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Restudy of the holotype of Operculodinium centrocarpum (Deflandre &

Cookson) Wall (Dinophyceae) from the Miocene of Australia, and the
taxonomy of related species

Article  in  Palynology · December 1997

DOI: 10.1080/01916122.1997.9989484

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Restudy of the Holotype of Operculodinium centrocarpum (Deflandre & Cookson) Wall

(Dinophyceae) from the Miocene of Australia, and the Taxonomy of Related Species
Author(s): Kazumi Matsuoka, Andrew McMinn, John H. Wrenn
Source: Palynology, Vol. 21 (1997), pp. 19-33
Published by: American Association of Stratigraphic Palynologists, Inc.
Stable URL: http://www.jstor.org/stable/3687586
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RESTUDY OF THE HOLOTYPEOF OPERCULODINIUM
CENTROCARPUM (DEFLANDRE& COOKSON)WALL
(DINOPHYCEAE)FROMTHE MIOCENEOF AUSTRALIA,
AND THE TAXONOMYOF RELATEDSPECIES

KAZUMIMATSUOKA
Departmentof Geology
Faculty of LiberalArts
NagasakiUniversity
Nagasaki, 852
Japan

ANDREW McMINN
Instituteof Antarcticand SouthernOcean Study
Universityof Tasmania
GPO Box 252C Hobart
Tasmania
Australia

JOHNH. WRENN
Centerfor Excellence in Palynology
Departmentof Geology & Geophysics
LouisianaState University
Baton Rouge, LA 70803, U.S.A.

Abstract

Operculodiniumcentrocarpum(Deflandre & Cookson) Wall This restudyalso confirmedthat 0. centrocarpum(Deflandre&

1967 is one of the most commonly reporteddinoflagellatecysts
(dinocysts) in samples from Tertiaryand Quaternarydeposits.
However,atleasttwo distinctdinocystmorphologiesareassigned
to this species in the literatureat the presenttime. In addition,two
separate species have been erected for one of these distinct
morphologies.
In order to define criteria with which to differentiatethese
morphologies,holotypeandtopotypematerialof 0. centrocarpum
(Deflandre& Cookson)Wall 1967 and 0. echigoense Matsuoka
1983wererestudiedandcomparedwithdinocystsofProtoceratium
reticulatum(Claparede& Lachmann)Biitschli 1885.
Re-investigation of the holotype and topotype specimens
of 0. centrocarpum (Deflandre & Cookson) Wall 1967
(=HystrichosphaeridiumcentrocarpumDeflandre & Cookson
1955) and 0. echigoense Matsuoka 1983 demonstratedthat 0.
echigoense Matsuoka 1983 is a junior synonym of the former.
Cookson)Wall 1967 (= H. centrocarpumDeflandre& Cookson
1955) is morphologicallydistinctfromthe Quaternarydinocysts
thatare widely attributedto this species in the literature.[These
Quaternaryformsarereferredto in this paperas 0. centrocarpum
(Deflandre& Cookson) sensu Wall 1967.] Viable cysts that are
morphologically identical to the Quaternaryforms [i.e., O.
centrocarpum(Deflandre & Cookson) sensu Wall 1967] have
been incubatedto producethemotiledinoflagellateP. reticulatum
(Clapar6de& Lachmann)Btitschli 1885.
We proposethat, 1) the lattername should be appliedonly to
large, morphologicallydistinct dinocysts comparableto those
originally describedfrom the Miocene of Australia,and 2) the
distinctiveQuaternaryforms [i.e., 0. centrocarpum(Deflandre&
Cookson) sensu Wall 1967] should be referredto as the cysts of
Protoceratiumreticulatum(Claparede& Lachmann)Biitschli
1885.

