Catalogue and systematics of
Pliensbachian, Toarcian and Aalenian
radiolarian genera and species
Špela Goričan
Elizabeth S. Carter
Paulian Dumitrică
Patricia A. Whalen
Rie S. Hori
Patrick De Wever
Luis O'Dogherty
Atsushi Matsuoka
Jean Guex
Špela Goričan, Elizabeth S. Carter, Paulian Dumitrică, Patricia A. Whalen, Rie S. Hori,
Patrick De Wever, Luis O’Dogherty, Atsushi Matsuoka & Jean Guex
Catalogue and systematics of Pliensbachian, Toarcian and Aalenian radiolarian genera and species
© 2006, Založba ZRC / ZRC Publishing
Izdajatelj / Issued by
Paleontološki inštitut Ivana Rakovca ZRC SAZU
Za izdajatelja / Represented by
Adrijan Košir
Založnik / Published by
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Oto Luthar
Glavni urednik / Editor-in-Chief
Vojislav Likar
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563.14”6152”
CATALOGUE and systematics of Pliensbachian, Toarcian and Aalenian radiolarian genera and species /
Špela Goričan ... [et al.]. - Ljubljana : Založba ZRC, ZRC SAZU = ZRC Publishing, 2006
ISBN-10 961-6568-65-5
ISBN-13 978-961-6568-65-4
1. Goričan, Špela
229850112
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Catalogue and systematics of
Pliensbachian, Toarcian and Aalenian
radiolarian genera and species
Špela Goričan
Paleontološki inštitut Ivana Rakovca ZRC SAZU, Ljubljana
Elizabeth S. Carter
Department of Geology, Portland State University
Paulian Dumitrică
Institut de Géologie et Paléontologie, Université de Lausanne
Patricia A. Whalen
Department of Geosciences, University of Arkansas, Fayetteville
Rie S. Hori
Department of Earth Sciences, Faculty of Science, Ehime University, Matsuyama
Patrick De Wever
Département Histoire de la Terre, Muséum National d’Histoire Naturelle, Paris
Luis O’Dogherty
Departamento de Geología, Facultad de Ciencias del Mar, Universidad de Cádiz
Atsushi Matsuoka
Department of Geology, Faculty of Science, Niigata University
Jean Guex
Institut de Géologie et Paléontologie, Université de Lausanne
Ljubljana, 2006
On the cover: Citriduma De Wever
The spatial vessel on the front cover reveals why radiolarians allow us to travel to pleasurable places which are like dreams
even though they are firmly anchored in the reality of the radiolarian world, where Paulian Dumitrica reigns as a master
(Citriduma is an anagram of his name).
Authors’ addresses
Špela Goričan
Paleontološki inštitut Ivana Rakovca
ZRC SAZU
Novi trg 2
SI-1000 Ljubljana
Slovenia
spela@zrc-sazu.si
Patricia A. Whalen
Department of Geosciences
University of Arkansas
118 Ozark Hall
Fayetteville
Arkansas 72701, USA
micropaw14@ipa.net
Elizabeth S. Carter
Department of Geology
Portland State University
Portland, Oregon 97207-0751, USA
mailing address: 17375 Jordan Road,
Sisters, Oregon 97759, USA
cartermicro@earthlink.net
Rie S. Hori
Department of Earth Sciences
Faculty of Science
Ehime University
Matsuyama 790-8577
Japan
shori@sci.ehime-u.ac.jp
Paulian Dumitrică
Institut de Géologie et Paléontologie
Université de Lausanne, BFSH 2
CH-1015 Lausannne
mailing address: Dennigkofenweg 33,
CH-3073 Guemligen, Switzerland
Paulian.Dumitrica@unil.ch
Patrick De Wever
Département Histoire de la Terre
Muséum National d’Histoire Naturelle
43 rue Buffon
F-75005 Paris
France
pdewever@mnhn.fr
Luis O’Dogherty
Departamento de Geología
Facultad de Ciencias del Mar
Universidad de Cádiz
Campus Río S. Pedro
11510 Puerto Real, Cádiz
Spain
luis.odogherty@uca.es
Atsushi Matsuoka
Department of Geology
Faculty of Science
Niigata University
Niigata 950-2181
Japan
matsuoka@geo.sc.niigata-u.ac.jp
Jean Guex
Institut de Géologie et Paléontologie
Université de Lausanne, BFSH 2
CH-1015 Lausannne
Switzerland
Jean.Guex@unil.ch
CONTENTS
Abstract 6
Povzetek 6
ACKNOWLEDGEMENTS 7
1. INTRODUCTION 9
1.1. Objectives of this publication 9
1.2. Organization of chapters 10
2. SYSTEMATICS 13
2.1. Concepts of systematics and limitations 13
2.2. Notes for user 14
2.3. Systematic description of genera and species 15
3. DESCRIPTION OF LOCALITIES 415
4. LISTING OF SPECIES 427
4.1. Alphabetical listing by genus 427
4.2. Alphabetical listing by species 431
4.3. Listing in ascending order of species/subspecies codes 436
REFERENCES 439
5
Abstract
This volume comprises a catalogue of 90 genera, 274 species and 13 subspecies of Pliensbachian, Toarcian and Aalenian
Radiolaria. Two genera, 37 species and 3 subspecies are new formal descriptions, 24 species are described in open
nomenclature. Each taxon is presented with a complete and up-to-date synonymy, original description and original
remarks (translated into English where necessary), subsequent emendations, remarks by the authors of this catalogue,
and etymology. Descriptions of species/subspecies further contain the original measurements, type locality, and data on
geographic distribution. Plates illustrate the holotype and one or several specimens from our material, from different
paleogeographic realms where possible. The material was collected from 30 measured sections in the Circum-Pacific belt
(Baja California Peninsula, Oregon, British Columbia, Japan) and the Tethyan realm (Oman, Turkey, Slovenia, Austria).
Abbreviated locality information and a list of all treated taxa are given in the last two chapters.
Povzetek
Knjiga je katalog 90 rodov, 274 vrst in 13 podvrst pliensbachijskih, toarcijskih in aalenijskih radiolarijev. Dva rodova,
37 vrst in 3 podvrste so formalno opisani novi taksoni, 24 vrst je opisanih v odprti nomenklaturi. Vsak takson je
predstavljen z vso dosedanjo sinonimiko, originalnim opisom in originalnimi opombami (v prevodu, če originalni jezik
ni angleščina), poznejšimi revizijami, pripombami avtorjev tega kataloga in etimologijo. Opisi vrst in podvrst vsebujejo še
originalne meritve, ime tipične lokalitete in podatke o geografski razširjenosti. Vsaka vrsta ali podvrsta je predstavljena s
samostojno tablo, na kateri so slike holotipa in več primerkov iz naših vzorcev. Kjer je mogoče, so ilustrirani primerki z
različnih paleogeografskih območij. Vzorci so bili pobrani na 30 profilih, posnetih v cirkumpacifiškem pasu (Kalifornijski
polotok, Oregon, Britanska Kolumbija, Japonska) in v območju Tetide (Oman, Turčija, Slovenija, Avstrija). Dodatek na
koncu knjige vsebuje kratek opis vzorčevanih profilov in seznam vseh obravnavanih taksonov.
6
ACKNOWLEDGEMENTS
The greater part of funding for this publication was provided
by the Ministry of Science and Education and Slovenian
Research Agency, Republic of Slovenia, who financed the
salary of Š. Goričan through research programmes P0-0521
and P1-0008, partially granted exchange visits of Š. Goričan
and E.S. Carter through bilateral project BI-US/04-05/46,
and covered printing expenses for the monograph. Muséum
National d’Histoire Naturelle, France, financed severalmonth fellowships to L. O’Dogherty and Š. Goričan, thus
enabling joint work with P. De Wever in Paris.
Š. Goričan further thanks French colleagues Jean
Marcoux, Cécile Robin, François Guillocheau and François
Béchennec, who invited her to participate in CNRS projects
in Oman, where the most complete Tethyan Pliensbachian
to Aalenian radiolarian-bearing successions are exposed.
She is also grateful to her students Petra Meglič, Nevenka
Šorli and Mojca Zega, who processed several hundred
radiolarian samples, some of which were used for this
publication.
E.S. Carter thanks the Geological Survey of Canada
(GSC,Vancouver) for providing field, sample processing and
SEM support, and for providing earlier collections made
by B.E.B. Cameron. She is especially grateful to Howard
Tipper (‘Tip’) (GSC, deceased 2005) whose passionate
interest in the paleontology of the Pliensbachian provided
encouragement and support for this work (contract
no. 23254-00532/001/XSB) together with ammonite
identifications and zonal assignments. She also thanks Paul
Smith (University of British Columbia) for continuing to
provide ammonite support.
P. Dumitrica thanks Kuei-Yu Yeh, Emile Pessagno and
Isamu Hattori for providing residues with extraordinarily
well-preserved radiolarians from Oregon and Japan. His
field trips in Oman for collecting samples and studies
of the radiolarian fauna from the Hamrat Duru Basin
(processing of samples and illustration of specimens with
SEM) were supported by the Swiss National Foundation
(project No. 2000 – 050681) and carried out at the Institute
of Geological Sciences at the University of Bern. He would
like to especially thank Ingo Blechschmidt, with whom he
collaborated during all field trips and the time of radiolarian
study.
Illustrations of holotypes and other specimens in our
previous works have been reproduced from original
publications. These are indicated in the plate captions by
reference to the author(s), year, plate and figure numbers
of the original publication. We are thankful for permissions
granted by the following journals/institutions:
Bulletins of American Paleontology, Paleontological
Research Institution, Ithaca, NY, USA;
Eclogae geologicae Helvetiae, Birkhäuser Verlag, Switzerland;
Geologisch-Paläontologische Mitteilungen Innsbruck,
University of Innsbruck, Austria;
Journal of Geosciences, Osaka City University, Japan;
Journal of Paleontology, University of Iowa, USA;
Marine Micropaleontology, Elsevier, The Netherlands;
Mémoires de Géologie (Lausanne), Institut de Géologie
et Paléontologie, Université de Lausanne, Switzerland;
Micropaleontology Press, New York, USA;
Mineral Research and Exploration Bulletin, Ankara,
Turkey;
Minister of Public Works and Government Services
Canada and Natural Resources Canada, Geological
Survey of Canada;
NRC (National Research Council) Research Press,
Canada;
National Museum of Natural Science, Taiwan;
News of Osaka Micropaleontologists (Dr. Katsuo Sashida,
editor of Special Volume 13), Japan;
Palaeontographica, E. Schweizerbart’sche Verlagsbuchhandlung (Nägele u. Obermiller), Science Publishers,
Stuttgart, Germany;
Revue de Micropaléontologie, Paris, France;
The Palaeontological Society of Japan.
We especially wish to thank Jean Pierre Caulet for
allowing us to use the RadWorld relational database, a
“work in progress” prepared by himself, Catherine Nigrini
and Annika Sanfilippo. He constantly provided us with
the latest version of this database containing original and
subsequent descriptions of numerous radiolarian genera
and their type species, thus sparing us much re-typing for
the present publication.
We are indebted to all Slovenian colleagues who
participated in the final preparation of this catalogue.
Dragica Turnšek and Simon Pirc read the manuscript and
gave valuable suggestions in the final stages of work. Tina
Zajc helped in preparation of plates, Robert Križmančič
designed the cover, and Adrijan Košir prepared the final
graphic design and page setting of the entire book.
7
8
1. INTRODUCTION
1.1. Objectives of this publication
The main goal of this catalogue is to present a set of
uniformly and precisely defined radiolarian species that
will serve as a coherent taxonomic base to establish a
global radiolarian zonation for three Jurassic stages:
the Pliensbachian, Toarcian, and Aalenian. Earliest
Jurassic (Hettangian and Sinemurian) and Middle to
Late Jurassic radiolarians have been studied extensively
but Pliensbachian to Toarcian faunas are less well known
taxonomically, and especially biochronologically. Aalenian
radiolarians are included in order to connect the top of the
studied interval with the base of the global low-latitude
radiolarian zonation of Baumgartner et al. (1995b).
Pliensbachian, Toarcian and early Aalenian radiolarians
have hitherto been systematically studied mostly from the
Circum-Pacific belt: western North America (Pessagno &
Whalen, 1982; Yeh, 1987a, b; Carter et al., 1988; Cordey,
1998; Whalen & Carter, 2002), Japan (Hori, 1988, 1990,
1997; Sashida, 1988; Hori & Otsuka, 1989; Matsuoka,
1991, 2004; Yao, 1997), and the Philippines (Yeh & Cheng,
1996, 1998). Only a few localities are described from the
Tethyan realm s.s.: Turkey (Pessagno & Poisson, 1981; De
Wever, 1981b, c, 1982a, b), Oman (De Wever et al., 1990),
and Slovenia (Goričan et al., 2003). It should also be noted
that most of the published taxonomic work is based on a
few excellently preserved but isolated samples (e.g. Yeh,
1987a, b; Pessagno & Poisson, 1981; De Wever, 1981b, c,
1982a, b) whereas few continuous sections suitable for
biochronological studies have been analyzed thus far. The
existing radiolarian zonations for the Pliensbachian to
Aalenian time interval (Pessagno et al., 1987b; Carter et
al., 1988; Hori, 1990) are local and mainly characterized
by a low-resolution potential (Fig. 1.1). The only highresolution range chart has been constructed for Queen
Charlotte Islands (Carter et al., 1988) but contains two
rather long discontinuities in the early Pliensbachian and
early Toarcian.
The construction of a global radiolarian zonation
for the Pliensbachian to Aalenian interval is essential
for future zonation of the entire Jurassic. Because of this
recognized need and because some of us were currently
studying rich assemblages of this age, we initiated a joint
international project under the framework of INTERRAD
(International Association of Radiolarian Paleontologists).
The Pliensbachian to Aalenian Working Group was formed
in 2000 during the 9th INTERRAD Meeting in Blairsden,
California. During two one-week meetings, held in 2001
and 2002 at the Ivan Rakovec Institute of Paleontology
9
Fig. 1.1. State of the art in Pliensbachian to Aalenian radiolarian biochronology. Radiolarian biozones are tied to standard
ammonite zones.
ZRC SAZU, Ljubljana, we agreed upon radiolarian taxa
and stratigraphic sections to be included in the zonation.
Moreover, a provisional range chart was calculated with
BioGraph computer program which is based on the
Unitary Association Method (UA) (Guex, 1977, 1991;
Savary & Guex, 1999). The preliminary information was
presented at two congresses - 6th International Symposium
on the Jurassic System (September 2002, Palermo) and
10th INTERRAD Meeting (September 2003, Lausanne).
Editing of systematics was compiled during September
2004 at Carters’ home in Sisters, Oregon, and concluded
during October 2005 in Ljubljana.
The present catalogue is the first publication of our collaborative research. It is a synthesis of previous knowledge on the taxonomy of Pliensbachian to Aalenian species complemented by actualized definitions and further
remarks, where necessary. In order to provide a complete
set of potentially important species, some newly described
species were added. The illustrated material comes from 30
measured sections in the Circum-Pacific belt (Baja California Sur, Oregon, British Columbia, Japan) and the Tethyan realm (Oman, Turkey, Slovenia, Austria) (Fig. 1.2).
10
The final goal of our joint project is to construct a radiolarian zonation that will span the missing interval between the well-established Hettangian to Sinemurian
(Carter et al., 1998) and Middle Jurassic to Lower Cretaceous (Baumgartner et al., 1995b) radiolarian biozones.
The range chart will be published in a separate paper.
1.2. Organization of chapters
The organization of our work and of this book was
inspired by INTERRAD Jurassic-Cretaceous Working
Group, whose publication (Baumgartner et al., 1995c) has
during ten years of application proven to be an extremely
useful reference tool, indispensable in systematic and
especially biostratigraphic studies.
The main part of this book is Chapter 2 Systematics,
arranged in alphabetical order of genera and species.
In addition to obligatory headings, such as synonymy,
description, and remarks, a paragraph with known
occurrences is added in order to increase the applicability
Fig. 1.2. World map with localities studied by the Pliensbachian to Aalenian Working Group.
1 – NBC. Northeastern British Columbia (Williston Lake)
2 – QCI. Queen Charlotte Islands (from north to south: Graham,
Maude, Moresby, Louise and Kunga islands)
3 – OR. Oregon (Izee - Paulina and Izee - John Day roads)
4 – BCS. Baja California Sur (Punta San Hipólito)
5 – SI. Slovenia: Julian Alps (Mt. Mangart)
6 – AT. Austria: Northern Calcareous Alps (Teltschengraben)
7 – TR. Turkey: Taurus Mountains, Gümüslu Allochthon
(Gümüslu village)
8 – OM. Oman, Hawasina Nappes: Hamrat Duru Group (Wadi
Mu’aydin, Jabal Safra, Al Sawad, Wadi Saal, Al Kashbah Mt.,
Sabt), Al Aridh Group (Al Aridh village, Jabal Buwaydah),
Umar Group (Humadiyin)
9 – JP. Japan: Mino Terrane (Inuyama, Nanjo, Mt. Norikuradake,
Gujo-Hachiman, Kamiaso areas), Chichibu Terrane (Kuma
area)
of the database for paleobiogeographic studies. The
number of Pliensbachian and Toarcian species included
here is far greater than that of the Aalenian representatives,
because many Aalenian species have already been included
in the catalogue of Baumgartner et al. (1995a). Only
those typical Middle Jurassic species needing revision,
completion of synonymy or additional illustrations
reappear in this book. The stratigraphic ranges of taxa are
not indicated because this portion of the project has not
been completed yet.
Chapter 3 contains basic data on all localities where
studied material was collected. When the Pliensbachian
to Aalenian Working Group was created in 2000, a few
localities integrated in this book were fully described
(Carter et al., 1988; De Wever, 1981b, c, 1982a, b) and
since that time some other papers have been published
(Whalen & Carter, 2002; Goričan et al., 2003; Matsuoka,
2004). Detailed descriptions of remaining localities and
their radiolarian inventory (e.g. Pliensbachian of Queen
Charlotte Islands, Pliensbachian to Aalenian of Oman)
are in preparation and will be published as individual
papers in the near future. Herein, only the location, a short
description of lithology and the overall stratigraphic range
of the studied successions are given. Stratigraphically
important co-occurring fossils are indicated and the
original publication is cited, if available.
Chapter 4 is an index of all taxa treated in the catalogue.
In order to allow an easy search, the list of species appears
in three different arrangements. The same list is sorted in
alphabetical order of genera (Chapter 4.1.), alphabetical
order of species (Chapter 4.2.), and ascending order of
alphanumerical species codes (Chapter 4.3.).
11
12
2. SYSTEMATICS
2.1. Concepts of systematics and
limitations
The main purpose of this catalogue is to provide precise
clues for doubtless identification of taxa that are useful for
global biochronology. The concept of species is basically
the same as that in Baumgartner et al. (1995a). Some
morphotypes with narrow delimitations were combined
into more broadly defined species. If morphological
characteristics of a taxon are narrowly defined but clearly
distinguishable in well-preserved material, we used such
a taxon as a subspecies. In this case the corresponding
species is denominated with sensu lato and has a separate
entry in the database, so that in poorly-preserved material
species identification is at least possible. On the other
hand, when a selected species contains a continuum of
variabilities, we called it species group without further
subdivison.
Generic level taxonomy was not the prime concern of
this book. Some long ago described genera (e.g. Stichocapsa
Haeckel 1881) certainly need thorough revision but
this would necessitate more detailed taxonomic and
phylogenetic studies, which were beyond the scope of this
catalogue. More modern descriptions were, nevertheless,
carefully examined and emended, if necessary. A clear
actualized definition was considered especially important
for those genera having their first or last occurrence in the
Pliensbachian to Aalenian interval and may therefore be
used for biochronology at generic level.
We recently discovered that some genera included
in this catalogue (Atalanta, Beatricea, Canutus, and
Thurstonia) are homonyms of other genera described long
ago. The resolution of this problem is not within the scope
of this publication and will be addressed in the future by
individual authors.
Suprageneric classification is deliberately ignored in
this book; for this information the reader is referred to
De Wever et al. (2001).
13
2.2. Notes for user
The included taxa are treated in alphabetical order of
genera and species. The following headings are used to
define a taxon:
Taxon code: Species and subspecies are coded. This code
has no significance in taxonomy but is required by the
BioGraph computer program for range-chart calculation.
Some species have been used previously in the calculation
of other range charts (Baumgartner et al., 1995b; Carter et
al., 1998); their codes are unchanged. The codes used by
Carter et al. (1998) and the ones newly designated in this
book are composed of three letters and two digits, while
the species codes defined by Baumgartner et al. (1995a, b)
have four digits.
Synonymy: All synonyms appear in chronological order.
Doubtful synonymy is preceded by “?”, taxa explicitly
excluded from a particular synonymy are preceded by
“not”.
Type designation: This heading designates holotype and
paratypes in the description of new species/subspecies.
Original description: This heading gives the original
description by the author of the taxon. Original descriptions
in languages other than English are translated. Because
the description is an authentic copy from the original
publication, this heading may be called Original diagnosis
or split into Original diagnosis plus Original description.
Emended definition (description): Gives subsequent emendations with the reference to the publication cited. If no
publication is cited, this means that the taxon is emended
herein.
Original remarks are the remarks by the author of the
taxon. If in the original publication only a reference to
remarks under another taxon is given, the remarks under
the latter taxon are also provided.
Further remarks are mostly remarks by the authors of this
catalogue. In a few cases, relevant previously published
comments are given and the source is cited.
Measurements are the measurements from the original
publication.
14
Etymology makes reference to the origin of the formal
name of the taxon.
Type locality states the type locality of individual species
and subspecies.
Included species/subspecies: This heading appears with
genera and species sensu lato, and gives a list of species or
subspecies treated in this catalogue. Names are preceded
by codes.
Occurrence: This heading gives a list of previously known
plus our ‘new’ localities for each species/subspecies. Only
broader regions and not individual locations are listed.
Names of lithostratigraphic units are indicated, when
available. The occurrence of the holotype always appears
as the first on the list.
Plates: Each species/subspecies has its plate that is
numbered with the code of the taxon. The plates show the
holotype (for formal taxa) and one or several illustrations,
from more than one locality if possible. Magnification is
indicated in the plate caption and scale bars are added.
The figure caption indicates for each figure, the following:
country code (e.g. QCI), sample number (e.g. 99-CNAMI-10), and photograph/specimen number (e.g. GSC
34567). The figure number of the holotype is followed by
(H), author, year, plate, and figure of original illustration.
The same indication is given for figures previously used
in other publications by the authors of this catalogue.
Permissions granted for reproduction of the illustrations
are gratefully acknowledged.
List of country codes used in the figure captions:
AT = Austria
BCS = Baja California Sur, Mexico
JP = Japan
NBC = Northeastern British Columbia, Canada
OM = Oman
OR = Oregon, USA
QCI = Queen Charlotte Islands, British Columbia, Canada
SI = Slovenia
TR = Turkey
Repository: The holotypes and paratypes of newly described species are stored in the authors’ collections. Repository numbers correspond to specimen numbers, indicated in plate captions.
2.3. Systematic description of genera and species
Genus: Acaeniotylopsis Kito & De Wever 1994
Type species: Acaeniotylopsis triacanthus Kito & De Wever 1994
Synonymy:
1994 Acaeniotylopsis n. gen. – Kito & De Wever, p. 130.
Original description: Test composed of spherical or subspherical shell with some radial spines, a medullary shell
and microsphere. Cortical shell comprises massive nodes
connected to each other by short bars. Outer medullary
shell is spherical and is connected to cortical shell by several strong triradiate radial beams and by numerous thin
secondary radial beams, which join nodes on cortical shell.
Primary beams merge into spines. Inner medullary shell
polyhedral.
Original remarks: The genus differs from Acaeniotyle
Foreman 1973 by the absence of perforated mammae on
the cortical shell, by a cortical shell composed of bars with
nodes and by the presence of the primary radial beams.
Further remarks: The genus also differs structurally from
Acaeniotyle in having a double medullary shell with a true
microsphere whereas the medullary shell of Acaeniotyle
is, by its size, closer to the macrosphere in the sense of
Hollande and Enjumet (1960). The genus differs essentially
from Acaeniotyle in having a double medullary shell, and
primary radial spines.
Etymology: Acaeniotyle + -opsis (masculine), in reference
to a similar morphology to Acaeniotyle Foreman.
Included species:
2001 Acaeniotylopsis ghostensis (Carter) 1988
4066 Acaeniotylopsis triacanthus Kito & De Wever 1994
15
Acaeniotylopsis ghostensis (Carter) 1988
Species code: 2001
Synonymy:
1988 Acaeniotyle (?) ghostensis Carter n. sp. – Carter et al., p. 33,
pl. 9, fig. 6.
1994 Acaeniotylopsis ghostensis (Carter) – Kito & De Wever,
p. 132, pl. 1, figs. 7-8.
1995a Acaeniotylopsis ghostensis (Carter) – Baumgartner et al.,
p. 56, pl. 2001, figs. 1-2.
1997 Acaeniotylopsis ghostensis (Carter) – Yao, pl. 3, fig. 102.
Original diagnosis: Test subspherical and slightly flattened
with 3 long, sturdy, tribladed spines. Surface of cortical
shell covered with strong, slightly perforate nodes.
Original description: Test subspherical, flattened in
plane of equatorial spines. Nodes on cortical shell strong,
moderately spaced with somewhat flattened distal surfaces
(tops); surfaces with fine perforations, some bearing
remnants of fine central spines. Nodes connected by
strong bars that form circular, elliptical and subtriangular
pores. Spines tribladed and long (entire ones greater than
3/4 diameter of test) carrying narrow rounded ridges
and wider grooves; complete spines are pointed. First
medullary shell has small irregular pore frames connected
to cortical shell by radial beams. Radial beams (3) are
strong, triradiate and continuous with each primary spine;
beams of lesser strength are attached to cortical shell at
base of nodes.
Original remarks: Genus queried; the form described
is doubtfully assigned to this genus because nodes are
smaller, knob-like rather than rounded, and have fewer
perforations, and all are much older.
Further remarks: By Kito & De Wever (1994): Our
specimens have longer radial spines than the specimens
of the original description. The microsphere was not
described in the original description, but external features
and internal structure of type specimens are completely
identical with our material.
Measurements (µm):
Based on 13 specimens.
Diameter of test
Length of longest spine
HT
146
121
Av.
145
108
Min.
175
145
Max.
139
82
Etymology: Named for Ghost Creek, north of the type locality.
Type locality: GSC locality C-080597. Toarcian of Phantom
Creek Formation, Graham Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Phantom Creek Formation, Queen Charlotte
Islands; Italy; Japan.
Acaeniotylopsis triacanthus Kito & De Wever 1994
Species code: 4066
Synonymy:
1989 Acaeniotyle (?) sp. 1 – Kito, p. 95, pl. 3, figs. 1-5, 8-9.
1991 Acaeniotyle sp. B. – Tonielli, p. 21, pl. 1, fig. 20.
1991 Acaeniotyle ? sp. A – Carter & Jakobs, p. 342, pl. 2, fig. 8.
1994 Acaeniotylopsis triacanthus n. sp. - Kito & De Wever,
p. 132, pl. 1, figs. 4-6, 9-11; pl. 3, figs. 5a-b, 6.
1995a Acaeniotylopsis variatus triacanthus Kito & De Wever
– Baumgartner et al., p. 58, pl. 4066, figs. 1-7.
1997 Acaeniotylopsis v. triacanthus Kito & De Wever – Yao,
pl. 3, fig. 104.
Original description: Test composed of a spherical or
subspherical cortical shell with three strong radial spines;
one outer and one inner medullary shell. Cortical shell
constituted of massive nodes connected by short bars.
Pores are small, polygonal or circular. Three radial spines
are arranged in a plane at about 120 degrees. Radial spines
possess three wide grooves (primary grooves) alternating
with three narrow grooves (secondary grooves). A short
spine on each ridge arises at the end of the radial spine and
provides a clove-like tip. Outer medullary shell is spherical
and connected to cortical shell by three triradate primary
beams and thin secondary beams. Inner medullary shell is
polygonal, composed of pentagonal pore frames.
16
Original remarks: This species differs from Acaeniotyle
diaphorogona variata Ozvoldova 1979 in the construction
of the cortical shell. The cortical shell lacks perforated
mammae. The species also differs from Acaeniotylopsis
ghostensis (Carter) in the aspect of its cortical shell and in
having branched spines.
Measurements (µm):
Based on 13 specimens.
Diameter of cortical shell
Length of spine A
Length of spine B
Length of spine C
HT
189
206
187
193
Av.
160
159
-
Min.
128
111
-
Max.
204
222
-
Etymology: From the Greek tri- (three) + acanthus
(spine).
Type locality: Sample S69, Contrada la Ferta, Sicily, Italy.
Occurrence: Italy; Japan; Phantom Creek Formation,
Queen Charlotte Islands.
Plate 2001. Acaeniotylopsis ghostensis (Carter). Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 9, fig. 6.
Fig. 2. QCI, GSC loc. C-080611, GSC 128704.
Plate 4066. Acaeniotylopsis triacanthus Kito & De Wever. Magnification x150. Fig. 1(H). Kito & De Wever 1994, pl. 1,
fig. 11. Fig. 2. Carter & Jakobs 1991, pl. 2, fig. 11. Fig. 3. Baumgartner et al. 1995a, pl. 4066, fig. 3.
17
Genus: Anaticapitula Dumitrica & Zügel 2003
Type species: Anaticapitula clauda Dumitrica & Zügel 2003
Synonymy:
2003 Anaticapitula n. gen. – Dumitrica & Zügel, p. 52.
Original description: Highly ovate dicyrtid test with bladed
apical horn and thorax prolonged into a thin-walled terminal tube. Cephalis and thorax continuous externally, without collar stricture. Initial spicule with A, V, D, two L, two
l, and a long Ax. Cephalis and thorax with a superimposed
network of strong ridges. With or without feet representing
external extensions of L and D. Feet, when present, with an
outer blade and two lateral blades.
Original remarks: By its general shape and the tubular
prolongation of the thorax this genus shows characters in
common with the genus Rhopalosyringium Campbell &
Clark, 1944. A comparable axobate was illustrated by De
Wever (1982) in the Lower Jurassic species Ovum pertusum De Wever, 1982. Jacus (?) anatiformis De Wever,
1982, described from the lower Pliensbachian of Turkey,
is also almost identical to A. pennata n. gen., n. sp. and,
although De Wever (1982) did not mention the presence
of Ax in his species, a small light-grey spot opposite to the
ventral spine, representing probably a small Ax, is visible
in a broken specimen figured by him (De Wever 1982;
pl. 11, fig. 13). Jacus (?) italicus Jud, 1994 described from
the Lower Cretaceous (Jud 1994) has a morphology similar
to J. (?) anatiformis and should also be assigned to the genus Anaticapitula n. gen.
From Napora, Anaticapitula n. gen. differs by having a
well developed Ax in the initial spicule, thorax continuous
with the velum, no crown of spines on the apical horn, and
a much larger cephalis.
Etymology: From the Latin anas: duck; and capitulus: small
head. Feminine gender.
Included species:
JAC02 Anaticapitula anatiformis (De Wever) 1982a
JAC04 Anaticapitula omanensis Dumitrica n. sp.
Anaticapitula anatiformis (De Wever) 1982a
Species code: JAC02
Synonymy:
1982 Bisphaerocephalina (?) sp. – Imoto et al., pl. 1, fig. 10.
1982a Jacus ? anatiformis n. sp. – De Wever, p. 205, pl. 11,
figs. 10-15.
1982b Jacus ? anatiformis De Wever – De Wever, p. 343, pl. 54,
figs. 1-5; pl. 58, figs. 1, 2, 6.
1984 Jacus sp. A – Murchey, pl. 2, fig. 29.
1984 Jacus sp. B – Murchey, pl. 2, fig. 28.
1987 Jacus sp. A – Hattori, pl. 11, fig. 7.
1987 Jacus sp. D – Hattori, pl. 11, fig. 8.
1989 Jacus sp. A – Hattori, pl. 5, fig. I.
1989 Jacus? sp. B – Hattori, pl. 5, fig. J.
1990 Jacus anatiformis De Wever – De Wever et al., pl. 3, fig. 10.
1998 Jacus ? anatiformis De Wever – Whalen & Carter, p. 74,
pl. 18, figs. 13, 14, 17, 18, 19, 27.
1997 Thetis sp. B – Yao, pl. 10, fig. 467.
1998 Jacus ? anatiformis De Wever – Yeh & Cheng, p. 32, pl. 6,
fig. 10.
2001 Jacus cf. anatiformis De Wever – Gawlick et al., pl. 2,
fig. 16.
2001 Jacus anatiformis De Wever – Gawlick et al., pl. 5, fig. 19.
2002 Jacus ? anatiformis De Wever – Hori & Wakita, pl. 3, fig. 7.
2002 Jacus ? anatiformis De Wever – Whalen & Carter, p. 138,
pl. 16, fig. 18.
2002 Anaticapitula anatiformis (De Wever) – Tekin, p. 191,
pl. 5, fig. 8.
2003 Jacus ? aff. anatiformis De Wever – Goričan et al., p. 296,
pl. 4, figs. 5-6.
2003 Jacus ? sp. – Goričan et al., p. 296, pl. 4, fig. 7.
2004 Anaticapitula (?) sp. – Matsuoka, fig. 144.
2004 Anaticapitula (?) anatiformis (De Wever) – Matsuoka,
fig. 145.
18
Original description: Form with two segments, a strong
apical horn, three diverging feet on thorax and a free subcylindrical velum.
Apical horn triradiate along most of length, but some
specimens with a rounded end. Hemispherical cephalis
imperforate, smooth, with small pustules or with strong
longitudinal wrinkles. A round pore, rather large, prolongs cephalic spine V. Thorax roughly tetrahedral in form
with lateral edges extending in three feet. Wall composed
of two superimposed irregular latticed networks: a delicate inner one and a coarse outer one. Thorax extends in
a subcylindrical velum with a thin wall and tiny irregular
pores.
Original remarks: This form is tentatively assigned to this
genus because of the relationship between the velum and
feet: whether the velum is free from the feet or bound to
them could be considered a generic criterion, but it seems
hasty to me in our present stage of knowledge.
This form resembles Lithomelissa amazon Foreman
(1968, p. 26) but is distinguished from it by its slender
appearance proximally. It differs also from all other species
of Lithomelissa by the lack of an axial spine (Ax).
The observation of different layers and their disposition
on the thorax and velum suggests the same growth pattern
as described by Petrushevskaya M.G. (1962) for Cenozoic
forms: development of thoracic wall, then proximal part
of velum, and lastly, the simultaneous development of a
proximal external layer on the thorax-velum suture zone
and distal part of velum.
Plate JAC02. Anaticapitula anatiformis (De Wever). Magnification x300. Fig. 1(H). De Wever 1982a, pl. 11, fig. 10.
Fig. 2. JP, IJIII12. Fig. 3. Whalen & Carter 2002, pl. 16, fig. 18. Fig. 4. QCI, GSC loc. C-080612, GSC 128815.
Fig. 5. OM, Haliw-038-R08-26. Fig. 6. OM, BR706-R05-01. Fig. 7. Goričan et al. 2003, pl. 4, fig. 5. Fig. 8. Goričan et al.
2003, pl. 4, fig. 6. Fig. 9. OM, BR706-R12-14. Fig. 10. QCI, GSC loc. C-175306, GSC 128816. Fig. 11. OM, BR706-R05-14.
19
This species differs from J. coronatus, J. clatratus and
J. isa by its free velum.
Further remarks: This species may easily be assigned to the
genus Anaticapitula (Dumitrica & Zügel, 2003) by the twolayered, thick-walled cephalis and thorax, and the tubular
distal extension of the thorax. This species is structurally
close to Anaticapitula clauda Dumitrica & Zügel.
A large variety of forms are now included in Anaticapitula
anatiformis. These forms differ in length of feet and
structure of thoracic wall, which can be two-layered and
irregular as in the holotype, or simple, composed of large
uniform polygonal pore frames.
Etymology: From Latin anas, -atis, duck and formis form.
In a form of a duck, by resemblence to the appearance of
duck in flight.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Sandilands and
Ghost Creek formations, Queen Charlotte Islands; San
Hipólito Formation, Baja California Sur; Dürrnberg Formation, Austria; Skrile Formation, Slovenia; Hocaköy Radiolarite, Turkey; Musallah Formation, Oman; Liminangcong Chert, Philippines; Japan.
Measurements (µm):
Based on 11 specimens.
Length of apical horn
Width of thorax
Total length
(including velum and apical horn)
Av.
82
84
Min.
67
75
Max.
100
100
238
200
290
Anaticapitula omanensis Dumitrica n. sp.
Species code: JAC04
Synonymy:
1989 Thetis spp. – Hattori, pl. 7, figs. E-G.
1997 Thetis sp. D – Yao, pl. 10, fig. 469.
2004 Anaticapitula (?) sp. – Matsuoka, fig. 142.
portion between top of cephalis and the level of the three
spines. Pores of thorax irregular in size, shape and arrangement; intervening bars of the outer layer usually forming
ribs in various directions.
Type designation: Holotype specimen BR1121-R08-17
from sample BR 1121, Guwayza Formation, Tawi Sadh
Member, Wadi Mu'aydin, Oman.
Remarks: Anaticapitula omanensis differs from A. anatiformis in having much shorter D and L spines; the conical
part of thorax is also shorter and inverted.
Diagnosis: Test small, subrhombic in outline with a threebladed pointed apical horn and short spines D, Lr and Ll.
Measurements (µm):
Based on 9 specimens.
Description: Test small, pyramidal in the upper half, inverted conical in the lower half. Cephalis globular, imperforate, lower part covered by ribs extending from outer layer
of thorax. Apical horn long, three-bladed, gently pointed.
Thorax much larger than cephalis, inflated in the middle
part where it bears three short laterally-downward directed
spines. Spines distally three-bladed, proximally multi-bladed with secondary blades representing prolongations of the
intervening bars of the external layer of the thoracic wall.
Collar stricture indistinct or slightly visible by a change in
outline. Beyond the level of the three spines, thorax decreases gradually in diameter and terminates irregularly.
This part of thorax approximally equal in length to the
20
Maximum length of shell
with apical horn
Length of apical horn
Diameter of thorax
at the level of the three spines
HT
Min.
Max.
170
170
209
53
50
70
87
85
88
Etymology: From its occurrence in Oman.
Type locality: Sample BR 1121, Guwayza Formation, Tawi
Sadh Member, Wadi Mu'aydin, Oman.
Occurrence: Guwayza Formation, Tawi Sadh Member,
Oman; Mino Terrane, Japan.
Plate JAC04. Anaticapitula omanensis Dumitrica n. sp. Magnification x300. Fig. 1(H). OM, BR1121-R08-17.
Fig. 2. OM, BR1121-R06-13. Fig. 3. OM, BR1121-R08-15. Fig. 4. OM, BR1121-R07-03. Fig. 5. OM, BR1122-R02-12.
Fig. 6. OM, BR1121-R08-08. Fig. 7. OM, BR1121-R10-01.
21
Genus: Archaeodictyomitra Pessagno 1976, emend. Pessagno 1977b
Type species: Archaeodictyomitra squinaboli Pessagno 1976
Synonymy:
1976 Archaeodictymitra n. gen. – Pessagno 1976, p. 49.
1977b Archaeodictyomitra Pessagno – Pessagno, p. 41.
1987b Combusta n. gen. – Yeh, p. 60.
1995a Archaeodictyomitra Pessagno – Baumgartner et al., p. 96.
Original description: Test conical, non-lobate, becoming
somewhat spindle-shaped in unbroken or mature forms;
cephalis, thorax, abdomen, and post-abdominal chambers
covered by linearly arranged continuous costae which
converge in the area of the cephalis and thorax; pores
distributed in single row between costae, entirely relict on
earlier chambers and observable only on etched or eroded
specimens.
Emended definition: By Pessagno (1977b): Definition as in
Pessagno (1976), but including forms with constrictions;
constrictions not occurring at joints.
Original remarks: Archaeodictyomitra n. gen., differs from
Dictyomitra Zittel by being non-lobate in outline and lacking well-developed strictures; and by possessing relict
pores and lacking primary pores. It differs from Diplostrobus Squinabol for the reasons cited above and by lacking an
apical horn.
Only a few of the many potential species known to be
assignable to this genus have been described from Jurassic
and Cretaceous strata. »Dictyomitra« margarita Aliev, 1961,
from the Lower Cretaceous of Russia is definitely assignable to Archaeodictyomitra.
Further remarks: Combusta Yeh is herein considered a
junior synonym of Archaeodictyomitra Pessagno because
there is no structural difference between the two genera.
Both lack an apical horn and constrictions or have only
very weak constrictions at joints; both possess longitudinal
costae with a row of pores in the intercostal depressions,
and a distal aperture.
Included species:
ADM01 Archaeodictyomitra munda (Yeh) 1987b
ADM02 Archaeodictyomitra sp. A
ADM03 Archaeodictyomitra sp. B
Archaeodictyomitra munda (Yeh) 1987b
Species code: ADM01
Synonymy:
1987b Combusta munda n. sp. – Yeh, p. 61, pl. 20, figs. 6-7, 11,
17; pl. 28, figs. 8, 25.
1987b Combusta sp. A – Yeh, p. 61, pl. 20, fig. 10.
1987b Combusta sp. B – Yeh, p. 61, pl. 28, figs. 9, 16.
2003 Parahsuum spp. – Goričan et al., p. 296, pl. 5, fig. 14 only.
2003 Archaeodictyomitra sp. sensu Kojima et al. 1991 –
Kashiwagi & Kurimoto, pl. 3, fig. 11.
2004 Archaeodictyomitra munda (Yeh) – Matsuoka, fig. 187.
Original description: Test as with genus, with seven to nine
post-abdominal chambers. Cephalis medium in size, domeshaped. Cephalis, thorax, and abdomen sparsely perforate.
Post-abdominal chambers with one row of small pore
frames between adjacent costae. Costae moderately thick,
merging apically. About twelve costae visible laterally.
Further remarks: Distally more inflated forms (Combusta
sp. A of Yeh, 1987b) and forms with slight constrictions
(Combusta sp. B of Yeh, 1987b) are regarded as intraspecific
variability of Archaeodictyomitra munda (Yeh).
22
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length of test (max.)
200
195
200
190
Width of test (max.)
94
90
94
85
Etymology: Mundus-a-um (Latin, adj.) = elegant.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Hyde and Snowshoe formations of eastcentral Oregon; Fannin Formation, Queen Charlotte
Islands; Skrile Formation, Slovenia; Japan.
Plate ADM01. Archaeodictyomitra munda (Yeh). Magnification x300. Fig. 1(H). Yeh 1987b, pl. 19, fig. 11.
Fig. 2. Matsuoka 2004, fig. 187. Fig. 3. Goričan et al. 2003, pl. 5, fig. 14.
23
Archaeodictyomitra sp. A
Species code: ADM02
Synonymy:
1989 Lupherium ? sp. C – Hattori, pl. 16, fig. D.
2004 Archaeodictyomitra sp. – Matsuoka, fig. 186.
Remarks: Test spindle-shaped with cephalis, thorax,
abdomen and an undetermined number of post abdominal
chambers that gradually increase in width as added.
Narrow, linearly arranged costae, 12-14 arranged over test;
costae converging apically. Single row of pores between
costae; pores relict on early chambers becoming more open
distally.
This is one of the oldest species of Archaeodictyomitra
(earliest Pliensbachian) and may be ancestral to A. munda
(Yeh). A. sp. A is smaller (length <150µm) than A. munda
and the final chamber/s are not constricted.
Occurrence: Ghost Creek Formation, Queen Charlotte Islands; Mino Terrane, Japan.
Archaeodictyomitra sp. B
Species code: ADM03
Remarks: Test large, narrowly conical apically becoming
almost tubular toward aperture. Coarse linearly arranged
costae (12-14) on exterior of test; costae with rounded
edges. Cephalis and thorax covered with a heavy layer of
microgranular silica.
Occurrence: Fannin Formation, Queen Charlotte Islands.
Genus: Archaeohagiastrum Baumgartner 1984
Type species: Archaeohagiastrum munitum Baumgartner 1984
Synonymy:
1984 Archaeohagiastrum n. gen – Baumgartner, p. 758
Original description: Test composed of four rays, placed at
right angles and of about equal length. The rays are formed
of a primary beam, three primary canals and six external
beams.
Original remarks: The rays of Archaeohagiastrum correspond to the medullary rays of the more evolved hagiastrins and represent the simplest possible hagiastrid
structure. It was referred to as ancestor of Hagiastrum
in Baumgartner (1980, Textfig. 7 and p. 284). Tetraporobracchia Kozur and Mostler 1979 has the same ray structure but rays are arranged along tetraedric or cubic axes.
24
Archaeotriastrum De Wever 1981b has a similar ray structure but has three rays. Because of its simple ray structure
this genus is tentatively included with the hagiastrins. It
should, together with Archaeotriastrum, be assigned to a
new subfamily ancestral to the Hagiastrinae.
Etymology: From the Greek archaeo = ancient, ancestral
form to Hagiastrum.
Included species:
3149 Archaeohagiastrum longipes Baumgartner 1995
3271 Archaeohagiastrum munitum Baumgartner 1984
HAG01 Archaeohagiastrum oregonense (Yeh) 1987b
HAG02 Archaeohagiastrum pobi Whalen & Carter 1998
Plate ADM02. Archaeodictyomitra sp. A. Magnification x300. Fig. 1. QCI, C-080612, GSC 128708. Fig. 2. QCI, GSC
loc. C-305388, GSC 128709. Fig. 3. Matsuoka 2004, fig. 186.
Plate ADM03. Archaeodictyomitra sp. B. Magnification x200. Fig. 1. QCI, GSC loc. C-140495, GSC 128710.
25
Archaeohagiastrum longipes Baumgartner 1995
Species code: 3149
Synonymy:
1982 Tetratrabs sp. - Kishida & Sugano 1982, pl. 6, fig. 11.
1987 Archaeohagiastrum sp. A – Hattori, pl. 3, figs. 3-4.
1988 Archaeohagiastrum sp. A – Hattori, pl. 5, fig. B.
1988 Tetratrabs sp. aff. T. gratiosa Baumgartner – Carter et al.,
p. 30, pl. 7, fig. 10.
1989 Archaeohagiastrum sp. – Hattori, pl. 4, fig. F.
1989 Archaeohagiastrum sp. A – Hattori, pl. 25, fig. F.
1991 Tetratrabs sp. aff. T. zealis (Ozvoldova) – Carter & Jakobs,
p. 344, pl. 2, fig. 7.
1995a Archaeohagiastrum longipes Baumgartner n. sp.
– Baumgartner et al., p. 106, pl. 3149, figs. 1-6.
1996 Archaeohagiastrum sp. A – Yeh & Cheng, p. 96, pl. 1, fig. 2;
pl. 8, figs. 6, 7, 12.
1997 Archaeohagiastrum longipes Baumgartner – Yao, pl. 7,
fig. 334.
Original description: Form with four smooth slender rays
of about equal length about at right angles, constructed as
with genus. One row of large circular pores between each
external beam. Beam cross-section hexagonal. Central area
small, smooth, with small, irregular pores or with 4-7 small
nodes. Lateral beams are continuous around the central
area. The external beams of rays are smooth or slightly
nodose. Ray tip sometimes slightly thickened, with short
three-bladed central spine.
Original remarks: This species differs from A. munitum
by distincly longer and slenderer rays and a generally less
nodose test surface. Central knobs are present but much
less developed than with A. munitum.
Measurements (µm):
Based on 7 specimens.
Length of rays AX
Length of rays BX
Length of rays CX
Length of rays DX
Width of rays
Width of central area
HT
208
198
195
41
70
av.
210
45
75
min.
192
33
65
max.
218
47
82
Etymology: Longipes, latin for "long-footed" named for its
long rays compared to the type species of this genus.
Type locality: Sample OR 554, Snowshoe Formation, EastCentral Oregon.
Occurrence: Snowshoe Formation, east-central Oregon;
Phantom Creek Formation, Queen Charlotte Islands; Italy;
Dürrnberg Formation, Austria; Tawi Sadh Member of the
Guwayza Formation, Oman; Liminangcong Chert, Philippines; Japan.
Archaeohagiastrum munitum Baumgartner 1984
Species code: 3271
Synonymy:
1982 Crucella sp. A – Sashida et al., pl. 1, fig. 9.
1982 Tetratrabs sp. B – Wakita, pl. 5, fig. 4.
1984 Archaeohagiastrum munitum n. sp. – Baumgartner, p. 759,
pl. 2, figs. 9-13.
1985 Archaeohagiastrum munitum Baumgartner – Nagai, pl. 2,
figs. 5-5a.
1985 Archaeohagiastrum munitum Baumgartner – Yamamoto et
al., p. 34, pl. 3, figs. 7a-b.
1988 Tetraditryma sp. B – Carter et al., p. 31, pl. 16, fig. 8.
1990 Archaeohagiastrum munitum Baumgartner – Kito et al.
1990, pl. 1, fig. 6.
1994 Archaeohagiastrum munitum Baumgartner – Goričan,
p. 62, pl. 5, fig. 14.
1995a Archaeohagiastrum munitum Baumgartner –
Baumgartner et al., p. 108, pl. 3271, figs. 1-6.
1997 Archaeohagiastrum munitum Baumgartner – Yao, pl. 7,
fig. 335.
Original description: Small form with four smooth to nodose rays of about equal length constructed as with genus.
Central area small, occupied by four to five broad, highly
raised, connected nodes, which alternate with four pores
placed at the proximal termination of the median beams.
The fifth node is central or slightly excentric and fused to
one of the corner nodes. A nearly centrally placed pore often occurs. Lateral beams are continuous around the central area. The external beams of rays are slightly to strongly
26
nodose, nodes increase in size towards central area and are
sometimes connected by a blade like ridge. Ray tip blunt or
with short central spine of round cross section.
Original remarks: A. munitum differs from other yet undescribed species of this genus by being distinctly smaller and
by having a strongly nodose test.
Measurements (µm):
Based on 7 specimens.
Length of rays AX:
Length of rays BX:
Length of rays CX:
Length of rays DX:
Width of rays:
Maximum length of spines:
Width of central nodose area:
min.
114
120
108
111
51
66
65
max.
95
42
48
60
av.
87
35
28
47
HT
120
51
66
76
Etymology: Munitum: fortified, protected (Latin), referring
to the nodose surface of test and central area.
Type locality: Blake Bahama Basin, West Atlantic (DSDP
Leg 71, Site 534).
Occurrence: Worldwide.
Plate 3149. Archaeohagiastrum longipes Baumgartner. Magnification x150. Fig. 1(H). Baumgartner et al. 1995a,
pl. 3149, fig. 4. Fig. 2. OM, BR871-R03-07. Fig. 3. AT, BMW21-31. Fig. 4. Carter et al. 1988, pl. 7, fig. 10.
Fig. 5. Carter & Jakobs 1991, pl. 2, fig. 7.
Plate 3271. Archaeohagiastrum munitum Baumgartner. Magnification x200. Fig. 1(H). Baumgartner 1984, pl. 2, fig. 9.
Fig. 2. Carter et al. 1988, pl. 16, fig. 8. Fig. 3. JP, MNA-10, MA11555. Fig. 4. JP, NK9-62.
27
Archaeohagiastrum oregonense (Yeh) 1987b
Species code: HAG01
Synonymy:
1987 Tetraditryma sp. A – Hattori, pl. 3, fig. 1.
1987b Higumastra oregonensis n. sp. – Yeh, p. 26, pl. 8 figs. 7, 15,
20; pl. 23, fig. 16; pl. 29, figs. 11, 18.
1990 Higumastra oregonensis Yeh – Nagai, pl. 5, fig. 2.
1998 Archaeohagiastrum sp. aff. A. pobi n. sp. – Whalen & Carter,
p. 45, pl. 10, figs. 1, 6, 10, 13, 17.
Original description: Rays relatively uniform in width,
elongate, with short massive primary spines. Primary spines
triradiate proximally, circular in cross section distally.
Central area small, square in outline. Rays nearly equal in
length, comprised of three rows of tetragonal pore frames.
Central area of cortical shell consisting of pentagonal and
hexagonal pore frames quadrilaterally arranged. Pore
frames larger on rays; all pore frames without prominent
nodes at vertices. Test with or without patagium.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length
of ray
92
95
101
78
Width
of ray
31
28
31
24
Width
of central area
61
58
61
54
Length
of spine
30
42
45
30
Type locality: Locality 600A, Hyde Formation along IzeePaulina road, east-central Oregon.
Occurrence: Hyde and Snowshoe formations, Oregon;
Sandilands Formation, Queen Charlotte Islands; Japan.
Archaeohagiastrum pobi Whalen & Carter 1998
Species code: HAG02
Synonymy:
1998 Archaeohagiastrum pobi n. sp. – Whalen & Carter, p. 44,
pl. 10, figs. 3, 4, 5, 9.
1998 Pseudocrucella sp. A – Cordey, p. 70, pl. 20, fig. 1.
Original description: Test composed of four short rays at
right angles, terminating in very long, massive, triradiate
spines. Each ray comprised of an internal primary beam,
three primary canals and six external beams. External spine
ridges and grooves part of integral geometry of four-rayed
test. Longitudinal beams developed on edges of each spine
ridge (two per spine, totaling six beams); transverse bars
connecting these beams creating linear rows of fairly regular pore frames, most tetragonal (four to fine horizontal
rows of pores per ray). Large raised elliptical to subrectangular nodes aligned perpendicular to beams; nodes located
at vertices of external beams and transverse bars. Three
hagiastrid canals formed by transverse bars spanning the
deeply grooved spine ridges. Central area large, composed
mostly of triangular pore frames with large subrounded
nodes at vertices. Spines very long with broad, rounded
ridges and deep rounded grooves.
Original remarks: Baumgartner (1980, p. 284) wrote, "An
early Sinemurian sample (QC 549) contains 3 types of hagiastrids. One of them, a possible ancestor of Hagiastrum, has
a central area similar to the Emiluvia-like forms" (=Udalia
in this paper) "and possesses 3 primary canals and 6 external beams. It seems possible that this form is the first
hagiastrid and has evolved from Emiluvia-like forms by developing transverse bars between raised ridges of primary
spines and thus enclosing primary grooves to form primary
28
canals". We believe the two species of Archaeohagiastrum
discussed and illustrated here conform to this statement.
Archaeohagiastrum pobi n. sp. differs from A. munitum
Baumgartner by the arrangement of pores and/or nodes
in the central area and by having much longer spines. See
A. sp. aff. A. pobi n. sp. for further comparison.
Measurements (µm):
Based on 9 specimens.
HT
Max.
Min.
Mean
Length
of longest ray
134
134
94
114
Width
of widest ray
50
64
41
50
Length
of longest spine
260
260
103
170
Etymology: Species name formed by an arbitrary
combination of letters (ICZN, 1985, p. 109, Appendix D,
pt. V, Recommendation 26). Species named in honour of
Dr. Peter O. Baumgartner (POB), Université de Lausanne
who investigated the early history of the hagiastrids
(Baumgartner, 1980) and whose ideas have led us to the
description of the earliest species of Archaeohagiastrum
and Hagiastrum.
Type locality: Locality 89-CNA-KUH-8, Sandilands Formation, Kunga Island, Queen Charlotte Islands, British
Columbia, Canada.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands; Bridge River Complex, British
Columbia.
Plate HAG01. Archaeohagiastrum oregonense (Yeh). Magnification x300, except Fig. 3b x600. Fig. 1(H). Yeh 1987b,
pl. 8, fig. 7. Fig. 2. OR600A-R03-11. Fig. 3. OR, OR600A, Fig. 3a. R1-1311b, Fig. 3b. R1-1311a.
Plate HAG02. Archaeohagiastrum pobi Whalen & Carter. Magnification x150. Fig. 1(H). Carter et al. 1998, pl. 10, fig. 3.
Fig. 2. QCI, GSC loc. C-305417, GSC 128801.
29
Genus: Archaeospongoprunum Pessagno 1973
Type species: Archaeospongoprunum venadoense Pessagno 1973
Synonymy:
1973 Archaeospongoprunum n. gen. – Pessagno, p. 57.
Original description: Test cylindrical, ellipsoidal, or ellipsoidal and lobate with two polar spines; polar spines triradiate or tetraradiate in axial section with longitudinally or
spirally arranged ridges alternating with grooves. Spongy
meshwork comprised of polygonal pore frames arranged in
concentric layers.
