: Up per Triassic and Jurassic Pan -
DE WEVER£"P., D UMITRIC, P.. CAULET, J.P., NIGRIN I, C & CARIDROIT, M. tane llidae from California, O rego n and British Co lumbia. Micro pale on-
2001: Radiolarian s in th e sed imen ta ry reco rd. 178-180 p., Gordon and tolo gy 26 (3), 225-273.
Breach Science Publ ish er s. RIEDEL, W.R. 1967: Subclass Ac ti no poda. In: H ARLAND, W.E. et a!' (Ed s.):
E HRENBERG, CG. 1838: U ber die Bildu ng de r Kreidefelsen und des Kr eid e- Prot ozoa. th e Fossil Record , 291-298. Geological Society of Lo ndon Pu b-
mer gels dur ch unsichtb are O rga nisme n. Ab ha nd lungen der Koeni glischen licati on.
A kademie der Wissen schaften , zu Berl in, Jahrgan g 1838, 59-147. SHANG, Q.H., CARIDROIT, M. & WANG Y.J . 2001: Rad iol ar ian s fro m th e
FOREMAN, H.P. 1963: U ppe r Devon ian Radiolaria fro m th e H uron member of Uppe rmos t Permian Cha ngxingia n of so uthern Gu ang xi. Ac ta Micro-
the O hio sha le. Micro pa leonto logy 9, 267- 304. palaeont ologica Sinica 18(3), 229- 240.
KAKUWA, K. 1996: Corre lation betw een the bedded chert seq uence of so uth- VISHNEVSKAYA, V. 1997: De vel opm en t of Paleozoic-Mesozoic Rad iolari a in
west J apan and BfuC exc ursion of ca rbo nte seq uence , and its significa nce the Northwes te rn Pacific Rim. Marine Microp aleontology 30, 79-95.
to the Per mia n-Tr iassic mass extinct ion . The Island Ar c 5, 194-202. YAO,J., YAD, A. & KUWAH ARA, K. 2001: Up per Permian biostr ati grap gic cor -
KOZUR, H . & MOSTLER, H . 1982: E ntac tinian suborde r, a new rad iolarian re lation between con od on t and rad iolar ian zones in the Tamb a-Min o Ter-
subo rder. Geologische-Palaeonto logische Mitt eilungen, Innsbruck 11. ra ne , Southwest Japan. J ou rn al of Ge osciences, Osaka City U nive rsity
399-414. 44(5) ,97-119.
KUWAHARA, K. & YAO, A. 1998: Div er sity of Late Permian radiolarian as-
se mblage s. News of Osaka Mic ro paleon tologist s, Special Volume I I,
33-46. (in Japanese with English abs tract)
KUWAH ARA, K., YAO, A. & YAMAKITA, S. 1998: Reexamination of Upper Per -
mian radiolarian bios tratigraph y. Ea rth Scienc e 52(5) , 391-404. Manu script received January 2004
Mekik, F.A . 2000: Earl y Cretaceo us Pantan ell iidae ( Ra diolaria) from No rth- Revisio n acce pted Mar ch 2005
west T urkey . Micro paleo nto logy. 46 (1) , 1-30.
A new genus of Lat e Permian Entactiniidae S71

Plate 1
Scanning electro n photomicrographs of the Changxingian radi olarians from the Dalon g Fo rma tion in southe rn G ua ngxi, Southw China (1- 14, 16, 17: bar =50!im;
15,1 8: bar= 20!im; 19: bar=lO!im ). 1-1 9 M egaporu s jini n. sp.; I . C6-411698; 2, C6-411 693; 3, C6-511723,; 4, C6-4/1696; 5, C6·4/1699; 6, C6-5/1722; 7, C6-5/1718; 8,
C6-4/1710; 9, C6-4/I 712; 10, C6-4/1700; I I, C6·511725; 12, C6-5 /2055; 13, C6-511727; 14, C6-3/ 1140, holot ype; 15, C6·3/1255, magn ification of holotype, sho wing
medull ar y shell; 16, C6-4/1716; 17, C6-4/1695; 18, C6-5/1717, magnifi cation of figur e 16, show ing medullary shell and initia l spicules in the medullary shell; 19,
C6·511 739, magnification of figure 16, sho wing initial spicules in the medullary shell.
S72 Q . Feng et al.

Plate 2
Scanning electron photomicrographs of the Changxingian radiolarians from the Dalong Formation in southern Guangxi, South China (1-3: bare lum; 4-8:
bare ltlum; 9-13: bar=50Jlm). 1-3,8,9,12 Megaporus jini n. sp.; 1-3, magnification of plate 1 figure 16, showing initial spicules in the medullary shell; 8, magnifi-
cation of the figure 12, showing initial spicules in the medullary shell; 9, C6-5/3116;12, C6-5/2500.4-7, 10, 11, Megaporus yini n. sp.; 4 and 5, magnification of the
figure 10, showing initial spicules in the medullary shell; 6 and 7, magnification of the figure 11, showing initial spicules in the medullary shell; 10, C6-5/3112; 11,
C6-5/2498. 13, Megaporus sp., C6-5/3105.
S74 Q. Feng et al.

Plate 3
Scanning electron photomicrographs of the Changxingian radiolarians from the Dalong Formation in southern Guangxi, South China (1-14, 16-20: bar=50llm; 15:
bar=I00llm). 1--6,12-14 Megaporus yangi n. sp.; 1, C6-3/1142, holotype; 2, C6-3/1145; 3, C6-5/1730; 4, C6-5/2057; 5, C6-5/1726; 6, C6-5/1737; 12, C6-3/1059, show-
ing thickness of cortical shell, and beams between cortical and medullary shells; 13, C6-3/1252, magnification of figure 2, showing medullary shell; 14, C6-3/1254,
magnification of the holotype, showing medullary shell. 7-9 Megaporus unicum n. sp.; 7, C6-5/2060, holotype; 8, C6-4/1686; 9, C6-3/1238. 10, 11 Megaporus sp.; 10,
C6-3/1127; 11, C6-5/2058. 15, Neoalbaillella optima Ishiga, Kito and Imoto, C6-3/1093. 16-20 Megaporus yini n. sp.; 16, C6-5/1728; 17, C6-4/1706; 18, C6-5/1720; 19,
C6-4/1713; 20, C6-4/1714.
S76 Q. Feng et al.

Plate 4
Scanning electron photomicrographs of the Changxingian radiolarians from the Dalong Formation in southern Guangxi, South China (1-5,7-9: bar=50/lm;
6: bar=20/lm). 1-9 Megaporus yini n. sp.; 1, C6-5/1732, holotype; 2, C6-5/1735;3, C6-4/171l; 4, C6-5/1731;5, C6-5/1729;6, C6-5/1738,magnification of the holotype,
showing medullary shell; 7, C6-311143; 8, C6-5/1736; 9, C6-5/1734.
S78 Q. Feng et al.

001 1O.1007/s00015-006-0604-3
Radiolarian faunal turnover through the Paleocene-Eocene transition,

Mead Stream, New Zealand
CHRISTOPHER 1. HOLLIS I
Key words: South Pacific, Paleogene. Radiolaria, diatoms, paleoecology, global change
ABSTRACT ZUSAMMENFASSUNG
The Mead Stream section, northern Clarence Valley, is the most complete Pa- Das Mead Stream Profil, nordliches Clarence Tal, ist die vollstandigste Palao-
leocene-early Eocene record of pelagic sedimentation in the mid-latitude zan-Fruh Eozan Aufzeichnung pelagischer Sedimentation in mittleren Breiten
(_55 0 S paleolatitude) Pacific Ocean. Integrated studies of sediments, siliceous (_55 0 S Palaobreite) des Pazifischen Ozeans. Integrierte Studien von Sedi-
and calcareous microfossils and carbon isotopes have shown that major global men ten, kieseligen und kalkigen Mikrofossilien und Kohlenstoffisotopen
climate events are recorded by distinct changes in lithofacies and biofacies. haben gezeigt, dass bedeutende globale Klimaereignisse durch deutliche Ver-
The consistent and often abundant occurrence of siliceous microfossils in the anderungen in Litho- und Biofazies gekennzeichnet sind. Das bestandige und
section provides a rare opportunity to undertake quantitative analysis of high- oft sehr haufige Vorkommen kieseliger Mikrofossilien im Profil bietet eine
latitude radiolarian population changes through the late Paleocene and early scltene Gelegenheit, quantitative Studien von Veranderungen innerhalb von
Eocene. Late Paleocene assemblages are dominated by spumellarians, al- Radiolarienpopulationen hoher Breiten wah rend des spaten Palaozans und
though the nassellarian species Buryella tetradica is the most abundant species. friihen Eozans durchzufiihren. Spatpalaozane Vergesellschaftungen werden
The Paleocene-Eocene boundary (= base of Paleocene-Eocene thermal maxi- von Spumellarien dominiert, obwohl die Nassellarienart Buryella tetradica die
mum) in the Mead Stream section is marked by major faunal turnover, includ- haufigste Art ist. Die Palaozan-Eozan Grenze (=Basis des Palaozan-Eozan
ing an abrupt decrease in B. tetradica. first occurrences of several low-latitude Warrncmaximums) im Mead Stream Profil ist durch bedeutende faunistische
species (e.g. Amphicraspedum pro/ixum s.s.. Lychnocanium auxilla, Podocyrtis Erneuerungen gekennzeichnet, einschliel3lich der abrupten Abnahme von B.
papalis, Phormocyrtis turgida, Theocorys'? phyzella) and increased abundance tetradica. dem ersten Auftreten zahlrcicher Arten niederer Breiten (z. B. Am-
of large, robust spumellarians relative to small actinommids. Above an 18-m phicraspedum prolixum S.S .• Lychnocanium auxilla, Podocyrtis papa/is, Phor-
thick, lowermost Eocene interval in which radiolarians are abundant to com- mocyrtis turgida, Theocorys'? phyzella) und der zunehmenden Haufigkeit von
mon, radiolarian abundance declines progressively, falling to <10 individuals groBen, robusten Spumellarien im Verhaltnis zu kleinen Actinommida. Ober-
per gram in the marl-dominated unit that is correlated with the early Eocene halb eines 18 Meter machtigen Intervals des untersten Eozans, in dem Radio-
climatic optimum. These trends in siliceous microfossil populations signal larien sehr zahlreich bis haufig sind, nimmt die Radiolarienhaufigkeit progres-
major changes in watermass characteristics along the northeastern New sive ab, bis auf eine Anzahl von weniger als zehn Individuen pro Gramm
Zealand margin in the earliest Eocene. Assemblages typical of cool, eutroph- innerhalb der Mergel-dominierten Einheit, die dem fruheozanen Klimaopti-
ic, watermasses that dominated the Marlborough Paleocene were replaced in mum entspricht. Diese Trends innerhalb von Populationen kieseliger Mikro-
the early Eocene by assemblages more characteristic of oligotrophic, strati- fossilien signalisieren bedeutende Vcranderungen in Wasserrnassencharakte-
fied, subtropical-tropical watermasses. ristika entlang des nordostlichen neuseelandischen Kontinentalrandes im fru-
hcsten Eozan. Vergesellschaftungen typisch fiir kiihle, eutrophische Wasser-
massen, die das Marlborough Palaozan dominierten, wurden im fruhen Eozan
durch Vergesellschaftungen ersetzt, die eher fiir oligotrophische, stratifizierte,
subtropische-tropische Wassermassen charakteristisch sind.
1. Introduction
Radiolaria are the most diverse and widely distributed group sedimentary successions have the potential to reveal much
of marine plankton preserved in the fossil record (Lipps 1993; about biosphere-geosphere interactions in the marine realm
De Wever et al. 2001). High evolutionary turnover and consid- during critical episodes in Earth history. Whereas quantitative
erable geographic variation in populations make radiolarian studies are common for late Cenozoic radiolarians (e.g.
fossils invaluable research tools in biostratigraphy and paleo- Boltovskoy 1987; Caulet et al. 1992), very few such studies
ceanography. Consequently, detailed census studies through have been made of early Cenozoic or Mesozoic assemblages.
1 Institute of Geological and Nuclear Sciences, PO Box 30-368, Lower Hutt, New Zealand. Email: [email protected]
Paleocene-Eocene radiolarians, Mead Stream S79

~ .., ii:'
~ !!!. N
~
i N
~
~ I
f-
Lithostratigraphy e
0 13 [bulk carb]
013 e
N
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"
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,~..-..
~
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.,
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~
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m ~~"'")1flH ' 50 ....
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0>
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56
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.., ~
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a.
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~ a. Waipawa formation
s
100 ."l 58
Fig. 1. Simplified geological map of eastern Marlborough (after Crampton et
al. 2003) showing location of Mead Stream and other stratigraphic sections 59
mentioned in the text.
c:
o 60
~
50 "c E
j., ~ Lower
" -n
61
_.
a. J: Cher1
Notable exceptions are studies of Cretaceous-Tertiary bound- Rock symbols
ary radiolarians in New Zealand and Ecuador (Hollis 1996; W

.,
>.
"C
't:l
'"
Q)
Mart ~
o 1 2
62
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Keller et al. 1997; Hollis et al. 2003a, b). In this article we re- '"'a.0:: ...
a.
li mes tone per mil PBC
z lime~ne ." ...... • .ramI

RP1-1 KIT boundary
ZBc;1los et I,
port the results of the first quantitative study of radiolarian as- """"one Rlbtlon et*'l
(200 1)
semblage variation through the Paleocene-Eocene (P-E) tran- '".!!! s:

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'"
0::
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-20
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sition (-60-50 Ma).

In addition, this is the first study of radiolarian faunal Fig. 2. Lithostratigraphy, biostratigraphy and carbon isotope stratigraphy of
change through the P-E transition in the mid-latitude South Upper Cretaceous-lower Eocene strata at Mead Stream (after Hollis et al.
Pacific; eastern Marlborough being situated at -55° S in the 2005). For age and biostratigraphy columns, intervals of uncertain age or bio-
zone are shaded. Three episodes of major global change are identified from
latest Paleocene (King et al. 1999; Sutherland et al. 2001). San-
bulk carbonate Ol3C stratigraphy: late Paleocene carbon isotope maximum
filippo & Nigrini (1998a) reviewed DSDP and ODP records of (PCrM), Paleocene-Eocene thermal maximum (PETM) and early Eocene cli-
late Paleocene-early Eocene radiolarians and established a matic optimum (EECO). For comparison, the global compilation of benthic
composite biostratigraphy from well-preserved, low-latitude foraminiferal 013C data of Zachos et al. (2001) is also shown; the plotted values
assemblages in 12 sites between 40 and 30 Most of these
0N 0S. are running five-point averages calibrated to the geochronometric time scale
of Berggren et al. (1995) following Hollis et al. (2005).
sites are in the western central and North Atlantic, with two in
the South Atlantic, one in the western central Indian Ocean
and one in the North Pacific. Subsequently, Nigrini & Sanfilip-
po (2000) and Sanfilippo & Blome (2001) reported on radio-
larian assemblages in well-constrained P-E boundary records Paleocene-Eocene (P-E) transition (Hollis et al. 2005). In the
in Caribbean and western North Atlantic ODP holes, respec- latter study, carbon isotope stratigraphy was integrated with
tively. radiolarian, foraminiferal and calcareous nannofossil bios-
The section exposed in the southern branch of Mead tratigraphy in order to relate lithofacies changes to major per-
Stream, Clarence River valley (Fig. 1), is the most complete turbations in the early Paleogene global carbon cycle and cli-
known record of South Pacific pelagic-hemipelagic sedimenta- mate system. Three of these episodes of perturbations have
tion from Late Cretaceous to middle Eocene (Strong et al. been identified in upper Paleocene and lower Eocene strata at
1995; Hollis et al. 2005). Rich radiolarian assemblages make it Mead Stream (Fig. 2): the late Paleocene carbon isotope maxi-
a key reference section for early Paleogene radiolarian bios- mum (PCIM - Corfield & Cartlidge 1992, Thompson &
tratigraphy and it is the type section for eight zones (RP6-13) Schmitz 1997; Kurtz et al. 2003), the Paleocene-Eocene ther-
in the South Pacific radiolarian zonation (Strong et al. 1995; mal maximum (PETM - Zachos et al. 2001, 2003), and the
Hollis 1997,2002; Hollis et al. 2005). Detailed stratigraphic and early Eocene climatic optimum (EECO - Zachos et al. 2001).
paleoenvironmental studies have been completed on the Cre- The base of the PETM corresponds with the Paleocene-
taceous/Tertiary (KIT) boundary (Hollis et al. 2003a) and the Eocene boundary (Zachos et al. 2003).The aim of the current
S80 c. J. Hollis
study is to use changes in radiolarian assemblages through the tail, as part of a paleoenvironmental study of this interval at
P-E transition at Mead Stream to further elucidate the influ- Mead Stream and nearby Branch Stream (Fig. 1). The KIT
ence of these global events on oceanic conditions in the South- boundary is defined as zero datum in the current study. As a
west Pacific. forerunner to the present study, Hollis et al. (2005) completed
a detailed stratigraphic study of upper Paleocene-lower
Eocene strata at Mead Stream, incorporating lithostratigraphy,
2. Material and Methods
foraminiferal, calcareous nannofossil and radiolarian bios-
This study is based on 42 radiolarian assemblages recovered tratigraphy and carbon isotope stratigraphy (Fig. 2).
from rock samples between 100 and 250 m above the well-con- Paleocene-Eocene strata exposed along Mead Stream con-
strained Krr boundary (Hollis et al. 2003a) in the southern and sist of well-bedded units of chert, siliceous limestone with chert
main branch of Mead Stream. Sampling methods are described nodules, siliceous mudstone, limestone and marl, which form
by Hollis et al. (2005). For radiolarian microfossil extraction, three formations within the Muzzle Group (Reay 1993). The
samples were crushed into 5-10 mm chips and leached in 15% Mead Hill Formation consists of dm-bedded chert and
hydrochloric acid (HCI) until reaction ceased. Siliceous rocks siliceous limestone. It extends from a faulted base 170 m below
with weak reaction to HCI digestion were then rinsed and the KIT boundary to a sharp contact with overlying the
leached in 5% hydrofluoric acid for 2-4 hours. Samples were Waipawa Formation at 108 m above the KIT boundary. The
then washed through a 63 urn screen and the residues cleaned Waipawa Formation is a 7.5 m thick interval (108-115.5 m)
by gentle heating in a 1:1 solution of 10% hydrogen peroxide that consists of lower and upper siliceous mudstone units with
and calgon, (NaP03)6. an intervening unit of siliceous limestone. The two mudstone
Taxonomic nomenclature is summarised in Table 1. Select- units carry the same distinctive geochemical signature of the
ed species are illustrated in Plates 1-3. Census data (Hollis et Waipawa Formation as in its type area in Hawkes Bay, North
al. 2005, supplementary data) have been derived from vertical Island (Killops et al. 2000).
traverses of strewn slides under transmitted light. For sparse The overlying formation, Amuri Limestone, has been sub-
samples (<300 radiolarians per slide), all radiolarians on one, divided into several lithotypes and members (Reay 1993; Hol-
two or more slides were counted. For richer samples, the fol- lis et al. 2005). The basal lithotype is Lower Limestone, which
lowing double count method was utilised. First, all specimens includes two members, Dee Marl and Upper Ribbon Chert.
were counted until a total count of -300 was achieved. The Lower Limestone (116-192 m) consists of dm-bedded siliceous
proportion of the slide examined to this point was determined limestone with thin marl interbeds and rare chert nodules. One
and the abundance of common taxa (>15 radiolarians or >5% unusually thick marl bed provides a useful reference point
of the total fauna) was estimated for the remainder of the (136.5-136.6 m) within Lower Limestone and is referred to as
slide. The remainder of the slide was then examined and oc- Mudstone C (following Hollis et al. 2005). Dee Marl is a 2.4 m
currences of rare taxa «5% in initial count) recorded. This thick unit (157-159.4 m) of dm-bedded alternating marl and
method has the advantage of establishing reliable abundance marly limestone. This unit marks the base of the Eocene at
estimates for rare taxa without expending great amounts of Mead Stream and at the type section, Dee Stream (Hancock et
time counting many hundreds of specimens. One disadvantage al. 2003). Upper Ribbon Chert, the highest occurrence of chert
is that the total count is highly variable, ranging in this dataset in the section, is a 12.6 m thick interval (159.4-172 m) of dm-
from <60 to >7000 individuals. Abundance of larger (>63/lm) bedded siliceous limestone with narrow chert stringers in the
diatoms was also recorded during the initial count (Hollis et al. centre of the beds. The abundance of chert decreases gradually
2005). up-section, whereas the abundance and thickness of marl in-
In order to obtain an accurate estimate of relative abun- terbeds increases. The top of Lower Limestone is placed at the
dance, all specimens have been assigned a taxonomic category. base of the first of a series of thick marl beds intercalated with
Although these counting groups range from the level of sub- limestone beds. The overlying lithotype, Lower Marl,consists
species to family, they are given equal weighting in diversity of 110 m of alternating marl and marly limestone beds, similar
analysis. to the lithology of Dee Marl. The stratigraphy of the upper
part of Amuri Limestone, which is not considered here, is de-
scribed by Strong et al. (1995).
3. Stratigraphy
4. Siliceous microfossil occurrence

The lithostratigraphy and foraminiferal, radiolarian and di-
noflagellate cysts biostratigraphy of Upper Cretaceous to mid-
dle Eocene strata at Mead Stream was described by Strong et Of 134 samples processed for radiolarians in upper Paleocene
al. (1995). Subsequently, the radiolarian biostratigraphy was and lower Eocene strata at Mead Stream (-75-300 m), 42 sam-
reviewed and revised by Hollis (1997). Hollis et al. (2003a) de- ples contained radiolarian assemblages of adequate preserva-
scribed the stratigraphy of the KIT boundary transition in de- tion and abundance for species level identification and quanti-
Paleocene-Eocene radiolarians. Mead Stream S81

} ~ a Radiolarian b . Radiolarian c. Diatom! d. Radiolarian e. Racnotanan f, FIrst and last
~:L abundanco
,se preserv ation Rad ralio divp.rsily HssemOlage occurrences
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Fig. 3. Variation (a) radiolarian pr eservation , (b)
~
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radiolarian abunda nce , (c) diatom/radiolari an
ratio, (d) radiolarian diversity (Fisher a Index).
~
.:: f- (e) relative ab unda nce of major radi olarian
=' groups, and (f) radi olarian first and last occur-
.. ,1 '" ... •
rences in upper Paleocene-lower Eocene siliceo us
microfossil assemblages at Mead Stream . Radio-
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~r- iF :""an • Mst C pre served ), Good (test morphology not affected
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4
interv al char acterised by peak radiolarian abun-
B P M G 10' 10' 10' 0 0 .2 0 .. 0 '1 8 12 0 50 HJO 0 70 40 60 80
Radiolarians!g Ratio Fisher Index "Ill Spumellaria No . of species dances (rad acme) .
tative analysis (Fig. 3) . Rad iolarians are generally common but lived increase in the marl bed (Mudstone C) near the ba se of
poorly preserve d in the upper Pa leocene where strong recrys- Z one RP7 . A rapid increase in diversity in the earliest Eocen e
ta llisation, includi ng siliceo us infilling and ove rgrowth, is evi- is followed by a gradual decline with relatively low ex values
den t in siliceo us limestone samples (Fig. 3a-b). In the first 21 through the Lower Marl.
m of Eocene strata, radiolarians are generally abundant and Paleocene radiolarian assemblages are dominated by
moderately preserved. R ecrystallisation is less prono unced in sp ume llarians (Fig. 3e) and, in orde r of abundance, the mo st
this int erval. T his probably reflects reduced silica content in common species and species groups are: Buryella tetradica,
Dee Marl. In the overlying Upper Ribbon Chert, chertifica- Haliomma spp. gr. b, Amphisphaera coronata gr, A . goruna,
tion is restricted to a central zone in each bed and samples Lithelius minor gr. and Buryella granulata. In the lowermost 21
tak en from the outer parts of beds contained relatively well- m of Eocene strata, spumell ari an dominance is greater than in
pre served siliceous microfossils. In the overlying marl-rich the Paleocene and the most common species are: Haliomma
Eocene interval, radiolarians are increasingly ra re but preser- gr. b., Amphisphaera coron ata gr. Lithelius minor gr. and
vation is moderate to good. Theocorys? d. phyzella. In overlying Eocene strata, spumel-
Di atoms are common in radiolarian resi due s in th e Pale- larians dominate to a similar extent as in the Paleocene and the
ocene but are rare to very rare in th e Eocene (Fig. 3c). The most common species are: Haliomma gr. b, Amphisphaera
orig ina l change in diatom abundance across the PIE bou ndary coronata gr. Lithelius minor gr. , Calocyclom a ampulla, Phor-
is like ly to have been even more pronounced tha n is described mocyrtis striata striata, Podocyrtis papalis, Sethocytris babylo-
here. Owing to their smaller average size and more delicate nis, Axoprunum pierinae and Periphaena heliasteriscus.
test s, diatoms are more prone to diagenetic destruction th an
radiolarians. Therefore, diatoms ar e likel y to be under-repre-
sented in Paleocene siliceous limestone samples relative to the
s. Ra dio larian fauna l turnover thro ugh the Paleocene-Eocene
transition
better pr eserved Eocene material.
Radiolarian di versity is highl y variable. Most assemblages In contrast to low-latitude regions where there is little chang e
contain 20---50 taxa whereas three assemblages have <20 taxa in radiolarian assemblages across the Paleocene-Eocene
and seven assemblages ha ve >50 ta xa. The Fisher ex Index is boundary (Sanfilip po & Nigrini 1998a; Nigrini & Sanfilippo
used to assess diversity trends in census data (Fig . 3d) . The di- 2000; Sanfilippo & Blome 2001), four episodes of significant
versity curve for all taxa is considered to be less re liable than faun al turnover are ob serv ed in the upper Pa leocene-lower
the curve for nassellarians. Mo st nassellarians have been iden- Eocene succession at Mead Stream (Fig . 3f, Table 2).
tified to species level , whereas many spume llarians remain un- Th e first episode occurs in th e lower Waipawa Formation
differentiated below family level. Thi s results in an underesti- and is associated with the RP5/RP6 zone boundary. The first
mate o f overall diversity in spumellarian-dominated samples, occurrences (FOs) of 22 species and the last occurrences (Las)
notably in the earliest Eocene. Nas sellarian ex values indicate of 12 species are recorded with in 3 m of strata (-108-111 m) .
relatively low diversity in the Paleocene, apart from a short- Some of the Las are isolated records of Cretaceous-early Pa-
S82 c. J. Hollis
Table 1. Annotated list of radiolarian species or species groups encountered in this study. Species ranges are based on the South Pacific zonation for Mead
Stream and other South Pacific ("Sth Pac") sections (Hollis. 1997,2002; Hollis et al. 1997). and on the tropical zonation (Sanfilippo and Nigrini, 1998a) for low lat-
itude ("Low Lat") records; LK = Late Cretaceous, UZP = unzoned Paleocene. EOC = Eocene. MIa = Miocene. Reference to taxon concept: CC42 = Clark and
Campbell, 1942;F73 = Foreman, 1973;H02 = Hollis. 2002; H97a = Hollis, 1997;H97b = Hollis et al. 1997;K99 = Kozlova, 1999;N92 = Nishimura, 1992;S85 = San-
filippo et al. 1985;SR92 = Sanfilippo and Riedel, 1992; S95 = Strong et al. 1995;SR73 = Sanfilippo & Riedel, 1973.
Taxon name Zonal range Taxon concept
Mead Sth Pac Low Lat. Ref. Strong et al, 1995:
Amphicraspedum murrayanum HAECKEL RP8 RP8 RP6-8 SR73