Palynology,21(1997):19-33
? 1997by AASPFoundation ISSN0191-6122
20
INTRODUCTION
Specimens of various species of OperculodiniumWall
1967 are a prominentcomponent of most Neogene and
Quaternarydinoflagellate cyst assemblages. Species of
Operculodiniumareusuallycharacterizedby a sphericalto
subsphericalcentral body, a precingulararcheopyle and
numerousintratabular or nontabular,solid processes.Neo-
gene species currentlyassignedto Operculodiniuminclude
0. centrocarpum(DeflandreandCookson)Wall,0. crassum
Harland,0. delicatum Kar, O. echigoense Matsuoka,O.
longispinigerumMatsuoka, 0. majus Jain & Dutta, O.
paucispinosum Kar, 0. wallii Matsuoka, 0.? erikianum
Headet al., O.janducheneiHeadet al., and 0. vacuolatum
Head et al.
The holotype of the genus, 0. centrocarpum(Deflandre
& Cookson) Wall 1967, was described from Australian
Miocene deposits by Deflandre and Cookson (1955) as
Hystrichosphaeridiumcentrocarpum.Wall (1967) subse-
quentlyreportedanddescribedspecimensthathe assigned
to thatspecies from Pleistocene deposits in the Caribbean
Sea, and simultaneously transferredthe species to the
genus Operculodinium,which he establishedin the same
paper.Wall and Dale (1968) successfully incubatedmod-
ern cysts that they identified as being identical to the
Pleistocene dinocysts thatWall (1967) had referredto O.
centrocarpum(Deflandre & Cookson) Wall 1967. The
incubatedmotile thecate dinoflagellate was identified as
Protoceratium reticulatum (Claparede & Lachmann)
Butschli 1885.
However, it has become evident since then that at least
two distinctdinocyst morphologieshave been assignedin
the literatureto 0. centrocarpum(Deflandre& Cookson)
Wall 1967. This apparentproblemwas addressedin round-
table discussions in 1989 and 1994 (Head and Wrenn,
1992; Head, 1995). Recent work on modern,Quaternary,
and Neogene dinoflagellate cyst assemblages by us has
demonstratedthatthereis a substantialdifferencebetween
the modern and Quaternary forms attributed to O.
centrocarpum(Deflandre& Cookson) Wall 1967 and the
holotype of that species. In this paper we redescribeand
illustrate the holotype and topotype material of O.
centrocarpum(Deflandre & Cookson) Wall 1967, and
compareit with 0. echigoenseMatsuoka1983 andcysts of
Protoceratium reticulatum (Claparede & Lachmann)
Butschli 1885.
PLATE 1
Holotypeof Operculodinium centrocarpum (Deflandre& Cookson)Wall 1967.Slide P15654of the NationalMuseumof Victoria,
Melbourne.Loc.GellibrandClayat BalcombeBay,Victoria,Australia.
Age:MiddleMiocene.
1-6 Ventralview of 1, ventralsurface,2-6, successively
lowerfoci.
21 - 1997
VOLUME21-1997
PALYNOLOGY,
TheholotypeofO. centrocarpum(Deflandre&Cookson)
Wall 1967 was recovered from the Middle Miocene
GellibrandClay [now the FyansfordFormation(Abele et
al., 1976)] fromBalcombeBay, on theeasternshoreof Port
Phillip, Victoria of Australia. The formation crops out
along a 2 km stretchof waterfrontandis mostly composed
of calcareous clayey siltstone that attains a maximum
thicknessof 50 m. It is not possible now to determinethe
precise location from which of the sample studied by
Delandreand Cookson (1955) was collected. The forma-
tioncontainsbotha richmegafaunaandmicrofauna,which
provide a date of late Early Miocene to early Middle
Miocene for the unit [N8 to N9 planktonicforaminiferal
zones of Blow (1966), accordingto Abele et al. (1976) and
Gostin(1966)]. The dinoflagellateassemblage,whichcon-
tains the age diagnostic species Dapsilidiniumpastielsii,
Heteraulacacysta campanula, Impagidiniumjaponicum
andReticulatosphaeraactinocoronata,indicatesa similar
age. Details of the dinoflagellate assemblage, obtained
from topotypematerial,are presentedin Appendix 1.
MATERIALSAND METHODS
The holotype of Hystrichosphaeridiumcentrocarpum
Deflandre & Cookson 1955 is on slide P 15654 at the
NationalMuseum of Victoria, Melbourne,and was made
availablefor the presentstudy.The holotypewas collected
from the Middle Miocene GellibrandClay at Balcombe
Bay; other specimens reported in the same paper were
recovered from Bore No.1 (514-516 feet) drilled near
Birregurra,Victoria.Topotypematerial,fromtheBalcombe
Bay location, also was providedby the NationalMuseum
of Victoriato supplementour study of the holotype.
The holotypeof 0. echigoenseMatsuoka1983 is located
on Slide NNA4-1(12.1/125.4; S26/2) and is curatedin the
Departmentof Geology, Facultyof LiberalArts,Nagasaki
University. The holotype was obtained from the early
MiddleMioceneNanataniFormationatMinami-Imogawa,
Shitada-Mura,Niigata Prefecture,Japan.
Modern specimens, previously assigned to O.
centrocarpum(Deflandre & Cookson) Wall 1967 were
examinedfrom Hachinoheoff the Pacific coast of Japan,
Lake Hirugaalong the coast of the Sea of Japan,the east
coast of Australia,and Tasmania.AdditionalQuaternary
specimenswereexaminedfromthePleistoceneNishiyama
K. Matsuoka,A. McMinn, J.H. Wrenn:Holotype of Operculodiniumcentrocarpum Plate 1