Original remarks: Archaeospongoprunum, n. gen., differs
from Spongoprunum Haeckel by possessing polar spines
with longitudinal grooves separated by longitudinal ridges.
A number of species of Archaeospongoprunum have grooves
and ridges that assume a spiral rather than a longitudinal
arrangement.
Included species:
ASP01 Archaeospongoprunum coyotense Whalen & Carter
2002
Archaeospongoprunum coyotense Whalen and Carter 2002
Species code: ASP01
Synonymy:
1987 ?Archaeospongoprunum spp. – Hattori, pl. 22, fig. 8.
1989 Archaeospongoprunum sp. – Hattori & Sakamoto, pl. 18,
fig. L.
2002 Archaeospongoprunum coyotense n. sp. – Whalen & Carter,
p. 110, pl. 4, figs. 1, 2, 6, 7, 10.
Original description: Test elongated along polar axis, cylindrical in shape; test surface planiform where it joins polar
spines. Meshwork moderately coarse, composed of large,
irregularly shaped and distributed pentagonal pore frames.
Polar spines massive, triradiate in axial section with narrow, rounded longitudinal ridges and steep-sided, longitudinal grooves; longitudinal ridges sometimes split along
proximal margin (see holotype).
Original remarks: Archaeospongoprunum coyotense n. sp.,
is distinguished from A. bipartitum Pessagno 1973, by having stronger triradiate spines, more irregular pore frames
and the absence of a sulcus. A. coyotense n. sp. is similar to
Archaeospongoprunum sp. B (pl. 5, no. 199) and A. sp. B2
30
(pl. 5, no. 201) of Yao 1997; it differs from the former in
having more tapering spines, and from the latter in having
a more cylindrical test.
Measurements (µm):
(Refer to text-figure 7 of Whalen & Carter, 2002).
(n) = number of specimens measured
AA'(9) A'S'(8)
120
180
165
206
120
143
142
165
AS(8)
53
195
105
141
BB'(9)
101
131
90
107
cc'(9)
38
38
23
31
dd'(9)
38
38
26
33
HT
Max.
Min.
Mean
Etymology: This species is named for Estero de Coyote
located to the southeast of the type area.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur, Mexico.
Occurrence: San Hipólito Formation, Baja California Sur;
Japan.
Plate ASP01. Archaeospongoprunum coyotense Whalen & Carter. Magnification x200. Fig. 1(H). Whalen & Carter
2002, pl. 4, fig. 1. Fig. 2. Whalen & Carter 2002, pl. 4, fig. 2.
31
Genus: Archaeotritrabs Steiger 1992, emend. Jud 1994
Type species: Archaeotritrabs gracilis Steiger 1992
Synonymy:
1992 Archaeotritrabs n. gen. – Steiger, p. 40.
1994 Archaeotritrabs Steiger emend. – Jud, p. 64.
1995a Archaeotritrabs Steiger emend. Jud – Baumgartner et al.,
p. 112.
Original description: Hagiastrid with three arms composed of 6 longitudinal ribs, which generate a hexagonal
cross section of the arms. The longitudinal ribs are noddy.
Between them 6 rows of simple pore frames occur. The arm
tips increase in width and have a rounded to trapezoidal
contour. The arm tips can have spines.
Emended description: By Jud (1994): Test three-rayed. Rays
of equal length, composed of 8 beams. Cross-section of rays
rectangular to octogonal. Beams connected with one another by bars forming rectangular pores on the upper and
lower sides of the test, and rectangular to trapezoidal pores
on the lateral sides. Ray tips inflated, with small, polygonal
pore-frames and usually with spines.
Original remarks: The genus Archaeotritrabs differs from
the genera of the subfamiliy of the Tritrabinae by having
simple pore rows between longitudinal ribs. After Baum-
gartner (1980) the Tritrabinae are defined by double pore
rows. It is questionable whether these forms can be related
to the Tritrabinae on the base of the hexagonal cross section
of the arms. The morphological range of the group should
be extended in the sense of having simple pore rows. Otherwise a new subfamily should be created to include simple
pore rows on the same level as double pore rows. Because
of the rare material this is actually impossible.
Further remarks: By Jud (1994): The genus was described
as possessing 6 longitudinal beams on each ray. This interpretation is a result of insufficient observation of the lateral
parts of the rays. Specimens unquestionably assignable to
A. gracilis Steiger occurring in our material prove that this
species has 8 beams and that the rays have a subrectangular cross-section. Moreover, cross-sections show that the
rays have 4 channels and not 3, as characteristic of Tritrabs
(P. Dumitrica, personal communication, and pl. 4, fig. 7).
Etymology: Greek: archaios – old. Designation signifying
that it is the probable ancestor of the genus Tritrabs.
Included species:
ATT01 Archaeotritrabs hattorii Dumitrica n. sp.
Archaeotritrabs hattorii Dumitrica n. sp.
Species code: ATT01
Synonymy:
1985 Homoeoparonaella sp. B – Nagai, pl. 1, figs. 5, 5a.
1987 Homoeoparonaella sp. O – Hattori, pl. 3, figs. 16, 17.
1987 Tritrabs (?) sp. C – Hattori, pl. 3, fig. 18.
1989 Tritrabs spp. – Hattori, pl. 38, fig. B.
Type designation: Holotype pl. ATT01, fig. 3, sample
BR871, chert of Tawi Sadh Member reworked in the Guwayza Formation, Al Khashbah Mountains, Oman.
Description: Rays equal in length, slender, spindle-shaped,
increasing slowly in diameter up to the distal third, then
decreasing to terminate in a pointed tip. Sometimes there
are 2 lateral spines in the equatorial plane originating in
the transverse ridges of lateral faces. Rays approximately
octagonal in cross-section, with 8 rows of pores separated
by 8 longitudinal beams. Pore frames rectangular but of two
types: those on oblique faces are simple whereas those on
upper, lower and lateral faces have high transverse ridges
and 2 pores in each rectangle separated by the branches of
the primary ray which form the 4 canals. One row usually
offset with pore frames of neighbouring rows. All vertices
of pore frames pointed. Central area large and triangular
outlined by three beams that connect the two longitudinal
beams on the face of each ray.
32
Remarks: Archaeotritrabs hattorii differs from A. gracilis
Steiger in having spindle-shaped rays with pointed tips,
and lacks nodes on rays. The triangular area in the center of
shell is a characteristic also known in Tritrabs casmaliaensis
(Pessagno). In the latter species the triangle has one cortical
beam in each corner, whereas A. hattorii has two. All other
characteristics, except the microsphere, differ in these two
species. The central part of the test of A. hattorii is very
thin, the cortical shell in the center of the triangular area
includes the top and the bottom of the microsphere as
illustrated for Tritrabs and the Tritrabidae (Dumitrica in
De Wever et al., 2001).
Measurements (µm):
Based on 4 specimens.
Length of rays
from the center to the distal end
Maximum diameter of rays
HT Min. Max.
140 115
140
32
47
22
Etymology: The species is named for Dr. Isamu Hattori,
Geological Laboratory, Fukui University, Japan, to honour
his valuable contribution to the knowledge of Jurassic
radiolarians of Japan.
Type locality: Sample BR871, chert of Tawi Sadh Member
reworked in the Guwayza Formation, Al Khashbah
Mountains, Oman.
Occurrence: Tawi Sadh Member of the Guwayza Formation, Oman; Mino Terrane, central Japan.
Plate ATT01. Archaeotritrabs hattorii Dumitrica n. sp. Magnification x300. Fig. 1. JP, Nanjo Massif, IH84120461.
Fig. 2. JP, Nanjo Massif, IH84120461. Fig. 3(H). OM, BR871-R09-12. Fig. 4. OM, BR871-R04-14.
Fig. 5. OM, BR871-R04-15.
33
Genus: Ares De Wever 1982a
Type species: Ares armatus De Wever 1982a
Synonymy:
1982a Ares n. gen. – De Wever, p. 202.
1986 Parares n. gen – Takemura, p. 46.
Original description: Form with three very strong spines,
recurved or not, corresponding to the extensions of the
cephalic spines A, V, and D. Six collar pores of variable
size; the largest are the cardinal pores, the smallest are
the jugular pores. Collar structure not plane, jugular and
especially cervical pores are oblique to the cardinal pores.
Spines A and V free, spine D attached to the shell by bridges.
Cephalis small, hemispherical. Thorax robust with two
spines as extensions of D and V. Pores of the flared postthoracic part (abdomen ?, velum ?) distributed in more or
less regular longitudinal rows.
Original remarks: This genus differs from Dictyoceras,
some species of which resemble A. armatus n. sp., in having two thoracic arms (extensions of V and D) instead of
three. It is distinguished from other genera by these two
characteristic arms.
Further remarks: Species lacking an apical horn or having only a very short one were assigned to Parares by
Takemura (1986). These species were later included in
the genus Ares, with Parares considered a junior synonym
(Baumgartner et al., 1995a), since the length of the apical
horn is not considered a generic character. In fact if we
take into account the range of the species as known thus
far and the morphology one can see that the evolution of
the genus followed a trend towards reduction of the apical
horn and eventually its disappearance together with exaggerated development of the ventral spine. During this
evolution the ventral spine changed its position and shape
from obliquely downward directed and straight (in the
Sinemurian species A. moresbyensis Whalen & Carter and
A. sutherlandi Whalen & Carter) to obliquely upward directed and recurved (in the Toarcian species A. avirostrum
n. sp. and the Aalenian-Bajocian species A. cylindricus
Takemura). Its exaggerated development pushed the cephalis to the dorsal side of the apex of shell. In this process
the apical horn changed also its position from practically
axial (in the Sinemurian species) to dorso-apical. During
the Pliensbachian the apical horn and ventral spine are already almost symmetrically curved and displaced from the
shell axis (A. cuniculiformis n. sp.) and during the Toarcian
the apical horn is already shorter than the ventral spine
(A. avirostrum n. sp.).
Etymology: Ares is the Greek War God, son of Zeus and
Hera, who was involved in the Trojan war when his daughter Penthesilea was killed by Achilles.
Included species and subspecies:
ARS03 Ares armatus De Wever 1982a
ARS07 Ares avirostrum Dumitrica & Matsuoka n. sp.
ARS06 Ares cuniculiformis Dumitrica & Whalen n. sp.
4061 Ares cylindricus s.l. (Takemura) 1986
3001 Ares cylindricus cylindricus (Takemura) 1986
4032 Ares cylindricus flexuosus (Takemura) 1986
ARS04 Ares mexicoensis Whalen & Carter 2002
ARS01 Ares moresbyensis Whalen & Carter 1998
ARS02 Ares sutherlandi Whalen & Carter 1998
ARS08 Ares takemurai Dumitrica & Matsuoka n. sp.
4008 Ares sp. A sensu Baumgartner et al. 1995a
Ares armatus De Wever 1982a
Species code: ARS03
Synonymy:
1982a Ares armatus n. sp. – De Wever, p. 203, pl. 10, figs. 1-4.
1982a Ares sp. 1 – De Wever, p. 203, pl. 10, figs. 5, 6.
1982b Ares armatus De Wever – De Wever, p. 335, pl. 51, figs. 2,
4, 5, 8.
1982b Ares sp. 1 – De Wever, p. 336, pl. 52, figs. 1, 2.
2004 Ares armatus De Wever – Matsuoka, fig. 131.
Original description: Cephalis small, spherical, not perforated, with an irregular, slightly pustulate surface. Apical
horn curved towards the frontal side, triradiate; grooves
correspond to the location of the secondary lateral (l) and
vertical (V) spines of the cephalic skeleton. Thorax has an
irregular surface, because of ribs between pores. Thoracic
pores and post-thoracic pores circular, closely similar in
34
size. Pores often subdivided by a star-shaped lattice. Vertical spine free, sub-rectilinear, triradiate; one ridge attached to the cephalis-thorax junction, prolongs the V
spine. Dorsal spines triradiate, slightly curved, and linked
to the shell by latticed bridges. Post-thoracic part (velum
or abdomen ?) forms a perforated veil which extends the
thorax. Circular pores quincuncially distributed on longitudinal rows. Post-thoracic part has a stricture before a
distal widening.
Original remarks: This species differs from Ares sp. by
having a partly vertical apical horn, well-differentiated
thorax with larger pores, a sub-rectilinear vertical spine,
and narrowing on the post-thoracic part close to the
thorax.
Measurements (µm):
total length (apical horn included)
length of cephalis + thorax
+ post-thoracic part
cephalis length
cephalis width
thorax length
thorax width
approx. length
of the curved apical horn
length of vertical spines
length of dorsal spine
HT
205
Av.
206
Min. Max.
200 220
140
135
100
160
20
30
30
70
21
29
30
72
20
25
30
70
24
33
32
80
155
150
120
130
160
140
125
130
Etymology: From the Latin armatus, -a, -um, adj. = armed.
Refering to martial look that evokes an Antiquity soldier.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Nanjo Massif,
Mino Terrane, central Japan.
Plate ARS03. Ares armatus De Wever. Magnification x250, except Fig. 4b x500. Fig. 1(H). De Wever 1982a, pl. 10,
fig. 4. Fig. 2. Matsuoka 2004, fig. 131. Fig. 3. De Wever 1982a, pl. 10, fig. 5. Fig. 4a, b. TR, 1662D-R11a, b.
35
Ares avirostrum Dumitrica & Matsuoka n. sp.
Species code: ARS07
Synonymy:
? 1987 Cuniculiformis sp. D – Hattori, pl. 20, fig. 14.
1989 Ares spp. – Hattori, pl. 34, figs. L, M.
2004 Ares sp. – Matsuoka, fig. 132.
Type designation: Holotype specimen MA 13750 from
sample MNA-10, Nanjo Massif, Mino Terrane, central
Japan.
Diagnosis: A species of Ares with apical horn much shorter than ventral spine; dorsal spine straight running along
thorax, and thorax campanulate with thick, undulate distal
end.
Description: Test conical with a slight constriction at the
middle of thorax resulting in a campanulate shape. Cephalis
small, poreless, displaced on the dorsal side of the apical
part of shell. Ventral spine long, curved, and bladed. Apical
horn much shorter than ventral spine; obliquely upward
directed and slightly curved when longer. Dorsal spine
straight, thin, circular in cross section beyond thorax; it is
longer than the thorax and tangential to it. Pores of thorax
quincuncially arranged, pore frames usually hexagonal,
increasing in size distally. Distal end of thorax expanded,
thick, and undulate.
Remarks: Ares avirostrum n. sp. differs from all the other
species of the genus so far known in that the distal part of
thorax is clearly delineated. It is morphologically interme-
36
diate between A. cuniculiformis n. sp. and A. cylindricus
Takemura. From the former it differs in having a much
shorter apical horn; from the latter it differs in having a
rather long apical horn and a straight dorsal spine, which
is adjacent to the wall of thorax. A. avirostrum is very close
to Ares takemurai n. sp. but differs in being slightly more
slender and the distal part of thorax is constricted and built
of intervening bars that become thinner and thinner. Ares
sp. A of Baumgartner et al. (1995a) differs from Ares avirostrum n. sp. in having a more robust and curved dorsal spine
and a narrower thorax.
Measurement (µm):
Based on 15 specimens.
Dimensions
Length of test without spines
Diameter of thorax
Length of apical horn
Length of ventral spine
HT
167
161
48
173
Min. Max.
160 200
140 197
30
63
100 173
Etymology: From the Latin avis – bird and rostrum – beak;
noun.
Type locality: Sample MNA-10, Nanjo Massif, Mino Terrane, central Japan (Matsuoka, 2004).
Occurrence: Mino Terrane, Japan; Tawi Sadh Member,
Guwayza Formation, Oman; Fernie Formation, NE British
Columbia.
Plate ARS07. Ares avirostrum Dumitrica & Matsuoka n. sp. Magnification x200, except Fig. 1c(H) x800. Fig. 1(H).
JP MNA-10. Fig. 1a(H). Matsuoka 2004, fig. 132. Fig. 1b(H). MA13749. Fig. 1c(H). MA13748. Fig. 2. JP, MNA-10,
MA13771. Fig. 3. JP, MNA-10, MA13778. Fig. 4. NBC, GSC loc. C-305813, GSC 111805. Fig. 5. OM, BR871-R07-20.
Fig. 6. OM, BR524-R05-09. Fig. 7. OM, BR528-R10-11.
37
Ares cuniculiformis Dumitrica & Whalen n. sp.
Species code: ARS06
Synonymy:
? 1987 Cuniculiformis sp. A – Hattori, pl. 20, fig. 11.
? 1987 Cuniculiformis sp. B – Hattori, pl. 20, fig. 12.
? 1989 Gen. 2, sp. 1 – Hattori, pl. 21, fig. K.
? 1997 Ares sp. A0 – Yao, pl. 8, fig. 382.
Type designation: Specimen figs. 1 and 1a, stub 1662DR03-13, sample 1662D, Gümüslü Allochthon, Taurus Mts.,
Turkey.
Diagnosis: A species of Ares with a conical test, A and V
spines relatively equal and recurved, and D spine long, over
twice length of test.
Description: Test high-conical with elliptical or rounded
polygonal pores. Cephalis small, poreless situated on the
dorsal side of the apical part of test, surface weakly pustulate.
Apical and ventral spines robust, bladed, recurved and
slightly dissimilar, usually slightly shorter than D spine
and may be more curved. D spine downwardly directed,
slightly curved, running along thorax and continuing a
long distance beyond.
Remarks: A. cuniculiformis n. sp. resembles A. armatus
De Wever, with which it partly co-occurs, but it differs in
having a narrower test, A and V spines are relatively equal
38
in length and both are recurved. There is a slight difference
between this species from the Pliensbachian (Turkey and
Baja California) and the species from the lower Toarcian
(Japan). In the former (and especially the holotype) the
apical horn is almost of the same length as the ventral horn,
whereas in the latter, the apical horn is shorter. They are
probably part of the phylogenetical lineage that gave rise to
A. cylindricus Takemura.
Measurements (µm):
Based on 4 specimens.
Length of test excluding spines
Length of ventral spine
Length of apical horn
Maximum diameter of thorax
HT Min. Max.
187 119 187
153 117 153
147 50
147
120 75
130
Etymology: From the Latin cuniculus – hare and forma–
shape; noun.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; San Hipólito
Formation, Baja California Sur.
Plate ARS06. Ares cuniculiformis Dumitrica & Whalen n. sp. Magnification x200, except fig. 1b(H) x400.
Fig. 1(H)a, b. TR, 1662D-R03-13a, b. Fig. 2. BCS, BPW30.
39
Ares cylindricus s.l. (Takemura) 1986
Species code: 4061
Synonymy:
1986 Parares cylindricus n. sp. – Takemura, p. 46, pl. 4, figs. 3-7.
1986 Parares flexuosus n. sp. – Takemura, p. 47, pl. 4, figs. 8-11.
? 1986 Nassellaria gen. et sp. indet. C in Yao et al. 1982 –
De Wever & Cordey, pl. 1, fig. 11.
1987 Parares (?) aff. P. cylindricus Takemura – Hattori, pl. 20,
fig. 9.
1995a Ares cylindricus (Takemura) – Baumgartner et al., p. 116.
See also subspecies.
Included subspecies.
3001 Ares cylindricus cylindricus (Takemura) 1986
4032 Ares cylindricus flexuosus (Takemura) 1986
Ares cylindricus cylindricus (Takemura) 1986
Species code: 3001
Synonymy:
1986 Parares cylindricus n. sp. – Takemura, p. 46, pl. 4, figs. 3-7.
1987 Parares (?) aff. P. cylindricus Takemura – Hattori, pl. 20,
fig. 9.
1989 Parares cylindricus Takemura – Kito, p. 204, pl. 23, fig. 11.
1995a Ares cylindricus cylindricus (Takemura) – Baumgartner
et al., p. 116, pl. 3001, figs. 1-4.
1997 Ares cylindricus cylindricus (Takemura) – Yao, pl. 8,
fig. 385.
Original description: Cephalis small, poreless and spherical, with well developed and triradiate vertical spine.
Vertical spine curved downward and in some specimens,
slightly curved upward distally. Thorax cylindrical and
long, with elliptical pores arranged longitudinally and
hexagonally. No apertural ring at the end of the thorax.
Dorsal spine strong and triradiate, about twice as long as
thorax. Dorsal spine curved slightly proximally and distally straight and slightly twisted anticlockwisely. Some bars
connecting dorsal spine and thoracic wall at several points
in some specimens.
Original remarks: Parares cylindricus n. sp. differs from
P. flexuous n. sp. in cylindrical thorax and distally straight
dorsal spine.
Measurements (µm):
Based on 10 specimens.
Length of shell including two spines
Height of cephalo-thorax
Maximum width of shell including two spines
Width of thorax
Min.
470
185
345
105
Max.
610
270
465
135
Etymology: The species name, cylindricus, cylindrical in
English is derived from the shape of thorax.
Type locality: Sample TKN-105, Komami, Yamato village,
Gifu Prefecture, central Japan.
Occurrence: Japan, Italy.
Ares cylindricus flexuosus (Takemura) 1986
Species code: 4032
Synonymy:
1986 Parares flexuous n. sp. – Takemura, p. 47, pl. 4, figs. 8-11.
1989 Parares flexuous Takemura – Kito, p. 204, pl. 23, fig. 12.
1995a Ares cylindricus flexuosus (Takemura) – Baumgartner et al.,
p. 118, pl. 4032, figs. 1-3.
1997 Ares cylindricus flexuosus (Takemura) – Yao, pl. 8, fig. 388.
Original description: Cephalis small, spherical and poreless, with stout and triradiate vertical spine, which is
curved downward distally. Thorax conical to subconical
and slightly narrow distally, without apertural ring. Thoracic pores spherical to ellipsoidal, usually arranged longitudinally and hexagonally. Dorsal spine which is twisted
anticlockwisely strong, triradiate and remarkably curved
downwardly.
Original remarks: Parares flexuous n.sp. is distinguished
from P. cylindricus by its conical to subconical thorax and
markedly curved dorsal spine.
40
Further remarks: By Baumgartner et al. (1995a): The name
flexuous is emended (I.C.Z.N., art.33a (I)) into flexuosus,
which is the correct Latin name.
Measurements (µm):
Based on 4 specimens.
Length of shell including two spines
Height of cephalo-thorax
Maximum width of shell including two spines
Width of thorax
Min.
280
145
360
100
Max.
335
175
420
110
Etymology: The species name, flexuosus, means bending,
derived from its curved dorsal spine.
Type locality: Sample TKN-105, Komami, Yamato Village,
Gifu Prefecture, central Japan.
Occurrence: Japan, Italy.
Plate 4061. Ares cylindricus s.l. (Takemura). Magnification x200. Fig. 1(H). Ares cylindricus cylindricus (Takemura)
1986, pl. 4, fig. 4. Fig 2(H). Ares cylindricus flexuosus (Takemura) 1986, pl. 4, fig. 8. Fig. 3. JP, MIN-1, MA09001.
Fig. 4. JP, MKM-1, MA10325. Fig. 5. JP,GUH39-14, RH676. Fig. 6. JP, GUH39-11, RH673. Fig. 7. JP, MIN-10, MA09917.
41
Ares mexicoensis Whalen & Carter 2002
Species code: ARS04
Synonymy:
thorax, and a steeper-sided thorax; with its sturdy arms and
horn Ares mexicoensis n. sp. is a more robust species than
Ares moresbyensis Whalen and Carter 1998.
Original description: Small, dome-shaped cephalis with
massive apical horn; cephalis mostly imperforate with
some small pores near base. Horn triradiate in axial section
with narrow, rounded, longitudinal ridges and broad,
longitudinal grooves. Thorax gradually increasing in
width distally; meshwork on thorax composed of irregular
pentagonal and tetragonal pore frames (mostly pentagonal)
becoming slightly larger distally. Thoracic arms massive,
gently curving downward with one arm slightly longer
than other; shorter arm positioned higher on thorax at
base of cephalis; arms triradiate in axial section with
narrow, longitudinal ridges alternating with steep-sided
longitudinal grooves; narrow, transverse thoracic ridges,
formed by a slight alignment of the pore frames, continue
onto arms, joining with longitudinal ridges.
Measurements (µm):
Based on 14 specimens.
1984 Ares sp. – Whalen & Pessagno, pl. 1, fig. 11.
2002 Ares mexicoensis n. sp. – Whalen & Carter, p. 140, pl. 15,
figs. 5, 12; pl. 18, figs. 6, 9.
Original remarks: Ares mexicoensis n. sp., is distinguished
from A. armatus De Wever 1982, in having a shorter
cephalic horn and thoracic arms, larger pore frames on the
Length
(excludes horn)
150
150
105
134
Length
of long arm
128 (broken)
210
135
169
Length
of short arm
128
150
105
133
HT
Max.
Min.
Mean
Etymology: This species is named for the United States of
Mexico.
Type locality: Sample BPW80-30, San Hipólito Formation,
Punta San Hipólito, Vizcaino Peninsula, Baja California.
Occurrence: San Hipólito Formation, Baja California Sur;
Fannin Formation, Queen Charlotte Islands; Tawi Sadh
Member, Guwayza Formation, Oman.
Ares moresbyensis Whalen & Carter 1998
Species code: ARS01
Synonymy:
Not 1984 Ares sp. – Whalen & Pessagno, pl. 1, figs. 11, 12.
1998 Ares moresbyensis n. sp. – Whalen & Carter, p. 75, pl. 21,
figs. 1, 2, 11; pl. 27, figs. 2, 8.
Original description: Test with small, dome-shaped cephalis and prominent tapering horn; horn approximately
equal in length to thorax, triradiate in axial section with
narrow, rounded longitudinal ridges and broad, rounded
longitudinal grooves; horn not aligned exactly with long
axis of test. Cephalis with small to medium polygonal pore
frames sometimes obscured by a layer of microgranular
silica. Thorax elongate, trapezoidal in outline with large,
pentagonal, hexagonal, and circular pore frames. Two
prominent spines attached to thorax at base of cephalis
at 45º angle with long axis of test; spines tapering distally,
triradiate in axial section with narrow, rounded longitudinal ridges and broad, rounded longitudinal grooves; near
base of cephalis, longitudinal ridges extend onto thorax
forming prominent transverse ridges. One spine slightly
longer and more massive than other; longer spines equal
to length of thorax.
42
Original remarks: The larger, more irregularly shaped
thoracic pore frames of Ares moresbyensis n. sp. distinguish it from A. sutherlandi n. sp.
Measurements (µm):
(n) = number of specimens measured.
Length
Width
(excluding horn) (6) of thorax (6)
105
71
120
75
90
71
75
112
Maximum length
of short arm (5)
90
120
68
92
HT
Max.
Min.
Mean
Etymology: This species is named for Moresby Island,
Queen Charlotte Islands, British Columbia, located to the
west of the type locality.
Type locality: Sample QC-675, Sandilands Formation,
Kunga Island - north side, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands Formation, Queen Charlotte
Islands.
Plate ARS04. Ares mexicoensis Whalen & Carter. Magnification x200. Fig. 1(H). Whalen & Carter 2002, pl. 15, fig. 5.
Fig. 2. QCI, GSC loc. C-304567, GSC 128706. Fig. 3. QCI, GSC loc. C-140495, GSC 128707. Fig. 4. OM, BR485-R20-10.
Plate ARS01. Ares moresbyensis Whalen & Carter. Magnification x250. Fig. 1(H). Carter et al. 1998, pl. 21, fig. 1.
43
Ares sutherlandi Whalen & Carter 1998
Species code: ARS02
Synonymy:
1984 Ares sp. – Whalen & Pessagno, pl. 1, fig. 12.
1998 Ares sutherlandi n. sp. – Whalen & Carter, p. 76, pl. 21,
figs. 3, 16; pl. 27, figs. 1, 7.
2002 Ares sp. A – Whalen & Carter, p. 142, pl. 15, figs. 6, 13.
Original description: Test with small, dome-shaped cephalis with prominent broad, tapering horn; horn approximately one-half length of cephalis and thorax combined,
triradiate in axial section with narrow, rounded longitudinal ridges and broad, rounded longitudinal grooves.
Cephalis with variably-sized pore frames usually partially
obscured by layer of microgranular silica. Thorax elongate,
cylindrical, with irregularly arranged small- to mediumsized polygonal (mostly pentagonal) pore frames; thoracic
pore frames sometimes partially masked by an outer layer
of irregular, polygonal pore frames. Two prominent spines
attached to thorax and base of cephalis; spines triradiate in
axial section with narrow, rounded longitudinal ridges and
broad, rounded longitudinal grooves. Larger, more robust
spine forming an approximate 55° angle with long axis of
test; larger spines bonded to thorax by approximately four
narrow, linear ridges which extend from longitudinal ridge
of spine and continue obliquely across proximal part of test;
smaller spine at 40° to long axis of test.
Original remarks: See remarks under A. moresbyensis n. sp.
Measurements (µm):
(n) = number of specimens measured.
Length
Width
(excluding horn) (6) of thorax (6)
109
120
83
97
75
75
75
75
Maximum
length
of short arm (5)
90
90
60
75
HT
Max.
Min.
Mean
Etymology: This species is named in honor of A. Sutherland
Brown (British Columbia Department of Mines and
Petroleum Resources, Vancouver, B.C.) who first mapped
the entire Queen Charlotte Archipelago and provided
a detailed account of the geology.
Type locality: Sample QC-675, Sandilands Formation,
Kunga Island - north side, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands Formation, Queen Charlotte
Islands; San Hipólito Formation, Baja California Sur.
Ares takemurai Dumitrica & Matsuoka n. sp.
Species code: ARS08
Synonymy:
? 1987 Parares (?) sp. A – Hattori, pl. 20, fig. 8.
1987 Parares (?) sp. C – Hattori, pl. 20, fig. 10.
1989 Ares sp. A – Hattori, pl. 3, fig. K.
1989 Parares (?) spp. – Hattori, pl. 3, fig. M, not fig. L.
2004 Ares sp. – Matsuoka, fig. 133.
Type designation: Holotype specimen MA 13779 from
sample MNA-10, Nanjo Massif, Mino Terrane, Japan.
Diagnosis: A species of Ares with thorax constricted
medially as well as distally. Apical horn much shorter than
V spine, D very long and straight.
Description: Test conical, two-segmented with a slight
constriction in the middle part of thorax. Cephalis small,
poreless, displaced on the dorsal side of the apical part of
shell. Ventral spine long, curved, and bladed. Apical horn
much shorter than ventral spine, obliquely directed upward and slightly curved. Dorsal spine straight, thin, very
long, one to two times length of thorax, circular in cross
section beyond thorax, three-bladed on the tangential portion. Pores of thorax quincuncially arranged, pore frames
usually hexagonal, increasing in size distally. Distal half of
thorax expanded, convex in outline, constricted terminally
and built of intervening bars that become ever thinner.
Distal border irregular.
44
Remarks: Ares takemurai n. sp. is very close to A. avirostrum n. sp.: the A, V and D spines and proximal half of
thorax are morphologically similar, the only major difference is the shape of the distal half of thorax. This part is
expanded with a clear-cut border in A. avirostrum, whereas in A. takemurai the distal half of thorax is also expanded, but the terminal part is constricted and made of intervening bars that become ever thinner.
Measurements (µm):
Based on 11 specimens
Dimensions
Length of test excluding spines
Length of ventral spine
Length of apical horn
Length of dorsal spine
Maximum diameter of thorax
HT Min. Max.
170 95
190
170 75
180
70 40
70
390 90
400
160 85
160
Etymology: The species is named for Dr. Atsushi Takemura,
Hygoyo University of Teacher Education, Japan, to honour
his valuable contribution to the taxonomy of Jurassic
Radiolaria.
Type locality: Sample MNA-10, Nanjo Massif, Mino
Terrane, Japan.
Occurrence: Nanjo Massif, Mino Terrane, Japan; Tawi Sadh
Member of the Guwayza Formation, Oman.
Plate ARS02. Ares sutherlandi Whalen & Carter. Magnification x300. Fig. 1(H). Carter et al. 1998, pl. 21, fig. 3.
Fig. 2. Whalen & Carter 2002, pl. 15, fig. 6.
Plate ARS08. Ares takemurai Dumitrica & Matsuoka n. sp. Magnification x200. Fig. 1. JP, MNA-10, MA13743.
Fig. 2. JP, MNA-10, MA13608. Fig. 3(H). Matsuoka 2004, fig. 133.
45
Ares sp. A sensu Baumgartner et al. 1995a
Species code: 4008
Synonymy:
1982 Nassellaria D (in Yao et al., 1982) – Matsuda & Isozaki,
pl. 1, fig. 21.
1986 Parares sp. – Takemura, p. 47, pl. 4, fig. 12.
1987 Parares (?) sp. A – Hattori, pl. 20, fig. 8.
1990 Ares ? sp. D – Hori, Fig. 8.25.
1995a Ares sp. A – Baumgartner et al., p. 118, pl. 4008, fig. 1.
1996 Ares sp. A – Yeh & Cheng, p. 118, pl. 2, fig. 8; pl. 7,
figs. 1, 2, 12.
1997 Ares sp. A Baumgartner et al. – Yao, pl. 8, fig. 387.
1997 Ares ? sp. D of Yao – Hori, pl. 1, fig. 11.
Original remarks: This species differs from A. cylindricus
by having a short triradiate apical horn.
Occurrence: Mino Terrane, Japan; Liminangcong Chert,
Philippines.
Genus: Atalanta Cordey & Carter 1996
Type species: Atalanta emmela Cordey & Carter 1996
Synonymy:
1996 Atalanta n. gen. – Cordey & Carter, p. 446.
Original diagnosis: Multicyrtid nassellarian. Proximal part
of test smooth, without ornamentation. Two rows of pore
frames per segment; pores regularly aligned transversely
and obliquely, not longitudinally.
Original description: Multicyrtid test conical and may be
constricted distally. Cephalis nonperforate with a horn.
Proximal part of test smooth without ornamentation.
Postabdominal segments have two transverse rows of circular pores arranged hexagonally. Pores regularly aligned
transversely and obliquely, not longitudinally. Proximal
segments show no development of transverse ridges. More
distally, segment junctions thicken to form transverse
ridges that show a slight to moderately exaggerated zig-zag
outline. Nodes more or less well developed at pore frame
intersections. Depending on species and (or) preservation,
the conical test may be constricted distally.
Original remarks: Atalanta n. gen. differs from Wrangellium Pessagno and Whalen (1982) by possessing hexagonal pore frames that are not aligned longitudinally. It
differs from Triversus Takemura (1986) in lacking an amphipyndax-type cephalic skeletal structure and in possessing two, rather than three, rows of pores per chamber. Atalanta n. gen. differs from Proparvicingula Carter (1993),
Parvicingula Pessagno (1977), and Ristola Pessagno &
Whalen (1982) by possessing two rows of pores between
46
circumferential ridges instead of three, and further differs
from Proparvicingula Carter in having a single rather than
double test wall. Atalanta n. gen. differs from Pseudoristola
Yeh (1987b) by possessing well-developed circumferential
ridges on postabdominal chambers, and the final chamber
is open rather than bulbous and closed. Comparisons with
Nitrader n. gen are developed under that genus.
Remarks under Nitrader n. gen.: Nitrader has clear external affinities with Atalanta n. gen., as both possess
two rows of offset pores set in hexagonal pore frames arranged between two circumferential ridges, and both
have an apical horn. Nevertheless, the pore arrangement
of Nitrader is different from Atalanta in that the pores
are H-linked on each side of transverse ridges. The structure of the proximal chambers also differs: Atalanta has
a smooth proximal portion with no clear signs of segmentation, whereas Nitrader develops an external ornamentation composed of a rugose network of segmentation
similar to Proparvicingula Carter, a new genus recently described from the Rhaetian of the Queen Charlotte Islands
(Carter 1993). This suggests that Nitrader n. gen. could be
an intermediate form between Proparvicingula Carter and
Atalanta n. gen.
Etymology: Atalanta is an arbitrary combination of letters
(ICZN 1985, article 11b (iii), p. 20).
Included species:
ATA02 Atalanta emmela Cordey & Carter 1996
Plate 4008. Ares sp. A sensu Baumgartner et al. Magnification x200. Fig. 1. JP, IYII 12-50. Fig. 2. JP, Hori 1990, fig. 8.25.
Fig. 3. Baumgartner et al. 1995a, pl. 4088, fig. 1.
47
Atalanta emmela Cordey & Carter 1996
Species code: ATA02
Synonymy:
1991 Gen. indet. Z sp. A – Tipper et al., pl. 8, fig. 8.
1996 Atalanta emmela n. gen., n. sp. – Cordey & Carter, p. 447,
pl. 1, figs. 1-3.
1998 Atalanta emmela Cordey & Carter – Cordey, p. 126, pl. 25,
fig. 1.
1998 Atalanta emmela Cordey & Carter – Whalen & Carter,
p. 67, pl. 24, fig. 13.
2001 Atalanta emmela Cordey & Carter – Gawlick et al., pl. 2,
fig. 22.
2002 Atalanta emmela Cordey & Carter – Whalen & Carter,
p. 128, pl. 16, figs. 1, 8.
2002 Atalanta emmela Cordey & Carter – Tekin, p. 190, pl. 4,
figs. 10, 11.
Original diagnosis: Multicyrtid nassellarian, conical. Cephalis with simple horn. Proximal part of test smooth,
without ornamentation.
Original description: Multicyrtid test, conical. Cephalis
nonperforate with a horn of medium length disposed asymmetrically. Proximal part of test smooth, without ornamentation. Post-abdominal segments have two transverse rows
of circular pores arranged in hexagonal pore frames. Proximal segments show no development of transverse ridges.
Transverse ridges begin to develop at one-third to one-half
of total length of test at junctions between segments; ridges
formed by a thickening of the pore frames. Nodes more or
less well-developed at pore frame intersections. Width of
segments increases regularly towards the distal aperture.
Original remarks: Atalanta emmela, n. sp., differs from
Atalanta epaphrodita, n. sp., in possessing a single horn,
more weakly developed thickening of transverse ridges,
and the test is not constricted distally.
Measurements (µm):
Based on 4 specimens.
Length of horn
Length of test, excluding horn
Maximum diameter of test
HT
35
220
125
Min.
30
200
105
Max.
40
260
140
Etymology: From the greek emmeles meaning harmonious.
Type locality: GSC loc. C-150155, Sandilands Formation,
Kunga Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands; chert clast from Voght Creek conglomerate, south
Intermontane Belt, British Columbia; San Hipólito Formation, Baja California Sur; Dürrnberg Formation, Austria;
Hocaköy Radiolarite, Turkey.
Genus: Bagotum Pessagno & Whalen 1982
Type species: Bagotum maudense Pessagno & Whalen 1982
Synonymy:
1982 Bagotum n. gen. – Pessagno & Whalen, p. 117.
Original description: Test ellipsoidal; final post-abdominal
chamber terminating in latticed, hemispherical cap. Cephalis lacking horn.
Original remarks: Bagotum n. gen., differs from Stichocapsa Rüst (1885, type species S. jaspidea Rüst, 1885) by displaying a thick, double-layered test, typically with more
irregular pore frames. Furthermore, whereas the proximal
part of the test of Bagotum is bluntly rounded or domeshaped, that of Stichocapsa is conical. Both genera possess
dome-shaped latticed caps on their final post-abdominal
chambers. Bagotum, n. gen., differs from Droltus n. gen.,
48
by lacking a horn, being ellipsoidal rather than conical and
by having a final post-abdominal chamber terminating in
a latticed, dome-shaped cap.
Etymology: Bagotum is a name formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, Pt. VI,
Recommendation 40, p.113).
Included species:
BAG01 Bagotum erraticum Pessagno & Whalen 1982
BAG03 Bagotum funiculum Whalen & Carter 2002
BAG02 Bagotum helmetense Pessagno & Whalen 1982
BAG04 Bagotum kimbroughi Whalen & Carter 2002
BAG05 Bagotum maudense Pessagno & Whalen 1982
BAG06 Bagotum modestum Pessagno & Whalen 1982
BAG07 Bagotum pseudoerraticum Kishida & Hisada 1985
Plate ATA02. Atalanta emmela Cordey & Carter. Magnification x250. Fig. 1(H). Cordey & Carter 1996, pl. 1, fig. 2.
Fig. 2. Whalen & Carter 2002, pl. 16, fig. 1.
49
Bagotum erraticum Pessagno & Whalen 1982
Species code: BAG01
Synonymy:
1982 Bagotum erraticum n. sp. – Pessagno & Whalen, p. 117,
pl. 1, fig. 10.
1988 Bagotum aff. erraticum Pessagno & Whalen – Li, pl. 1, fig. 3.
1995 Bagotum erraticum Pessagno & Whalen – Suzuki, pl. 8,
fig. 1.
1998 Bagotum sp. cf. B. erraticum Pessagno & Whalen
– Kashiwagi, pl. 2, figs. 6, 7.
2001 Bagotum erraticum Pessagno & Whalen – Gawlick et al.,
pl. 5, fig. 7.
Original description: Test inflated, broader distally than
proximally, usually with four post-abdominal chambers.
Cephalis hemispherical; remaining chambers trapezoidal
in outline; final post-abdominal chamber with broad,
dome-shaped cap. Post-abdominal chambers increasing
slowly in length; increasing moderately rapidly in width
proximally; final one or two post-abdominal chambers
decreasing slightly in width. Proximal one-third of test with
vermicular appearance due to presence of irregular, often
elongate polygonal pore frames in outer layer. Remainder
of test with more regular tetragonal (frequently square or
rectangular) and pentagonal pore frames which sometimes
are aligned longitudinally in rows.
Original remarks: Bagotum erraticum, n sp., is compared
with Bagotum maudense, n. sp., under the latter species.
Further remarks: The more irregular pore frames and
inflated distal portion of test distinguish this species from
Bagotum maudense Pessagno & Whalen.
Measurements (µm):
Based on 11 specimens.
Length
212.5
225
175
194.7
Width (maximum)
100
112.5
87.5
99.7
HT
Max.
Min.
Mean
Etymology: Erraticus-a-um (Latin, adj.) = erratic.
Type locality: Sample QC-549, Sandilands Formation
(Kunga Formation in Pessagno & Whalen, 1982), Queen
Charlotte Islands, British Columbia.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands; Dürrnberg Formation, Austria;
Tawi Sadh Member of the Guwayza Formation and Musallah Formation, Oman; Dengqen area, Tibet; Japan.
Bagotum funiculum Whalen & Carter 2002
Species code: BAG03
Synonymy:
Original remarks: Bagotum funiculum n. sp. is distinguished from B. erraticum Pessagno and Whalen 1982, by
the more massive linear pore frames on the distal postabdominal chambers.
Original description: Test with approximately four postabdominal chambers. Cephalis hemispherical with small
spine; post-abdominal chambers trapezoidal in outline,
gradually increasing in width; final post-abdominal
chamber abruptly decreasing in width, terminating in
closed dome-like cap. Post-abdominal chambers gradually
increasing in height. Cephalis and thorax composed
of small, irregularly shaped pore frames (circular and
tetragonal) on outer latticed layer. Pore frames of outer
latticed layer on first few post-abdominal chambers very
irregularly distributed and shaped (circular, elongate,
pentagonal, tetragonal). Pore frames of outer latticed
layer on distal half of test composed of regular, square
to rectangular pore frames with bars strongly aligned
longitudinally.
Measurements (µm):
(n) = number of specimens measured
1984 Bagotum spp. – Whalen & Pessagno, pl. 2, figs. 7, 8, 11.
2002 Bagotum funiculum n. sp. – Whalen & Carter, p. 114, pl. 9,
figs, 5, 6, 9.
50
Length (4)
240
255
225
236
Width (max.) (5)
150
150
86
98
HT
Max.
Min.
Mean
Etymology: Funiculus, i (Latin, m) = thin rope, cord, string.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur, Mexico.
Occurrence: San Hipólito Formation, Baja California Sur;
Fannin Formation, Queen Charlotte Islands.
Plate BAG01. Bagotum erraticum Pessagno & Whalen. Magnification x300. Fig. 1(H). Pessagno & Whalen 1982,
pl. 1, fig. 10. Fig. 2. QCI, GSC loc. C-305388, GSC 128712. Fig. 3. OM-99-83, 011401. Fig. 4. OM-251, 021530.
Fig. 5. JP, MNA-10, MA12232.
Plate BAG03. Bagotum funiculum Whalen & Carter. Magnification x300. Fig. 1(H). Whalen & Carter 2002,
pl. 9, fig. 5. Fig. 2. QCI, GSC loc. C-175310, GSC 128713.
51
Bagotum helmetense Pessagno & Whalen 1982
Species code: BAG02
Synonymy:
1982 Bagotum? helmetense n. sp. – Pessagno & Whalen, p. 118,
pl. 1, fig. 11; pl. 12, fig. 23.
1984 Bagotum spp. – Whalen & Pessagno, pl. 2, figs. 1, 2.
1993 Bagotum (?) helmetense Pessagno & Whalen – Kashiwagi
& Yao, pl. 1, fig. 6.
1998 Bagotum helmetense Pessagno & Whalen – Whalen &
Carter, p. 61, pl. 15, fig. 1; pl. 26, fig. 1.
1998 Bagotum (?) helmetense Pessagno & Whalen – Kashiwagi,
pl. 2, fig. 4, not fig. 5.
2002 Bagotum helmetense Pessagno & Whalen – Whalen &
Carter, p.114, pl. 10, figs. 1, 14.
2002 Bagotum helmetense Pessagno & Whalen – Tekin, p. 186,
pl. 3, fig. 6.
Original description: Test as with genus, usually with six
post-abdominal chambers. Cephalis hemispherical with
small spine; post-abdominal chambers trapezoidal in outline. Proximal post-abdominal chambers gradually increasing in width; distal post-abdominal chambers gradually decreasing in width; domelike cap on final post-abdominal
chamber with irregular tetragonal and pentagonal pore
frames. Post-abdominal chambers gradually increasing in
height. Cepahlis and thorax with small, irregularly shaped
pore frames (circular and elongate) in outer latticed layer;
larger, irregularly shaped polygonal pore frames (mostly
elongate) in outer latticed layer of post-abdominal chambers. Pore frames of outer latticed layer not aligned on any
portion of test.
Original remarks: Bagotum (?) helmetense, n. sp., differs
from all other species of Bagotum by showing no linear
arrangement of pore frames. Peculiar circular structure
(diameter approximately 1/3 width of test) formed by pore
frames of outer latticed layer obscured on some specimens;
inner latticed layer of test exposed in center of circular
structure.
Further remarks: The circular structure formed by pore
frames of the outer latticed layer, that was first observed
on upper Sinemurian bagotids from the Queen Charlotte
Islands, is also observed on some specimens from Baja
California Sur.
Measurements (µm):
Based on 10 specimens.
Length
175
198
142
167.8
Width (max.)
80
90
63
78.3
HT
Max.
Min.
Mean
Etymology: Bagotum helmetense, n. sp., is named for
Helmet Island, northwest of its type locality.
Type locality: Sample QC 590A, Sandilands Formation
(Kunga Formation in Pessagno & Whalen, 1982), Queen
Charlotte Islands, British Columbia.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands; San Hipólito Formation, Baja
California Sur; Hocaköy Radiolarite, Turkey; Japan.
Bagotum kimbroughi Whalen & Carter 2002
Species code: BAG04
Synonymy:
1984 Bagotum spp. – Whalen & Pessagno, pl. 2, fig. 3-6.
1987 Bagotum aff. B. maudense – Hattori, pl. 15, fig. 3.
? 1987b Drulanta (?) sp. A – Yeh, pl. 4, fig. 27.
? 1987b Drulanta (?) sp. C – Yeh, p. 74, pl. 4, fig. 26.
1992 Bagotum? sp. – Sashida, pl. 1, fig. 18.
1998 Bagotum sp. C – Whalen & Carter, p. 62, pl. 15, fig. 3.
2002 Bagotum kimbroughi n. sp. – Whalen & Carter, p. 114,
pl. 9, figs. 7, 8, 12-15; pl. 17, figs. 4, 5.
Original description: Test large, strong, usually with four
to five post-abdominal chambers. Cephalis hemispherical; abdomen and post-abdominal chambers trapezoidal
in outline. All post-abdominal chambers rapidly increasing in width distally till large, final post-abdominal chamber which terminates in dome-like cap; post-abdominal
chambers gradually increasing in height distally. Cephalis
and thorax with small, irregularly shaped pore frames in
outer latticed layer; larger, regularly shaped tetragonal pore
frames (square and rectangular) in outer latticed layer of
post-abdominal chambers aligned in rows. Circular struc52
ture (diameter approximately one third width of test) observed in outer latticed layer of test; inner latticed layer exposed in center of circular structure.
Original remarks: The larger pore frames and the eccentric,
ellipsoidal test of Bagotum kimbroughi n. sp., with its final
post-abdominal chamber almost twice as wide as the thorax
and abdomen, distinguish it from B. modestum Pessagno
and Whalen. Circular structures similar to those reported
on specimens of Bagotidae and Canutidae from the
Sandilands and Ghost Creek formations, Queen Charlotte
Islands, British Columbia.
Measurements (µm):
Based on 11 specimens.
Length
203
248
180
203
Width (max.)
120
135
120
127
HT
Max.
Min.
Mean
Plate BAG02. Bagotum helmetense Pessagno & Whalen. Magnification x300. Fig. 1(H). Pessagno & Whalen 1982,
pl. 1, fig. 11. Fig. 2. Whalen & Carter 2002, pl. 10, fig. 1. Fig. 3. QCI, GSC loc. C-305386, GSC 128714.
Etymology: This species is named for Dr. David
Kimbrough (San Diego State University) a noted student
of the Mesozoic rocks of the Vizcaino Peninsula and
Cedros Island.
Type locality: Sample BPW80-30, San Hipólito Formation,
Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Sandilands Formation, Queen Charlotte Islands; Japan.
Plate BAG04 Bagotum kimbroughi Whalen & Carter. Magnification x300. Fig. 1(H). Whalen & Carter 2002, pl. 9, fig. 7.
Fig. 2. Whalen & Carter 2002, pl. 9, fig. 8.
53
Bagotum maudense Pessagno & Whalen 1982
Species code: BAG05
Synonymy:
1982 Bagotum maudense n. sp. – Pessagno & Whalen, p. 118,
pl. 3, figs. 6, 11, 20.
1984 Bagotum sp. aff. B. modestum Pessagno & Whalen –
Murchey, pl. 1, fig. 29.
1987b Bagotum sp. aff. B. maudense Pessagno & Whalen – Yeh,
p. 53, pl. 9, fig. 12; pl. 28, fig. 13.
1989 Bagotum sp. aff. B. maudense Pessagno & Whalen –
Hattori, pl. 10, fig. K.
? 1996 Bagotum modestum Pessagno & Whalen – Pujana,
p. 137, pl. 1, fig. 11.
1997 Parahsuum maudense (Pessagno & Whalen) – Yao, pl. 13,
fig. 637.
1997 Parahsuum sp. NC2 – Yao, pl. 13, fig. 643.
1998 Bagotum sp. aff. B. maudense Pessagno & Whalen –
Kashiwagi, pl. 1, fig. 4.
2001 Bagotum maudense Pessagno & Whalen – Gawlick et al.,
pl. 5, fig. 8.
Original description: Test relatively elongate, central portion nearly cylindrical. Cephalis hemispherical; remaining
chambers trapezoidal in outline. Post-abdominal chambers
gradually increasing in length; first one or two post-abdominal chambers increasing moderately rapidly in width;
increasing more gradually in width medially in cylindrical
portion of test; final one or two post-abdominal chambers
decreasing in width. Cephalis and thorax with small irregular tetragonal and pentagonal pore frames; pore frames of
other chambers a mixture of large pentagonal and tetragonal pore frames. Domelike cap on final post-abdominal
54
chamber with relatively large, irregular tetragonal and
pentagonal pore frames. Test typically with seven postabdominal chambers.
Original remarks: Bagotum maudense, n. sp., differs from
B. erraticum, n. sp., by having a slender, less inflated test
that is centrally more cylindrical in character. Furthermore,
B. maudense possesses a greater preponderance of linearly
arranged square to rectangular pore frames.
Measurements (µm):
Based on 10 specimens.
Length
250
287.5
210
219.25
Width (max.)
100
130
90
108
HT
Max.
Min.
Mean
Etymology: This species is named for Maude Island, its
type locality.
Type locality: Sample QC 534, Fannin Formation (Maude
Formation in Pessagno & Whalen, 1982), Queen Charlotte
Islands, British Columbia.
Occurrence: Fannin Formation, Queen Charlotte Islands;
Nicely and Hyde formations and Warm Springs member
of the Snowshoe Formation, Oregon; Franciscan Complex,
California; Sierra Chacaicó Formation, Argentina; Dürrnberg Formation, Austria; Japan.
Plate BAG05. Bagotum maudense Pessagno & Whalen. Magnification x300. Fig. 1a(H). Pessagno & Whalen 1982,
pl. 3, fig. 6. Fig. 1b(H). Pessagno & Whalen 1982, pl. 3, fig. 11. Fig. 2. QCI, GSC loc. C-080611, GSC 128715.
Fig 3. GSC loc. C-080612, GSC 128716. Fig. 4. JP, MNA-10, MA13268.
55
Bagotum modestum Pessagno & Whalen 1982
Species code: BAG06
Synonymy:
1982 Bagotum modestum n. sp. – Pessagno & Whalen, p. 120,
pl. 3, fig. 7, 16, 17.
1984 Bagotum spp. – Whalen & Pessagno, pl. 2, fig. 9, 10.
1990 Bagotum modestum Pessagno & Whalen – Hori, Fig. 8.29.
1993 Bagotum modestum Pessagno & Whalen – Kashiwagi &
Yao, pl. 1, fig. 8.
1998 Bagotum modestum Pessagno & Whalen – Kashiwagi,
pl. 1, fig. 13
2002 Bagotum modestum Pessagno & Whalen – Whalen &
Carter, p. 116, pl. 10, figs. 9, 11, 12.
2003 Bagotum modestum Pessagno & Whalen – Goričan et al.,
p. 296, pl. 5, fig. 22.
2004 Lantus? sp. – Hori, pl. 1, fig. 62 only.
2004 Bagotum modestum Pessagno & Whalen – Matsuoka,
fig. 193.
Original description: Test broader distally than proximally,
having six post-abdominal chambers. Cephalis moderately
broad, hemispherical; remaining chambers trapezoidal in
cross section. Post-abdominal chambers all with linearly
arranged square to rectangular pore frames; chambers
gradually increasing in length; all but last one or two postabdominal chambers increasing moderately rapidly in
width as added; final one or two post-abdominal chambers
decreasing slightly in width as added. Dome-shaped cap
covering final post-abdominal chamber with irregular
polygonal pore frames.
56
Original remarks: Bagotum modestum, n. sp., differs from
all other new species of Bagotum described herein by showing linearly arranged pore frames on all post-abdominal
chambers.
Further remarks: Some specimens (pl. BAG06, figs. 5-9)
have a well-differentiated proximal part, separated from
the rest of the shell by a distinct constriction.
Measurements (µm):
Based on 9 specimens.
Length
200
250
200
223.6
Width (max.)
100
125
100
108.6
HT
Max.
Min.
Mean
Etymology: Modestus-a-um (Latin, adj.) = moderate,
orderly, restrained.
Type locality: Sample NSF 960, Franciscan Complex,
California.
Occurrence: Franciscan Complex, California; San Hipólito
Formation, Baja California Sur; Skrile Formation, Slovenia;
Musallah Formation, Oman; Japan .
Plate BAG06. Bagotum modestum Pessagno & Whalen. Magification x300. Fig. 1(H). Pessagno & Whalen 1982, pl. 3,
fig. 7. Fig. 2. Matsuoka 2004, fig. 193. Fig. 3. Whalen & Carter 2002, pl. 10, fig. 9. Fig. 4. Goričan et al. 2003, pl. 5, fig. 22.
Fig. 5. Hori 1990, fig. 8.29. Fig. 6. OM-00-252, 021730. Fig. 7. OM-00-118, 000611. Fig. 8. OM-00-252, 022015.
Fig. 9. OM-00-252, 021725.
57
Bagotum pseudoerraticum Kishida & Hisada 1985
Species code: BAG07
Synonymy:
? 1983 Bagotum (?) sp. – Hattori & Yoshimura, pl. 8, fig. 6.