Amphicraspedum prolixum form. RP8-9 RP8-9 K99
gracilis (LIPMAN)
Amphicraspedum prolixum RP7-JO RP7-10 RP7-13 SR73
SANFILIPPO & RIEDEL gr.
Amphicraspedum prolixum RP8-JO RP8-10 RP7-9 RP7-9 208
SANFILIPPO & RIEDEL .1'•.1'.
Amphipternis alamedaensis (CAMPB. & CL.) RP3-9 RK9-RP9 LK-RP7 H97a
Amphipyndax stocki (CAMPB. & CLARK) RK9-RP7 RK9-RP7 LK-UZP H97a
Amphisphaera aft magnaporulosa RP5-9 RP4-6 ? H02
CL. & CAMPB.
Amphisphaera coronata s.l. (EHRENBERG) RP4-9 RP4-6 UZP-EOC H97a 208 (A. radiosa)
Amphisphaera goruna (SANFILIPPO & RIEDEL) RP3-8 RP2-6 UZP-RP7 H97a 208. fig. 8G. 9A
Amphisphaera kina HOLLIS RP2-7 RP2-6 H97a 208. fig. 8F
Amphisphaera macrosphaera (NISHIMURA) RP3-9 RP3-14 UZP-RP6 H97a 208 (A. radiosa)
Amphymenium splendiarmatum RP6-8 RP2-6 RP8-20 H97a
CLARK & CAMPB.
Artostrobus pusillus (EHRENBERG) RK9-RP8 RK9-RP8 H97a
Aspis murus NISHIMURA RP6 RP5-6 RP6 N92
Axoprunum pierinae (CLARK & CAMPB.) RP5-9 RP6-14 UZP-EOC SR73 208, fig. JOC
Axoprunum ? aff. bispiculum (POPOFSKY) RP6-9 RP4-6 ? H02
Bathropyramis magnifica (CLARK & CAMPB.) RP5-l3 RP4-13 ? H02
Bathropyramis sanioaquinesis RK9-RP8 RK9-RP8 LK-UZP H97a
(CAMPB. & CLARK)
Bekoma bidartensis RIEDEL & SANFILIPPO RP8 RP8 RP7-8 S95 208. fig. 9G
Bekoma campechensis FOREMAN RP6 RP6 RP6-7 S95 208. fig. 9F
Bekoma divaricata FOREMAN RP6-8 RP6-8 RP7 S95 208. fig. 9H
Buryella dumitricai PETRUSHEVSKAYA RP4-7 RP4-7 H97a 208. fig. 8L-M
Buryella foremanae PETRUSHEVSKA YA RP4-7 RP4-7 H97a 208. fig. 8K
Buryella granulata (PETRUSHEVSKA VA) RP3-8 RP4-8 H97a 208. fig. 8J
Buryella helenae O'CONNOR RP6 RP4-6 ? H02
Buryella kaikoura HOLLIS RP5-6 RP5-6 ? H97a
Buryella pentadica FOREMAN RP6 RP6 UZP-RP7 N92
Buryella petrushevskayae O'CONNOR RP6 RP5-6 ? H02
Buryella tetradica tetradica FOREMAN RP5-9 RP5-8 UZP-RP9 H02 208. fig. 8N, 9Q-R
Buryella tetradica tridica O'CONNOR RP6 RP5-6 H02
Calocycloma ampulla (EHRENBERG) RP5-10 RP5-10 RP6-14 S95 208. fig. 9V-X
Cassideus aft mariae NISHIMURA RP5-9 RP5-9 H02 209. fig. 9C tMicrosciadiocapsa ? sp.)
Clathrocyclas australis HOLLIS RP3-6 RP2-6 ? H97a P. 208. 8P (Clathrocycloma sp. A)
Clathrocyclas universa gr. RP5-13 RP5-13 ? H97b
Conoactinomma stilloformis (LIPMAN) RP9 RP9 ? K99 206 ( Conocaryomma sp.)
Cornutella calijornica CAMPB. & CLARK RK9-RP13 RK9-RP13 LK-UZP H97a
Corythomelissa adunca (SANFILIPPO & RIEDEL) RP5-8 RP4-8 RP6-8 H02 208. fig. 9B
Corythomelissa spp. RP6-8 RP6-8 ? H02
Cryptocarpium ? d. ornatum (EHRENBERG) RP6-13 RK9-RP13 H97a
Cycladophora aff. cosma LOMBARI AND & LAZARUS RP6-8 RP4-8 H02
Cycladophora d. bicornis (POPOFSKV) RP6-8 RP6-8 H02 208. fig. 90 (Clathrocycl. humerus)
Dictyomitra andersoni (CAMPB. & CLARK) RK9-RP6 RK9-RP6 LK-UZP H97a
Haliommagr. B RP4-9 H97a
Lamptonium fabaeforme fabaeforme (KRASHEN.) RP8-9 RP8-9 RP7-9 S85
Lamptonium pennatum FOREMAN RP7-9 RP7-9 UZP-RP8 S95 208, fig. 91
Lamptonium ? aff. colymbus FOREMAN RP9 F73
l.ithelius ? minor gr. JORGENSEN RK9-RP9 RK9-RPI4 H97a
Lithocampe wharanui HOLLIS RP3-6 RK9-RP6 ? H97a
Lithomespilus coronatus SQUINABOL RK9RP7 RK9-RP8 UZP-RP8 H97a
Lithostrobus longus GRIGORJEVA RPI-7 RPI-6 ? H02 209. fig. 8H-I iSuchomitra wero)
Lophophaena mugaica (GRIGORJEVA) RP5-8 RP2-8 H02

Table 1. (cont.)
Taxon name Zonal range Taxon concept
Mead Sth Pac Low Lat. Ref. Strong et aI. 1995:
Lychnocanium aff. carinatum EHRENBERG RP8 RP8 RP6-7 N92 209, fig. 9M (Rhopa/ocanium? sp.)
Lychnocanium aft. tripodium EHRENBERG RP7-8 RP7-8 K99
Lychnocanium amphitrite (FOREMAN) RP9-13 RP9-I4 RP8-20 F73
Lychnocanium auxilla (FOREMAN) RP8 RP8 RP6-7 F73
Lychnocanium bellum CLARK & CAMPB. RP9-13 RP9-I4 RP8-I4 F73 209, fig. IIl-1 (as Lychnocanoma)
Lychnocanium satelles (KOZLOVA) RP5-9 RP5-9 H02 208, fig. 9L (L. auxilla)
Lychnocanium sp. B RP7-8 S95 209, fig. 91
Lychnocanium zhamoidai KOZLOVA RP7-9 K99 209, fig. 9K (Lychnocanoma sp. A)
Middourium regu/are RP6-8 RP6-8 RP6-7 K99
Mita regina (CAMPB. & CLARK) RK9-RP6 RK9-RP6 LK-UZP H97a
Mita sp. A (? = Siphocampe? "cilizabethae") RP9-10 RP9-RPI5 ? S95 209, fig. lOX
Monobrachium irregu/are RP7-8 RP7-8 RP6-7 K99
Myllocercion acineton FOREMAN RK9-RP6 RK9-RP4 LK-UZP H97a
Orbiculiforma renillaeformis gr. (CAMPB. & CL.) RK9-RP6 RK9-RP6 LK-UZP H02
Patagospyris confluens (EHRENBERG) RP4-RP8 RP4-8 UZP-RPlO K99
Periphaena a/veo/ata (LIPMAN) RP5-9 RP5-9 K99
Periphaena heliasteriscus (CLARK & CAMPB.) RP8-9 RP6-9 UZP-EOC SR73
Peritiviator? dumitricai NISHIMURA RP5-7 RP5-6 UZP-RP6 N92
Peta/ospyris fovea/ata EHRENBERG RP5-9 RP5-9 K99 209, fig. 80-R
Peta/ospyris senta (KOZLOVA) RP6-9 RP6-9 K99 209, fig. 8S
Phacostaurus? quadratus NISHIMURA RP6-7 RP6 RP6-7 N92
Phormocyrtis cubensis (RIEDEL & SANFILIPPO) RP8 RP8 RP6-8 F73
Phormocyrtis striata exquisita (KOZLOVA) RP7-8 RP7-8 RP6-9 S95 209, fig. 9N
Phormocyrtis striata striata (BRANDT) RP7-l0 RP7-IO RP7-I4 S95 209, fig. 90-P
Phormocyrtus turgida (KRASHENINNIKOV) RP8 RP8 RP6-8 F73
P/ectodiscus circu/aris (CLARK & CAMPB.) RP6-8 RP6-8 RP6-EOC PK72
Podocyrtis acalles SANFILIPPO & RIEDEL RP8-9 RP8-9 RP8-9 S92 209, fig. 9T-U (P. aphorma)
Podocyrtis papalis EHRENBERG RP8-10 RP8-I4 RP7-I8 S92 209, fig. 9S
Pterocodon pocu/um NISHIMURA RP6-7 RP6-7 RP6-7 H02
Saturnalis kennetti DUMITRICA RP3-8 RPI-8 ? H97a
Sethochytris baby/onis (CLARK & CAMPB.) gr. RP6-13 RP6-I5 RP6-20 H02 208, fig. llA-B (as Lychnocanoma)
Siphocampe nodosaria (HAECKEL) RP6-13 RP4-I5 RP6-I4 H02 209 (S. arachnaea)
Siphocampe quadrata (PETRUSHEV. & KOZLOVA) RP8-13 RP5-I5 RP6-20 H02 209, fig. lOR
Spongurus bi/obatus gr. CLARK & CAMPB. RP5-9 RP4-9 ? H02
Stichomitra carnegiense CAMPB. & CLARK RK9-RP6 RK9-RP6 LK-UZP H97a
Sty/odictya targaeformis CLARK & CAMPB. RP5-9 RP5-9 ? CC42
Sty/osphaera minor CLARK & CAMPB. RP3-9 RP3-6 UZP-MIO H97a
Theocampe urceo/us (CAMPB. & CLARK) RP9-13 RP9-I5 RP8-20 S95 209, fig. lOQ
Theocampe vanderhooft (CAMPB. & CLARK) RK9-RP6 RK9-RP6 LK-UZP H97a
Theocorys? aft. phyzella FOREMAN RP6-8 RP6-8 ? H02 208, fig. 9E (Ca/ocyclas asperum)
Theocorys? cf. phyzella FOREMAN RP7-8 ? RP7 SBOI (as Theocorys aff. phyzella)
Theocorys? phyzella FOREMAN RP8 RP7-8 F73
Tholodiscus densus (KOZLOVA) RP6-8 RK9-RP8 ? H97a
S84 c. 1. Hollis
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Fig. 4. Ranges of radiolarian index species in upper Paleocene-lower Eocene strata at Mead Stream. For age and biostratigraphy columns, intervals of uncertain
age or biozone are shaded, Low-latitude ranges are based on (Sanfilippo & Nigrini 1998a, b) and Sanfilippo & Blome (2001).
leocene species (e.g. Uthocampe wharanui, Mita regina, Myl- LOs are the last record of Cretaceous-Paleocene species that
locercion acineton) that may be reworked into the siliciclastic are generally very rare in the upper Paleocene but are present
Waipawa Formation. Other events are isolated records of taxa in one or more of three relatively rich samples in this interval
that are largely restricted to two particularly rich and diverse (P20/flOn, fl076, fl081); two of these species may be re-
assemblages (P30/f318, f452), e.g. Buryella helenae, B. petru- worked into Mudstone C (Amphipyndax stoch, Theocampe
shevskayae, Cryptocarpium cf. ornatum. As most of the FOs vanderhoofi). Three species with FOs in this interval are key
recorded in this interval relate to species that are known to index species that have much earlier FOs in low latitudes
have earlier local FOs (Hollis 1997, 2002), this episode is not (Lamptonium pennatum, Phormocyrtis striata exquisita, P. s.
thought to reflect a time of high evolutionary turnover. In- striata - Fig. 4). All are very rare in overlying Paleocene strata
stead, the large number of events at the base of the Waipawa but occur consistently in the lowermost Eocene.
Formation suggests that either the unit is highly condensed or The third faunal turnover episode occurs at the base of the
that there is a basal unconformity. Dee Marl (Fig. 3f, Table 2). The FOs of 13 species are record-
The second episode occurs within or directly below the 10 ed within the first 3 m of Eocene strata at Mead Stream, with 9
cm thick marl bed (Mudstone C). The FOs of 6 species and of these species in the basal Eocene sample. Four of these
the LOs of 5 species occur within 5 m of strata. Most of the species appear to be global markers for the PIE boundary

\/J
00
0-
(1
~ Table 2. Radiolarian species in order of (A) first and (B) last occurrence at Mead Stream. Species with first occurrences lower in the section (Hollis et al. 2003a) or last occurrences higher in the section
::c (Strong et al. 1995) are not shown. (MHF = Mead Hill Formation, WF = Waipawa Formation, OM = Dee Marl, URC = Upper Ribbon Chert)
~ A. IN ORDER OF FIRST OCCURRENC E
International Age Late Paleocene Early Eocene
South Pacific Radiolarian Zone 5 RP6 RP7 RP8 RP9
Lithologic Unit MHF I WF I Lower Limestone DMI URC I Lower Lst Lower Marl
r- N '-0-"'1" 0. 0 ...... 0 t.I) t.I) N ,..., on 0. N 00 \0 t> ...... M
'-0 r--OOMNVlOO O\r-r-OOo\o\OOO ...... O ............ N N N NMNMM,,"'o:::tlrlN V)O\\Ot-ooooO\r-
("'fj ........................ ,...,
;;; "'1" ,...,
N N O O O O O ...... M ................... ............ M - M - - - N
NZ Fossil Record No. P30/f t.I)-_ .................. t.I) ..................
"'1" on ~;:;:;~~:2~ _V)-_ . . . . . . ........... t.I)-t.I)---- on on ~r;:;~~r;:;r;:;~
r-NMMOO-.:::tV) O\oo\N\oo\NO'o:::t-MOOQOO\O\O MO\ONV)-OOt.l)t.I) 0 0 V)oaOOOlrl
0 r--: I.d""': N~\Cir-..:o\,....;N~v)c;:.,o,....;ooi..,fv)'..ci ooo\o\.....:~r...:oci.....:v)
000000 -D a ~06v)OON
0; O O O - - ...... M MMMMM'o:::t'o:::t""1" " T " T V ) V ) V ) V) V) V) V)t.I)V)\O'O\O\Or--r- r-. co O\NM'o:::j"VlOOO\
DEPTH (metres above KTB) 0. ----- ............. - - - - - - - ........................ ----- - - - .................................... --NNNNNN
Axoprunum pierinae
* * * * * * * * * * * * * * * * * * * * * * * * * *
-*
* * * * * * * *
Axoprunum? aff. bispicu/um
Bathropyramis magnifica
* * * ** * * * * * * * * * * * * * * * * * * * * * *
* * * *
Buryella tetradica
* * * * ** * * * * * * * * * * * * * * * * * * * * *
* * * * * * *
Cassideus afJ. mariae
* * * * ** * * * * * * * * * * * * * * * * * * * * * * * * * * *
* * * * * *
Conocaryomma stilloformis
* * * * ** * * * * * * * * * * * * * * * *
Corythome/issa adunca
* *
He/iosty/us spp.
* * * * * * * * * * * * * *
Peritiviator? dumitricai
* * * * *
Periphaena a/veo/ata
* * *
Sty/odictya targaeformis
* * * * ** * * ** * * * * * * * * * * * * * * * * * * * * *
Ca/ocyc/oma ampulla
* * * * *
Cyc/adophora aff. cosma
* * * ** * * * * * * * * * * * * * * * * *
Lophophaena mugaica
* * * *
Patagospyris conjluens
* * * * *
Peta/ospyris fovea/ata
* * * * * * * * * * * *
Amphisph. aff. magnaporu/osa
* * * * * * * * * * * * * * * *
Amphymenium sp/endiarmatum
* * ** * * * * * * * * * * * * * * * * * * * * * * * * *
Aspis murus
* * * * * * * * * * *
Bekoma campechensis
*
Burella pentadica
* *
Buryella tridica
* *
Cyc/adophora cf. bicornis
* *
Peta/ospyris senta
* * * * * * *
Siphocampe nodosaria
* * * * * * * * * * * *
Amphicraspedum pro/ixum gr.
* * * * * *
Buryella he/enae
* * * * * * * * * * * * * *
Buryella petrushevskayae
*
Cryptocarpium cf. ornatum
*
* * * * *
Table 2 (cont.)
South Pacific Radiolarian Zone 5 RP6 RP7 RP8 RP9
Lithologic Unit MHF I WF I Lower Limestone DMI URe I Lower Lst Lower Marl
t- N\O--"¢O\O -Olf)lf) NMV'l O\N 00 \Or- ...... (""')
- '0 r--OOMNtr)OO O\r---r--OOO\O"IOOO ...... O - - N N N NMNMM"'1"..q-lJiN ~ If)O\\Or-ooooO\r-
M
NZ Fossil Record No. P30/f ~ ~ ~~~~S~ N O O O O O - M - - - - M - - - --('f')-M---N v; ~I~;;:~~r;::r;::~
r-NMMOO""'1" V)
trl------tr'l----lJi--- --1£)-'£)----
O\OO\N\oo\NO"'1"-MOOQOO\O\O MO\oNt.n-OOV)V) 0 Olr)OOOOOlll
0 r-:oooo66-.o"",: NIJ:3~~~,....;N...fv)o\O.....:...f~v)"O 000\ 0\""':..0 r-: 00""': v) -D ci..q:6ov)OON
~ OOO---M lr)lr)lf)\O\O\O\Or--r--
MMMMM"'1"~-.:t""T"'i'"V)l.r)lJilr)V)V) t- OOCf\NM"'i'"lJiOOO\
DEPTH (metres above KTB) o- ------- ---------------- --------- --NNNNNN
Bekoma divaricata * * * *
-*
Lychnocanium zhamoidai
* *
Middourium regulare
* * * * * * * * * * * * *
Phacostaurus? quadratus
* * * * * * * * * * * * * * * * * * * * *
Plectodiscus circularis
* * *
Sethochytris babylonis gr.
* * * * * * *
Theocorys ? aff. phyzella
* * * * * * * * * * * * * * * * * * * * * * * * * * **
Theocorys? cf. phyzella
* * * * * * *
Tholodiscus densus
* * * * * * * * * * * * * * * * * * *
Lychnocanium sp. B
* * * * *
Phormocyrtis striata exquisita
* * * * * * * *
Lychnocanium aff. tripodium
* * * * * * * * * * * * * * * * * *
Phormocyrtis striata striata
* * * * * *
Pterocodon poculum
* * * * * * * * * * * * * * * * * * * *
Lamptonium pennatum
* * *
Monobrachium irregulare
* * * * * * * * * * *
Amphicraspedum murrayanum
* * * * * * * * * * * * * *
Amphicraspedum prolixum s.s.
* *
A. prolixum f. gracilis
* * * * * * * * * *
* * * * * * *
Lamptonium.f fabaeformae * * * * * * * *
Lychnocanium auxilla * * * * * * * * * *
Phormocyrtis cubensis * * * * *
-e
Phormocyrtis turgida * * * *
;:;
'" Theocorys ? phyzella
o Podocyrtis papilis
* * * * * * * * * * *
n
(l) * * * * * * * * * * * * * * *
(l)
Clathrocyclas universa gr. * * *
"
'in Lychnocanium aff. carinatum
* * * * * * *
g Podocyrtis acelles
* * *
(l) * * * * * *
(l) Bekoma bidartensis
..,'"
Siphocampe quadrata
* *
0- * * *
s'" Periphaena heliasteriscus * * * * * * **
0;
::l. Mita sp. A * *
Lamptonium? aff. columbus
*
~'"
Theocampe urceolus
* * *
~ * * *
(l)
Lychnocanium bellum * *
'"0- Lychnocanium amphitrite * *
~
(l) First Occurrences 10 5 9 4 0 9 0 0 2 3 I 0 0 0 0 0 0 0 0 0 0 I 0 0 9 3 I 0 I 0 0 0 I I 0 I 0 2 0 I 0 0
'"a
(/)
00
-.J
CIl
00
00
o,.... Table 2 (cont.)

B. IN ORDER OF LAST OCCURRENC E
::r:
~ South Pacific Radiolarian Zone 5 RP6 RP7 RP8 RP9
Lithologic Unit MHF I WFI Lower Limestone DMI URe I LowerLst Lower Marl
r- N"-O"-'o::1"O\O .- 0 l£) VI N ("0') V") 0'0 N 00 1..0 r- .- M
\0 t-OOMC".! V) 00 O \ r - r - O O O \ O \ O O O - O - - N N N NMNMM"T"TV)N lIiO\\Ot-OOOOO\t-
N O O O O O - M - - - - M ............ - - - M ...... M - - - N ("0')
NZ Fossil Record No. P30/f ~ ~ ~~~~~~ Vl- .................. --Vl-- . . . . -tr) ............ - ...... - t r l - l £ ) . . - ................... V")
""" ~~~~~~r;::~
r-NMMOO"TV) O\CO\N\OO\MO-.:::t-MOOQOO\o\o MO\OMV')-OOV)l.(') 0 Ol()OOOOOtn
0 r----:aOoooci~,...; NI,cj'-Cir...:o\"";N~V-)o\O"";-.i~~-.o oOa\o\"",:",or:-..:cx:i"";ar) o~OOv)OON
~
0\ 0 0 0 ................... ("1') MMMMM"T"T"T..q-"TV)V')l.r)V)V)V) tr)lr)lf)\O\OI"Q\Or-r- r- OOO'\NM"Ttr)OOO'\
DEPTH (metres above KTB) 0'0 ........................ ..- ............ -_ ...... .- ............ ..- __ .................................... - .................. _----- - --NNNNNN
Aspis murus
Lithocampe wharanui
*
Mita regina
* *
Buryella helenae
*
Buryella kaikoura
*
Burella pentadica
* * *
Buryella petrushevskayae
* *
Buryella tetradica tridica
*
Myllocercion acineton
* *
Bekoma campechensis
* *
Clathrocyclas australis
* *
Dictyomitra andersoni
* * * *
Stichomitra carnegiense
* * * *
Amphipyndax stocki
* * * *
* * ** * * ?
Theocampe vanderhoofi
Phacostaurus? quadratus
* * *
Peritiviator? dumitricai
* * *
Pterocodon poculum
* * *
Buryella dumitricai
* * *
Buryella foremanae
* * * ** * * * * *
Amphisphaera kina
* * * ** * * * * * *
Lithomespilus coronatus
* * * ** * * * * * * * * * * * *
Theocorys ? aff. phyzella
* * * * * * * * * * * * * * * * *
Lithostrobus longus
* * * * * * *
Amphicraspedum murrayanum
* * * * * * * * * * * *
Cycladophora aff. cosma
* *
Lychnocanium aff. tripodium
* * * *
Bekoma bidartensis
* * * * * *
Lychnocanium sp. B
* *
Plectodiscus circularis
* * * * * * * *
Lophophaena mugaica
* * * * * * *
Patagospyris conjluens
* * * * *
Cassideus aff. mariae
* * * * * * * * * * * *
Orbiculiforma renillaeformis gr.
* * * * * * * * * * * * * * * * * * * * * *
Phormocyrtis cubensis
* * * *
Siphocampe quadrata
* * * * *
* * *
Table 2 (cont.)
South Pacific Radiolarian Zone 5 I RP6 RP7 RP8 RP9
Lithologic Unit MHF I WF I Lower Limestone DMI URC I Lower Lst Lower Marl
r-. NI..O-"::to\O -OV"'l1fl NMt.(") O\N 00 \Or--r<")
O\r--r--OOO\O\OOQ-O ............. N N N NMN("'I")("'I")'o::T"::tIflN
_ _ ('I")_("I") _ _ _ N V) 0\ \0 r--oo OOO"lt-
- '0 r--OO("'f"")NlIJOO Ifl __ _ _ _ t.rl _ _ ___ _ _ If'l_tr) _ _ _ _
NO O_ O O O -M - -_
- -_
(" f 'l£}
) ............ - """
NZ Fossil Record No. P30/f ~ ~ ~;;;~~S~ <r)
'" r;:;2:r;:;r;;~r;:;r;:;~
r--N ("f") M 00 "::t If) O\OO\N\oo\NO"::t-MooOoo\.O\O MO\oNV"'l-OOlflV) 0 OlflOOOOOl.rl
Cl r---=oOoocici~"""': N0v:3r-:o\,......:N~v)o\ci,......:~~v)~ 00 0.; 0\""": 0 r-:oci"""': v-; 0 o-¢ooV-:oo<"i
OOo\NM7V)OOQ'\
o-. o O O - - - r < " ) ("f'jMM("'I"")r+")"::t "::t"::t"::t"::t 1 f l V ) V ) V ) V ) t . ( " ) V) l£) V) \0 \0 \0\0 r--r-- r-
DEPTH (metres above KTB) ...- ..- ...... ...- ........................ ............ ...... - N N N N N N
...... ............ ............ ...- ...... ......
o- --_ - __ -_ -----_ -
Artostrobus pusil/us * * * * * * * * * *
* * * * * *
Buryella granulata * * * * * * * * * * * * *
* * * * * * * * * * * * * * *
Cycladophora cf. bicornis * * * *
Lychnocanium aff. carinatum
* * *
* * *
Middourium regulare * * * * * * * * * * * * * * * * * * * * *
Phormocyrtis striata exquisita * * * * * * * * * * * * * * * * * *
Phormocyrtis turgida * * * *
Saturnalis kennetti * * * * * * *
Siphocampe nodosaria * * * * * *
Tholodiscus densus * * * * *
Amphisphaera goruna * * * * * * * * * * * * * * * * * * * * * * * * * * * * * *
Amphymenium splendiarmatum * * * * * * * * * * *
Bekoma divaricata * * * * * * *
Corythomelissa adunca * * * * * * * * * * *
Lychnocanium auxil/a
* * *
* * * * * * * * * *
Theocorys? cf. phyzella * * * * * * * * * * * *
Theocorys ? phyzella
* * * * * * *
* * * * * * * * * * *
Amphipternis alamadaensis * * * * * *
Axoprunum? aff. bispiculum * * * * * * * * * * * * * * * * * * * * *
Lamptonium pennatum
* * * * * * * * * *
* * * * * * * * * * *
Petalospyris fovealata * * * * * * * * * * * * * * * *
Petalospyris senta * * * * * * * * * * * *
Amphisph. aff. magnaporulosa * * * *
"0 * * * * * * * * * * * * * * * * * * * * * * * * *
eo. Monobrachium irregulare
C1> * * * * * * * * * * * * * *
o Clathrocyclas universa gr. * *
* * * * * * * *
Ii::J Lamptonium? aff. columbus
C1> * * *
m Lychnocanium satelles * * * * * * ** * * * * * * * * * * * * * * * *
g Stylodictya targaeformis * * * * *
::J Buryella tetradica
C1> * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * * *
..., Cryptocarpium cf. ornatum
0> * * *
0- * *
o' Lamptonium f fabaeformae * * * * * * * *
::l. Lychnocanium amphitrite * *
'"
§ Lychnocanium zhamoidai * * * * * * *
.'" Periphaena alveolata
* * * * * *
~
C1>
* * * * * * * * * * * * * * * * * * * * * * * * * * * * * * *
0> Podocyrtis acelles * * * * * *
0- 0 2 1 0 3 I I 0 4 10 7 5 2 3 I 1 5 I
Last Occurrences 10 0 3 6 0 3 I 0 0 2 I 2 0 0 0 0 0 0 0 I 0 2 0 I
iq
C1>
0>
a
C/l
00
\0
mtll·es
La te Paleocene Earty Eocene
RPSI RP6 I RP7 RP8 I RP9
....
Latest Paleocene (56 Ma)
. >I ; ~:-..-; If > I
1
',-I
Radiolarian Biofacies A B I Radiolarian Biofacies C
:p' • Amphisphaera goruna gr.
:J V "" .'" Buryella granulata gr. >:

'"5 '
-<
SO'S
Buryella tetradica
Cassideus aft. mariae

• SO'S
I
m~
• gif
°rJ----.,-.. -..-~.-~-~------------
;; Axoprunum spp.
~ ..~ .. Cool surface currents (eut rophi c reg ime)
Fig. 6. Paleo geographic reconstru ction of New Zealand region durin g the lat-
Haliomma? gr. b est Paleocene (adapted from King et al. 1999) showing major current systems
based on coupled climate model simulations for the Eocen e (Huber 2(0 2)
a
:.,.r--- - --- - -+!- - " " " Periphaena
":"--:-:-- -spp.- - - - -
] ........ !* PMiddouriumIMonobrachium ,
of the Waipawa Formation, the low number of LOs relative to
: .,.... - P/ectopy~mid';' ! • • FO s in the PIE boundary interval indicates that th e faunal
:J Ca/ocycJoma ampulla ! turnover is not due to a str atigraphic discontinuity. The abr upt
OJ -=oj , '4l '-< appe arance of many low-latitude species indicates that warm-
s Phormocyrtis spp. i m >:
ing at the PETM promoted pol e-ward migration of warm -
J "7
Am-p-;h"7"ic-ra-sp-e-:;"
du--m:-p--ro
:-::/i""
xu--m--g--r·-+
i --""""'--====----======-- ~ i
.
: T
water radiolar ians. Similar, but less extre me, shor t-lived warm-
oJ] Cit
Theocorys? phyzella gr.
', .--_ - --
'a 4JI"
:.,
- - - -- -_
ing ma y explain the FOs of severa l low-latitude species around
the time of deposition of Mud stone C (Fig. 3f, Table 2).
~1 i
J Podocyrtis papalis The final episode of marked faunal change in the studied
~ section occurs -20 m abo ve the PIE boundary (Fig. 3f, Table
2). Within the same 5 m interval in which radiolarian abun-
Fig. 5. Relative abundance of commo n radiolarian species and species group s dance begins to decrease (175-1 80 m), the LOs of 21 species
in upper Paleocene-lower Eocene strata at Mead Stream.
are recorded. These speci es include many that thrived through
much of the late Paleocene as well as many that appeared in
th e ea rliest Eocene (Fig. 4). Only two FOs are recorded in this
(Sanfilippo & Nigrini 1998a; Sanfilippo & Blome 2001): Lamp-
int erval, one of these being th e FO of the common E ocene
tonium fabaeforme fabaeforme, Pod ocyrtis papalis, Phorm o-
ph acodiscid Peripha ena heliasteriscus. The high number of
cyrtis turgida and Theo corys? ph y zella (Fig. 4). Five of the
LO s, coupled with th e onset of long term declin e in radiolarian
species are index species with late Paleocene FOs in low lati -
abunda nce and diversity, ind icat es that this episode of faun al
tudes (Sanfilippo & Nigrini 1998a, b): A mphicraspe dum mur-
change refle cts a oceanographic changes unfavourabl e to all
rayanum , A prolixum s.s., Bekoma bidartensis, Lychnocanium
rad iolari ans.
auxilla, Phormocyrtis cub ensis (Fig. 4).
There ar e relatively few significant LOs associated with the
PIE boundary (Fig. 3f) . The LOs of Lithomespilus coronatus, 6. Radiolarian biofacies
Lith ostrobu s longus and Theocorys? aff. ph yz ella occur within
To simplify the discussion and presentation of census dat a
th e upp ermost 3 m of Paleocene stra ta . Of the three LOs that
(Fig. 5) in th is section, some species are combined into larger
occur with in the basal Eocene De e Marl , two are species that
gro ups that reflect a close ph ylogen et ic relationship or mor-
are restricted to the unit (A mphicraspedum murrayanum, Ly-
ph ologic al similarity:
chnocanium aff. tripodium ) while one is the LO of a sporadi-
cally occurring late Paleocen e sp ecie s (Cycladophora aff , J. A mphisphaera coronata gr. (A. macrosphaera, A. coronata
cosma). In contrast to the faun al changes observed at the base s.l, A. aff, magnaporulosa).
S90 c. J. Hollis
2. Amphisphaera goruna gr. (A. kina, A. goruna) -137 and -142 m, and overlying Eocene marl layers (Fig. 5).
3. Axoprunum spp. (Axoprunum pierinae, A. aft bispiculum) Despite the significant lithologic change from Dee Marl to
4. Buryella granulata gr. (B. granulata, B. foremanae, B. du- Upper Ribbon Chert, there is little change in siliceous micro-
mitricai) fossil assemblages. A slight increase in biosiliceous productivi-
5. Middourium/Monobrachium (Middourium regulare, Mo- ty in Upper Ribbon Chert may be inferred from the increase in
nobrachium irregulare) silica content and a small increase in diatom abundance, al-
6. Periphaena spp. (Periphaena alveolata, P. heliasteriscus) though overall radiolarian abundance and diversity declines
7. Phormocyrtis spp. (Phormocyrtis cubensis, P. striata ex- (Fig.3b-d).
quisita, P. striata striata, P. turgida) Biofacies B has unusual characteristics that make interpre-
8. Plectopyramidins (Bathropyramis and Cornutella spp.) tation difficult. The presence of many low-latitude species in-
9. Theocorys? phyzella gr. (Theocorys? d. phyzella, Theoco- dicates relatively warm climatic conditions. The low abun-
rys? phyzella) dance of diatoms and of radiolarian taxa characteristic of the
eutrophic Paleocene Biofacies A, suggests conditions of rela-
Three radiolarian biofacies are identified in the Paleocene- tively low productivity. However, radiolarian abundance and
Eocene section at Mead Stream: (A) upper Paleocene diversity maxima are recorded in Dee Marl and in underlying
(100-157 m), (B) lowermost Eocene (157-178 m) and (C) Mudstone C, which also contains an assemblage similar to Bio-
lower Eocene (178-250 m). facies B. A very similar assemblage is present in lowermost
Biofacies A is characterised by abundant Buryella tetradica, Eocene carbonate ooze at high-latitude DSDP Site 277, south-
common to abundant Amphisphaera coronata gr. and Haliom- west Campbell Plateau (Hollis et al. 1997). The three features
ma gr. b, and few to common Amphisphaera goruna gr., of high total abundance, high diversity and high numbers of
Buryella granulata gr. and Cassideus aft. mariae (Fig. 5). Di- warm-water species are consistent with late Cenozoic tropical
atoms are common in this biofacies (Fig. 3c) and sediments are radiolarian assemblages (Casey 1993). This suggests that the
moderately siliceous, with a highly siliceous facies of the or- PETM may have promoted the expansion of weakly eutrophic
ganic-rich Waipawa Formation (Killops et al. 2000) occurring subtropical water into the Southwest Pacific (Fig. 6). Further
in the lower part of the interval. Faunal similarities with di- biogeographic study of the distinctive spumellarians that dom-
atom-rich, biosiliceous Paleocene lithofacies lower in the Marl- inate this biofacies is required to confirm this interpretation.
borough sequence (Hollis et al. 1995, 2003a, b) and on the Biofacies C occurs in the upper part of the lower Eocene
eastern margin of Campbell Plateau (Hollis 2002) suggest sequence (178 to 250 m). It is characterised by abundant Am-
moderately eutrophic conditions, consistent with weak to mod- phisphaera coronata gr., Haliomma gr. b, common Calocyclo-
erate upwelling in outer shelf to upper slope depths (Fig. 6). ma ampulla, few to common Phormocyrtis striata striata and
This biofacies spans an interval correlated with the PCIM, few to rare Amphicraspedum prolixum gr., Axoprunum spp.,
which is interpreted as a 3 m.y. episode of enhanced global car- Buryella tetradica, Periphaena heliasteriscus, plectopyramidins,
bon burial, either through increased marine or terrestrial pro- Podocyrtis papalis and Sethochytris babylonis gr. An upwards
ductivity (Corfield & Cartlidge 1992; Thompson & Schmitz decrease in radiolarian abundance and diversity, coupled with
1997; Kurtz et al. 2003). The regional occurrence of organic- low diatom abundance, indicates that this biofacies represents
rich mudstone in the lower part of the interval (Killops et al. relatively oligotrophic conditions. Similar assemblages are
2000; Hollis et al. 2005) and abundant siliceous microfossils found in lower Eocene carbonate ooze at DSDP site 277 (Hol-
(Hollis 2002) indicates that this event is associated with en- lis et al. 1997). As this biofacies is correlated with the EECO, it
hanced marine productivity in the New Zealand region. seems likely that global warming associated with this event
Biofacies B occurs in the lowermost 21 m of Eocene strata promoted the expansion of oligotrophic subtropical water-
(157-178 m). It is characterised by abundant Haliomma gr. b masses into the Southwest Pacific.
and few to common Phormocyrtis spp. and Theocorys? phyzel-
la gr. (Fig. 5). The base of the biofacies coincides with the PIE
7. Conclusions
boundary and the onset of the PETM. A marked increase in
radiolarian abundance and diversity is associated with a sharp The first quantitative study of radiolarian assemblages from an
decline in diatom abundance and in the abundance of Buryella upper Paleocene-lower Eocene bathyal succession has identi-
tetradica - the dominant radiolarian species in the upper Pale- fied three distinct biofacies that can be used to determine the
ocene (Fig. 3b-d, f). This species and B. granulata s.s. are rare influence of major changes in the global carbon cycle and cli-
in this interval, apart from two samples in which both species mate system on Southwest Pacific oceanic conditions.
are common (P30/f533, fl223). The high abundance of large, Biofacies A is associated with the PCIM, a 3 m.y. episode of
robust spumellarians, namely Haliomma gr. b, Middouri- maximum 013C values that is inferred to represent high rates of
um/Monobrachium gr. and Periphaena alveolata, is a distinc- carbon burial due to enhanced marine or terrestrial productivi-
tive feature of Biofacies B. These taxa tend to be most abun- ty (Corfield & Cartlidge 1992; Thompson & Schmitz 1997;
dant in marl samples of Dee Marl and are also common to Kurtz et al. 2003). Hollis et al. (2005) argued that Mead Stream
abundant in underlying Paleocene marly layers, notably at lithofacies, including organic mudstone and siliceous limestone,

are indicative of enhanced marine productivit y in the Marlbor- REF ERE NCES
ough basin during the PCIM. Siliceous microfossil assemblages BERGGREN W., KENT D., SWISHER C. & A UBRY M.-P . 1995: A rev ised Ceno-
also indicate enhanced marine productivity. Biofacies A is asso- zoic geochro no logy and chronostratigraphy. In : BERGGREN, W. et al.
ciated with common diatom s and is dominated by the same or (E ds) : Geochro no logy, ti me sca les and globa l stratigr aphic co rre latio n,
129- 212, SE PM Spe cial Pu blicati o n 54.
similar species that are found in biogenic silica-rich lithologies
8 0 LTOVSKY D . 1987: Sedimen tar y record of radiolarian bio geograph y in the
in the lower Paleocene in Marlb orou gh (Hollis et al. 2oo3a, b) eq ua torial to Antar ctic wes te rn Pacific O cean. Micropaleontology 33,
and in the upper Paleocene of the Campbell Plateau margin 267-281.
(Hollis 2002). Deposition of radiolarian and diatom-rich sedi- CASEY R. 1993: Radi olar ia. In: LIPPS, J.H . (Ed.): Fossil Prok aryote s and Pro-
tists, 249- 284. Blackw ell Scientific Pu blicat ion s, O xford/London , U K.
ments in the Marlborough basin, as well as on the northwest
CAULET J ., VENEC-PEYRE M.-T. , VERGNA UD-GRAZZINI C. & NIGRINI C. 1992:
and southeast continental margin of New Zealand, in the late Va riatio n of South So ma lian upw elling du ring the last 160 ka: rad iolarian
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which at Mead Stream is associated with the transition from earliest Tertiary of Marlbo rou gh , New Zealand. Geology (Boulder) 23.
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characterised by declining radiol arian abundance and diversity HOLLIS c, WAGHORN D ., STRONG C. & CROUCH E. 1997: Integrated Paleo-
ge ne biostratigraphy of DSD P site 277 ( Leg 29): foramin ifer a, cal care ou s
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well as high-latitude radiolarian species disappear at the onset HOLLIS c, RODGERS K., STRONG c., FIELD B. & ROGERS K. 2oo3a : Paleoenvi-
of the EE CO . With similar early Eocene radiolarian assem- ro nme nta l chan ges ac ross the CretaceouslTertiary boundary in th e north -
ern Clare nce Va lley, so uthea stern Ma rlbo rou gh , New Zea land. New
blages recorded from the southern Campbell Plateau (Hollis et Zealand Journ al of Geology and G eophysics 46, 209-234.
al. 1997), progressive warming is inferr ed to have promoted in- HOLLIS C; STRONG C , RODGERS K. & R OGERS K. 2oo3 b: Paleoen vironment al
creasingly oligotrophic oceanic conditions around New Zealand. cha nges across the Cre taceouslT erti ar y bo und ary at F laxbourne R iver
and Wo od side Cr eek , eas te rn Marlborough, New Z ealand. New Z ealand
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Acknowledgements Eoce ne tran sitio n at Mead Stre am, New Z ealand: a southe rn Pacif ic
Sa mp le collec tio n and stra tigr a phic integration was ca rrie d o ut wit h the assi- reco rd of ea rly Ce nozo ic globa l change. Pal ae ogeography, Palaeoclima-
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versity Press, New York.

Plate 1
Scanning electron micrographs of spumellarian radiolarians from lower Eocene strata, Mead Stream. Scale bars = 100 urn.
1. Amphisphaera coronata EHRENBERG gr. CH12076, P30/f534, 176 rn, RP8.

2. Axoprunum aft. bispiculum (POPOFSKY). CH12086, P30/f534, 176 m, RP8.
3. Axoprunum pierinae (CLARK AND CAMPBELL). CH12079, P30/f534, 176 rn, RP8.
4. Stylosphaera min or CLARK AND CAMPBELL. CH12078, P30/f534, 176 m. RP8.
5. Haliomma gr. b. (cf. Theocospha erella rotunda (BORISSENKO» . CH 12005, P30/fl1 29, 158.3 m, RP8.
6. Haliomma gr. b. CH l 2001. P30/fl 129, 158.3 m, RP8.
7. Periphaena? duplu s (KOZLOVA). CH12020, P30/fl1 29, 158.3 m, RP8.
8. Periphaena? duplu s (KOZLOVA). CH12019, P30/fl1 29, 158.3 m, RP8.
9. Periphaena? duplus (KOZLOVA). CH12016. P30/fl1 29, 158.3 m, RP8.
10. Periphaena? duplus (KOZLOVA). CH12018, P30/f1129. 158.3 m, RP8.
11. Midd ourium regulare (BORISSENKO). CH12301, P30/fl1 29, 158.3 m, RP8.
12. Monobrachium irregulare (NISHIMURA) . CH12012, P301f1129, 158.3 m, RP8.
13. Monobrachium irregulare (NISHIMURA). CH I2011, P30/fl1 29, 158.3 m. RP8.
14. A mphiscraspedum prolixum SANFILIPPO AND RIEDEL CH12094, P30/f534. 176 m. RP8.
15. A mphicraspedum gracilis (LIPMAN). CH12093, P30/f534, 176 m, RP8.
16. A mphicraspedum gracilis (LIPMAN) . CH 12092, P30/f534, 176 m, RP8.
17. Amphicraspedum gracilis (LIPMAN). CH12091, P30/f534, 176 m, RP8.
18. A mphicraspedum murrayanum HAECKEL. CH12030, P30/fl1 29, 158.3 m, RP8.
19. A mphicraspedum murra yanum HAECKEL. CHI 2064. P30/fl129, 158.3 rn, RP8.
20. Amphiscraspedum prolixum SANFILIPPO AND RIEDEL CH12059. P30/f1129. 158.3 m, RP8.
21. A mphiscraspedum pro/ixum SANFILI PPO AND RIEDEL CH12089. P30/f534, 176 m, RP8.
22. Spumellarian gen. et sp. indet. CH 12085, P30/f534, 176 m, RP8.
S94 c. J. Hollis
Plate 2
Scanning electron micrograph s of nassellarian radiolarians from upper Paleocene (Fig. 4) and lower Eocene stra ta, Mead Stream.
Scale bar = 100 11m.
1. Patagopsyris confluens (EHRENBERG). CH12074, P30/fl129, 15S.3 m, RPS.

2. Petalospyris foveolata (EHRENBERG). CH12075, P30/f1129, 15S.3 m, RP8.
3. Clathrocyclas universa CLARK AND CAMPBELL. CH12103, P30/f534, 176 m, RP8.
4. Buryella tetradica FOREMAN. CHOI028, P30/f318, 108.3, RP6.
5. Buryella tetradica FOREMAN. CH12040, P30/fl129, 158.3 m, RPS.
6. Buryella tetradica FOREMAN. CH12065, P30/fl129 , 158.3 m, RP8.
7. Bathropyramis magnifica (CLARK AN DCAMPBELL). CH12101, P30/f 534, 176 m, RP8.
8. Bathr opyram is m agnifica (CLARKAND CAMPBELL). CH12100, P30/f534, 176 m, RP8.
9. Lychnocanium pon derosa (KOZLOVA). CHI 2067, P30/fl 129, 158.3 m, RP8.
10. Lychnocanium satelles (KOZLQVA). CH I 2098, P30/f534, 176 m, RPS.
11. Phormocyrtis striata exq uisita (KOZLQVA). CH12070, P30/fl 129, 158.3 m, RP8.
12. Phormocyrtis striata striata BRANDT. CH12038, P30/fl1 29, 158.3 m, RP8.
13. Phormocyrtis turgida (KRASHENINNIKOV). CH12023, P30/fl1 29, 15S.3 m, RP8.
14. Phormocyrtis turgida (KRASHENINNIKOV). CH12056, P30/fll 29, 158.3 m, RP8.
15. Phormocyrtis cub ensis (RIEDEL AND SANFILIPPO). CH12057, P30/fl1 29, 158.3 rn, RP8.
16. Phormocyrtis cubensis (RIEDEL AND SANFILIPPO). CH12029, P30/fl1 29, 158.3 m, RP8.
17. Phormocyrtis cuben sis (RIEDEL ANDSANFILIPPO). CH12028, P30/fl1 29, 158.3 m, RP8.
IS. Lamptonium fabaefo rme fabaeforme (KRASHENINNIKOV). CH12055, P30/fl1 29, 158.3 m, RP8.
19. Lamptonium faba eforme faba eform e (KRASHENINNIKOV). CH1204S, P30/fl 129, 158.3 m, RP8.
20. Theocorys? phy zella FOREMAN . CH12073, P30/f1129, 158.3 m, RPS.
21. Th eocorys? ph yzella FOREMAN . CH12042, P30/f1129, 158.3 m, RP8.
22. Theocorys? ph yz ella Foreman. CH12058, P30/fl1 29, 158.3 rn, RPS.
23. Theocorys? ph y zella FOREMAN. CH12045, P30/fl1 29, 158.3 m, RP 8.
24. Theocorys? phy zella FOREMAN. CH 12043, P30/fl129 , 158.3 rn, RPS.
S96 c. J. Hollis
Plate 3
Transmitted light micrographs of radiolarians from upper Paleocene and lower Eocene strata, Mead Stream. Scale bars = 100 urn.
1. Haliomma gr. A-K47/4, P30/f451, 99 m, RP5.