.1 i2
22
FormationandtheHolocene sedimentsof the Sea of Japan;
ODP sites 765 and 820 from northernAustralia,cores PC-
7 andNB-3b from easternAustralia;surfacesedimentsof
moderncontinentalslope of the easternU.S.A. andmodern
surfacesedimentsin CastleHarbor,Bermuda,in the Atlan-
tic Ocean.
Re-examinationof the holotype of 0. centrocarpum
(Deflandre& Cookson)Wall 1967 was carriedoutby K.M.
and A.M. at the Instituteof Antarcticand SouthernOcean
Studies,Universityof Tasmania,withZeiss Axioscope and
Nikon Labophotomicroscopeswith phase contrastoptics.
Photographyof the holotype was carriedout on a Zeiss
Photomicroscope II at the N.S.W. Geological Survey,
Sydney.Theholotypeof 0. echigoenseMatsuoka1983and
otherJapanesespecimenswerere-examinedby K.M. atthe
Departmentof Geology, Nagasaki University, using a
Nikon Biophoto microscope with interferenceand phase
contrastoptics.
SYSTEMATICPALYNOLOGY
Includedbelow are results of our study of the holotype
andtopotypesof Operculodinium centrocarpum(Deflandre
& Cookson) Wall, the holotype and paratypes of
Operculodiniumechigoense Matsuokaand on cysts of P.
reticulatum(Claparede& Lachmann)Bitschli. The genus
Operculodiniumis emendedand 0. echigoense Matsuoka
1983 is synonymized with the senior synonym O.
centrocarpum(Deflandre & Cookson) Wall 1967 (= H.
centrocarpumDeflandre & Cookson 1955). In addition,
the latter species is differentiatedfrom 0. centrocarpum
(Deflandre& Cookson)sensu Wall 1967, the cyst stage of
P. reticulatum(Claparede& Lachmann)Butschli.
Class DINOPHYCEAEPascher 1914
OrderGonyaulacalesTaylor 1980
Family GonyaulacaceaeLindemann1928
Genus OperculodiniumWall 1967 emend.
Type Species. Operculodinium centrocarpum
(Deflandre & Cookson) Wall 1967 = Hystricho-
sphaeridiumcentrocarpumDeflandre & Cookson 1955
Original Diagnosis. Sphericalto ovoid cysts possessing
simple, dorsal precingulararcheopyles (reflecting plate
3"), and lacking polar structures.A weakly defined girdle
andventralsulcaldepressionoftenpresent.Cell walldouble,
the inner thin and the outer thickerand microgranularor
microreticulate.Elementsof ornamentationvariable,from
VOLUME21 -1997
PALYNOLOGY,
small cones to long spines, but all with circular,minutely
striatedbasesandoftencapitateextremities.Spinearrange-
ment intratabularwith several spines to each plate-area,
commonlyarrangedimmediatelywithinthereflectedplate-
areamargins."(Wall, 1967, page 110.)
Emended Diagnosis. Sphericalto ovoid cysts possess-
ing a 3" precingulararcheopyle and lacking polar struc-
tures.A weaklydefinedgirdleandventralsucaldepression
may or may not be present.Cyst wall composed of a thin,
smooth endophragm and a thicker microgranular or
microreticulateperiphragm.Positive sculpturing varies
from small cones to long processes with circularbases.
Processesbases maybe striateormicroreticulate.A variety
of distalprocessshaftterminationsmay be present,includ-
ing capitate,recurved,truncatedorpatulate,thoughusually
one type per species. Processes distributionnontabular
and/or intratabular,with several processes per paraplate
area.
Remarks. Operculodiniumis emendedhereinto include
nontabularprocesses and a widervarietyof processtermi-
nationsthanwas includedin the originalgenericdiagnosis.
Operculodiniumcentrocarpum
(Deflandre& Cookson) Wall 1967
Plate 1, figs. 1-6; Plate 2, figs. 1-9
Hystrichosphaeridium centrocarpum Deflandre &
Cookson, 1955, p. 272-273, pl. 8, figs. 3-4.
OperculodiniumechigoenseMatsuoka,1983, p. 126, pl. 7,
figs. 1-5, 8.
Operculodiniumwallii Matsuoka;Mudie, 1987, p. 805, pl.
2, fig. 1; Mudie, 1989, pl. 2, fig. 6.
Non 0. echigoense;Mudie, 1987, p. 804-805, pl. 2, fig. 7.
Diagnosis. Choratecyst with a sphericalto subsphical
centralbody, and microreticulatesurfacesculpturingthat
extends onto, and is coarser on, each process base. Pro-
cesses are nontabularand have solid, slender, shafts of
variablelength.Ususally unbranched,butrarelybranched.
Shafts taperdistally, though slightly widened on the end,
and process terminationsare recurvedor patulate.
Holotype. Slide P 15654, illustratedby Deflandreand
Cookson (1955), pl. 8, figs. 3,4; lodged at the National
Museum of Victoria, Melbourne, Australia. Topotype.
P-81529-A02 SPL #6 15614, illustratedherein,P1.1, figs.
1-6;lodgedin thePaleontologicalCollectionof theUniver-
sity of Tasmania.
Original Description. "Shell globular, more or less
ellipsoidal (probablyby deformation),covered with nu-
merous straight,slender processes placed irregularlyor
with a certainalignment.Processes solid, showing radiat-
K. Matsuoka,A. McMinn, J.H. Wrenn:Holotype of Operculodiniumcentrocarpum
I I
ing fibrils at theirpoint of intersectionon the shell, which
is clearly ornamentedwith a fine reticulum.The slightly
widenedapicesof theprocessesarefringedwith 10 ormore
small curved spines (this structureis only visible with a
high-poweredobjective)."(Delandreand Cookson, 1955,
pages 272-273.)
Redescription of the Holotype. The original shape of
the centralbody was probablysubsphericalbuthas subse-
quentlybeen deformedby compressionandshrinkage.(A
second specimen present on the same slide also is de-
formedbut was probablysubsphericalin shape too. ) The
holotype cyst wall is composed of a 2 ,jm thick, fibrous
periphragmand a thin, smooth endophragm.The central
body surface ornamentationis microreticulate;around
and on the process bases it is coarser than the rest of the
cyst surface. The solid, slender shafts of the nontabular
processes are smooth, distally tapered, and sometimes
flexible. Process shafts vary in length and are rarely
branched.Distal terminationsare slightly expanded and
either recurvedor patulate.The archeopyleis not clearly
visible but is presumed to be precingular.No accessory
archeopyle sutures were observed. The paracingulumis
undifferentiatedbut may be markedby a few relatively
short processes that are present opposite the presumed
archeopyle opening. The position of the parasulcus is
probablyreflected by a numberof shorter,more slender
processes, also opposite the archeopyle.
Dimensions.
Deflandre&
Cookson (1955) Measuredherein
Cyst diameter 54-80 gim 75 gm
Length of processes 13-18 gim ca. 17 im
Remarks. Specimens of other dinoflagellate species
on the holotype slide include Spiniferites bulloideus (=
Hystrichosphaera bulloidea Deflandre & Cookson),
Spiniferitessp.indet.,andOperculodinium longispinigerum
Matsuoka.Theholotypeslide ofH. centrocarpumis in very
poor condition; it contains many small air bubbles, and
most of the glycerinejelly is dehydrated.Pollen, spores,
andotherdinoflagellatesthatoccur on the slide arelightly