1985 Bagotum pseudoerraticum n. sp. – Kishida & Hisada,
p. 113, pl. 2, figs. 1-5.
1992 Bagotum aff. pseudoerraticum Kishida & Hisada – Sashida,
pl. 1, figs. 13-15, 17.
1998 Bagotum (?) helmetense Pessagno & Whalen – Kashiwagi,
pl. 2, fig. 5, not fig. 4.
1998 Bagotum pseudoerraticum Kishida & Hisada – Kashiwagi,
pl. 2, figs. 8, 9.
2002 Bagotum sp. A – Hori & Wakita, pl. 3, fig. 3.
Original diagnosis: Test inflated, usually made up of 7
chambers. Final post-abdominal chamber with domeshaped cap. Outer layer of test with vermicular appearance.
Original description: Test inflated, usually with 4 postabdominal chambers, 5th or 6th chamber broadest. Cephalis hemispherical. Remaining chambers trapezoidal in
outline. Final post-abdominal chamber with dome-shaped
cap. Thorax, abdomen and first post-abdominal chambers
increasing slowly in height and increasing rapidly in width
as added; distal 2 chambers decreasing slightly in height
and width as added. Test with vermicular appearance due
to presence of irregular, often elongate polygonal frames in
outer layer. Inner layer of test with pores aligned longitudinally in rows.
Original remarks: Bagotum pseudoerraticum n. sp. differs
from Bagotum erraticum Pessagno and Whalen 1982, in
having vermicular appearance through the outer layer and
more inflated test. It is likely that the latter gave rise to the
former in early Early Jurassic.
Measurements (µm):
Based on 10 specimens.
Length
166
166
153
161
Width
92
105
88
97
HT
Max.
Min.
Av.
Type locality: Locality 230, Ueno-mura area, Kanto Mountains, Central Japan.
Occurrence: Kanto Mountains, Japan; Ghost Creek Formation, Queen Charlotte Islands; Musallah Formation,
Oman.
Genus: Beatricea Whalen & Carter 1998
Type species: Beatricea christovalensis Whalen & Carter 1998
Synonymy:
1998 Beatricea n. gen. – Whalen & Carter, p. 57.
Original description: Test small, circular to sub-rectangular
in outline with four long spines in the same plane 90°
apart. Cortical shell thick with planiform upper and lower
surfaces each with variably sized central cavity; sides of
cortical shell straight. Shell composed of numerous layers
of small, irregularly shaped pore frames lacking concentric
arrangement. Layers of pore frames much thicker on
margins of test. Spines usually triradiate in axial section,
rarely circular.
Original remarks: Beatricea n. gen. differs from Praeorbiculiformella Pessagno by always possessing four strong
primary spines at 90˚ and in lacking concentrically
58
arranged pore frames. Beatricea n. gen. differs from Sophia
n. gen. and Udalia n. gen. in possessing irregularly arranged
pore frames as well as a central cavity; Beatricea n. gen. further differs from Sophia n. gen. in lacking a central spicular
network.
Etymology: This genus is named for the ship C. P. R. Princess
Beatrice a sailing vessel in the Queen Charlotte Islands in
the early 1900s.
Included species:
ORB04 Beatricea? argescens (Cordey) 1998
PDC01 Beatricea? baroni (Cordey) 1998
SPI03 Beatricea christovalensis Whalen & Carter 1998
ORB07 Beatricea sanpabloensis (Whalen & Carter) 2002
CRU18 Beatricea? sp. A
Plate BAG07. Bagotum pseudoerraticum Kishida & Hisada. Magnification x300. Fig. 1(H). Kishida & Hisada 1985,
pl. 2, fig. 1. Fig. 2. QCI, GSC loc. C-305388, GSC 128717. Fig. 3. QCI, GSC loc. C-305388, GSC 128718. Fig. 4. QCI, GSC
loc. C-304281, GSC 128719. Fig. 5. OM-00-254, 022133.
59
Beatricea? argescens (Cordey) 1998
Species code: ORB04
Synonymy:
1988 Orbiculiforma sp. A – Carter et al., p. 45, pl. 1, fig. 9.
1989 Emiluvia ? spp. – Hattori, pl. 2, fig. J.
1996 Orbiculiforma sp. A of Carter in Carter et al. – Hori et al.,
pl. 1, fig. 19.
1998 Orbiculiforma argescens n. sp. – Cordey, p. 94, pl. 21,
figs. 6, 9, 11.
Original diagnosis: Orbiculiforma possessing four strong
spines.
Original description: Thick test circular to subsquare in
outline with four arms (one at each corner). Lower and upper surfaces planar. Test sides vertical to slightly concave.
Cortical cavity deep and with smaller pores than those of
external crown. Test with polygonal pore frames (pentagonal to hexagonal) with stout nodes at vertices. Four coplanar spines orthogonally disposed and circular in cross-section: sometimes elongated proximal pores, extending up to
two-third length of spine.
Original remarks: This form resembles to O. quadrata
Pessagno by its subsquare shape and four spines, but differs
from it by its more massive skeleton and pore-frames and
much longer, massive radial spines.
Further remarks: This species differs from Beatricea
christovalensis Whalen & Carter by having a more squareshaped test and shorter spines. See also remarks under
B.? baroni (Cordey) and Beatricea? sp. A.
Etymology: From Greek argo- shiny.
Type locality: Locality GSC C-300407, Bridge River Complex, Lake Carpenter, British Columbia.
Occurrence: Bridge River Complex, British Columbia; Fannin Formation, Queen Charlotte Islands; Fernie Formation, NE British Columbia; Japan; Newcastle Group, New
Zealand.
Beatricea? baroni (Cordey) 1998
Species code: PDC01
Synonymy:
1998 Pseudocrucella? baroni n. sp. – Cordey, p. 70, pl. 20, fig. 5-7.
2002 Pseudocrucella? baroni Cordey – Whalen & Carter, p. 105,
pl. 1, figs. 5, 6, 9, 10, 13, 14.
Original diagnosis: Pseudocrucella? with wide central zone,
broad, short latticed arms, and long distal spines.
Original description: Test almost square, built with a latticed zone whose corners are prolonged with massive triradiate spines. Square zone corresponds to the development
of four latticed arms externally constructed with cortical
shell; the latter comprising several longitudinal beams
(from three to four per lateral face), connected by transverse bars. Nodes observed at the intersection of bars and
beams. Cortical shell absent on central part of test, revealing the medullary shell formed by small orthogonal pore
frames. Latticed faces of arms concave. Long triradiate
spines extend from arms.
Original remarks: Pseudocrucella? baroni differs from all
other species of Pseudocrucella by a broad development of
central zone, short latticed parts on arms and long distal
spines.
This morphotype is questionably assigned to the genus
Pseudocrucella because the latticed zones on the arms of
60
Pseudocrucella are usually longer. The concavity of lateral
faces suggests that this form cannot be assigned to the
genus Higumastra.
Further remarks: This species differs from Beatricea
argescens (Cordey) because the outer edges of the test
between spines are concave, and the spines are longer and
more developed.
Measurements (µm):
Based on 2 specimens.
Total length of ray
Maximum width of rays
Diameter of central cavity
Min.
80
45
45
Max.
110
75
55
Av.
95
50
50
Etymology: Arbitrary combination of letters (ICZN, 1985,
art. 11b(iii), p. 20).
Type locality: Locality GSC C-300407, Bridge River Complex, Lake Carpenter, British Columbia.
Occurrence: Bridge River and Hozameen complexes, British Columbia; San Hipólito Formation, Baja California
Sur.
Plate ORB04. Beatricea? argescens (Cordey). Magnification x200. Fig. 1(H). Cordey 1998, pl. 21, fig. 6. Fig. 2. QCI,
GSC loc. C-C-080611, GSC 111799. Fig. 3. Carter et al. 1988, pl. 1, fig. 9. Fig. 4. QCI, GSC loc. C-305417, GSC 111800.
Fig. 5. QCI, GSC loc. C-080612, GSC 111801. Fig. 6. NBC, GSC loc. C-305208, GSC 111802.
Plate PDC01. Beatricea? baroni (Cordey). Magnification x200. Fig. 1(H). Cordey 1998, pl. 20, fig. 5. Fig. 2. Whalen &
Carter 2002, pl. 1, fig. 5. Fig. 3. Whalen & Carter 2002, pl. 1, figs. 6, 10.
61
Beatricea christovalensis Whalen & Carter 1998
Species code: SPI03
Synonymy:
1998 Beatricea christovalensis n. sp. – Whalen & Carter, p. 57,
pl. 11, figs. 13, 14, 16-20, 22.
2002 Beatricea christovalensis Whalen & Carter – Suzuki et al.,
p. 175, figs. 6 G-H.
2002 Beatricea christovalensis Whalen & Carter – Tekin, p. 185,
pl. 3, fig. 1.
Original description: Test small, sub-rectangular in outline with four, long prominent spines. Test thick with planiform upper and lower surfaces and straight sides. Cortical shell with irregularly shaped tetragonal and polygonal
pore frames with very small nodes at pore frames vertices.
Central cavity variable in size but usually about one-half
diameter of cortical shell. Central area of test thinner than
margins and often missing. Spines usually triradiate (rarely
circular) in axial section with narrow longitudinal ridges
and broad grooves sometimes showing torsion.
Original remarks: This is the first species of Beatricea
n. gen. described from the Lower Jurassic of Queen Charlotte Islands. Several morphological features of this species are quite variable: the size and shape of cortical shell,
the width of the central cavity, and the length of primary
spines. For the present we have included all Hettangian
and Sinemurian specimens in one widely variable species,
Beatricea christovalensis n. sp. We recognize a possible relationship between B. christovalensis and Spumellarian indet. B but owing to our incomplete knowledge of the inner
structure of both forms, their differences are not addressed
in this paper.
Measurements (µm):
Based on 12 specimens.
Diameter of
corticall shell
225
225
97
150
Diameter of
central area
124
124
38
67
Length of
primary spines (max.)
237
329
84
193
HT
Max.
Min.
Mean
Etymology: This species is named for the San Christoval
Range in Queen Charlotte Islands.
Type locality: Sample 89-CNA-KUG-1A, Sandilands Formation, Kunga Island, north side; Queen Charlotte Islands,
British Columbia.
Occurrence: Sandilands, Ghost Creek and Fannin formations, Queen Charlotte Islands; Hocaköy Radiolarite, Turkey; Pucara Group, Peru.
Beatricea sanpabloensis (Whalen & Carter) 2002
Species code: ORB07
Synonymy:
1984 Orbiculiforma sp. – Whalen & Pessagno, pl. 1, fig.18.
2002 Orbiculiformella sanpabloensis n. sp. – Whalen & Carter,
p. 109, pl. 1, figs. 1-2.
Original description: Test small, circular in outline with
vertical sides and upper and lower surfaces of rim rounded.
Test thick in proportion to diameter. Four to six primary
peripheral spines, medium length, triradiate in axial section; very small subsidiary spines or spinules located between principal peripheral spines. Central cavity shallow
almost one half diameter of test, with raised center. Meshwork primarily composed of large, irregularly shaped polygonal pore frames; meshwork distinctly finer in central
cavity area.
Original remarks: The rounded top, bottom and rim surfaces of Orbiculiformella sanpabloensis n. sp. distinguish it
from O. trispinula (Carter 1988) while the much shorter
spines and larger central cavity distinguish it from O. trispinosa (Yeh 1987).
Measurements (µm):
Based on 7 specimens.
Width (Max.)
158
225
158
194
Width of central cavity (Max.)
90
135
75
103
HT
Max.
Min.
Mean
Etymology: This species is named for Punta San Pablo
located to the northwest of the type area.
Type locality: Sample SH-412-14, San Hipolito Formation,
Baja California Sur, Mexico.
Occurrence: San Hipolito Formation, Baja California Sur;
Fannin Formation, Queen Charlotte Islands.
Beatricea? sp. A
Species code: CRU18
Synomnymy:
2002 Crucella? sp. A – Whalen & Carter, p. 107, pl. 1, figs. 4, 8, 12.
Remarks: The test outline of Beatricea? sp. A appears to be
intermediate between Beatricea? argescens (Cordey) and
Beatricea? baroni (Cordey).
Occurrence: San Hipólito Formation, Baja California Sur.
62
Plate SPI03. Beatricea christovalensis Whalen & Carter. Magnification: Fig. 1 x150 (scale bar A), Figs. 2, 3 x200 (scale
bar B). Fig. 1(H). Carter et al. 1998, pl. 11, fig. 14. Fig. 2. QCI, GSC loc. C-304566, GSC 128886. Fig. 3. QCI, GSC loc.
C-304566, GSC 128887.
Plate ORB07. Beatricea sanpabloensis (Whalen & Carter). Magnification x200. Fig. 1(H). Whalen & Carter 2002,
pl. 1, figs. 1-2. Fig. 2. QCI, GSC loc. C-080611, GSC 128850.
Plate CRU18. Beatricea? sp. A. Magnification x200. Fig. 1. Whalen & Carter 2002, pl. 1, fig. 4.
63
Genus: Bernoullius Baumgartner1984
Type species: Eucyrtis (?) dicera Baumgartner, in Baumgartner et al. 1980
Synonymy:
1984 Bernoullius n. gen. – Baumgartner, p. 759.
Original description: Spongodiscid spumellarian with
distinct bilateral symmetry: A delicate, finely spongy main
body of flattened egg-shape carries on the narrow end two
symmetric, strongly developed, usually triradiate lateral
spines and sometimes one central spine.
Original remarks: Because of the clear bilateral symmetry,
the spines were interpreted as cephalic horns of a nassellarian by Baumgartner in Baumgartner et al. 1980. Well
preserved specimens from DSDP Site 534A show that the
spines are attached to a finely spongy body lacking any
resemblance to nassellarian morphology. For most specimens, the spongy body is not as poorly preserved as spongy
round mass at the base of the spines. Kozur & Mostler
(1979, pl. 21, fig. 2) illustrated a Triassic form which possibly belongs to this genus.
Etymology: Dedicated to Daniel Bernoulli, Zurich, Switzerland, in honour of his contribution to the understanding of ancient passive continental margins in the AlpineMediterranean realm.
Included species:
3222 Bernoullius delnortensis Pessagno, Blome & Hull 1993
BER01 Bernoullius saccideon (Carter) 1988
Bernoullius delnortensis Pessagno, Blome & Hull 1993
Species code: 3222
Synonymy:
1987 Bernoullius sp. A – Goričan, p. 181, pl. 1, fig. 17.
1993 Bernoullius delnortensis Pessagno, Blome & Hull n. sp.
– Pessagno et al., p. 120, pl. 1, figs. 4, 15, 26.
1994 Bernoullius rectispinus Kito, De Wever, Danelian & Cordey
s.l. – Goričan, p. 63, pl. 8, figs. 7, 8, ?9, 11, 12 only.
1995a Bernoullius rectispinus delnortensis Pessagno, Blome &
Hull – Baumgartner et al., p. 126, pl. 3222, figs. 1-4.
1997 Bernoullius delnortensis Pessagno, Blome & Hull – Hull,
p. 16, pl. 1, fig. 2.
2004 Bernoullius delnortensis Pessagno, Blome & Hull
– Matsuoka, fig. 2.
Original description: Test relatively slender, flaring slightly
laterally away from spines. Primary spines straight, rather
short and massive, triradiate in axial section with three
longitudinal ridges alternating with three longitudinal
grooves. Longitudinal grooves, narrow, deeply incised,
gradually decreasing in width in a distal direction. Ridges
wide proximally, becoming progressively narrower in a
distal direction.
Original remarks: This form greatly resembles Bernoullius
sp. A of Goričan (1987). It possesses straight, short, subequal spines with parallel sided, deeply incised grooves
separating wide, longitudinal ridges which wedge out distally. Bernoullius sp. A of Gorican, however, possesses short
spines which are nearly equal in length and are somewhat
shorter than those of B. delnortensis. B. delnortensis differs from B. sp. A (herein) by having considerably shorter,
64
wider, and more massive primary spines. B. delnortensis differs from B. cristatus Baumgartner (1984) by having spines
which are straight and lack curved tips.
Further remarks: By Baumgartner et al. (1995a): This subspecies differs from Bernoullius rectispinus rectispinus by
having smaller size. The species also differs from Bernoullius dicera and B. cristatus by having straight spines.
Measurements (µm):
Based on 4 specimens (see Pessagno et al., 1993 for explanation of system of measurements for this species).
AA':
Sx:
S'x':
BB':
SS':
HT
210
165
135
-
av.
206
130
136
135
187
min.
195
105
123
105
180
max.
225
165
150
180
195
Etymology: This species is named for Del Norte County,
California.
Type locality: Volcanopelagic strata above Josephine ophiolite, Smith River subterrane, Klamath Mountains, northwestern California.
Occurrence: Worldwide.
Plate 3222. Bernoullius delnortensis Pessagno, Blome & Hull. Magnification x200. Fig. 1(H). Pessagno et al. 1993, pl. 1,
fig. 4. Fig. 2. OM, BR871-R02-11. Fig. 3. OM, BR871-R02-15. Fig. 4. OM, BR871-R02-16. Fig. 5. Matsuoka 2004, fig. 2.
65
Bernoullius saccideon (Carter) 1988
Species code: BER01
Synonymy:
1988 Spongiostoma saccideon Carter n. sp. – Carter et al., p. 46,
pl. 12, figs. 4, 7, 10.
? 1988 Spongiostoma sp. A – Carter et al., p. 47, pl. 12, figs. 8, 9.
Original diagnosis: Test subcircular in outline, composed
of two spongy concentric layers which gape open. Short
hinge on one edge marked by two strong triradiate spines;
periphery with a few very fine secondary spines.
Original description: Test as for genus; subcircular in outline. Hinge normally short and straight with a well defined,
triradiate primary spine on either end. A few short and very
fine, secondary spines radiating from the circular periphery have been noted on some tests.
Original remarks: Compared to Spongiostoma sp. A under
that species. Common to very abundant in middle/upper
Toarcian samples.
Measurements (µm):
Measurements are treated in a very preliminary manner as
many of these specimens appear incomplete.
Based on 11 specimens.
Maximum diameter of test
Length of hinge
(between centres at margin)
Length of longest spine
HT
121
Av.
207
Max.
250
Min.
160
109
100
120
76
109
124
165
80
Etymology: Latin, saccus (n.), bag or pouch; saccideon =
of sac-like appearance.
Type locality: GSC locality C-080583, Phantom Creek Formation, Graham Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Phantom Creek Formation, Queen Charlotte
Islands.
Genus: Bipedis De Wever 1982a
Type species: Bipedis calvabovis De Wever 1982a
Synonymy:
1982a Bipedis n. gen – De Wever, p. 192.
Original description: Form with two segments, with a
strong apical horn and two or four feet. Cephalic skeleton
with spines A, V, MB, Ll, Lr, ll and lr; spine D is absent.
Apical horn is an extension of A spine. A cephalic opening
exists at the prolongation of V spine. Two feet correspond
to Ll and Lr, sometimes two smaller feet occur as external
extensions of ll and lr.
Original remarks: When only two feet are present, they are
not separated from each other by an angle of 180°, due to
the respective locations of Ll and Lr spines on the cephalic
skeleton. For practical reasons, I have placed this genus in
the Pylentonemidae, although the absence of the D spine
makes this placement uncertain.
Etymology: From the Latin bi = two, and pes, pedis = foot
(form with two feet).
Included species:
BPD13 Bipedis calvabovis De Wever 1982a
BPD05 Bipedis diadema Whalen & Carter 1998
BPD14 Bipedis fannini Carter 1988
BPD15 Bipedis japonicus Hori n. sp.
BPD16 Bipedis yaoi Hori n. sp.
Bipedis calvabovis De Wever 1982a
Species code: BPD13
Synonymy:
1982a Bipedis calvabovis n. sp. – De Wever, p. 193, pl. 2,
figs. 7-11.
1982b Bipedis calvabovis De Wever – De Wever, p. 337, pl. 52,
figs. 5-9.
1982 Bipedis calvabovis De Wever – De Wever & Origlia-Devos,
pl. 1, fig. J, ? figs. K, L.
2002 Bipedis sp. aff. B. calvabovis De Wever – Tekin, p. 192,
pl. 5, fig. 10.
Original description: Bipedis with a strong apical horn
and two lanceolate feet, triradiate in cross- section along
their length. Cephalis smooth and imperforate proximally,
smooth or longitudinally ribbed, perforate or not, distally.
Cephalis with a small lateral opening protected by a perfo66
rate hood at the end of the V spine (Pl. 2, fig. 7, 8, 10, 11).
The change in outline between the cephalis and thorax is
sometimes very clear (Pl. 2, fig. 8) and sometimes not (Pl.
2, fig. 10). Cephalis and thorax cone-shaped. Thorax with
small irregular pores in different specimens: large (Pl. 2, fig.
7) or small (Pl. 2, fig. 10). Cephalis and thorax more robust
in specimens with smaller pores than in specimens with
large pores. This could result from a more or less important
(ontogenetical ?) development of an external silica layer as
is the case in other forms.
Original remarks: A form close to B. calvabovis n. sp. was
found but it has four feet instead of two.
Plate BER01. Bernoullius saccideon (Carter). Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 12, figs. 4, 10.
Fig. 2. Carter et al. 1988, pl. 12, fig. 7.
Measurements (µm):
Based on 10 specimens.
Length apical horn
Cephalo-thorax length
Cephalo-thorax width
Length of feet
bovis = ox. For similarity with skull of bovids, when the
apical horn is turned downwards.
HT
100
100
100
153
Av.
90
103
100
118
Min.
75
90
90
100
Max.
100
125
109
153
Etymology: From the Latin calva, –ae =skull; and bos,
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon and Hocaköy Radiolarite, Turkey; Drimos Formation, Greece; Haliw (Aqil) Formation, Oman.
Plate BPD13. Bipedis calvabovis De Wever. Magnification x200. Fig. 1(H). De Wever 1982a, pl. 2, fig. 7. Fig. 2. OM,
Haliw-038-R08-05.
67
Bipedis diadema Whalen & Carter 1998
Species code: BPD05
Synonymy:
1998 Bipedis diadema n. sp. – Whalen & Carter, p. 76, pl. 21, figs.
7-10, 13, 17; pl. 22, fig. 1; pl. 27, figs. 11, 12.
Original description: Test hemi-elliptical in outline with
small hemispherical cephalis and medium-sized horn;
horn usually triradiate in axial section proximally with
narrow, rounded longitudinal ridges and broad grooves
becoming rounded in axial section distally. Cephalis
mostly smooth, imperforate, covered with thick, irregular
layer of microgranular silica; some relict pores observed on
cephalis where layer of microgranular silica is thinner. Both
cephalis and thorax compressed in plane of feet. Thorax
with mostly small, irregularly shaped pore frames with
slight development of orientation transverse to long axis of
the test. Test with two medium-sized feet, triradiate in axial
section with narrow rounded ridges and broad grooves;
feet curved slightly inward towards center of test; mouth
elliptical in outline with prominent, broad imperforate
band.
Original remarks: Bipedis diadema n. sp., is distinguished
from all other species of Bipedis by the very wide imperforate band which rims the mouth, the hemi-elliptical outline, and the compressed thorax and cephalis.
Measurements (µm):
Based on 10 specimens.
Length
(excluding horn)
105
105
90
101
Width
(max.)
90
109
75
94
Length of feet (max.)
75
75
45
65
HT
Max.
Min.
Mean
Etymology: Diadema, atis (Latin; neuter) = a royal headband, diadem.
Type locality: Sample QC 675. Sandilands Formation,
Kunga Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands.
Bipedis fannini Carter 1988
Species code: BPD14
Synonymy:
1988 Bipedis fannini Carter n. sp. – Carter et al., p. 61, pl. 2,
figs. 7, 8.
Original diagnosis: Dicyrtid with small cephalis, globose
thorax with roughened surface, long triradiate horn and
two downward curving terminal feet.
Original description: Bilaterally symmetrical dicyrtid
test. Cephalis medium-sized, spherical, sparsely perforate
basally with long, tapering apical horn. Horn triradiate
with deep grooves on basal half. Thorax large and globose
with constricted circular aperture and two strong, triradiate
downward curving terminal feet. Thorax with roughned
surface composed of low nodes or tubercles surrounded by
small circular to elliptical pores.
Original remarks: This species appears to be entirely new
and differs significantly from all other known species of Bipedis. Rare at type locality, abundant in older Pliensbachian
samples.
68
Measurements (µm):
Based on 9 specimens.
Height of cephalis and thorax
Maximum width of thorax
Length of apical horn
Length of feet
HT
132
139
60
-
Av.
148
121
62
105
Max.
170
140
70
120
Min.
132
120
52
75
Etymology: Named in honour of John Fannin, curator of
the provincial museum in Victoria, British Columbia, in
the late 1800's.
Type locality: GSC locality C-080577, Fannin Formation,
Creek locality, Maude Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands and Williston Lake, north-east British
Columbia.
Plate BPD05. Bipedis diadema Whalen & Carter. Magnification x300. Fig. 1(H). Carter et al. 1998, pl. 21, fig. 7.
Fig. 2. Carter et al. 1998, pl. 21, fig. 8. Fig. 3. QCI, GSC loc. C-080612, GSC 128711. Fig. 4. Carter et al. 1998, pl. 21,
fig. 10. Fig. 5. Carter et al. 1998, pl. 21, fig. 9. Fig. 6. QCI, GSC loc. C-080612, GSC 128720.
Plate BPD14. Bipedis fannini Carter. Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 2, figs. 7-8. Fig. 2. QCI, GSC
loc. C-080611, GSC 128721. Fig. 3. QCI, GSC loc. C-080613, GSC 128889. Fig. 4. QCI, GSC loc. C-304566, GSC 128890.
Fig. 5. QCI, GSC loc. C-080611, GSC 128891. Fig. 6. NBC, GSC loc. C-305813, GSC 128892.
69
Bipedis japonicus Hori n. sp.
Species code: BPD15
Synonymy:
1982 Nassellaria gen. and sp. indet. B – Yao et al., p. 41, pl. 2,
fig.13.
1986 Bipedis sp. – Hori, p. 52, fig. 6-12.
1990 Bipedis sp. A – Hori, p. 581, fig. 8-12.
1993 Bipedis sp. – Fujii et al., p. 87, pl. 2, fig. 4.
1994 Bipedis sp. A – Matsuoka et al., pl. 5, fig. 14.
1997 Bipedis horiae n. sp. – Sugiyama, p. 145, fig. 39.10,
not fig. 28.7.
Type designation: Holotype specimen no.Kb05-18 (pl.
BPD15, fig. 1), sample Kb05, Katsuyama Section.
Description: Test dicyrtid with large dome-shaped cephalis
and a long massive horn, triradiate in axial section. Cephalis
smooth, imperforate at base of horn. Thorax large, relatively
inflated, subspherical with fairly regular polygonal pore
frames. Two wing-like feet attached at base of thorax;
feet inwardly curving, triradiate in axial section. Narrow
longitudinal ridges and broad grooves visible along side of
thorax for most part. Large aperture at base of thorax.
Remarks: Bipedis sp. A of Hori (1990) was placed in synonymy with Bipedis horiae by Sugiyama (1997), but these
two species are quite different in the shape of shell and feet.
Bipedis sp. A of Hori (1990) is described herein as Bipedis
japonicus n. sp., which is distinguished from Bipedis horiae
Sugiyama by having very long stout feet and a strong horn.
B. japonicus n. sp. differs from Bipedis rotundus Whalen and
Carter by having long well-developed feet and horn, and a
wider thorax with more regularly arranged pore frames.
Measurements (µm):
Based on 5 specimens.
Height of cephalis and thorax
Maximum width of thorax
Length of apical horn
Length of feet
HT
147
146
107
134
Av. Max. Min.
130 147 120
144 153 133
84
99 113
140 153 133
Etymology: This species of Bipedis occurs mainly in Lower
Jurassic strata of Japan.
Type locality: Kb05 (UFI3+183cm) UF (Katsuyama)
section, Inuyama, Mino terrane, Japan.
Occurrence: Mino terrane, Japan.
Bipedis yaoi Hori n. sp.
Species code: BPD16
Synonymy:
1990 Bipedis sp. B – Hori, p. 581, fig. 8-13.
1990 Bipedis sp. A – Yao, p. 341, pl. 2, fig. 10.
1994 Bipedis sp. B – Matsuoka et al., p. 53, pl. 5, fig. 21.
2001 Bipedis sp. – Gawlick et al., pl. 6, fig. 9.
2005 Bipedis sp. – Hori, pl. 9, fig. 27.
Type designation: Holotype specimen no. IYII24-28 (pl.
BPD16, fig. 1), sample IYII24, IY Section.
Description: Cephalis small and spherical with a long horn;
horn triradiate or solid circular in axial section. Thorax
large and spherical with fairly regularly arranged polygonal
pore frames, a circular aperture and two long feet. Feet
asymmetrical, straight or sometimes curving outward
distally. Feet thin, blade-like in axial section proximally
terminating with a solid spine.
Remarks: This species is distinguished from all other species of Bipedis by having a large almost spherical thorax. It
is similar to Bipedis hannai Whalen and Carter 1998, but
differs by having longer feet, a longer stout horn and larger
70
shell. B. yaoi Hori n. sp. differs from Bipedis japonicus Hori
n. sp. and B. horiae Sugiyama 1997 by having thin feet that
are circular in cross section terminally, and a small circular
aperture.
Measurements (µm):
Based on 12 specimens.
Height of cephalis and thorax
Maximum width of thorax
Length of apical horn
Length of feet
Diameter of aperture
HT
147
150
102
131
77
Av. Max. Min.
143 157 130
145 152 132
96 105
72
94
112 139
79
89
68
Etymology: This species is named in honor of Akira Yao for
his pioneering works on Lower Jurassic radiolarian fossils.
Type locality: IYII24 (IY Section), bedded chert sequences,
Inuyama, Mino terrane, southwest Japan.
Occurrence:
Mino terrane, Japan; Dürrnberg Formation, Austria.
Plate BPD15. Bipedis japonicus Hori n. sp. Magnification x200. Fig. 1(H). JP, Kb05-18, RH(1) 1834.
Fig. 2. JP, Kb05-12, RH(1) 1832.
Plate BPD16. Bipedis yaoi Hori n. sp. Magnification x200. Fig. 1(H). Hori 1990, fig. 8.13. Fig. 2. JP, IYII24-29, RH(1) 1683.
71
Genus: Bistarkum Yeh 1987b
Type species: Bistarkum rigidium Yeh 1987b
Synonymy:
Original remarks: The name Bistarkum is introduced to
avoid assigning species to Amphibrachium whose definition
is obscured by poor descriptions and illustrations of its type
species.
Original description: Test medium to large in size, with
two rays linearly aligned. Rays nearly equal in length, often
terminated with expanded tips. Tips subcircular; elliptical
in outline, or bifurcated. Meshwork of test comprised
of sponge layers or regular (i.e., triangular) or irregular
polygonal pore frames. Cross section of rays ellipsoidal,
rectangular; or subrectangular in outline. Rays with or
without spines at distal surface of tips or along sides of
rays.
Etymology: Bistarkum is a name formed by an arbitrary
combination of letters (ICZN, 1985, Appendix D, Pt. I.4
Recommendation 40, p.201).
1971 Amphibrachium Haeckel emend. – Pessagno, p. 20.
1980 Amphibrachium Hertwig emend. – Baumgartner, p. 300.
1987b Bistarkum n. gen. – Yeh, p. 42.
Included species:
BIS04 Bistarkum mangartense Goričan, Šmuc &
Baumgartner 2003
BIS02 Bistarkum phantomense (Carter) 1988
BIS01 Bistarkum rigidium Yeh 1987b
BIS03 Bistarkum saginatum Yeh 1987b
Bistarkum mangartense Goričan, Šmuc & Baumgartner 2003
Species code: BIS04
Synonymy:
1997 Bistarkum sp. A – Yao, pl. 6, fig. 265.
? 1997 Bistarkum sp. C0 – Yao, pl. 6, fig. 269.
2003 Bistarkum mangartense n. sp. – Goričan, Šmuc &
Baumgartner, p. 293, pl. 2, figs. 7-10.
2004 Bistarkum mangartense Goričan, Šmuc & Baumgartner
– Matsuoka, fig. 27.
Original description: Test ellipsoidal; cylindrical through
most of its length and terminating with hemispherical ray
tips. Rays circular in cross-section, central area not differentiated externally. Spongy meshwork very fine. Short
spines, circular in cross-section, occur on the surface of
well-preserved specimens. Spines more numerous at ray
tips than in the middle part of the shell.
Original remarks: Bistarkum mangartense n. sp. differs
from other Bistarkum species by its ellipsoidal shape without enlargement at ray tips. From Bistarkum phantomense
72
(Carter) it differs also by being circular in cross-section.
Bistarkum sp. C0 of Yao (1997) is questionably assigned to
Bistarkum mangartense, because the shell seems more flattened, i.e. elliptical and not circular in cross-section.
Measurements (µm):
Based on 21 specimens.
Total length (excluding spines) L
Width (across the center) W
W/L ratio
HT
153
88
0.57
Max.
225
123
0.70
Min. Av.
126 190
74 100
0.36 0.53
Etymology: Named after type locality.
Type locality: Sample MM 21.70, Skrile Formation, Mt.
Mangart in the Julian Alps, Slovenia.
Occurrence: Skrile Formation, Slovenia; Mino Terrane,
Japan.
Plate BIS04. Bistarkum mangartense Goričan, Šmuc & Baumgartner. Magnification x250. Fig. 1. Matsuoka 2004,
fig. 27. Fig. 2. SI, MM6.76, 000532. Fig. 3. Goričan et al. 2003, pl. 2, fig. 10. Fig. 4. Goričan et al. 2003, pl. 2, fig. 8.
Fig. 5(H). Goričan et al. 2003, pl. 2, fig. 7.
73
Bistarkum phantomense (Carter) 1988
Species code: BIS02
Synonymy:
1988 Amphibrachium (?) phantomensis n. sp. – Carter et al.,
p. 39, pl. 12, fig. 1; figure 9.
1997 Bistarkum sp. C – Yao, pl. 6, fig. 267.
2003 Bistarkum phantomense (Carter) – Goričan et al., p. 295,
pl. 2, figs. 13-16.
Original diagnosis: Two-rayed form with very fine, spongy
meshwork. One or both rays variably bilobed.
Further remarks: We include also specimens with both rays
almost symetrically developed. An indentation in the lobe
of the ray tips can be very indistinct or absent. The rays are
very short so that the shell sometimes appears only slightly
constricted in the middle part. A weakly differentiated
circular central area is observed in some specimens.
Bistarkum phantomense differs from Bistarkum saginatum
Yeh by having wider rays.
Original description: Two-rayed patulibracchiid with
very fine, layered spongy meshwork. Tips of rays widely
expanded and rounded; may form either one large lobe
or bifurcate to form two smaller lobes. On some bilobed
specimens lobes become elongate, giving the test an almost
three-rayed appearance. Rays equal in length, very short
and wide, no defined central area. Spines on ray tips vary
from a single central spine to numerous fine ones.
Measurements (µm):
Based on 11 specimens.
Original remarks: Genus is tentatively placed with
Amphibrachium and queried because of the bilobed nature
of ray tips. It is conceivable, however, that it is an extreme
variant of Paronaella spongiosa n. sp. It is postulated that
in the early late Toarcian a variant form of Paronaella
had appeared (see Pl. 11, fig. 7); by the late Toarcian the
figured two-layered bilobed form (A. (?) phantomensis)
had evolved through enlargement of the primary ray
and reduction of the secondary and tertiary rays to form
a single bilobed ray.
Etymology: Named for Phantom Creek, south of type
locality.
Length of ray
Width of ray
AX
BX
cc'
dd'
HT
130
135
166
209
Av.
137
140
185
232
Max.
170
170
280
280
Min.
125
125
140
180
Type locality: GSC locality C-080597, Phantom Creek
Formation, Graham Island.
Occurrence: Phantom Creek Formation, Queen Charlotte
Islands, Guwayza Formation, Oman; Skrile Formation,
Slovenia; Japan.
Bistarkum rigidium Yeh 1987b
Species code: BIS01
Synonymy:
1987b Bistarkum rigidium n. sp. – Yeh, p. 43, pl. 1, figs. 5, 17,
pl. 22, figs. 1, 3, 7, 11.
1987b Gorgansium rigidum n. sp. – Yeh, pl. 30, fig. 13.
1987b Bistarkum bifurcum n. sp. – Yeh, p. 43, pl. 1, fig. 10; pl. 21,
fig. 5; pl. 22, figs. 5-6.
1987b Bistarkum sp. cf. G. bifurcum n. sp. – Yeh, p. 43, pl. 9,
fig. 13; pl. 21, fig. 11.
2004 Bistarkum rigidium Yeh – Matsuoka, fig. 22.
Original description: Rays about equal in length, medium
of width, one ray usually slightly wider than the other;
both rays terminating in large ellipsoidal tips. Width of
tips about equal to length of ray shafts. Test comprised of
small irregularly arranged polygonal pore frames without
prominent small nodes at vertices. Several spines with
circular cross-section and variable length occurring at
distal surface of tips and sides of rays.
Original remarks: Bistarkum rigidium Yeh, n. sp., differs
from G. bifurcum n. sp., by lacking bifurcate tips and by
having a wider test.
Further remarks: B. rigidium Yeh and B. bifurcum Yeh are
synonymized because a morphological continuum of forms
74
with ellipsoidal to distictly bifurcating ray tips generally occurs in a single sample. Specimens with only moderately
developed bifurcation (determined as Bistarkum cf. bifurcum by Yeh, 1987b) are the most common.
Measurements (µm):
System of measurement shown in text-figure 7 of Yeh
(1987b). Ten specimens measured.
HT
Mean
Max.
Min.
LR
230
225
230
184
WR
90
77
90
64
WT
150
158
172
150
LT
80
75
80
54
Etymology: Rigidius-a-um (Latin, adj.) = rigid.
Type locality: Sample OR-589D, Warm Springs member,
Snowshoe Formation, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs m+ember of the Snowshoe Formation, Oregon;
Mino Terrane, Japan; Tawi Sadh Member of the Guwayza
Formation, Oman.
Plate BIS02. Bistarkum phantomense (Carter). Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 12, fig. 1.
Fig. 2. OM, BR871-R06-24. Fig. 3. SI, MM5.00, 010111. Fig. 4. SI, MM11.76, 010128. Fig. 5. Goričan et al. 2003, pl. 2,
fig. 16. Fig. 6. Goričan et al. 2003, pl. 2, fig. 14a. Fig. 7. Goričan et al. 2003, pl. 2, fig. 15. Fig. 8a. Goričan et al. 2003, pl. 2,
fig. 13. Fig. 8b. SI, MM11.76, 010127.
Plate BIS01. Bistarkum rigidium Yeh. Magnification x100. Fig. 1(H). Yeh 1987b, pl. 22, fig. 11. Fig. 2. Matsuoka 2004,
fig. 22. Fig. 3. OM, BR1123-R05-03. Fig. 4. OM, BR1123-R05-02.
75
Bistarkum saginatum Yeh 1987b
Species code: BIS03
Synonymy:
? 1885 Heliodiscus inchoatus n. sp. – Rüst, p. 293 (23), pl. 29 (4),
fig. 13.
1987b Bistarkum saginatum n. sp. – Yeh, p. 44, pl. 22, figs. 13, 16.
1988 ?Heliodiscus inchoatus Rüst – Carter et al., p. 38, pl. 12,
figs. 2, 5.
2004 Bistarkum saginatum Yeh – Matsuoka, fig. 24.
Original description: Rays extremely short, wide, subellipsoidal in cross-section, with large subtriangular to hemispherical tips. Test comprised of nearly uniformly sized irregular polygonal pore frames. Pore frames without prominent nodes at vertices. Short spines occurring on distal
surface of tips and sides of rays.
Original remarks: This species differs from other Bistarkum spp. in this report by having extremely large tips and
by having short, massive rays.
Further remarks: ?Heliodiscus inchoatus Rüst, illustrated
by Carter et al. (1988) appears to be the same species but
with more expanded ray tips.
Measurements (µm):
System of measurements shown in text-figure 7 of Yeh
(1987b). Ten specimens measured.
HT
Mean
Max.
Min.
LR
183
181
187
170
WR
70
72
75
70
WT
158
154
161
150
LT
107
105
107
98
Etymology: Saginatus-a-um (Latin, adj.) = flattened.
Type locality: OR-589D, Warm Springs Member, Snowshoe
Formation, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, east-central Oregon; Phantom Creek Formation, Graham Island,
Queen Charlotte Islands; Mino Terrane, Japan; Tawi Sadh
Member of the Guwayza Formation, Oman.
Genus: Broctus Pessagno & Whalen 1982
Type species: Broctus selwynensis Pessagno & Whalen 1982
Synonymy:
1982 Broctus n. gen. – Pessagno & Whalen, p. 120.
Original description: Test as with family: spindle-shaped.
Final post-abdominal chamber terminating in narrow,
tubular structure (pl. 2, fig. 20) extending from aperture;
cephalis lacking horn. Pore frames regular to irregular,
tending to be more regular distally than proximally.
Original remarks: Broctus n. gen., differs from Bagotum
n. gen. by being spindle-shaped instead of ellipsoidal and
76
differs from Bagotum, Droltus, and Noritus by having a final
post-abdominal chamber terminating in a narrow tubular
structure.
Etymology: Broctus is a name formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, pt. VI,
Recommendation 40, p.113).
Included species:
BRO02 Broctus kuensis Pessagno & Whalen 1982
BRO03 Broctus ruesti Yeh 1987b
BRO01 Broctus selwynensis Pessagno & Whalen 1982
Plate BIS03. Bistarkum saginatum Yeh. Magnification x200. Fig. 1(H). Yeh 1987b, pl. 22, fig. 13. Fig. 2. Matsuoka
2004, fig. 24. Fig. 3a,b. Carter et al. 1988, pl. 12, figs. 2, 5. Fig. 4. OM, BR706-R02-13a. Fig. 5. OM, BR871-R01-01.
Fig. 6. OM, BR871-R01-02. Fig. 7. OM, BR871-R02-06. Fig. 8. OM, BR706-R13-08.
77
Broctus kuensis Pessagno & Whalen 1982
Species code: BRO02
Synonymy:
1982 Broctus kuensis n. sp. – Pessagno & Whalen, p. 120, pl. 1,
fig. 7; pl. 2, figs. 17, 21.
2002 Broctus kuensis Pessagno & Whalen – Tekin, p. 186, pl. 3,
fig. 7.
Original description: Test as with genus, usually having six
to seven post-abdominal chambers. Cephalis conical; thorax, abdomen, and post-abdominal chambers trapezoidal
in cross section. Post-abdominal chambers increasing rapidly in width; last two post-abdominal chambers rapidly
decreasing in width. Cephalis and thorax with small irregular, polygonal pore frames almost completely obscured by
cover of microgranular silica. Abdomen and post-abdominal chambers composed of irregular tetragonal and pentagonal pore frames gradually increasing in size distally;
final two post-abdominal chambers composed of larger,
aligned, tetragonal (rectangular) pore frames.
Original remarks: See remarks under B. selwynensis, n. sp.
Measurements (µm):
Based on five specimens.
Length
245.0
245.0
185.0
207.6
Width (max.)
100.0
100.0
78.0
91.0
HT
Max.
Min.
Mean
Etymology: Broctus kuensis, n. sp., is named for Kue
Passage, west of its type locality.
Type locality: Sample QC 590A, Sandilands Formation
(Kunga Formation in Pessagno & Whalen, 1982), north
shore of Kunga Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands Formation, Queen Charlotte
Islands; Hocaköy Radiolarite and Gümüslü Allochthon,
Turkey.
Broctus ruesti Yeh 1987b
Species code: BRO03
Synonymy:
1987b Broctus ruesti n. sp. – Yeh, p. 54, pl. 4, figs. 1-3, 7, 21.
1987b Broctus sp. aff. B. ruesti n. sp. – Yeh, p. 54, pl. 4, fig. 6.
1987b Broctus sp. A – Yeh, p. 54, pl. 4, figs. 13, 25.
1987 Canutus (?) sp. A – Hattori, pl. 15, fig. 13.
1997 Parahsuum sp. B – Yao, pl. 14, fig. 657.
2004 Broctus ruesti Yeh – Ziabrev et al., Fig. 5-9.
Original description: Test spindle-shaped (conical when
broken), with five to seven post-abdominal chambers.
Cephalis small, hemispherical without horn. Chambers
increasing gradually in length, rapidly in width as added.
Final post-abdominal chamber terminating in narrow
tubular extension. Test wall double-layered. Inner latticed
wall consisting of regularly aligned square to rectangular
pore frames. Outer layer of cephalis imperforate, covered
with layer of microgranular silica. Thorax and subsequent
chambers with massive costae and thin transverse bars
forming rectangular pore frames overlapping on inner layer.
Bars of outer layer fragile, often broken and leaving short
remnants at costae. About eleven to thirteen costae visible
laterally. Pore frames gradually increasing in size from
thorax to final post-abdominal chamber then decreasing in
size distally.
Original remarks: Broctus ruesti n. sp., differs from B. sp. A
by having a narrower test with regular rectangular pore
frames throughout the test rather than irregular pore
78
frames on earlier chambers and regular pore frames on
final post-abdominal chambers.
Further remarks: Broctus ruesti Yeh is similar to Broctus
selwynensis Pessagno & Whalen in general shape of the test
and linear arrangement of pores. These two species differ in
the shape of the first segments: B. ruesti is pointed apically
whereas B. selwynensis is rounded.
Note that our specimens of. B. ruesti (pl. BRO03, figs. 2-4)
are considerably smaller than the type material.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length (max.)
400
350
400
270
Width (max.)
230
200
230
180
Etymology: This species is named for Dr. D. Ruest, in honor
of his contribution to the study of Radiolaria.
Type locality: Sample OR-536J, Nicely Formation, southeast side of Morgan Mountain, east-central Oregon.
Occurrence: Nicely Formation, Oregon; Fannin Formation,
Queen Charlotte Islands; Musallah Formation, Oman;
Japan; Bainang Terrane, Tibet.
Plate BRO02. Broctus kuensis Pessagno & Whalen. Magnification Fig. 1 x200, Fig. 2 x300. Fig. 1. TR, 1662D-R01-07.
Fig. 2(H). Pessagno & Whalen 1982, pl. 1, fig. 7.
Plate BRO03. Broctus ruesti Yeh. Magnification Fig. 1 x 150 (scale bar A), Figs. 2-4 x200 (scale bar B). Fig. 1(H). Yeh
1987b, pl. 4, fig. 21. Fig. 2. QCI, GSC loc. C-304565, GSC 128893. Fig. 3. QCI, GSC loc. C-080610, GSC 128894.
Fig. 4. OM-00-254, 022204.
79
Broctus selwynensis Pessagno & Whalen 1982
Species code: BRO01
Synonymy:
1982 Broctus selwynensis n. sp. – Pessagno & Whalen, p. 121,
pl. 1, fig. 6; pl. 2, figs 18, 20; pl. 12, fig. 10.
Original description: Test as with genus, usually with seven
or eight post-abdominal chambers. Cephalis hemispherical, imperforate. Abdomen and post-abdominal chambers increasing rapidly in width; last two post-abdominal
chambers rapidly decreasing in width. Thorax, abdomen
and post-abdominal chambers trapezoidal in cross section.
Thorax, abdomen and proximal post-abdominal chambers
with small, irregular polygonal pore frames in outer latticed
layer; central part of test composed of more regular tetragonal (mostly rectangular) pore frames in outer latticed layer
aligned in rows; final post-abdominal chamber and tubular
structure composed of less regularly aligned tetragonal and
pentagonal pore frames.
Original remarks: The aligned pore frames over the central
portion of the test distinguishes B. selwynensis, n. sp., from
B. kuensis, n. sp. The latter species does not show such a
strong alignment of pore frames, except for the final two
post-abdominal chambers.
Further remarks: See remarks under Broctus ruesti Yeh.
Measurements (µm):
Based on 10 specimens.
Length
237.5
260.0
210.0
238.8
Width (max.)
107.5
120.0
90.0
106.3
HT
Max.
Min.
Mean
Etymology: Broctus selwynensis is named for Selwyn Inlet,
northwest of its type locality.
Type locality: Sample QC 590A, Sandilands Formation
(Kunga Formation in Pessagno & Whalen, 1982), north
shore of Kunga Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands, Ghost Creek and Fannin formations, Queen Charlotte Islands; Williston Lake, NE British
Columbia; Tawi Sadh Member of the Guwayza Formation
and Haliw Formation, Oman.
Genus: Canoptum Pessagno 1979
Type species: Canoptum poissoni Pessagno, in Pessagno et al. 1979
Synonymy:
1979 Canoptum Pessagno n. gen. – Pessagno et al., p. 182.
1987a Paracanoptum n. gen. – Yeh, p. 67.
1987a Neowrangellium n. gen. – Yeh, p. 65.
Original description: Test spindle-shaped (often conical
when broken) with dome-shaped cephalis lacking horn.
Thorax and abdomen trapezoidal in outline. Post-abdominal segments subtrapezoidal in outline, separated from
each other by rather broad, slightly perforate, circumferential ridges at the joints; pores on ridges circular to elliptical in shape, not set in pore frames. Ridges of inner layer
considerably narrower. Area between a given ridges imperforate or sparsely perforate. Segments somewhat constricted between joints and circumferential ridges. Each
postabdominal segment separated by partitions with large,
circular apertures.
80
Original remarks: Canoptum n. gen., differs from Spongocapsula Pessagno in having a two-layered test wall lacking
spongy meshwork. It differs from Parvicingula Pessagno in
possessing a two-layered test with a microgranular outer
layer lacking discrete pore frames.
Etymology: Canoptum is an arbitrary combination of letters
(see ICZN, 1964. p. 113. recommendation 40).
Included species:
CAN12 Canoptum anulatum Pessagno & Poisson 1981
CAN13 Canoptum artum Yeh 1987b
CAN08 Canoptum columbiaense Whalen & Carter 1998
CAN09 Canoptum dixoni Pessagno & Whalen 1982
CAN11 Canoptum margaritaense Whalen & Carter 1998
CAN14 Canoptum rugosum Pessagno & Poisson 1981
Plate BRO01. Broctus selwynensis Pessagno & Whalen. Magnification x200. Fig. 1(H). Pessagno & Whalen 1982,
pl. 1, fig. 6. Fig. 2. QCI, GSC loc. C-140418, GSC 111806. Fig. 3. OM, BR1121-R10-14. Fig. 4. OM, Haliw-R03-05.
Fig. 5. QCI, GSC loc. C- 080611, GSC 128722. Fig. 6. QCI, GSC loc. C-080612, GSC 128723. Fig. 7. NBC, GSC loc. C305208, GSC 128724. Fig. 8. QCI, GSC loc. C-140495, GSC 128725.
81
Canoptum anulatum Pessagno & Poisson 1981
Species code: CAN 12
Synonymy:
1981 Canoptum anulatum n. sp. – Pessagno & Poisson, p. 60,
pl. 9, figs. 6-9, pl. 10, figs. 1-9.
1982 Dictyomitrella (?) sp. – Imoto et al., pl. 1, fig. 5.
1982 Canoptum anulatum Pessagno & Poisson – Pessagno &
Whalen, p. 123, pl. 6, figs. 1-2.
1982 Canoptum anulatum Pessagno & Poisson – De Wever &
Origlia-Devos, p. 1, fig. X.
1984 Canoptum anulatum Pessagno & Poisson – Murchey,
pl. 1, fig. 32.
1984 Canoptum anulatum Pessagno & Poisson – Whalen &
Pessagno, pl. 3, fig. 7.
1985 Canoptum anulatum Pessagno & Poisson – Kishida &
Hisada, pl. 2, figs. 21-22.
1987a Paracanoptum anulatum (Pessagno & Poisson) – Yeh,
p. 67, pl. 1, figs. 12, 13.
1987b Paracanoptum anulatum (Pessagno & Poisson) – Yeh,
p. 58, pl. 4, fig. 28; pl. 15, fig. 4; pl. 27, fig. 1, 9, 11.
1987 Canoptum anulatum Pessagno & Poisson – Hattori, pl. 18,
fig. 9.
1988 Canoptum anulatum Pessagno & Poisson – Carter et al.,
p. 50, pl. 5, figs. 9-10, 14.
1989 Canoptum sp. aff. C. anulatum Pessagno & Poisson
– Hattori, pl. 14, fig. A, not fig. B.
1990 Canoptum anulatum Pessagno & Poisson – De Wever et
al., pl. 4, fig. 5.
1992 Paracanoptum anulatum (Pessagno & Poisson) – Pessagno
& Mizutani, pl. 99, fig. 3.
1997 Canoptum anulatum Pessagno & Poisson – Yao, pl. 12,
fig. 574.
1998 Paracanoptum anulatum (Pessagno & Poisson) – Cordey,
pl. 24, fig. 1-2, 11.
1998 Canoptum anulatum Pessagno & Poisson – Kashiwagi,
pl. 1, fig. 15.
2002 Canoptum anulatum Pessagno & Poisson – Whalen &
Carter, p. 118, pl. 10, fig. 5.
2004 Canoptum anulatum Pessagno & Poisson – Matsuoka,
fig. 245.
2004 Canoptum anulatum Pessagno & Poisson – Ishida et al.,
pl. 5, fig. 9, 10.
Original description: Cephalis dome-shaped, lacking a
horn. Subsequent chambers trapezoidal in outline, numerous; closely spaced except for final chambers. Post-abdominal chambers eleven to fifteen in number, separated
by prominent circumferential ridges; ridges with short,
discontinuous costae; approximately fifteen costae visible
on a given ridge laterally. Short costae at right angles to
82
circumferential ridges, forming linked-H pattern. Single
small, circular pores occurring between two given costae
and adjacent to ridge. Pores, ridges and costae usually buried by microgranular outer layer of shell material except
when specimen is excessively etched. Pores in area between
ridges usually elliptical in shape, set in linearly arranged,
rectangular pore frames; usually buried by outer layer of
shell material. Final two post-abdominal chambers (segments) decreasing in width, increasing in height; penultimate chamber often with tubular extension.
Original remarks: Canoptum anulatum, n. sp., possesses
circumferential ridges that are significantly different from
those of the type species of Canoptum, C. poissoni Pessagno
(1979). The linked-H circumferential ridge structure displayed by C. anulatum is shared by a number of yet undescribed forms from the Lower Jurassic. Forms with this sort
of structure have not been observed below the Hettangian.
C. anulatum is tentatively included in Canoptum in this report. However, it may be desirable in the future to include
it under a new genus.
C. anulatum also differs from C. poissoni by having a
slender, more elongate test with more closely spaced postabdominal chambers (segments).
Measurements (µm):
Based on seven specimens.
Length
Width
HT
310
95
Min.
310
90
Max.
435
100
Etymology: Anulatus-a-um (Latin, adj.): beringed, ornamented with rings.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Bridge River
Complex, and Fannin and Whiteaves formations, Queen
Charlotte Islands; Nicely and Hyde formations, Warm
Springs member of the Snowshoe Formation, Oregon;
Franciscan Complex, California; San Hipólito Formation,
Baja California Sur; Drimos Formation, Greece; Tawi Sadh
Member of the Guwayza Formation, Oman; Japan.
Plate CAN12. Canoptum anulatum Pessagno & Poisson. Magnification x250, except Figs. 2b and 7b x500. Fig. 1(H).
Pessagno & Poisson 1981, pl. 9, fig. 6. Fig. 2a,b. TR, 1662D-R06-17. Fig. 3. JP, MNA-10, MA10776. Fig. 4. OM, BR1122R01-04. Fig. 5. OM, BR682-R09-02. Fig. 6. QCI, GSC loc. C-140495, GSC 128726. Fig. 7a,b. OM, BR523-R01-10a, b.
Fig. 8. OM, BR1122-R02-07. Fig. 9. Whalen & Carter 2002, pl. 10, fig. 5.
83
Canoptum artum Yeh 1987b
Species code: CAN13
Synonymy:
1987b Canoptum artum Yeh n. sp. – Yeh, p. 56, pl. 5, fig. 23; pl. 6,
figs. 5, 19-20; pl. 14, fig. 9; pl. 27, fig. 4.
1987b Canoptum sp. aff. C. artum Yeh n. sp. – p. 57, pl. 27,
figs. 2, 10., 24-25.