2. Haliomma gr. A-L36/3, P30/f533, 166.5 m, RP8.
3. Haliomma gr. A-S28/0, P30/f533, 166.5 rn, RP8.
4. Haliomma gr. A-D29/3, P30/f533, 166.5 m, RP8.
5. Haliomma gr. A-R46/l, P30/f533, 166.5 m, RP8.
6. Periphaena? duplus (KOZLOVA). A-032/2, P30/f451 , 99 m, RP5.
7. Periphaena? duplus (KOZLOVA). A-S31/3, P30/f451, 99 m, RP5.
8. Periphaena? duplus (KOZLOVA). A-G29/1, P30/f451 , 99 m, RP5.
9. Middourium regulare (BORISSENKO). A-W53/0, P30/fl076, 136.9 m, RP7.
10. Dorcadospyris foveolata (EHRENBERG). A-L49/3, P30/f527, 108.2 rn, RP6.
11. Dorcadospyris foveolata (EHRENBERG). A-P42/0, P30/f527, 108.2 m, RP6.
12. Dorcadospyris confluens (EHRENBERG). A-W50/1, P30/f527, 108.2 m, RP6.
13. Lophophaena mugaica (GRIGORJEVA). A-B42/l, P30/f527, 108.2 m, RP6.
14. Corythomelissa adunca (SANFILIPPO AND RIEDEL). A-?, P30/f451, 99 m, RP5.
15. Corythomelissa sp. A. B-F36/0, P30/f527, 108.2 m, RP6.
16. Buryella granulata (PETRUSHEVSKAYA). D-P34/O, P30/f451 , 99 m, RP5.
17. Buryellaforemanae (PETRUSHEVSKAYA). B-U42/O, P30/f527, 108.2 m, RP6.
18. Buryella tetradica FOREMAN. A-F48/0, P30/fl076, 136.9 m, RP7.
19. Buryella tetradica FOREMAN. D-P32/0, P30/f451, 99 m, RP5.
20. Buryella pentadica FOREMAN. C-034/2, P30/f527, 108.2 m, RP6.
21. Pterocodon poculum (NISHIMURA). A-047/2, P30/flOn, 136.6 m, RP7.
22. Pterocodon poculum (NISHIMURA). A-L37/3, P30/fl076, 136.9 m, RP7.
23. Theocorys? aff. phyzella FOREMAN. C-G39/0, P30/f452, 110.8 m, RP6.
24. Theocorys? aff. phyzella FOREMAN. C-E41/3, P30/f452, 110.8 m, RP6.
25. Theocorys? ct. phyrella FOREMAN. D-U29/3, P30/f452, 110.8 m, RP6.
26. Theocorys? phyzella FOREMAN. B-V33/l, P30/flOn, 136.6 m, RP7.
27. Theocorys? phyzella FOREMAN. A-H34/4, P30/flon, 136.6 m, RP7.
S98 c. J. Hollis
DOl 1O.1007/s00015-006-0606-1
Upper Permian to Middle Jurassic radiolarian assemblages of

Busuanga and surrounding islands, Palawan, Philippines
EDANJARLO 1. MARQUEZ1,2, JONATHAN C. AITCHISON 2 & LAWRENCE R. ZAMORAS 3
Key words: North Palawan Block, radiolarians, biostratigraphy, Triassic, Permian, Jurassic, East Asian accretion
ABSTRACT
The North Palawan Block is regarded as the southernmost continuation of a mian, Lower Jurassic, and Middle Jurassic assemblages were also found. The
Late Mesozoic accretionary complex, which developed along the length of the presence of benthonic and planktonic foraminifers at some localities indicates
East Asian margin. It records a long (up to - 100 My) period of pelagic depo- that portions of the Liminangcong Formation were deposited in environments
sition on an oceanic plate from Late Permian to Late Jurassic when subduction above the carbonate compensation depth (CCD). Manganese deposits found
resulting in the disappearance of the plate by the Early Cretaceous began. in some areas (e.g. Busuanga and Dabatonay) suggest that parts of the deposi-
Subduction-accretion resulted in the development of three lithotectonic belts. tional area experienced a very low average sedimentation rate. Examination of
From thirteen localities within these belts, radiolarian investigations yielded the North Palawan accretionary complex reveals the ghosted history of the
173 species belonging to 92 genera and 45 families. Most of the samples con- Izanagi Plate and constrains the timing of subduction beneath the eastern mar-
tain Middle to Upper Triassic faunas. Several sections containing upper Per- gin of Asia.
Introduction
Cenozoic geology of the North Palawan Block. The Calamian
Palawan is a NE-SW trending ridge located between the South Island Group is an integral part of this block and has been the
China Sea and the Sulu Sea. The northern part of Palawan, to- subject of such investigations (Isozaki et al. 1988; Cheng 1989,
gether with the Calamian Island Group, Cuyo Island Group, 1992; Tumanda et al. 1990; Tumanda 1991a, 1991b, 1994; Yeh
Reed Bank, Spratly Islands, Dangerous Grounds, western 1992; Yeh and Cheng 1996a, 1996b, 1998; Tumanda-Mateer et
Mindoro, Tablas and northwest Panay, is interpreted as a mi- al. 1996; Zamoras & Matsuoka 2000, 2001 and Zamoras 2001).
crocontinent, called the North Palawan Block (NPB) (Figure These studies produced radiolarian biostratigraphic zonations
1). The block is inferred to have rifted from the eastern margin of the Middle Permian to Jurassic chert sequences. Zamoras
of Eurasia as a result of the opening of the South China Sea at (2001) and Zamoras & Matsuoka (2001) established that the
around 32 Ma (Hamilton 1979; Taylor & Hayes 1980; Hol- chert-clastic succession youngs from north to south, and dis-
loway 1982; Briais et al. 1993; Almasco et al. 2000) and collided criminated three different belts - the Northern Busuanga
with the Philippine mobile belt during the middle Miocene (NBB), Middle Busuanga (MBB) and Southern Busuanga
(Hall 2002). Several investigations (e.g., Isozaki et al. 1988; Ko- (SBB) belts. They reported that the NBB accreted to the
jima 1989; Faure & Ishida 1990) considered the NPB as the Asian margin in the late Middle Jurassic, whereas, the MBB
southernmost continuation of the accretionary complex along and SBB accreted in Late Jurassic and Early Cretaceous, re-
the Eurasian margin that extends southwards from the Russian spectively.
Far East, through NE China, NE and SW Japan and the The present research was undertaken in order to attempt
Ryukyu islands. to locate the Permo-Triassic boundary in the region and to ex-
In the last 30 years, radiolarian research has contributed pand the knowledge of the Triassic radiolarians in Southeast
tremendously in the deciphering and understanding of the pre- Asia, in particular those from the Calamian Island Group, in-
I Department of Physical Sciences and Mathematics, College of Arts and Sciences, University of the Philippines-Manila, Padre Faura St., Errnita, Manila,
Philippines. Email: [email protected];[email protected]
2 Department of Earth Sciences, The University of Hong Kong, Pokfulam Road, Hong Kong SAR, China. Email: [email protected]
3 Graduate School, University of the East, Manila, 1008 Philippines
Upper Permian - Middle Jurassic radiolarians, Philippines S101

120"E
o
~ 15 km
Fig.1. Geologic map of Busuanga and surround-

ing islands modified after Zamoras (2001). SBB =
<:r Sampling locat ion Southern Busuanga Belt, MBB = Middle Busuan-
ga Belt, NBB = Northern Busuanga Belt. Loca-
' SO
l'hilippinc ""- Fault
Stt,d.v un·u••• • • ~:n:~~ Sro
tions of the sections where the cherts belonging to
" Belt boun dary
the Liminangcong Formation were collected are
indicated. Numbers refer to the following areas:
[J)] Guin lo Formation (clast ic unit s) 1. Calauit-Illultuk Bay, 2. Dabatonay, 3. Mapa-
D Liminangcong Form at ion (chert unit s) dolo, 4. Dipuyay-Decalangwang River, 5. Lusong,
6. Marily, 7. Mayanpayan, 8. Dimanglet, 9. Road
SO
~ Lim esto ne unit s
cut, township of Coron, 10. Calindo, 11.Sitio Tingil,
12. Mabudyen, 13. Mabintangin River.
eluding Busuanga and surrounding islands. Results presented prising the islands together with the north Palawan mainland
herein provide information from many new localities, which to be a subduction-related complex and named this suite the
complement the previous works in the description of Upper North Palawan Complex. Faure & Ishida (1990) considered
Permian and Middle Jurassic radiolarians. The data presented these rocks to be olistostromes. Zamoras (2001) and Zamoras
herein help to constrain the age of the oceanic materials ac- & Matsuoka (2001) referred to the accretionary complex on
creted along the eastern margin of Eurasia during the Late Busuanga as the Malampaya Sound Group.
Mesozoic. Several investigators considered the chert as part of the
Liminancong Formation (Hashimoto & Sato 1973; Bureau of
Mines and Geosciences 1984; Wolfart et al. 1986; Zamoras
Geologic Background
2001; Zamoras & Matsuoka 2001) and assigned various ages:
Busuanga is the largest island in the Calamian Island Group Middle Triassic (Hashimoto & Sato 1973; Fontaine 1979; Wol-
(Figure 1). As with the rest of the group, the island is part of a fart 1984); Lower to Middle Jurassic (Bureau of Mines and
Late Mesozoic subduction complex that extended from the Geosciences 1984) and Middle-Upper Permian to Upper
Russian Far East southwards at least as far as the north Jurassic (Wolfart et al. 1986; Isozaki et al. 1988; Tumanda
Palawan mainland. The subduction complex mainly consists of 1991a; 1991b; Zamoras 2001; Zamoras and Matsuoka 2001).
accreted limestones and cherts intercalated with trench-fill Japan International Cooperation Agency-Metal Mining
elastic sediments. The location of the sampling sites and a geo- Agency of Japan (1990) subdivided the chert into Liminang-
logic map of the study area are given in Figure 1. cong Chert (Triassic) and Busuanga Chert (Jurassic). Wolfart
Various authors have proposed different stratigraphic sub- et al. (1986) adopted the name Bacuit Formation for the Per-
divisions. Isozaki et al. (1988) interpreted the rock suite com- mian sequence of chert, elastic rocks and limestone.
S102 E. J. Marquez, J. C. Aitchison & L. R. Zamoras

Assemblage Zanes
after Kuwahara, K., Yao,A. Interval Zones
and Yamoklta, S.(1998) after Tumanda,F. (1991)
Rhaetian Livarel/a
UPPER Norian
TRIASSIC Capnodoce
Carnian Capnuchosphaera
Muelferitortiscochfeata
Ladinian tnossocorooe deweveri
MIDDLE Pseudostylosphaera japonico
TRIASSIC Anisian Hozmadia altipedaria
Pactarentinia kaikei
Spathian
LOWER Narnmalian
TRIASSIC
Griesbachian
Neoalbaillella optima
UPPER
Changxingian NeoaJbaillelJa ornithoformis
PERMIAN
Longtanian
FoJlicucullus charveri-Albaillellayamakitai Fig. 2. Permian and Triassic radiolarian zonation
Fol/icucullus ventricosus-Foflicucullus scholosticus schemes used inthis study.
The limestone is distributed sporadically. Based on fora- nas were recovered from the chert on Busuanga and surround-
minifers, algae, conodonts, Porifera and Cnidaria (Fontaine ing islands (Calauit, Dabatonay, Mapadolo, Lusong, Marily,
1979; Hashimoto et al. 1980; Fontaine et al. 1983,1986; Wol- Mayanpayan and Dimanglet). Foraminifers were also noted in
fart et al. 1986), they range from Middle Permian to Upper samples from Mabudyen and on the islands of Marily, Diman-
Jurassic. The Bureau of Mines and Geosciences (1984) dis- glet, Mapadolo and Calauit. The conodont Epigondollela sp.
criminated two limestone units: the Malajon Limestone was found in one sample collected along the Concepcion-
(Upper Triassic) and the Coron Formation (Upper Triassic Dipuyay road on Busuanga.
to Lower Jurassic). Kiessling & Flligel (2000) described the In order to establish the age of the radiolarian assemblages,
limestone outcrops on and around Busuanga, assigning them we used the zonation schemes proposed by Kuwahara et al.
to the Coron Formation. They interpreted the carbonates as (1998) for the Upper Permian, and by Tumanda (1991a,
reefs (Malajon) and carbonate platforms (Kalampisanan, 1991b) for the Triassic (Figure 2). The Jurassic biostratigraphic
Busuanga and Coron). ranges of the radiolarian species discussed in this study have
On geologic maps prepared by the Bureau of Mines and been determined from synthesis of occurrences recorded in the
Geosciences, clastic rocks are separated into the King Ranch literature.
and Liminangcong formations. Several workers used the term In the Mabintangin River section (sample location 13),
Guinlo Formation for the clastic succession (Hashimoto & Sato three late Permian zones have been recognized. The presence
1973; Zamoras 2001; Zamoras & Matsuoka 2001). Wolfart et al. of Albaillella cavitata KUWAHARA was used to delineate the
(1986) subdivided the clastic rocks into the Coron Formation upper Longtanian interval in the Liminangcong Formation. A.
(Triassic) and the Guinlo Formation (Jurassic). Stratigraphic cavitata KUWAHARA is a common species occurring in the Fol-
assignments vary from Middle Jurassic (Tumanda 1991a, licucullus charveti-Albaillella yamakitai assemblage zone
1991b), Middle Jurassic to Upper Jurassic (Tumanda-Mateer et (Kuwahara et al. 1998). Most of the Mabintangin samples also
al. 1996) and Middle Jurassic to Lower Cretaceous (Zamoras contain radiolarian assemblages belonging to the succeeding
2001; Zamoras & Matsuoka 2001). The reported ages of the Neoalbaillella ornithoformis assemblage zone. Follicucullus sp.
clastic rocks were established based on radiolarians. d. F. charveti CARIDROIT & DE WEVER, Foremanhelena
Zamoras (2001) and Zamoras &Matsuoka (2001) intro- musashiensis (SASHIDA & TONlsHI), Albaillella? protolevis
duced the name Bicatan melange for the minor melange bod- KUWAHARA and Nazarovella gracilis DE WEVER & CARIDROIT
ies units they mapped at Buyod Point, Bicatan Peninsula. They are some of the species found in these samples.
recognized two types; limestone-basalt-chert melange and F. musashiensis (SASHIDA & TONISHI) and Albaillella trian-
sandstone-chert-mudstone melange. gularis ISHIGA, KITO & IMOTO are found in one sample. F.
musashiensis (SASHIDA & TONISHI) is a common species of the
Neoalbaillella ornithoformis assemblage zone and the Neoal-
Radiolarian biostratigraphy
baillella optima assemblage zone whereas Albaillella triangu-
Ninety-four samples yielding identifiable radiolarians comprise laris ISHIGA, KITO & IMOTO, considering its occurrence, be-
part of a suite of 261 samples collected from the Calamian longs to the Neoalbaillella optima zone. Based on this, the sam-
Group of Islands, Palawan. Moderately to well-preserved fau- ple is assigned to the Neoalbaillella optima zone.

Table 1. Radiolarian assemblages from the Liminangcong Form ation samples. Th e samples were collected from the following sections: Dabatonay and Mapad o-
10.Calauit- Illultuk Bay and Dipuyay-Decalangwang River.
Sam ple Locatio n
Species
_A:_c q~.I~~.sp'h{]_e."-C1 .(? J ,!,:q.c.k ( K_q ~!J_R..~ _ ~9.~~~.~ ~ ..~..,