stained with Safranin O. Coiled and biserial
microforaminifers,radiolarianfragments,and sponge spi-
cules areabundant,as arelight-coloredmineralgrains.The
thickness of the preparationpreventsexaminationwith a
lOOxoil objective.
Description of the Topotype. Because of the poor
condition of the holotype, a topotype is designated and
described herein. The materials containing the topotype
werecollected fromthe late Earlyto earlyMiddleMiocene
GellibrandClay at Balcombe Bay of Victoria, Australia,
23
and its slide, P-81529-A02 SPL #6 15614, is storedin the
PalaeontologicalCollectionof theUniversityof Tasmania.
A list of species co-occurring on the topotype slide is
presentedin Appendix 1.
The topotype cyst is subspherical, and slightly de-
formed.The surfaceof the cyst is microreticulate,with the
reticulationextending onto and becoming coarser at the
base of the processes. Processes are nontabular,solid,
slender, tapering, and sometimes flexible. The process
shaftsaresmoothexcept at the base andthe distal ends are
patulate or rarely recurved. The precingular(type P) 3"
archeopyleis pentagonalin outline with roundedcorers.
Low septa appearto indicate the paracingulum,and the
parasulcus is probably reflected by a cluster of 3 to 5
relatively shortandmore slenderprocesses presenton the
ventral surface.
Dimensions (five specimens measured).
Cyst diameter 56-76 jim (average68 jm)
of
Length processes 11-17 gum(average 13 gm)
Discussion.
"0. echigoense Matsuoka1983"
Re-examinationof the holotype and paratypesof "O.
echigoense Matsuoka 1983" were conducted. "0.
echigoense Matsuoka 1983" is intermediate in size,
subsphericalto ellipsoidal in shape and its surfaces are
microreticulate,or more rarely,microgranulate.The base
of eachprocessis microreticulateratherthanmicrogranulate,
andfibrous.Thesolid andslenderprocessesarenontabular,
tapered,and relatively long and flexible. The distal ex-
tremities are apparentlycapitate, bifid, or recurved.The
Type P precingulararcheopyleis formedby the loss of the
3" paraplate.The paracingulumand parasulcusare not
clearly indicated, though the parasulcus may be repre-
sented by a cluster of relatively slender and shorterpro-
cesses on the oppositeside of the cyst fromthe archeopyle.
This re-examination indicates that "0. echigoense
Matsuoka1983"is a junior synonym of 0. centrocarpum
(Deflandre & Cookson) Wall 1967. Forms formerly as-
signed to "O. echigoense Matsuoka 1983 ," but now in-
cluded in 0. centrocarpum(Deflandre& Cookson) Wall
1967 are shown in Plate 3, figs. 1-12.
Dimensions.
Holotype Otherspecimens
Cyst diameter 72 gim 67-86 gim
Lengthof processes 12 im 12-21 gm
24
0. centrocarpum(Deflandre& Cookson)
sensu Wall 1967
The Quaternaryspecimens recovered by Wall (1967)
from cores in the Caribbean Sea and assigned to O.
centrocarpum(Deflandre& Cookson) Wall 1967 are re-
ferredto in this paper as 0. centrocarpum(Deflandre&
Cookson) sensu Wall 1967. Specimens of this dinocyst
morphologyare illustratedon Plate 4 of this paper.
These cysts have a subsphericalcentral body, whose
surfaceis microgranularsurfaceanddensely coveredwith
radiatingprocesses. The solid processes are slender and
have conical bases bearing minute striations.The distal
extremitiesof the processes bear small hooklets or minute
platforms.The Type P precingulararcheopyleis large.The
paracingulumand parasulcusare undifferentiated.
The cysts recovered by Wall (1967) are identical to
cysts producedby Protoceratiumreticulatum(Claparede
& Lachmann) Biitschli 1885 (see below), not to
Operculodiniumcentrocarpum(Deflandre & Cookson)
Wall 1967 = Hystrichosphaeridium centrocarpum
Deflandre & Cookson 1955.
Dimensions. (fromWall, 1967).
Cyst diameter 40-56 gm
Process length 8-16 im
Protoceratiumreticulatum
(Claparede& Lachmann)Butschli 1885
Operculodiniumcentrocarpum
(Deflandre& Cookson) sensu Wall 1967
Remarks. Braarud(1945) originallydescribedcysts of
P. reticulatum(Claparede& Lachmann)Butschli 1885
from unialgal cultures. Evitt and Davidson (1964) de-
scribedspecimenscollected in planktontows fromDr0bak
sound, Oslo Fjord,and discussed theresignificanceof the
cysts. Wall and Dale (1968) incubated cysts that they
identified as 0. centrocarpum(Deflandre & Cookson)
sensuWall(1967), producingmotilecells of P. reticulatum.
PLATE2
Topotype of Operculodiniumcentrocarpum(Deflandre& Cookson) Wall 1967. Slide: P-81529-A02 SPL #6 15614, late Early to early
Middle Miocene. Palynology ResearchCenter;storedat the PalaeontologicalCollection of the Universityof Tasmania.
1-6 Different focus levels from dorsal to ventral surface
underbrightfield.
PALYNOLOGY, 21 - 1997
VOLUME21-1997
Severalotherauthorshave subsequentlydescribedthe cyst
of P. reticulatumbased on cyst incubationstudies (Bolch
and Hallegraeff, 1990; Matsuoka, 1990). Specimens de-
scribed by Wall and Dale (1968) were from the Atlantic
Ocean,whereasthose describedby Matsuoka(1990) were
collected from the Pacific Ocean. The former authors
report that "the central body is spherical and densely
ornamentedwithmoderatelylong (6 to 16 gim)intratabular
or notabularspineswith capitateandminutelyhookedtips.
The archeopyleis a mid-dorsal,precingular(3"is spherical
to subsphericalwithmicrogranulatesurface,andpossesses
partlyintratabularandmostly nontabular,slenderlycylin-
drical to tubiformshafts with capite extremities distally
closed, andhave the precingulararcheopyleformedby the
loss of the 3"paraplate.These authorsdescribedthe cysts
of P. reticulatumas being approximatelyspherical with
microgranulatesurfaceornamentanddenselycoveredwith
moderatelylong, intratabularor nontabularprocesses, the
extremitiesof which are either capitateor bearingminute
platforms.Theprecingulararcheopyleis formedby theloss
of the 3"paraplate.
Modern Cysts Dimensions.
Cyst diameter Process length
Atlantic specimens
(Wall and Dale, 1969) 33-37 gm 6-16 gm
Pacific specimens
(Matsuoka,1990) 28-50 ,um 4-8 im
Occurrence. Cysts of P. reticulatumhave been widely
reportedfrommost partsof the world.A totalsynonymyis
notpresentedhere.Rather,a summary,by regions,of many
of the reportsthat include illustrationsof the cysts of P.
reticulatumis presentedbelow:
North Pacific including some marginal seas: Matsuoka
(1976a, 1976b, 1985,1987)
South Pacific: McMinn(1991, 1992)
CaribbeanSea: Wall (1967)
North Sea: Harland(1983), Matthiessen(1991), Nehring
(1994)
NorthAtlantic:Wall et al. (1971)
7-9 Different focus levels from dorsal to ventral surface
underinterferencecontrast.
K. Matsuoka,A. McMinn, J.H. Wrenn:Holotype of Operculodiniumcentrocarpum Plate 2