2004 Canoptum artum Yeh – Matsuoka, fig. 246.
2004 Canoptum sp. – Hori et al., pl. 6, figs. 21, 22.
Original description: Test wide, conical, with six to eight
post-abdominal chambers. Cephalis conical to domeshaped without horn. Cephalis, thorax, and abdomen imperforate, covered with layer of microgranular silica and
separated from each other by one row of small pores. All
post-abdominal chambers lobated, closely spaced, gradually increasing in width and length as added. Abdomen and
subsequent chambers separated from each other by smooth,
perforated circumferential ridges. Ridges usually with two
to four rows of small irregular polygonal pore frames. Inner
latticed layer of test comprised of larger, irregular polygonal
pore frames, outer layer of microgranular silica imperforate at constricted median band of earlier post-abdominal
chambers, perforated with small polygonal pore frames at
final three or four post-abdominal chambers.
Original remarks: Canoptum artum, n. sp., differs from
Canoptum poissoni Pessagno by having a wider test with
84
a larger cephalis and more closely spaced post-abdominal
chambers, and by having an outer layer of test wall more
perforate on final post-abdominal chambers.
Further remarks: Note that the size of this species is
quite variable. The length of the test ranges from 190 µm
(pl. CAN13, fig. 8) to 450 µm (pl. CAN13, fig. 5).
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length (max.)
300
295
310
240
Width (max.)
150
150
150
108
Etymology: Artus-a-um (Latin, adj.) = close, tight.
Type locality: OR-589D, Warm Springs member of the
Snowshoe Formation, near Izee, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, Oregon;
Ghost Creek and Fannin formations, Queen Charlotte
Islands; Mino Terrane, Japan; Tawi Sadh Member of the
Guwayza Formation and Musallah Formation, Oman.
Plate CAN13. Canoptum artum Yeh. Magnification Figs. 1-5 x200 (scale bar A), Figs. 6-8 x300 (scale bar B). Fig. 1(H).
Yeh 1987b, pl. 6, fig. 5. Fig. 2. OM, BR1121-R08-11. Fig. 3. QCI, GSC loc. C-304281, GSC 128727. Fig. 4. QCI, GSC loc.
C-080612, GSC 128728. Fig. 5. QCI, GSC loc. C-080612, GSC 128729. Fig. 6. OM-00-117, 021134. Fig. 7. JP, MNA-10,
MA13392. Fig. 8. OM, BR1122-R02-07.
85
Canoptum columbiaense Whalen & Carter 1998
Species code: CAN08
Synonymy:
1998 Canoptum columbiaense n. sp. – Whalen & Carter, p. 64,
pl. 15, figs 6, 10, 11, 15, 19.
1998 Canoptum sp. – Kashiwagi, pl. 2, fig. 12.
2002 Canoptum dixoni Pessagno & Whalen – Suzuki et al.,
p. 181, figs. 8 C-E, J-K.
? 2002 Canoptum columbiaense Whalen & Carter – Tekin,
p. 189, pl. 4, fig. 5.
Original description: Test conical, usually with 10 to 11
post-abdominal chambers. Cephalis and thorax combined
steeply conical, almost knob-like, with distinct break in
slope from abdomen. Abdomen and post-abdominal
chambers trapezoidal in outline, gradually increasing in
width and height as added. Cephalis and thorax smooth,
imperforate, covered by layer of microgranular silica.
Post-abdominal chambers separated from each other
and abdomen by moderately wide circumferential ridges
alternating with constrictions. Inner latticed layer of
post-abdominal chambers consisting of small, irregular
polygonal pore frames exposed on circumferential ridges;
layer of microgranular silica in constrictions mostly
covering polygonal pore frames on proximal part of test;
many pore frames exposed within constrictions on distal
part of test.
Original remarks: Canoptum columbiaense n. sp. differs
from C. margaritaense n. sp. by the absence of pronounced
nodes on the circumferential ridges; and from C. unicum
Pessagno and Whalen by the development of a thicker layer
of microgranular silica in the constrictions and the shape of
the cephalis.
Measurements (µm):
Based on 11 specimens.
Length
218
259
195
232
Width (max.)
98
120
90
108
HT
Max.
Min.
Mean
Etymology: This species is named for the Province of
British Columbia.
Type locality: Sample QC-676, Sandilands Formation,
Kunga Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands, Ghost Creek and Fannin
formations, Queen Charlotte Islands; Pucara Group, Peru;
Japan.
Canoptum dixoni Pessagno & Whalen 1982
Species code: CAN09
Synonymy:
1982 Canoptum dixoni Pessagno & Whalen n. sp. – Pessagno &
Whalen, p. 124, pl. 2, fig. 1-2, 8-9, 14; pl. 12, fig. 2.
1988 Canoptum anulatum Pessagno & Poisson – Li, pl. 1, fig. 2.
1998 Canoptum dixoni Pessagno & Whalen – Whalen & Carter,
p. 64, pl.17, fig. 1; pl. 26, fig. 5.
2002 Canoptum dixoni Pessagno & Whalen – Whalen & Carter,
p. 118, pl. 10, figs. 3, 4.
Original description: Test as with genus; conical, usually
with 12 to 14 post-abdominal chambers. Cephalis conical;
thorax, abdomen, and post-abdominal chambers increasing
gradually in width and height as added. Thorax, abdomen,
and post-bdominal chambers trapezoidal in outline.
Cephalis and thorax smooth, imperforate, covered by layer
of microgranular silica. Cephalis, thorax, and abdomen
together forming prominent area of test. Post-abdominal
chambers separated from each other and abdomen by
narrow circumferential ridges alternating with constrictions.
Inner latticed layer of post-abdominal chambers consisting
of small, irregular polygonal pore frames exposed on
circumferential ridges; layer of microgranular silica in
constrictions mostly covering polygonal pore frames,
particularly on proximal portion of test (many more pores
within constrictions remaining open distally). Distinctive
beadlike structures on circumferentail ridges formed by
86
raised areas of microgranular silica surrounded by small
polygonal pores.
Original remarks: Canoptum dixoni, n. sp. is distinguished
from other species of Canoptum by the beaded nature of its
narrow circumferential ridges.
Measurements (µm):
Based on 7 specimens.
Length
312.5
360.0
312.5
339.7
Width (maximum)
112.5
120.0
100.0
110.5
HT
Max.
Min.
Mean
Etymology: This species is named for Captain George
Dixon who explored the Queen Charlotte Islands in 1787.
Type locality: QC 590A, Sandilands Formation (Kunga
Formation in Pessagno & Whalen, 1982), north shore of
Kunga Island, Queen Charlotte Islands.
Occurrence: Sandilands formation, Queen Charlotte
Islands; San Hipólito Formation, Baja California Sur;
Dengqen area, Tibet; Tawi Sadh Member of the Guwayza
Formation and Musallah Formation, Oman.
Plate CAN08. Canoptum columbiaense Whalen & Carter. Magnification x200. Fig. 1(H). Carter et al. 1998, pl. 15,
fig. 6. Fig. 2. Carter et al. 1998, pl. 15, fig. 11. Fig. 3. Carter et al. 1998, pl. 15, fig. 10. Fig. 4. QCI, GSC loc. C-080613, GSC
128730. Fig. 5. QCI, GSC loc. C-304566, GSC 128731. Fig. 6. QCI, GSC loc. C-304565, GSC 128732. Fig. 7. QCI, GSC
loc. C-304281, GSC 128733. Fig. 8. QCI, GSC loc. C-140495, GSC 128734. Fig. 9. QCI, GSC loc. C-175309, GSC 128735.
Plate CAN09. Canoptum dixoni Pessagno & Whalen. Magnification x200. Fig. 1(H). Pessagno & Whalen 1982, pl. 2,
fig. 1. Fig. 2. Pessagno & Whalen 1982, pl. 2, fig. 2. Fig. 3. Whalen & Carter 2002, pl. 10, fig. 3. Fig. 4. Whalen & Carter
2002, pl. 10, fig. 4. Fig. 5. OM-00-252, 021824.
87
Canoptum margaritaense Whalen & Carter 1998
Species code: CAN11
Synonymy:
1987 Canoptum preanulatum Pessagno & Whalen – Hattori,
pl. 18, figs. 7, 8.
1987a Neowrangellium pessagnoi n. sp. – Yeh, p. 66, pl. 1, figs. 3,
4, ?10, 22, 23, ; not pl. 1, figs. 5-7, 11, 14.
1998 Canoptum margaritaense n. sp. – Whalen & Carter, p. 64,
pl. 17, figs. 2, 3.
? 2002 Canoptum margaritaense Whalen & Carter – Tekin,
p. 189, pl. 4, figs. 6, 7.
Original description: Test conical, lobulate, usually with 11
to 13 post-abdominal chambers. Cephalis conical; thorax,
abdomen and post-abdominal chambers trapezoidal in
outline, increasing gradually in width and height as added.
Cephalis and thorax smooth, imperforate, covered by
layer of microgranular silica. Post-abdominal chambers
separated from each other and abdomen by broad, nodose
circumferential ridges alternating with constrictions;
constrictions covered by thick layer of microgranular silica.
Inner latticed layer of post-abdominal chambers usually
obscured but when exposed, consisting of small, polygonal
pore frames. H-linked pattern on circumferential ridges
formed by raised nodes of microgranular silica surrounded
by small polygonal pores.
Original remarks: The H-linked pattern on the circumferential ridges distinguish Canoptum margaritaense n. sp.
from C. dixoni Pessagno and Whalen and C. columbiaense
n. sp.
Further remarks: In this species we include forms without
pores in the constrictions that are identical to the paratypes
of Neowrangellium pessagnoi Yeh but differ from the holotype by lacking a horn.
Measurements (µm):
Based on 6 specimens.
Length
368
368
225
287
Width (max.)
128
128
83
101
HT
Max.
Min.
Mean
Etymology: This species is named for Cape St. Margarita,
the northern tip of the Queen Charlotte Islands.
Type locality: Sample QC-675, Sandilands Formation,
Kunga Island - north side, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands, Ghost Creek and Fannin formations, Queen Charlotte Islands; Hyde Formation, Oregon.
Canoptum rugosum Pessagno & Poisson 1981
Species code: CAN14
Synonymy:
1981 Canoptum rugosum n. sp. – Pessagno and Poisson, p. 61,
pl. 11, figs. 5-9; pl. 13, fig. 3; pl. 14, figs. 1-2.
1982 Canoptum rugosum Pessagno & Poisson – Pessagno and
Whalen, p. 125, Pl. 6, Fig. 7.
1987 Canoptum rugosum Pessagno & Poisson – Hattori, pl. 18,
figs. 10-12.
1988 Canoptum rugosum Pessagno & Poisson – Sashida, p. 23,
pl. 2, figs. 13, 14, 22, 23.
1988 Canoptum rugosum Pessagno & Poisson – Li, pl. 1, fig. 1.
1989 Canoptum rugosum Pessagno & Poisson – Hattori, pl. 13,
figs. F, G, H, I.
1995 Canoptum rugosum Pessagno & Poisson – Suzuki, pl. 8,
fig. 2.
1998 Canoptum rugosum Pessagno & Whalen – Kashiwagi,
pl. 1, fig. 16, pl. 2, fig. 11.
2003 Canoptum rugosum Pessagno & Poisson – Goričan et al.,
p. 297, pl. 5, fig. 11.
2003 Canoptum cf. rugosum Pessagno & Poisson – Kashiwagi &
Kurimoto, pl. 3, fig. 14.
2004 Canoptum rugosum Pessagno & Poisson – Matsuoka,
fig. 244.
2005 Canoptum sp. cf. C. rugosum Pessagno & Poisson
– Kashiwagi et al., pl. 5, fig. 1.
Original description: Test as with genus. Cephalis hemispherical; post-cephalic chambers trapezoidal in ouline
increasing relatively rapidly in width and height as added.
Circumferential ridges absent between cephalis and thorax
88
and between abdomen and thorax; present between abdomen and first post-abdominal chamber and each of subsequent four or five post-abdominal chambers. Circumferential ridge when stripped of outer layer of microgranular
shell material displaying linked-H pattern identical to that
described for C. anulatum, n. sp. Post-abdominal chambers
constricted medially, giving rise to lobulate test outline. Inner layer of post-abdominal chambers comprised of two
rows of massive tetragonal pore frames between circumferential ridges. Outer microgranular layer on well preserved
specimens with rugose surface; rugosities probably a reflection of massive of inner layer.
Original remarks: Canoptum rugosum, n. sp., differs from
C. anulatum, n. sp., (1) by having a shorter, broader test
with one half to one third the number of post-abdominal
chambers; (2) by having widely rather than closely spaced
circumferential ridges; (3) by having post-abdominal
chambers with a rugose surface; and so forth. Both species
share the same linked-H circumferential ridge structure.
Measurements (µm):
Based on eight specimens.
Length
Width
HT
140
75
Min.
140
65
Max.
165
95
Plate CAN11. Canoptum margaritaense Whalen & Carter. Magnification x200. Fig. 1(H). Carter et al. 1998, pl. 17,
fig. 2. Fig. 2. Carter et al. 1998, pl. 17, fig. 3. Fig. 3. QCI, GSC loc. C-080611, GSC 128736. Fig. 4. QCI, GSC loc. C080611, GSC 128737. Fig. 5. QCI, GSC loc. C-080610, GSC 128738.
Etymology: Rugosus-a-um (Latin, adj.): wrinkled.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Ghost Creek
Formation, Queen Charlotte Islands; Skrile Formation,
Slovenia; Dengqen area, Tibet; Japan; Musallah Formation,
Oman.
Plate CAN14. Canoptum rugosum Pessagno & Poisson. Magnification x300. Fig. 1(H). Pessagno & Poisson 1981, pl.11,
fig. 5. Fig. 2. OM-00-117, 021130. Fig. 3. JP, MNA-10, MA13381. Fig. 4. SI, MM6.76, 010301. Fig. 5. Goričan et al. 2003,
pl. 5, fig. 11. Fig. 6. QCI, GSC loc. C-304281, GSC 128739.
89
Genus: Canutus Pessagno & Whalen 1982
Type species: Canutus tipperi Pessagno & Whalen 1982
Synonymy:
1982 Canutus n. sp. – Pessagno & Whalen, p.127.
Original description: Test spindle-shaped to subconical;
when spindle-shaped, often quite inflated. Cephalis without horn. Abdomen and post-abdominal chambers with
two or three layers of fragile polygonal pore frames.
Original remarks: Canutus n. gen., differs from Archaeodictyomitra Pessagno by having a test with several latticed
layers of pore frames, by developing pillar-like nodes, and
by lacking costae.
Further remarks: Only the inflated spindle-shaped forms
are considered to belong to Canutus Pessagno & Whalen,
the subconical forms are now assigned to Parahsuum Yao.
See also further remarks under genus Parahsuum.
Etymology: Canutus is a name formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, Pt. VI,
Recommendation 40, p.113).
Included species:
CTS06 Canutus baumgartneri Yeh 1987b
CTS08 Canutus diegoi Whalen & Carter 2002
CTS09 Canutus hainaensis Pessagno & Whalen 1982
CTS10 Canutus nitidus Yeh 1987b
CTS15 Canutus rennellensis Carter n. sp.
CTS03 Canutus rockfishensis Pessagno & Whalen 1982
CTS12 Canutus tipperi gr. Pessagno & Whalen 1982
CTS16 Canutus sp. O
Canutus baumgartneri Yeh 1987b
Species code: CTS06
Synonymy:
1987b Canutus baumgartneri n. sp. – Yeh, p. 59, pl. 19, figs. 3-5,
9, 16-17, 20-21.
Original description: Test spindle-shaped, large, inflated,
usually with five to six post-abdominal chambers. Cephalis
hemispherical, covered with layer of microgranular
silica. Thorax and subsequent chambers trapezoidal in
outline, gradually increasing in width, final three postabdominal chambers gradually decreasing in width. Test
wall consisting of three layers. Inner latticed layer and
intermediate (second) latticed layer comprised of medium
size square to rectangular pore frames (pl. 19, figs. 4-5, 16,
21), outermost latticed layers consisting predominantly of
triangular pore frames. Distal post-abdominal chambers
often lacking outermost layer of meshwork and showing
rectangular pore frames of inner latticed layer(s) (pl. 19,
figs. 3, 20).
Original remarks: Canutus baumgartneri, n. sp. differs
from C. blomei Pessagno & Whalen by having a less inflated
test with less massive pore frames. It can be distinguished
from C. tipperi Pessagno and Whalen by having a test with
less massive pore frames and a less pointed cephalis.
90
Further remarks: Canutus tipperi and C. blomei are synonymized herein as the C. tipperi group. C. baumgartneri differs from this group mainly by having less massive, more
irregularly-shaped pore frames. Lowest Pliensbachian representatives of this genus are more strongly spindle-shaped
(almost closed at the bottom) and pore frames are even
more irregular.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length (max.)
367
370
380
363
Width (max.)
227
228
230
225
Etymology: This species is named after Dr. P. O.
Baumgartner, in honor of his studies on the Mesozoic
Radiolaria.
Type locality: Sample OR-600D, Hyde Formation at IzeePaulina road, east-central Oregon.
Occurrence: Hyde Formation, Oregon; Ghost Creek and
Fannin formations, Queen Charlotte Islands.
Plate CTS06. Canutus baumgartneri Yeh. Magnification x150. Fig. 1(H). Yeh 1987b, pl. 20, fig. 3. Fig. 2. QCI, GSC loc.
C-304568, GSC 128770. Fig. 3. QCI, GSC loc. C-175306, GSC 128771. Fig. 4. QCI, GSC loc. C-304567, GSC 128772.
Fig. 5. QCI, GSC loc. C-175311, GSC 128773. Fig. 6 QCI, GSC loc. C-080611, GSC 128774.
91
Canutus diegoi Whalen & Carter 2002
Species code: CTS08
Synonymy:
1984 Canutus? sp. – Whalen & Pessagno, pl. 3, fig. 5, 6.
? 1987 Canutus (?) sp. K – Hattori, pl. 15, fig. 10.
2002 Canutus diegoi n. sp. – Whalen & Carter, p. 120, pl. 10,
figs. 6, 10, 13, 16, 17; pl. 17, figs. 6, 7.
Original description: Test spindle shaped, slightly inflated
with approximately six post-abdominal chambers; large,
dome-shaped cephalis covered by layer of microgranular
silica; thorax, abdomen and post-abdominal chambers
trapezoidal in outline, gradually increasing in width; last
few post-abdominal chambers gradually decreasing in
width and terminating in a short, irregular tubular extension. Chambers gradually increasing in height till central
post-abdominal chamber and then gradually decreasing
in height. Cephalis, thorax, abdomen and post-abdominal
chambers on proximal half of test with irregularly shaped
pore frames in outer latticed layer; distal post-abdominal
chambers composed of slightly more aligned tetragonalpentagonal pore frames. Inner latticed layer of entire test
composed of strong, rectangular pore frames.
Original remarks: The less inflated test and strongly tapering distal chamber distinguish this species from Canutus
rockfishensis Pessagno and Whalen 1982.
Measurements (µm):
Based on 9 specimens.
Length
225
225
180
203
Width (Max.)
120
135
120
124
HT
Max.
Min.
Mean
Etymology: Canutus diegoi n. sp., is named for Diego
de Becerra, one of the early explorers of the Baja California
Peninsula.
Type locality: Sample BPW80-30, San Hipólito Formation,
Punta San Hipólito, Vizcaino Peninsula, Baja California
Sur.
Occurrence: San Hipólito Formation, Baja California Sur.
Canutus hainaensis Pessagno & Whalen 1982
Species code: CTS09
Synonymy:
1982 Canutus hainaensis n. sp. – Pessagno & Whalen, p. 128,
pl. 4, figs. 3-4; pl. 5, figs. 1, 13, 14, 16-18, 20; pl. 12, fig. 9.
1988 Canutus hainaensis Pessagno & Whalen – Carter et al.,
p. 51, pl. 3, figs. 10-11.
1992 Canutus hainaensis Pessagno & Whalen – Pessagno &
Mizutani, pl. 99, figs. 5, 14, 19.
Original description: Test elongate, spindle-shaped, often
large, with seven to nine post-abdominal chambers usually
present. Cephalis conical, rounded apically; remaining
chambers trapezoidal in cross section. Thorax, abdomen,
and all but final two post-abdominal chambers increasing
moderatly rapidly in width and more gradually in length
as added. Final two post-abdominal chambers decreasing
somewhat in width. Inner latticed layer of post-abdominal
chambers consisting of square to rectangular pore frames
with nodes at vertices; pore frames gradually increasing
in size, becoming large on final post-abdominal chamber.
Outer latticed layer (second layer) consisting of regular
to irregular, commonly tetragonal (rectangular) and
triangular pore frames. Outer latticed layer not developed
on final chamber of well-preserved specimens; pore frames
of inner layer with rudimentary nodes.
92
Original remarks: Canutus hainaensis, n. sp., is considerably more elongate and much less inflated than C. tipperi,
n. sp. It also differs from C. tipperi by having a cephalis
rounded rather than pointed apically and by having two
rather than three layers of meshwork on its post-abdominal chambers.
Measurements (µm):
Based on 6 specimens.
Length
375.0
400.0
300.0
357.5
Width (max.)
200.0
200.0
160.0
177.9
HT
Max.
Min.
Mean
Etymology: This species is named for Haina, an abandoned
Indian village on the east shore of Maude Island.
Type locality: Sample QC 534, Ghost Creek Formation
(Maude Formation in Pessagno & Whalen, 1982), Queen
Charlotte Islands, British Columbia.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands.
Plate CTS08. Canutus diegoi Whalen & Carter. Magnification Fig. 1a x200, Fig. 1b x300. Fig. 1(H)a, b. Whalen &
Carter 2002, pl. 10, figs. 10, 13.
Plate CTS09. Canutus hainaensis Pessagno & Whalen. Magnification x150. Fig. 1(H). Pessagno & Whalen 1982, pl. 4,
fig. 3. Fig. 2. QCI, GSC loc. C-304567, GSC 128775. Fig. 3. QCI, GSC loc. C-304567, GSC 128776. Fig. 4. QCI, GSC loc.
C-175311, GSC 128895.
93
Canutus nitidus Yeh 1987b
Species code: CTS10
Synonymy:
1987b Canutus nitidus n. sp. – Yeh, p. 59, pl. 6, figs. 1, 17; pl. 19,
figs. 1-2, 6, 11, 18-19.
1987b Canutus sp. aff. C. nitidus n. sp. – Yeh, p. 60, pl. 4, figs. 4-5,
22.
1988 Canutus nitidus Yeh – Carter et al., p. 50, pl. 3, figs. 5, 8, 12.
1988 Canutus sp. aff. C. nitidus Yeh – Carter et al., p. 51, pl. 3,
fig. 6.
Original description: Test spindle-shaped, large, very inflated, usually with five to six post-abdominal chambers.
Cephalis small, hemispherical, usually without rudimentary
spine. Abdomen, thorax to second post-abdominal chamber rapidly increasing in width, remaining post-abdominal
chambers rapidly decreasing in width. Cephalis covered
with layer of microgranular silica, remaining chambers
comprised of two inner layers of variable size of irregular
polygonal pore frames (pl. 19, fig. 6), outer most layer of
triangular pore frames. Pore frames thin in rims and thick
in sides, with largest pores at middle portion of test, decreasing in size apically and distally. Final post-abdominal
chamber terminating in narrow tubular extension.
Original remarks: Canutus nitidus, n. sp., differs from
C. baumgartneri n. sp., by possessing a very inflated test
94
with extremely small cephalis, and by having a test with two
inner layers of variable sized irregular pore frames.
Further remarks: By Carter et al. (1988): The narrow tubular extension terminating the distalmost post-abdominal
chamber, as described by Yeh (1987b, p. 59), has not been
observed in any specimens.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length (max.)
324
315
325
302
Width (max.)
248
243
250
237
Etymology: Nitidus-a-um (Latin, adj.) = sleek, glittering.
Type locality: Sample OR-600M, Hyde Formation at IzeePaulina road, east-central Oregon.
Occurrence: Hyde Formation, Oregon; Fannin Formation,
Queen Charlotte Islands; Fernie Formation, NE British
Columbia.
Plate CTS10. Canutus nitidus Yeh. Magnification x150. Fig. 1(H). Yeh 1987b, pl. 20, fig. 1. Fig. 2. Carter et al. 1988,
pl. 3, fig. 5. Fig. 3. NBC, GSC loc. C-305208, GSC 128896. Fig. 4. QCI, GSC loc. C-175309, GSC 128897.
Fig. 5. QCI, GSC loc. C-304567, GSC 128898. Fig. 6. QCI, GSC loc. C-080612, GSC 128899. Fig. 7. QCI, GSC loc. C175306, GSC 128900. Fig. 8. QCI, GSC loc. C-304567, GSC 128777. Fig. 9. QCI, GSC loc. C-304567, GSC 128778.
Fig. 10. QCI, GSC loc. C-304567, GSC 128901. Fig. 11. QCI, GSC loc. C-304567, GSC 128902.
95
Canutus rennellensis Carter n. sp.
Species code: CTS15
Synonymy:
1982 Canutus blomei n. sp. – Pessagno & Whalen, p. 127, pl. 3,
fig. 14 only.
1996 Canutus sp. A - Pujana, p. 138, pl. 1, figs. 18, 19.
1996 Canutus sp. A of Carter – Pujana, p. 138, pl. 1, fig. 20.
2004 Canutus sp. – Matsuoka, fig. 211.
Type designation: Holotype GSC 111711 and paratype
GSC 111712 from GSC loc. C-080612; Ghost Creek
Formation (lower Pliensbachian).
Description: Test spindle-shaped, large, inflated, usually
with five or six post-abdominal chambers. Cephalis hemispherical; thorax, abdomen and first few abdominal chambers rapidly increasing in width, last few post-abdominal
chambers strongly decreasing in width. Apical portion of
test fairly pointed and covered with a layer of microgranular silica. Pore frames on early chambers relatively small,
irregularly-shaped and arranged; on medial chambers
pore frames mainly tetragonal and aligned in rows, some
with costae-like ridges between. Pore frames on distalmost
chamber(s) smaller and irregular.
Remarks: Canutus rennellensis n. sp. differs from the
C. tipperi group in having finer, more irregular meshwork,
and distal chambers are more constricted. It differs from
C. baumgartneri Yeh in having much finer meshwork and
pore frames on distal post-abdominal chambers are aligned
rather than irregular.
Measurements (µm):
Based on 6 specimens.
Length (excl. horn)
Maximum width
HT
326
195
Max.
353
205
Min.
316
179
Mean
328
191
Etymology: Species named for the type locality at Rennell
Junction, the confluence of logging roads leading north to
Masset and west to Rennell Sound; Graham Island, Queen
Charlotte Islands.
Type locality: Sample CAA-79-Ren-Phant, lms 1 (GSC
loc. C-080611), Ghost Creek Formation, Rennell Junction
section, central Graham Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Sierra Chacaicó Formation, Argentina;
Mino Terrane, Japan; Haliw (Aqil) and Musallah formations, Oman.
Canutus rockfishensis Pessagno & Whalen 1982
Species code: CTS03
Synonymy:
1982 Canutus rockfishensis n. sp. – Pessagno & Whalen, p. 129,
pl. 2, figs. 4, 12, 15, 19; pl. 12, fig. 22.
1998 Canutus rockfishensis Pessagno & Whalen – Whalen &
Carter, p. 65, pl. 17, fig. 18; pl. 26, fig. 8.
2002 Canutus rockfishensis Pessagno & Whalen – Suzuki et al.,
p. 184, fig. 9 I.
Original description: Test as with genus, spindle-shaped,
large, moderately inflated, usually with seven postabdominal chambers. Cephalis and thorax conical, mostly
imperforate, composed of inner latticed layers of pore
frames covered by an outer layer of microgranular silica.
Abdomen and proximal post-abdominal chambers rapidly
increasing in width, central two or three post-abdominal
chambers gradually increasing in width; last two postabdominal chambers rapidly decreasing in width; thorax,
abdomen and post-abdominal chambers trapezoidal in
cross section. All post-abdominal chambers increasing
in height distally. Abdomen and most post-abdominal
chambers composed of irregular pentagonal to tetragonal
pore frames in outer latticed layer; last two post-abdominal
chambers composed of aligned tetragonal (rectangular)
pore frames. Inner latticed layer composed of rectangular
pore frames.
Original remarks: Canutus rockfishensis, n. sp., differs from
C. blomei, n. sp., by having a test which is not nearly so
96
broad and by having post-abdominal chambers with two
rather than three layers of latticed meshwork. Furthermore,
most of the outer latticed layer of C. rockfishensis possesses
irregular pore frames, whereas that of C. blomei possesses
triangular pore frames.
Further remarks: The holotype of Canutus blomei Pessagno
& Whalen is now assigned to C. tipperi Pessagno & Whalen
while its paratype is assigned to C. rennellensis Carter n. sp.
C. rockfishensis Pessagno & Whalen differs from both these
species by having a much less inflated test.
Measurements (µm):
Based on 10 specimens.
Length
222.5
270.0
220.0
243.0
Width (maximum)
125.0
150.0
125.0
138.5
HT
Max.
Min.
Mean
Type locality: Sample QC 590A, Sandilands Formation
(Kunga Formation in Pessagno & Whalen 1982), Queen
Charlotte Islands, British Columbia.
Occurrence: Sandilands and Fannin formations, Queen
Charlotte Islands; Nicely Formation, east-central Oregon;
Pucara Group, Peru.
Plate CTS15. Canutus rennellensis Carter n. sp. Magnification x150. Fig. 1(H). QCI, GSC loc. C-080612, GSC 111711.
Fig. 2. QCI, GSC loc. C-080612, GSC 128783. Fig. 3. QCI, GSC loc. C-304281, GSC 128784. Fig. 4. QCI, GSC loc. C080612, GSC 111712. Fig. 5. JP, MNA-10, MA13048. Fig. 6. OM-00-118, 000604.
Plate CTS03. Canutus rockfishensis Pessagno & Whalen. Magnification x250. Fig. 1(H). Pessagno & Whalen 1982,
pl. 2, fig. 4.
97
Canutus tipperi gr. Pessagno & Whalen 1982
Species code: CTS12
Synonymy:
1982 Canutus tipperi n. sp. – Pessagno & Whalen, p. 129, pl. 4,
figs 7-9, 11, 12, 14-17; pl. 12, fig. 21.
1982 Canutus blomei n. sp. – Pessagno & Whalen, p. 127, pl. 3,
figs. 13, 15 (not fig. 14); pl. 12, fig. 20.
1988 Canutus tipperi Pessagno & Whalen – Carter et al., p. 51,
pl. 3, fig. 3.
Original description: Test spindle-shaped, pointed apically, large, inflated, usually with seven to nine post-abdominal chambers. Cephalis hemispherical; remaining
chambers trapezoidal in cross section; cephalis and thorax
with apically converging ridges on better preserved specimens. Abdomen and first three post-abdominal chambers
rapidly increasing in height as added. Inner latticed layer
and intermediate (second) latticed layer comprised of large
square to rectangular pore frames (pl. 4, figs. 15, 17); outer
latticed layer comprised predominantly of triangular pore
frames. Final post-abdominal chamber of well-preserved
specimens often lacking outer two layers of meshwork and
showing only rudimentary development of nodes at pore
frame vertices.
Queen Charlotte Islands indicate that test morphology is
quite variable. Basal Pliensbachian forms are small, less than
half the size of the type species, others are elongate and less
inflated, but all retain the typical massive rectangular pore
structure of C. tipperi. The shape of the apical portion of the
test is also variable, ranging from the fairly pointed shape
of the holotype to the more hemispherical shape of the
original holotype of C. blomei (the latter now synonymized
with C. tipperi).
C. tipperi was originally recorded from the Rennell
Junction member of the Fannin Formation (upper lower
Pliensbachian) but is now known to appear in basal beds
of the lower Pliensbachian Ghost Creek Formation and
ranges throughout the entire Pliensbachian.
Measurements (µm):
Based on 6 specimens.
Length
475.0
475.0
280.0
313.0
Width (max.)
200.0
200.0
180.0
195.4
HT
Max.
Min.
Mean
Original remarks: Canutus tipperi n. sp. is compared to
C. blomei n. sp. under the latter species.
Original remarks under C. blomei: Canutus blomei, n. sp.,
appears closely related to C. tipperi, n. sp. It can be distinguished from C. tipperi by the less pointed and more rounded nature of the apical portion of the test. Furthermore, the
test of C. blomei is more inflated and ellipsoidal in character
than that of C. tipperi. In addition, its pore frames are more
massive with less prominent nodes.
Type locality: QC 532, Fannin Formation (Maude Formation in Pessagno & Whalen, 1982), Skidegate Inlet, Maude
Island, Queen Charlotte Islands, British Columbia.
Further remarks: Canutus tipperi is an extremely large
multicyrtid with distinctive pore structure, but recent
studies of Canutus in over 80 Pliensbachian samples from
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands, and Williston Lake, northeastern British
Columbia.
98
Etymology: This species is named for Dr. Howard W. Tipper,
Geological Survey of Canada, in honor to his contributions
to the study of the Jurassic ammonite biostratigraphy of the
Queen Charlotte Islands.
Plate CTS12. Canutus tipperi gr. Pessagno & Whalen. Magnification x150. Fig. 1(H). Pessagno & Whalen 1982, pl. 4,
fig. 7. Fig. 2. QCI, GSC loc. C-175311, GSC 128779. Fig. 3. QCI, GSC loc. C-175309, GSC 128780. Fig. 4. QCI, GSC loc.
C-080612, GSC 128781. Fig. 5. QCI, GSC loc. C-175309, GSC 128782. Fig. 6. Carter et al. 1988, pl. 3, fig. 3.
Fig. 7. QCI, GSC loc. C-127867, GSC 128903.
99
Canutus sp. O
Species code: CTS16
Remarks: Test large, subconical to slightly spindle-shaped.
Cephalis hemispherical, thorax and abdomen trapezoidal,
post abdominal chambers gradually increasing in width as
added Cephalis and thorax sparsely perforate covered with
a layer of microgranular silica. Pore frames on initial postabdominal chambers mostly irregular, on distal portion
pore frames mainly tetragonal and aligned vertically. This
species differs from C. hainaensis in having a more broadly
conical shape with prominently aligned distal chambers.
Occurrence: Ghost Creek Formation, Queen Charlotte
Islands; Musallah Formation, Oman.
Genus: Carterwhalenia Dumitrica n. gen.
Type species: Saitoum (?) minai Whalen & Carter 2002
Description: Test monocyrtid, hemispherical to subglobular with an initial spicule consisting of apical (A),
dorsal (D), ventral (V), and primary lateral spines (Lr, Ll)
originating in a short median bar (MB), and arches LL,
LD, LV, AV, AL, and AD. A spine extended outside into
apical horn, and D and L into feet. Apical horn and feet
four-bladed and practically equal, at least in type species. Blades with a row of three or more pores decreasing
in size distally. Pores aligned between the axis of apical
horn and feet and the external border of blades. Ventral
spine of initial spicule extended outside cephalic wall into
a short bladed spine.
Remarks: Carterwhalenia n. gen. externally resembles Saitulpus Dumitrica & Zügel from which it differs structurally
especially in missing the secondary lateral spines of the initial spicule. By this character it also resembles very much
the Cenozoic genera Euscenium Haeckel, Archiscenium
Haeckel, and Pteroscenium Haeckel. From each it differs in
having four-bladed rather than three-bladed spines. Carterwhalenia is close to the Middle and Upper Jurassic genus
Turriseiffelus Dumitrica & Zügel, in the structure of the
100
initial spicule, apical horn, feet, and cephalic wall, but differs in having four-bladed, short apical horn and feet.
The pores of the blades of the apical horn and feet of this
genus have the same origin as similar pores of the genera
Turriseiffelus, Pteroscenium (e.g. Pteroscenium pinnatum
Haeckel 1887, pl. 53, figs. 14-16), and Arachnoplecta Dumitrica & Zügel 2003. They appear as meshes among the
axis of these spines, a bar forming the external margin of
the blades and branches arising practically perpendicular
from the axis of the spines. Usually these branches may extend into thorns outside the margin of the blades.
Until present the genus is represented only by its type
species.
Etymology: The genus is named for my radiolarian
colleagues E. S. Carter and P. A. Whalen to honour their
valuable contribution to the knowledge of Upper Triassic
and Lower Jurassic radiolarians.
Included species:
SUM03 Carterwhalenia minai (Whalen & Carter) 2002
Plate CTS16. Canutus sp. O. Magnification x200. Fig. 1. QCI, GSC loc. C-175311, GSC 128785. Fig. 2. QCI, GSC loc.
C-080612, GSC 128786. Fig. 3. OM-00-115, 023003. Fig. 4. OM-00-115, 023010. Fig. 5. QCI, GSC loc. C-175306, GSC
128787. Fig. 6. QCI, GSC loc. C-175306, GSC 128904. Fig. 7. QCI, GSC loc. C-175309, GSC 128905.
101
Carterwhalenia minai (Whalen & Carter) 2002
Species code: SUM03
Synonymy:
2002 Saitoum? minai n. sp. – Whalen & Carter, p. 130, pl. 12, figs.
7-9, 16, 17; pl. 13, fig. 10.
Original description: Test monocyrtid, sub-spherical in
shape with massive apical horn and three feet. Horn triradiate in axial section with narrow, longitudinal ridges and
broad grooves for most of length; elongated pores occasionally located at base of grooves of horn; horn circular in
axial section distally, separated from triradiate portion by
three to four short verticils; verticils aligned at right angles
to long axis of horn; horn located off center of highest point
on cephalic dome. Three massive feet triradiate in axial
section for most of length, circular in axial section distally;
triradiate and circular portions of feet separated by verticils
aligned at right angles to long axis of feet; proximal part
of feet sometimes with small pores piercing grooves. Imperforate border collar continuous with ridges of feet. Pore
frames irregular, triangular to circular, sometimes appearing to radiate from a central point. V-spine prominently exposed on exterior of test at point half way between border
collar and apex of test.
Original remarks: Saitoum? minai n. sp. is distinguished
from other species of Saitoum by the massive, spiny triradiate
apical horn and feet with verticils. This species differs from
species of Saitulpus (Dumitrica and Zügel, in press) in that
the V spine does not connect with the border collar.
Further remarks: An attentive look at the apical horn and
feet of the type specimens (holotype and two paratypes)
and of the two specimens herein presented from the
Pliensbachian of Turkey (pl. SUM03, figs. 4a-b, 5a-b) shows
that these external spines are four-bladed rather than threebladed as originally mentioned and that the secondary
lateral spines are completely missing.
Measurements (µm):
(n) = number of specimens measured.
HT
Max.
Min.
Mean
Length (6)
(excludes horn)
98
105
83
92
Width (5)
(Max.)
90
113
90
104
Length of foot
(7)
90
105
75
87
Etymology: This species is named for F. Mina (Geólogos
de Petróleos Mexicanos), one of the first geologists to study
the rocks of the Vizcaino Peninsula, Baja California Sur.
Type locality: Sample BPW80-30, San Hipólito Formation,
Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Gümüslü Allochthon, Turkey.
Genus: Charlottea Whalen & Carter 1998
Type species: Charlottea amurensis Whalen & Carter 1998
Synonymy:
1998 Charlottea n. gen. – Whalen & Carter, p. 37.
Original description: Test with three prominent spines in
same plane, equally spaced or with two spines closer together;
spines triradiate in axial section, tapering distally. Cortical
shell spherical to sub-spherical, sometimes triangular in
outline with flattened upper and lower surfaces. Outer layer
of pore frames on cortical shell irregularly shaped (usually
tetragonal, triangular, pentagonal) and sized, with nodes at
pore frame vertices; larger pores on cortical shell sometimes
observed at base of spines.
Original remarks: The external morphology of Charlottea
n. gen. and Ferresium Blome is very similar but the inner
structure is different: Charlottea n. gen. contains an eccentric spicular meshwork occupying a large part of the central
area of the test; Ferresium Blome contains a small microsphere surrounded by a loosely constructed inner spongy
meshwork of bars and small arches (see Family Ferresidae
of Carter 1993, p. 68). Charlottea n. gen., is distinguished
from all other genera of the Subfamily Charlotteinae by
having three straight spines in the same plane.
102
Further remarks: The families Perispyridiidae, Ferresiidae
and the subfamily Charlotteinae are considered junior
synonyms of the family Eptingiidae because they have a
similar initial spicule, and three spines of the spicule extend
outside of the test (De Wever et al., 2001).
Etymology: Charlottea n. gen., is named for the British ship
Queen Charlotte, for which Captain George Dixon named
the Queen Charlotte Islands in August of 1787.
Included species:
CHA02 Charlottea amurensis Whalen & Carter 1998
CHA09 Charlottea hotaoensis Carter n. sp.
CHA10 Charlottea penderi Carter n. sp.
CHA03 Charlottea proprietatis Whalen & Carter 1998
CHA05 Charlottea triquetra Whalen & Carter 1998
CHA07 Charlottea sp. A sensu Whalen & Carter 2002
CHA08 Charlottea sp. B
CHA11 Charlottea sp. C
XNM01 Charlottea? sp. Y
Plate SUM03. Carterwhalenia minai (Whalen & Carter). Magnification x300. Fig. 1(H). Whalen & Carter 2002,
pl. 12, fig. 7. Fig. 2. Whalen & Carter 2002, pl. 12, fig. 9. Fig. 3. Whalen & Carter 2002, pl. 12, fig. 8.
Fig. 4a,b. TR, 1662D-R07-10. Fig. 5a,b. TR, 1662D-R01-08.
103
Charlottea amurensis Whalen & Carter 1998
Species code: CHA02
Synonymy:
? 1989 Acaeniotyle spp. – Hattori, pl. 1, fig. G.
1998 Charlottea amurensis n. sp. – Whalen & Carter, p. 37, pl. 2,
figs. 8, 9, 10; pl. 3, figs. 1, 2, 9.
2001 Charlottea amurensis Whalen & Carter – Gawlick et al.,
pl. 2, fig. 5.
2002 Charlottea amurensis Whalen & Carter – Suzuki et al.,
p. 168, figs. 4 H-J.
Original description: Test with medium-sized cortical shell
and three moderately long spines. Cortical shell subspherical
and somewhat compressed in plane of spines; cortical shell
composed of medium-sized, irregularly shaped, tetragonal
and triangular pore frames with prominent elongated
nodes at pore frame vertices; pore frame bars much thinner
in Y direction than Z direction (refer to Pl. 4, fig. 11 for
measurement system); large pores sometimes located on
cortical shell at base of spines. Internal spicular network
composed of delicate pore frames with no difference in
thickness between Y and Z direction and no apparent
pattern or orientation. Spines tapering distally, usually
shorter than diameter of cortical shell, triradiate in axial
section with narrow, rounded longitudinal ridges and
broad, rounded longitudinal grooves; spines evenly spaced
around cortical shell.
Original remarks: The more delicate, elongated nodes and
triradiate spines of Charlottea amurensis n. sp. distinguish
it from C. johnsoni n. sp.
Measurements (µm):
Based on 7 specimens.
Maximum diameter
of cortical shell
Maximum length
of primary spines
HT
Max.
Min.
Mean
146
150
138
147
105
120
75
95
Etymology: This species is named for Amur Rocks in Dana
Passage, located to the northwest of the type locality.
Type locality: Sample QC-676, Sandilands Formation,
Kunga Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands; Dürrnberg Formation, Austria; Pucara Group, Peru.
Charlottea hotaoensis Carter n. sp.
Species code: CHA09
Synonymy:
1989 Protoperispyridium ? spp. – Hattori, pl. 3, fig. G.
1997 Perispyridium sp. A02 – Yao, pl. 15, fig. 707.
2004 Perispyridium sp. – Matsuoka, fig. 184.
Type designation: Holotype GSC 111713 and paratype
GSC 111714 from GSC loc. C-304566; Rennell Junction member of the Fannin Formation (upper lower
Pliensbachian).
Description: Test with small cortical shell, triangular to
subtriangular in outline and three medium-sized spines
of equal length. Surface of cortical shell slightly convex
with large subtriangular to irregularly shaped pore frames
and deeply incised pores, sub-round in shape. Pore frame
vertices usually with small to medium sized nodes; some
groups of pore frames partly surrounded by massive raised
ridges. Spines assymetrically arranged; two spines closer
together than the third. Spines stout, triradiate and strongly
tapering with narrow ridges and wide grooves.
Remarks: Charlottea hotaoensis n. sp. differs from
C. triquetra Whalen & Carter in having a strongly triangular rather than subtriangular shell and less convex shell
104
surfaces. It is likely that C. hotaoensis n. sp. is derived from
C. triquetra. In our material, C. triquetra ranges from upper Sinemurian to basal Pliensbachian; C. hotaoensis n. sp.
does not range below the Pliensbachian. This new species
may also represent the transition between the two eptingiid
genera, Charlottea and Perispyridium.
Measurements (µm):
Based on 5 specimens.
Diameter of cortical shell
Length of longest spine
HT
132
80
Max.
153
126
Min.
123
56
Mean
132
93
Etymology: This species is named for the Haida village
of Hotao on the southwest side of Maude Island, Queen
Charlotte Islands, British Columbia.
Type locality: Sample 99-CNA-MI-9 (GSC loc. C-304566),
Rennell Junction member of the Fannin Formation; Fannin
Bay, south side of Maude Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Mino Terrane, Japan.
Plate CHA02. Charlottea amurensis Whalen & Carter. Magnification x200. Fig. 1(H)a,b. Carter et al. 1998,
pl. 2, figs. 8, 9.
Plate CHA09. Charlottea hotaoensis Carter n. sp. Magnification x200. Fig. 1(H). GSC loc. C-304566, GSC 111713.
Fig. 2. QCI, GSC loc. C-304566, GSC 128742. Fig. 3. QCI, GSC loc. C-175306, GSC 128743. Fig.4. QCI, GSC loc. C304566, GSC 111714. Fig. 5. QCI, GSC loc. C-305388, GSC 128744. Fig. 6. QCI, GSC loc. C-305388, GSC 128745.
Fig. 7. QCI, GSC loc. C-175311, GSC 128746.
105
Charlottea penderi Carter n. sp.
Species code: CHA10
Type designation: Holotype GSC 111715 from GSC loc.
C-080611, Ghost Creek Formation (lower Pliensbachian).
Paratype GSC 111716 from GSC loc. C-304566, Rennell
Junction member of the Fannin Formation (upper lower
Pliensbachian).
Description: Cortical shell generally subtriangular,
somewhat fluted and irregular in outline with convex
upper and lower surfaces and three medium-sized,
strongly tapering spines. Surface of shell double layered;
inner layer of pore frames variably-sized and irregular
in shape. Outer layer of pore frames extremely variable
in size (some quite enormous), irregular in shape; outer
layer of pore frames with very large raised nodes some
coalescing to form strong raised ridges. Pores subequal in
size, irregular in shape. Primary spines equally distributed
around shell; spines composed of very narrow ridges and
wide deep grooves.
Remarks: Charlottea penderi differs from Charlottea hotaoensis n. sp. in having an irregularly-shaped cortical
shell with more convex surfaces, large raised nodes some
forming ridges, and primary spines are more equally distributed around shell.
Measurements (µm):
Based on 6 specimens.
Diameter of cortical shell
Length of longest spine
HT
137
84
Max.
167
100
Min.
110
73
Mean
140
85
Etymology: This species is named after Captain Pender who
originally named Maude Island during the 1866 survey.
Type locality: Sample 99-CNA-MI-9 (GSC loc. C-304566),
Rennell Junction member of the Fannin Formation; Fannin
Bay, south side of Maude Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Ghost Creek Formation, and Rennell Junction
member of the Fannin Formation, Queen Charlotte Islands;
Fernie Formation, northeastern British Columbia.
Charlottea proprietatis Whalen & Carter 1998
Species code: CHA03
Synonymy:
1998 Charlottea proprietatis n. sp. – Whalen & Carter, p. 39,
pl. 4, figs. 1, 6, 7, 8, 9, 10, 13.
Original description: Test with large, inflated cortical shell
with three long, broad spines. Cortical shell subspherical
in shape and subrectangular in outline with irregularly
sized and shaped tetragonal and triangular pore frames
with prominent, elongated nodes at pore frame vertices; pore frame bars thin in Y direction, much thicker in
Z direction; larger pores usually located at base of spines.
Internal spicular network composed of thin, coarsely interwoven pore frames with no apparent pattern or orientation.
Spines triradiate in axial section with narrow, rounded longitudinal ridges and broad, rounded longitudinal grooves;
spines evenly spaced around cortical shell.
Original remarks: The distinctive subrectangular outline of
Charlottea proprietatis n. sp. distinguishes it from all other
species of Charlottea.
106
Measurements (µm):
HT Max. Min. Mean
Maximum diameter of cortical shell
180 180 150 161
(8 specimens measured)
Maximum length of primary spines
120 221 120 188
(5 specimens measured)
Etymology: Proprietas, atis (Latin; noun) = a property,
pecularity.
Type locality: Sample QC-675, Sandilands Formation,
Kunga Island - north side, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands.
Plate CHA10. Charlottea penderi Carter n. sp. Magnification x200. Fig. 1(H). GSC loc. C-080611, GSC 111715.
Fig. 2. NBC, GSC loc. C-305208, GSC 128747. Fig. 3. NBC, GSC loc. C-305208, GSC 128748. Fig. 4. GSC loc. C-304566,
GSC 111716. Fig. 5. QCI, GSC loc. C-080611, GSC 128749. Fig. 6. QCI, GSC loc. C-080612, GSC 128750.
Plate CHA03. Charlottea proprietatis Whalen & Carter. Magnification x200. Fig. 1(H). Carter et al. 1998, pl. 4, fig. 1.
107
Charlottea triquetra Whalen & Carter 1998
Species code: CHA05
Synonymy:
Original remarks: The shorter, less massive spines and less
prominent nodes of C. triquetra n. sp. distinguish it from
Charlottea sp. C.
Original description: Test with small, slightly elongated
cortical shell compressed in plane of spines and three
moderately long spines. Cortical shell with large, irregularly
shaped and sized pentagonal and hexagonal pore frames
with low, rounded nodes at pore frame vertices; pores
irregularly sized (large and small), usually subcircular in
outline. Internal spicular network composed of very delicate
pore frames with no pattern or orientation. Spines triradiate
in axial section with narrow, rounded longitudinal ridges
and broad, rounded longitudinal grooves; spines usually
shorter than or equal to long dimension of cortical shell;
two spines closer together than third, sometimes curving
inwards toward long axis of cortical shell.
Measurements (µm):
1998 Charlottea triquetra n. sp. – Whalen & Carter, p. 39, pl. 4,
figs. 2, 3, 4, 5, 11, 14, 15.
Charlottea sp. A sensu Whalen & Carter 2002
Species code: CHA07
Synonymy:
2002 Charlottea sp. A – Whalen & Carter, p. 122, pl. 7, figs. 4, 5.
Original remarks: The pronounced torsion of the short,
triradiate spines of Charlottea sp. A distinguishes it from
C. amurensis Whalen and Carter 1998 and C. harbridgensis
Whalen and Carter 1998.
Occurrence: San Hipólito Formation, Baja California Sur.
Charlottea sp. B
Species code: CHA08
Remarks: Cortical shell similar in size and shape to
C. amurensis Whalen & Carter, but spines are shorter and
more robust with thin rounded ridges and deep grooves;
distal ridges of spines with numerous small thorn-like
protuberances.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands; Baja California Sur.
108
Maximum diameter of cortical
shell (6 specimens measured)
Maximum length of primary
spines (5 specimens measured)
HT
Max.
Min.
Mean
75
90
75
84
71
75
68
73
Etymology: Triquetrus, a, um, (Latin, adj.) = three cornered,
triangular.
Type locality: Sample QC-677, Sandilands Formation,
Kunga Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands.
Plate CHA05. Charlottea triquetra Whalen & Carter. Magnification x300. Fig. 1(H)a,b. Carter et al. 1998, pl. 4, figs. 2, 3.
Plate CHA07. Charlottea sp. A sensu Whalen & Carter. Magnification x200. Figs. 1-2. Whalen & Carter 2002, pl. 7, figs. 4-5.
Plate CHA08. Charlottea sp. B. Magnification x200. Fig. 1. QCI, GSC loc. 304281, GSC 128740. Fig. 2. QCI, GSC loc.
305417, GSC 128741. Fig. 3. BCS, SH-412-14.
109
Charlottea sp. C
Species code: CHA11
Remarks: Cortical shell subtriangular in shape, multilayered.
Outer layer composed of densely packed triangular pore
frames with large highly raised nodes at vertices. Spines
short, thin and circular in axial section. Internal structure
unknown.
Occurrence: Rennell Junction member of the Fannin Formation, Queen Charlotte Islands; Fernie Formation, northeastern British Columbia.
Charlottea? sp. Y
Species code: XNM01
Synonymy:
1988 Tripocyclia (?) sp. A – Carter et al., p. 27, pl. 1, fig. 4.
Remarks: Large shell with three long straight spines. Shell
thick, comprised of several layers of latticed pore frames;
outer layer made up of medium-sized, irregularly shaped
and arranged pore frames (mostly triangular, tetragonal
and pentagonal); pore frames with small rounded nodes at
vertices. Spines very long, usually broken; spines triradiate
with wide rounded ridges and narrow deep grooves. Spines
prolonged internally to the edges of a small microsphere.
This species is questionably assigned to Charlottea
because pore frames on the thick-walled outer shell are
smaller, more irregularly arranged, and lack the typical
triangular pattern of Charlottea. Further study of the inner
structure is needed to confirm genus identity.
Occurrence: Fannin Formation, Queen Charlotte Islands.
110
Plate CHA11 Charlottea sp. C. Magnification x200. Fig. 1. QCI, GSC loc. C-304566, GSC 111791. Fig. 2. QCI, GSC loc.
C-304566-, GSC 111792. Fig. 3. NBC, GSC loc. C-305208, GSC 111793.
Plate XNM01. Charlottea? sp. Y. Magnification x 150. Fig. 1. QCI, GSC loc. 304566, GSC 128906. Fig. 2. QCI, GSC loc.
304566, GSC 128907. Fig. 3. QCI, GSC loc. 304566, GSC 128908. Fig. 4. QCI, GSC loc. 304567, GSC 128909.
Fig. 5. QCI, GSC loc. 304566, GSC 128910.
111
Genus: Citriduma De Wever 1982a
Type species: Citriduma radiotuba De Wever 1982a
Synonymy:
lower part giving a discoidal shape to this genus; thorax
bears lateral tubes.
Original description: Neosciadiocapsidae with a cephalic
structure including 8 spines (A, V, D, MB, Lr, Ll, lr and ll).
Location of cephalis on one side distinguishes the upper
and a lower face. Cephalis hemispherical, with an apical
horn and a lateral horn, respectively outgrowing from A
and V spines of the cephalic skeleton. Thorax, closed in the
Etymology: Anagram of P. Dumitrica (Bucarest), in honor
to his meticulous work upon Mesozoic radiolarians.
1982a Citriduma n. gen – De Wever, p. 202.
Included species:
4033 Citriduma hexaptera (Conti & Marcucci) 1991
CIT05 Citriduma radiotuba De Wever 1982a
Citriduma hexaptera (Conti & Marcucci) 1991
Species code: 4033
Synonymy:
1987 Gn. sp. indet. – Hattori, pl. 23, figs. 18.
? 1989 Gen. sp. indet. – Hattori, pl. 17, fig. L.
1991 Podocapsa (?) hexaptera n. sp. – Conti and Marcucci,
p. 803, pl. 3, figs. 12, 13, 14, 16, 17, 18.
1992 Unnamed 6-rayed livarellids – Yeh, pl. 3, fig. 6.
1995a Podocapsa (?) hexaptera Conti & Marcucci
– Baumgartner et al., p. 428, pl. 4033, figs. 1-5.
1997 Citriduma sp. A – Yao, pl. 12, fig. 558.
2003 Citriduma hexaptera (Conti & Marcucci) – Goričan et al.,
p. 297, pl. 5, fig. 4.
2004 Citriduma hexaptera (Conti & Marcucci) – Matsuoka,
fig. 128.
Original description: The shell shows two distinct parts:
a hemispherical small proximal part without apparent
segmental division but possibly including cephalis and
thorax, and a large and flat abdomen with six porous wings,
lacking a terminal tube. None of the available specimens
show a horn on the proximal part referable to cephalis.