:A..':.t;.~':!!!~~!!i!sp~.a.er.q__spP-._..-. ..x _..~ . ~ _ .x. __
~ ~ .: ~.. ,._~ I .. __. ~. 1 x I I. X . X ;.. ~ ; x ! ! x._•.~. : x __~-. ~ ~.
!!1!_~~':!9:'J.!~"-'_~PP:. .. _..
·C:ap~.'!.chpSP~_C1~~"_ ~~~'~\I.e!:_~ ~9:Z_~ ~_ .&..,~.9.~TL~R.. _ e~~~.d ...B_~.()~E ...
Capnu chosph aera__missionensis c:ORDEY
~~~/~':.f!'!.~a.__
~ p_~
~C;ru.~t!/~" .sP~ __
,c.ryp~p.s.tep~{1_,!ic!~Il.~ , ~().~11.ige~~ p~ ~ !!.~ ~<;=A
.Ept/,!gill'!!_ '!'_~ '}fre.c!.i.p~.l\1r:rR.lC::A_
~p.lit'g!'!"!:I1.a:~~e.koi .~9?~R_ ~.T A.L: .
~P.~!~gt~.'!!.~P- . ..~ x
J!~~~.~.e.I!t:Y':.~!Il_~~_.sP~ _
H~/~o"!rna ~p .
.1j(~t!~.~ ~P~.C1f!~". ~p!n~.t~~.C1 _ (N"t\K~~~.~9..~ .~I.S_~_I~lJ_~) ..
lft?z.~{]_t!~~t~~L~'!!a.t~_.P.~.M.I.T.~_~t\._:t<;Q~':.l~ .~_ .~lg.s.~~~ ... .x
!!.o.~'!!:~tjJ~ 'J!~!!.'!~'!:.~~ ~:S.~~9 ~. ~~~Ii.~~~~) : x...
!!,~;~~~,:!!.,!: spi"Jfe.':.a., ~ygIY.~ ~.~ .
/(o.;"~.C!.f:Fa. ~pp·
.I,ac!Jn,f!.c.q_'!lP.ejt:lpo..t!.i~~..CN.,:'~~~.~O _~. ,N.:!.S}I.J~l}_~.A)
L.'!~~!.~9~'!.'!1Pf! .~p.
9~!!.I!~fO."gusJn.t}~q~isp.!t!.os.llS.. ~lJ,~ !TRI~~ . ~~~lJ-'~· .~ .~ ~~r'-:E.R.
f'.~!'.~t:l!!.~'.~i!J./1J..~p. ...
!,(1r:.':l.~~~!.i~y~t!~",spP...,. ~, . ~. . .X: . I
.Par:.~.~p~t:lgo.fl . ~P.P:. .. . . ; .1 ~ .. .,. ~ ., . ... ..... ~ X
~p.a.r:.t?~.':!/~fJl!.'-!lP!.s..~PP: ..
.!'.~ntC!£!.~~~c.ap!i~.'~.l!:a.i!."~.~ .~.~~~~.~9 .~ .~ I~_"- ~.~lJ.R!tJ , .
f~~.~0:~.t.~~9.c.a.rp.,!sfl!.sif.'!.r:.rt!.~~. P~_~.I:r.~~.~ " . ~...._~ .....~ x x
fe.~.~ q~,~ {'!l!c.q.rp.u.s..t.et':.~~.q'!!.h.lfs..I?lJ,~rr.~ ~~ _ ..
Pentqc.t!n.orbis.kozuri , D.uMIl"~rc:,'\
P ou lp us spp, x. x..
.f':~e.c.?"-l!t:;!!'f),!!~'!.'q .SP :.. ..
/~,~_~ ':J. ~l!~I)·!C!~P~." e.!:t}. ~.<?~p~c:!t}. ~~.~_~.~~g. ~.~.~~!:,.~~.y..~). .. . . . _~ ... .. ; .. ~ ~ . • J:... ~ .x. . ~..: . ~..
f.s.~~qo.s.~'.'o.sp.h.a.~~(1j(1P2t!.!~t:l .(t'lt\.~~~,~ 9 .~..~ ~.~.J:I_I.~.lJ.~.>. . X ~ _.1.. .
fs.e.~r!o.~ry.?o.!pb.'1.~!.~ _t'!':lg.~sP~·'!.(}s.q..19??~~.& .¥9~~~~ ... x ~ X:... .. ~ -.:
.!'.s.e.'!.c!.f!.~ty/o!ph(1~!.~a .",~g~~sp!'!~~_l! :'(~~t ....
.f'~~lf.4'!.~ty(l!sp~~~':.C!.. ~pp=. . .... ,~ ... ,x. . x x..
Pseudostylospha ero timor ensis..~A S H I DA ,&. KAMAl A X: . x
lli~i.~~at.t!,l/qspp. . .x ,. . x x
~q':.lq..sp:
Spi~,C!~~.~":r.~.C!~q.'!!P.e..<;'.'!, '!.!~/a.I,! CN~~S~.~~.~.. ~.I.~.~.I.~l!.~)" , .
~p~~C!!~,~a.s~l!':~ ,!}P!. .~.u.'}g,! rica . ~{)~UR ..
SJX!ngo~.i!icq.'!'!.ige.': .~P: ...
.SP'C!.':.gl!S/~p~q.~i£!i~,!! jap~~i~u."' .lJ'!"~~~.~ E:!<:()..8l:..~.I~~!~~~)
~PC?fJgl!~t.ep~.'!.nJd.i!!.'!i..!.o.!lgisp'~n~~.,!", .~ A.~~~~A . . x ..
Staura corul um .sPP: I . ..x ....
s.la,u.':l!/9t1~hi!flu(!ge/i. .~<:l~~~. ~ ~OS!~~R.. ...~. ;x
~tCl.If~~*?rl~,he~pp... . .x , ..~ , ', . x x .
.S./~lf~9/o.",c;h~. !':.~spi!l.o.~", {~()ZlJ.~.~..~9~.!~.~~) .x
.Tiborelia florida .(NAKASEKO.& N ISHI>AU RA) .
:!ibu,':.e./!a..magfliclen.ta{q DY,M~l"~C:A ~ . ~.O~~~..~ MgsTLE.~ .
Tioo!ellas p,
Triassocampecoronota..B.~c:J ~ I .. ~ I .. .X. .
T,.i~,s '?, c.af71~.~e"~·t:ve~,( .~.~.'0~~.~9..~. ,~!.~ ,':f~.~Y.~) . X I I
l~i~:!.,?£Cl!!'P.~..~ PP~ . . •.. ~ .. I X
,!r!'!~~~{1,!,p_l!Jr/~~~." fJ~.n.s.is . TlJ~t~.N.[)!.' .

Trilonche sp..cf . T . c~i~kensis (SASH.I.DA.8<.IGO)
Trilo.l1che spp. x
...
~
... -e -e ..... ..... .. ..
... ...
e- -e -e ... ... ... -e ... ... ... ... ... ...
"i ~ "i ..l
;l ;l ;l ;l
'i ;l ~ ~ ~ -l -l
~ ~
'i U ~.:.e . ..l ..l

~ ~ ~ ~ ~ . ~ ~
~
~ ~ ~ ~ ~
1 ~ ·r ofe . 1 ....~
~ ~
-l -l ..l ..l
Range
C
-e ~ ~ ~
..l
~
C
~ ~ .
..l
~ -t ..l
~ ~
..l
~
~ : ~ -t ;l
c
J,
«c ,. c
c. C C C C C C C C '
« « -e ~,; « « -e -e
C C C C C C ..l
« « z -e « C C C
-e « < -e -e « -e
c c -e «
~ «,; ,; «,; «,;
Co
«
,; ,; ,; ,; =, ,; ,; ,; ,; =,
,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,;
=
S104 E. J. Marquez, J. C. Aitchi son & L. R. Zamoras

Table 2. Radiolarians occurring at Dipuyay-Decalangwang River.
Sample Location Dipuyay-Decalangwang River (sample location 4)
Species
"
~Q..c
To.
'"
<: '"
r-
<: <:
ee
<: '"
<:
<:
<:
;;;
<: '"
<: '"
-e
'" <:'" r-<:
<: <: <:
ee
<: '" :; <: '"
<: <: '"
. . . . .. .. . '" '"
<: <: '"
<: '"
-e
r-. ee ;;:: '" 00 <:
.e e= '"'"'" '"'"'" '"'"'" '"'" '"'" '"'"'" '"'" '"'"'" '"'"'" '"'"'" '"'"'" '"'"'" '"'"'" '"'"'" '"'"'" '"'" '"'" '"'" '"'" '"'" '"'" '"'" '"'" '"'" '"'"'" '"'"'" '"'"'" '"'"'" '"'"'" '"'"'"
'" '" <: <: <: <: <: <: <: <: <: <:
rfJ;Z
<: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <: <:
'" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '" '"
Acanthosphaera (?) mocki KOZUR & MOSTLER x x
Archaeocenosphaeraspp. x x x x x x x x x x x x x x x x x x x x x x x x x. x x x x
Cryptamphorella sp.
Cryptostephanidium cornigerum DUMITRICA
Eptingiwn manfredi DUMITRICA x x X x x x x x x x x x x x x x x x x x x x x
Eptingium rarnovsi KOZUR ET AL.
Eptingium spp. x x
Haliomma spp.
Hexalonche sp.
Hindeosphaero spinulosa (NAKASEKO & NISHIMURA) . x
Hozmadia reticulata DUMITRICA, KOZUR & MOSTLER X X X
Hozmadia rotunda (NAKASEKO & NISHIMURA) x x x x
Hozrnadia sp. cf. H. reticula/a DUMITRICA, KOZUR &
MOSTLER
Hozmadia spinifera SUGIYAMA
Hozmadia spp. x x x x x x x x x x x x x x
Katorella bifurcata KOZUR & MOSTLER
Ladinacampejaponica (NAKASEKO & NISHIMURA)
Ladinocarnpe_sp.
Muelleritortis sp.
Napora spp.
Nassellariagen. et. sp. indet,
Pantanellium sp.
Pararuesticyrtium spp. x x x x x x x x x x x x X
Parasepsagon spp. x x x x x x x
Parasepsagon variab/lis (NAKASEKO & NISHIMURA)
Pentactinocapsa awaensis (NAKASEKO & NISHIMURA) x x x x x x x x x x x x x x
Pent(Jctinocarpus fusiformis DUMITRICA x X x x x x x x x x
Pentactinocarpus tetracanthus DUMITRICA
Pentaspongodiscus spp. x x x
Plajkerium abboti PESSAGNO
Poulpus spp. x
Pseudosty/osphaera compacta (NAKASEKO & NISHIMURA) x x x x x x x x x x x x x x x x x x x x
Pseudostylo~phaera,goricanae KOZUR ET AL.
Pseudostylosphaerajaponica (NAKASEKO & NISHIMURA) X X X X X X X X X X X X X X X X x
Pseudostylosphaera longispinosa KOZUR & MOSTLER x x x x x x x x x x x x x
Pseudostylosphaeraspp. x x x x x x x x x x x x x x x x x x
Pseudostylosphaera timorensis SASHIDA & MMATA x x x
Rikivatella spp.
Sarla spp. x
Sepsagon sp.
Spinotriassocampe annu/ata (NAKASEKO & NISHIMURA) X X x
Spongostephanidium japonicum (NAKASEKO &
NISHIMURA) x X x
Stauracontium spp. x
Stauf'olonche spp. x x x
Staurolonche trispinosa (KOZUR & MOSTLER) x x x x x
Thaisphaera spp.
Tiborellaflorida (NAKASEKO & NISHIMURA)
Tiborella sp. cf. T anisica KOZUR & MOSTLER
riborella spp. x x x
Triassistephanidium laticome DUMITRICA
Triassocampe campcmilis (KOZUR & MOSTLER)
Triassocampe coronata BRAGIN x x x x x x x x x x x x x x x x x x x
Triassoeampedeweveri (NAKASEKO & NISHIMURA) X X X X X X X X X X
Triassocampe sea/oris DUMITRICA, KOZUR & MOSTLER
Triassoeampe spp. x x x x x x x x x x x x x X X
Trilonche spp x x X x x x x
.... .... .... .... .... .... .... .... .... .... .... ..... ..... ..... ..... ..... .... .... .... .... .... .... .... .... .... ..... ..... .... .... ....
..l ..l ..l ..l
Range
"i "i "i "i "i ..lJ, ..lJ, ..lJ, ..lJ, "J: ..l~ ..lJ, "i= "i= "i "i "J: "J: "i "i ..lJ, ..lJ, ..lJ, ..l=J, ..l=J, "i "i "i ..lJ,
=========== =
0( 0( 0( 0( 0( 0( 0( 0(
~
0(
0(
0(
=========
0( 0( 0( 0( 0( 0(
0( 0(
====
0( 0( 0( 0( 0( 0( 0( 0( 0( 0( 0(
,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,; ,;
Upper Permian - Middle Jurassic radiolarians. Philippines Sl05

T riassic ra d io la ria ns were obse rved from samples co llecte d lith ology present is recr yst allized th inl y bedded greenis h-
from severa l localities. A number of spe cies is common in th e gray che rt. Only two of these sa m ples (H K U- PLWN02031904
Pseudostylosphaera japonica int erval zo ne (T um and a 1991a , and HKU-PLWN02031905 ) yie lde d ide ntifiable radiolarian s
1991b ). T his zone, consid ered as late Anisian- early Ladinian in (Table 1). The assemblage present is upper Anisian-Ladinian .
age, is defined by the suc cessive low est occur rences of Hin-
deosphaera sp in ulosa (NAKASEKO & N ISHIMURA) and
Dab otona y (samp le location 2)
Eptingi um gro up at th e ba se of the zone and Triassocampe
deweveri (NAKASEKO & N ISHIMU RA) at th e top. Some of th e Samples HKU-PLWN02031109 to HKU-PLWN02031120 we re
faunas present in our assemblage , which are comm only found collect ed from a coastal sec tio n (12°6.186'N/119°50.758'E) com-
in th is zo ne, include Hindeosphaera spinulosa (NAKASEKO & posed of steeply dipping lam inat ed to thinly bedded red che rt.
NISHIMURA), Eptingium spp., Pseudostylosphaera mag- The beds are co mplex ly fo lded and faulted . Some parts o f th e
nisp inosa YEH, Tib orella fl orida (NAKASEKO & NISHIMURA), sectio n contain manganese nodules. O ccurrences of radi olar i-
Katorella bifurcata KOZUR & MOSTLER, Pentactinocarp us tetra- an s in the area are shown on T abl e 1. Samples HKU-PLWN
canthus D UMITRICA, Penta ctinorbis k ozuri D UMITRICA, Stau- 0203 1109 to HKU-PLWN02031112, HKU-PLWN0203 1114 ,
rolon che trispinosa (KOZUR & M ostler), Parasepsagon vari- HKU-PLWN02031116 and HKU-PLWN02031117 contain
abilis (NAKASEKO & NISHIM URA) and Tiborella magnidentata upper Anisian-Ladinian radiol ari ans.
DUMITRICA, KOZUR & MOSTLER.
Th e Triassocampe deweveri interval zone (Tumanda 1991a,
Mapadolo (sample location 3)
1991b) is represented by samples from Dipuyay-Decalang-
wan g R iver (sample location 4) and Calind o (sample loc ation Samples HKU-PLWN02031101 to HKU-PLWN020311 08
10). The following species common to thi s zone were found in were co llec ted from a coast al sectio n (12°06.849'N/I19°
o ur sa mples: Triassocampe deweveri (NAKASEKO & NISHIMU- 52.006'E) of co mple xly fo lde d, steeply dipping laminat ed to
RA), Spo ngostep hanidium japonicum (NAKASEKO & NISHIMU- th in ly bedded chert. The chert is re cr ystallized. Three sa mples
RA), Oertlispongus inaequispinosus D UMITRICA, KOZUR & yie lde d rare radiolarian s. Samples HKU -PLWN02031101 and
MOSTLER, Staurolonche trisp inosa (KOZUR & MOSTLER), Ho z- HK U -PLWN0 20311 03 co ntai n upper Anisian-Lad ini an and
m adia rotunda (N AKASEKO & NISHIMURA), Pentactin orb is upper Norian-Rhaetian asse mblages, respecti vel y. Sample
k ozuri D UMITRICA, Pseudostylosph aera jap on ica (N AKASEKO HKU-PLWN0 203ll02 yie lde d Ca rn ia n radiolarians (T a ble I).
& NISHIMURA), Pseudostylosph aera com pacta (NAKASEKO &
NISHIMURA), Ep tingium m anfr edi D UMITRICA, Hindeosphaera
D ipu yay-Decalangwan g River (sam ple locati on 4)
spi nulosa (NAKASEKO & NISHIMURA), Cryptostephanidium
cornigeru m D UMITRICA a nd Parasepsagon variabilis Samples HKU-PLWN02032007 to HKU-PLWN02032060 were
(NAKASEKO & NISHIMURA). collect ed from a section on th e Conce pcion-Di puyay road
T he Ladinian to probably lowe r Ca rnian inte rval (Muelleri- (12°0l.848' N/ll 9°59.792'E). Th e lith ology is predominantl y chert
tortis cochleata interval zone ) is rep re sented by sample HKU- with minor siliceous mudston e. Beds strike - 030 and dip steeply
PLWN 02031402. This sample is characterized by the presence to th e NW . Three radiolarian assemblages are recognized. The
of the diagnostic species Muelleritortis cochleata (NAKASEKO & assemblages are diverse and contain well-preserved radiolarians.
NISHIMURA). The Carnian and upper Ca rn ian-middle Norian Samples HKU-PLWN02032008 to HKU-PLWN02032010,
were established based on the pr esence of Capnuchosphaera HKU-PLWN020320l6, HKU·PLWN02032018 to HKU-PLWN
de weveri KOZUR & MOSTLER, eme nd. BLOME and Capn od oce 02032021, HKU-PLWN02032023 to HKU-PLWN02032032, HKU-
anapetes D E W EVER, respectivel y. Based on the presence of Li- PLWN02032034, HKU-PLWN02032036 to HKU-PLWN
varella spp., th e upper Norian-Rhaet ian was identified. 02032039, HKU-PLWN02032041, HKU-PLWN02032042, HKU-
Jurassic radiolarians we re a lso o bserved. D iagnost ic PLWN02032045 to HKU-PLWN02032052, HKU-PLWN
species are very rare. Howe ver, based o n th e reported ran ges 02032054, HKU-PLWN02032055 and HKU-PLWN02032058 to
of so me spec ies, th e samples are ass igne d Jurassic ages. These HKU-PLWN02032060 contain upper A nisia n-La dinian rad iolar-
include Kat roma westerm anni WHALEN & CARTER (upper ia ns. Samples HKU-PLWN02032033, HKU-PLWN02032035,
Sin emurian), Plessus sp. aff. P. aptus YEH (lower Sinemurian ) HKU-PLWN02032040, HKU-PLWN02032043 , HKU-PLWN
and Tran shsuum m edium (U ni ta ry A ssociation Zones 1-7 , 02032044 and HKU-PLWN02032057 yielded upper Anisian -
early-m idd le A alenian to late Bathonian -e arl y Callov ian) . lower Ladinian radiolari an s. Sample HKU-PLWN02032022 con -
Followin g is th e de scription of th e localities in vestigated: tains Ladinian radiolari an s. Tabl es 1 and 2 show radiolarian oc-
currences in the ar ea .
Calauit-Illultuk Bay (sam ple location 1)

Luso ng (sam p le location 5)
Samples HKU-PLWN02031901 to HKU-PLWN02031908
were co llected from two coastal se ctions located at Samples HKU-PLWN02031802 to HKU-PLWN02031 805
12°16.078'N/ 119°52.080' E and 12°17.484'NIl 19°51.960'E. Th e were collected from th e western coa st of Lusong (11 °58.899'NI

Table 3. Radiolarian assemblages within the Liminangcong Formation. The samples were collected from Lusong, Marily, Mayanpayan and Dimanglet.
Sample Location
Species
Acanthosphaera? sp.
Archaeocenosphaera spp. x x x x x x x x
Archaeodictyomitra spp. x
Archicapsa sp.
Betraccium deweveri PESSAGNO & BLOME x x
Betraccium mac/earni PESSAGNO & BLOME
Betraccium sp.cf. B. inornatum BLOME x
Busuangat'') sp. x
Canoptum spp. x x
Capnodoce anapetes DE WEVER
Capnodoce e~tent(lBLOME x
Capnuchosphaera lea DE WEVER x x x
Complexapora sp. x
Corum sp. cf. C. de/gada SUGIYAMA x
Eptingium sp. x
Gorgansium sp. x x x
Haeckelicyrtium'! sp. x
Hagiastrum sp.? x
Haliomma sp.
Japonocampe sp. x
Katroma coliforme (HORi)
Katroma sp. x x
Katroma sp. cf. K. irvingi WHALEN & CARTER
Katroma westerrnanni WHALEN & CARTER
Livarella magna TEKIN x x x
Minocapsa sp.
Muelleritortis cochleata (NAKASEKO & NISHIMURA)
Napora sp.
Pachus multinodosus TEKIN
Pantanellium sp. x
Pantanelliurn ultrasincerum BLOME x
Parahsuum simplum Y AO x
Parahsuum spp. x x x
Pararuesticyrtium sp.
Paroertlispongus sp. x x
Paronaella sp. x
Parvicingula sp.
Pentaspongodiscus sp.
P1eesus sp. cf. P. aptus YEH
Podobursa sp. x
Praemesosaturnalis huxleyensis (CARTER) x
Praemesosaturnalis rugosus (YEH)
Praemesosaturnalis sandspitense (BLOME)
Praemesosaturnalis sp. cf. P.finchi (PESSAGNO) x x
Pseudostylosphaera compacta (NAKASEKO & NISHIMURA) x
Pseudostylosphaerajaponica (NAKASEKO & NISHIMURA) x
Pseudostylosphaera spp.
Sarla sp.
Sethocapsa spp. x x x
Silicarmiger sp.
Spongostephanidiumjaponicum (NAKASEKO & NISHIMURA) x
Staurolonche spp. x x
Stichocapsa spp. x x x x x
Triassocampe deweveri (NAKASEKO & NISHIMURA)
Trilonche sp. x
Tripocyc/ia sp.
Vinassaspongus subsphaericus KOZUR & MOSTLER
Williriedellum sp. x
Xipotheca sp, cf. X. rugosa BRAGIN, emend. TEKIN x
. .
o
Z
0
z ~.:
=U
.
=
.00
Range
== . Ci3~
uu~ "
..l
== ::i ::i ::i ::i ::i ::i ::i ::i
::i ==
Upper Permian - Middle Jurassic radiolarians, Philippines SI07

Table 4. Radiolarians occurring at Sitio Tingil, Mabudyen, a road cut 35 km east of the township of Coron (11°59.625'N/120013.637'E) and Calindo.
... =
Sample Location
--=~
= .'" '0= ~'"

~ <= <0 .>.= =
~
:::
<0
.
<0 <0
c. <=
~ ..2'"
= ..
~
<0 .S
'0
-=
S
...
~ ,S Q, G\ .S ..2'"
.S ..2
~ c.
. ..2
~ ""
::; ~
c.
~ !=
";j
u ~
c.
!
s
!= ..'" ..=
'0 <0
!=
~ U
<0
Species
...
..!! .. :::; <:>
- ...
.... "" ..,:::l~ ..,:::l .....,:::l ..,:::l ..,:::l ..,...~
<:> <:>
...~ ..,
~ "" ~ '~ "' Si :::l :::; ~ -
..,...<:> .....,<:> ..,...<:> ..,...<:> ..,...<:> ..,...<:> ..,...<:> .....,<:>
== Z= ... ... ... ... ... ... ... ... ... ... ... ... ... ...
rIO <:> <:> <:> <:> <:> <:>
Archaeocenosphaera spp. x
Astrocentrus sp. cfA.pulcherKozuR& MOSTLER
Bulboc),rtiumsp:.
Canoptumsp.aff. Cunicum
Canoptumspp.
Capnodocesp:.cf..C .anapetesDEWEvER
Cellurontasp.
OeJlandrec),rtium .carterae.YEH&.CHENG
Eptingiummanfredi DUMITRICA
Eptingium sp.
flexalonchesp.
Hindeosphaeraspinulosa (NAKASEKO & NISHIMURA)
Hozmadia reticulata DUMITRICA, KOZUR & MOSTLER
Hozmadia
" "
rotunda.. (NAKASEKO AND NISHIMURA)
......... ... .............
-".,,,',.,
Hozmadiaspp.
Ladinocampe sp:
Lina.re.siasp:
LivareliamagnaTEK IN
Oertlispongus diacanthus Sugiyama
Paleosaturnalis sp. cf. P dotti (BLOME)
'Pararuesticyrtiumsp,
Parasepsagonsiip.
Parasepsagonvariabilis (NAKASEKO&' NISHIMURA)
f'aronaellasp.
-f~!l._~t}_~~ifl.9E_qp~q__q_~(J~,!§i~
f'entaspongodiscus.spp: x x x
Praemesosalurnalisgracilis(KozUR.&MoSTLER)
Praemesosalurnalis huxleyensis (CARTER)
j>raern~so;~I~~~~li;l1ii~;~;YEH x
Praemesosaturnalis spp. . ........ x
Pseudostylosphaera compacla (NAKASEKO.&' NISHIMURA) x x x x x
Pseudosl)'losphaerajaponica(NAKASEKO&.NISHIMURA) x x x
Pseudostylosphaera longispinosaKOzuR&MOSTLER x x x
['s~ud~sl)'losphaeramagnispinosa. YEH x
Pseudosl)'losphaerasI'I"
Pseudostylosphaera timorensis SASHIDA & KAMATA
Sarlasp.
'Spinotriassocampeannulata (NAKASEKO. &.NISHIMURA)
Spongosilicarmiger sp.
Spongostephanidiumjaponicum.(NAKASEKO & NISHIMURA)
Spongosl)'lustricostalus.KOZUR,.KRAINER.& MOSTLER x
Spumellariagen.et. sp.indet.
Stauracontium__ ~p.
SlaurolonchespI': ..... . .......
Staurolonchetrispinosa (KOZUR&IvI0STLER)
Stichocapsa sP:..
Thaisphaerasp. . ........................
Tfborellaflorida (NAKASEKO & NISHIMURA)
Tiborellasp.
Transhsuum medi~mT AI<EMURA x
Triassocampe coronata BRAGIN x x x
Triassocampedeweveri(NAKASEKO.&.NISHIMU RA )
Triassocampe sPI': x
Trilonche spp. x x x
......c:
..
.
'0 ...
'" ~ Z .::~ .::~ ~ .::~ -:l'" '"
..c:
<0 '0 '0
.'" 'i'" .'" =
'0 '0 '0
Range J, .:.
:p .:. .:. .:. .:.
..
-<= Z:i u= Z:i
<0 0 <0 0 <0 '" 1 Z Z Z
:i :i :i
-<= -<=
:i :i
.'" = .'" = e
'0 J,
-<= -<
:i :i
'0 J,
:p
-< ';'
:i =:l
:i :i
:i
Sl08 E. J. Marquez, J. C. Aitchison & L. R. Zamoras

120000.509'E).
The chert present at this locality is dull red and 02032119, were collected from this roadside section
thinly bedded. Bedding is oriented 030/50NW. Two samples, (11°59.625'N/120°13.63TE). The chert isweathered and thinly
HKU-PLWN02031802 and HKU-PLWN02031803, yielded bedded. Bedding strikes 160 and dips gently to the SW. The
rare upper Carnian-lower Norian radiolarians (Table 3). two samples contain upper Norian-Rhaetian radiolarians
(Table 4).
Marily (sample location 6)
Calindo (sample location 10)
Samples HKU-PLWN02031302 to HKU-PLWN02031305
were collected from a coastal section consisting of reddish Samples HKU-PLWN02032102 to HKU-PLWN02032117
brown chert beds. The section is situated at the SE side of the were collected from a roadside section located at 12°00.939'NI
island, 11°56.848'N/120000.942'E. Bedding strikes 165 and dips 120°15.681'E. The outcrop is composed of uniformly bedded
steeply to the east. Three samples (HKU-PLWN02031302 to reddish brown chert. Bedding strikes 030 and dips gently to
HKU-PLWN02031304) contain Lower Jurassic? (Hettangian- the NW. Upper Anisian-Ladinian radiolarians occur in sam-
Sinemurian or younger) radiolarians. The occurrences of radi- ples HKU-PLWN02032102, HKU-PLWN02032103, HKU-
olarians are shown in Table 3. PLWN02032109, HKU-PLWN02032110 and HKU-PLWN
02032117 whereas Ladinian radiolarians are present in samples
HKU-PLWN02032106 and HKU-PLWN02032112 (Table 4).
Mayanpayan (sample location 7)
A spot sample (HKU-PLWN02032101) was collected
The coastal outcrop where samples HKU-PLWN02031709 to from the top of a small hill (100 m high) located at
HKU-PLWN02031715 were collected is located on the SE side 12°01.477'N/120016.232'E. The sample, a chert, is upper
of the island, 11°58.816'N/120007.382'E. It consists of folded Bajocian to uppermost middle Bathonian? (Table 4).
thinly to medium bedded reddish brown chert. Samples HKU-
PLWN02031709 and HKU-PLWN02031710 contain Lower
Sitio Tingil, 2.5 km SE of Coconogon Point
Jurassic? (Sinemurian or younger) assemblages. Sample HKU-
(sample location 11)
PLWN02031713 contains Sinemurian radiolarians whereas
sample HKU-PLWN02031715 yielded upper Norian-Rhaetian Samples HKU-PLWN02031209 to HKU-PLWN02031220 and
radiolarians (Table 3). HKU-PLWN02031618 to HKU-PLWN02031621 were col-
lected from a coastal section (12°12.570'NI120013.282'E) com-
posed of brown, dark gray to black, thinly bedded to laminated
Dimanglet (sample location 8)
chert. The beds are folded and locally highly sheared. Sample
Samples HKU-PLWN02031401 to HKU-PLWN02031435 and HKU-PLWN0203121O yielded upper Anisian-lower Ladinian
HKU-PLWN02031701 to HKU-PLWN02031708 were collect- radiolarians whereas samples HKU-PLWN02031217 and
ed from a continuous chert section exposed along the SE coast HKU-PLWN02031220 contain upper Norian-Rhaetian and
of the island (12°01.848'NI119°59.792). The reddish brown upper Carnian - upper Norian radiolarians, respectively
chert is thinly to medium bedded. It is highly folded and fault- (Table 4).
ed. Some horizons are massively recrystallized and appear to
be thickly bedded. Four radiolarian assemblages are recog-
Mabudyen, 2 km west of Malawig (sample location 12)
nized. Samples HKU-PLWN0203141O, HKU-PLWN02031413
and HKU-PLWN02031701 yielded upper Carnian-Norian ra- The coastal section (12°11.941'NI120013.447) where samples
diolarians. Ladinian - lower Carnian faunas occur in samples HKU-PLWN02031201 to HKU-PLWN02031207 were col-
HKU-PLWN02031402 and HKU-PLWN02031702 whereas lected consists of thinly bedded black-brown chert. The chert
upper Carnian - lower Norian radiolarians are found in sam- is recystallized. Bedding strikes 030 and dips steeply to the
ples HKU-PLWN02031405, HKU-PLWN02031407 and HKU- NW. Only two samples, HKU-PLWN02031201 and HKU-
PLWN02031408. Samples HKU-PLWN02031409, HKU-PLWN PLWN02031202, contain identifiable forms (Table 4). The as-
02031414, HKU-PLWN20301418 to HKU-PLWN02031420, semblage is upper Norian-Rhaetian.
HKU-PLWN02031425, HKU-PLWN02031427, HKU-PLWN
02031705 and HKU-PLWN02031706 contain upper Norian-
Mabintangin River (sample location 13)
Rhaetian faunas. Lower to Middle Jurassic radiolarians occur
in samples HKU-PLWN02031430 (Sinemurian or younger) Samples HKU-PLWN0203100l to HKU-PLWN02031Ol7
and HKU-PLWN02031707 (Bajocian or younger). Table 3 were collected from a section along the Mabintangin River.
shows the radiolarians occurring in this area. The outcrop is composed of folded thinly to medium bedded
black chert. Twelve samples yielded identifiable Upper Permi-
an radiolarians. Table 5 shows the radiolarian assemblage in
Road cut, 3.5 km east ofthe township ofCoron (sample location 9)
this section.
Two samples, HKU-PLWN02032118 and HKU-PLWN
Upper Permian - Middle Jurassic radiolarians, Philippines Sl09

Table 5. Distribution of Upper Permian radiolar-
ians within the Liminangcong Formation exposed SampleLocation MabtntangtnRlver
along the Mabintangin River. ~ ~,~ N ~ ~ ~ ~ ~;~ ~ ~ ~:~
- ~- 0 Q <:> => <= 0 => <:> 0 ,..... ,...: ~
Q,,CQQQQQQQQQQQQ
Species ~ §:;:;,;;;;;;;;;;;;;::;;;;;; ;;;:;:;;:;-;;

rJJ:Z ;:.;:;:;:;:;:;:;: ;:;:;:;:
?Latentifistulasp.Fsensu KUWAHARA AND Y AO
?Stigmosphaerostylus spp, x
,Albaillella?protolevisKUWAHARA x .x
Albaillella cavitata KUWAHARA
,Albaillella sp, x
,Albaillella triangularis ISHIGA,KJTO & IMOTO x
brokenFollicucullus spp. x
Copicyntra spp.
Copicyntroides? sp. Asensu KU\VAHARA AN])yAO x
Copiellintra?sp.A sensuKuwAHARAAND Yi\O x
peflandrellasp. x