I
26
The relationshipof processlengthto cyst diameteron O.
centrocarpum(Deflandre & Cookson) Wall is shown in
Text-Figure1. This figureis based on specimenscollected
from within and aroundJapan,and referredto either "O.
echigoense" or 0. centrocarpum(Deflandre& Cookson)
Wall 1967 by Matsuoka (1983). The cyst diameter and
process length are positively correlated. However, two
differentclusters are clearly present. One consists of the
smallercysts (approximately30 gimto 60 gimin diameter)
with shorterprocesses (4 gLmto 12 gim),and the other,of
largercysts (approximately70 gimto 100 gm in diameter)
withlongerprocesses(15 gimto 25 gm). These two clusters
are also differentiatedstratigraphically;namely most Mi-
ocene andPliocene specimenshave a greaterdiameterand
longerprocesses thando Pleistocene, Holocene, andmod-
em specimens. These data suggest that "0. echigoense"
and 0. centrocarpumare taxonomically independentof
one anotherbut phylogeneticallyrelated.
DISCUSSION
A re-investigationof the holotype of 0. centrocarpum
(Deflandre & Cookson) Wall 1967 has revealed several
importantmorphologicalfeaturesthatclearlydifferentiate
it fromthe Quaternaryandmodem cysts of P. reticulatum.
The holotype of 0. centrocarpum(Deflandre& Cookson)
Wall 1967 is largerand has a microreticulateratherthan
microgranulatesurface ornamentation.In addition, the
microreticulationis coarser around and on the process
bases, and the distal process terminationsare patulateor
recurvedratherthan capitatelike those on the cysts of P.
reticulatum.These differencesaresufficientto separateO.
centrocarpum(Deflandre& Cookson)Wall 1967 fromthe
Quaternaryand modern cysts of P. reticulatum[i.e., O.
centrocarpum(Deflandre& Cookson)sensuWall (1967)].
PLATE3
Opercutodiniumcentrocarpum (Deflandre& Cookson)Wall1967(= "Operculodinium
Referencesaregivenin brackets[ ].
1-3 "Holotype",Nanatani Formation,loc. Minami-
Imogawa,Shitada-Mura,NiigataPrefecture,Japan.
Age;earlyMiddleMiocene;slideNNA4-1,12.1/125.4
[S26/2].
4-5 "Topotype",Nanatani Formation,loc. Minami-
Imogawa,Shitada-Mura,NiigataPrefecture,Japan.
Age;earlyMiddleMiocene;slideNNA4-5[C25/3].
PALYNOLOGY,VOLUME21 - 1997
PALYNOLOGY,VOLUME 1997 -
"0. echigoenseMatsuoka1983"is ajunior synonymof O.
centrocarpum(Deflandre & Cookson) Wall 1967 (= H.
centrocarpumDeflandre& Cookson 1955).
The cyst of P. reticulatumis probablythe most abundant
dinoflagellatecyst in Quaternarysedimentsand it is also
common in many Neogene, particularlyPliocene, assem-
blages. It has been widely recordedfrom estuaries,conti-
nentalshelves, anddeep ocean sites frommost continents.
We thereforerecognize the confusion that renamingthis
species may cause and do not recommend this change
lightly. 0. centrocarpum(DeflandreandCookson)Wall (=
H. centrocarpumDeflandre& Cookson 1955) andcysts of
P. reticulatum,however,arenotendmembersof a morpho-
logical spectrum.Theycanbe clearlydifferentiatedon size,
wall structure,and process form.
They also appear to have different time-stratigraphic
ranges with 0. centrocarpum(Deflandre and Cookson)
Wall (= H. centrocarpumDeflandre & Cookson 1955)
being reported only from Pliocene or older deposits
(Deflandre and Cookson, 1955; Matsuoka 1983, as "O.
echigoense", McMinn in press a, b, as "0. echigoense").
ThecystsofP. reticulatum[i.e., 0. centrocarpum(Deflandre
& Cookson) sensu Wall (1967)] are common only in
deposits of Quaternaryage.
The dual system of nomenclaturefor dinoflagellates,
with one namefor the motile form andanotherfor the cyst,
is unacceptableto us. Wherethereis generalagreementthat
themotile andcyst formsrepresentthe sametaxa,the name
of the species thathas priority(cyst or motile form)should
be followed under the InternationalCode of Botanical
Nomenclature.However,if thereremainssome doubtthat
thecyst andmotileformsarethe sametaxathenbothnames
should be retained.Harland(1982) applied a similar ap-
proach for species of Protoperidiniumand his revised
nomenclature,based on the motile form,has gained wide-
spreadacceptance.
Matsuoka1983").EnglandFinder
echigoense"
7-8 "Topotype", Nanatani Formation, loc. Minami-
Imogawa,Shitada-Mura,
NiigataPrefecture,Japan.
Age;earlyMiddleMiocene;slideNNA4-6[M30/4].
10-12 "Topotype", Nanatani Formation, loc. Minami-
Imogawa,Shitada-Mura,
NiigataPrefecture,Japan.
Age;earlyMiddleMiocene;slideNNA4-2[C34/1].
K. Matsuoka,A. McMinn, J.H. Wrenn:Holotype of Operculodiniumcentrocarpum Plate 3