The proximal part presents loosely scattered pores smaller
than those of the abdomen and wings. The abdomen shows
circular uniformly distributed pores. Six conical wings are
seated along the equatorial zone of the abdomen: they show
pores similar to those of abdomen.
Original remarks: This species differs from P. amphitreptera
in the flat rather than globose shape of the distalmost
112
segment, in having six equatorial wings and lacking a
terminal tube. Its tentative assignment to genus Podocapsa
is based on the presence of porous wings and of a broad
distalmost segment.
Further remarks: By Goričan et al. (2003): Citriduma
hexaptera differs from the Rhaetian Citriduma asteroides
Carter, 1993 and Citriduma sp. C (Carter, 1993) by having
a constant number of tubes (always six), longer tubes and
smaller pores.
Measurements (µm):
Based on 5 specimens.
Length wings
Width wings
Diameter abdomen
Min.
90
30
125
Max.
160
50
200
HT
100
35
135
Etymology: Greek, hexa = six plus pteron = wing.
Type locality: Ponte di Lagoscuro, Liguria, Italy.
Occurrence: Liguria, Italy; Skrile Formation, Slovenia;
Japan; Warm Springs member of the Snowshoe Formation,
Oregon; Tawi Sadh Member of the Guwayza Formation,
Oman.
Plate 4033. Citriduma hexaptera (Conti & Marcucci). Magnification x200. Fig. 1(H). Conti & Marcucci 1991, pl. 3,
fig. 12. Fig. 2. Matsuoka 2004, fig. 128. Fig. 3. OM, BR706-R12-16. Fig. 4. Goričan et al. 2003, pl. 5, fig. 4.
113
Citriduma radiotuba De Wever 1982a
Species code: CIT05
Synonymy:
1982a Citriduma radiotuba n. sp. – De Wever, p. 202, pl. 8,
figs. 10-11; pl. 9, figs. 1-8.
1982b Citriduma radiotuba De Wever – De Wever, p. 285, pl. 39,
figs. 1-5; pl. 40, figs. 1-7.
Original description: Discoidal dicyrtid fringed by numerous radial tubes. Location of cephalis on one side
distinguishes the upper and a lower face. Cephalis hemispherical with an apical horn and a lateral horn, respectively outgrowing from A and V spines of the cephalic
skeleton. Cephalis imperforate, covered with numerous
spiny irregularities.
Thorax forms the largest part of the test. It is a biconvex
disc laterally extended by twelve to twenty tubes. These
tubes are situated on prolongation of ridges which originate at the base of the cephalis and may sometimes protrude outside the tubes (Pl. 9, fig. 3). Tubes are distally
narrower and terminally closed, ending in a point.
One specimen shows, on the upper side of the disc near
the cephalis and under the lateral spine, a horizontal opening whose significance is unknown. Near the cephalis, the
thoracic wall is made of two latticed layers, closely linked
by small pillars. Distally, these two latticed layers are interconnected and constitute a two-layered wall.
Interior of thorax hollow in the center but pillars exist
between the upper and the lower face laterally. These pillars
are in line with the radial ridges that are visible outside and
inside the test.
Cephalis has spines typical of nassellarians; A and
V extend outside as horns. D, Ll, Lr, ll and lr extended
peripherally.
Measurements (µm):
Based on 6 specimens.
Width of thorax
including tubes
609
609
333
468
Width of thorax
without tubes
418
418
233
307
HT
Max.
Min.
Mean
Etymology: From the Latin radiare = to radiate, and tuba,
-ae = tube, pipe.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Fannin Formation, Queen Charlotte Islands.
Genus: Crubus Yeh 1987b
Type species: Crubus chengi Yeh 1987b (subsequent designation by Carter, in Carter et al., 1988)
Synonymy:
1987b Crubus n. gen – Yeh, p. 69.
1988 Crubus Yeh – Carter et al., p. 53.
Original description: Test as with family, conical to
subcylindrical, with constrictions between joints. Cephalis
conical, with horn. Cephalis and thorax usually sparsely
perforate, covered with layer of microgranular silica. Outer
layer of abdomen and first one or two post-abdominal
chambers covered with small irregular polygonal pore
frames, remaining chambers with costae superimposed
between each row of pore frames.
Original remarks: Crubus, n. gen., differs from Drulantus,
n. gen., by having a horn, and by having a lobated test with
outer layer of test wall comprised of small irregular polygonal pore frames on the apical portion of the test.
114
Further remarks: By Carter et al. (1988): Yeh (1987)
designated Crubus robustus Yeh, 1987 as the type species
of the genus Crubus. However, in the original description
of this species and in all Yeh's subsequent references to this
taxon, the binominal name appears as Crubus (?) robustus.
In the original description Yeh states »this species is
questionably assigned to Crubus n. gen. because it lacks
a horn«. Querying generic assignment of a type species
invalidates that species as type (see Article 67C, ICZN,
1985). A new type species is herein designated that more
clearly conforms to the generic description and is better
documented.
Included species:
CRB01 Crubus chengi Yeh 1987b
Plate CIT05. Citriduma radiotuba De Wever. Magnification x150 (scale bar A) except Figs. 3b-3c x500 (scale bar B).
Fig. 1(H)a,b. De Wever 1982a, pl. 9, figs. 1, 3. Fig. 2. TR, 1662D-R02-05. Fig. 3a,b,c. TR, 1662D-R02-05.
115
Crubus chengi Yeh 1987b
Species code: CRB01
Synonymy:
1987b Crubus chengi n. sp. – Yeh, p. 69, pl. 18, figs. 13-15, 19-20,
24; pl. 19, figs. 7, 15.
1987b Crubus firmus n. sp. – Yeh, p. 69, pl. 18, figs. 12, 18.
1987b Crubus (?) robustus n. sp. – Yeh, p. 70, pl. 3, fig. 18; pl. 18,
figs. 9, 10, 22.
1987b Crubus sp. A – Yeh, p. 70, pl. 18, figs. 11, 17.
Original description: Test wide, subcylindrical, usually
with eight to ten post-abdominal chambers. Cephalis conical, with short massive rudimentary horn and covered with
layer of microgranular silca. Thorax to second post-abdominal chambers covered by layer of massive, irregular
polygonal pore frames. Costae moderately thick, about ten
to twelve visible laterally.
Original remarks: Crubus chengi Yeh, n. sp., differs from
C.(?) robustus, n. sp., by having a long, massive horn, and
having a test with apical portion more pointed in nature.
Crubus chengi, n. sp., differs from C. firmus, n. sp., by
having a more cylindrical test with more massive horn on
a smaller cephalis.
Further remarks: We consider that Crubus chengi,
C.? robustus, C. firmus, and C. sp. A represent variation in
a single species.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length (max.)
287
280
287
272
Width (max.)
143
140
143
130
Etymology: This species is named for Dr. Yen-Nien Cheng
for his help on this project.
Type locality: Sample OR-600M, Hyde Formation at IzeePaulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, Oregon; Fannin
Formation, Queen Charlotte Islands.
Genus: Crucella Pessagno 1971
Type species: Crucella messinae Pessagno 1971
Synonymy:
1971 Crucella n. gen. – Pessagno, p. 52.
Original description: Test as with subfamily. Four rays,
elliptical to rectangular in cross-section with polygonal
meshwork arranged linearly to sublinearly; rays equal in
length; tapering distally; terminating in centrally placed
spines. Central area with polygonal (often triangular)
meshwork; sometimes with a lacuna, with or without
patagium.
Original remarks: Crucella n. gen., differs from Hagiastrum
Haeckel (1) by possessing rays of nearly equal length; (2) by
possessing rays with tapered rather than bulbous tips; and
(3) by having prominent spine at the tip of each ray.
Etymology: From the Latin crux = cross.
116
Included species and subspecies:
CRU21 Crucella angulosa s.l. Carter 1988
CRU11 Crucella angulosa angulosa Carter 1988
CRU12 Crucella angulosa longibrachiata Carter n. ssp.
PDC02 Crucella beata (Yeh) 1987b
CRU22 Crucella cavata s.l. Whalen & Carter 1998
CRU10 Crucella cavata cavata Whalen & Carter 1998
CRU20 Crucella cavata giganticava Carter n. ssp.
CRU19 Crucella cavata intermedicava Carter n. ssp.
PDC05 Crucella jadeae Carter & Dumitrica n. sp.
CRU13 Crucella mijo De Wever 1981b
CRU14 Crucella mirabunda Whalen & Carter 2002
CRU15 Crucella spongase De Wever 1981b
CRU16 Crucella squama (Kozlova) 1971
3131 Crucella theokaftensis Baumgartner 1980
Plate CRB01. Crubus chengi Yeh. Magnification x200. Fig. 1(H). Yeh 1987b, pl. 18, fig. 15. Fig. 2. QCI, GSC loc. C304566, GSC 128752. Fig. 3. QCI, GSC loc. C-304567, GSC 128753. Fig. 4. QCI, GSC loc. C-304566, GSC 128754.
117
Crucella angulosa s.l. Carter 1988
Species code: CRU21
Synonymy:
1988 Crucella angulosa Carter n. sp. – Carter et al., p. 43, pl. 4,
figs. 11, 12.
See also subspecies.
Included subspecies:
CRU11 Crucella angulosa angulosa Carter 1988
CRU12 Crucella angulosa longibrachiata Carter n. ssp.
Crucella angulosa angulosa Carter 1988
Species code: CRU11
Synonymy:
1988 Crucella angulosa Carter n. sp. – Carter et al., p. 43, pl. 4,
fig.11 only.
1998 Pseudocrucella carpenterensis n. sp. - Cordey, p. 69, pl. 19,
figs. 3, 4.
Original diagnosis: Test cruciform. Rays medium to long
and of uniform width, with long sturdy central spines.
Original description: Test cruciform with medium to long
rays terminated by long central spines. Rays uniform in
width, of more or less equal length, diverging abruptly from
the central area. Pore frames irregular in size, shape and
arrangement; composed of thin bars with small rounded
pores at vertices. Rays rectangular in cross-section. Central
spines have (three?) wide, rounded, longitudinal ridges
alternating with wide, strong grooves.
Original remarks: This form, although extremely variable
in ray length, bears no resemblance to any described species of Crucella. Indeed, the longer-rayed forms (e.g., Pl. 4,
fig. 12), having finer bars and smaller nodes, may be found
to represent another species when additional, better preserved specimens are found.
Measurements (µm):
Based on 10 specimens.
Length of ray
Width of ray
Length of longest spine
HT
111
39
95
Av.
124
51
71
Max.
191
66
119
Min.
111
39
45
Etymology: Latin, angulosus (adj.), full of corners.
Type locality: GSC locality C-080577, Fannin Formation,
Maude Island.
Occurrence: Ghost Creek, Fannin, Whiteaves and Phantom
Creek formations, Queen Charlotte Islands; Fernie Formation, Williston Lake, NE British Columbia; Bridge River
Complex, British Columbia.
Crucella angulosa longibrachiata Carter n. ssp.
Species code: CRU12
Synonymy:
1988 Crucella angulosa Carter n. sp. – Carter et al., p. 43, pl. 4,
fig. 12, not fig. 11.
Type designation: Holotype GSC 111717 from GSC loc. C305208, Fernie Formation (lower Pliensbachian).
Measurements (µm):
Based on 9 specimens.
Length of longest ray
Max. width of ray tips
Length of longest spine
HT
126
65
203
Max.
158
75
214
Min.
112.5
47
150
Mean
136
59
221 (7)
Description: Test cruciform with long rays terminated by
long central spines. Rays uniform in width, of more or
less equal length. Pore frames irregular in size, shape and
arrangement; composed of thin bars with small rounded
pores at vertices. Rays rectangular in cross-section. Central
spines with three wide, rounded, longitudinal ridges alternating with wide, strong grooves.
Etymology: From the Latin: longus, -a, -um = long, and
brachium, i = arm; longibrachiatus, -a,- um = with long
arms; adjective.
Remarks: This species differs from C. angulosa angulosa in
always having much longer arms. Crucella angulosa Carter
originally consisted of two distinctive morphotypes: one
with short arms and one with much longer arms; these are
now separated into subspecies.
Occurrence: Fernie Formation, Williston Lake, NE British
Columbia; Fannin Formation, Queen Charlotte Islands.
118
Type locality: Sample 00-TD-HALL (GSC loc. C-305208),
Fernie Formation, Black Bear ridge, Williston Lake, British
Columbia
Plate CRU11. Crucella angulosa angulosa Carter. Magnification x150. Fig. 1(H). Carter et al. 1988, pl. 4, fig. 11.
Fig. 2. QCI, GSC loc. C-080611, GSC 128757. Fig. 3. QCI, GSC loc. C-080611, GSC 128755. Fig. 4. QCI, GSC loc. C304566, GSC 128756. Fig. 5. NBC, GSC loc. C-305208, GSC 128758.
Plate CRU12. Crucella angulosa longibrachiata Carter n. ssp. Magnification x150. Fig. 1(H). NBC, GSC loc. C-305208,
GSC 111717. Fig. 2. Carter et al. 1988, pl. 4, fig. 12.
119
Crucella beata (Yeh) 1987b
Species code: PDC02
Synonymy:
1987b Pseudocrucella beata n. sp. – Yeh, p. 28, pl. 2, figs. 11-12;
pl. 23, figs. 10, 25.
Original description: Test thick with four wide rays. Rays
subellipsoidal in cross section, medium in length, with
five to six external beams and four to five sublinearly arranged rows of rectangular pore frames. Central area large
with irregularly arranged subtriangular or rectangular pore
frames. Primary spines medium in length, moderately
thick, circular in axial section. Test with or without patagium.
Original remarks: Pseudocrucella beata, n. sp., differs from
P. jurassica, n. sp., by having a wider test with rays which
are ellipsoidal in axial section and by lacking a central
cavity.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length
of ray
108
105
108
103
Width of ray
Width
at base
of central area
84
130
82
131
84
134
80
130
Length
of spine
86
88
103
83
Etymology: Beatus-a-um (Latin, adj.) = happy.
Type locality: Sample OR-536J, Nicely Formation, southeast side of Morgan Mountain, east-central Oregon.
Occurrence: Nicely Formation, Oregon; Fannin Formation,
Queen Charlotte Islands.
Crucella cavata s.l. Whalen & Carter 1998
Species code: CRU22
Synonymy:
See subspecies.
Included subspecies:
CRU10 Crucella cavata cavata Whalen & Carter 1998
CRU20 Crucella cavata giganticava Carter n. ssp.
CRU19 Crucella cavata intermedicava Carter n. ssp.
Crucella cavata cavata Whalen & Carter 1998
Species code: CRU10
Synonymy:
1998 Crucella cavata n. sp. – Whalen & Carter, p. 49, pl. 12,
figs. 15, 18, 19, 21, 22.
Original description: Test with large central area, four short
rays expanded distally, each with a moderately long central
spine. Rays wide, subrectangular in axial section with upper and lower planiform surfaces. Rays gradually widening
to ray tips. Each ray with one long, massive spine, circular
in axial section. Rays with irregularly sized and shaped polygonal pore frames with no development of external lineation; small nodes at pore frame vertices. Prominent circular lacuna in central area variable in size with sides sloping
towards center.
Original remarks: The distinctive lacuna distinguishes
Crucella cavata n. sp. from all other species of Crucella
Pessagno in the Sandilands fauna.
120
Measurements (µm):
Number of specimens measured = (n)
Length of Maximum width Width of ray Length of
longest
of central area
tips (Max.)
longest
ray (12)
(12)
(12)
spine (7)
124
82
97
111
225
82
122
150
124
54
79
94
180
69
101
128
HT
Max.
Min.
Mean
Etymology: Cavatus, a, um (Latin; adj.) = hollowed out.
Type locality: Sample 86-OF-KUC-8, Sandilands Formation, north side of Kunga Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte
Islands.
Plate PDC02. Crucella beata (Yeh). Magnification x200. Fig. 1(H). Yeh 1987b, pl. 23, fig. 10. Fig. 2. QCI, GSC loc. C080611, GSC 128759.
Plate CRU10. Crucella cavata cavata Whalen & Carter. Magnification x150. Fig. 1(H). Carter et al. 1998, pl. 12, fig. 15.
121
Crucella cavata giganticava Carter n. ssp.
Species code: CRU20
Type designation: Holotype GSC 111718 from GSC loc. C140413; Rennell Junction member of the Fannin Formation
(upper lower Pliensbachian).
Description: Large cruciform test with massive deep lacuna
and a short central spine on each ray tip. Rays wide, subrectangular in axial section, upper and lower surfaces planiform. Rays slightly expanded towards tips; tips rounded,
not tapering. Pore frames surrounding and within central
lacuna large, mostly triangular in shape with moderately
large nodes at vertices; pore frames smaller towards ray
tips. Lacuna occupying most the central area of test. Primary spines at ray tips short and rod-like.
Remarks: Crucella cavata giganticava n. ssp. differs from C.
cavata cavata in having much larger, more regularly shaped
pore frames with stronger nodes at vertices, and primary
spines are smaller. Lacuna in C. cavata giganticava n. ssp.
larger than in all other subspecies.
Measurements (µm):
Based on 6 specimens.
Length of longest ray
Max. width of ray tips
Max. width of central cavity
Length of longest spine
HT
153
111
116
broken
Max.
184
135
116
111
Min.
111
80
56
58
Mean
148
102
78
78 (3)
Etymology: From Latin: giganteus, -a, -um = giant, gigantic
and cavus, -a, -um = caved; adjective.
Type locality: North side Cumshewa Inlet, Moresby Island,
Queen Charlotte Islands, British Columbia.
Occurrence: Ghost Creek Formation and Rennell Junction
member of the Fannin Formation, Queen Charlotte
Islands.
Crucella cavata intermedicava Carter n. ssp.
Species code: CRU19
Type designation: Holotype GSC 128888 from GSC loc. C080612; Ghost Creek Formation (lower Pliensbachian).
Description: Test with relatively small central area and
four moderately long rays each with a strong central
spine. Rays constant in width or just slightly expanded
distally, subrectangular in axial section with upper and
lower surfaces planiform. Each ray with one massive spine
(usually broken), triradiate in axial section. Rays with
small irregularly arranged pore frames, mostly triangular
and tetragonal in shape; small nodes at pore frame vertices.
Lacuna in central area variable in size, deep with steep
sides.
Remarks: Crucella cavata intermedicava Carter n. ssp.
differs from C. cavata cavata Whalen & Carter in possessing
slimmer rays with slightly larger, more regularly arranged
pore frames, and spines are triradiate rather than circular
in axial section. Differs from C. cavata giganticava n. ssp.
in having narrower rays, less massive pore frames, and a
smaller lacuna.
122
Measurements (µm):
Based on 7 specimens.
Length of longest ray
Max. width of ray tips
Max. width of central area
HT
108
72
55
Max.
168
72
61
Min.
112
47
37
Mean
133
62
48
Etymology: From Latin: intermedius, -a, -um = intermediate
and cavus, -a, -um = caved; adjective.
Type locality: Sample CAA-80-T-7, lms. (GSC loc. C080612), Ghost Creek Formation, Rennell Junction, Yakoun River area, central Graham Island, Queen Charlotte
Islands.
Occurrence: Ghost Creek Formation, Queen Charlotte
Islands; Musallah Formation, Oman.
Plate CRU20. Crucella cavata giganticava Carter n. ssp. Magnification x150. Fig. 1(H). QCI, GSC loc. C-140413, GSC
111718. Fig. 2. QCI, GSC loc. C-304566, GSC 128760. Fig. 3. QCI, GSC loc. C-304566, GSC 128761.
Plate CRU19. Crucella cavata intermedicava Carter n. ssp. Magnification x150. Fig. 1(H). QCI, GSC loc. C- 080612,
GSC 128888. Fig. 2. QCI, GSC loc. C-175311, GSC 128762. Fig. 3. OM-00-251, 021518.
123
Crucella jadeae Carter & Dumitrica n. sp.
Species code: PDC05
Synonymy:
Whalen & Carter but differs in having longer arms and
mostly triradiate rather than circular spines.
Type designation: Holotype, pl. PDC05, fig. 1 (Yeh 1987b,
pl. 2, fig. 18); paratype, fig. 2, GSC 128877; Ghost Creek
Formation, Queen Charlotte Islands.
Measurements (µm):
Based on 6 specimens.
1987b Pseudocrucella sp. E – Yeh, p. 30, pl. 2, fig. 18; pl. 3, fig. 14.
Description: Test flat with long and narrow rays; rays
increasing very slowly in width distally and terminating in
more or less expanded tips. Each ray with a three-bladed,
pointed distal spine the length of which is about half the
length of ray. Central area flat, small. Surface of rays and
central area with small, dense, irregularly arranged pores.
Sides of rays vertical to slightly concave.
Remarks: As written by Yeh (1987b) this form differs from
Pseudocrucella magna Blome (1984b) by having rays with
expanded tips and with less massive spines. Crucella jadeae
is very close to the Sinemurian species C. kaisunensis
Length of rays
Width of rays at base
With of rays at tip
HT
170-200
50
70-88
Max.
390
70
140
Min.
180
50
68
Mean
230
56
91
Etymology: The species is named for Kuei-Yu Yeh (Jade)
who illustrated the first specimens.
Type locality: OR-536J, southeast side of Morgan Mountain, east-central Oregon.
Occurrence: Nicely Formation, east-central Oregon; Ghost
Creek Formation, Queen Charlotte Islands; Tawi Sadh
Member of the Guwayza Formation, Oman.
Crucella mijo De Wever 1981b
Species code: CRU13
Synonymy:
1981b Crucella mijo n. sp. – De Wever, p. 35, pl. 4, figs. 1, 2.
1982b Crucella mijo De Wever – De Wever, p. 253, pl. 28,
figs. 1-3.
1996 Crucella sp. A – Pujana, p. 136, pl. 1, fig. 14.
2002 Crucella mijo De Wever – Suzuki et al., p. 176, fig. 7 C.
Original description: Patulibracchiinae with four wineskin-shaped arms terminating in a spine triradiate in axialsection along its length. Fine spongy network visible, in a
relic stage, mainly at base of arms and between them where
it is probably more protected. Here and there one or several
tiny spines arise radially from this web (Pl. 4, fig. 1). At base
of terminal spine on arms, ridges, which separate grooves
(corresponding to prolongation of a pore), are carved by
secondary grooves. Pores tend to be disposed in an orthogonal network in center, oblique relative to axis of arms,
and sometimes slightly aligned on arms. Nodes are often
present at intersection of bars on the network.
Original remarks: This species differs from Crucella messinae Pessagno (1971, p. 55) by its wineskin-shaped arms and
spines triradiate along their length with grooves carved at
base. It is distinguished from C. plana Pessagno (1971,
p. 56) and C. espartoensis Pessagno (1971, p. 54) which
have different pores, less visible nodes and primary spines
rounded in cross-section; from C. irwini Pessagno (1971,
124
p. 55) by the shape of the arms. Stauralastrum euganeum
Squinabol (1903, p. 123) has a more massive shape and
does not show aligned pores.
Measurements (µm):
Based on 8 specimens.
Overall length (2 arms +
center, without spines)
Maximal width of arm
Diagonal of central part
Av.
Min.
Max.
HT
366
350
380
380
100
115
85
85
116
140
110
85
Length of terminal spines sometimes reach 115 µm, as on
holotype.
Etymology: Arbitrary combination of letters (ICZN, Append. D, V, 26)
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Rennell Junction member of the Fannin Formation; Fernie Formation,
NE British Columbia; Sierra Chacaicó Formation, Argentina; Pucara Group, Peru.
Plate PDC05. Crucella jadeae Carter & Dumitrica n. sp. Magnification x100. Fig. 1(H). Yeh 1987b, pl. 2, fig. 18.
Fig. 2. QCI, GSC loc. C-305388, GSC 128877. Fig. 3. QCI, GSC loc. C-305386, GSC 111807. Fig. 4. OM, BR524-R04-07.
Fig. 5. OM, BR523-R03-19.
Plate CRU13. Crucella mijo De Wever. Magnification x150. Fig. 1(H). De Wever 1981b, pl. 4, fig. 2. Fig. 2. QCI, GSC loc.
C-304566, GSC 128763. Fig. 3. NBC, GSC loc. C-305208, GSC 128764. Fig. 4. QCI, GSC loc. C-304566, GSC 128765.
125
Crucella mirabunda Whalen & Carter 2002
Species code: CRU14
Synonymy:
1987 Crucella sp. B - Hattori, pl. 4, fig. 8.
1988 Pseudocrucella sp. A – Carter et al., p. 29, pl. 7, figs. 8-9.
2002 Crucella mirabunda n. sp. – Whalen & Carter, p. 106, pl. 1,
figs. 7, 11; pl. 2, figs. 1, 8.
Original description: Rays with irregularly sized and
shaped triangular and tetragonal pore frames with no linear
arrangement; pore frames with medium-sized nodes at
vertices. Rays subcircular in axial section gradually widening
toward distal part of ray. Each ray with one massive spine,
triradiate in axial section with broad, rounded longitudinal
ridges and narrow longitudinal grooves. Central area broad,
flat with no lacuna; pore frames in central area with similar
construction and arrangement as rays.
Original remarks: Crucella mirabunda n. sp. differs from
C. kaisuensis Whalen and Carter 1998 by having shorter
triradiate spines; it differs from C. mijo De Wever 1981b by
having shorter, broader rays and more robust spines.
Measurements (µm):
Based on 10 specimens.
Length of ray
(Max.)
105
120
83
98
Length of spine
when entire (Max.)
75
116
60
88
HT
Max.
Min.
Mean
Etymology: Mirabundus, a, um (Latin, adj.) = full of
wonder.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Phantom Creek Formation, Queen Charlotte Islands;
Japan.
Crucella spongase De Wever 1981b
Species code: CRU15
Synonymy:
1981 Crucella sp. A – Pessagno & Poisson, pl. 2, figs. 6, 8.
1981b Crucella spongase n. sp. – De Wever, p. 36, pl. 5, figs. 1-3.
1982b Crucella spongase De Wever – De Wever, p. 254, pl. 29,
figs. 1-3.
Original description: Patulibracchiinae with four arms orthogonally disposed, each ending in a spine, with a patagium. Arms massive, sometimes bearing very small spines,
and covered with a thin spongy network, often residual.
Terminal spines triradiate in cross section, but one of them
without a 3 part symmetry in cross section. Indeed, one of
the basal pores, open in a groove, is much larger than the
other two. This large pore suggests a bracchiopyle (Pl. 5,
fig. 2).
Arms inflated in center of test, which is flat, and does
not show a depressed part (lacuna). Network is looser
in center, where pores are triangular, than distally where
pores are rectangular. Nodes exist at bar intersections.
126
Original remarks: This form differs from Histiastrum
valanginica Aliev (1965) by presence of secondary spines,
irregularly arranged pores without alignment, and absence
of lacuna.
Measurements (µm):
10 specimens measured.
Total length of both arms
without terminal spines
Length of primary spines
HT
Max.
Min.
Mean
336
400
297
336
59
32
59
47
Etymology: Anagram of E. A. Pessagno Jr. in honor of his
pioneer work on Mesozoic Radiolaria.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Dürrnberg
Formation, Austria.
Plate CRU14. Crucella mirabunda Whalen & Carter. Magnification x200. Fig 1(H). Whalen & Carter 2002, pl. 1, fig. 7.
Fig. 2. Whalen & Carter 2002, pl. 2, fig. 1.
Plate CRU15. Crucella spongase De Wever. Magnification x150. Fig. 1(H). De Wever 1981b, pl. 5, fig. 3.
Fig. 2. AT, BMW-21-24.
127
Crucella squama (Kozlova) 1971
Species code: CRU16
Synonymy:
1971 Hagiastrum squama n. sp. – Kozlova, p. 1175, pl. 1, fig. 10.
1973 Hagiastrum squama n. sp. – Kozlova, p. 59, pl. 18, fig. 8.
1981b Crucella squama (Kozlova) – De Wever, p. 38, pl. 5, fig. 7.
1982b Crucella squama (Kozlova) – De Wever, p. 255, pl. 29, fig. 4.
1988 Crucella sp. aff. C. squama (Kozlova) – Carter et al., p. 43,
pl. 12, figs. 11, 12.
2002 Crucella squama (Kozlova) – Whalen & Carter, p. 106,
pl. 2, figs. 2, 5.
Original description: Skeleton small, cross-shaped, thick.
Four similar, short, triangular arms ending in thick, threebladed spines. Structure of central part of the disk and arms
similar, skeleton consisting of several porous layers that
pass in one another. Pores rounded, oval and reniform.
Original remarks: By the outline of skeleton and shape
of arms Hagiastrum squama Kozlova, sp. nov. is close to
Stauralastrum (?) sp. Holmes from which it differs only in
having less long arms; the comparison with this species is
difficult because due to the poor preservation of specimens
W. M. Holmes did not describe the structure of skeleton.
Further remarks: Crucella squama is very similar to Crucella beata (Yeh) but differs from the latter in having threebladed spines.
Measurements (µm):
Length of arms from centre (without spines)
width of arms at base
Width of spines at base
Diameter of pores
Min.
105
75
6
Max
120
105
27
12
Etymology: Squama (Lat.) – scale.
Type locality: Sample Timano-Ural region, Pizhma river,
Lower Kimmeridgian Marls.
Occurrence: Timano-Ural region, Russia; Gümüslü Allochthon, Turkey; San Hipólito Formation, Baja California Sur;
Fannin, Whiteaves and Phantom Creek formations, Queen
Charlotte Islands.
Crucella theokaftensis Baumgartner 1980
Species code: 3131
Synonymy:
1980 Crucella theokaftensis n. sp. – Baumgartner, p. 308, pl. 8,
figs 19-22; pl. 12, fig. 1.
1982 Crucella theokaftensis Baumgartner – Aita, pl. 3, fig. 12.
? 1985 Crucella theokaftensis Baumgartner – Nagai, pl. 5,
figs. 5, 5a.
1987 Crucella theokaftensis Baumgartner – Aita, p. 63, pl. 1,
fig. 8; pl. 8, fig. 3.
1987 Crucella theokaftensis Baumgartner – Kito, pl. 1, fig. 10.
1989 Crucella sp. A – Hattori, pl. 25, fig. G.
1995a Crucella theokaftensis Baumgartner – Baumgartner et al.,
p. 158, pl. 3131, figs. 1-3.
1997 Crucella theokaftensis Baumgartner – Hull, p. 20, pl. 4,
figs. 6, 12, 14.
2003 Crucella theokaftensis Baumgartner – Goričan et al.,
p. 293, pl. 1, fig. 19.
2004 Crucella theokaftensis Baumgartner – Matsuoka, fig. 44.
Original description: Test as with genus, central area inflated subspherical on both sides raised over rays. Rays
slender conical tapering into long triradiate spines. Central
area with small, irregular pore frames, ray with lengthened
pores becoming larger toward the base of the spines, sometimes weakly linearly arranged.
128
Original remarks: This species is related to C. messinae but
differs in having an inflated central area with smaller pores
and slenderer conical rays. The specimen from the lowest
sample of the Argolis Peninsula POB 899 (pl. 8. fig. 19; pl.
12, fig. 1) differs from the topotypic material (POB 986) in
having much shorter spines and a smaller test; see measurements.
Measurements (µm):
Based on 7 specimens.
Length of rays AX
Length of rays BX
Length of rays CX
Length of rays DX
Width of rays at base
L. longest spine
HT
140
210
200
70
150
Av.
119
65
61
Min.
97
50
50
Max.
210
80
150
Etymology: Named for the type locality.
Type locality: Locality D of Baumgartner (1980); Argolis
Peninsula (Peloponnesus, Greece).
Occurrence: Worldwide.
Plate CRU16. Crucella squama (Kozlova). Magnification x150. Fig. 1(H). Kozlova 1971, pl. 1, fig. 10.
Fig. 2. De Wever 1981b, pl. 5, fig. 7. Fig. 3. Whalen & Carter 2002, pl. 2, fig. 2.
Plate 3131. Crucella theokaftensis Baumgartner. Magnification x200. Fig. 1(H). Baumgartner 1980, pl. 8, fig. 22.
Fig. 2. Matsuoka 2004, fig. 44. Fig. 3. Goričan et al. 2003, pl. 1, fig. 19.
129
Genus: Cyclastrum Rüst 1898
Type species: Cyclastrum infundibuliforme Rüst 1898
Synonymy:
1898 Cyclastrum n. gen – Rüst, p. 28.
Original description: Three rays are linked at their distal
ends by a band of patagium.
Included species:
CYC01 Cyclastrum asuncionense Whalen & Carter 2002
CYC02 Cyclastrum scammonense Whalen & Carter 2002
CYC03 Cyclastrum veracruzense Whalen & Carter 2002
CYC04 Cyclastrum sp. A
Cyclastrum asuncionense Whalen & Carter 2002
Species code: CYC01
Synonymy:
2002 Cyclastrum asuncionense n. sp. – Whalen & Carter, p. 110,
pl. 4, figs. 8, 9, 14; pl. 17, fig. 1.
Original description: Test subtriangular in outline, margins
gently convex between peripheral spines; test thin with
rounded edges. Spines medium-sized, slender, triradiate in
axial section with rounded, longitudinal ridges, and grooves.
Meshwork along triangular margin of test composed of
medium sized tetragonal and pentagonal pore frames with
no distinctive alignment. Central portion of test depressed,
broad, triangular in outline, with three-rayed area defined
by smaller, more delicate pore frames than on remainder of
test; three-rayed structure raised above central cavity and
each ray aligned with peripheral spines.
distinguishes C. asuncionense from all other species of
Cyclastrum.
Measurements (µm):
(n) = number of specimens measured.
Diameter of
cortical shell (Max.) (8)
236
274
225
245
Length of
primary spine (Max.) (6)
90
90
41
60
HT
Max.
Min.
Mean
Etymology: This species is named for Punta Asuncion
located to the northwest of the type area.
Original remarks: See remarks under Cyclastum veracruzense n. sp. and Cyclastrum scammonense n. sp.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur, Mexico.
Further remarks: The shape of the test, moderately
compressed, subtriangular in outline with rounded edges,
Occurrence: San Hipólito Formation, Baja California Sur.
Cyclastrum scammonense Whalen & Carter 2002
Species code: CYC02
Synonymy:
? 1998 Orbiculiforma silicatilis n. sp. – Cordey, p. 93, pl. 21, fig. 7
(not figs. 5, 8).
2002 Cyclastrum scammonense n. sp. – Whalen & Carter, p. 111,
pl. 4, figs. 3-5, 11-13, 15; pl. 5, figs. 1, 2, 9.
Original description: Test outline a nearly straight-sided
equilateral triangle; test very thick with vertical sides. Recessed area sometimes girdles edge of test. Spines mediumsized, triradiate in axial section with broad, rounded longitudinal ridges and narrow longitudinal grooves, becoming
circular in axial section distally. Pore frames irregularly
shaped, pentagonal, tetragonal and circular showing an
indistinct lineation subparallel to margin of test. Central
area of test broad, triangular in outline and slightly depressed. Subspherical area, in center of test, with smaller
pore frames, connecting with poorly defined rays aligned
with peripheral spines.
Original remarks: Cyclastrum scammonense n. sp. is distinguished from C. veracruzense n. sp. and C. asuncionense n.
sp. by having a straight-sided test.
130
Measurements (µm):
(n) = number of specimens measured
Diameter of
Length of
cortical shell (max.) (17) primary spine (max.) (14)
225
71
255
94
195
45
222
62
HT
Max.
Min.
Mean
Etymology: This species is named for Scammon's Lagoon
(a haven for migrating gray whales) located to the northeast of the type area.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur, Mexico.
Occurrence: San Hipólito Formation, Baja California Sur;
Tawi Sadh Member of the Guwayza Formation, Oman.
Plate CYC01. Cyclastrum asuncionense Whalen & Carter. Magnification x200. Fig. 1(H)a, b. Whalen & Carter 2002,
pl. 4, figs. 8-9.
Plate CYC02. Cyclastrum scammonense Whalen & Carter. Magnification x200. Fig. 1a,b(H). Whalen & Carter 2002,
pl. 4, figs. 3-4. Fig. 2. Whalen & Carter 2002, pl. 5, fig. 2. Fig. 3a,b. Whalen & Carter 2002, pl. 4, figs. 5, 12.
Fig. 4. OM, BR1121, 15928.
131
Cyclastrum veracruzense Whalen & Carter 2002
Species code: CYC03
Synonymy:
1984 unidentified Radiolaria – Whalen & Pessagno, pl. 1, fig. 14.
2002 Cyclastrum veracruzense n. sp. – Whalen & Carter, p. 111,
pl. 5, figs. 3, 4, 13; pl. 17, fig. 2.
Original description: Test outline a nearly straight-sided
equilateral triangle; on some specimens (including holotype), margins curved gently inward between peripheral spines. Test very thin with gently rounded edges.
Spines short, stout, triradiate in axial section with narrow,
rounded longitudinal ridges and broad shallow longitudinal grooves. Meshwork along triangular margin of test
composed of irregular, medium-sized tetragonal and pentagonal pore frames with no distinctive alignment. Central
area of test slightly depressed, broad, triangular in outline,
with three-rayed area defined by smaller pore frames with
slightly more alignment than on remainder of test; threerayed structure aligned with peripheral spines.
Original remarks: The shorter, broader peripheral spines
and distinctive shape of Cyclastrum veracruzense n. sp.,
distinguish it from C. asunsionense n. sp.
Cyclastrum sp. A
Species code: CYC04
Description: This species is subtriangular in outline and
very thick with near vertical margins. Upper and lower
surfaces of test with a broad outer rim composed mostly of
subrectangular pore frames, and a narrow poorly defined
central cavity made up of smaller pore frames. Test has
a single spine at each corner of test; spines very small,
triradiate at base becoming circular towards tips.
Remarks: This species differs from Cyclastrum asuncionense
Whalen & Carter in having a much thicker test, a smaller
central area and short minuscule spines.
Occurrence: Ghost Creek Formation and Rennell Junction
member of the Fannin Formation, Queen Charlotte
Islands.
132
Further remarks: The very thin, compressed test and short
peripheral spines distinguish Cyclastrum veracruzense
n. sp. from all other species of Cyclastrum.
Measurements (µm):
Based on 6 specimens.
Diameter
of cortical shell (max.)
225
225
195
218
Length
of primary spine (max.)
53
79
30
59
HT
Max.
Min.
Mean
Etymology: This species is named for Pico Vera Cruz
located to the north of the type area.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Fannin Formation, Queen Charlotte Islands.
Plate CYC03. Cyclastrum veracruzense Whalen & Carter. Magnification x200. Fig. 1a,b(H). Whalen & Carter 2002,
pl. 5, figs. 3-4. Fig. 2. QCI, GSC loc. C-304567, GSC 128766.
Plate CYC04. Cyclastrum sp. A. Magnification x200. Fig. 1. QCI, GSC loc. C-305386, GSC 128767. Fig. 2. QCI, GSC loc.
C-304566, GSC 128768. Fig. 3. QCI, GSC loc. C-304566, GSC 128769.
133
Genus: Danubea Whalen & Carter 1998
Type species: Danubea howardi Whalen & Carter 1998
Synonymy:
1998 Danubea n. gen. – Whalen & Carter, p. 40.
Original description: Test with two prominent spines in
the polar positions. Cortical shell inflated, sub-elliptical in
outline with slightly planiform surfaces adjacent to spines;
meshwork composed of tetragonal and triangular pore
frames with prominent nodes at pore frame vertices; large
pores sometimes located on cortical shell at base of spines.
Spines triradiate in axial section and tapering distally.
Original remarks: The bipolar spines of Danubea n. gen.,
distinguish it from all other genera of the Subfamily
Charlotteinae. Danubea n. gen. differs from Pantanellium
Pessagno by having an inner eccentric spicular network.
Protopsium Pessagno and Poisson differs from Danubea
n. gen. by having spongy meshwork.
Etymology: Danubea n. gen., is named for the steamer
Danube, a well known trading ship in the Queen Charlotte
Islands in the late 1800s.
Included species:
DAN02 Danubea sp. A sensu Whalen & Carter 2002
Danubea sp. A sensu Whalen & Carter 2002
Species code: DAN02
Synonymy:
2002 Danubea sp. A – Whalen & Carter, p. 112, pl. 7, figs. 7, 8.
Original remarks: The much smaller cortical shell with
smaller pore frames but with more prominent nodes at pore
frame vertices and the proportionally longer, more massive
polar spines distinguish this species from D. howardi
Whalen and Carter.
Occurrence: San Hipólito Formation, Baja California Sur;
Rennell Junction member of the Fannin Formation, Queen
Charlotte Islands.
Genus: Droltus Pessagno & Whalen 1982
Type species: Droltus lyellensis Pessagno & Whalen, 1982
Synonymy:
1982 Droltus n. gen – Pessagno & Whalen, p. 120
Original description: Test conical to cylindrical, lacking
strictures at joints. Cephalis with short to long horn. Aperture of final post-abdominal chamber open, not enclosed
by latticed dome-shaped cap.
Original remarks: Droltus is compared to Bagotum, n. gen.,
under the latter genus.
Further remarks: Droltus differs from Broctus by lacking
a narrow tubular structure on the final postabdominal
chamber.
134
Etymology: Droltus is a name formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, Pt. VI,
Recommendation 40, p.113).
Included species:
DRO07 Droltus eurasiaticus Kozur & Mostler 1990
DRO02 Droltus hecatensis Pessagno & Whalen 1982
DRO03 Droltus laseekensis Pessagno & Whalen 1982
DRO06 Droltus lyellensis Pessagno & Whalen 1982
DRO08 Droltus sanignacioensis Whalen & Carter 2002
Plate DAN02. Danubea sp. A sensu Whalen & Carter. Magnification Fig. 1 x200, Fig. 2 x400. Fig. 1. Whalen & Carter
2002, pl. 7, fig. 7. Fig. 2. Whalen & Carter 2002, pl. 7, fig. 8.
135
Droltus eurasiaticus Kozur & Mostler 1990
Species code: DRO07
Synonymy:
1982 Parahsuum (?) sp. A – Yao, pl. 3, fig. 6.
1990 Droltus eurasiaticus n. sp. – Kozur & Mostler, p. 223, pl. 17,
fig. 3-4.
1998 Droltus eurasiaticus Kozur & Mostler – Yeh & Cheng,
p. 20, pl. 12, fig. 1.
2002 Droltus eurasiaticus Kozur & Mostler – Whalen & Carter,
p. 116, pl. 16, figs. 5, 6.
Original description: Test conical, multicyrtid, with 6-7
postabdominal segments lacking strictures at joints. Cephalis rounded conical, imperforate, with prominent apical
horn. Cephalis covered by a layer of microgranular silica.
Thorax and subsequent chambers trapezoidal in cross section. Pores arranged in vertical lines. In the thorax they
are closed by layer of microgranular silica. In the remaining chambers the pores are open and become increasingly
larger toward the final postabdominal chamber. Outer latticed layer indistinct, with large pore frames, arranged in
vertical lines and with small to distinct nodes at pore frame
vertices.
Original remarks: The other Droltus species of our material have tricarinate spines.
Measurements (µm):
Length of test
Maximum width
Min.
200
83
Max.
214
100
Etymology: According to its occurrence in Eurasia.
Type locality: Kirchstein Limestone, Kirchstein, Bavaria,
Germany.
Occurrence: Kirchstein Limestone, Germany; Várhegy
Limestone, Hungary; San Hipólito Formation, Baja California Sur; Liminangcong Chert, Philippines.
Droltus hecatensis Pessagno & Whalen 1982
Species code: DRO02
Synonymy:
1982 Droltus hecatensis n. sp. – Pessagno & Whalen, p. 121; pl. 1,
fig. 12, 13, 18, 22; pl. 4, figs. 1, 2, 6, 10; pl. 12, figs. 18-19.
1988 Droltus sp. – Sashida, p. 24, pl. 3, figs. 7, 16, 17.
1989 Droltus hecatensis Pessagno & Whalen – Hattori, pl. 12,
fig. F.
1996 Droltus hecatensis s.l. Pessagno & Whalen – Pujana, p. 138,
pl. 1, figs. 6, 16, 17.
1996 Bagotidae gen. et sp. indet. – Pujana, p. 138, pl. 1, fig. 10.
1998 Droltus hecatensis Pessagno & Whalen – Whalen &
Carter, p. 63, pl. 15, fig. 14.
2001 Droltus hecatensis Pessagno & Whalen – Gawlick et al.,
pl. 5, fig. 13.
2002 Droltus hecatensis Pessagno & Whalen – Suzuki et al.,
p. 181, figs. 8 G, L-M, not fig. 8 H.
2002 Droltus hecatensis Pessagno & Whalen – Tekin, p. 186,
pl. 3, fig. 9.
Original description: Test conical with six or seven postabdominal chambers which are about 5 times as wide as
long. Cephalis small, hemispherical, having small horn
with subsidiary spine. Cephalis and thorax imperforate.
Thorax and subsequent chambers trapezoidal in cross section. Outer latticed layer of abdomen and first several postabdominal chambers with irregularly sized and shaped
polygonal (predominantly tetragonal and pentagonal) pore
frames; pore frames of last two or three post-abdominal
chambers larger, more uniformly sized, predominantly tetragonal (square to rectangular) and aligned in rows.
136
Original remarks: This species differs from D. lyellensis,
n. sp., by having a larger, more massive horn with a
subsidiary spine, by being more pointed apically, and by
having more aligned and more uniformly sized tetragonal
pore frames on its final post-abdominal chambers.
Measurements (µm):
Based on 9 specimens.
Length excluding horn
250.0
260.0
225.0
243.33
Width (maximum)
125.0
150.0
117.5
132.2
HT
Max.
Min.
Mean
Etymology: D. hecatensis, n. sp., is named for Hecate
Straight east of its type locality.
Type locality: Sample QC 534, Rennell Junction member
of the Fannin Formation (Maude Formation in Pessagno
& Whalen, 1982), Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands, Ghost Creek and Fannin formations, Queen Charlotte Islands; Sierra Chacaicó Formation,
Argentina; Pucara Group, Peru; Dürrnberg Formation,
Austria; Hocaköy Radiolarite, Turkey; Musallah Formation
and Tawi Sadh Member of the Guwayza Formation, Oman;
Japan.
Plate DRO07. Droltus eurasiaticus Kozur & Mostler. Magnification x200. Fig. 1(H). Kozur & Mostler 1990, pl. 1, fig. 3.
Fig. 2. Whalen & Carter 2002, pl. 16, fig. 6. Fig. 3. Whalen & Carter 2002, pl. 16, fig. 5.
Plate DRO02. Droltus hecatensis Pessagno & Whalen. Magnification x200. Fig. 1(H). Pessagno & Whalen 1982, pl. 4,
fig. 1. Fig. 2. QCI, GSC loc. C-080611, GSC 128788. Fig. 3. QCI, GSC loc. C-175310, GSC 128789. Fig. 4. QCI, GSC loc.
C-304567, GSC 128790. Fig. 5. OM-00-251, 021431. Fig. 6. OM, BR1122-R02-10. Fig. 7. JP, MNA-10, MA13212.
137
Droltus laseekensis Pessagno & Whalen 1982
Species code: DRO03
Synonymy:
1982 Droltus laseekensis n. sp. – Pessagno & Whalen, p. 122,
pl. 2, fugs. 5, 6, 11, 16; pl. 12, fig. 8, 15.
1998 Droltus laseekensis Pessagno & Whalen – Whalen & Carter,
p. 63, pl. 15, fig. 8; pl. 26, fig. 4.
2004 Droltus laseekensis Pessagno & Whalen – Matsuoka,
fig. 199.
Original description: Test as with genus, conical, usually
with seven or eight post-abdominal chambers. Abdomen
and most post-abdominal chambers rapidly increasing
in width; final three post-abdominal chambers gradually
increasing in width. Cephalis conical with small horn.
Cephalis and thorax sparsely perforate, covered by veneer
of microgranular silica; outer latticed layer of abdomen
and most post-abdominal chambers with irregularly sized
and shaped polygonal (tetragonal and pentagonal) pore
frames; pore frames of last two or three post-abdominal
chambers slightly larger, more uniformly sized and shaped
(rectangular) and aligned in rows.
Measurements (µm):
Based on 10 specimens.
Length excluding horn
257.5
280.0
190.0
232.3
Width (maximum)
142.5
142.5
95.0
122.6
HT
Max.
Min.
Mean
Etymology: D. laseekensis, n. sp., is named for Laseek Bay,
north of its type locality.
Type locality: Sample QC 590A, Sandilands Formation
(Kunga Formation in Pessagno & Whalen 1982), Queen
Charlotte Islands, British Columbia.
Occurrence: Sandilands and Ghost Creek formations and
Rennell Junction member of the Fannin Formation, Queen
Charlotte Islands; Mino Terrane, Japan.
Original remarks: D. laseekensis, n. sp., differs from
D. hecatensis, n. sp., by having pore frames on its final postabdominal chambers that are more irregular in shape and
disposition.
Droltus lyellensis Pessagno & Whalen 1982
Species code: DRO06
Synonymy:
1982 Droltus lyellensis n. sp. – Pessagno & Whalen, p. 122, pl. 2,
figs. 3, 10; pl. 12, fig. 7.
1998 Droltus lyellensis Pessagno & Whalen – Whalen & Carter,
p. 63, pl. 16, fig. 9.
2002 Droltus lyellensis Pessagno & Whalen – Suzuki et al.,
p. 182, fig. 8 I.
Original description: Test conical to subcylindrical with
five or six post-abdominal chambers over 5 times as wide
as long. Cephalis relatively small, hemispherical with
asymmetrically oriented short horn. Cephalis and thorax
sparsely perforate. Outer latticed layer of abdomen and postabdominal chambers with somewhat irregular tetragonal,
pentagonal, and hexagonal pore frames. Tetragonal pore
frames often aligned in rows, tending to be more uniformly
sized.
Original remarks: Droltus lyellensis, n. sp., is compared to
D. hecatensis n. sp., under the latter species.
138
Measurements (µm):
Based on 9 specimens.
Length excluding horn
170.0
210.0
125.0
162.0
Width (maximum)
87.5
100.0
75.0
90.5
HT
Max.
Min.
Mean
Etymology: This species is named for Lyell Island south of
its type locality in the Queen Charlotte Islands.
Type locality: Sample QC 550, Sandilands Formation
(Kunga Formation of Pessagno & Whalen, 1982), north
shore of Kunga Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands and Ghost Creek formations and
Rennell Junction member of the Fannin Formation, Queen
Charlotte Islands; Pucara Group, Peru.
Plate DRO03. Droltus laseekensis Pessagno & Whalen. Magnification x200. Fig. 1(H). Pessagno & Whalen 1982,
pl. 2, fig. 6. Fig. 2. QCI, GSC loc. C-305386, GSC 128791. Fig. 3. QCI, GSC loc. C-175311, GSC 128792.
Fig. 4. QCI, GSC loc. C-304566, GSC 128878. Fig. 5. Matsuoka 2004, fig. 199.
Plate DRO06. Droltus lyellensis Pessagno & Whalen. Magnification x250. Fig. 1(H). Pessagno & Whalen 1982,
pl. 2, fig. 3. Fig. 2. QCI, GSC loc. C-175310, GSC 128793.
139
Droltus sanignacioensis Whalen & Carter 2002
Species code: DRO08
Synonymy:
1984 Bagotum sp. – Whalen & Pessagno, pl. 2, figs. 12-14.
1990 Droltus (?) sp. – De Wever at al., pl. 4, fig. 6.
1998 Droltus sp. – Kashiwagi, pl. 1, fig. 12; pl. 2, figs. 2, 3.
2002 Droltus sanignacioensis n. sp. – Whalen & Carter, p. 116,
pl. 10, figs. 7, 8, 15.
2003 Parahsuum sp. – Kashiwagi & Kurimoto, pl. 3, fig. 5.
Original description: Test conical, with approximately five
post-abdominal chambers. Cephalis hemispherical without
a horn. Thorax, abdomen and most post-abdominal
chambers gradually increasing in width till last postabdominal chamber which slightly decreases in width.
Outer latticed layer on proximal half of test composed of
narrow, irregularly shaped pore frames elongated parallel
to long axis of test; pore frames of outer latticed layer on
distal half of test tetragonal (rectangular) in outline and
aligned in rows.
Measurements (µm):
(n) = number of specimens measured.
Length (10)
165
180
135
158
Width (Max.) (11)
90
105
90
95
HT
Max.
Min.
Mean
Etymology: Droltus sanignacioensis is named for the town
of San Ignacio located to the east of the type area.
Type locality: Sample BPW80-30, San Hipólito Formation,
Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Ghost Creek Formation, Queen Charlotte Islands; Williston
Lake, north-east British Columbia; Musallah Formation,
Oman; Japan.
Original remarks: The elongated, irregularly shaped pore
frames on the proximal portion of the test distinguish this
species from Droltus lyellensis Pessagno and Whalen 1982.
Genus: Ducatus Whalen & Carter 2002
Type species: Ducatus hipolitoensis Whalen & Carter 2002
Synonymy:
2002 Ducatus n. gen. – Whalen & Carter, p. 132.
Original description: Test multicyrtid, composed of cephalis, thorax, abdomen, and swollen post-abdominal
chamber. Cephalis with horn. Post-abdominal chamber
always much larger than cephalis, thorax, and abdomen,
terminating in long, gently tapering, closed tubular extension. Post-abdominal chamber with two prominent,
porous arms, gently tapering to closed distal tips; arms
circular in axial-section, attached at mid-point of spherical portion of post-abdominal chamber, at right angles to
long axis of test.
140
Original remarks: The presence of circumferential porous
arms or wings rather than solid spines distinguishes
Ducatus n. gen., from Katroma Pessagno and Poisson 1981,
and Podobursa Wisniowski 1889, emend. Foreman 1973.
Ducatus is distinguished from Podocapsa Rüst l885, by
having only two circumferential wings, rather than three.
Etymology: Ducatus is a name formed by an arbitrary
combination of letters (ICZN 1985, Appendix D, pt. VI,
Recommendation 40, p. 201).
Included species:
DUC01 Ducatus hipolitoensis Whalen & Carter 2002
Plate DRO08. Droltus sanignacioensis Whalen & Carter. Magnification x250. Fig. 1(H). Whalen & Carter 2002,
pl. 10, fig. 7. Fig. 2. OM-00-252, 021731. Fig. 3. OM-00-118, 000629. Fig. 4. NBC, GSC loc. C-305208, GSC 128911.
Fig. 5. QCI, GSC loc. C-305417, GSC 128794. Fig. 6. QCI, GSC loc. C-080611, GSC 128795. Fig. 7. QCI, GSC loc. C305417, GSC 128912.
141
Ducatus hipolitoensis Whalen & Carter 2002
Species code: DUC01
Synonymy:
1984 unidentified Radiolaria – Whalen & Pessagno, pl. 1,
fig. 13.
2002 Ducatus hipolitoensis n. sp. – Whalen & Carter, p. 132,
pl. 13, figs. 3, 5-7, 11-13, 15; pl. 18, figs. 3, 4.
Original description: Test large with cephalis, thorax,
abdomen, and post-abdominal chamber. Cephalis large,
dome-shaped with horn; cephalis and thorax covered
with layer of microgranular silica. Moderately sized single
horn, circular in cross-section, tapering distally. Abdomen
trapezoidal in outline; noticeable change in slope between
chamber wall of thorax and abdomen; abdomen partially
covered by layer of microgranular silica. Sub-spherical
post-abdominal chamber, large, inflated; hexagonal
pore frames larger on medial portion of post-abdominal
chamber becoming smaller towards abdomen and terminal
tube. Two large porous arms extending at right angles from
post abdominal chamber at medial position and 180° apart;
arms as long as distal closed tube and covered with pores
aligned with long axis of arms.
Original remarks: Ducatus hipolitoensis n. sp. is a monospecific genus at this time and is not compared to any other
species.
Measurements (µm):
(n) = number of specimens.
Length (9)
(excludes horn)
375
375
251
300
Length of arms
(Max.) (10)
143
143
79
106
HT
Max.
Min.
Mean
Etymology: Ducatus hipolitoensis n. sp. is named for Punta
San Hipólito, type locality of the species.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur.
Occurrence: San Hipólito Formation, Baja California.
Genus: Dumitricaella De Wever 1982a, emend. Dumitrica herein
Type species: Dumitricaella pauliani De Wever 1982a
Synonymy:
1982a Dumitricaella n. gen. – De Wever, p. 197.
Original description: Form with two segments bearing
a strong apical horn, two lateral spines and three feet.