FollicucullusporrectusRuDENKO
pollicucullussp. cfT charvetic:ARIDROIT&DEVVEVER x x x
Follicucullusso. cf,F. scholasticusORMISTON&BOBCOCK x
Foremanhelena musashiensis (SASHIDA & TONISHI) x x
lshigaumnicolasensis TUMANDA x x
lshigaum sp, cf.1. trifustisDE VVEVER & CARIDROIT x
Ishigaum spp. x x x
Latentibifistulasp.cf.LasperspongiosaSAsHIDA & TONISHI x
Latentibifistula spp.
Latentifistulasp. Esensu TUMANDA x
Latentifistula spp. x x x .x x x
~azarovellagracilisDE \VEVER&;CARIDROIT x x
Nazarovellaso. x
Polyfistula sp.
Pseudotormentussp. x
Ruzhencevispongus (?)sp, B sensuTuMANDA x x x
Spumellariagen. et.sp. indet. x x x x x x x x x
Stauraxon F sensu YAO& KUWAHARA x
Stigmosphaerostylussp. cf. S. modesta (SASHIDA &TONISHI) x x
Trilonche sp.cf. T. cimelia(NAZARoV & ORMISTON) x x x x
Trilonche spp, x x x
Range Upper Permian
Summary and Conclusion

A previous study of Busuanga (Zamoras 2001; Zamoras &
This research has expanded the known geographic extent of Matsuoka 2001) recognized three structural divisions: North-
studied sections and has contributed towards refinement of ern Busuanga Belt (NBB), Middle Busuanga Belt (MBB) and
calibration of the Triassic in Northern Palawan. Samples col- Southern Busuanga Belt (SBB). Samples collected during this
lected from several sections on Busuanga and surrounding study were incorporated into the composite stratigraphy of the
small islands yielded 173 species distributed among 92 genera chert-clastic sequence described by Zamoras (2001). The sec-
in 45 families. Most of the samples are Triassic, spanning the tions from Calauit-Illultuk Bay, Sitio Tingil and Mabudyen are
time interval, Anisian to Rhaetian. Four of the sections also from the NBB. The Dabatonay, Mapadolo and Calindo sec-
yield Jurassic faunas; three of which include Lower Jurassic ra- tions lie within the MBB. The rest of the sections are from the
diolarians whereas two sites contain Bajocian-Bathonian? as- SBB. Figure 3 shows the data from this study together with the
semblages. Upper Permian radiolarians were recovered only Triassic data of Zamoras (2001) and Zamoras & Matsuoka
from sample location 13 at the Mabintangin River in Busuan- (2001), and earlier results of Yeh (1992) and Tumanda (1991a,
gao No other Permian outcrops were encountered on the sur- 1991b). In the SBB stratigraphy, Zamoras (2001) noted that
rounding islands investigated during this study. Lowermost the Upper Triassic cherts represent the lower part of the suc-
Triassic (Griesbachian and Nammalian) sections are not docu- cession. However, the inclusion of data from the present study
mented herein and they have not been reported from earlier clearly demonstrates the presence of a Middle Triassic interval
investigations. It remains unclear whether the absence of the in the SBB. Tumanda (1991a, 1991b) also reported an upper
lowermost Triassic outcrops is related to sampling density, Spathian assemblage in Decalangwang.
poor radiolarian preservation or a possible hiatus coinciding The chert-clastic succession in North Palawan documents
with the worldwide latest Permian catastrophic event. almost 100 My of the history of a subducted oceanic plate.

Northern Busuanga Belt Middle Busuanga Belt Southern Busuanga Ben
""' .
CRETACEOUS
YaJaiIglf.'ian
BemBsioan .~ ~ ~
E~8~
~ ~~:;
E ga~
~
f-
a>
~Q
5
c
;::
.,
g
e
R ~~
m oo,"" o ~ E
Ucnft1 E
1IJ
~ ~ iD
R
'" ~ s
~
u.... 8fa c c R
~
~
~
K.mmen~""an N
u- E =->- ~
JURASSIC
Oltoroaafl "'~ ~
s f- _
M ~ j !
E
~
~
'"
c
Ca IfO ~\M
r-t-
z: s ~ g
c
~ ~
!
E
-a'"
~
c 8
~ ~
§ ~ E
~
I- II
BJrhonian
B !
-
E E
~
. IDDLE ;;;- U
JURASSIC a.;oo., Icc
~ '" ~ N ! c ! '0 s mill:-: :~
... f
-a -a'"
c
~ g
~ ~
§ §' G;
~
0.
~
~ t E
~
i? :s;
~~
~
E l' ~
~
;5. c a: ~
~
raa_ E ! ! ! g g>
:l'
• ~
! .'" g
B ~. '"
c 9
" "E
~
~ fi
i ~
PJ!enso.ac.h,ian c '5. :l' <5
j r ~
I
LOWE' ~
E ~;;.
JUIU.SSIC 5.lrtemlJl7l3n "
F
.,g
.," 00 t 8 '"
c ..... " ~ ~
~ !E ~
al
~ ! ;5. ! "C
~
Herra~n '".3 iii

""" ~~:
~ g'"
(i
~
~
.,. ~ 0
~
..... I ~ !l ri ~*i
Rh(3ell~
~ ~¥.
~~ ::..;:
~ '~" ~
*I
..J
~ oOJ"
'5 ?-£~ Q.
~
Nr;JI1(3n "..
~
.0
TJIIASS~ :~'". OJ
I#. 8
~~
~
eam." o <5
~ ~
Wi; ~ ~ ~
~ ~~ ~ ~ ~
• m.<
Ladm~
~~ ' ~. ~~ ~ :~. ;- .'.
f"lIASStC A"""" ~ c
H~ .~ ,,-
"'-
LOWE'
TJHA$SIC _ 'an
Spatman Ii;i;
~
e ll '"
E~
i3- -g~
s
G!i!Jro.,Ii."
mp .[E
coangOnfl'" f¥.i ~" ...~
loogran.ian r&. ~ ... ;~ .go .
g~ D
~~
~ Terrigeneous ClastICS
u.... "'~
PERMWf
Cap(taniM
Wordrt11'l
~
@.1' ."
l~
El Siliceous Mudstone
~
Fig. 3. Stratigraphic ranges of lithologies in the
UlirniM ~ .. a ~
••••• C hert
three accretionary belts of Busuanga as con-
~..
~
1000EPflI.JM KLLPgBf'I8I'1 ' ~ .... o
strained by radiolarian faunas.
This plate traveled from a deep sea environment to a conver- continent/continental fragments to the northwest. He added
gent margin, which developed along the eastern margin of that this spreading episode produced a NNW-directed plate
Asia. However, the paleolatitudes of the block during this motion with the oceanic lithosphere being consumed at a NW-
journey are still under investigation. Kiessling and Fltigel dipping subduction zone (East Asian Subduction Zone;
(2000) suggested that the facies and ages of the carbonates Zamoras 2001) on the southeastern margins of the Indochina
from the islands of Malajon, Kalampisanan, Busuanga and and South China blocks.
Coron do not support a close paleogeographical connection The pelagic sediments in Busuanga and surrounding is-
between the NPB and South China Block. They suggested lands records up to 100 My of deposition. The presence of ben-
that during the Carboniferous and Permian, the NPB was part thonic and planktonic foraminifers at some localities probably
of the Indochina Block from which it separated during the indicates that not all of the chert was deposited below the
Middle Permian. The NPB collided with the South China CCD. Manganese deposits found on Busuanga and Dabatonay
Block during the Late Cretaceous. Paleomagnetic studies car- indicate an oceanic setting affected by a low average sedimen-
ried out by Almasco et al. (2000) on the NPB show that the tation rate. The limestone bodies presently found in fault con-
paleomagnetic directions fail regional fold tests and have al- tact with the chert-clastic succession can be interpreted as shal-
ternating field (AF) demagnetisation features consistent with low water facies or carbonate build-ups on seamounts. The fa-
secondary magnetisation. However, they pointed out that the cies change from cherts through hemipelagic clastics to terrige-
Cretaceous paleolatitude is still comparable to regions of per- nous clastics suggests that the ocean was being influenced by
vasive Cretaceous remagnetisation in the South China border- subduction zone related sedimentation by the Middle to Late
land. They suggested that this may reflect similar rernagneti- Jurassic (Aalenian-Bajocian in the NBB, early-late Bathonian
sation and is consistent with the NPB's proposed South China in the MBB and early-late Tithonian in the SBB).
origin.
Based on the tectonic model of Maruyama et al. (1997),
Zamoras (2001) interpreted the NPB to have been part of the
Acknowledgements
NNW-traveling Izanagi Plate from Late Permian to Late
Jurassic-Early Cretaceous. He suggested that the Middle Car- The authors gratefully acknowledge the financial support of The University of
Hong Kong-Committee on Research and Conference Grants towards this reo
boniferous limestone described by Kiessling & Fltigel (2000)
search. We also thank Dr. Jason R. Ali and Prof. Graciano P. Yumul, Jr. for
probably originated from the Farallon Plate. Holloway (1982) their company and stimulating discussions while we were undertaking our field
noted that during the Middle Jurassic to Late Jurassic an investigations. Valuable suggestions and comments by our reviewers, Dr.
episode of spreading commenced between Australia and a Spela Gorican and Dr. Martial Caridroit are greatly appreciated.

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Palawan, Philippines. ESAP-CCOP Newsletter 6,40-47. TUMANDA-MATEER, F., SASHIDA, K. & IGo, H. 1996: Some Jurassic radiolari-
FONTAINE, H., DAVID, P., PARDEDE, R. & SUWARNA, R. 1983: Marine Jurassic ans from Busuanga Island, Calamian Island Group, Palawan, Philippines.
in Southeast Asia. ESCAP-CCOP Technical Bulletin 16, 1-30. In: NODA, H. & SASHIDA K. (Eds.): Prof. Hisayoshi Igo Commemorative
FONTAINE, H., DEN, N.D. & VACHARD, D. 1986: Discovery of the Permian Volume on Geology and Paleontology of Japan and Southeast Asia,
limestone south of Tara Island in the Calamian Islands, Philippines. 165-192.
ESCAP-CCOP Technical Bulletin 18, 161-166. WOLFART, R. 1984: Stratigraphy of Palawan Island, Philippines. Rep. Federal.
HALL, R. 2002: Cenozoic geological and plate tectonic evolution of SE Asia Inst. Geosci. Nat. Resources, Archive no. 97074,40 p.
and the SW Pacific: computer-based reconstructions, models and anima- WOLFART, R., CEPEK, P. GRAMANN, F., KEMPER, E & PORTH, H. 1986: Stratig-
tions. J. Asian Earth Sci. 20, 353-431. raphy of Palawan Island, Philippines. Newsl. Strat. 16/1,19-48.
HAMILTON, W. 1979: Tectonics of the Indonesian Region. U.S. Geol. Surv. YEH, K.-Y. 1992: Triassic Radiolaria from Uson Island, Philippines. Bull. Natl.
Prof. Paper 1078, 1-338. Mus. Nat. Sci. 3, 51-91.
HASHIMOTO, W. & SATO, T. 1973: Geological structure of North Palawan, and YEH, K.-Y. & CHENG, Y.-N. 1996a: An Upper Triassic (Rhaetian) radiolarian
its bearing on the geological history of the Philippines. In: KOBAYASHI, T. assemblage from Busuanga Island, Philippines. Bull. Natl. Mus. Nat. Sci.
& TORIYAMA, R. (Eds.). Geol. Paleont. Southeast Asia 13, 145-161.
7,1-43.
HASHIMOTO, W., TAKIZAWA, S., BALCE, G.R., ESPIRITU, E.A. & BAURA, CA. YEH, K.-Y. & CHENG, Y.-N. 1996b: Jurassic radiolarians from the northwest
1980: Discovery of Triassic conodonts from Malajon Island Group, coast of Busuanga Island, North Palawan Block, Philippines. Mi-
Palawan Province, the Philippines and its geological significance. Proc. croplaeontology 42/2, 93-124.
Jap. Acad. 56 (Ser. B), 69-73. YEH, K.-Y. & CHENG, Y.-N. 1998: Radiolarians from the Lower Jurassic of
HOLLOWAY, N.H. 1982: North Palawan Block, Philippines - Its relation to the Busuanga Island, Philippines. Bull. Natl. Mus. Nat. Sci. 11, 1--65.
Asian Mainland and role in evolution of South China Sea. Bull. Amer. ZAMORAS, L. 2001: Jurassic-Early Cretaceous accretionary complex in the
Assoc. Petroleum Geol. 66,1355-1383. Calamian Islands, North Palawan Block (Philippines). Ph.D. Thesis, Ni-
ISOZAKI, Y., AMISCARAY, EA. & RILLON, A. 1988: Permian, Triassic and igata Univ., 120 p.
Jurassic bedded radiolarian cherts in North Palawan Block, Philippines: ZAMORAS, L.R. & MATSUOKA, A. 2000: Early Late Jurassic radiolarians from
Evidence of late Mesozoic subduction-accretion. In: ICHIKAWA, K. (Ed.): the clastic unit in Busuanga Island, North Palawan, Philippines. Sci. Rep.,
Report No.3 of the IGCP Project 224: Pre-Jurassic evolution of eastern Niigata Univ. 15 (Ser. E, Geol.), 91-109.
Asia. Osaka, 99-115. ZAMORAS, L.R. & MATSUOKA, A. 2001: Malampaya Sound Group: A Jurassic-
JAPAN INTERNATIONAL COOPERATION AGENcy-METAL MINING AGENCY OF Early Cretaceous accretionary complex in Busuanga Island, North
JAPAN 1990: Report on the mineral exploration, mineral deposits and tec- Palawan Block (Philippines). J. geol. Soc. Japan 107/5, 1-336.
tonics of two contrasting geologic environments in the Republic of the
Philippines, terminal report. 121 p, Metal Mining Agency, Tokyo.
KIESSLING, W. & FLOGEL, E. 2000: Late Paleozoic and Late Triassic limestones
from North Palawan Block (Philippines): Microfacies and paleogeograph- Manuscript received January 2004
ical implications. Facies 43, 39-78. Revision accepted February 2005
S112 E J. Marquez, J. C Aitchison & L. R. Zamoras

Plate 1
Upper Permian radiolarians from the Mabintangin River (sample location 13), Busuanga, Philippines. Scale bars = 100 urn.
Figs. 1-5 from HKU·PLWN02031001, Upper Permian:

1. spherical radiolarian; 2. ?Copicyntroides sp. A sensu KUWAHARA & YAO;3. ? Nazarovella gracilis DE WEVER & CARIDROIT; 4, 5. broken Follicucullus
Figs. 6--11 from HKU·PLWN02031002, Upper Permian:
6. Trilonche sp. d. T. cimelia (NAZAROV & ORMISTON); 7, 8. Follicucullus porrectus RUDENKO; 9-11. Follicucullus charveti CARIDROIT & DE WEVER
12. spherical radiolarian; 13. ?Copicyntra sp.; 14. Ishigaum sp. d. I. trifustis DE WEVER & CARIDROIT; 15. Latentifistula sp. F sensu KUWAHARA AND YAO?;
16. Deflandrella sp.; 17. Nazarovella sp.; 18, 19. Latentibifistula spp.
Figs. 20, 21 from HKU·PLWN02031004, Upper Permian:
20. Copicyntroides? sp. A sensu KUWAHARA AND YAO;21. Ishigaum sp.
22. ?Stigmosphaerostylus sp.; 23. Trilonche sp. cf. T. cimelia (NAZAROV & ORMISTON); 24. Copicyntra sp.?; 25. Copiellintra? sp. A sensu KUWAHARA AND
YAO; 26. Albaillella sp.; 27. Albaillella sp. d. A. cavitata KUWAHARA
28,29. Albaillella cavitata KUWAHARA; 30. Ishigaum nicolasensis TUMANDA; 31. Latentibifistula sp.
32. Foremanhelena musashiensis (SASHIDA & TONISHI); 33. Albaillella triangularis ISHIGA, KITO & IMOTO
34. ? Stigmosphaerostylus sp.; 35. Polyfistula sp.
Fig. 36 from HKU·PLWN02031011, Upper Permian:
36. Ruzhencevispongus (?) sp. B sensu TUMANDA
Fig. 37 from HKU·PLWN02031016, Upper Permian:
37. Albaillella? protolevis KUWAHARA
Figs. 38--41 from HKU·PLWN02031017, Upper Permian:
38. Ishigaum nicolasensis TUMANDA; 39. Trilonche sp.; 40. Nazarovella gracilis DE WEVER & CARIDROIT; 41. Foremanhelena musashiensis (SASHIDA & TONISHI)

Plate 2
Middle to Upper Triassic radiolarians from Mapadolo (sample location 3) (figs. 1-4), Dabatonay (sample location 2) (figs.5-35) and Mabudyen (sample location 12)
(figs. 36-40), Philippines. Scale bars = 100 urn,
Fig. 1 from HKU-PLWN02031101, upper Anisian-Ladinian

Crucella sp.
Figs. 2,3 from HKU-PLWN02031102, Carnian:
2. Capnuchosphaera deweveri KOZUR & MOSTLER, emend. BLOME; 3. Capnuchosphaera missionensis CORDEY
Fig. 4 from HKU-PLWN02031103, upper Norian-Rhaetian?
Haeckelicyrtium'l sp.
Figs. 5-15 from HKU-PLWN02031109, uppermost Anisian-Ladinian:
5. Pentactinocarpus [usiformis DUMITRICA; 6. Archaeocenosphaera (?) sp.; 7. Triassocampe sp.; 8. Spinotriassocampe hungarica KOZUR; 9. Eptingium man-
[redi DUMITRICA; 10. Spongostephanidium longispinosum SASHID; 11. Tiborella sp.; 12. Parasepsagon sp.; 13. Pseudostylosphaera longispinosa KOZUR &
MOSTLER; 14. Pseudostylosphaera compacta (NAKASEKO & NISHIMURA); 15. Pseudostylosphaera magnispinosa YEH
Figs. 16-22 from HKU-PLWN02031110, upper Anisian-Ladinian:
16. Archaeocenosphaera (?) sp.; 17, 18. Poulpus spp.; 19. Pentactinocarpus fusiformis DUMITRICA; 20. Oertlispongus inaequispinosus DUMITRICA, KOZUR &
MOSTLER; 21. Triassocampe deweveri (NAKASEKO & NISHIMURA); 22. Triassocampe coronata BRAGIN
23. ?Archaeocenosphaera sp.; 24. Pentactinocarpus fusiformis DUMITRICA; 25. Hozmadia sp.; 26. Pseudostylosphaera magnispinosa YEH
27. Triassocampe coronata BRAGIN; 28. Spinotriassocampe hungarica KOZUR; 29. Pseudostylosphaera japonica (NAKASEKO & NISHIMURA); 30. Pentactinocar-
pus tetracanthus DUMITRICA; 31. Pentactinocarpus [usiformis DUMITRICA; 32. Eptingium manfredi DUMITRICA; 33. Parasepsagon sp.; 34. Poulpus sp.
Fig. 35 from HKU-PLWN 02031116, upper Anisian-Ladinian:
35. Spinotriassocampe annulata (NAKASEKO & NISHIMURA)
Figs. 36-40 from HKU-PLWN02031202, upper Norian-Rhaetian:
36. Praemesosaturnalis huxleyensis (CARTER); 37. Praemesosaturnalis gracilis (KOZUR & MOSTLER); 38-40. Praemesosaturnalis spp.

Plate 3
Middle Triassic to Lower Jurassic radiolarians from Mabudyen (sample location 12) (figs. 1,2), Sitio Tingil (sample location 11) (figs. 3--{j), Marily (Figs. 7-15) and
Dimanglet (sample location 8) (figs. 16-37), Philippines. Scale bars = 100 urn.
Figs. 1,2 from HKU-PLWN02031202, upper Norian-Rhaetian:

1. Praemesosaturnalis rugosus (YEH); 2. Paleosaturnalis sp. cf. P. dotti (BLOME)
Figs. 3-5 from HKU-PLWN0203121O, upper Anisian-lower Ladinian:
3,4. Oertlispongus diacanthus SUGIYAMA; 5. Astrocentrus sp. ct. A. pulcher KOZUR & MOSTLER
Fig. 6 from HKU-PLWN02031220, upper Carnian-upper Norian:
6. Capnodoce sp. cf. C. anapetes DE WEVER
Figs. 7-9, from HKU-PLWN02031302, Hettangian-Sinemurian?:
7, 8. Parahsuum simplum YAO; 9. Sethocapsa sp.;
Figs. 10, 11 from HKU-PLWN02031303, Hettangian-Sinemurian?:
10, 11. Archicapsa sp.
Figs. 12-15 from HKU-PLWN02031304, Hettangian-Sinemurian?:
12. Pantanellium ultrasincerum BLOME; 13. Haliomma sp.; 14. Trilonche sp.; 15. Sethocapsa sp.
Figs. 16-19 from HKU-PLWN02031402, Ladinian-lower Carnian:
16. Napora sp.; 17. Silicarmiger sp.; 18. Muelleritortis sp.; 19. Pararuesticyrtium sp.
Figs. 20--23 from HKU-PLWN02031405, upper Carnian-lower Norian:
20. Haeckelicyrtium'i sp.; 21. Latium sp.; 22. Capnuchosphaera lea DE WEVER;23. Pachus multinodosus TEKIN
Fig. 24 from HKU-PLWN02031407, upper Carnian-lower Norian
24. Capnuchosphaera lea DE WEVER
Fig. 25 from HKU-PLWN02031413, upper Carnian-Norian
25. Capnodoce extenta BLOME
Figs. 26-28 from HKU-PLWN02031414, upper Norian-Rhaetian:
26. Betraccium maclearni PESSAGNO & BLOME; 27. Betraccium deweveri PESSAGNO & BLOME; 28. Praemesosaturnalis sandspitense (BLOME)
Fig. 29 from HKU-PLWN02031418, upper Norian-Rhaetian:
29. Betraccium deweveri PESSAGNO & BLOME
Fig. 30 from HKU-PLWN02031419, upper Norian-Rhaetian:
30. Canoptum sp.
Figs. 31, 32 from HKU-PLWN02031420, upper Norian-Rhaetian:
31. Acanthosphaera sp.; 32. Livarella magna TEKIN
Figs. 34-37 from HKU-PLWN02031430, Sinemurian
33. Archaeodictyomitra sp.; 34. Parahsuum sp.; 35. Katroma westermanni WHALEN & CARTER; 36. Podobursa sp.; 37. Sethocapsa sp.
Sl18 E. J. Marquez, J. C. Aitchison & L. R. Zamoras

Plate 4
Middle Triassic to Lower Jur assic radiolarians from Dimanglet (sample location 8) (figs. 1- 15), Mayanpa yan (sample location 7) (figs. 16, 17), Lusong (sample lo-
cation 5) (figs. 18,1 9), Calauit-I llul tuk Bay (sample location I) (figs. 20-2 1) and Dipuyay-Decalangwang River, Busuanga (sample location 4) (figs. 22-44) , Philip-
pines. Scale bars = 100 urn.
Figs. 1-3 from HKU-PLWN02031701, upper Carnian-Norian:

I. Eptin gium sp.; 2. Jap on ocamp e sp.: 3. Corum sp. cf. C. delgado SUGIYAMA
Figs. 4-7 from HK U-PLWN02031702, Ladinian-lower Carnian:
4. Busuanga (?) sp.; 5. Pseudostylosph aera compacta (NAKASEKO & NISHIMURA); 6. Vinassaspo ng us sub spha ericus KOZUR & MOSTLER; 7. Mu elleritortis
cochleata (NAKASEKO & NISHIMURA)
Fig. 8 from HKU -PLWN02031705, upper Norian-Rh aetian:
8. Paroertlispongus sp
Figs. 9, 10 from HKU-PLWN02031706, upper Norian-Rh aetian:
9, 10. Praem esosaturnalis sp. cf. P. finchi (Pessagno)
Figs. 11-15 from HKU-PLWN02031707, Bajocian?, could be younger:
11,1 2. Williriedellwn sp.; 13-1 5. Parahsuum sp.
Figs. 16, 17 from HKU-PLWN0 2031713, Sinemurian:
16. Pleesus sp. cf. P. aptus YEH: 17. Katr oma colif orm e (HORI)
Fig. 18 from HKU-P LWN02031802. upper Carnian-lower Norian:
18. Canoptum [arawayense BLOME
Fig. 19 from HKU-P LWN02031803, upper Carnian-lower Norian:
19. Xip otheca sp. d . X. rugosa BRAGIN, emend. TEKIN
Fig. 20 from HKU- PLWN02031904, uppe r Anisian-Ladinian:
20. Riki vatella sp.
Fig. 21 from HKU -PLWN02031905, uppe r Anisian-lower Ladinian:
21. Triassocampe tulbuanensis T UMANDA
Figs. 22-24 from HKU -PLWN02032013, upper Anisian-Ladini an:
22. Paroertlispongus sp.; 23. Triassocampe sp.; 24. Triassocampe de weveri (NAKAsEKo & NISHIMURA)
Fig. 25 from HKU -PLWN02032018, uppe r Anisian-Ladinian:
25. Praecon ocaryomma sp.
Figs. 26-28 from HKU -PLWN02032020, upper Anisian-Ladinian:
26. Pseud ostylosphaera tim orensis SASIlIDA & KAMATA; 27. Hindeospha era sp inulosa (NAKASEKO & NISHIMURA); 28. Sp on gostephanidium [aponicum
(NAKASEKO & NISHIMURA)
29. Pararuesticyrtium sp.; 30. Tiborella fl orida (NAKASEKO & NISHIMURA); 31. Eptingi um nakasekoi KOZUR & MOSTLER; 32. Cryptostephanidium cornigerum
DUMITRICA
Figs. 33-39 from HKU-PLWN02032022, LADINIAN:
33. Pentactino capsa awaensis NAKASEKO & NISHIMURA; 34. Ho zm adia rotunda NAKASEKO & NISHIMURA; 35. Spon gostephanidium japonicum (NAKASEKO &
NISHIMURA); 36. Pseudostylospha era japon ica (NAKASEKO & NISHIMURA); 37. Parasepsagon sp.; 38. Pseud ostylospha era com pacta (NAKASEKO & NISHIMURA);
Triassocampe coron ata BRAGI N
Figs. 4Q-44from HKU-PLWN02032023, upper Anisian-Ladinian:
40. Hindeosphaera spinulosa (NAKASEKO & NISHIMURA); 41. Pseud ostyl osph aera ma gnispinosa YEH; 42. Tibo rella m agn identata Du mitrica, KOZUR &
MOSTLER:43. Stau rolonche trispin osa (KOZUR & MOSTLER); 44. Tiborella fl orida (NAKAsEKo & NISHIMURA)
S120 E. J. Marq uez, J. C. Aitchison & L. R. Zamoras

Plate 5
Middle to Uppe r Triassic radiolarians from Dipuyay-Decalangwang River (sample location 4), Busuanga, Philippines. Scale bars = 100 11m.
Figs. 1-4 fro m HK U-PLWN02032025, upper Anisia n-Ladinian:

I. A eanth ospha era (?) mocki KOZUR & MOSTLER;2. Thaisphaera sp.; 3. Pararuesticyrtium sp.; 4. Eptin gium ram o vsl KOZUR et al.
Figs. 5-9 from HKU-PLWN0203202 9, upper Anisian-Ladinian:
5. Triassocampe coronata BRAGIN; 6. Staurolonche sp.; 7. Stauralon ehe trispinosa (KOZUR & MOSTLER); 8. Sarla sp.; 9. Triassistephanidium laticorn e
DUMITRICA
Figs. 10- 18 from HK U-PLWN02032027, upper Anisian-Ladinian:
10. Pentactinoc apsa awaensis NAKASEKO & NISHIMURA; 11, 12. Ho zm adia reticulata DUMITRICA, KOZUR & MOH LER; 13. Hexalon ehe sp.; 14. Pseudosty-
losphaera go ricanae KOZUR ET AL.; 15. Epigondollela sp.; 16. Tibo rella sp.; 17. Plafk erium abb oti PESSAGNO; 18. Parasepsagon variabilis NAKASEKO &
NISHIMURA
Figs. 19-26 from HK U-PLWN02032031, upper Anisian-Ladinian:
19. Pentactinocapsa awaensis NAKASEKO & NISHIM URA; 20. Hindeosphaera spinu losa (NAKASEKO & NISHIMURA); 21. Pseudostylosphaera timorensis
SASIIlDA & KAMATA; 22. 23. Pseudostylospha era compacta (NAKASEKO & NISH IM URA); 24. Pseudostylosphaera japonica (NAKASEKO & NISHIM URA); 25. Tri-
assoeampe dewe veri (NAKASEKO & NISHI MURA); 26. Triassocampe coronata BRAGI!'!
Figs. 27,28 from HKU-PL WN02032037, upper Anisian-Ladinian:
27. Triassocamp e scalaris D UMITRICA, KOZUR & MOHLER; 28. Spinotriassocamp e annulata (NAKAsEKo & NISHIMURA)
Figs. 29-31 from HK U-PLW N02032041, upper Anisian-Ladinian:
29, 30. Ho zm adia reticulata DUMITRICA, KOZUR & MOHLER; 31. Napora sp.
Figs. 32, 33 from HKU-PLWN02032049, upper Anisian-Ladinian:
32. Cryp tostephanidium cornigerum DUMITRI CA; 33. La dinocampe japonica (NAKASEKO & NISHIMURA)
Fig. 34 from HKU-PLWN02032050, upper Anisian-Ladinian:
34. Bulbo cyrtium sp.
Figs. 35-41 from HK U-PLWN02032052, upper Anisian-Ladinian:
35. Sepsagon sp.; 36. 38. Ho zm adia sp.; 37. A rchaeocenosphaera sp.; 39. Ho zm adia rotunda (NAKAsEKo & NISHIMURA); 40. Tibore lla sp.; Katorella bifur cate
KOZUR & MOSTLER
Figs. 42,43 from HK U-PLWN02032057, upper Anisian-lower Ladinian:
42. A rehaeoce no sp haera sp.: 43. Pentactinoco rbis k oz uri DUMITRICA
SI 22 E. J. Marquez, J. C. Ait chison & L. R. Zamoras

Plale6
Middle Triassic to Middle Jurassi c radiolarians from Dipuyay-Decalangwang River (sample location 4) (figs. 1-9 ), Calindo (sample location 10) (figs. 10-4 1) and