8
28
28 21 - 1997
PALYNOLOGY,VOLUME 21-1997

Operculodinium
centrocarpum &Cookson)
(Deflandre
= Hystrichosphaeridium &Cookson1955
centrocarpum-Deflandre
= Operculodinium
echigoense Matsuoka1983 0
A A
A Middle Miocene A
to Pliocene

20- o
Pleistocene alX
* Holocene A A
*~ Modern A* A A
0
A e

E
3 _
s 0
G.) AA
ow
0-
0
0
cL
10- o 6o *

* e- 0

~?',..~~"-- I
Cystof Protoceratium
reticulatum
(Clapaede &Lachmann)
= Operculodinium
centrocarpum(Deflandre&Cookson)
sensuWall1967

?n ,-- -- - I - .
u )
50 Cyst diameter (pm) 100

Text-Figure1. Distributionof process length andcyst diameterof Operculodiniumcentrocarpumincludingformer0. echigoense

and the cysts of Protoceratium reticulatumcollected from Japanese Neogene, Quaterary, and modem sediments. Middle
Miocene to Pliocene specimens are attributedto 0. centrocarpum(Deflandre& Cookson) Wall, and Pleistocene, Holocene and
modem specimensare the cysts of Protoceratiumreticulatum(Clapar6de& Lachmann)Butschli.

PLATE 4

The cyst ofProtoceratiumreticulatum(Claparede& Lachmann)Biitschlicollected frommodem sedimentsof the AtlanticOcean.England

FinderReferences are given in brackets[ ].

1-4 Loc. Modemcontinentalslope, easternU.S.A.; slide U-
E6(3) [029/-]; centralbody diameter=20gm, overall
diameter=34gum.
5-7 Loc. Modem continentalslope, easternU.S.A.; slideU-
E6 [D36/3]; central body diameter=24 gm, overall
diameter=31gm.
8-9 Loc. Modem surface sediment in Castle Harbor,Ber-
muda;slide 5, [037/4 ]; centralbody diameter22 jlm,
overall diameter40 gm.
10-13 Loc. Modem continentalslope off of U.S.A.; slide U-
E6 [025/-]; centralbody diameter23 gm, overalldiam-
eter 32 glm.
K. Matsuoka,A. McMinn, J.H. Wrenn:Holotype of Operculodiniumcentrocarpum Plate 4
30 PALYNOLOGY,
PALYNOLOGY,VOLUME21 - 1997
VOLUME21-1997

CONCLUSIONS ACKNOWLEDGMENTS

1. "0. echigoense"Matsuoka1983 is thejuniorsynonym We aregreatlyindebtedto David Wall (Houston,Texas)

of 0. centrocarpum(Deflandre& Cookson)Wall 1967
(= H. centrocarpumDeflandre& Cookson 1955).
2. 0. centrocarpum(Deflandre& Cookson)Wall 1967 (=
H. centrocarpumDeflandre & Cookson 1955) should
be applied only to large, morphologically distinct
dinocystscomparableto thoseoriginallydescribedfrom
the Miocene of Australia.
3. The distinctive Quaternarydinocysts discussed herein
as 0. centrocarpum(Deflandre& Cookson)sensuWall
1967 were produced by Protoceratium reticulatum
(Claparede& Lachmann)Butschli 1885 and shouldbe
referredto as the cysts of that motile species, not 0.
centrocarpum(Deflandre& Cookson) Wall 1967.
for the loan of his slide of Operculodinium'centrocarpum'
and also to MartinJ. Head (the Universityof Toronto)for
his importantandcriticaldiscussion andcommentswhich
areveryusefulforimprovementtheearliermanuscript.We
alsothankto Laurentde Verteuil(theUniversityof Toronto)
for useful commentson the taxonomyfor Operculodinium
centrocarpumandrelatedspecies.Theseniorauthor(K.M.)
acknowledges The JapanSociety for Science Promotion
for financialsupportto visit to the Universityof Tasmania.