Cephalic skeleton consists of actines A, V, D, MB, Ll, Lr,
ll and lr. At the extremity of V is a large pore (sometimes
double) on the cephalic wall. The actines ll and lr give rise
to two lateral spines; the actines D, Ll and Lr to the three
distal feet. Wall of test composed of several layers as for
Jacus n. gen.
Emended description: Skeleton two-segmented with
an initial skeleton consisting of spines A, V, D, Ll, Lr, ll, lr
originating in a short MB. A spine extended into a strong,
three-bladed apical horn. V and ll and lr, or only l spines
extended outside wall into horizontally directed horns.
Wall of cephalis and possibly of thorax two layered. D, Ll
and Lr extended into three, three-bladed feet. Cephalis
relatively large, more or less separated from thorax by
a constriction. Thorax short with a wide aperture and
a distinct rim.
142
Original remarks: Dumitricaella differs from Jacus by the
presence of the two horizontal lateral spines.
Further remarks: The inclusion of Dumitricaella trispinosa
n. sp. in this genus makes it necessary to emend the genus
specifying that the V spine may also be prolonged outside
the cephalic wall. In fact, the V spine is easily prolonged
outside the cephalis in many genera, whereas the two
l spines commonly stop in the cephalic wall. By most its
characters Dumitricaella seems to be closely related to Napora Pessagno from which it differs essentially by the presence of the two secondary lateral spines on the cephalis.
Etymology: Dedicated to P. Dumitrica (Romania) for his
excellent, very meticulous work on Triassic radiolarians.
Included species:
JAC05 Dumitricaella trispinosa Dumitrica n. sp.
Plate DUC01. Ducatus hipolitoensis Whalen & Carter. Magnification x200. Fig. 1(H). Whalen & Carter 2002, pl. 13,
fig. 3. Fig. 2. Whalen & Carter 2002, pl. 13, fig. 5.
143
Dumitricaella trispinosa Dumitrica n. sp.
Species code: JAC05
Synonymy:
1989 Dumitricaella? spp. – Hattori, pl. 19, fig. L.
Type designation: Specimen R20-02 (pl. JAC05, fig. 1)
from sample BR 485, Tawi Sadh Member of the Guwayza
Formation, Oman.
Diagnosis: Dumitricaella with three equal V and l spines.
Description: Test small, pyramidal with a thick, pointed,
three-bladed apical horn. Blades of horn aligned with
spines V and the two l. Cephalis relatively large, perforate,
thick-walled with three short, broad, laterally directed,
three-bladed spines. Spines sometimes replaced by a lobelike prolongation of cephalis that becomes trilobate. Pores
of cephalis with rounded triangular or quadrangular raised
pore frames that can form ridges with different orientations.
Collar stricture relatively well marked by a constriction.
Thorax slightly longer and broader than cephalis, pyramidal.
Dorsal and primary lateral spines prolonged from the base
of cephalis in the wall of thorax, their outer blade forming
a high rib, and extended into three divergent, curved,
pointed feet. Distal end of thorax wide, open and bordered
distally by an imperforate peristome. Pores of thorax
small, rounded; pore frames aligned to form longitudinal,
transversal or oblique ribs.
Remarks: Dumitricaella trispinosa n. sp. differs from
D. pauliani De Wever in having V and l spines equally developed, in having these spines shorter, and a rather well
marked collar constriction.
Measurements (µm):
Based on 7 specimens.
Total length of skeleton
Length of apical horn
Length of cephalis
Length of thorax
Diameter of cephalis with spines or lobes
Diameter of thorax
Min.
160
45
26
46
50
72
Max.
195
60
35
60
74
100
Etymology: From the three spines (V, lr and ll) laterally
extended from the cephalis.
Type locality: Sample BR 485, Guwayza Formation, Tawi
Sadh Member, Jabal Safra, Oman.
Occurrence: Tawi Sadh Member of the Guwayza Formation, Oman; Skrile Formation, Slovenia; Japan.
Genus: Elodium Carter 1988
Type species: Elodium cameroni Carter 1988
Synonymy:
1988 Elodium Carter – Carter et al., p. 56.
1996 Elodium Carter – Yeh & Cheng, p. 118.
Original description: Test conical and large, with well developed horn and numerous closely spaced post-abdominal chambers separated by nodose circumferential ridges.
Three rows of longitudinally aligned circular to subcircular pores in polygonal (mostly tetragonal) pore frames,
between circumferential ridges. Lateral pore rows flanking ridges slope steeply away from ridges. Post-abdominal
chambers constricted between ridges. Pores in constricted
area may be irregular to absent on distalmost chambers of
test. Cephalis and thorax sparsely perforate to imperforate,
covered with outer layer of microgranular silica; this covering may extend onto earliest post-abdominal chambers.
144
Original remarks: Elodium n. gen. possesses three rows of
primary (open) pores between circumferential ridges; it
differs from Parvicingula Pessagno in that these pores are
longitudinally aligned rather than offset.
Further remarks: Elodium Carter can be distinguished
from Parahsuum Yao by having prominent circumferential
ridges and much less pronounced longitudinal costae.
Etymology: Elodium is formed by an arbitrary combination
of letters (ICZN, 1985, Appendix D, Pt. VI, Recommendation
40, p. 201).
Included species:
3411 Elodium cameroni Carter 1988
PHS08 Elodium? mackenziei Carter n. sp.
ELD02 Elodium pessagnoi Yeh & Cheng 1996
ELD03 Elodium wilsonense (Carter) 1988
Plate JAC05. Dumitricaella trispinosa Dumitrica n. sp. Magnification x300. Fig. 1(H). OM, BR485-R20-02.
Fig. 2. OM, BR871-R05-22. Fig. 3. BR871-R03-01. Fig. 4. BR871-R08-09. Fig. 5. OM, BR871-R03-03.
Fig. 6. OM, BR871-R09-14. Fig. 7. SI, MM6.76, 000506.
145
Elodium cameroni Carter 1988
Species code: 3411
Synonymy:
1988 Elodium cameroni n. sp. – Carter et al., p. 56, pl. 13, figs. 1,
2, 6, 9.
1991 Elodium cameroni Carter – Tipper et al., pl. 9, fig. 12.
1991 Elodium cameroni Carter – Carter & Jakobs, p. 342, pl. 3,
fig. 18.
1995a Elodium cameroni Carter – Baumgartner et al., p. 194,
pl. 3411, figs. 1-2.
1996 Elodium sp. aff. E. pessagnoi n. sp. – Yeh & Cheng, p. 120,
pl. 11, fig. 7 only.
1997 Elodium aff. cameroni Carter – Yao, pl. 13, fig. 635.
Not 1997 Elodium cameroni Carter – Yao, pl. 13, fig. 636.
Original diagnosis: Large conical-cylindrical test with 10
to 14 post-abdominal chambers and a strong asymmetric
apical horn. All pores large, primary and circular; three rows
on proximal chambers, two rows on distalmost chambers.
Original description: Test large with 10 to 14 strongly
constricted post-abdominal chambers separated by nodose
circumferential ridges; nodes low and rounded. Cephalis
and thorax trapezoidal in external outline, partially
perforate, covered by veneer of microgranular silica.
Cephalis has strong, asymmetric apical horn. All pores on
post-abdominal chambers circular and primary (open);
those within constricted areas smaller, disappearing on
distalmost chambers. Earliest post-abdominal chambers
trapezoidal, increasing gradually in width and height, distal
chambers almost cylindrical with slight decrease in height.
Original remarks: Elodium cameroni is compared to
E. nadenensis n. sp., under the latter. Elodium cameroni is
very abundant, in all middle/upper Toarcian samples.
Original remarks under Elodium nadenensis Carter in
Carter et al. (1988): Differs from Elodium cameroni n. sp. by
having a more conical, apically pointed test with a heavier
coating of microgranular silica. In addition, the horn is
shorter, more symmetrical and circumferential ridges
are wider and more rounded. Differs from Lupherium (?)
sp. B by having more prominent circumferential ridges.
Abundant.
Measurements (µm):
Based on 20 specimens.
Length (excluding horn)
Maximum width
HT
369
161
Av.
352
159
Max.
450
185
Min.
280
147
Etymology: This species is named in honour of B.E.B.
Cameron for his important contribution to the Mesozoic
stratigraphy and foraminiferal biostratigraphy of the Queen
Charlotte Islands, B.C.
Type locality: GSC locality C-080597. Phantom Creek
Formation. Yakoun River, Graham Island, Queen Charlotte
Islands, British Columbia.
Occurrence: Queen Charlotte Islands, British Columbia;
Liminangcong Chert, Philippines; Japan.
Elodium? mackenziei Carter n. sp.
Species code PHS08
Synonymy:
? 1982 Lupherium sp. A – Pessagno & Whalen, p. 136, pl. 6, fig. 4.
1987b Lupherium sp. G – Yeh, p. 68, pl. 23, fig. 5.
1988 Lupherium (?) sp. B – Carter et al., p. 54, pl. 5 fig. 11; pl. 13,
figs. 5, 10, 12.
? 1990 Parahsuum simplum Yao – De Wever at al., pl. 4, fig. 9.
? 2004 Archaeodictyomitra? sp. – Hori, pl. 1, fig. 55.
2004 Lupherium sp. – Matsuoka, figs. 217, 218.
Type designation: Holotype GSC 80758 (Carter et al. 1988,
pl. 13, figs. 5, 10, 12), from GSC loc. C-080583; Phantom
Creek Formation (upper Toarcian).
Description: Test elongate, quite pointed apically, usually
with ten to twelve post-abdominal chambers. Cephalis
conical, possibly including a short apical horn. Remaining
chambers in apical half of the test trapezoidal, increasing
more in height than width as added; next few chambers
cylindrical, final two chambers slightly constricted.
Slightly raised circumferential ridges visible between postabdominal chambers on the distal portion of the test.
Costae fine, narrowly spaced throughout length of test.
Pores subcircular to subelliptical in shape.
146
Remarks: Elodium? mackenziei n. sp. differs from Elodium
nadenense Carter (1988) in having less prominent
circumferential ridges between post abdominal chambers
and an absent to poorly developed apical horn. It differs
from E. wilsonense Carter (1988) in having a more narrowly
conical shape and less differentiated apical horn. Genus
Elodium is queried because the three rows of aligned pores
between chambers that characterize the genus are not well
developed. E.? mackenziei n. sp. may be ancestral to all other
species of Elodium in the upper Toarcian and Aalenian of
Queen Charlotte Islands and it may also represent the link
between Parahsuum and Elodium.
Measurements (µm):
Based on 11 specimens.
Length (excl. horn)
Maximum width
HT
310
118
Max.
343
131
Min.
210
92
Mean
290
120
Etymology: This species is named for J.D. MacKenzie
(Geological Survey of Canada) who first mapped central
Graham Island in 1913-1914.
Plate 3411. Elodium cameroni Carter. Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 13, fig. 2.
Fig. 2. Carter & Jakobs 1991, pl. 3, fig. 18. Fig. 3. Carter et al. 1988, pl. 13, fig. 1.
Type locality: Sample GSC loc. C-080583, Phantom Creek
Formation, Yakoun River, Graham Island, approximately
2 km south of Ghost Creek; east side of the river, Queen
Charlotte Islands, British Columbia.
Occurrence: Fannin member of the Fannin Formation,
Whiteaves and Phantom Creek formations, Queen Charlotte Islands; Nicely and Hyde formations, Oregon; Mino
Terrane, Japan.
Plate PHS08. Elodium? mackenziei Carter n. sp. Magnification x250. Fig. 1(H). Carter et al. 1988, pl. 13, fig. 5.
Figs. 2, 3. Matsuoka 2004, figs. 217, 218.
147
Elodium pessagnoi Yeh & Cheng 1996
Species code: ELD02
Synonymy:
1982 Parahsuum? sp. – Matsuda & Isozaki, pl. 1, figs. 18, 19.
1989 Parahsuum (?) sp. aff. P. (?) magnum Takemura – Hori &
Otsuka, p. 182, pl. 3, figs. 13-15.
1990 Parahsuum (?) aff. P. magnum Takemura – Hori, Fig. 9.37.
1995a Parahsuum sp. M - Baumgartner et al., p. 384, pl. 2015,
figs. 1, ?2.
1996 Elodium jurassicum n. sp. – Yeh & Cheng, p. 118, pl. 11,
figs. 1, 2, 6, 12, 13.
1996 Elodium pessagnoi n. sp. – Yeh & Cheng, p. 120, pl. 4,
figs. 5, 7-8, 10-13; pl. 11, figs. 3, 4, 9, 14.
1996 Elodium sp. aff. E. pessagnoi n. sp. – Yeh & Cheng, p. 120,
pl. 6, fig. 13; pl. 11, figs. 5, 11, 15, not fig. 7.
1996 Elodium sp. cf. E. pessagnoi n. sp. – Yeh & Cheng, p. 120,
pl. 4, fig. 6.
1996 Elodium sp. B – Yeh & Cheng, p. 122, pl. 12, figs. 1, 6, 8, 12,
not fig. 4.
1996 Elodium sp. C – Yeh & Cheng, p. 122, pl. 12, fig. 2, 7, 9, 10.
1996 Elodium sp. D – Yeh & Cheng, p. 122, pl. 12, figs. 3, 15.
1996 Elodium sp. E – Yeh & Cheng, p. 122, pl. 12, figs. 5, 11.
1997 Parahsuum (?) aff. P. magnum Takemura – Hori, pl. 1, fig. 7.
2004 Parahsuum (?) sp. aff. magnum Hori & Otsuka – Ishida et
al., pl. 5, fig. 21.
Original description: Test medium in size, conical in shape,
multicyrtid with six to seven post-abdominal chambers.
Cephalis and thorax moderately broad, hemispherical with
massive horn. Horn tapered, circular in axial section. Outer
test layer of cephalis and thorax covered with raised irregular
pore frames. First postabdominal chamber separated
from abdomen and subsequent postabdominal chambers
separated from each other by nodose circumferential ridges.
Abdomen and postabdominal chambers trapezoidal in
shape; each chamber having three rows of tetragonal pore
frames. Pore frames longitudinally aligned. Postabdominal
chambers increasing very slowly in height and gradually in
width as added.
Further remarks: In Elodium pessagnoi we include all
forms with a long straight apical horn, circular in crosssection. The test is conical or subcylindrical in shape; the
apical portion of the test is covered by simple circular
pores or possesses an additional layer of raised irregular
nodes. Elodium jurassicum is herein synonymized with
E. pessagnoi. It should be noted that holotypes of these
two species (and most specimens of informal species)
illustrated from Busuanga Island (Yeh & Cheng, 1996) are
from a single sample.
Measurements (µm):
Based on 7 specimens.
Max.
Max.
test width test length
HT
148
284
Mean
145
264
Max.
150
291
Min.
138
239
Length
No. of
of horn postabdom. chambers
49
6
55
5.5
74
6
41
5
Etymology: This species is named for Prof. E. A. Pessagno,
UT-Dallas, U.S.A., in honor of his great contribution to
Mesozoic radiolarian studies.
Type locality: Liminangcong Chert near Ocam Ocam village, Busuanga Island, Philippines.
Occurrence: Liminangcong Chert, Philippines; Japan.
Elodium wilsonense (Carter) 1988
Species code: ELD03
Synonymy:
1988 Crubus wilsonensis Carter n. sp. – Carter et al., p. 53, pl. 5,
fig. 12.
1996 Elodium sp. cf. E. wilsonense (Carter) – Yeh & Cheng,
p. 122, pl. 12, figs. 13, 14.
Original diagnosis: Test large, broadly conical with short
horn. Eighteen costae visible laterally; three longitudinally
aligned pores per chamber between adjacent costae. Ridges
slightly raised with small nodes superimposed on costae.
Original description: Test broadly conical, rounded apically
with short cylindrical horn. Cephalis hemispherical,
all other chambers trapezoidal in outline. Cephalis
imperforate, thorax and abdomen sparsely perforate; all
three covered with a layer of microgranular silica. Usually
7 to 9 postabdominal chambers. All but final chamber
increase gradually in width and height. Fifteen continuous
narrow costae visible laterally, nodose along ridges. Single
148
rows of longitudinally aligned pores (three per chamber, set
in square pore frames) alternate with costae.
Original remarks: Differs from Crubus firmus Yeh in having three rows of linearly arranged pore frames per chamber, larger pores on initial chambers, a stouter horn, and
stronger costae.
Further remarks: Elodium wilsonense differs from
E. cameroni Carter in having fewer postabdominal chambers and much less prominent circumferential ridges. It
differs from E. pessagnoi Yeh & Cheng in having a more
broadly conical shape, two rows of pore frames on early
postabdominal chambers and a shorter apical horn.
Measurements (µm):
Based on 10 specimens.
Length (excluding horn)
Maximum width
HT
334
188
Av.
290
171
Max.
334
190
Min.
210
155
Plate ELD02. Elodium pessagnoi Yeh & Cheng. Magnification x250. Fig. 1(H). Yeh & Cheng 1996, pl. 11, fig. 3.
Fig. 2. JP, NK86050720. Fig. 3. Hori 1990, fig. 9-37.
Etymology: Named for Wilson Creek on Graham Island,
site of one of the early coal mines.
Occurrence: Whiteaves Formation, Queen Charlotte Islands; Liminangcong Chert, Philippines.
Type locality: GSC locality C-080579, Whiteaves Formation, Creek locality, Maude Island, Queen Charlotte Islands, British Columbia.
Plate ELD03. Elodium wilsonense (Carter). Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 5, fig. 12.
149
Genus: Eospongosaturninus Kozur & Mostler 1990
Type species: Spongosaturnalis protoformis Yao 1972
Synonymy:
1990 Eospongosaturninus n. gen. – Kozur & Mostler, p. 211.
Original description: Shell large, spongy, consisting of
several concentric layers. Microsphere latticed. The shell
reaches on the ridge around the base of the peripolar
spines. Ring transversally elongated elliptical, narrow to
moderately broad, in parts of the ring undifferentiated, in
other parts with ridge near the inner side of the ring. Outer
margin of ring smooth or with one peripolar spine on the
long axis poles. Near the base of the peripolar spines tiny
auxiliary spines may be present.
Original remarks: Eospongosaturninus n. gen. is the forerunner of Spongosaturninus Campbell and Clark, 1944b
that has more distinct ridges along the whole inner margin of the ring, a latticed second medullary shell and a
rather large third medullary shell (transitional to a cortical
shell) covered by a still rather thick spongy layer. Pseudacanthocircus n. gen. has an undifferentiated ring and the
shell reaches never until the ring. Spongosaturnalis Campbell and Clark, 1944b displays more distinct ridges along
the whole inner margin of the ring and the shell does not
reach the ring. Eospongosaturninus n. gen. lies in the transition field between the genera Pseudacanthocircus n. gen.,
Spongosaturninus Campbell and Clark, 1944b and Spongosaturnalis Campbell and Clark, 1944b. The type species
is surely the forerunner of Spongosaturninus Campbell
and Clark, 1944b. In Eospongosaturninus? bispinosus (Yao,
1972) the overreach of the shell on the ring is not so dis-
tinct and in several specimens the inner ridge on the ring
is indistinct, partly even missing (Yao, 1972, pl. 2, fig. 9).
The specimen, figured by Yao (1972, pl. 2, fig. 8) as Spongosaturnalis bispinosus belongs to an other species and is
probably the first representative of the genus Spongosaturnalis Campbell and Clark, 1944b. The genus Acanthocircus
Squinabol, 1903b has strong outer ridges on the ring or the
ridges cover the whole ring. A deep furrow on the lateral
outer side of the ring is always present. This genus is only
homoeomorphic to the Pseudacanthocircidae n. fam. (and
in it to Eospongosaturninus n. gen.) and evolved from the
parasaturnalinid stock.
Further remarks: In disagreement with Kozur and Mostler (1990) we consider that in spite of its name Eospongosaturninus has nothing to do with Spongosaturninus and is
not »in the transition field between the genera Pseudacanthocircus Kozur and Mostler, Spongosaturninus Campbell
and Clark and Spongosaturnalis Campbell and Clark«. In
the Middle Jurassic the genus should be restrained to the
type species. Its main generic character is the three-bladed
ring in the middle part and twisting of the ring, the result
being the change of the position of the blades from the
middle part to the distal part.
Etymology: Forerunner of Spongosaturninus Campbell &
Clark 1944.
Included species:
2021 Eospongosaturninus protoformis (Yao) 1972
Eospongosaturninus protoformis (Yao) 1972
Species code: 2021
Synonymy:
1972 Spongosaturnalis protoformis n. sp. – Yao, p. 27, pl. 1, figs.
2-7; pl. 10, figs. 1-2.
1995a Acanthocircus protoformis (Yao) – Baumgartner et al.,
p. 64, pl. 2021, figs. 1-3.
1996 Acanthocircus protoformis (Yao) – Yeh & Cheng, p. 108,
pl. 2, fig. 11.
Original description: Spongosaturnalis with simple ring,
where no spine is developed. Shell approximately spherical,
spongy, composed of irregular meshes which become
denser centrally. Polar spines short, smooth, not always
distinguished when shell extends completely across ring.
Polar spines change to sturdy spines inside shell. When
shell is not preserved, numerous fragmentary thorns
are observed on sturdy spines and rarely on ring where
each of polar spines bifurcates. Ring generally bilaterally
symmetrical or ovoidal, simple, with ridges on both edges
near polar spines. Ridge on outer edge extends across ends
150
of polar spines, and another one on inner edge disappears
at polar spines. Both ridges become obsolete on terminal
end of ring. No spine on ring.
Original remarks: This species may be similar to Saturnalis
simplex Squinabol (1914, p. 286-287, pl. 22, fig. 2; Jurassic,
Fontanafredda (Euganei), Italy) in the shape of the saturnalin ring, but the generic assignment of S. simplex is doubtful
because the nature of the shell is not known, namely the
shell is not preserved and the fragmentary thorns are not
observed on the polar spines. Spongosaturnalis protoformis
differs from S. bispinus (described below) in lacking spine
on each terminal end of the ring, and in having ridges on
both edges of the ring.
Further remarks: In spite of the similarity to Saturnalis
simplex Squinabol the species differs from the latter by the
character of the blades.
Measurements (µm):
Based on 6 specimens.
HT
Av.
Diameter of ring along polar spines 170 191
Diameter of ring transversaly
335 362
153
Diameter of shell
110
Length of polar spine
20
15
Breadth of ring
11-21 12-20
Min. Max.
170 210
330 420
110 180
7
20
9
29
Type locality: Manganese carbonate ore, Mino Belt, river
side of the Kiso, east of Unuma, Kagamihara City, Gifu
Prefecture, Central Japan.
Occurrence: Inuyama area, Japan; Italy; Tawi Sadh Member
of the Guwayza Formation, Oman; Liminangcong Chert,
Philippines; Snowshoe Formation, Oregon.
Plate 2021. Eospongosaturninus protoformis (Yao). Magnification x200. Fig. 1(H). Yao 1972, pl. 1, fig. 2.
Fig. 2. JP, IYII-17. Fig. 3. OM, BR871-R01-08. Fig. 4. OM, BR871-R01-11. Fig. 5. BR871-R01-05. Fig. 6. OM, BR871R01-12. Fig. 7. OM, BR871-R02-12. Fig. 8. JP, Nanjo Massif, IH84120462-R05-08. Fig. 9. OR555-R07-07.
151
Genus: Eucyrtidiellum Baumgartner 1984
Type species: Eucyrtidium (?) unumaensis Yao 1979
Synonymy:
1984 Eucyrtidiellum n. gen. – Baumgartner, p. 764.
1986 Monosera n. gen – Takemura & Nakaseko, p. 1021.
1986 Eucyrtidiellum Baumgartner – Takemura, p. 66.
1990 Eucyrtidiellum Baumgartner – Nagai & Mizutani, p. 593.
Original description: Test composed of four segments.
Cephalis small, spherical, poreless with variably developed
straight or slightly oblique apical horn, rare forms with
apical and vertical horn. A sutural pore is present at collar
stricture or on proximal portion of thorax. Thorax domeshaped, poreless, with irregular ornamentation consisting
of ridges and nodes leaving depressions (“closed pores” of
some authors) or with plicae. One or two rows of pores may
occur at stricture between thorax and abdomen. Abdomen
inflated annular to hemispherical, poreless, except for the
distal quarter, where one or two irregular rows of pores may
occur. Ornamentation of abdomen varying with species.
One row of large pores marks the joint with fourth segment.
Fourth segment delicate, mostly cylindrical, covered with
circular pores in loose diagonal rows, with a distal poreless
constriction.
Original remarks: The Mesozoic species hitherto
questionably assigned to Eucyrtidium are assigned to this
new genus, because they bear no resemblance to the type
species E. acuminatum (Ehrenberg).
Further remarks: The stratigraphic record and structural
features prove that Eucyrtidiellum is derived from the genus
Thetis De Wever, and especially T. oblonga De Wever. In
fact, the two genera seem to be synonymous: both have
four segmented tests where the first three are thickerwalled and the fourth is thin-walled and usually not
preserved; a practically imperforate cephalis; and usually
an apical horn that is always circular in cross section. What
would differentiate T. oblonga De Wever, the type species
of the genus Thetis, from the other species of the genus
Eucyrtidiellum would be the presence of three thoracic
spines representing probably the dorsal and the two primary
lateral spines, and the large ventral pore aligned with the
ventral spine. However, a ventral pore is present with
Thetis, or at least the type species. T. oblonga? (De Wever,
1982a, pl. 4, fig. 16) seems to show it, and a specimen we
illustrate herein from the type sample shows clearly such a
pore (see under T. oblonga, pl. THT01, fig. 4). The fact that
the holotype of T. oblonga shows no ventral pore is because
it is illustrated in the dorsal position.
Included species:
EUC09 Eucyrtidiellum disparile gr. Nagai & Mizutani 1990
EUC10 Eucyrtidiellum gujoense (Takemura & Nakaseko)
1986
EUC03 Eucyrtidiellum gunense gr. Cordey 1998
EUC06 Eucyrtidiellum nagaiae Dumitrica, Goričan &
Matsuoka n. sp.
EUC07 Eucyrtidiellum omanojaponicum Dumitrica,
Goričan & Hori n. sp.
EUC04 Eucyrtidiellum ramescens Cordey 1998
Eucyrtidiellum disparile gr. Nagai & Mizutani 1990
Species code: EUC09
Synonymy:
1986 Eucyrtidiellum sp. a and sp. a1 – Nagai, pl. 1, figs. 5, 6.
1986 Monosera unumaensis (Yao) – Takemura & Nakaseko,
p. 1022, pl. 4, fig. 9.
1987 Eucyrtidiellum sp. A – Hattori, pl. 12, figs. 1, 2.
1987 Eucyrtidiellum aff. E. unumaensis Yao – Hattori, pl. 12,
figs. 5, 6.
1988 Eucyrtidiellum sp. a – Nagai, pl. 1, figs. 1a-d, 2.
1988 Eucyrtidiellum sp. A – Hattori, pl. 8, fig. D.
1988 Eucyrtidiellum sp. aff. E. unumaensis Yao – Hattori, pl. 8,
fig. J.
1989 Eucyrtidiellum spp. – Hattori, pl. 7, fig. H.
1989 Eucyrtidiellum unumaensis (Yao) - Hattori, pl. 28, Fig. H.
1990 Eucyrtidiellum disparile n. sp. – Nagai & Mizutani, p. 594,
fig. 3. 6-8a,b,c.
1991 Eucyrtidiellum sp. cf. E. disparile – Kojima et al., pl. 1, fig. 17.
1993 Eucyrtidiellum disparile Nagai & Mizutani – Fujii et al.,
pl. 1, fig. 13.
1993 Eucyrtidiellum sp. a – Fujii et al., pl. 1, fig. 14.
1995 Eucyrtidiellum disparile Nagai & Mizutani – Nagai, pl. 4,
fig. 3; pl. 5, fig. 8.
1997 Eucyrtidiellum aff. unumaense (Yao) – Yao, pl. 10, fig. 458.
152
1998 Eucyrtidiellum disparile Nagai & Mizutani – Kashiwagi,
pl. 1, fig. 18.
Not 2001 Eucyrtidiellum disparile Nagai & Mizutani
– Matsuoka et al., pl. 3, fig. 16.
2003 Eucyrtidiellum disparile Nagai & Mizutani – Goričan et
al., p. 296, pl. 5, figs. 2, 3.
2004 Eucyrtidiellum disparile Nagai & Mizutani – Hori, pl. 3, fig. 42;
pl. 5, figs. 52-54, not fig. 55; pl. 6, figs. 4-8, pl. 10, fig. 27.
2004 Eucyrtidiellum sp. – Ishida et al., pl. 5, fig. 13.
? 2004 Eucyrtidiellum disparile Nagai & Mizutani – Matsuoka,
fig. 177.
2004 Eucyrtidiellum disparile Nagai & Mizutani – Suzuki &
Ogane, pl. 8, fig. 2.
2005 Eucyrtidiellum aff. disparile Nagai & Mizutani – Hori,
pl. 8, fig. 20.
2005 Eucyrtidiellum disparile Nagai & Mizutani – Hori, pl. 8,
fig. 21; pl. 12, figs. 18-20, 49; pl. 13, figs. 24-25.
2005 Eucyrtidiellum disparile Nagai & Mizutani – Kashiwagi et
al., pl. 6, fig. 11.
Original diagnosis: Abdomen has circular pores regularly
arranged along two diagonal lines on its whole surface.
Original description: This species has a test which is composed generally of three segments, cephalis, thorax and
abdomen. Cephalis is small, spherical with a mediumsized apical horn. Sutured pores are arranged at a stricture
between thorax and abdomen. Thorax truncated-conical
with closed pores and irregular hexagonal meshworks on
the whole surface. Abdomen is relatively large and inflated-hemispherical, with circular pores regularly arranged
along two diagonal lines on its whole surface.
Original remarks: External shell form of Eucyrtidiellum
disparile is almost identical with that of E. unumaense, but
differs in having pores on its entire abdomen. Many specimens of Eucyrtidiellum have been provisionally described
as Eucyrtidium (?) sp. or Eucyrtidiellum (?) sp., just because of the presence of opened pores in their abdomen.
Further remarks: Although initially we were tempted to
separate this species into two subspecies, one with shorter
apical horn, the other with longer apical horn, we decided
against this because, except for the length of this horn,
the test morphology is similar in almost all the other
respects. The specimen determined by Matsuoka (2004)
as E. disparile is questionably included in the synonymy
of this species because it lacks pores on the middle part
of the abdomen. Accordingly, this specimen is closer to
E. unumaense (Yao) and looks intermediary between the
two species. This would prove that the evolutionary trend
giving rise to E. unumaense started in the early Toarcian.
Measurements (µm):
Based on 10 specimens.
Height of apical horn
Height of entire body including
cephalis, thorax and abdomen
Height of cephalis
Width of cephalis
Height of thorax
Width of thorax
Height of abdomen
Width of abdomen
Min.
3
Max.
13
Av.
8
67
100
79
5
8
14
31
41
48
19
21
23
38
66
77
13
17
18
35
48
71
Etymology: Derived from Latin adjective disparilis, -e,
which means unlike or dissimilar.
Type locality: Sample MNHK05, Kamiaso, Mino Terrane,
central Japan.
Occurrence: Japan; Skrile Formation, Slovenia; Tawi Sadh
Member of the Guwayza Formation, Oman.
Plate EUC09. Eucyrtidiellum disparile gr. Nagai & Mizutani. Magnification x400. Fig. 1(H). Nagai & Mizutani 1990,
fig. 3-8c. Fig. 2. Goričan et al. 2003, pl. 5, fig. 2. Fig. 3. Goričan et al. 2003, pl. 5, fig. 3. Fig. 4. OM, BR1121-R06-10.
Fig. 5. OM; BR1121-R06-24. Fig. 6. OM, BR706-R13-03. Fig. 7. JP, MNA-10, MA13437. Fig. 8. JP, MNA-10, MA11896.
153
Eucyrtidiellum gujoense (Takemura & Nakaseko) 1986
Species code: EUC10
Synonymy:
1986 Monosera gujoensis n. sp. – Takemura & Nakaseko,
p. 1022, figs. 4.10, 4.11, 5.1-5.3.
1986 Eucyrtidiellum gujoensis (Takemura & Nakaseko)
– Takemura, p. 67, pl. 12, figs. 13-15.
1986 Eucyrtidiellum sp. e – Nagai, p. 14, pl. 1, figs. 3a-c.
1987 Eucyrtidiellum gujoensis – Hattori, pl. 12, figs. 14, 15.
1989 Eucyrtidiellum sp. aff. E. gujoensis Takemura – Hattori &
Sakamoto, pl. 10, fig. B.
1988 Eucyrtidiellum gujoensis (Takemura & Nakaseko) – Nagai,
pl. 2, figs. 5, 9.
1988 Eucyrtidiellum sp. f – Nagai, pl. 2, fig. 8.
1989 Tethys oblonga De Wever – Hattori, pl. 7, figs. C, D.
1989 Eucyrtidiellum gujoensis Takemura & Nakaseko – Nagai,
pl. 1, fig. 7; pl. 2, fig. 7a-b; pl. 3, fig. 7; pl. 4, fig. 5.
1995 Eucyrtidiellum gujoensis Takemura – Nagai, pl. 5, fig. 9.
1995 Eucyrtidiellum sp. f – Nagai, pl. 5, fig. 10.
1997 Eucyrtidiellum sp. D0 – Yao, pl. 10, fig. 452.
2001 Eucyrtidiellum disparile Nagai & Mizutani – Matsuoka
et al., pl. 3, fig. 16.
2002 Eucyrtidiellum gujoense (Takemura & Nakaseko) – Hori &
Wakita, pl. 3, fig. 4.
2004 Eucyrtidiellum gujoense (Takemura & Nakaseko)
– Matsuoka, fig. 178.
2004 Eucyrtidiellum sp. D0 sensu Yao – Hori, pl. 2, fig. 8.
Original diagnosis: A species of Monosera characterized by
the existence of pore frames on abdominal surface and by
irregular or longitudinal arrangement and dense distribution of abdominal pores.
Original description: Cephalis small, spherical to subspherical, poreless with a strong rod-like apical horn. Thorax
truncated conical to subhemispherical with usually irregularly arranged pores or relict pores. Abdomen subspherical
with pores on surface. Abdominal pores with pore frames,
longitudinally or irregularly distributed in great density.
Fourth segment inflated-cylindrical with irregularly distributed pores. Cephalic structure same as genus.
154
Original remarks: Monosera gujoensis n. sp. is distinguished from M. unumaensis by the presence of abdominal
pore frames and the distribution and density of abdominal
pores. Monosera gujoensis differs from Eucyrtidium ptyctum Riedel and Sanfilippo in the absence of plicae on the
surface of the abdomen and in the presence of abdominal
pores and pore frames.
Further remarks: Morphologically E. gujoense is very close
to Thetis oblonga De Wever from which it differs in having a
shorter, conical apical horn, and lacks spines on the thorax,
and shoulders on the upper third of the abdomen.
Measurements (µm):
Based on 15 specimens.
Length of apical horn
Height of cephalis
Height of thorax
Height of abdomen
Width of cephalis
Width of thorax
Width of abdomen
Min.
10
15
20
40
20
40
70
Max.
50
20
30
50
25
50
90
Etymology: The trivial name, gujoensis, is derived from the
Gujo County, Gifu Prefecture, Japan.
Type locality: Manganese carbonate ore deposits (sample
TKN-105) from the Gujo-hachiman area of the Mino
Terrane, Japan.
Occurrence: Japan; Tawi Sadh Member of the Guwayza
Formation and Musallah Formation, Oman.
Plate EUC10. Eucyrtidiellum gujoense (Takemura & Nakaseko). Magnification x400. Fig. 1(H). Takemura &
Nakaseko 1986, fig. 4-10. Fig. 2. Matsuoka 2004, fig. 178. Fig. 3. OM, BR1121-R08-28. Fig. 4. OM-00-252, 021905.
Fig. 5. OM-00-258, 022630.
155
Eucyrtidiellum gunense gr. Cordey 1998
Species code: EUC03
Synonymy:
1982 Eucyrtidium (?) sp. C – Imoto et al., pl. 1. fig. 9.
1984 Eucyrtidiellum sp. A – Murchey, pl. 2, fig. 25.
1984 Eucyrtidiellum sp. B – Murchey, pl. 2, fig. 26.
1986 Eucyrtidiellum sp. C group – Nagai, p. 12, pl. 2, fig. 10,
not 11-12.
1987 Eucyrtidiellum sp. C – Hattori, pl. 12, fig. 10.
1990 Eucyrtidiellum sp. C3 – Nagai, pl. 4, fig. 2, 3.
1992 Eucyrtidiellum sp. – Sashida, pl. 1, figs. 19, 20.
1995 Eucyrtidiellum sp. C – Nagai, pl. 4, fig. 6.
1997 Eucyrtidiellum sp. Q0 – Yao, pl. 10, fig. 453.
1998 Eucyrtidiellum gunensis n. sp. – Cordey, p. 109, pl. 25,
figs. 8-9.
2004 Eucyrtidiellum sp. Q0 sensu Yao – Hori, pl. 2, fig. 7.
2004 Eucyrtidiellum gunense Cordey – Matsuoka, fig. 180.
2005 Eucyrtidiellum sp. C sensu Nagai – Hori, pl. 8, figs. 16-18,
not fig. 19.
Original diagnosis: Eucyrtidiellum with a long and massive
apical horn.
Original description: Shell with four segments. Cephalis
cylindrical and seems incorporated into the proximal part
of the long, stout apical horn. Diameter of base of horn
same as cephalis; decreasing progressively to the generally
sharp tip. Sutural pore observed at base of horn. Thorax
small, without pores proximally and smooth or with small
costae. Abdomen wide, sphaeroidal, porous and without
costae. One row of large pores separates abdomen form
fourth segment which is seldom preserved. Fourth segment
made of thick pore frames with a loose network.
Original remarks: This form is close to E. gujoensis
(Takemura and Nakaseko), but differs mainly by having
a much stouter and longer apical horn (horn includes
cephalis). E. gujoensis is known only from the Middle
Jurassic while E. gunense n. sp. appears during the Lower
Jurassic. This new morphotype shows apparent affinities
with species attributed to genus Thetis De Wever (Thetis
undulata De Wever, 1982a, p. 196, pl. 5, fig. 8, 9, ?Thetis
oblonga De Wever, 1982a, p. 196, pl. 4, fig. 15-16); cephalic
structure of Thetis has not been observed by its author, but
this genus possesses three external cephalic spines, which
are probably external extensions of inner cephalic spines
(De Wever, personal communication, 1987). However, the
cephalic structure of Eucyrtidiellum does not have these
three spines; this implies true phylogenetic differences
156
between Eucyrtidiellum and Thetis in spite of the common
external resemblance of certain species.
Further remarks: Most specimens from Oman and Japan
included in this species do not correspond perfectly to
the original diagnosis and differ significantly from the
holotype, but to a lesser extent from the paratype. The
apical horn is shorter and conical in our specimens, and
the thorax is larger, well marked by the cortical and lumbar
constrictions and is either truncate conical or convex in
outline. The thorax is also completely porous whereas the
original description mentions that it is poreless proximally.
The abdomen is also different: the holotype shows a
sphaeroidal abdomen but the abdomen of the paratype
is also truncated conical. The original description also
mentions a row of large pores separating the abdomen from
the postabdominal segment. Our study shows that there
are no such pores at the postlumbar boundary, but false
pores (when existing) result from some ribs superposed
on the test in different directions, most in a longitudinal
direction. The fourth segment is wide open and inverted
truncate conical, in some specimens (pl. EUC03, figs. 3, 4)
it is almost closed distally.
One of the characteristic features of this species, besides
those mentioned in the original description or in the
paragraph above, is that the abdominal pores are commonly
aligned in oblique rows and, consequently, most are
rhombically framed. Also the surface of the abdomen may
have some oblique ribs representing thickened intervening
bars.
E. gunense is very close to Thetis oblonga but differs in
having the cephalis less marked, and lacking thoracic
spines, and in having thick ribs on the entire postabdominal
segment or only proximally around the postlumbar
stricture.
Etymology: From the name of a local stream Gun.
Type locality: Locality GSC C-300407, Bridge River
Complex, Carpenter Lake, British Columbia.
Occurrence: Bridge River Complex, British Columbia;
Hyde Formation, Oregon; Franciscan Complex, California;
Skrile Formation, Slovenia; Tawi Sadh Member of the
Guwayza Formation, Oman; Japan.
Plate EUC03. Eucyrtidiellum gunense gr. Cordey. Magnification x300. Fig. 1(H). Cordey 1998, pl. 25, fig. 8.
Fig. 2. OR600A-R03-01. Fig. 3. SI, MM11.76, 010211. Fig. 4. OM-99-83, 011312. Fig. 5. OM, BR1121-R06-25.
Fig. 6. OM, BR1121-R08-27. Fig. 7. OM, BR1121-R08-03. Fig. 8. OM; BR524-R04-19. Fig. 9. OM, BR1122-R02-1.
Fig. 10. OM, BR1122-R02-14. Fig. 11. OM; BR1121-R08-21. Fig. 12. Matsuoka 2004, fig. 180.
Fig. 13. OM, BR523-R03-08. Fig. 14. OM, BR1121-R08-24. Fig. 15. OM, BR1121-R08-06.
157
Eucyrtidiellum nagaiae Dumitrica, Goričan & Matsuoka n. sp.
Species code: EUC06
Synonymy:
1986 Eucyrtidiellum sp. C2 – Nagai, pl. 2, fig. 12.
1987 Eucyrtidiellum sp. C2 – Hattori, pl. 12, fig. 12.
1989 Eucyrtidiellum spp. – Hori & Otsuka, pl. 4, fig. 3, not figs. 1, 2.
1990 Eucyrtidiellum sp. C2 – Nagai, pl. 4, figs. 1a-c.
1995 Eucyrtidiellum sp. C2 – Nagai, pl. 4, figs. 7a-b, 8.
1997 Eucyrtidiellum sp. X – Yao, pl. 10, fig. 456.
1997 Eucyrtidiellum sp. Y – Yao, pl. 10, fig. 457.
2003 Eucyrtidiellum sp. C2 sensu Nagai – Kashiwagi & Kurimoto,
pl. 4, figs. 8, 9.
2004 Eucyrtidiellum sp. – Matsuoka, fig. 179.
Type designation: Holotype
specimen
MA13547
(pl. EUC06, figs. 1a,b) from sample MNA-10, Nanjo Massif, Mino Terrane, Japan.
Diagnosis: Eucyrtidiellum with stout apical horn and strong
longitudinal abdominal costae separated by 2-4 longitudinal rows of pores.
Description: Test composed of three segments. Cephalis
smooth, poreless, incorporated into a long, stout, pointed
apical horn, circular in cross-section throughout length.
Sutural pore situated in the proximal part of the cephalis.
Thorax larger, trapezoidal in outline, with a regular meshwork of polygonal pore frames. Abdomen inflated, porous,
ornamented with 7-9 strong, continuous vertical costae visible laterally. Pores on abdomen circular; number of pore
rows between adjacent costae may vary from two (rarely
one) to four on a single specimen.
Remarks: Eucyrtidiellum nagaiae n. sp. differs from
E. ramescens Cordey by having well-individualized costae
throughout the abdomen. The wall of the abdomen is
otherwise smoother, perforate but the pore frames do not
form coalescent thickenings as in E. ramescens and the
apical horn is shorter. The stout apical horn and the rows
of intercostal pores also differentiate this species from
E. ptyctum (Riedel & Sanfilippo).
Measurements (µm):
Based on 8 specimens.
Height of cephalis and apical horn
Width of cephalis at base
Height of thorax
Maximum width of thorax
Height of abdomen
Maximum width of abdomen
HT Min. Max. Mean
86 37
86
57
40 20
43
29
24 20
32
25
58 40
58
48
80
107 47
114
118
95
112 77
Etymology: Named for Hiromi Nagai, Nagoya University,
to honour her contribution to the knowledge of Mesozoic
Radiolaria and especially of Eucyrtidiellum.
Type locality: Sample MNA-10, Nanjo Massif, Mino
Terrane, Japan.
Occurrence: Nanjo Massif, Mino Terrane, Japan; Hyde
Formation, Oregon; Musallah Formation, Oman.
Eucyrtidiellum omanojaponicum Dumitrica, Goričan & Hori n. sp.
Species code: EUC07
Synonymy:
1990 Eucyrtidiellum (?) sp. C group – Hori, fig. 8. 27.
1997 Eucyrtidiellum sp. C group of Nagai – Hori, pl. 1, fig. 12.
Type designation: Holotype specimen 15862 (pl. EUC07,
fig. 1) from sample BR1122, Tawi Sadh Member of the
Guwayza Formation, Wadi Mu’aydin, Oman.
Diagnosis: Eucyrtidiellum with numerous thin, longitudinal abdominal costae and a row of pores in each intercostal
depression.
Description: Test composed of four segments of which only
the first three are completely preserved. Cephalis integrated
into the proximal part of the long, stout conical apical horn
and visible as a slightly inflated portion at the base of the
latter. Diameter of horn at base the same as cephalis, decreasing progressively to the sharp tip. Thorax small, truncate conical with convex sides and numerous small pores
disposed irregularly. Abdomen wide, subglobular and porous; pores circular aligned in longitudinal rows, each row
separated by thin costae. Fourth segment preserved only
as remains.
158
Remarks: The species is very rare and is described based
on 3 specimens only. E. omanojaponicum n. sp. resembles
E. nagaiae n. sp. in having longitudinal abdominal costae,
but the costae are weaker and separated by a single row
of pores whereas E. nagaiae has 2-4 rows of pores in the
intercostal intervals.
Measurements (µm):
Based on 3 specimens.
Length of apical horn + cephalis
Length of thorax
Length of abdomen
Diameter of cephalis
Diameter of thorax
Diameter of abdomen
HT Min. Max.
59 38
84
27 26
30
65 54
86
24 24
27
55 45
55
93 83
105
Etymology: From its occurrence in Oman and Japan.
Type locality: Sample BR 1122 from theTawi Sadh Member
of the Guwayza Formation, Wadi Mu’aydin, Oman.
Occurrence: Tawi Sadh Member of the Guwayza Formation,
Oman; Japan.
Plate EUC06. Eucyrtidiellum nagaiae Dumitrica, Goričan & Matsuoka n. sp. Magnification x300. Fig. 1a,b(H).
JP, MNA-10, MA13547, MA13548. Fig. 2. JP, MNA-10, MA12501. Fig. 3. OM-00-255, 022326. Fig. 4. JP, MNA-10,
MA12499. Fig. 5. OM-00-254, 022126. Fig. 6. Hori & Otsuka 1989, pl. 4, fig. 3.
Plate EUC07. Eucyrtidiellum omanojaponicum Dumitrica, Goričan & Hori n. sp. Magnification x300.
Fig. 1(H). OM, BR1122-R04-04. Fig. 2. Hori 1990, fig. 8-27. Fig. 3. OM-00-92, 011110.
159
Eucyrtidiellum ramescens Cordey 1998
Species code: EUC04
Synonymy:
1982 Eucyrtidium (?) sp. C – Yao et al., pl. 2, fig. 12.
1982 »Eucyrtidium« sp. – Nishizono et al, pl. 2, fig. 13.
1986 Eucyrtidiellum sp. C1 – Nagai, pl. 2, fig. 11.
1987 Eucyrtidiellum sp. C1 – Hattori, pl. 12, fig. 11.
1995 Eucyrtidiellum sp. C1 – Nagai, pl. 4, fig. 5.
1998 Eucyrtidiellum ramescens n. sp. – Cordey, p. 110, pl. 25,
figs. 7, 11-13.
2004 Eucyrtidiellum sp. C1 sensu Nagai – Hori, pl. 5, fig. 24.
2005 Eucyrtidiellum sp. C sensu Nagai – Hori, pl. 8, fig. 19, not
figs. 16-18.
Original diagnosis: Eucyrtidiellum with a very long apical
horn and massive, porous abdominal costae.
Original description: Test composed of four segments.
Cephalis cylindrical and appears integrated into the
proximal part of the long, stout apical horn. Diameter of
horn at base, the same as cephalis, decreasing progressively
to the sharp tip. Sutural pore situated in proximal part
or near the collar edge. Thorax small, without pores in
proximal part and smooth or with small costae. Abdomen
wide, sphaeroidal and porous; pores circular without
specific alignment. Some pore-frames reinforced near
proximal part of abdomen and become thicker distally
giving rise to longitudinal costae, sometimes transversally
connected. One row of large pores separates abdomen
from fourth segment which is seldom preserved. Fourth
segment composed of thick pore frames with a loose
network.
Original remarks: This form differs from E. gunensis n. sp.
by the presence of thick abdominal and post-abdominal
costae.
Etymology: From Latin ramus (branch).
Type locality: Locality GSC C-300407, Bridge River Complex, lake Carpenter, British Columbia.
Occurrence: Bridge River Complex, British Columbia;
Musallah Formation, Oman; Japan.
Genus: Farcus Pessagno, Whalen & Yeh 1986
Type species: Farcus graylockensis Pessagno, Whalen & Yeh 1986
Synonymy:
1986 Farcus n. gen. – Pessagno, Whalen & Yeh, p. 23.
Original diagnosis: Test as with family but possessing
a single, massive apical horn that is attached to the apical
bar. Thorax lacking tubular, velum-like structure distally.
Original description of family Farcidae (Pessagno, Whalen
& Yeh 1986, p. 22): Test dicyrtid with single layer of latticed
meshwork on both cephalis and thorax. Latticed layer of
cephalis and occasionally proximal portion of thorax covered by thin outer layer of microgranular silica. Cephalis
large, hemispherical with one horn (e.g., Farcus n. gen.), or
two horns (e.g., Rolumbus n. gen.), which are triradiate in
axial section. Cephalic skeletal elements cyrtoid, including
vertical bar, primary left lateral bar, primary right lateral
bar, median bar, secondary left lateral bar, secondary right
lateral bar, and apical bar (dorsal bar absent). Thorax large,
inflated, with four (rarely five) feet that are triradiate in
axial section. Four feet opposed to two primary lateral and
160
two secondary lateral bars; fifth foot, if present, opposed to
vertical bar. Base of thorax hemispherical with centrallyplaced circular aperture (mouth) that has an imperforate
rim. Thorax with (e.g., Rolumbus, n. gen.) or without (e.g.,
Farcus, n. gen.) fragile tubular, velum-like structure extending distally from aperture (mouth) of well-preserved
specimens.
Original remarks: Farcus n. gen., is compared to Rolumbus
n. gen., under the latter genus.
Etymology: Farcus (masc.) is a name formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, Pt. VI,
Recommendation 4, p. 113).
Included species:
FAR02 Farcus asperoensis Pessagno, Whalen & Yeh 1986
FAR04 Farcus graylockensis Pessagno, Whalen & Yeh 1986
FAR03 Farcus kozuri Yeh 1987b
Plate EUC04. Eucyrtidiellum ramescens Cordey. Magnification x300. Fig. 1(H). Cordey 1998, pl. 25, fig. 7.
Fig. 2. OM-00-118, 000702.
161
Farcus asperoensis Pessagno, Whalen & Yeh 1986
Species code: FAR02
Synonymy:
1986 Farcus asperoensis n. sp. – Pessagno, Whalen & Yeh, p. 23,
pl. 3, figs. 12, 16, 17, 21; pl. 11, fig. 9.
2002 Farcus asperoensis Pessagno, Whalen & Yeh – Whalen &
Carter, p. 124, pl. 11, figs. 3, 11, 15.
Original diagnosis: Cephalis medium-sized, hemispherical, with single, massive triradiate horn; cephalis commonly covered by layer of microgranular silica. Horn triradiate in axial section with rounded, longitudinal ridges and
narrow grooves. Thorax with small, polygonal pore frames,
commonly partially covered by a thin layer of microgranular silica. Four feet, medium-sized, triradiate in axial section with narrow, rounded longitudinal ridges and broad
grooves. Four feet commonly attached to base of thorax,
although on some specimens two of the feet are attached
part way up the thorax. Circular mouth surrounded by imperforate rim.
Original remarks: Farcus asperoensis n. sp., differs from
other species of Farcus by the nature of the distinctive
imperforate rim surrounding the mouth. An undescribed
species of Farcus (Pl. 11, fig. 14) from the Hyde Formation
of east-central Oregon possesses an elongated thorax and
massive horn similar to those of F. asperoensis, n. sp.
Measurements (µm):
Numbers of specimens measured are in parentheses.
length of cephalis
length of thorax
width of thorax at top
width of thorax at base
length of horn
width of horn at base
length of foot (maximum)
HT
20
80
48
50
48
13
32
Mean
22.3 (9)
80 (9)
50.5 (9)
61.2 (8)
64.6 (9)
20.4 (9)
61.3 (9)
Max.
30 (9)
90 (9)
64 (9)
70 (8)
80 (9)
25 (9)
80 (9)
Min.
18 (9)
70 (9)
43 (9)
50 (8)
48 (9)
13 (9)
32 (9)
Etymology: This species is named for Pico Aspero, which is
located east of its type area.
Type locality: Sample SH-412-14, San Hipólito Formation,
Vizcaino Peninsula, Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Musallah Formation, Oman.
Farcus graylockensis Pessagno, Whalen & Yeh 1986
Species code: FAR04
Synonymy:
1986 Farcus graylockensis n. sp. – Pessagno, Whalen & Yeh,
p. 24, pl. 2, figs. 4, 6-8, 12, 15.
1987b Farcus graylockensis Pessagno, Whalen & Yeh – Yeh,
p. 76, pl. 1, fig. 7.
1996 Farcus graylockensis Pessagno, Whalen & Yeh – Pujana,
p. 139, pl. 1, fig. 7.
1997 Farcus graylockensis Pessagno, Whalen & Yeh – Yao, pl. 8,
fig. 395.
1997 Farcus aff. kozuri Yeh – Yao, pl. 8, fig. 396.
2002 Farcus graylockensis Pessagno, Whalen & Yeh – Tekin,
p. 189, pl. 4, fig. 2.
Original diagnosis: Single horn wide, massive, triradiate in
axial section; horn comprised of three wide, wedge-shaped
grooves alternating with three wide, rounded ridges. Thorax with massive, uniformly-sized tetragonal and pentagonal pore frames. Feet moderately long, triradiate in axial
section; three longitudinal grooves deep, wider proximally
than distally, wedging out distally; three longitudinal ridges
rounded, becoming progressively narrower distally.
Original remarks: This species differs from similar forms
among the Hilarisiregidae Takemura and Nakaseko, 1982,
by possessing a dicyrtid test, and considerably different
wall structure (see Pl. 1, figs. 4, 5, 10).
162
Measurements (µm):
Based on 10 specimens.
length of cephalis
length of thorax
width of thorax at top
width of thorax at base
length of horn
width of horn at base
length of foot (maximum)
HT
25
100
62.5
112.5
70
25
95
Mean
25.7
92.5
63.7
106.2
72.7
23.5
96.5
Max.
30
100
87.5
120
87.5
25
125
Min.
20
75
50
100
62.5
20
75
Etymology: Farcus graylockensis n. sp., is named for Graylock Butte, which is located north of its type locality.
Type locality: OR-536, Nicely Formation, southeast side of
Morgan Mountain, east-central Oregon.
Occurrence: Nicely Formation, Oregon; Sierra Chacaicó
Formation, Argentina; Hocaköy Radiolarite, Turkey; Tawi
Sadh Member of the Guwayza Formation, Oman; Japan.
Plate FAR02. Farcus asperoensis Pessagno, Whalen & Yeh. Magnification x300. Fig. 1(H). Pessagno, Whalen & Yeh
1986, pl. 3, fig. 12. Fig. 2. Pessagno, Whalen & Yeh 1986, pl. 3, fig. 16. Fig. 3. OM-00-251, 021607.
Plate FAR04. Farcus graylockensis Pessagno, Whalen & Yeh. Magnification x300. Fig. 1(H). Pessagno, Whalen & Yeh
1986, pl. 2, fig. 6. Fig. 2. OM, BR1122-R01-09.
163
Farcus kozuri Yeh 1987b
Species code: FAR03
Synonymy:
1986 Farcus sp. A – Pessagno, Whalen & Yeh, p. 24, pl. 3, fig. 4.
1986 Farcus sp. B – Pessagno, Whalen & Yeh, p. 24, pl. 3, fig. 13.
1987b Farcus kozuri n. sp. – Yeh, p. 75, pl. 1, figs. 2, 6, 13.