a road cut, 3.5 kms east of the township of Coron (sample location 9) (figs. 42-4 5). Scale bars = 100 urn.
Figs. 1--6from HKU-PLWN02032057, upp er Anisian-lower Ladinian:

l a. Hozmadia spinifera SUGIYAMA; lb. Hind eosphaera spinulosa (NAKASEKO & NISHIMURA); 2, 3. Paratriassocamp e sp.; 4. Triassocampe eoronata BRAGIN;
5, 6. Tiborella florida (NAKASEKO & NISHI MURA)
Figs. 7- 9 from HKU-PLWN02032060, upp er Anisian-Ladinian:
7. Staurolonche trispinosa (KOZUR & MOSTLER); 8. Hozmadia rotunda (NAKASEKO & NISHIMURA); 9. Hind eospha era spinulosa (NAKASEKO & NISHIMURA)
Figs. 10-13 from HKU-PLWN0203210 1, upp er Bajocian-uppermost middle Bat honian:
10. Sriehocapsa sp.; 11. Lina resia sp.; 12. Staurolonche sp.; 13. Transhsuum medium TAKEMURA
Figs. 14-22 from HKU -PLWN02032lO2, upper Anisian-Ladinian:
14. Paratriassoeamp e sp.; 15. Pseudosty/ospha era /ongispinosa KOZUR & MOSTLER; 16. Pseudosrylospha era compacta (NAKASEKO & NISHIMURA); 17. Spon-
gostylus tricostatus KOZUR, KRAINER & MOSTLER; 18. Eptingium manfredi DUMITRICA; 19. Hozmadia reticulata DUMITRICA, KOZUR & MOSTLER; 20. Hoz-
madia rotunda (NAKASEKO & NISHIMURA; 21. ? Hexalonche sp.; 22. Staurolonche sp.
Figs. 23-25 from HKU-PLWN02032106, Ladinian :
23. Eptingium m anfredi DUMITRICA; 24. Stauracontium sp.; 25. Pentaspongodiscus sp.
Figs. 26- 32 from HKU-PLWN0203 2109. upper Anisian-Ladinian:
26. Hozmadia rotunda (NAKASEKO & NISHIMURA); 27. Spon gosilicarmiger sp.; 28. Hind eosphaera spinulosa (NAKAsEKo & NISHIMURA); 29. Hozmadia
rotunda (NAKAsEKo & NISHIMURA ); 30. Triassocampe caronara BRAGIN; 31. I'ararriassocampe sp.; 32. Parasepsagon variabilis (NAKASEKO & NISHIMURA)
Figs. 33-36, 38 from HKU -PLWN0 2032110, upper Anisian-Ladinian :
33. Pentaetinoeapsa awaensis NAK AsEKO & NISHIMURA; 34. ?Trilonehe sp.: 35. Celluronta sp.; 36. Bulb ocyrtium sp.: 38. Tiborella flor ida (NAKASEKO &
NISHIM URA)
Figs. 37, 39-4 1 from HK U-PLWN02032117, upper Anisian-Ladinian:
37. ? Ladin ocampe sp.; 39. Spin otriassocamp e annulata (NAKAsEKo & NISHIM URA); 40. Spongo stephanidium japonicum (NAKASEKO & NISHIM URA); 41. Hoz -
madia sp.
Figs. 42-45 from HKU -PLWN02032118, upper Norian-Rhaetian:
42. Canoptum sp.; 43. Canoptum sp. aff. C. unicum PESSAGNO & WHALEN; 44. Deflandrecyrtium earterae YEH & CHENG; 45. Livarella magna TEKIN

DOl 10.1007/s00015-006-0601-6
Influence of the Frasnian-Famennian event on radiolarian faunas

YU-JING WANG 1, HUI Luo! & JONATHAN C. AITCHISON 2
Key words: Radiolaria, Frasnian. Famennian. extinction event. South China
ABSTRACT
The impact of the Frasnian-Famennian biotic mass extinction event (F-F flourished during the Famennian. The F-F event may not have had any drama-
event) on radiolarian faunas is examined in 13 sections, which belong to two tic effect on the radiolarian faunas in deep water and the disappearance of Fa-
types of cherty basin in South China. No appreciable decrease in radiolarian mennian radiolarians in shallow water platform sections may simply be a result
biodiversity is observed across the F-F boundary. Indeed, radiolarian faunas of sea level change rather than any biotic crisis.
tional basin and the characteristics of the fossil assemblages,

Introduction
these 13 stratigraghic sections may be divided into two basin
The F-F event is one of the great biotic mass extinction events types, namely the open-ocean cherty basin type and the cherty
that occurred during the Phanerozoic. It took place between platform basin type (Fig. 2). These two cherty basins experi-
late Frasnian and Famennian time in the Late Devonian and enced different developmental histories during the F-F biotic
its scale was of only marginally lesser magnitude than the mass extinction event.
largest mass extinction, which occurred at the PIT boundary. The open-ocean cherty basin indicates deep-water basin
The F-F event caused a great extinction or rapid diminution in deposition except for minor platform and the slope sediments.
the abundance of marine invertebrate animals, such as stro- This is a basin of compensative lack that experienced the same
matoporoids, brachiopods, rugose corals, tabulate corals, trilo- deep-water anoxic conditions for a long time (possibly from
bites, ammonoids, tentaculitids, ostracods, conodonts and so Silurian to Permian or Triassic). The Devonian sections at the
on. Over 60% of total taxa of these animals disappeared at the Shiti Reservoir, Bancheng, Qinzhou, Guangxi and in the
end of the Frasnian (Xian et al. 1995). Based on Frasnian- Changning-Menglian terrane, west Yunnan are considered to
Famennian radiolarian materials collected from South China be the typical representatives of this kind of basin. They con-
and those reported from elsewhere, we discuss the effects of tain continuous radiolarian zones composed of the Frasnian
the F-F biotic mass extinction event on radiolarian faunas. Helenifore laticlavium and Helenifore robustum zones and the
Famennian Holoeciscus foremanae zone. The cherty platform
basin developed as a result of siliceous pelagic sedimentation
Stratigraphy and lithologic characteristics
within a platform basin. This basin experienced slow sedimen-
Our radiolarian samples were gathered from thirteen strati- tation in an anoxic environment during periods of the plat-
graphic sections containing cherty rocks distributed in Yunnan forms development (e.g. during the early or late Frasnian),
(6 sections), Guizhou (1 section) and Guangxi (6 sections) re- owing to sea level rise caused by transgression in the basin.
spectively in South China (Fig. 1). The general characteristics With the exception of the sections in the Qinzhou area,
of these sections include an age range from Frasnian to Fa- Guangxi and in Changning-Menglian terrane, Yunnan, the
mennian, rocks composed of thin-bedded cherts associated other stratigraphic sections cited-above belong to this basin
with mudstone and shale, and some rich radiolarian faunas. type. In these sections, there are no continuous radiolarian
According to the developmental background of the deposi- zones. Generally, only one fossil zone, such as, the lower or
I Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008,China. Email: [email protected]
2 Department of Earth Sciences, University of Hong Kong, Pokfulam Road, Hong Kong SAR, China
Influence of the F-F event on radiolarian faunas S127

100 105 110 tion , west bank of th e Shiti Reservoir , Bancheng countryside ,
Qinzhou, Guangxi (Wa ng et al. 2(03). It is considered to be
•
301---+----I-2J"ft-~---r__--_ru l
lower Frasnian. Helenifore robustum (Boundy-Sanders &
Murchey 1999) is characterized by an elliptical shell with a nar-
rower and thicker ring (Plate 1, Figs . 6-10). This is an upper
Frasnian species, which is distributed worldwide. It has been
reported fro m eastern Australia (Ishiga 1988; Ishiga et al. 1988;
, Stratford & Aitchison 1997) , Thailand (Sa shida et al. 1998),
f 0 5
, ." , ... , .- ,t.., Ne vad a (Boundy-Sanders & Murchey 1999) and Guangxi,
~-- j - I., ',
a, Guizhou, Yunnan in Ch ina (Wang et a l. 1998; Wan g et al.
'., '\
'_ .- . Guang i 11; ,. 2003). Therefore, we subdivide the Upper Devonian radiol ari -
I
10 .' an fauna into 3 zones: Two zones (He lenifo re laticlavium zone
and H. robustum zon e) are Frasnian and one zon e (Ho loecis-
cus f oremanae zon e) is Famennian.
150
I
Correlation between Upper Devonian radiolarian and
conodont zones
1. East section of Shiti Reservoir, Qinzhou, Guangxi
2. West section of Shiti Reservoir, Qinzhou, Guangxi At pr esent 15 typical Upper Devonian conodont zon es are
3. Wuxiangling section, Nanning, Guangxi erected (7 zones belonging to Frasnian and 8 zon es to Fa-
4. Yangdi section, Guilin, Guangxi mennian , respectively) (Fig. 3) (Wang Cheng-yuang 2000).
5. Bazhai section, Ziyun, Guangxi
6. Shaijingpo section, Xiangyun, Yunnan The Helenifore laticlavium zone from th e Gogo Formation of
7. Lila section, Lancang, Yunnan western Australia is associated with conodonts th at include
8. Taierbu section, Lancang, Yunnan Polygnathus asymmetrica and Ancyrodella rotundilobata ro-
9. Huiku section, Menglian, Yunnan tundilobata . It is comparable to the lower-middle part of the
10. Niuyunling section, Yulin, Guangxi
11. Etang section, Hexian, Guangxi Polygnathus asymmetrica zone (Nazarov & Ormiston 1983).
12. Nanya section, Menglian, Yunnan The Helenifore robustum zone occurs in the Slaven Chert of
13. Ali section, Lancang, Yunnan the Shoshone Range, Nevada, USA (Boundy-Sanders and
Murchey 1999) and in th e Tarbu area, Lachang county, west-
Fig. 1. Map showing Ihelocations of sections studied through Upper Devon- ern Yunnan, Ch ina (Wang et al. 2000) . It is associated with
ian chertystrata in South China.
conodonts that include Palmatolepis eurek a, P. rhenana masu-
ta, P. aubr ecta, Polygnathus brevilaminus, P. aff. timanicus
(Boundy-Sanders and Murchey 1999) or Palmatolepis rhenana
upper Frasnian Helenifore laticlavium or Helenifore robustum rhenana, P. rhenana nasuta , P. hasii and P. sp. (W ang et al.
zone can be distinguished. Moreover, Famennian radiolarian s 2000). Consequently, th e radiolarian Helenifore robu stum zone
and radiolarian zones have never been discovered. From mid- may be correlated with the conodont Palmatolepis hasii zone
dle Frasnian to Famennian, the radiolarian-bearing cherts in to Palmatolepis rhenana zone. The Holoeciscus foremanae ra-
the cherty platform basin were gradually replaced by pha- diolarian zone from the Frankenwald are a, northern Bavaria,
coidal lime stone due to regression. The timing of th is facies Germany is associated with the con odonts Palmatolepis ru-
change differed between areas. It occurred in the middle gosa, P. glabra lepta, P. gracilis gracilis, P. per/obata schinde-
Frasnian in SE Guangxi and in the Famennian in SW Guizhou. wolfii, Branmmehla bohlenana, Pseudop olygnathus granulasus
and occurs in the Palmatolepis trachytera zone to lower P.
po stera zon e in the middle Famennian (Kiessling & Tragelehn,
Age assignment
1994). The same rad iolarian zone from th e Shiti Reservoir For-
mation in Bancheng, Q inzhou, Guangxi is also associated with
Upper Devonian radiolarian faunas
the conodonts Palmatolepis glabra acuta, P. per/obata schinde-
Our recent study suggests that the species Helenifore laticlavi- wolfii. It is correlated with the Famennian Palmatolepis crepi-
um identified by various scholars may be separated into two da zone to P. marginifera zone (Wang et al. 1998). Therefore,
stratigraphically and morphologically distinct spe cies (W ang et the Holoeciscus forem anae zone can be correlated with the
al. 2(03). Helenifore laticlavium NAZAROV & ORMISTON (1983) Palmat olepis crepida zone to P. postera zone (Fig. 3) .
is characterized by a shell with a thinner, ne arl y circular plat e-
ring , two spines (top and base spines) and an opening on one
Upper Devonian radiolarian diversity
extremity of the ring (Plate 1, Figs. 1-5). This species occurs in
the Gogo Formation, Ca nning Basin , western Australia According to our Upper Devonian radiolarian materi als col-
(Nazarov & Ormiston 1983) and the Shiti Reservoir Forma- lect ed from South Ch ina (Wang 1997; Wang et al. 2003) and
S128 Y.-J. Wang, H. Luo & J. C. Aitchison

Op en -o cean cherty basin Platform cherty basin
Shi ti Reser vo ir
East Section Ali
:::
~
:::
:::
v
E E
c ~ t.l.
~ u, l-
'ee 0
>
>
~
I-
v
Q V
<Il
J- ~
ell
Q. ....
:::
::l
Q.
~ -=
U1
e<Il
~
I-
t.l.
...
~
::: C
o
~ ell
:c ·ce
~
0
:0=
Q)
<::
0
"0
.- > .:2: 0
:E Q
ell (9 .s
><
HI=Helenifore laticlavium fauna EI=Eoalbaillella lilaensis fau na

@=Radiolarian fauna without Helenifore laticlavium Hf=Holoeciscus f oremanae fauna
Hr=Helenifore rob ustum fauna A=Albaillella fauna
~ Mudstone i55 Limestone m Phacoidal li mestone E Cherts a Siliceo us shale
Fig. 2. Two different basin types containing cherty facies.
other published materials on Upper Devonian radiolarian fau- 10 genera and 12 species are same in both zones indicating a
nas (Aitchison 1993; Boundy-Sanders et al. 1999; Cheng 1986; close affinity. However, in the Famennian Holoeciscus forem-
Foreman 1963; Holdsworth et al. 1978; Ishiga et al. 1988; anae zone, 154 species, belonging to 28 genera and 10 families,
Kiessling & Tagelehn 1994; Li & Wang 1991; Nazarov 1973, are described. 18 species and 13 genera, as well as 8 families
1975; Nazarov & Ormiston 1983; Nazarov et al. 1982; Sashida continued through from two Frasnian zones. 2 new families
et al. 1993; Schwartzapfel & Holdsworth 1996; Stratford & (Holoeciscidae and Pylentonemidae) and 14 new genera ap-
Aitchison 1997), we find that there are 77 and 35 species with- pear. The number of newborn genera is almost double the
in the Frasnian Helenifore laticlavium zone and H. robustum total number of radiolarian genera in the Frasnian. Although
zone respectively. They all belong to 15 genera and 8 families. the species number decreased in Upper Frasnian compared to
Although these two faunas have the same number of families that of Lower Frasnian, the diversity of Famennian radiolarian
and genera, the number of species recognized in the Helenifore families, genera and species are obviously higher than that of
robustum zone is less half than that in the H. laticlavium zone. whole Frasnian (Fig. 4, Fig. 5).

Early Frasnian LateFrasnian Famennian
conodont zones radiolarian zones
Helenifore
praeslliLvrJa
Ceratoikiscum
c exPIJII!fIJ
-- --- -- --- - - - ------ ---- -- ------ Stigmosphaerostylus
'2" poston
»Entactinia
s tracln;tera
margin era
HoloeciscusjoremanlU
Trtlonche
§ rho'lnboil!ea »Entactinosphaera
~
crepida
trianRuJari.~
-
- ---- - - -- - - - - - - - - - - - - - - - - - - - -- Soonzentactinia
linguifornti.v
-- -- -- - - ------ - - - - - ---- - - - -- ---
Polyentactinia
Astroentactinta
------
C rhmonn
Jlf!/Lnijo" robustum Helioentactinia
.S "ilmUfJe
C
e
.....
hassi
panLttUll
tran.utallS
-
- - - - - - - - - - - - - --- - - - - - - - - - - --
H,/mijore laticlJlvillm
-- Somohoentactinia
Spongentactineila
Secuicollacta
"lsiovo/is
Palaeoscenidium
Palaeotripus
Fig. 3. Correlation of Upper Devonian radiolarian and conodont zonations. Haplentactinia
Bissvlentoctinia
Palacantholithus
Protoalbaillella
Archocyrtium
Lower Frasnian .lJPperFrasnian Famennian Trtaenospbaera
Popofskyetlum
Holoeciscus
Pvlentonema
Quadrapesus
Cerarchocyrtium
Cynentactinia
Huasha
Cyrtisphaeractentum
Lapidopiscum
Totollum
Kantollum
Stourentactinia
Tetrentacunia
Deflandrellium
Robotium
Fig. 5. Ranges of Upper Devonian radiolarian genera.
mennian radiolarians are present. The disappearance of radio-

• Family numbers ~ Genera numbers ~ Species numbers
larians from the platformal sections is'obviously related to fa-
cies change rather than extinction. We note that in the Early
Fig. 4. Diversity changes of the radiolarian family, genera and species in
Carboniferous, when deposition in the platform basin changed
Upper Devonian.
from limestone facies into a cherty facies again an albaillellid
radiolarian fauna re-appeared in this platform basin.
Consequently, we conclude:
Influence ofthe F-F event on radiolarians
1. The F-F event may have only resulted in a major biotic
The materials mentioned above show that the F-F biotic mass mass extinction for biotas associated with shallow water
extinction event did not have any marked negative effect upon reef facies and parts of the pelagic biota. It did not have a
the open ocean radiolarian fauna. In the open ocean cherty great influence on the radiolarian fauna in deep water and
basin 8 upper Frasnian radiolarian families continue to flourish indeed radiolarians flourished during the Famennian.
and only 2 out of 15 genera become extinct in the Famennian. 2. The disappearance of radiolarians from shallow water plat-
No families and only 13% of genera become extinct. These fig- form sections may simply be a result of sea level change
ures are well below the minimum level of extinction (11 % at rather than any biotic crisis. In deeper water successions,
family level and 20-25% of genera) regarded as indicative of a radiolarians were less affected by sea level change. There
biotic mass extinction (Raup 1982). Elsewhere, in the cherty was no appreciable decrease in radiolarian biodiversity
platform basin, the radiolarians belonging to Helenifore lati- across the F-F boundary.
clavium and H. robustum zones thrived during the early and
late Frasnian. However, as the radiolarian-bearing cherts were In order to test this hypothesis we need to find and examine
replaced by phacoidallimestone because of regression, no Fa- more Famennian platformal cherty successions.
S130 Y.-J. Wang, H. Luo & J. C. Aitchison

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Microp al aeont ologica Sinica 8(4) , 395-4 04. Wu , H.. XIAN, X. & KUANG, G . 1994: Late Paleozoic Radi olari an asse mblages
NAZAROV, B.B . 1973: Per vye na khod ki radiolyari i Entactiniidae i Ce ra- of so uthe rn G ua ngxi and its geo logical significance. Scie ntia Geologic a
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Plate 1
Index species of Late Devonian radiolarian zonations from South China. All scales = 100 11m: A for figure 1; B for figures 2-10; C for figures 11-15.
Figs. 1-5, Helenifore laticlavium NAZAROV & ORMISTON. 1-5 from sample 96shiti159, Shiti Reservoir Formation, Shiti, Bancheng, Guangxi.
Figs. 6-10, Helenifore robustum (BOUNDY-SANDERS & MURCHEY). 6-10 from sample 92J9, Shiti Reservoir Formation, Shija, Bancheng, Guangxi.
Figs. 11-13, Holoeciscus foremanae CHENG. 11-13 from sample 96shiti185, Bancheng Formation, Shiti, Bancheng, Guangxi.
Figs. 14-15, Holoeciscus elongates KIESSLING & TRAGELEHN. 14-15 from sample 96shiti185, Bancheng Formation, Shiti, Bancheng, Guangxi.
S132 Y-J. Wang, H. Luo & J. C. Aitchison

DOl lO.1007/s00015-006-0610-5
Birkhauser Verlag, Basel , 2006
Middle to Upper Permian radiolarian faunas from chert blocks in

Pai area, northwestern Thailand
NUTIHAWUT WONGANAN I & MARTIAL CARIDROIT2
Key words: Radiolaria, chert, biostratigraphy, Permian, north westernThailand

Mots cles: Radiolaire, «chert», biostratigraphie. Perrnien, Thailande du Nord-Ouest
ABSTRACf RESUME
Well-preserved Permian radiolarians have been discovered from chertblocks Des assemblages de radiolaires bien preserves ont ete extraits de blocks de
embedded in a unitin the Paidistrict. Mae Hong Son province. northwestern «chert» presents dans la region de Pai (Province de Mae Hong Son. Thailand
Thailand that was recently mapped as Carboniferous. Twenty-four taxa be- du Nord). recernment cartographies en tantque roches du Carbonifere. Vingt-
longingto ten genera have been recognized and somesignificant forms of Fol - quatre especes appartentnant a 10 genres sont reconnues, comprenant plu-
licucullus are present. Two upper Maokouan (Capitanian) to lower Wuchi- sieure Follicucullidae tres significatifs pour la stratigraphie. Deux zones d'as-
apingian radiolarian assemblages iFol licucullus monacanthus, Follicucullus semblages i Fotlicu cullus monacanthus et Fo/li cucullus charveti-F. porrectusi
charveti - F. por rectus assemblages) are reported. Follicucullus charvet i sont presentes et datent Ie Maokouien superieur (Capitanien) et Ie Wuchia-
CARIDROIT & DEWEVER is interpreted as a possible provincialism rather than pingien inferieur; elles sont similairs a celles presentes en Chie du Sud et au
being endemic like other members of the same genus. These results provide Japon. La presence de Follicucullu s charveti CARIDROIT& DEWEVER, espece
more data to demonstratethat in northernThailand.the Devonian to Triassic interpretee comme une forme d'eau chaude, est discutee, Ces resultas sont
was of a zone of deep siliceous marine sedimentation.The resultant deposits fondamentaux et sont une donnee supplernentaire pour prouver que l'en-
are one of the longest witnesses of continuousdeposition in an oceanic setting. semble des temps Devonien a Trias superieur est representepar des depots si-
liceux de mer profonde; ceci etant un des plus longs temoins connus d'une se-
dimentation oceanique continue.
1. Introduction
from Thailand. Radiolarian assemblage zones described from
Th e study of Permian radiolarians in Thailand, has become a Japan and China are identified and briefly discussed. Po ssible
popular topic since radiolarian research started (Caridroit et provincialism is observed with the occurrence of Follicu cullus
aI., 1990). Occurrences of Permian radiolarians are widespread charveti CARIDROIT & D E WEVER at few localities, compared
and th ey have now been studied in many regions; northern with the endemic nature of othe r members of this genus.
Th ailand (Chiang Mai: e.g . Caridroit 1991, 1993; Caridroit et
al. 1992; Sashida et al. 1993; Wongan an et al. 2002) , north-cen-
2. General Geology and Occurrence of Radiolarians
tr al T ha iland (Sukhothai: Sash ida & Nak ornsri 1997), north-
eastern Thailand (Loei: e.g. Sash ida et al. 1993; Sashida & Igo The district of Pai is located in NW northern Thailand, ap-
1999) , eastern Thailand (Sra Kh aew : e .g. Hada et al. 1999; proximately 110 km from Chiang Mai . According to Bunopas
Sashida et al. 20(0). The goals of th ese studies are both to im- (1981) the main Palaeozoic sedime ntary rocks widel y exposed
pr ove the biostratigraphy and to tr ack radiolarite s as remnants in the area is the Ma e Hong Son Formation. Age assi gnm ents
of oceanic palaeoenvironments. Only lower to middle Lower for thi s unit range from Siluro-D evonian (Bunop as 1981) to
and uppermost Permian faunas have been reported from Ca rbo nifero us-Permian (Chua viroj et al. 1985; Intawong e t al.
northern Thailand so far (e .g. Ca ridroit 1993; Sashida et al. 1997). The formation is dominat ed by unmetamorphosed
1993). The aim of thi s paper is to pr esent details of a new Palaeozoic sedimentary rocks, which ar e extensively dist rib-
upper Midd le to lower Upper Permian radiolarian locality uted in northwest Thailand. It mainl y consists of ma ssive or
I Department of Geological Sciences, Faculty of Science. Chiang Mai University,ChiangMai 50200 Thailand. E-mail: [email protected]
2 Laboratoirede Paleontologic et Paleogeographie du Paleozoique (LP3). UMR 8014 et UFR 1818 du C.N.R.S.,
Sciences de laTerre: Universitedes Sciences et Technologies de Lille, F-59655 Villeneuve d'Ascq cedex France. E-mail: [email protected]
Middle-Upper Permian radiolarians, Thailand S133

• I
:-•v
"",
Figure 2 \h:nll ll .... n ~

....., ~l'
~
.l.,""',~..
...
;!"~:
:
IO~ [3 ,....,
.. , Cb,.... ll.Io
•
~.
I
C M.a:II,,,,i' '' ''
.' ~
o
~
e
Radiolarian locality
Studied area
Provincia1capital
City
I"'
""
• Khun "",urn
-
-... ,
C
~~
'
~
-,
.,;
t
.;...
.,,~
'
•.
" ~I '
Il,I....
,
Fig. 1. Index map of northwestern Thailand
showing the studied area and the radi olarian lo-
calit y.
bedded sandstone, shale, chert and slate (Fig. 2). Limestones, beds intercalated within the Upper Silurian to Carboniferous
forming isolated karstic hills are widespread and contain abun- Mae Hong Son Formation (Bunopas 1981), or newly estab-
dant Carboniferous and Permian foraminifers and other fos- lished Carboniferous-Permian of Chauviroj et al. (1985). Ac-