TABLE1. Morphological charactersof Hystrichosphaeridium

centrocarpum, echigoense,and the cyst of
Operculodinium
reticulatum.
Protoceratium

Operculodinium Operculodinium Restingcyst of Restingcyst of

centrocarpum echigoense Protoceratium Protoseratium
(Deflandre& Matsuoka1983 reticulatumin modern reticulatum
Cookson1955) sediments Incubated
(Thisstudy) specimens(Wall&
Dale1968)
Cyst diameter 54-80,m 12-21gm 32.5-42.5gm 33-37um
Surfaceornament microreticulate microreticulate microgranular (microgranular)
Lengthof 13-18 im 12-21gm 8-16 im 8-10I.m
processes
Proximalbase of conicalwithrough reticulate? conicalwithminute not clear
processes reticulation (fenestrate) striations
(fenestrate)

Shape of process slenderlytapering taperingto cylindrical cylindrical

Distalend of fibrous,patulate fibrous,rarely minutelyhookedor minutelyhookedor
processes rarelyrecurved recurved,capitateor platformedshape platformedshape
rarelybifid

Archeopyle not clear,probably (3P)

precingular (3P)
precingular (3P)
precingular
precingular (trapezoidal) (trapezoidal) (trapezoidal)
not clear,probably notclear,probably probablyreflected
Paracingulum reflectedby rowof notclear reflectedby rowof by rowof
intratabularprocesses intratabular intratabular
processes processes
clusterof relatively clusterof relatively
Parasulcus shortand more shortandmore not indicated not indicated
slender processes slender processes
K. Matsuoka,A. McMinn, J.H. Wrenn:Holotype of Operculodiniumcentrocarpum
I
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1955 Fossil microplanktonfromAustralianlate Mesozoic
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1966 Tertiary stratigraphyof the Mornington district,
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HARLAND, R.
1982 A review of Recent and Quaternaryorganic-walled
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1983 Distributionmaps of Recent dinoflagellatecysts in
bottomsedimentsfromtheNorthAtlanticOceanand
adjacentseas. Palaeontology, 26: 321-387.
HEAD, M.J., and WRENN J.H. (eds.)
1992 A forumonNeogene-Quaterarydinoflagellatecysts:
The editedtranscriptof a roundtablediscussionheld
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1993 Fossil Dinoflagellates:Indexto GeneraandSpecies
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Palynologists, ContributionSeries, 28; 856 pgs.
MATSUOKA,K.
1976 Recent thecate and fossilized dinoflagellates off
Hachinohecoast, northeastJapan.Publicationof the
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1983 Late Cenozoic dinoflagellatesand acritarchsin the
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Oceanographie.GeomarReport,7: 104 pgs.
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1990 Recent dinoflagellate cyst distributionin eastern
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of Deep
1987 Palynologyanddinoflagellatebiostratigraphy
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WALL,D., andDALE,B.
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200.

APPENDIX 1.

Otherspecies observed in the same slide containingtopotypesof Operculodiniumcentrocarpum(Deflandre& Cookson)

Wall.

Lingulodiniummachaerophorum(Deflandre& Cookson) C
(forms with both normaland shortprocess are present)
Rottonestiaborusica Deflandre& Cookson C
Spiniferitesbulloideus(Deflandre& Cookson) A
(only specimens lacking membraneousantapicalprecesses)
OperculodiniumlongispinigerumMatsuoka R
OperculodiniumjanducheneiHead et al. C
(smallercyst body, shorter,dense processes in
comparisonwith 0. longispinigerum)
ImpagidiniumjaponicumMatsuoka VR
Spiniferitesramosus(Ehrenberg)Mantell A
Systematophoraplacacantha Deflandre& Cookson VR
Reticulatosphaeraactinocoronata(Benedek) R
Dapsilidiniumpastiersii (Davey & Williams) VR
HetelaulacacystacampanulaDrugg & Loeblich C
MelitasphaeridiumchoanophorumHarland& Sharp R
NematosphaeropsislemniscataBujak VR
Spiniferitesmirabilis (Rossignol) VR
Spiniferiteshyperacanthus(Deflandre& Cookson) VR
CordosphaeridiumminutumMatsuoka& Bujak VR
Spiniferitesspp. indet. A

A: abundant,C: common, R: rare,VR: very rareon the slide.

The co-occurrenceof such dinoflagellatecysts s Dapsilidiniunpastielsii, Heteraulacacystacampanula,Impagidinium

japonicum and Reticulatosphaeraactinocoronatain this dinoflagellatecyst assemblagesuggests thatthe geologic age for
the GellibrandClays is late Early to early Middle Miocene.
K. Matsuoka,A. McMinn, J.H. Wrenn:Holotype of Operculodiniumcentrocarpum 33
I

APPENDIX2.

References in Appendix2 are listed in Lentin and Williams (1993).

Synonymy for the cyst of Protoceratiumreticulatum(Clapar6de& Lachmann)Butschli;referenceschecked hereinare

those only dealing with late Pleistocene and Recent sedimentsand with microphotographs.