Original description: Test dicyrtid, cephalis medium in
size, hemispherical, imperforate, covered with layer of
microgranular silica. Single horn moderately massive,
triradiate with three narrow ridges alternating with three
wide grooves. Thorax hemispherical, with slightly variable
size of massive polygonal pore frames. Distal portion of
thorax with three to four prominent transverse ridges,
ridges continuous or slightly offset, parallel to each other
and merging with those of feet. Feet massive, medium in
length, tapering distally, triradiate in axial section with
three wide, deep longitudinal grooves alternating with
three narrow longitudinal ridges.
Original remarks: Farcus kozuri, n. sp., differs from F. graylockensis Pessagno, Whalen, and Yeh (1986) by possessing
three to four well-developed transverse ridges on the distal
portion of thorax.
Measurements (µm):
Based on 10 specimens.
Length of cephalis
Length of thorax
width of thorax at top
width of thorax (maximum)
length of apical horn
length of foot (maximum)
HT
29
78
43
114
78
100
Mean
29
84
42
112
65
90
Max.
30
78
43
114
78
100
Min.
28
90
40
108
36
78
Etymology: This species is named for Dr. H. Kozur, in
honor of his studies on Mesozoic Radiolaria.
Type locality: Sample OR-536J, Nicely Formation, southeast side of Morgan Mountain, east-central Oregon.
Occurrence: Nicely Formation, Oregon; Tawi Sadh Member of the Guwayza Formation, Oman.
Genus: Foremania Whalen & Carter 1998
Type species: Foremania sandilandsensis Whalen & Carter 1998
Synonymy:
1998 Foremania n. gen. – Whalen & Carter, p. 79.
Original description: Test elongate, cylindrical with singlelayered wall. Cephalis large, dome shaped with irregularly
branching horn. Post-abdominal chambers generally
rectangular in outline; first few chambers very gradually
increasing in size as added; post-abdominal chambers
gradually decreasing in width producing an open tube.
Regularly shaped pore frames on most post-abdominal
chambers; pore frames more regular in shape (tetragonal)
and aligned in distinct horizontal and vertical rows on distal
portion of test; pore frames smaller on more proximal part
of test, larger and more regularly shaped on distal part of
test.
164
Original remarks: The narrow but open tubular extension
of the test and the very regularly shaped and aligned pore
frames on the distal post-abdominal chambers distinguish
Foremania n. gen. from Pseudoeucyrtis Pessagno. The
single-layered wall distinguishes Foremania n. gen. from
Canutus Pessagno and Whalen and Droltus Pessagno and
Whalen.
Etymology: This genus is named in memory of Helen
Foreman a noted scholar of fossil Radiolaria.
Included species:
FRM01 Foremania sandilandsensis gr. Whalen & Carter
1998
Plate FAR03. Farcus kozuri Yeh. Magnification x200. Fig. 1(H). Yeh 1987b, pl. 1, fig. 6. Fig. 2. Pessagno, Whalen & Yeh
1986, pl. 3, fig. 4. Fig. 3. Pessagno, Whalen & Yeh 1986, pl. 3, fig. 13.
165
Foremania sandilandsensis gr. Whalen & Carter 1998
Species code: FRM01
Synonymy:
1982a gen. et sp. indet. 1 – De Wever, p. 220, pl. 13, figs. 13-14.
1982b gen. et sp. indet. 1 – De Wever, p. 354, pl. 56, figs. 10-11.
? 1990 Nassellaria gen. et sp. indet. – De Wever et al., pl. 4, fig. 2.
1990 Gen. sp. indet. 1 in De Wever 1982 – De Wever et al.,
pl. 4, fig. 3.
1998 Foremania sandilandsensis n. sp. – Whalen & Carter, p. 79,
pl. 24, figs. 14, 15, 18-21, 24-26.
Original description: Test elongate, cylindrical, commonly
with ten postabdominal chambers. Cephalis large, domeshaped with horn; horn irregularly shaped, branching,
usually with two or three prongs varying in size. Cephalis
and thorax mostly imperforate, almost completely
covered by layer of microgranular silica. Thorax and
abdomen trapezoidal in outline. Most postabdominal
chambers subrectangular to square in outline, very
gradually increasing in width and height as added; last
few postabdominal chambers gradually decreasing in
width. On well preserved specimens, test terminating in a
narrow, open tube. Pore frames on initial chambers of test
small and irregularly shaped; pore frames on test gradually
increasing in size as added, becoming much more regularly
tetragonal and aligned in distinct horizontal and vertical
rows on distal part of test; small rounded nodes at vertices
of all pore frames.
Original remarks: Foremania sandilandsensis is the only
species of Foremania yet recognized. Since the test is
very large and seldom complete, the observed number of
postabdominal chambers depends on preservation.
Further remarks: The size of this species is very variable.
The length of the specimens illustrated herein varies from
140µm (pl. FRM01, fig 1(H)) to 530µm (pl. FRM01, fig.
8a).
Measurements (µm):
Based on 5 specimens.
Length
140
227
140
200
Max. width
26
103
26
80
HT
Max.
Min.
Mean
Etymology: This species is named for Sandilands Island,
located in Skidegate Inlet, north of South Bay, Queen
Charlotte Islands, British Columbia.
Type locality: Sample QC 590A. Kunga Island, north side,
Sandilands Formation, Queen Charlotte Islands, British
Columbia, Canada.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands; Dürrnberg Formation, Austria;
Gümüslü Allochthon, Turkey; Tawi Sadh Member of the
Guwayza Formation and Musallah Formation, Oman;
Japan.
Genus: Gigi De Wever 1982a
Type species: Gigi fustis De Wever 1982a
Synonymy:
1982a Gigi n. gen. – De Wever, p. 194.
1988 Gigi De Wever – Hori, p. 558.
Original description: Two-segmented test with a long porous closed tube. Apical horn small, simple or forked. On
the cephalis, or at the collar stricture, one (or several ?)
spines corresponding to the external prolongation of cephalic spines are present. Cephalic skeleton not in collar
plane.
Revised description: By Hori (1988): Test comprising 2
to 3 chambers, cephalis, thorax or abdomen, or both and
closed tube, without radial spines. The proximal portion
subspherical, having smooth surface and circular pores
with a crown-like apical horn and thorn(s). Tube porous
and elongated cylindrically, sometimes weakly expanded
near the distal end.
166
Original remarks: This genus differs from Katroma by having only two segments, very small cephalic horns (except
the apical horn) and smaller pores. It differs from Podobursa
by the number of segments and the structure of the pores.
This genus could be considered as older than Katroma (P.
Dumitrica, pers. com.).
Further remarks: Contrary to the assumption in the original remarks, our new data show that Katroma appears prior
to Gigi.
Etymology: Name formed by an arbitrary combination of
letters (ICZN, art. 40, p. 113).
Included species:
GIG01 Gigi fustis De Wever 1982a
Plate FRM01. Foremania sandilandsensis gr. Whalen & Carter. Magnification Figs. 1-7 x300 (scale bar A), Figs. 8-16
x200 (scale bar B), except Fig. 8b x400. Fig. 1(H). Carter et al. 1998, pl. 24, fig. 20. Fig. 2. Carter et al. 1998, pl. 24, fig. 19.
Fig. 3. Carter et al. 1998, pl. 24, fig. 18. Fig. 4. OM; BR1121-R10-06. Fig. 5. OM, BR1121-R10-11. Fig. 6. OM, BR1121R08-18. Fig. 7. QCI, GSC loc. C-305417, GSC 111808. Fig. 8a,b. TR, 1662D-R03-12. Fig. 9. TR, 1662D-R02-13.
Fig. 10. JP, IYII8-85. Fig. 11. OM-00-252, 022012. Fig. 12. AT, BMW21-17. Fig. 13. OM-00-118, 000612.
Fig. 14. OM, BR528-R10-07. Fig. 15. OM, BR524-R04-24. Fig. 16. OM, BR706-R12-03.
167
Gigi fustis De Wever 1982a
Species code: GIG01
Synonymy:
1982a Gigi fustis n. sp. – De Wever, p. 195, pl. 4, figs. 1-8.
1982b Gigi fustis De Wever – De Wever, p. 340, pl. 57,
figs. 1-6, 12.
1982 Gigi fustis De Wever – De Wever & Origlia-Devos, pl. 1,
fig. F.
1988 Gigi sp. aff. G. fustis De Wever – Hori, p. 559, fig. 9.1, 2.
1988 Katroma dengqenesis n. sp. – Li, p. 328, pl. 1, fig. 18,
not fig. 9.
1993 Katroma sp. – Kashiwagi & Yao, pl. 1, fig. 10.
1993 Gigi fustis De Wever – Kashiwagi & Yao, pl. 1, fig. 11.
1994 Gigi fustis De Wever – Goričan, p. 70, pl. 16, figs. 11-13.
1995 Gigi fustis De Wever – Suzuki, pl. 8, fig. 5.
1997 Gigi cf. G. fustis De Wever – Hori, pl. 1, fig. 20.
1997 Gigi sp. cf. G. fustis De Wever – Hori et al., fig. 2.3.
1998 Gigi fustis De Wever – Kashiwagi, pl. 2, figs. 13, 14.
2001 Gigi fustis De Wever – Kashiwagi, fig. 6.4.
2004 Gigi fustis De Wever – Matsuoka, fig. 116.
Original description: Two-segmented test with a long
closed tube and a very small clove-shaped apical horn.
Horn sometimes simple and not branched when poorly
preserved. Cephalis conical, porous. Cephalic skeleton
massive, especially at the level of median bar. One spine of
the cephalic skeleton (probably V) outgrowing as a small
horn. Thorax globular with small pores. Thorax extending
in a long tube with widely open proximal pores and smaller
distal pores; the latter are aligned along the tube and show
a tendency towards helicoidal distribution. Two keels seem
to be developed distally.
Original remarks: This species differs from Katroma neagui,
K. bicornus and K. sp. A, K. sp. B, by having a simpler
outline, smaller horns (apical and lateral), and especially by
its two-segmented test.
Measurements (µm):
Based on 14 specimens.
Total length
(including the apical horn)
Tube length
Tube width
Length of cephalis plus thorax
Maximum width of thorax
HT
Max.
Min.
Mean
450
490
360
443
320
24
107
79
390
33
170
110
260
24
90
72
315
28
120
88
Etymology: From the Latin fustis, -is, m. = bludgeon,
referring to general shape.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Budva Zone,
Montenegro; Pindos Zone, Greece; Haliw (Aqil) Formation,
Oman; Dengqen area, Tibet; Japan; New Zealand.
Genus: Gorgansium Pessagno & Blome 1980
Type species: Gorgansium silviesense Pessagno & Blome 1980
Synonymy:
1980 Gorgansium n. gen. – Pessagno & Blome, p. 234.
Original description: Cortical shell typically elliptical
with 3 primary spines of unequal length usually occurring
in same plane. Primary spines asymmetrically arranged;
2 spines closer together, often considerably shorter than
third spine. Cortical shell usually compressed in plane of 3
primary spines. First medullary shell small, spherical with
fragile pore frames.
Original remarks: Gorgansium, n. gen. differs from Betraccium Pessagno, (Pessagno et al., 1979), in the asymmetri168
cal arrangement and unequal length of its primary spines.
Whereas Betraccium has symmetrically arranged, more or
less equidistant spines of equal length, Gorgansium has its 2
shorter spines situated close together.
Etymology: Gorgansium is a name formed by an arbitrary
combination of letters (ICZN, 1964. Appendix D. Pt. 6.
recommendation 40. p.113). The gender of this genus is
neuter.
Included species:
GOR02 Gorgansium gongyloideum Kishida & Hisada 1985
GOR03 Gorgansium morganense Pessagno & Blome 1980
Plate GIG01. Gigi fustis De Wever. Magnification x300, except Fig. 1b(H) x500. Fig. 1a,b(H). De Wever 1982a, pl. 4,
figs. 1, 2. Fig. 2. TR, 1662D-R02-08. Fig. 3. Matsuoka 2004, fig. 116. Fig. 4. JP, Nanjo chert, NA-16, RH(1)394.
Fig. 5. JP, Nanjo chert, NAI-76, RH(1)442. Fig. 6. JP, Nanjo chert, NA-11-12, RH(1)391. Fig. 7. OM, Haliw-039-R02-05.
Fig. 8. OM, Haliw-038-R09-23. Fig. 9. OM, Haliw-038-R08-08. Fig. 10. OM, Haliw-039-R02-06. Fig. 11. OM, Haliw038-R08-19. Fig. 12. OM, Haliw-039-R06-20. Fig. 13. OM, Haliw-039-R02-08. Fig. 14. OM, Haliw-039-R06-28.
169
Gorgansium gongyloideum Kishida & Hisada 1985
Species code: GOR02
Synonymy:
1982 Gorgansium sp. A – Kishida & Sugano, pl. 4, fig. 8.
1984 Gorgansium sp. aff. morganense Pessagno & Blome
– Whalen & Pessagno, pl. 1, figs. 15-16.
1985 Gorgansium gongyloideum n. sp. – Kishida & Hisada,
p. 116, pl. 1, figs. 21-22.
1986 Gorgansium gongyloideum Kishida & Hisada – Kishida &
Hisada, Fig. 4.4.
1990 Gorgansium gongyloideum Kishida & Hisada – Hori, Fig. 8.6.
1994 Gorgansium gongyloideum Kishida & Hisada – Goričan,
p. 70, pl. 1, fig. 6.
1998 Gorgansium gongyloideum Kishida & Hisada – Yeh &
Cheng, p. 12, pl. 1, fig. 1.
2002 Gorgansium gongyloideum Kishida & Hisada – Whalen &
Carter, p. 105, pl. 6, figs. 3-5, 9-11.
2002 Gorgansium gongyloideum Kishida & Hisada – Tekin,
p. 179, pl. 1, fig. 4.
2003 Gorgansium spp. – Goričan et al., p. 291, pl. 1, fig. 7.
Original diagnosis: Cortical shell spherical, with three
three-bladed primary spines. Two spines somewhat shorter than third spine. Third spine as long as diameter of
cortical shell.
Original description: Cortical shell spherical, with predominantly hexagonal pore frames lacking well-developed nodes at vertices. Thickness of pore frame bars in
Z direction about two times as thick as in Y direction (Textfig. 5). Seven pore frames visible on test surface along AB,
six to seven pore frames visible along CD (Text-fig. 5). Primary spines triradiate in axial section; composed of three
narrow grooves alternating with three moderately wide
ridges longitudinally. Two primary spines nearly equal in
length; third spine somewhat longer. Third spine as long as
diameter of cortical shell.
Original remarks: Gorgansium gongyloideum n. sp. differs
from Gorgansium crassum n. sp., in having cortical shell
without compression, more numerous pore frames and
primary spines with wider ridges.
Measurements (µm):
Based on 5 specimens.
System of measurements shown in Text-fig. 5 of Kishida &
Hisada (1985).
AB
78
81
78
79
CD
81
83
81
82
AT
80
88
78
82
EF
61
69
56
62
GH
59
68
57
61
HT
Max.
Min.
Av.
Etymology: The name is derived from the Latin adjective
gongylis + deus, meaning spherical.
Type locality: Locality 230 of Kishida & Hisada (1985),
Ueno-mura area, Kanto Mountains, Central Japan.
Occurrence: Japan; Fernie Formation, northeastern British Columbia; San Hipólito Formation, Baja California
Sur; Skrile Formation, Slovenia; Budva Zone, Montenegro;
Hocaköy Radiolarite, Turkey; Liminangcong Chert, Philippines.
Gorgansium morganense Pessagno & Blome 1980
Species code: GOR03
Synonymy:
1980 Gorgansium morganense n. sp. – Pessagno & Blome,
p. 234, pl. 6, figs. 10, 18, 23.
Original description: Cortical shell circular in outline
compressed somewhat in plane of spines and comprised
of large pore frames with poorly developed nodes at the
vertices. Bars of pore frames of medium thickness along Y;
thicker along Z (text-fig. 5). Five to 6 pore frames visible on
test surface along AB; 5 pore frames visible along CD (see
pl. 8, fig. 16). Two primary spines nearly equal in length;
third spine longer. All 3 spines triradiate in axial section,
comprised of 3 massive, wide ridges alternating with 3
moderately wide grooves longitudinally; grooves somewhat
wider than ridges.
Original remarks: This species differs from G. silviesense,
n. sp., in having a cortical shell that is circular rather than
elliptical in outline and in having much longer primary
spines.
170
Measurements (µm):
Based on 7 specimens.
System of measurement shown in plate 8, figure 16 of
Pessagno & Blome (1980).
AB
94
96
106
88
CD
75
81
113
75
AT
94
93
106
81
EF
69
55
75
56
GH
75
65
88
63
HT
Av.
Max.
Min.
Etymology: G. morganense is named for Morgan Mountain
near its type locality.
Type locality: Sample OR 536, Nicely Formation, northeast
side of Morgan Mountain, Oregon.
Occurrence: Nicely Formation, Oregon; Fannin Formation,
Queen Charlotte Islands.
Plate GOR02. Gorgansium gongyloideum Kishida & Hisada. Magnification x300. Fig. 1(H). Kishida & Hisada 1985,
pl. 1, fig. 21. Fig. 2. JP, MNA-10, MA11018. Fig. 3. Whalen & Carter 2002, pl. 6, fig. 3. Fig. 4. Whalen & Carter 2002,
pl. 6, fig. 5. Fig. 5. Whalen & Carter 2002, pl. 6, fig. 4.
Plate GOR03. Gorgansium morganense Pessagno & Blome. Magnification x300. Fig. 1(H). Pessagno & Blome 1980,
pl. 6, fig. 10. Fig. 2. QCI, GSC loc. C-304566, GSC 128796.
171
Genus: Haeckelicyrtium Kozur & Mostler 1979, emend. Carter 1993
Type species: Haeckelicyrtium austriacum Kozur & Mostler 1979
Synonymy:
1979 Haeckelicyrtium n. gen. – Kozur & Mostler, p. 98.
1993 Haeckelicyrtium Kozur & Mostler – Carter, p. 96.
1997 Haeckelicyrtium Kozur & Mostler – Sugiyama, p. 154-155.
Original description: Cephalis imperforate without apical
horn. Thorax very wide, cap-shaped, with rough pores,
which can be completely closed in the proximal part by
an imperforate layer. Abdomen short, very strongly and
abruptly flaring into a disc distally. Distal border smooth
in the type species, but with short, wide and blunt spines
distributed in the disc plane in Haeckelicyrtium? spinosum
n. sp. Aperture wide, circular. Internal spicule as in family.
Emended description: Carter (1993): Includes forms with
or without an apical horn. Distal rim of abdomen may be
spinose as well as smooth.
Original remarks: Dreyericyrtium n. gen. and Deflandrecyrtium n. gen. have an apical horn. Dreyericyrtium is
however considerably more slender, and Deflandrecyrtium
has a two-segmented conical cephalis as well as laterally
and downwardly directed spines at the distal end of the
abdomen.
Further remarks: Sugiyama writes that based on
evidence from Hori (1992) and from diverse Lower and
Middle Jurassic assemblages of Yao (1997), the genus
Haeckelicyrtium disappears at the end of the Triassic.
However, Carter et al. (1998) found Haeckelicyrtium sp.
A in Hettangian/Sinemurian strata of the Sandilands
Formation, Queen Charlotte Islands, Whalen and Carter
(2002) found Haeckelicyrtium sp. B in Baja California Sur,
and abundant Haeckelicyrtium (H. crickmayi described
herein) are present in Pliensbachian formations of Queen
Charlotte Islands. Rare specimens of Haeckelicyrtium are
also known from the Fernie Formation of northeastern
British Columbia.
Etymology: Named for E. Haeckel, the famous pioneer of
radiolarian research.
Included species:
HCK05 Haeckelicyrtium crickmayi Carter n. sp.
HCK04 Haeckelicyrtium sp. B sensu Whalen & Carter 2002
Haeckelicyrtium crickmayi Carter n. sp.
Species code: HCK05
Type designation: Holotype GSC 111719 and paratype GSC
111720 from GSC loc. C-304281, Ghost Creek Formation
(lowermost Pliensbachian).
in having a less distinct boundary between the thorax and
cephalis and in possessing long, net-like arms on the edge
of the skirt.
Description: Cephalis small and globular, imperforate,
without horn. Thorax short, funnel-shaped, with small circular to subcircular pores. Abdomen abruptly flaring to a
wide, downwardly directed net-like skirt composed mostly
of large subcircular to circular pore frames becoming larger
towards periphery. Periphery of skirt most irregular with
outer portions of skirt extending as flared perforate arms of
variable width. All arms of skirt terminating in one or more
short circular spines; narrower arms commonly terminate
with bifurcating spines.
Measurements (µm):
Based on 9 specimens.
Remarks: Haeckelicyrtium crickmayi n. sp. differs from
H. karcharos Carter (1993) in lacking a horn and in possessing long, net-like arms of variable width on the edge
of the skirt that terminate in one or more spines. It differs
from Haeckelicyrtium sp. B of Whalen and Carter (2002)
Diameter of cephalis
Max. diameter of thorax
(excl. arms and spines)
HT
52
Max.
66
Min.
32
Mean
51 (7)
464
469
352
429
Etymology: Named for Colin H. Crickmay, who contributed significantly to the knowledge of Late Triassic and Early
Jurassic ammonoids of British Columbia
Type locality: Ghost Creek Formation, South side of Maude
Island, several hundred metres west of Ells Bay, Skidegate
Inlet, Queen Charlotte Islands, British Columbia.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Fernie Formation, NE British Columbia.
Haeckelicyrtium sp. B sensu Whalen & Carter 2002
Species code: HCK04
Synonymy:
1998 Haeckelicyrtium karcharos Carter – Yeh & Cheng, p. 32,
pl. 12, fig. 8.
2002 Haeckelicyrtium sp. B – Whalen & Carter, p. 122, pl. 16, fig. 9.
172
Original remarks: The pore frames of this species are more
massive than the pore frames of Haeckelicyrtium sp. A of
Whalen & Carter 1998.
Occurrence: San Hipólito Formation, Baja California Sur;
Liminangcong Chert, Philippines.
Plate HCK05. Haeckelicyrtium crickmayi Carter n. sp. Magnification x100. Fig. 1(H). GSC loc. C-304281, GSC 111719.
Fig. 2. QCI, GSC loc. C-304567, GSC 128803. Fig. 3. QCI, GSC loc. C-304281, GSC 111720. Fig. 4. QCI, GSC loc. C304567, GSC 128913. Fig. 5. NBC, GSC loc. C-305208, GSC 128802.
Plate HCK04. Haeckelicyrtium sp. B sensu Whalen & Carter. Magnification x150. Fig. 1. Whalen & Carter 2002,
pl. 16, fig. 9.
173
Genus: Hagiastrum Haeckel 1881
Type species: Hagiastrum plenum Rüst 1885 (subsequent designation by Campbell, 1954)
Synonymy:
arranged subsidiary beams to cortical shell. Medullary shell
as with family, rays circular in cross section, composed of
several medullary beams comprising 3 (sometimes up to
6) primary canals. Central area of medullary shell with
internal vertical beamlets.
Original description: 3c. Tribe: Euchitonida. Porodiscida
with arms, arms chambered situated in the equatorial
plane and radiating from the margin of the disk (often with
terminal spines on the arms, and often with the arms joined
by a patagium or chambered web).
C. With four arms, arranged in the form of a rectangular
cross.
CI. With simple arms.
Ia. Without patagium.
Further remarks: By Baumgartner (1980): Included are
only those four-rayed forms displaying linear arrangement
of beams and pore rows and having an inner structure as in
the subfamily. Species with more irregular pore arrangement
assigned to this genus by Kozur and Mostler (1978) and
Pessagno, Finch and Abbott (1979), should be assigned to
the genus Crucella Pessagno under the Patulibracchiidae
Pessagno, emend.
This volume: Although Hagiastrum is the type genus of
the family Hagiastridae the cross-section of rays of the type
species of Hagiastrum is not yet known. In revising the family, Baumgartner (1980) made no remarks on the number
of beams and rows of pores because of the difficulty to find
a specimen confidently assigned to the type species. Herein
we assign to this genus all species answering more or less
the emended definition without taking into account the ray
structure. However, we note that Hagiastrum macrum De
Wever (see below) has rays with 3 primary and 9 secondary
canals.
1881 Hagiastrum n. gen. – Haeckel, p. 460.
1887 Hagiastrum Haeckel – Haeckel, p. 542.
1971 Hagiastrum Haeckel emend. – Pessagno, p. 52.
1977b Hagiastrum Haeckel emend. – Pessagno, p. 72.
1980 Hagiastrum Haeckel emend. – Baumgartner, p. 289.
Emended description: By Baumgartner (1980): Test as
with subfamily, composed of 4 arms approximately at right
angles. Rays slender elongate usually with bulbous tips,
with or without spines.
Emendation of the subfamily Hagiastrinae Riedel 1971
(Baumgartner, 1980, p. 288): Test as with family, composed
of 2 to 4 rays extending from a central area which is simply
formed by the convergence of the rays. Cortical rays
composed of numerous (8-12) longitudinal external beams,
connected by bars regularly in transverse rows forming
single rows of circular, rectangular or parallelogram-shaped
pores between beams. Cross-section of ray circular or
elliptical. Central area of cortical shell usually with smaller,
more irregular pore frames, nodes may be developed.
Medullary shell centrally placed, about one-third the
diameter of cortical shell, leaving a cylindrical cortical space
around it. Medullary shell connected by numerous radially
Etymology: From the Greek agion = holy, and astron =
starrulet.
Included species:
HAG06 Hagiastrum macrum gr. De Wever 1981b
HAG03 Hagiastrum majusculum Whalen & Carter 1998
HAG04 Hagiastrum rudimentum Whalen & Carter 1998
Hagiastrum macrum gr. De Wever 1981b
Species code: HAG06
Synonymy:
1981b Hagiastrum macrum n. sp. – De Wever, p. 29, pl. 1, figs. 7-9.
1982b Hagiastrum macrum De Wever – De Wever, p. 232,
fig. 75, pl. 20, figs. 7-8; pl. 21, fig. 1.
1987b Tetratrabs imlayi n. sp. – Yeh, p. 31, pl. 21, figs. 8, 10, 12.
2004 Tetraditryma macra (De Wever) – Matsuoka, fig. 41.
Original diagnosis: Hagiastrum with four long and thin
rays ornamented with long secondary spines.
Original description: Straight arms, composed of stout
longitudinal beams with regularly distributed connecting
bars that form quadrangular pores. This network exists only
on arms, while pores are randomly arranged in center.
Original remarks: This species is distinguished from other
species of Hagiastrum by its skeleton-like shape and welldeveloped secondary spines.
174
Arm structure is slightly different from other Hagiastrinae. In this subfamily the medullary shell is surrounded
by a cortical space, crossed by bars. For H. macrum, the
large number of bars and their distinctive shape seem
to form porous canals that fill the cortical space (pl. 1,
fig. 9). This is in agreement with P. O. Baumgartner’s observations (1981, pers. comm.) who noticed that the cortical
space is more and more restricted in early forms.
Further remarks: Although not mentioned in the original description, the view along one broken ray (De Wever
1981b, pl. 1. fig. 9) shows 3 primary canals surrounded by
9 secondary canals and 9 or more external beams; such
structure is similar to that of Homoeoparonaella elegans
(see Baumgartner, 1980).
Measurements (µm):
Based on 7 specimens.
HT Min. Max. Mean
Total length
442 430 496 458
(of two rays without terminal spines)
40 40
54
44
Width of rays
Length of terminal spines
70 62
70
66
Etymology: From Latin macer, -a, -crum, adj. = with little
substance, thin, stunted. By analogy with the emaciated
shape of this form.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Fernie Formation, NE British Columbia; Hyde Formation and Warm
Springs member of the Snowshoe Formation, Oregon;
Mino Terrane, Japan; Tawi Sadh Member of the Guwayza
Formation, Oman.
Plate HAG06. Hagiastrum macrum gr. De Wever. Magnification 150x. Fig. 1(H). TR, De Wever 1981b, pl. 1, fig. 7.
Fig. 2. TR, 1662D-R07-11. Fig. 3. NBC, GSC C-305208, GSC 128807. Fig. 4. OM, BR871-R04-04. Fig. 5. OM, BR525R08-10. Fig. 6. OM; BR1121-R07-18. Fig. 7. OM; BR706-R12-15. Fig. 8. OM-00-231, 020405.
175
Hagiastrum majusculum Whalen & Carter 1998
Species code: HAG03
Synonymy:
1987b Tetratrabs sp. E – Yeh, p. 32, pl. 11, fig. 12; pl. 22, fig. 2.
1988 Hagiastrum sp. cf. H. egregium Rüst – Carter et al., p. 29,
pl. 7, figs. 11, 12.
1988 Hagiastrum sp. A – Carter et al., p. 29, pl. 2, fig. 2.
1990 Hagiastrum sp. – Nagai, pl. 5, fig. 7.
1991 Hagiastrum sp. cf. H. egregium Rüst – Carter & Jakobs,
p. 342, pl. 2, fig. 10.
1996 Hagiastrum cf. H. egregium Rüst – Tumanda et al., p. 172,
fig. 6.6.
1996 Tetraditryma sp. A – Yeh & Cheng, p. 97, pl. 1, fig. 5.
1996 Hagiastrum sp. A – Yeh & Cheng, p. 96, pl. 1, fig. 12.
1997 Tetraditryma sp. F – Yao, pl. 7, fig. 331.
1998 Hagiastrum sp. A – Cordey, p. 67, pl. 19, figs. 5, 7, 9-10 only.
1998 Hagiastrum majusculum n. sp. – Whalen & Carter, p. 45, pl.
10, figs. 11-12, 14-16.
2002 Hagiastrum majusculum Whalen & Carter – Whalen &
Carter, p. 103, pl. 8, figs. 3, 11, 13.
2004 Hagiastrum rudimentum Whalen & Carter – Matsuoka,
fig. 38.
Original description: Test composed of four long, nodose
rays with bulbous tips terminating in moderately long
triradiate spines. Rays usually composed of eight external
nodose longitudinal beams; beams display strong linearity
frequently becoming slightly twisted. Beams connected by
transverse bars forming single, longitudinal rows of square
to tetragonal pore frames; pores circular to subcircular.
Nodes at vertices of pore frames elliptical to subrectangular, strongly raised and highly distinctive. Central area
small, composed of triangular and rectangular pore frames
with large nodes at vertices. Bulbous ray tips composed
of square pore frames with weak nodes at vertices. Spines
moderate in length, triradiate.
Original remarks: This form is likely derived from
Hagiastrum rudimentum n. sp. It differs from the latter
species in having longer rays with more strongly pronounced
linearity, stronger nodes on both rays and central area, and
in developing bulbous tips at the ends of the rays.
Further remarks: This species now includes forms with six
to eight external beams and a slightly larger, more nodose
central area.
Measurements (µm):
Based on 14 specimens.
Length of
longest
ray
249
395
205
292
Width of
widest
ray
43
52
37
43
Width of
bulbous
tips
99
74
85
Length of
longest
spine
131
75
92
HT
Max.
Min.
Mean
Etymology: Name from the Latin majusculus, um (m.)
meaning somewhat larger or greater.
Type locality: Sample 86-OF-KUB-2, Sandilands Formation, Kunga Island - north side, Queen Charlotte Islands,
British Columbia.
Occurrence: Sandilands, Ghost Creek, Fannin and Phantom
Creek formations, Queen Charlotte Islands; Bridge River
Complex and Williston Lake, British Columbia; San
Hipólito Formation, Baja California Sur; Hyde Formation,
Oregon; Japan; Liminangcong Chert, Philippines; Skrile
Formation, Slovenia; Tawi Sadh Member of the Guwayza
Formation, Oman.
Plate HAG03. Hagiastrum majusculum Whalen & Carter. Magnification Figs. 1-5 x100 (scale bar A), Figs. 6-10 x150
(scale bar B), except Fig. 8c x300. Fig. 1(H). Carter et al. 1998, pl. 10, fig. 11. Fig. 2. QCI, GSC loc. C-304567, GSC
128806. Fig. 3. QCI, GSC loc. C-304567, GSC 128804. Fig. 4. QCI, GSC loc. C-175306, GSC 128805. Fig. 5. Whalen &
Carter 2002, pl. 8, fig. 3. Fig. 6. OR600A-R01-03. Fig. 7. OR600A-R01-07. Fig. 8a,b,c. OM, BR682-R10-10.
Fig. 9. JP, MNA-10, MA10864. Fig. 10. SI, MM5.00, 010102.
176
177
Hagiastrum rudimentum Whalen & Carter 1998
Species code: HAG04
Synonymy:
1987 Hagiastrum sp. C – Hattori, pl. 3, fig. 14.
1998 Hagiastrum sp. A – Cordey, p. 67, pl. 19, figs. 6, 8 only.
1998 Hagiastrum rudimentum n. sp. – Whalen & Carter, p. 46,
pl. 10, figs. 2, 7, 8, 18, 19.
2002 Hagiastrum rudimentum Whalen & Carter – Suzuki et al.,
p. 178, figs. 7 H-I.
2002 Hagiastrum rudimentum Whalen & Carter – Whalen &
Carter, p. 103, pl. 8, fig. 2.
Original description: Test with four long rays, almost
cylindrical in cross-section, sometimes broadening slightly
near tips. Rays terminating in long triradiate spines having
broad ridges and grooves. Rays usually composed of eight
external longitudinal beams separated by a single row of
pore frames and connected by transverse bars to from
a single linear row of pores between two beams. Nodes
at vertices of pore frames round and moderately raised.
Central area large, composed mostly of triangular and
tetragonal pore frames with strongly raised circular nodes
at vertices.
Original remarks: This is the earliest species of Hagiastrum
to appear in our samples. It has a few irregularly arranged
pores on the rays but in all other aspects seems to conform
to the definition of Hagiastrum as emended by Baumgartner
(1980). See. H. majusculum n. sp. for comparisons.
Measurements (µm):
Based on 10 specimens.
Length of
longest ray
181
244
161
206
Width of
widest ray
38
56
38
46
Length of
longest spine
43
146
43
62
HT
Max.
Min.
Mean
Etymology: Name from the Latin rudimentum meaning
first principle, beginning.
Type locality: Sample QC-675, Sandilands Formation,
Kunga Island - north side, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands; Bridge River Complex, British
Columbia; San Hipólito Formation, Baja California Sur;
Pucara Group, Peru; Japan.
Genus: Helvetocapsa O’Dogherty, Goričan & Dumitrica 2006
Type species: Tricolocapsa matsuokai Sashida, in Sashida et al. 1999
Synonymy:
2006 Helvetocapsa O’Dogherty, Goričan & Dumitrica n. gen.
– O’Dogherty et al., p. 450.
Original diagnosis: Test spindle shaped, multisegmented,
composed of three, four, or possibly more segments. Last
segment inversely conical with a small constricted aperture at its base. Segmental divisions generally not well pronounced externally, only faint strictures present in some
species. Small circular pores on surface arranged in longitudinal rows. Numerous longitudinal plicae are generally
well developed on the entire test. One row of pores present
between neighbouring plicae.
Original remarks: The genus is similar to Striatojaponocapsa Kozur, from which it differs by having an inversely
conical last segment with a simple aperture but no appendage or eccentric porous depression. It differs from
Protunuma Ichikawa and Yao by having only one row of
pores between adjacent plicae.
A relatively continuous stratigraphic record can be
reconstructed for Helvetocapsa n. gen. from previously
178
known occurrences. The genus ranges from the early
Toarcian (Matsuoka 1991a, Goričan et al. 2003), through
Aalenian and Bajocian (Yao 1997; Sashida et al. 1999) to
the Bathonian (Yamamoto et al. 1985; this study). Species
of this genus have previously been assigned to different
genera, depending on the number of segments. We
grouped them together because the available data suggest
that the number of pores (one vs. two or more) between
adjacent plicae is an important taxonomic character but
the number of segments varies through time in the same
phylogenetic lineage.
Etymology: Referring to the occurrence in Switzerland.
Included species and subspecies:
TPS03 Helvetocapsa minoensis (Matsuoka) 1991
SCP03 Helvetocapsa nanjoensis (Matsuoka) 1991
SCP06 Helvetocapsa plicata s.l. (Matsuoka) 1991
SCP04 Helvetocapsa plicata plicata (Matsuoka) 1991
SCP05 Helvetocapsa plicata semiplicata (Matsuoka) 1991
Plate HAG04. Hagiastrum rudimentum Whalen & Carter. Magnification x150. Fig. 1(H). Carter et al. 1998, pl. 10, fig. 2.
Fig. 2. QCI, GSC loc. C-080612, GSC 128808.
179
Helvetocapsa minoensis (Matsuoka) 1991
Species code: TPS03
Synonymy:
1987 Tricolocapsa sp. A – Hattori, pl. 13, fig. 1.
1987 Tricolocapsa sp. B – Hattori, pl. 13, fig. 2.
1989 Tricolocapsa sp. A – Hattori, pl. 10, fig. A.
1989 Tricolocapsa spp. – Hattori, pl. 10, fig. D.
1990 Tricolocapsa sp. – Nagai, pl. 4, figs. 5a-b.
1991 Tricolocapsa minoensis n. sp. – Matsuoka, p. 723,
Fig. 2. 1a-5b.
? 1991 Tricolocapsa sp. cf. T. plicarum – Kojima et al., pl. 1, fig. 10.
1997 Tricolocapsa minoensis Matsuoka – Yao, pl. 9, fig. 422.
2003 Tricolocapsa minoensis Matsuoka – Goričan et al., p. 297,
pl. 4, figs. 13a-b, 14a-b.
2004 Tricolocapsa minoensis Matsuoka – Matsuoka, fig. 81.
Original description: Shell of three segments, drop-like
shaped. Cephalis hemispherical, poreless. Thorax truncate
conical. Abdomen large, subspherical with constricted aperture. Collar and lumber strictures slightly recognizable
or indistinct externally. Outer surface of shell ornamented
with continuous longitudinal plicae. Eleven to 15 moderately spaced plicae visible on outer shell. One row of pores
present between longitudinal plicae. Pores small to moderate in size, circular and uniform in shape. Aperture moderate in size, circular.
Original remarks: Pores vary in size among specimens,
from small (Figures 2-1a,b) to moderate (Figure 2-3).
Tricolocapsa minoensis, n. sp., differs from T. plicarum
Yao by lacking a dish-like basal appendage, by having more
spaced plicae and by its smaller size.
Measurements (µm):
Numbers of specimens measured are in parentheses.
Total height of shell
Maximum width of shell
Diameter of aperture
HT
115
94
13
Max.
115
94
14
Min. Mean
93
105
72
83
12
13
(15)
(15)
(6)
Etymology: This species is named for the Mino Terrane
which includes the type area, Nanjo Massif.
Type locality: Sample MNA-10, Nanjo Massif, Mino Terrane, central Japan.
Occurrence: Japan; Hyde Formation, Oregon; Skrile Formation, Slovenia; Tawi Sadh Member of the Guwayza Formation, Oman.
Helvetocapsa nanjoensis (Matsuoka) 1991
Species code: SCP03
Synonymy:
1991 Stichocapsa nanjoensis n. sp. – Matsuoka, p. 733,
Fig. 9. 1 – 4b.
1997 Stichocapsa nanjoensis Matsuoka – Yao, pl. 9, fig. 440.
2004 Stichocapsa nanjoensis Matsuoka – Matsuoka, fig. 88.
Original description: Shell of four segments, spindleshaped. Cephalis hemispherical, poreless. Thorax truncate
conical, abdomen barrel-shaped and fourth segment
inverted conical with a constricted, circular aperture.
Collar stricture pronounced externally. Other segmental
joints indistinct externally. Eight to 10 weakly developed
longitudinal plicae run from thorax to the distal end
in lateral view. One row of pores arranged between the
longitudinal plicae. Pores circular to subcircular, small and
uniform in size.
Original remarks: Stichocapsa nanjoensis, n. sp. is distinguished from S. biconica, n. sp. by its small size, by consisting of four segments rather than five and by having longi-
180
tudinal plicae. It also differs from Cyrtocapsa (?) kisoensis
Yao by having longitudinal plicae and by lacking an apical
horn.
Measurements (µm):
Numbers of specimens measured are in parentheses.
HT
Total height of shell
104
Maximum width of shell 62
Diameter of aperture
-
Max.
104
63
8
Min. Mean
90
96
(10)
54
59
(10)
7
8
(5)
Etymology: The species is named for the Nanjo Massif, its
type locality.
Type locality: MNA-10, Nanjo Massif, Mino Terrane,
central Japan.
Occurrence: Mino Terrane, Japan; Tawi Sadh Member of
the Guwayza Formation and Musallah Formation, Oman.
Plate TPS03. Helvetocapsa minoensis (Matsuoka). Magnification x400. Fig. 1(H). Matsuoka 1991, Fig. 2.1a.
Fig. 2. OM, BR1122-R02-04. Fig. 3. OM, BR1122-R04-16. Figs. 4a-b, 5a-b. Goričan et al. 2003, pl. 4, figs. 14a-b, 13a-b.
Plate SCP03. Helvetocapsa nanjoensis (Matsuoka). Magnification x500. Fig. 1(H). Matsuoka 1991, Fig. 9.1.
Fig. 2. OM, BR1122-R04-15. Figs. 3a, b. OM-00-254-022212, 022213.
181
Helvetocapsa plicata s.l. (Matsuoka) 1991
Species code: SCP06
Synonymy:
1991 Stichocapsa plicata n. sp. – Matsuoka, p. 727, Fig. 5-1a-5b:
6-1a-6b.
See also subspecies.
Included subspecies:
SCP04 Helvetocapsa plicata plicata (Matsuoka) 1991
SCP05 Helvetocapsa plicata semiplicata (Matsuoka) 1991
Original description: Shell of four to six segments, droplike or spindle shaped. Cephalis hemispherical, poreless.
The last segment large, inverted hemispherical or inverted
conical with a constricted aperture. Remaining segments
including thorax and abdomen truncate conical. Strictures between segments indistinct externally. Fifteen to 22
densely spaced longitudinal plicae visible on outer shell except for cephalis. The longitudinal plicae distinct or partly obscure. One row of pores present between the plicae.
Pores uniform, small and circular. Aperture small, circular,
occasionally with a short protruding rim.
Original remarks: Two subspecies are included under
this species; these are S. plicata plicata, n. subsp. and
S. plicata semiplicata, n. subsp. S. plicata, n. sp. differs from
S. convexa Yao by having longitudinal plicae between which
one row of pores is present.
Measurements (µm):
Numbers of specimens measured are in parentheses.
HT
Total height of shell
154
Maximum width of shell 108
Diameter of aperture
8
Max.
170
124
8
Min.
110
80
6
Mean
142
96
7
(29)
(29)
(12)
Etymology: This specific name comes from the Latin
plicatus-a-um (= plicate).
Type locality: MNA-10, Nanjo Massif, Mino Terrane, central Japan.
Occurrence: See subspecies.
Helvetocapsa plicata plicata (Matsuoka) 1991
Species code: SCP04
Synonymy:
1989 Tricolocapsa sp. B. – Hattori, pl. 10, fig. B.
1989 Tricolocapsa spp. – Hattori, pl. 10, fig. F.
1989 Tricolocapsa sp. D. – Hattori, pl. 29, fig. D.
1989 Tricolocapsa sp. – Hattori & Sakamoto, pl. 19, fig. B.
1991 Stichocapsa plicata plicata n. subsp. – Matsuoka, p. 729,
fig. 5. 1a – 5b.
1997 Stichocapsa plicata Matsuoka – Yao, pl. 9, fig. 437.
2003 Stichocapsa plicata plicata Matsuoka – Goričan et al.,
p. 297, pl. 4, fig. 17a-b.
2004 Stichocapsa plicata plicata Matsuoka – Matsuoka, fig. 84.
n. subsp. differs from S. plicata semiplicata, n. subsp. by
having distinct longitudinal plicae that extend from the
thorax to the distal end.
Further remarks: In comparison to the type material, some
specimens have less numerous plicae and much larger pores
with thicker pore frames (pl. SCP04, figs. 6-9). They are included in Helvetocapsa plicata plicata, because transitional
forms (pl. SCP04, figs. 5a-b) also exist.
Measurements (µm):
Numbers of specimens measured are in parentheses.
Original description: Shell of four to five segments, droplike or spindle shaped. Cephalis hemispherical, poreless.
The last segment large, inverted hemispherical or inverted
conical with a constricted aperture. Remaining segments
including thorax and abdomen truncate conical. Strictures
between segments indistinct externally. Fifteen to 22
densely spaced longitudinal plicae visible on outer shell.
The plicae distinct on whole shell except for cephalis. One
row of pores present between plicae. Pores uniform, small
and circular. Aperture small, circular, occasionally with
a short protruding rim.
Type locality: MNA-10, Nanjo Massif, Mino Terrane,
central Japan.
Original remarks: Shape varies among specimens; some
are drop-like shaped (Figure 5-1a, 3a, 4, 5a, b) and others
spindle shaped (Figure 5-2a). Stichocapsa plicata plicata,
Occurrence: Japan; Skrile Formation, Slovenia; Musallah
Formation and Tawi Sadh Member of the Guwayza
Formation, Oman.
182
HT
Total height of shell
154
Maximum width of shell 108
Diameter of aperture
8
Max.
170
124
8
Min.
110
82
6
Mean
142
102
7
(20)
(20)
(9)
Etymology: This subspecies is the nominotypical subspecies of Stichocapsa plicata, n. sp.
Plate SCP04. Helvetocapsa plicata plicata (Matsuoka). Magnification x400. Fig. 1(H) a, b. Matsuoka 1991,
Figs. 5.1a-b. Figs. 2, 3. Matsuoka 1991, Fig. 5.3a, 4. Fig. 4. Goričan et al. 2003, pl. 4, fig. 17a. Figs. 5a, b. OM-99-89011416, 011415. Fig. 6. OM-00-263-021330. Figs. 7a, b. OM-00-252-022010, 022011. Fig. 8. OM-00-251-021609.
Fig. 9. OM-00-115-023022.
183
Helvetocapsa plicata semiplicata (Matsuoka) 1991
Species code: SCP05
Synonymy:
1991 Stichocapsa plicata semiplicata n. subsp. – Matsuoka,
p. 729, Fig. 6. 1a – 6b.
1997 Stichocapsa semiplicata Matsuoka – Yao, pl. 9, fig. 438.
2004 Stichocapsa plicata semiplicata Matsuoka – Matsuoka, fig. 85.
2005 Stichocapsa plicata semiplicata Matsuoka – Kashiwagi et al.,
pl. 5, fig. 8.
Original description: Shell of five to six segments, drop-like
shaped. Cephalis hemispherical, poreless. The last segment
large, truncate subspherical with a constricted aperture.
Remaining segments including thorax and abdomen
truncate conical. Strictures between segments indistinct
externally. Fifteen to 22 densely spaced longitudinal plicae
visible on outer shell except for cephalis and the middle part
of shell. Pores small, circular and arranged longitudinally.
One row of pores present between the plicae. Aperture
small, circular, occasionally with a short protruding rim.
184
Original remarks: Stichocapsa plicata semiplicata, n. subsp.
is compared to S. plicata plicata, n. subsp. under the latter
subspecies.
Measurements (µm):
Numbers of specimens measured are in parentheses.
Total height of shell
Maximum width of shell
Diameter of aperure
HT
139
95
7
Max.
149
99
8
Min.
123
80
7
Mean
142
94
7
(9)
(9)
(3)
Etymology: The subspecific name comes from the Latin
semi (=half) and plicatus-a-um (=plicate).
Type locality: Sample MNA-10, Nanjo Massif, Mino Terrane, central Japan.
Occurrence: Japan; Tawi Sadh Member of the Guwayza
Formation, Oman.
Plate SCP05. Helvetocapsa plicata semiplicata (Matsuoka). Magnification x400. Fig. 1(H)a, b. Matsuoka 1991,
Fig. 6.1a-b. Figs. 2, 3. Matsuoka 1991, Figs. 6.3, 6.5. Fig. 4. OM, BR524-R05-16.
185
Genus: Hexasaturnalis Kozur & Mostler 1983
Type species: Spongosaturnalis? hexagonus Yao 1972
Synonymy:
1983 Hexasaturnalis n. gen. – Kozur & Mostler, p. 28.
1983 Yaosaturnalis n. gen. – Kozur & Mostler, p. 31.
Original description: Ring and outer spines strongly bladed. Outline of ring hexagonal to octagonal or subquadratically rounded. 4-8 very strong outer spines. Two massive
polar spines opposite to interspine spaces on the outer
margin of the ring. No auxiliary spines. Ring often a little
constricted in the polar spine attachment region. Cortical
shells spongy, widely separated from the inner margin of
the ring. Medullary shell latticed.
Original remarks: By increase of the number of marginal
spines the hexagonal to octagonal outline of the ring is
transformed to a polygonal to subcircular one. In this manner the genus Spongosaturnalis Campbell and Clark, 1944b,
evolved in the Cretaceous from Hexasaturnalis n. gen.
Praehexasaturnalis n. gen. from the Norian has the same
outline of ring, but the polar spines are still situated opposite to the marginal spines and the narrow ring is still flat
to shallow oval in cross section. Yaosaturnalis n. gen. has
the same outline and structure of ring as Hexasaturnalis
n. gen., but auxiliary spines are present.
Further remarks: Herein we follow De Wever et al. (2001)
who synonymized Yaosaturnalis Kozur & Mostler with
Hexasaturnalis Kozur & Mostler.
Etymology: According to the outline.
Included species:
3502 Hexasaturnalis hexagonus (Yao) 1972
SAT11 Hexasaturnalis octopus Dumitrica & Hori n. sp.
3089 Hexasaturnalis tetraspinus (Yao) 1972
Hexasaturnalis hexagonus (Yao) 1972
Species code: 3502
Synonymy:
1972 Spongosaturnalis ? hexagonus n. sp. – Yao, p. 31, pl. 6,
figs. 1-3; pl. 11, figs. 3a-c.
1982 Spongosaturnalis ? hexagonus Yao – Wakita & Okamura,
pl. 5, fig. 2.
1982 Spongosaturnalis ? hexagonus Yao – Matsuda & Isozaki,
pl. 1, fig. 20.
1982 Spongosaturnalis ? hexagonus Yao – Wakita, pl. 4, fig. 11.
1982 Spongosaturnalis (?) tetraspinus Yao – Kishida & Sugano,
pl. 6, fig. 9, not 10.
1982 Acanthocircus hexagonus (Yao) – Kido, pl. 3, fig. 10.
Not 1986 Hexasaturnalis hexagonus (Yao) – Grill & Kozur, pl. 2,
fig. 5.
1987 Acanthocircus hexagonus (Yao) – Hattori, pl. 1, fig. 2.
not 1988 Mesosaturnalis hexagonus (Yao) – Carter et al., pl. 47,
pl. 9, figs. 11-12.
1988 Acanthocircus hexagonus (Yao) – Hattori, pl. 1, fig. K.
1989 Acanthocircus hexagonus (Yao) – Hattori & Sakamoto,
pl. 2, fig. B, not fig. C.
1989 Acanthocircus hexagonus (Yao) – Hattori, pl. 18, fig. B,
pl. 35, fig. F.
1989 Mesosaturnalis sp. cf. M. hexagonus (Yao) – Hori & Otsuka,
pl. 4, fig. 9.
1990 Mesosaturnalis hexagonus (Yao) – Hori, Fig. 9.42.
1991 Mesosaturnalis hexagonus (Yao) – Carter & Jakobs, p. 343,
pl. 2, fig. 15.
1993 Acanthocircus hexagonus Yao – Fujii et al., pl. 1, fig. 3.
1995a Hexasaturnalis hexagonus (Yao) – Baumgartner et al.,
p. 252, pl. 3502, figs. 1-3.
1996 Mesosaturnalis hexagonus (Yao) – Tumanda et al., p. 173,
fig. 5.19.
1996 Acanthocircus hexagonus (Yao) – Yeh & Cheng, p. 106,
pl. 2, fig. 9; pl. 7; fig. 4.
1997 Mesosaturnalis hexagonus Yao – Hori, pl. 1, figs. 10a-c.
186
1997 Hexasaturnalis hexagonus (Yao) – Yao, pl. 5, fig. 218.
? 1997 Hexasaturnalis aff. hexagonus (Yao) – Yao, pl. 5, fig. 222.
2002 Hexasaturnalis hexagonus (Yao) – Hori & Wakita, pl. 3,
fig. 16.
2004 Hexasaturnalis hexagonus (Yao) – Hori, pl. 4, fig. 38; pl. 6,
fig. 2; pl. 9, fig. 6; pl. 13, fig. 59; pl. 23, fig. 21.
2005 Hexasaturnalis hexagonus (Yao) – Hori, pl. 12, fig. 21;
pl. 13, fig. 36.
Original description: Spongosaturnalid with subhexagonal ring, and with six strong spines on ring. Shell not
preserved, but believed to be wholly spongy because numerous fragmentary thorns, which may be connected with
spongy shell, are clearly observed on sturdy spines. Polar
spines short, thick, with no ridge. Ring bilaterally symmetrical, subhexagonal, strong, with clear ridge on outer
edge. Inner edge of ring curves rather smoothly, while
outer edge is subhexagonal, with spine at each vertex. Ring
which joins with polar spine bends slightly toward inside.
Spines, situated diagonally on ring, strong, somewhat long,
of sharp tip, with clear ridges which continue to one on
outer edge of ring.
Original remarks: This species differs from Spongosaturnalis ? septispinus in the number of the spine, and from
S. ? minoensis in lacking auxiliary spines on the inner margin of the saturnalin ring. Spongosaturnalis ? sp., reported
by Foreman (1971, pl.1, fig.4; Cretaceous sediments core,
Site 61, west margin of East Mariana Basin, through the
Deep Sea Drilling Project), is similar to this species, but
the former has slender spines on which there is no ridge.
Measurements (µm):
Based on 6 specimens.
Diameter of ring longitud.
Diameter of ring trans.
Diameter of shell
Lenght of polar spine
Lenght of spine
Breadth of ring
HT
243
278
136
23
126
36
Av.
198
245
104
19
101
27
Min.
156
188
75
10
62
17
Max.
243
282
136
25
130
39
Type locality: Manganese carbonate ore, Mino Belt, river
side of the Kiso, east of Unuma, Kagamihara City, Gifu
Prefecture, Central Japan.
Occurrence: Worldwide.
Plate 3502. Hexasaturnalis hexagonus (Yao). Magnification x150. Fig. 1(H). Yao 1972, pl. 6, fig. 2. Fig. 2. Hori 1990,
fig. 9-42.
187
Hexasaturnalis octopus Dumitrica & Hori n. sp.
Species code: SAT11
Synonymy:
1972 Spongosaturnalis ? sp. a – Yao, pl. 8, figs. 6, 7.
1997 Hexasaturnalis sp. A – Yao, pl. 5, fig. 221.
Type designation: Holotype pl. SAT11, fig. 1, sample BR871,
chert of Tawi Sadh Member reworked in the Guwayza
Formation, Al Khashbah Mountains, Oman.
Diagnosis: Hexasaturnalis with eight-spined ring.
Description: Shell spongy, ellipsoidal when preserved. Ring
symmetrical, subcircular or elliptical, rounded on the inner
margin, octagonal on outer margin, with a spine at each
corner. Inner margin without auxiliary spines. Middle part
of ring more or less constricted in the vicinity of contact
with polar rays. Outer blades of ring well developed
especially at proximal part of ring. Spines usually strong,
four-bladed, pointed; peripolar spines longer than distal
spines, the latter rarely extremely short. Commonly distal
spines closer to each other than to peripolar spines.
188
Remarks: This new species differs from the other species of
the genus Hexasaturnalis in the number of spines.
Measurements (µm):
Based on 6 specimens.
Longitudinal size of ring
Transversal size of ring
Length of shell
Length of polar rays
Length of peripolar spines
Length of distal spines
Breadth of ring bar
HT
213
220
90
67+
47-60
27-33
20
Av.
195
236
56
56
28
21
Min.
180
220
47
40
20
17
Max.
213
267
73
67
47
27
Etymology: From the Greek octo - eight and pous, podos
– foot, referring to the number of spines; noun.
Type locality: Sample BR871, Tawi Sadh Member, Guwayza
Formation, Al Khasbah Mountains, Oman.