sils. In general mos t Thai geologists correlate these limestones cording to field observations, no outcrops were found which
with the Ratburi G roup. Limestones in this area are lithologi- expose the strata underlying the chert.
cally variable, but mainly consist of massive limestones, with Under the microscope, the chert from lower part consists
local occurrences of dolomitic limestone with lenticular or of mainly cryptocrystalline to microcrystalline quartz associat-
nodular cherts. The relationship between limestone units and ed with very fine clay minerals. Radiolarians are rather well
Pa laeozoic siliceous sedimentary units is not clear. Field ob- preserved and abundant and filled by chalcedony. Further up
servations indicate the limestones unconformably overlie the lithologic column the cherts consist of cryptocrystalline
siliceous sediments, and where faulted the basal contact is lo- quartz with more abundant fine clay mineral and fine volcanic
cally marked by breccias. grains, without any traces of fossils . This probably indicates
The studied section is located approximately 7- 8 km north that there was volcanic activity within or nearby the site of de-
of Pai (Fig. 2), or at position 438455 on Thai topographic map position area conditions did not favour the preservation of ra -
sheet 4647 I Amphoe Pai, in the area of Ban Tan Jed Ton vil- diolarian skeletons.
lage. The section is exposed for approximately 850 m along a Samples from the main chert section (PAI-391 through
gravel road connecting Pai and Wieng Haeng that has been cut PAI-400) yielded the following Upper Permian radiolarian
through a high mountainous area. The section consists of gray faunas: Follicucullus ventricosus ORMISTON & BABCOCK, F.
to greenish gray , partiy dark gray, well-bedded chert with 4 to scholasticus ORMISTON & BABCOCK, F. orthogonus CARIDROIT
10 em thick beds intercalated with thin shale and clay (mm - & DE WEVER, F. porrectus RUDENKO, F. charveti CARIDROIT
few em) layers. The main and lowermost exposed section is & D E WEVER, F. sp. cf. F. bipartitus CARIDROIT & D E WEVER,
about 150 m long , before being obscured by strongly weath- Hegleria mammilla (SHENG & WANG) and others. The associa-
ered yellow volcanic rocks for about 200 m. The second section of F. charveti and F. porrectus RUDENKO is diagnostic for
tion, which is believed to be a separate block, is composed of assignment to the lower Wuchiapingian. Sample PAI-403 con -
greenish gray well-bedded chert an d about 10 m long. Ten tains fewer radiolarians, however the occurrence of F.
chert samples were collected from the main section (PAI-391 monacanthus ISHIGA & IMOTO is the diagnostic of the Midd le
to PAI-400) and three more were taken from the second (PAI- Permian F. monacanthus Assemblage (Capitanian), therefore
401 to PAI-403). The next three samples (PAI-404 to PAI-406) these chert beds are assigned to the Capitanian.
were taken from a small chert outcrop exposed on the other
side of 200 m of weathered tuffaceous material. Four chert
3. Radiolarian biostratigraphy
samples were collected (PAI-407 to PAI-412) from the last
section which ends at a summit approximately 150 m from the Twenty-two chert samples were collected, and nine samples
previous section. This chert has long been regarded as chert yielded Permian radiolarians (Plate 1). The studied chert sec-
S134 N. Wonganan & M. Caridroit

LEG EN D 10 I' m
kaJiol.uilCdI,·P....ih
c l~c [ )(' \ ,)l'Ii ~ n 1&.) I ';llc 1\'1'm ian '~ )
M.:t<4ITMll'phk rtd ,,,

t Silun ~ I)c: \ (",ian',
"
"",'
.... I nf,"m:d filull
Fig. 2. Simplified geologic map of Pai area, Mae
,~ mi-4.1 ,"'*.,til1t.nJ
C1('njal) ~ 1 1.
d ..:~l k unit
~ I
\ k liUrM lfJ'hk
lCamhrian'!)
n'll,:"~ .... Hong Son province, northwestern Thailand (see
~+ +
++ ++
...
In
Gr.uwtk rnc.....
(Iria ..!>ic?J / hUIt Fig. 1 for location); shows the studied sections.
(after Geologic map 1:50,000 Sheet Amphoe Pai
l"la..tic ~irncnl<I" unit
I
~
l lr
" Q
H.a..ic HlIC3nic n-.:h Quadrangle, Geological Survey Division, Depart-
tC'.oU'hl,nil'(Rlu... l \: m u Oln ). :./1'=1. .. lL.alC'Cartnnifefuu....·l.arl} I\'nnian!,
ment of Mineral Resources, Bangkok, Thailand
1985; Intawong et al. 1997, modified)
tion extends over two radiolarian assemblage zones: the Fol- 1990), Oregon in North America (Blome & Reed 1992), South
licucullus monacanthus, and Follicuculllus charveti - F. porrec- China (Wang et al. 1994), and is also reported in studies by
tus assemblage zones. These zones have been reported by Ishi- Caridroit (in De Wever et al. 2001, fig. 202, p. 313).
ga (1986, 1990) from Japan, Wang et al. (1994) from China and Follicuculllus charveti - F. porrectus assemblage zone
Caridroit in De Wever et al. (2001) and are known to occur in This assemblage zone is found within green chert samples
the middle Maokouan (Capitanian) to lower Wuchiapingian. PAI-393, 396, 398 & 399. It is defined by the co-occurrence of
Follicucullus monacanthus assemblage zone. Follicuculllus charveti CARIDROIT & DE WEVER and F. porrec-
This assemblage zone is found within sample PAI-403. It is tus RUDENKO. The range of this assemblage is considered as a
characterized by the occurrence of Follicucullus monacanthus total range of F. charveti CARIDROIT & DE WEVER. The other
ISHIGA & IMOTO, which is the characteristic taxon. Other radi- characteristic species are Follicuculllus venticosus ORMISTON
olarians species that occur in this assemblage include F. & BABCOCK, Follicucullus scholasticus ORMISTON & BABCOCK,
scholasticus ORMISTON & BABCOCK, Pseudotormentus sp. ct. P. Follicucullus sp. ct. F. bipatitus CARIDROIT & DE WEVER, Ishi-
kamigoriensis CARIDROIT & DE WEVER, F. orthogonus gaum similicutis CARIDROIT & DE WEVER, Triplanospongus
CARIDROIT & DE WEVER, F. ventricosus ORMISTON & BAB- musashiensis SASHIDA & TONISHI, Latentifistula texana
COCK, and Ormistonella robusta CARIDROIT & DE WEVER. The NAZAROV & ORMISTON, Hegleria mammilla (SHENG &
Follicucullus monacanthus assemblage is known from Japan WANG), and others. The assemblage contains the same radio-
(e.g. Ishiga 1986), Far East Russia (Rudenko & Panasenko larian species reported from Japan (e.g. Ishiga 1990), North

~
= lh03-J12
-Th03-..J.l1
= Th03-.ws
III
-Th03-107
=
=
=
Th03-399
Th03-398 ,.
II'
I
I
I
I
1"11
I I I
I I I
.,,:
=
=
Th03--106
Th03-405
ThOJ-404
,
~
",
-s,
~
I
I
I
I
c: ~
,
I I
'§'"
I I
~ I I
~ I
~ ~
~ ...:
='1'1103-396
II
..J .~ I
t
~
...:
= Th03-395
I
I
I
= Th03-394
I
I
I I
I I
• • Th03-393' I 1IIIIIIIIIil
:..,.v....
:v~....
=
",. Tb03-392
I I
I I I
I """"I
II • •
I
I =
-
/V
ThOJ-.w1
ThO.'-402 •••••••• Fig. 3. Columnar sections of the studied bedded
chert section exposed on road between Pai and
Th03391 .1'1 ...v ..;
_ Presence _ _ lnfercd Wieng Hang cities in Ban Tan Jed Ton area (Pai
I
city, Mae Hong Son province), showing distri-
2m Legend: rurtaccccsrocks Cover part bution of characteristic taxa of radiolarians ob-
I
tained.
America (e.g. Murchey 1990), South China (e.g. Wang et al. by Caridroit & De Wever (1986) and from the Prymorye re-
1994) and New Zealand (e.g. Caridroit & Ferriere 1988). It is gion, Far East Russia by Rudenko & Panasenko (1990) and is
comparable to the F. scholasticus - F. venticosus zone from here regarded as F porrectus RUDENKO. Specimens of Follicu-
China (Wang et at. 1994) and the F. ventricosus - Ps. cullus ventricosus ORMISTON & BABCOCK (PI. 1; Fig. 9) from
fusiformis assemblage of North America (Murchey 1990). the study area are closely similar to those reported from N.
America (Ormiston & Babcock 1979; Blome & Reed 1992)
and from S. China (Wang et al. 1994) having a slightly curved
4. Discussion: Follicucullus faunas and palaeobiogeography
apical cone and big pseudothorax, with short pseudoabdomen;
Follicucullus charveti CARIDROIT & DE WEVER is character- both ventral and dorsal flap are very short.
ized by a long apical cone, which is slightly curved to the ven- Materials presented in this study also contains some bro-
tral side (PI. 1; Fig. 8). Specimens from the area we studied ken pieces that have been assigned to species such as Guste-
have a strongly inflated pseudothorax and a short pseudoab- fana obliqueannulata KOZUR (PI. 1; Fig. 30) and Nazarovella
domen with small, disk-like, rather short ventral flap compared phlogidea WANG (PI. 1; Fig. 31). These forms in fact have little
to those reported from Japan and S. China (e.g. Caridroit & taxonomic value and in our collections all of these broken
De Wever 1986; Wang & Li 1994). F. scholasticus ORMISTON & pieces appear to be the ending part of an arm of already de-
BABCOCK m. I (e.g. Ishiga 1985; Blome & Reed 1992; Wang et fined radiolarian species.
al. 1994), and F scholasticus ORMISTON & BABCOCK m. II (e.g. It is not yet proven that siliceous skeleton zooplanktonic
Ishiga 1985) are considered as two different species. The first is faunas are useful indicators for palaeo-ecology and/or palaeo-
the most abundant form and found in all levels of the section. geographic position. Nevertheless, the possible palaeogeo-
This form has a smooth, undulating shell, however, somewhat graphie implications of the occurrence of a particular taxon
obscure separated portions can be observed in some speci- with distinctive form at a few localities merit some discussion.
mens. Small very short flaps, extending at apertural margin of The Upper Permian Follicucullus charveti - F porectus radio-
the dorsal and ventral sides, are well preserved on most speci- larian assemblage recognized in this study is believed to be
mens. The second morphotype, on the other hand, has a bul- equivalent to the F bipartitus- F charveti assemblage known
bous shell characteristic similar to those reported from Japan from southwest Japan (e.g. Caridroit & De Wever 1984, 1986),

I\liddl e PERMIA N
(la te Ca pita nia n)
"'IN
'/:'---
,vf11l1h C(J(I/n..,,.,horinu
... . tb.l.....
lOS
-------------~~~~
Pal_T"lh,'l
Fig. 4. Palaeogeographic reconstruction during

Middle Permian (late Capitanian) showing rela-
tive palaeogeographic positions and possible Fol-
licucullus charveti province discussed in the text.
Shallow Sea (Palaeogeographic map and terrane positions are
. . Follicucullus char veti palaeogeogruphic positions ........ Subduction Zone
based on Scotese 1997; Metcalfe 1996, and water
province areas are after Henderson & Mei 2003,
F. cha rveti province , Radiolarite depos it Land Deep Sea
where ST: Shan-Thai; T: Tarim; WB: West
Burma).
northern New Zealand (e.g. Caridroit & Ferriere 1988; Take- CARIDROIT, M. 1993: Permian Radiolarian from NW Thailand. In: THANA-
mura et al. 1999), and south & southwest China (e.g. Wang & SUTHIPITAK, T. (Ed.): Proceedings of the International Symposium on
Biostratigraphy of Mainland Southeast Asia Facies and Paleontology 1,
Li 1994). According to the known literature reported so far, Chiang Mai, Thailand, 83-96.
this assemblage has only been discovered from the four differ- CARIDROIT, M. & DE WEVER, P. 1984: Description de quelques nouvelles es-
ent geographic areas mentioned above. Regarding Follicucul- peces de Follicucullidae et d'Entactinidae (Radiolaires polycystines) du
Ius charveti CARIDROIT & DE WEVER, a diagnostic taxon of Permien du Japon (Description of some new species of Follicucullidae
and Entactinidae (Polycystin radiolarians) from the Permian of Japan).
this assemblage, it is not clear whether it occurs in a restricted
Geobios 17. 639-644.
area with short period (early Wuchiapigian). The occurrence CARIDROIT, M. & DE WEVER. 1'.1986: Some Late Permian Radiolarians from
of this fauna in a restricted area as plotted on a palaeogeo- pelitic rocks of the Tatsuno Formation (Hyogo Prefecture), Southwest
graphic map may suggest that its geographic range was proba- Japan. Marine Micropaleontology 11, 55-90.
CARIDROIT, M. & FERRIERE, J. 1988: Premieres datations precises du Paleo-
bly confined to a special part in the tropical palaeoequator
zoique par radiolaires en Nouvelle Zealande: interet geologique et pale-
(low-latitude) realm within the equatorial warm water ontologique, Compte rendus Academie des Sceances de l'Academie des
province. As this species can likely be defined as a warm-water Sciences (Paris), series II 306, 321-326.
taxon the palaeogeographic positions of terranes containing CARIDROIT, M., FONTAINE, H., JONGKANJAMASOONTORN, Y., SUTEETHORN, V.
radiolarites with this fauna were probably at low latitudes not & VACHARD, D. 1990: First results of a paleontological study of North-
west Thailand. CCOP Technical Secretariat, 337-350.
far from one another (Fig. 4). CARIDROIT, M., V ACHARD, D. & FONTAINE, H. 1992: Datations par radiolaires
(Carbonifere, Permien et Trias) en Thailande nord-occidentale. Mise en
evidence de nappe de charriage et d'olisostromes. Academie des Sciences
Acknowledgements Compte Rendu Paris 314, 515-520.
CHAUVIROJ, S., CHATURONGKAWANICH, S. LEEVONGCHAROEN, S. & SOPON-
Our field survey in January 2003 was financial supported by the Chiang Mai
PONGPIPAT, P. 1985: Hydrothermal Research Project, Geological Survey
University (Thailand), the Laboratoire de Pale ontologie et Paleogeographie
Division, Department of Mineral Resources, Bangkok, Thailand. 66.
du Paleozoique (CNRS-UMR 8014) and the Universite des Sciences et Tech-
DE WEVER, 1'., DUMITRICA, 1'., CAULET, J.P., NIGRINI, C & CARIDROIT, M.
nologies de Lille (France). The Faculty of Science, Chiang Mai University for
2001: Radiolarians in the Sedimentary Record. 533 p., Gordon and
supporting field-working facilities is credited here. Fabrice Corday and
Breach Science Publishers.
Frances Spiller are thanked for reviews that helped to improve the manuscript.
HADA. S., BUNoPAs, S., ISHII, K. & YOSHIKURA, S. 1999 Rift-drift history and
the amalgamation of Shan-Thai and Indochina/East Malaya blocks In:
METCALFE, I. (Ed.): Gondwana Dispersion and Asian Accretion. IGCP
REFERENCES
321, Sp. Publ., 67-88, A.A. Balkema.
BLOME, C D. & REED, K. M. 1992: Permian and Early (?) Triassic radiolarian HENDERSON, CM. & MEl, S. 1990: Stratigraphic versus environment signifi-
faunas from the Grindstone terrane, central Oregon. Journal of Paleon- cance of Permian serrated conodonts around the Cisuralian-Guadalupian
tology 66, 351-383. boundary: new evidence from Oman. Palaeogeography, Palaeoclimatol-
BUNoPAs,S. 1981: Palaeogeographic history of western Thailand and adjacent ogy, Palaeoecology 191, 301-328.
parts of South-east Asia: a plate tectonic interpretation. Geological Sur- INTAWONG, A., KUNMANEE, J. & WONGANAN, N. 1997: Geological report in
vey paper vo!. 5, Department of Mineral Resources, Bangkok, Special the area of Ban Tan Jed Ton, Tambol Wieng Nue, Amphoe Pai, Chang
Issue, 810. Wat Mae Hong Son. Department of Geological Sciences, Chiang Mai
CARIDROIT. M. 1991: Taxonomic study on Carboniferous and Permian Radio- University, 82.
laria from NW Thailand. Paleontologic. stratigraphic and tectonic signifi- ISHIGA, H. 1985: Discovery of Permian radiolarians from Katsumi and Oi For-
cances. Abstracts. Sixth Meeting, International Association of Radiolari- mation along south of Maizuru Belt, Southwest Japan and its significance.
an Paleontologists (INTERRAD VI), p. 21. Ibid 39, 175-185.

ISHIGA, H. 1986: Late Carboniferous and Permian radiolarian biostratigraphy SASHIDA, K., IGo, H., HISADA, K., NAKORNSRI, N. & AMPORNMAHA, A. 1993:
of southwest Japan. Journal of Geoscience, Osaka City University 29, Occurrence of Paleozoic and early Mesozoic Radiolaria in Thailand (pre-
89-100. liminary report). Journal of Southeast Asian Earth Science 8, 97-108.
ISHIGA, H. 1990: Paleozoic radiolarians. In: ICHIKAWA, K., MIZUTANI, SASHIDA, K., IGo, H., UENO, K., NAKORNSRI, N. & SARDSUD, A. 1998: Late
S.,HARA, 1., HADA, S. & YAO, A. (Eds.): Pre-Cretaceous terranes of Paleozoic radiolarian fauna from northern and northeastern Thailand.
Japan. Publication of IGCP Project No. 224: Pre-Jurassic Evolution of Science Reports of the Institute of Geoscience, University of Tsukuba,
Eastern Asia, Nippon Insatsu Shuppan, Co. Ltd, Osaka, 285-295. Section B 19, 1-27.
METCALFE, 1. 1996: Gondwana dispersion, Asian accretion and evolution of SASHIDA, K., IGo, H., ADASHI, S., UENO,K., KAJIWARA, Y., NAKORNSRI, N. &
Eastern Tethys. Australian Journal of Earth Sciences 43, 605-623. SARDSUD, A. 2000: Late Permian and Middle Triassic radiolarian faunas
MURCHEY, B.L. 1990: Age and Depositional setting of siliceous sediments in from northern Thailand. Journal of Paleontology 74, 789-811.
the Upper Paleozoic Havallah sequence near Battle Mountain, Nevada: SCOTESE, K. 1997: Paleogeographic Atlas Paleomap Progress Report 90-0497.
implications for the paleogeography and structural evolution of the west- Paleomap Project ed., Univ. of Texas at Arlington, Arlington, Texas, 45.
ern margin of North America. Geological Society of America, Special TAKEMURA, A., MORIMOTO, T., AlTA, Y., HORI, R.S., HIGUCHI, Y., SPORLI,
paper 255, 137-155. K.B., CAMPBELL, H.L., KODAMA, K. & SAKAI, T. 1999: Permian Albaillel-
ORMISTON, A. & BABCOCK, L. 1979: Follicucullus, new radiolarian genus from laria (Radiolaria) from a limestone lens at the Arrow Rocks in the Waipa-
the Gaudalupian (Permian) Lamar Limestone of the Delaware Basin. pa Terrane, Northland, New Zealand. In: DE WEVER, P. & CAULET, J.-P.
Journal of Paleontology 53, 328-334. (Eds.): InterRad VII, Paris/Bierville, September 1997. Geodiversitats
RUDENKO, V. & PANASENKO, E.S. 1990: A new findings of the Upper Permian 21(4),789-811.
radiolarians in Prymorye region. In: ZAKHAROV, Y.D., BELYAEVA, G.V. WANG, Y. & LI, J. 1994: Discovery of the Follicucullus bipartitus -F. charveti
& NIKITINA, A.P. (Eds.): New data on Paleozoic and Mesozoic biostratig- radiolarian assemblage zone and its geological significance. Acta Mi-
raphy of the South Far East. Far Eastern Branch of the USSR Acad. of cropalaeontologica Sinica 11, 201-212.
Sci., Vladivostok, 117-124. WANG, Y., CHENG, Y.N. & YANG, Q. 1994: Biostratigraphy and systematics of
SASHIDA, K. & IGo, H. 1999: Occurrence and tectonic significance of Paleozoic Permian radiolarians in China. Paleoworld 4,172-202.
and Mesozoic Radiolaria in Thailand and Malaysia. In: METCALFE, 1. WONGANAN, N., CARIDROIT, M. & RANDoN, C. 2002: Radiolarians and con-
(Ed.): Gondwana Dispersion and Asian Accretion Final Results Volume odonts from radiolarites in NW-Thailand; witnesses of a 140 my (at least)
for IGCP Project 321, A.A. Balkema, Rotterdam, 175-196. oceanic realm. In: The 64 th Palaeontological Association Annual Meeting,
SASHIDA, K. & NAKORNSRI, N. 1997: Lower Permian radiolarian faunas from Cambridge, abstract. The Palaeontology Newsletter 51, 73.
the Khanu Chert Formation distributed in the Sukhothai Area, northern
central Thailand. In DHEERADILOK, P., C. HINTHONG, P. CHAODUMRONG,
P. PUTTHAPIBAN, W. TANASTHIEN, C. UTHA-AROON, N. SATTYARAK, N.
NUCHANONG & T. TECHAWAN (Eds.): Proceedings of the International Manuscript received January 2004
Conference on Stratigraphy and Tectonic Evolution of Southeast Asia Revision accepted March 2005
and the South Pacific, Bangkok, Thailand, 101-108.
Plate 1
Permian radiolarians from Pai area, northwestern Thailand. All figures are scanning electronic micrographs. Scale bar 100 urn applies to all specimens. 1-6. Fol-
licucullus scholasticus ORMISTON & BABCOCK, PAI-421; 7. Follicucullus porreetus RUDENKO, PAI-421; 8-10. Follicucullus charveti CARIDROIT & DE WEVER, PAI-
421; 11. Follicucullus monacanthus ISHIGA & IMOTO, PAI-421; 12-13. Follicucullus ventricosus ORMISTON & BABCOCK, PAI-419; 14. Follicucullus orthogonus
CARIDROIT & DE WEVER, PAI-421; 15-17. Pseudoalbaillella sp. A, PAI-421; 18. Neoalbaillella sp. ct. N. grypa ISHIGA ETAL., PAI-421; 19. Albaillella sp. aff. A.
levis ISHIGA ET AL., PAI-421; 20. Nazarovella gracilis DE WEVER & CARIDROIT, PAI-402; 21. Quadricaulis femoris CARIDROIT & DE WEVER, PAI-393; 22. Ishi-
gaum (?) similicutis CARIDROIT & DE WEver, PAI-393; 23. Ishigaum trifustis DE WEVER & CARIDROIT, PAI-393; 24. Trianospongos musashiensis SASHIDA &
TONISHI, PAI-393; 25. Latentifistula texana NAZAROV & ORMISTON, PAI-398; 26. Ormistonella robusta DE WEVER & CARIDROIT, PAI-393; 27. Pseudotormentus
sp. ct. P. kamigoriensis DE WEVER & CARIDROIT, PAI-402; 28. Copicyntra sp., PAI-393; 29. Hegleria mammila (SHENG & WANG),PAI-393; 30--31. broken part of
latentifistulinid (30: ct. Gustefana obliqueannulata KOZUR; 31: ct. Nazarovella phlogidea WANG), PAI-393.


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Supplement 1:

Eclogae Geologicae Radiolaria

Zeitschrift der I Revue de la

BAUMGARTNER, P. O. FENG, Q., MENG,Y., HE, W., Gu, S.:

DOLVEN, J. K., SKJERPEN, H. A.:

UNIL I Univers ite de Lausanne

Radiolaria - Siliceous plankton through time

INTERRAD X, held at the University of Lausanne, Switzerland, September 2003

Turonian Radiolarians from Karnezeika, Argolis Peninsula,

Turonian Radiolarians from Karneze ika SI

Dhidh lm i·Trapezona Composite Unit

S2 A.N . Band ini, P.O. Baumgartner & M. Caron

UPPER CRET. ALL OCHTHONOUS SERI ES

DHIDHIMI·TRAPEZONA COMPOSITE UNIT

Pelag ic limestones & marls

Migdhalitsa Ophiolite Unit

Basa l Seq uence

~ Pantokrato r Li mes ton e

Turo nian Radiolarians from Karn ezeik a S3

Stratigra phic log of the studied part

Dolom itic breccias blok s of several meters

Oph iol ite-carbonate breccias with ol istoliths of

S4 A.N. Bandini, P.O. Baumgartner & M. Caron

Acan thocircus hueyi

Archaeocenosphaera (7) mellifera

Turonian Radiolarians from Karn ezeika S5

Rtedet and SanfiJi 1974

Patel/ufa he/ios Durmrnca 1975

S6 A,N, Bandini , P.O. Baumgartner & M. Caron

Pseudoa woph a cus p umh en~s

Tu roni an Radiolarians from Karnezeika S7

Triactome cellulose Dumitrica 197 5

Triectom e hexeris Dumrtnca 1 75

S8 A.N. Bandini , P.O. Baumgartner & M, Ca ron

IPseud08ulophaeus seulptus <I> ? i

IArehaeoeenosphaera (?) mellifera l

I Triaetoma cettutose I 21 I <t--I> ?I I I I I I I I I I

Aeanthocircus venetus 10 -e > ? I I

IMarginotruncana marianosi CKAR7

Diearinella primitiva I CKAR 14

Margino truneana sigali , CKAR 13 & All6 1350

Marginotruneana sehneegansi CKAR 13 & 14

Marginotruneana corona ta > CKAR 13

Marginotruneana pseudolinneiana > CKAR 14 & AIl6 1350

IMarginotruneana sinuosa 1 I I I I I I I CKAR 14 I

Tu roni an Radiolarians from Karnezeik a S9

Family P ARVIVACCIDAE PESSAGNO & YANG in PESSAGNO

Acaeniotyle sp, B Pseudoacanthosphaera galeata Q 'DOGHERTY 1994

Family XIPHOSTYLIDAE HAECKEL 1881 Pseudoacanthosphaera superba (SQUINABOL 1904)

S10 A.N. Bandini , P.O. Baumgart ner & M. Caron

Genus PRAECONOCARYOMMA P ESSAGNO 1976

Praeconocaryomma californiaensis PE SSAGNO 1976 Genus HALESIUM PESSAGN O 1971

Genus PESSAGNOBRACCHIA K O WR & MosTLER 1978

Tu ron ian Radiolarians from Karnezeika S11

Sup erfamily SPONGURACEA H AECKEL 1862

S1 2 A .N. Bandini, P.O. Baumgartner & M . Caron

Dietyom itra formosa SQ UIN ABOL 1904

Stichomitra communis SQ UINABOL 1903

Turoni an Radiolarians from Karnezeika S 13

Genus AFENS RIEDEL & SANFILIPPO 1974 Acknowledgements

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