1945 Cyst of Protoceratiumreticulatum(Claparede& Lachmann)Bitschli; Braarud,p. 15-17, pi. IV, figs. d, e, Text-fig. 6
1954 Cyst of Protoceratiumreticulatum(Clapar6de& Lachmann);Erdtmann,fig. iB.
1964 Cyst of Protoceratium reticulatum(Clapar6de& Lachmann);Evitt & Davidson, p. 5, pl. 1, fig. 12
1966 Cyst of Protoceratium reticulatum(Clapar6de& Lachmann);Wall, pl. 2, fig. 6
1966 Baltisphaeridiumcentrocarpum(Deflandre & Cookson); Wall & Dale, fig. 4.
1967 Operculodiniumcentrocarpum(Deflandre& Cookson) Wall, p. 11, pl. 16, figs. 1, 2, 5.
1967 Operculodiniumcentrocarpum(Deflandre & Cookson); Wall & Dale, p. 352, pl. 1, fig. M.
1968a Operculodiniumcentrocarpum(Deflandre& Cookson); Wall & Dale, pl. I, fig. 12
1968b Resting spore of Protoceratiumreticulatum(Clapar6de& Lachmann);Wall & Dale, p. 272, pl. 1, fig. 20
1968 Operculodiniumcentrocarpum(Deflandre & Cookson); Harland,p. 546-548, figs. 11, 25, 26.
1969 Hemicystodiniumzoharyi (Rossignol) Wall; Harland& Downie, pl. 7, fig. 10.
1969 Hemicystodiniumzoharyi (Rossignol); Downie & Singh, figs. 1, 2.
1971 Cyst of Protoceratium reticulatum(Clapar6de& Lachmann);Wall, pl. 2, fig. 6.
1973 Cyst of Protoceratiumreticulatum(Clapar6de& Lachmann);Harland,p. 236-239, p. II, figs. 4,7-16, pi. III, figs. 1-6, pi.
IV, figs. 1, 2.
1974 Operculodiniumcentrocarpum(Deflandre & Cookson); Reid, p. 594-595, pl. 2, figs. 10, 11.
1974 Operculodiniumcentrocarpum(Deflandre & Cookson); Jux, pl. 1, fig. 4, pl. 4, figs. 1-3, pl. 5, figs. 1-4.
1975 Operculodiniumcentrocarpum(Deflandre & Cookson); Davey & Rogers, pl. 5, figs. 1-4.
1976a Operculodiniumcentrocarpum(Deflandre & Cookson); Matsuoka,p. 364-365, pl. II, fig. 10.
1976b Operculodiniumcentrocarpum(Deflandre & Cookson); Matsuoka,p. 112-113, pl. 27, figs. 3-8.
1976 Operculodiniumcentrocarpum(Deflandre & Cookson); Dale, pl. 1, fig. 3
1977 Operculodiniumcentrocarpum(Deflandre & Cookson); Harland,p. 96-97, pl. 1, fig. 19, pl. 4, figs. 9, 10.
1979 Operculodiniumcentrocarpum (Deflandre & Cookson); Morzadec-Kerfour, p. 226, pi. 33, figs. 5,6,11, pi. 35, figs.
10,12,14,15.
1983 Operculodiniumcentrocarpum(Deflandre & Cookson); Harland,p. 330, pl. 43, figs. 9-10.
1984 Operculodiniumcentrocarpum(Deflandre& Cookson);Bradford& Wall, p. 32-33, PI. 1, figs. 10-11.
1985 Operculodiniumcentrocarpum(Deflandre & Cookson); Matsuoka,p. 41, pl. 7, figs. 1-6.
1986 Operculodiniumcentrocarpum(Deflandre & Cookson); Mudie, p. 804, pl. 2, fig. 4.
1986 Operculodiniumcentrocarpum(Deflandre & Cookson); Wrenn& Kokinos, pl. 8, figs. 5,8.
1987 Operculodiniumcentrocarpum(Deflandre & Cookson); Matsuoka,p. 48, pl. 4, figs. 1-12.
1988 Operculodiniumcentrocarpum(Deflandre & Cookson); Harland,fig. 3b.
1990 Operculodiniumcentrocarpum(Deflandre & Cookson); He Cheng-quan,p. 145, p. 59, fig. 3.
1990 Gonyaulaxgrindleyi Reinecke; Bolch & Hallegraeff, p. 175, figs. 2a-d.
1990 Operculodiniumcentrocarpum(Deflandre & Cookson); Duane & Harland,pl. 1, fig. 4.
1990 Gonyaulaxgrindleyi Reinecke; Bolch & Hallegraeff, p. 175, figs. 2a-d.
1991 Operculodiniumcentrocarpum(Deflandre & Cookson); McMinn, pl. 2, figs. 6,9.
1991 Operculodiniumcentrocarpum(Deflandre & Cookson); McMinn, p. 5,
1991 Operculodiniumcentrocarpum(Deflandre & Cookson); He & Sun, p. 282-283, pl. 5, figs. 6-11.
1992 Operculodiniumcentrocarpum(Deflandre & Cookson) Wall; Matsuoka,pl. 1, figs. 7-9.
1992 Operculodiniumcentrocarpum(Deflandre & Cookson) Wall; Mudie, pl. 3, figs 11-13.
1992 Spinose cysts of Protoceratium reticulatum(Clapar6de& Lachmann);Hallegraeff & Bolch, figs. 22, 23.
1992 Operculodiniumcentrocarpum(Deflandre & Cookson); Sun & Song, p. 47-48, pl. 1, figs. 1, 4-6.
1993 Protoceratium reticulatum(Clapar6de& Lachmann)Biitschli; Nehring, fig. 5I.
1994 Protoceratium reticulatum(Claparede& Lachmann)Butschli; Nehring, p. 211, fig. 4-1N,O; 4-3B; 7-iH; 7-3C,F.
1994 Operculodiniumcentrocarpum(Deflandre& Cookson) Wall; Matsuoka,pl. 1, figs. 5-6.
1994 Operculodiniumcentrocarpum(Deflandre & Cookson) Wall; McMinn & Sun, p. 46, pl. 3, figs. 8, 12.

var.
minrus
Operculodiniumcentrocarpum Morzadec-Kerfourn,p. 226, p. 33, fig. 7 is probablynot Operculodinium,
because no precingulararcheopylebeing.

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