Occurrence: Tawi Sadh Member of the Guwayza Formation,
Oman; Japan; Snowshoe Formation, Oregon.
Plate SAT11. Hexasaturnalis octopus Dumitrica & Hori n. sp. Magnification x150. Fig. 1(H). OM, BR871-R01-10.
Fig. 2. OM, BR871-R01-07. Fig. 3. JP, IYII10-143. Fig. 4. JP, IYII14-41. Fig. 5. JP, IYII14-65. Fig. 6. JP, IYII14-44.
Fig. 7. JP, IYII14-43. Fig. 8. JP, IYII10-142. Fig. 9. JP, IYII10-11. Fig. 10. JP, IYII-144. Fig. 11a,b. OR555-R07-13.
189
Hexasaturnalis tetraspinus (Yao) 1972
Species code: 3089
Synonymy:
1972 Spongosaturnalis ? tetraspinus n. sp. – Yao, p. 29, pl. 4,
figs. 1-6; pl. 11, figs. 1-2.
1982 Spongosaturnalis ? tetraspinus Yao – Wakita, pl. 4, fig. 12.
not 1982 Spongosaturnalis ? tetraspinus Yao – Kishida &
Sugano, pl. 6, figs. 9-10.
? 1984 Mesosaturnalis squinaboli – Carayon et al., pl. 1, fig. 2.
? 1986 Hexasaturnalis hexagonus (Yao) – Grill & Kozur, pl. 2,
fig. 5.
1987 Mesosaturnalis tetraspinus (Yao) – Goričan, p. 184, pl. 3,
fig. 1.
1988 Acanthocircus tetraspinus Yao – Hattori, pl. 2, fig. B.
? 1988 Mesosaturnalis hexagonus (Yao) – Carter et al., pl. 47,
pl. 9, figs. 11-12.
1989 Acanthocircus tetraspinus Yao – Hattori, pl. 35, fig. G.
1990 Mesosaturnalis tetraspinus (Yao) – Yao, pl. 3, fig. 24.
1991 Mesosaturnalis tetraspinus (Yao) – Carter & Jakobs, p. 343,
pl. 2, fig. 16.
1991 Hexasaturnalis tetraspinus (Yao) – Tonielli, p. 23, pl. 1,
fig. 5.
1995a Hexasaturnalis tetraspinus (Yao) – Baumgartner et al.,
p. 254, pl. 3089, figs. 1-3.
1996 Acanthocircus tetraspinus (Yao) – Yeh & Cheng, p. 106,
pl. 2, fig. 10.
1997 Hexasaturnalis tetraspinus (Yao) – Yao, pl. 5, fig. 219.
? 1997 Hexasaturnalis aff. tetraspinus (Yao) – Yao, pl. 5, fig. 220.
2004 Hexasaturnalis tetraspinus (Yao) – Hori, pl. 4, fig. 39,
pl. 11, fig. 43, pl. 13, fig. 60.
2005 Hexasaturnalis tetraspinus (Yao) – Hori, pl. 12, fig. 22.
Original description: Spongosaturnalid with four strong
spines on proximal part of ring. Shell not preserved, but
judged from numerous fragmentary thorns attached to tip
of polar spines and on sturdy spines, it is most probably
190
spongy. Polar spines extend and bifurcate to form a subcircular ring with distinct indentation proximally. Ring bilaterally symmetrical, strong, with clear ridge on outer edge.
Four spines are present symmetrically on proximal part of
ring. Spines strong, slightly curved, with sharp tip, and with
clear ridges. In some specimens, short spine is present on
terminal end of ring.
Original remarks: This species is distinguished from other
species by the strong spines on the proximal part of the ring.
There is little variation in the shape of the ring, excluding
the presence of a short spine at the terminal end.
Complete specimen with the shell was not found and
fragmentary rings are common. Although the generic
assignment of this species is slightly doubtful, it may belong
to the genus Spongosaturnalis because of its morphological
feature.
Measurements (µm):
Based on 6 specimens.
Diameter of ring longit.
Diameter of ring; trans.
Diameter of shell
Length of polar spine
Length of spine
Breadth of ring
HT
360
485
105
95
90
37
Av.
294
374
102
63
75
30
Min.
220
220
80
40
58
16
Type locality: Inuyama area, central Japan.
Occurrence: Worldwide.
Max.
360
485
120
95
100
37
Plate 3089. Hexasaturnalis tetraspinus (Yao). Fig. 1(H). Magnification x100. Fig. 1(H). Yao 1972, pl. 4, fig. 6.
Fig. 2. JP, IYII-14. Fig. 3. JP, IYII5-65. Fig. 4. OR555-R07-15. Fig. 5. OR555-R07-14.
191
Genus: Higumastra Baumgartner 1980
Type species: Higumastra inflata Baumgartner 1980
Synonymy:
1980 Higumastra n. gen. – Baumgartner, p. 290.
Original description: Test composed of 4 rays at right
angles. Cortical rays composed of thin external beams
connected by regular bars forming large circular pores in
longitudinal rows. Ray tips with central or 2 lateral and
central spines. Inner structure in rays and medullary shells
always visible in transmitted light observation. Centrally
placed shells (1 or 2) are on both sides joined to the cortical
shell. Vertical septae lying below the median pore row
extend from the innermost medullary shell and divide the
inner space of the rays into 2 main canals of semicircular
cross section. Vertical septum composed of primary beam
and primary lamellae penetrated by large lamellar pores.
Vertical septum with 1 or 2 channels below the median
pore row on each side. Patagium may be well developed,
present as remnants, or absent.
Original remarks: Higumastra n. gen. differs from all other
four-armed hagiastrids by the easily visible inner structure
and in having large pore frames in longitudinal rows with a
distinct median pore row.
Etymology: Higumastra is an anagram of Hagiastrum.
Included species:
HIG01 Higumastra laxa Yeh 1987b
HIG04 Higumastra lupheri Yeh 1987b
HIG03 Higumastra transversa Blome 1984b
Higumastra laxa Yeh 1987b
Species code: HIG01
Synonymy:
1987b Higumastra laxa n. sp. – Yeh, p. 25, pl. 8, figs. 13, 18;
pl. 29, fig. 20.
1987b Higumastra splendida n. sp. – Yeh, p. 26, pl. 8, figs. 2, 4,
11-12, 16, 18, 25-27.
1987b Higumastra sp. aff. H. splendida n. sp. – Yeh, p. 27, pl. 8,
figs. 5, 28.
1987b Higumastra sp. A – Yeh, p. 27, pl. 8, figs. 3, 17.
1997 Higumastra laxa Yeh – Yao, pl. 7, fig. 317.
? 2003 Higumastra laxa Yeh – Goričan et al., p. 293, pl. 1, fig. 18.
2004 Higumastra laxa Yeh – Matsuoka, fig. 48.
Original description: Test with large central area and
short, wide rays. Rays tapering distally with short, massive
triradiate spines. One spine slightly longer than other
three. Rays comprised of large tetragonal and pentagonal
pore frames with five pores visible laterally at proximal
end and three pores visible distally. Central area of cortical
shell consisting of concentrically arranged pentagonal and
hexagonal pore frames without prominent nodes at vertices.
Test with or without patagium.
Original remarks: Higumastra laxa, n. sp., differs from
H. splendida, n. sp., by lacking massive nodes at pore frame
vertices and by having wider rays with shorter primary
spines.
192
Remarks: In this work Higumastra laxa Yeh and Higumastra splendida Yeh are synonymized.
Measurements (µm):
Ten specimens measured.
Length
of ray
61
73
60
65
Width of ray
at base
61
73
58
65
Width of
central area
122
128
118
123
Length
of spine
61
61
30
45
HT
Max.
Min.
Mean
Etymology: Laxus-a-um (Latin, adj). = wide.
Type locality: OR-600M, Hyde Formation at Izee-Paulina
road, east-central Oregon.
Occurrence: Hyde Formation and Warm Springs member
of the Sowshoe Formation, east-central Oregon; Tawi Sadh
Member of the Guwayza Formation, Oman; Japan.
Plate HIG01. Higumastra laxa Yeh. Magnification x250. Fig. 1(H). Yeh 1987b, pl. 8, fig. 13. Figs. 2a, b. OR600A,
13151a,b. Fig. 3. OR600A, 13126. Fig. 4. OM, BR682-R09-21. Fig. 5. OM, BR1122-R02-19. Fig. 6. OM, BR524-R05-27.
193
Higumastra lupheri Yeh 1987b
Species code: HIG04
Synonymy:
1987b Higumastra lupheri n. sp. – Yeh, p. 25, pl. 8, figs. 8, 24.
1988 Higumastra sp. A – Carter et al., p. 29, pl. 10, fig. 6.
2004 Higumastra lupheri Yeh – Matsuoka, fig. 48.
Original description: Rays medium in length, wide proximally, slightly tapering distally with long massive triradiate spines, one spine slightly longer than other three. Test
comprised of large, nearly uniform size of linearly arranged
tetragonal pore frames on rays, small pentagonal and hexagonal concentrically arranged pore frames at central area.
Central area medium in size. All pore frames lacking prominent nodes at vertices. Test with or without patagium.
Original remarks: Higumastra lupheri, n. sp., differs from
H. oregonensis, n. sp., by having wider and shorter rays with
longer primary spines and by concentrically arranged pore
frames on the cortical shell of central area.
Measurements (µm):
Ten specimens measured.
Length
of ray
165
170
150
160
Width of
ray at base
110
116
105
108
Width of
central area
220
225
215
220
Length
of spine
110
118
105
110
HT
Max.
Min.
Mean
Etymology: This species is named for Dr. R. L. Lupher in
honor of his early contribution to the geology of east-central Oregon.
Type locality: Sample OR-600A, Hyde Formation at IzeePaulina road, east-central Oregon.
Occurrence: Hyde Formation and Warm Springs member
of the Sowshoe Formation, east-central Oregon; Phantom
Creek and Graham Island formations, Queen Charlotte
Islands; Mino Terrane, Japan; Tawi Sadh Member of the
Guwayza Formation, Oman.
Higumastra transversa Blome 1984b
Species code: HIG03
Synonymy:
1984b Higumastra transversa n. sp. – Blome, p. 350, pl. 1,
figs. 3-5, 8-13, 16-19; pl. 15, fig. 4.
1988 Crucella sp. A – Carter et al., p. 43, pl. 15, figs. 9, 12.
1991 Higumastra sp. cf. H. transversa Blome – Carter & Jakobs,
p. 342, pl. 2, fig. 3.
Original description: Test as with genus. Rays usually short,
of equal length; rays usually slightly twisted (nodose dorsal
surface of ray slightly offset with respect to ventral surface);
rays inflated, subrectangular in axial section, usually thickest medially; rays terminating in relatively long central
spines, circular in axial section. Meshwork of cortical shell
(both rays and central area) consisting of two visible layers of pentagonal and hexagonal pore frames: nodose outer
layer composed of three to four rows of nodose pore frames
visible on both dorsal and ventral surfaces, pore frames
possessing massive, subspherical nodes at the pore frame
vertices; outer layer extending linearly from distal end of
rays onto middle of central area; inner layer visible on lateral portions of central area, pore frames more regular in
size, remnants of subsidiary beams (pillars which connect
inner layer to outer layer) sometimes observable at pore
frame vertices (see pl. 1, fig. 12). Some specimens exhibit
remnants of patagium on lateral surfaces between rays (see
pl. 1, fig. 16). Medullary shell as with genus.
Original remarks: Higumastra transversa differs from
H. inflata by having slightly twisting, subrectangular rays, a
194
cortical shell with massive nodes at the pore frame vertices,
and longer central spines on the ray tips.
Further remarks: Forms with slightly tapering rays disposed in an X-shaped pattern (i.e. not at 90˚) are also included.
Measurements (µm):
Based on 7 specimens.
Ray
length
100, 105,
106, 112
120
105
115
Ray width
72, 75, 78,
79
87
60
74
Cortical
shell width
147
168
133
147
Spine
length
87, 90,
112, 118
114
65
96
HT
Max.
Min.
Av.
Etymology: Transversus-a-um (latin, adj., f.) = transverse,
oblique, in an oblique direction.
Type locality: Sample 80AJM 8A, Shelikof Formation,
Puale Bay, southern Alaska.
Occurrence: Shelikof Formation, Alaska; Lonesome Formation, Oregon; Phantom Creek and Graham Island formations, Queen Charlotte Islands.
Plate HIG04. Higumastra lupheri Yeh. Magnification x250. Fig. 1(H). Yeh 1987b, pl. 8, fig. 8. Fig. 2. OR600A, 13162.
Fig. 3. OM, BR1121, 15921. Fig. 4. OM, BR706, 15780. Fig. 5. OM, BR1121, 15931. Fig. 6. JP, MNA-10, MA10707.
Plate HIG03. Higumastra transversa Blome. Magnification x150. Fig. 1(H). Blome 1984b, pl. 3, fig. 3.
Fig. 2. Carter et al. 1988, pl. 15, fig. 9. Fig. 3. Carter & Jakobs 1991, pl. 2, fig. 3.
195
Genus: Homoeoparonaella Baumgartner 1980
Type species: Paronaella elegans Pessagno 1977a
Synonymy:
1980 Homoeoparonaella n. gen – Baumgartner, p. 288.
Original description: Test as with subfamily, composed of
3 rays with equal to subequal interradial angles lacking a
bracchiopyle and a patagium. Cortical rays composed of
numerous longitudinal external beams connected by short
bars in transverse rows forming small pore frames. Nodes
well developed. Ray tips bulbous with or without central
spines. Medullary shell composed of centrally placed
medullary rays merging in central area. Medullary rays
composed of 3 (sometimes 5) primary canals arranged
around primary beams. Medullary shell connected by
numerous radially arranged subsidiary beams to cortical
shell.
Original remarks: Homoeoparonaella, n. gen. differs from
Paronaella Pessagno, 1971 (placed in Patulibracchiidae
herein) by its regular linear arrangement of pores and external beams and by its differentiation into cortical and medullary shells. It is distinguished from all other three-armed
hagiastrid genera in having numerous external beams and
in lacking a bracchiopyle.
Etymology: Homoeoparonaella is named for its external
homeomorphy with Paronaella Pessagno.
Included species:
HOM01 Homoeoparonaella lowryensis Whalen & Carter
2002
HOM02 Homoeoparonaella reciproca Carter 1988
Homoeoparonaella lowryensis Whalen & Carter 2002
Species code: HOM01
Synonymy:
1998 Homoeoparonaella sp. A – Whalen & Carter, p. 46, pl. 13,
fig. 13, 17.
2002 Homoeoparonaella lowryensis n. sp. – Whalen & Carter,
p. 104, pl. 3, figs. 5, 6, 13, 15.
Original description: Elongate slender rays, sub-circular
in axial section, all approximately same length. Expanded
ray tips with slightly planiform top and bottom surfaces.
Meshwork on rays composed of irregularly shaped triangular and tetragonal pore frames with distinct nodes at pore
frame vertices; alignment of pore frames parallel to long
axis of each ray. Meshwork on expanded ray tips composed
of irregularly shaped and distributed rectangular to circular
pore frames with slight development of nodes at pore frame
vertices. Each ray with one small spine, usually broken.
Original remarks: Homoeoparonaella hydensis Yeh 1987
and H. reciproca Carter 1988 appear to be distinctly different
species from H. lowryensis n. sp. because their pore frames
are much more regular.
196
Measurements (µm):
Based on 10 specimens.
Length of ray
225
225
173
193
HT
Max.
Min.
Mean
Etymology: This species is named for Pico Lowry located to
the northeast of the type area.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur, Mexico.
Occurrence: San Hipólito Formation, Baja California Sur;
Sandilands Formation, Queen Charlotte Islands; Dürrnberg
Formation, Austria; Tawi Sadh Member of the Guwayza
Formation, Oman.
Plate HOM01. Homoeoparonaella lowryensis Whalen & Carter. Magnification x150. Fig. 1(H). Whalen & Carter 2002,
pl. 3, fig. 5. Fig. 2. Whalen & Carter 2002, pl. 3, fig. 6. Fig. 3. AT, BMW21-27. Fig. 4. OM, BR1121, 15892.
197
Homoeoparonaella reciproca Carter 1988
Species code: HOM02
Synonymy:
much smaller. The alternating pore frame pattern batween
beams is diagnostic of H. reciproca but observable on well
preserved specimens only.
Original diagnosis: Test has three rays of moderate (near
equal) length with strongly expanded ray tips terminated
by numerous short, fine spines. Pore frames and beams are
aligned longitudinally, producing a pattern of single rows
of square pore frames that alternate with double rows of
triangular pore frames.
Measurements (µm):
Based on 11 specimens.
1988 Homoeoparonaella reciproca Carter – Carter et al., p. 28,
pl. 7, figs. 2-3.
Original description: Three-rayed test. Rays of moderate
length, interradial angles subequal. Rays composed of 8-10
longitudinal beams with transverse bars oriented both perpendicular and oblique to the beams, forming single rows
of square pore frames that alternate with double rows of
triangular pore frames. Rays circular in axial section. Pore
frames on ray tips are irregularly distributed, polygonal in
shape.
Original remarks: Rays are short and stout compared
with those of Homoeoparonaella argolidensis Baumgartner.
Homoeoparonaella reciproca differs from H. hydensis Yeh
in having double rows of triangular pore frames alternating with single rows of square pore frames; H. hydensis
has only linearly arranged square pore frames and is also
198
Lengths of rays
AX
BX
CX
Width of rays
Width of ray tips
Length of longest spine
HT
185
202
202
46-49
122-133
23
Av.
191
Max.
210
Min.
150
56
128
29
62
140
43
46
95
22
Etymology: Latin, reciprocus (adj.), alternating. Refers to
the alternating pattern of rows of square, and rows of triangular, pore frames between longitudinal beams.
Type locality: GSC locality C-080584, Phantom Creek
Formation, Yakoun River, Graham Island, Queen Charlotte
Island, British Columbia.
Occurrence: Whiteaves and Phantom Creek formations,
Queen Charlotte Island; Japan; Tawi Sadh Member of the
Guwayza Formation, Oman.
Plate HOM02. Homoeoparonaella reciproca Carter. Magnification x150. Fig. 1(H). Carter et al. 1988, pl. 7, fig. 2.
Fig. 2. OM, BR706, 15800. Fig. 3. JP, MNA-10, MA11574.
199
Genus: Hsuum Pessagno 1977a
Type species: Hsuum cuestaense Pessagno 1977a
Synonymy:
1977a Hsuum n. gen – Pessagno, p. 280.
1986 Hsuum Pessagno 1977a, emend. – Takemura, p. 49.
1986 Transhsuum n. gen – Takemura, p. 51.
Original description: Test multicyrtoid, conical lacking
strictures. Cephalis conical, with small horn and sparse irregularly dispersed pores. Thorax trapezoidal with sparse
irregularly displaced pores. Abdomen and post-abdominal chambers with massive, continuous to discontinuous,
diverging costae; three to six rows of small square pore
frames with circular pores between costae. Costae of some
species with irregular branches that link adjoining costae
and obscure linearly arranged pore frames beneath. Pores
of all post-thoracic chambers tending to remain open during ontogeny and to be primary pores.
Original remarks: Hsuum n. gen., appears to build its test
by secreting costal projections each time a new chamber
is formed; linearly arranged square pore frames are then
secreted between costal projections. Because it shares the
same mode of test building as the Archaeodictyomitridae
Pessagno, it is tentatively placed in this family.
Hsuum differs from Archaeodictyomitra Pessagno
in having several rows of pores between costae and by
possessing primary rather than relict pores (cf. Pessagno,
1976).
Further remarks: For purposes of this catalogue we include Hsuum Pessagno and Transhsuum Takemura together, although we recognize morphological difference.
Etymology: This genus is named for Dr. Kenneth J. Hsu
(Swiss Federal Institute of Technology, Zurich, Switzerland) to honor his contributions to the study of the Franciscan complex.
Included species:
HSU01 Hsuum altile Hori & Otsuka 1989
HSU02 Hsuum arenaense Whalen & Carter 2002
HSU03 Hsuum busuangaense Yeh & Cheng 1996
HSU04 Hsuum exiguum Yeh & Cheng 1996
HSU05 Hsuum lucidum Yeh 1987b
3195 Hsuum matsuokai Isozaki & Matsuda 1985
3278 Hsuum medium (Takemura) 1986
HSU06 Hsuum mulleri Pessagno & Whalen 1982
HSU07 Hsuum optimum Carter 1988
HSU08 Hsuum philippinense Yeh & Cheng 1996
HSU11 Hsuum plectocostatum Carter n. sp.
HSU10 Hsuum sp. A sensu Carter 1988
Hsuum altile Hori & Otsuka 1989
Species code: HSU01
Synonymy:
1982 “Lithostrobus” sp. b – Kido, pl. 4, figs. 9, 10.
1982 Hsuum sp. – Matsuda & Isozaki, pl. 1, figs. 1, 2.
1984 Hsuum sp. B – Murchey, pl. 1, fig. 23.
1985 Hsuum sp. A – Kishida & Hisada, pl. 4, figs. 11, ?13, 14.
1988 Hsuum (?) matsuokai Isozaki & Matsuda – Hattori, pl. 13,
fig. E.
1989 Hsuum altile n. sp. – Hori & Otsuka, p. 180, pl. 1, figs. 1-6.
1990 Hsuum altile Hori & Otsuka – Hori, Fig. 9.33.
1996 Hsuum altile Hori & Otsuka – Yeh & Cheng, p. 108, pl. 10,
figs. 1, 2, 6, 10, 11.
1997 Hsuum altile Hori & Otsuka – Hori, pl.1, fig. 2.
2001 Hsuum altile Hori & Otsuka – Matsuoka et al., pl. 3, fig. 8.
2004 Hsuum altile Hori & Otsuka – Hori, pl. 5, fig. 41; pl. 22,
figs. 52-53.
? 2004 Hsuum altile Hori & Otsuka – Ishida et al., pl. 5, fig. 15.
2005 Hsuum altile Hori & Otsuka – Kashiwagi et al., pl. 6, fig. 1.
Original description: Test multi-segmented, exact number
of chambers unknown, possibly less than 8 or 9. Outline
of test gourd-shaped with a weak to strong stricture in
proximal 1/3 portion. Cephalis hemispherical with an apical horn; horn solid and mostly polygonal in cross-section.
The proximal portion of test, above stricture, possessing
irregularly arranged pores and circular to polygonal pore
frames; surfaces of pore frames smooth to rough, occasionally spiny (Pl. 1, Fig. 1c). The distal portion below stricture
200
inflated and having longitudinally and transversally aligned
pores and 14 to 19 longitudinal continuous costae; longitudinal costae mostly long, developed at an interval of 2 or 3
rows of pores, and frequently having branches. Immediately below stricture, some discontinuous costae occasionally
observed. In complete specimens, costae disappeared at the
distal end of test (Pl. 1, Figs. 3a, b, 5).
Original remarks: Hsuum altile sp. nov. is very similar to
Hsuum (?) matsuokai Isozaki and Matsuda, 1985b on its
form and costal arrangement. The former, however, differs
from the latter by lacking robust massive apical horn which
is tetraradiate cruciform in cross section and possessing
irregular arranged pores on proximal portion of test. On
the basis of morphological resemblances and stratigraphic
positions, H. sp. α (= the provisional name of H. altile) is
regarded as the ancestor of H. (?) matsuokai.
This species also resembles Hsuum sp. B of Takemura
(1986) and Hsuum parvulum Yeh, 1987 but differs in having
irregular pore frames on proximal part of test and in being
larger and gourd-shaped.
Hsuum sp. aff. H. mirabundum of Pessagno and Whalen
(1982) and Hsuum sp. A of Carter in Carter et al. (1988) are
similar to H. altile. However, the former two species can
be distinguished from the latter by possessing costae on
proximal portion of test.
Etymology: The name is derived from the Latin adjective
altilis, meaning stout.
Measurements (µm):
Based on 9 specimens.
HT
Av.
Max.
Min.
Height
306+
275
326
236+
Width
141
145
165
130
H/W
2.2+
1.9
2.2
1.6+
Type locality: The Mt. Norikuradake area, Azumi village,
Azumi-gun, Nagano Prefecture, central Japan.
Occurrence: Japan; Liminangcong Chert, Philippines;
Xialu chert, Tibet; Franciscan Complex, California.
Plate HSU01. Hsuum altile Hori & Otsuka. Magnification x200. Fig. 1(H). Hori & Otsuka 1989, pl. 1, fig. 1a. Figs. 2-4.
Hori & Otsuka 1989, pl. 1, figs. 2a, 3a, 5.
201
Hsuum arenaense Whalen & Carter 2002
Species code: HSU02
Synonymy:
1984 Hsuum sp. – Whalen & Pessagno, pl. 4, figs. 1-4.
2002 Hsuum arenaense n. sp. – Whalen & Carter, p. 124, pl. 12,
figs. 1, 2, 11, 15; pl. 17, figs 10, 11.
Original description: Conical test with approximately seven
to eight post-abdominal chambers. Dome-shaped cephalis
covered by layer of microgranular silica and terminating in
a very small horn. Thorax and post-abdominal chambers
mostly trapezoidal in outline (sub-rectangular distally),
increasing gradually in width and height as added. Costae,
about twice as high as wide, with lateral branches much
more irregularly developed in proximal parts of test; costae
becoming shorter, narrower with fewer lateral branches
and more linearly arranged in distal portions of test. Inner
layer consisting of large pore frames, square to rectangular,
with round to elliptical pores. Pore frames appear smaller
in proximal parts of test due to more extensive development
of costae and lateral branches.
Original remarks: The development of very irregularly
branching costae on the proximal portions of the test
(more developed on some specimens than others) distinguish Hsuum arenaense n. sp from H. mulleri Pessagno
and Whalen 1982, H. parvulum Yeh 1987 and from other
Lower and Middle Jurassic species of Hsuum.
Measurements (µm):
Based on 11 specimens.
Length
240
240
195
220
Width (Max.)
105
128
105
114
HT
Max.
Min.
Mean
Etymology: This species is named for Isla Arena located to
the southeast of its type area.
Type locality: Sample BPW80-30, San Hipólito Formation,
Vizcaino Peninsula, Baja California Sur, Mexico.
Occurrence: San Hipólito Formation, Baja California Sur.
Hsuum busuangaense Yeh & Cheng 1996
Species code: HSU03
Synonymy:
at distal portion). This form differs from H. altile Hori
and Otsuka (1989) by having a relatively inflated postabdominal chambers and by having apical portion with
short, irregularly arranged costae rather than with dense
polygonal pore frames at the outer layer test wall.
Original description: Test multicyrtid, subspindle-shaped,
pointed apically and terminating in a moderately long horn.
Cephalis subconical in shape. Cephalis, thorax, and abdomen chambers relatively narrower, outer layer of test wall
covered by short, irregularly arranged costae. Post-abdominal chambers inflated, outer layer of test wall covered with
long, continuous longitudinal costae. Two or three longitudinal rows of tetragonal pore frames between every two
post-abdominal costae. Post-abdominal chambers lacking
strictures. Final portion of post-abdominal chambers decreasing in width prominently.
Measurements (µm):
Five specimens measured.
1982 Hsuum sp. – Matsuda & Isozaki, pl. 1, fig. 2.
1996 Hsuum busuangaense n. sp. – Yeh & Cheng, p. 110, pl. 3,
figs. 5, 9, 13.
2004 Hsuum aff. altile Hori & Otsuka – Hori, pl. 4, fig. 19.
Original remarks: This form is characterized by having
a subspindle-shaped inflated test and by having two types
of costae structure (short, irregularly arranged costae at
apical portion and long, continuous longitudinal costae
202
HT
Mean
Max.
Min.
Max. test width
148
153
158
148
Max. test length
303
296
308
279
Length of horn
37
34
39
27
Etymology: This form is named for its type locality, the
Busuanga Island, Philippines.
Type locality: Sample CR91-30B, Liminangcong Chert,
Busuanga Island, Philippines.
Occurrence: Liminangcong Chert, Philippines; Japan.
Plate HSU02. Hsuum arenaense Whalen & Carter. Magnification Fig. 1a(H) x 200, Fig. 1b(H) x400. Fig. 1a,b(H).
Whalen & Carter 2002, pl. 12, figs. 1, 11.
Plate HSU03. Hsuum busuangaense Yeh & Cheng. Magnification x200. Fig. 1(H). Yeh & Cheng 1996, pl. 3, fig. 5.
Fig. 2. JP, UFI (22).
203
Hsuum exiguum Yeh & Cheng 1996
Species code: HSU04
Synonymy:
1989 Parahsuum (?) sp. B – Hori & Otsuka, p. 183, pl. 3,
figs. 11-12.
1989 Parahsuum (?) sp. Y – Hori & Otsuka, p. 182, pl. 3,
figs. 6-7.
1990 Parahsuum (?) sp. B – Hori, Fig. 9.32.
1997 Parahsuum (?) sp. B – Hori, pl. 1, fig. 5.
1996 Hsuum exiguum n. sp. – Yeh & Cheng, p. 110, pl. 3, figs. 1, 2,
6, 10; pl. 10, fig. 3.
2004 Hsuum sp. – Hori, pl. 3, fig. 30 only; pl. 22, figs. 57-60, 62.
2004 Hsuum sp. X sensu Hori & Otsuka – Hori, pl. 4,
figs. 20-23.
2004 Hsuum sp. Y sensu Hori & Otsuka – Hori, pl. 4, figs. 24-26.
2005 Hsuum exiguum Yeh & Cheng – Kashiwagi et al., pl. 6,
fig. 2.
Original description: Test relatively short, bell-shaped,
cephalis hemispherical in outline, with a moderately
long, stout horn. Cephalis and thorax covered by a layer
of medium-sized irregular polygonal pore frames. Costae
of outer latticed layer short, discontinuous and poorly
developed. Post-abdominal chambers slightly increasing in
width distally. Final post-abdominal chamber terminated
with a rim-like structure.
204
Original remarks: This form is characterized by its small,
short test and by having a stout long horn at cephalis and a
rim-like structure at its final post-abdominal chamber.
Measurements (µm):
Five specimens measured.
HT
Mean
Max.
Min.
Max. test width
124
130
142
124
Max. test length
213
216
238
203
Length of horn
52
55
43
66
Etymology: Exiguus-a-um (Latin, adj.) = short
Type locality: Sample CR91-30B, Liminangcong Chert,
Busuanga Island, Philippines.
Occurrence: Liminangcong Chert, Philippines; Tawi Sadh
Member of the Guwayza Formation, Oman; Japan..
Plate HSU04. Hsuum exiguum Yeh & Cheng. Magnification x300. Fig. 1(H). Yeh & Cheng 1996, pl. 3, fig. 1.
Fig. 2. Hori 1990, fig. 9-32. Fig. 3. JP, Nanjo mudstone-1. Fig. 4. JP, Nanjo mudstone-2. Fig. 5. JP, NKII19-14.
Fig. 6. OM, BR871-R08-30. Fig. 7. OM, BR871-R08-04. Fig. 8. OM, BR871-R07-08. Fig. 9. OM, BR871-R06-15.
Fig. 10. OM, BR871-R06-16. Fig. 11. OM, BR871-R06-23. Fig. 12. OM, BR828-2-R12-03.
205
Hsuum lucidum Yeh 1987b
Species code: HSU05
Synonymy:
1987b Hsuum (?) lucidum n. sp. – Yeh, p. 64, pl. 16, figs. 4, 8, 16.
1987b Hsuum validum n. sp. – Yeh, p. 66, pl. 3, fig. 26; pl. 5, figs.
17, 22; pl. 17, fig. 12; pl. 28, fig. 1.
1987b Hsuum sp. E – Yeh, p. 67, pl. 16, fig. 9; pl. 28, fig. 12.
2003 Transhsuum lucidum (Yeh) – Goričan et al., p. 296, pl. 5,
figs. 12, 13.
2004 Hsuum lucidum Yeh – Matsuoka, fig. 225.
Original description: Test conical, lobate, pointed apically,
usually with six to seven post-abdominal chambers. Cephalis small, conical, without rudimentary horn. Cephalis
and thorax covered with layer of microgranular silica. Abdomen and post-abdominal chambers rapidly increasing
in width, gradually increasing in length as added. Post-abdominal chambers with inner layer of massive tetragonal
pore frames with small circular pores; outer layer of mesh
work with short, moderately massive costae. Costae mostly
occurring on joints of chambers.
Original remarks: Hsuum (?) lucidum, n. sp., differs from
H. parasolensis Pessagno and Whalen by lacking a horn on
cephalis, and by having a more conical test with less massive costae.
Further remarks: According to Yeh’s (1987b) original description Hsuum validum differs from Hsuum lucidum by
having a subconical instead of a conical test. We consider
that this difference is not expressed well enough to distinguish two different species, therefore both species are synonymized.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length (max.)
245
253
270
241
Width (max.)
153
148
155
140
Etymology: Lucidus-a-um (latin, adj.) = bright.
Type locality: Sample OR-600A, Hyde Formation at IzeePaulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, Oregon; Fannin
and Phantom Creek formations, Queen Charlotte Islands;
Skrile Formation, Slovenia; Tawi Sadh Member of the Guwayza Formation and Musallah Formation, Oman; Mino
Terrane, Japan.
Plate HSU05. Hsuum lucidum Yeh. Magnification x300. Fig. 1(H). Yeh, 1987b, pl. 16, fig. 4. Fig. 2. QCI, GSC loc. C304568, GSC 128809. Fig. 3. QCI, GSC loc. C-175309, GSC 128810. Fig. 4. QCI, GSC loc. C-304568, GSC 111809.
Fig. 5. SI, MM5.00, 000103. Figs. 6-7. Goričan et al. 2003, pl. 5, figs. 12-13. Fig. 8. JP, MNA-10, MA13145.
Fig. 9. OM-00-117, 021128. Fig. 10. OM-00-254, 022202. Fig. 11. OM, BR1123-R05-06.
206
207
Hsuum matsuokai Isozaki & Matsuda 1985
Species code: 3195
Synonymy:
1982 Hsuum sp. C – Hattori & Yoshimura, pl. 3, fig. 8.
1982 Hsuum sp. B – Kishida & Sugano, pl. 7, figs. 14-16.
1982 Unnamed nassellaria – Wakita & Okamura, pl. 7, fig. 3.
1984 Hsuum sp. – Yao, pl. 1, figs. 6-7.
1985 Hsuum sp. – Ishida, pl. 1, fig. 3.
1985 Hsuum (?) matsuokai n. sp. – Isozaki & Matsuda, p. 438,
pl. 3, figs. 1-14.
1985 Hsuum maxwelli Pessagno – De Wever & Miconnet, pl. 4,
fig. 3.
1986 Hsuum primum n. sp. – Takemura, p. 50, pl. 5, figs. 17-21.
1986 Hsuum sp. – Matsuoka, pl. 2, figs. 1, 3.
1987 Hsuum aff. mclaughlini Pessagno & Blome – Goričan,
p. 183, pl. 2, fig. 11.
1987 Hsuum primum Takemura – Hattori, pl. 17, figs. 11-13, not
figs. 8-9.
Not 1988 Hsuum (?) matsuokai Isozaki & Matsuda – Hattori,
pl. 13, fig. E.
1988 Hsuum (?) matsuokai Isozaki & Matsuda – Sashida, p. 19,
pl. 4, figs. 16-18.
1989 Hsuum primum Takemura – Hattori & Sakamoto, pl. 15,
figs. I-J.
1989 Hsuum (?) matsuokai Isozaki & Matsuda – Hattori &
Sakamoto, pl. 16, fig. I.
1990 Hsuum matsuokai Isozaki & Matsuda – Hori, Fig. 9.53.
1991 Hsuum matsuokai Isozaki & Matsuda – Yao, pl. 2, fig. 18.
1992 Ogivus falloti n. sp. – El Kadiri, p. 46, pl. 2, figs. 3-4.
1992 Hsuum matsuokai Isozaki & Matsuda – Sashida, pl. 2,
fig. 4.
1994 Hsuum matsuokai Isozaki & Matsuda – Goričan, p. 73,
pl. 19, figs. 9, 11-13.
1995a Hsuum matsuokai Isozaki & Matsuda – Baumgartner et
al., p. 284, pl. 3195, figs. 1-5(H).
2001 Hsuum matsuokai Isozaki & Matsuda – Matsuoka et al.,
pl. 3, fig. 4.
2004 Hsuum matsuokai Isozaki & Matsuda – Hori, pl. 2, fig. 56,
pl. 9, fig. 36, pl. 10, fig. 4.
2004 Hsuum aff. matsuokai Isozaki & Matsuda – Hori, pl. 9,
fig. 37, pl. 10, figs. 5-6, 8.
2004 Hsuum matsuokai Isozaki & Matsuda – Suzuki & Ogane,
pl. 9 fig. 20.
2004 Hsuum primum Takemura – Suzuki & Ogane, pl. 9,
figs. 22, 23.
2005 Hsuum matsuokai Isozaki & Matsuda – Hori, pl. 13, fig. 6.
Original description: Shell of 7 segments, possibly more,
long, spindle-shaped; slenderly conical in proximal 3 segments; broad, barrel-shaped in distal half. Cephalis conical
with robust apical horn, coated by outer microgranular layer,
on which sparse irregularly dispersed pores remain open. Apical horn variously ornamented with thick blades or narrow
grooves, having transverse section typically of tetraradiate cruciform with 4 blades at base, almost circular at tip. Internally,
6 collar pores, divided by median bar, D-bar, V-bar, 2 L-bars
and 2 l-bars. Post-cephalic segments free from the outer microgranular layer, trapezoidal in longitudinal section; each
segment becoming wide distally except for the distal-most
one, which is reversely trapezoidal in longitudinal section.
Average ratio of height to width of a single segment approximately 1:3 for thorax and abdomen, approximately 1:4 for
208
post-abdominal segments. Wall of segment, thin; its longitudinal section flat in proximal half, slightly convex outward in
distal half. Pores circular, uniform in size. Square pore frames
aligned longitudinally and transversely; in 2 to 3 longitudinal
rows of pores between every neighbouring pairs of costae, in
4 transverse rows for each segment. 16-19 continuous costae developing on post-abdominal segments. Weak irregular
transverse bars rarely present, linking adjoining costae. Internal partitions rudimentary, circular in outline with a large
centrally placed aperture.
Original remarks: Ornamentation on cephalis varies considerably from specimen to specimen. Generallly, larger specimens tend to have slenderer shell and apical horn of more
completely tetraradiate cruciform section (pl. 3, figs. 1-2). Most
of the specimens possess 4 rudimentary ornamenting blades
around apical horn.
This species is distinguished from other species of the genus
Hsuum Pessagno by its extraordinarily conspicuous apical
horn with various ornamentation and restricted development
of thick costae within distal half of the shell. Furthermore,
bifurcation of costae or distally widening silhouette of the shell
cannot be recognized in H. (?) matsuokai n. sp. although they
are common features among most of the species belonging to
Hsuum Pessagno.
Pessagno & Whalen (1982) established some new multicyrtoid nassellarian genera of Early to Middle Jurassic age, such as
Droltus and Canutus, which are essentially characterized by linear arrangement of square pore frames. H. (?) matsuokai n. sp.
is not referable to them in wall structure mentioned above. On
the other hand, the proximal half of the shell of this species
looks rather like that of genus Parahsuum Yao, except for its
robust apical horn. In these circumstances, this species is here
provisionally classified under genus Hsuum Pessagno.
Further remarks: By Baumgartner et al. (1995a): This species differs from its ancestor Hsuum altile Hori & Otsuka
1989 by having a robust massive apical horn which is tetraradiate cruciform in cross-section.
Measurements (µm):
Based on 13 specimens.
HT
Mean
Max.
Min.
Height
380
310
410
240
Width
150
140
160
120
Etymology: This species is named for Dr. Matsuoka in honor
to his contribution to Jurassic radiolarian biostratigraphy
in southwest Japan.
Type locality: Sample 140, Hisuikyo, Kamiaso area, Gifu
Prefecture, central Japan.
Occurrence: Worldwide.
Plate 3195. Hsuum matsuokai Isozaki & Matsuda. Magnification 200x. Fig. 1(H). Isozaki & Matsuda 1985, pl. 3,
fig. 1. Fig. 2. JP, HM1-11, RH623. Fig. 3. OM, BR292-4-R10-01. Fig. 4. OM, BR292-4-R10-03. Fig. 5. OM-99-137,
020710. Fig. 6. OM-99-137, 000827.
209
Hsuum medium (Takemura) 1986
Species code: 3278
Synonymy:
1986 Transhsuum medium n. sp. – Takemura, p. 51, pl. 6,
figs. 1-2; not pl. 5, figs. 25-26.
1987 Hsuum sp. – Hattori, pl. 17, fig. 16.
1990 Transhsuum medium Takemura – Hori, fig. 9.43.
1997 Transhsuum medium Takemura – Hori, pl. 1, fig. 4.
1995a Transhsuum medium Takemura – Baumgartner et al.,
p. 582, pl. 3278, figs. 1-3.
Original description: Shell conical to cylindrical, with 10
to 15 segments, with strictures at joints of distal segments.
Cephalis conical and poreless, with or without conical apical horn. Thorax truncated-conical usually with a single
transverse row of small pores. In some specimens some
longitudinal ridges covering on the surface of cephalo-thorax. Abdomen and post-abdominal segments cylindrical
with small pores, which are usually rectangularly arranged
on the inner surface. Each post-abdominal segment bearing four to five transverse rows of pores. Indistinct discontinuous costae, their length is equal to the height of one
segment, lying on distal segments. In mature specimens,
small spines arising on the shell surface.
Original remarks: The shell structure of the proximal part
of Transhsuum medium resembles that of the distal part
of Parahsuum cruciferum (pl. 5, figs. 9, 11). Therefore it
suggests that this new species represents the initial stage
of the formation of Transhsuum-type discontinuous costae
and that it is the intermediate form between Parahsuumlike form and Transhsuum.
Further remarks: Herein we follow Baumgartner et al.
(1995a) who assigned forms with a strong apical horn to
this species.
Measurements (µm):
Based on 8 specimens.
Length of shell
Maximum width of shell
Min.
265
100
Max.
385
135
Etymology: The name medium, derived from medius,
means intermediate.
Type locality: Sample TKN-105, Komami, Yamato Village,
Gifu Prefecture, central Japan.
Occurrence: Japan; Sogno Formation, Lombardy, Italy.
Hsuum mulleri Pessagno & Whalen 1982
Species code: HSU06
Synonymy:
1982 Hsuum mulleri n. sp. – Pessagno & Whalen, p. 133, pl. 5,
figs. 6, 8, 9; pl. 12, figs. 16-17.
1982 Hsuum sp. D – Pessagno & Whalen, p. 134, pl. 5, fig. 5.
Original description: Test conical, elongate for genus, usually with seven post-abdominal chambers. Cephalis hemispherical with short rudimentary horn; other chambers,
except for later post-abdominal chambers, trapezoidal in
outline; later post-abdominal chambers subrectangular in
outline. Cephalis and thorax imperforate to sparsely perforate; pores buried by layer of microgranular silica. Abdomen and post-abdominal chambers with inner latticed
layer of fragile, thin, linearly arranged, square pore frames
and outer latticed layer of moderately massive, discontinuous costae which are inserted between rows of pore frames.
Costae with few ramifications, more or less equally developed over abdomen and post-abdominal chambers. Postabdominal chambers increasing slightly in length and more
rapidly in width as added; final two or three chambers show
little increase in width.
Original remarks: Hsuum mulleri, n. sp., differs from
H. sp. D by having a narrower, more elongate test and less
massive, more evenly distributed costae which show few
branches laterally.
210
Measurements (µm):
Based on 6 specimens.
Length excluding horn
325.0
325.0
230.0
270.8
Width (max.)
137.5
145.0
100.0
126.3
HT
Max.
Min.
Mean
Etymology: This species is named for Dr. Jan E. Muller
(Geological Survey of Canada, Vancouver) in honor of his
contributions to the geology of British Columbia.
Type locality: Sample QC 534, Fannin Formation (Maude
Formation in Pessagno & Whalen, 1982), Queen Charlotte
Islands, British Columbia.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Fernie Formation, Williston Lake, northeastern British Columbia.
Plate 3278. Hsuum medium (Takemura). Magnification 200x. Fig. 1(H). Takemura 1986, pl. 6, fig. 1. Fig. 2. Hori 1990,
fig. 9-43.
Plate HSU06. Hsuum mulleri Pessagno & Whalen. Magnification x200. Fig. 1(H). Pessagno & Whalen 1982, pl. 5,
fig. 6. Fig. 2. QCI, GSC loc. C-080611, GSC 128811. Fig. 3. NBC, GSC loc. C-305208, GSC 128812. Fig. 4. QCI, GSC loc.
C-080611, GSC 128914. Fig. 5. QCI, GSC loc. C-080611, GSC 128813.
211
Hsuum optimum Carter 1988
Species code: HSU07
Synonymy:
1988 Hsuum optimus Carter n. sp. – Carter et al., p. 51, pl. 5, fig. 6.
Original diagnosis: Test large, slender conical with short
cylindrical horn. Chamber boundaries marked by rows of
distinct elongate nodes. Pores small and circular.
Original description: Test large, elongate conical, pointed
apically with short, slender, cylindrical horn and as many as
11 postabdominal chambers. Cephalis hemispherical and
imperforate; all remaining chambers perforate, trapezoidal
in outline. Chamber width increases gradually throughout
entire test length; chamber height fairly constant. Test has
inner layer of small, linearly arranged, square to rectangular pore frames; pores circular to subcircular. Outer layer
consists, in the first three or four chambers, of discontinuous costae. In subsequent chambers, the surface is marked
by nonlinear, elongate nodes at chamber joints.
Original remarks: Differs from all other species of Hsuum
in having elongate nodes (rather than raised costae) superimposed on the longitudinal bars of the inner latticed layer.
Differs from Hsuum sp. B, in having a more conical shape
with more numerous, closely spaced elongate nodes. Extremely abundant.
Further remarks: Hsuum optimum differs from H. philippinense Yeh & Cheng by having a hemispherical cephalis
and a short horn, circular in cross section.
Measurements (µm):
Based on 14 specimens.
Length excluding horn
Maximum width
HT
346
190
Av.
293
165
Max.
346
190
Min.
210
125
Etymology: Latin, optimus (adj.), best.
Type locality: GSC locality C-080579, Whiteaves Formation, Creek locality, Maude Island, Queen Charlotte Islands, British Columbia.
Occurrence: Whiteaves and Phantom Creek formations,
Queen Charlotte Islands; Tawi Sadh Member of the Guwayza Formation, Oman.
Hsuum philippinense Yeh & Cheng 1996
Species code: HSU08
Synonymy:
1996 Hsuum philippinense n. sp. – Yeh & Cheng, p. 112, pl. 3,
figs. 3, 4, 8, 12.
1996 Hsuum sp. aff. H. philippinense n. sp. – Yeh & Cheng,
p. 112, pl. 9, figs. 3, 4, 10, 11, 12.
Original description: Test subcylindrical, pointed apically and terminating in a short horn. Cephalis and thorax
covered by a layer of microgranular silica, costae present
at apical portion, often arranged in radial pattern. Costae
of outer latticed layer short, discontinuous, and staggered.
Costae quite well-developed, nodose in lateral view. Final
post-abdominal chambers slightly decreasing in width as
added.
Original remarks: This form is characterized by having a
triangular-shaped cephalis with a short pointed horn, and
by having short, discontinuous, nodose costae throughout
the post-abdominal chambers.
212
Further remarks: See remarks under Hsuum optimum
Carter.
Measurements (µm):
Four specimens measured.
HT
Mean
Max.
Min.
Max. test width
121
124
132
121
Max. test length
271
281
313
259
Length of horn
25
28
36
20
Etymology: This species is named for the country of its type
locality, the Philippines.
Type locality: Busuanga Island (CR91-30B), Philippines.
Occurrence: Busuanga Island, Philippines; Japan; Tawi
Sadh Member of the Guwayza Formation and Musallah
Formation, Oman.
Plate HSU07. Hsuum optimum Carter. Magnification x200. Fig. 1(H). Carter et al. 1998, pl. 5, fig. 6.
Fig. 2. OM, BR871-R08-07.
Plate HSU08. Hsuum philippinense Yeh & Cheng. Magnification x250. Fig. 1(H). Yeh & Cheng, 1996, pl. 3, fig. 3.
Fig. 2. OM-00-256, 022423. Fig. 3. OM-00-256, 022525. Fig. 4. OM, BR871-05. Fig. 5. JP, NK9-50.
213
Hsuum plectocostatum Carter n. sp.
Species code: HSU11
Synonymy:
1988 Hsuum sp. B – Carter et al., p. 52, pl. 5, figs. 7, 8.
1989 Hsuum (?) sp. Z – Hori & Otsuka, p. 182, pl. 3, fig. 8.
1991 Hsuum sp. B – Carter & Jakobs, p. 343, pl. 3, fig. 15.
1996 Hsuum sp. cf. H. philippinense n. sp. – Yeh & Cheng, p. 114,
pl. 9, figs. 5, 13.
Type designation: Holotype GSC 99415 (Carter & Jakobs
1991, pl. 3, fig. 15; late early Aalenian), from GSC loc. C156399, Phantom Creek Formation.
Description: Test large, conical, pointed apically, with eight
or nine post-abdominal chambers and a short, tapering
cylindrical horn. Cephalis hemispherical and imperforate
with several heavy outer costae penetrating almost on to
horn; all remaining chambers trapezoidal, expanding
gradually in width as added; final post-abdominal
sometimes decreasing in width (see holotype, pl. HSU11,
fig. 1). Inner layer of pore frames linearly arranged, square
to subrectangular in shape, pores subrounded. Outer layer
of test consisting of irregularly twisted costae with small
rounded nodes superimposed on circumferential ridges.
Remarks: Hsuum plectocostatum n. sp. differs from Hsuum
optimum Carter in having a more broadly conical shape and
Hsuum sp. A sensu Carter 1988
Species code: HSU10
Synonymy:
1988 Hsuum sp. A – Carter et al., p. 52, pl. 5, fig. 2.
Remarks: The distal post-abdominal chambers of this
pyritized form are similar to Hsuum altile Hori & Otsuka
but initial chambers are quite different: they lack the typical
porous structure of H. altile and instead have discontinuous
costae that appear almost node-like.
Occurrence: Whiteaves Formation, Queen Charlotte Islands.
214
the costae, although irregularly fluted/twisted, are more
continuous and possess small rounded nodes at ridges. It
differs from H. infirmum Sashida (1988) in having a much
larger test with a lobate outline, and stronger costae.
Measurements (µm):
Based on 13 specimens.
HT
Mean
Max.
Min.
Max. test width
170
150
237
112.5
Max. test length
329
286
416
229
Length of horn
15
24
37
15
Etymology: From the Latin: plecto + costatus, -a, -um referring to twisted costae.
Type locality: Sample GSC loc. C-156399, Phantom Creek
Formation Yakoun River, 2.0 km south of Ghost Creek, east
side of river; Graham Island; Queen Charlotte Islands, British Columbia.
Occurrence: Whiteaves and Phantom Creek formations,
Queen Charlotte Islands; Liminangcong Chert, Philippines;
Japan.
Plate HSU11. Hsuum plectocostatum Carter n. sp. Magnification x200. Fig. 1(H). Carter & Jakobs 1991, pl. 3, fig. 15.
Fig. 2. Carter et al. 1988, pl. 5, fig. 7. Fig. 3. Carter et al. 1988, pl. 5, fig. 8.
Plate HSU10. Hsuum sp. A sensu Carter. Magnification x250. Fig. 1. Carter et al. 1988, pl. 5, fig. 2.
215
Genus: Katroma Pessagno & Poisson 1981, emend. Whalen & Carter 1998
Type species: Katroma neagui Pessagno & Poisson 1981
Synonymy:
Original description: Test multicyrtid, comprised of
cephalis, thorax, abdomen, and with type species one postabdominal chamber. Post-abdominal chamber terminating
in long, cylindrical, open, tubular extension. Cephalis
hemispherical with horn; thorax and abdomen trapezoidal
in outline. First post-abdominal chamber subspherical,
considerably larger than previous chambers and with
variable number of medially arranged circumferential
spines.
closed, while on others it remains open. Since this tube is
often broken distally, it is not considered morphologically
diagnostic whether it is open or closed.
This volume: Since the open or closed tube is not a
diagnostic character, there seems to be no difference between
Katroma and Podobursa, in which case Katroma should be
considered a junior synonym of Podobursa. For the time
being, we retain the generic name Katroma but restrict it to
Early Jurassic species, because a phylogenetic relationship
between Early and Middle Jurassic Syringocapsidae is
not yet fully understood. It is interesting to note that no
Katroma, Syringocapsa or Podobursa have been recorded
in a well preserved early-middle Aalenian HK 140 sample
from Japan, although they are diverse in older and younger
assemblages (see Yao, 1997).
Original remarks: Katroma differs from Podobursa Wisniowski by having an open tube on its final post-abdominal
chamber.
Etymology: The name Katroma is formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, Pt. IV,
Recommendation 40, p.113). Its gender is feminine.
Further remarks: By Whalen & Carter (1998): The small
cephalic spines noted by De Wever (1982a) on specimens
of Katroma from Turkey were not observed on either the
Sinemurian or upper Pliensbachian specimens of Katroma
from British Columbia or Baja California Sur (Whalen,
1985). Due to the vagaries of preservation, these spines are
not considered a diagnostic feature of this genus. Transmitted light photography has revealed that on some species of
Katroma, the final chamber is the postabdominal chamber,
while on others it is the abdominal chamber. The genus
Katroma is hereby emended to accommodate both conditions. In addition, the terminal tube on some species is
Included species:
KAT07 Katroma angusta Yeh 1987b
KAT08 Katroma aurita Whalen & Carter 2002
KAT09 Katroma bicornus De Wever 1982a
KAT12 Katroma brevitubus Dumitrica & Goričan n. sp.
KAT10 Katroma clara Yeh 1987b
KAT17 Katroma elongata Carter n. sp.
KAT13 Katroma neagui Pessagno & Poisson 1981, emend.
De Wever 1982a
KAT14 Katroma ninstintsi Carter 1988
KAT16 Katroma? sinetubus Carter n. sp.
KAT18 Katroma sp. 4
1981 Katroma n. gen. – Pessagno & Poisson, p. 62.
1982a Katroma Pessagno & Poisson emend. – De Wever, p. 193.
1988 Katroma Pessagno & Poisson – Hori, p. 551.
1998 Katroma Pessagno & Poisson, emend. – Whalen & Carter,
p. 69.
Katroma angusta Yeh 1987b
Species code: KAT07
Synonymy:
1984 Katroma sp. – Whalen & Pessagno, pl. 1, fig. 3.
1987b Katroma angusta n. sp. – Yeh, p. 79, pl. 23, fig. 8; pl. 30,
fig. 10.
1987b Katroma inflata n. sp. – Yeh, p. 81, pl. 9, fig. 11; pl. 10,
figs. 12-13, 22.
1998 Katroma megasphaera n. sp. – Yeh & Cheng, p. 28, pl. 5,
fig. 18; pl. 7, figs. 16, 18, 21, not figs. 9, 20; pl. 9, figs. 15, 16,
25; pl. 10, figs. 6, 11, 19.
2002 Katroma angusta Yeh – Whalen & Carter, p. 134, pl. 14,
figs. 1-3, 9, 10; pl. 18, figs. 7, 8.
? 2004 Katroma angusta Yeh – Hori et al., pl. 6, fig. 4.
Original description: Cephalis small, dome-shaped, with
short, well-developed horn. Horn circular in axial section.
Thorax and abdomen closely spaced, trapezoidal in outline.
First post-abdominal chamber inflated in larger subellipsoidal outline and terminating in a narrow tubular extension.
Tubular extension tapering distally. Cephalis, thorax, and
abdomen sparsely perforate, covered with layer of microgranular silica. Meshwork of first post-abdominal chamber
216
and tubular extension comprised mainly of pentagonal and
hexagonal pore frames. Size of pore frames decreasing apically and distally with largest pore frames at central portion
of post-abdominal chamber. Pore frames thin in rims and
sides.
Original remarks: Katroma angusta, n. sp., differs from
K. inflata, n. sp., by having a smaller, narrower test with
shorter horn.
Measurements (µm):
Ten specimens measured.
HT
Length of apical horn
37
Length of proximal
37
conical portion
Width of conical portion at base 73
146
Length of inflated segment
154
Width of inflated segment
Mean
40
Max.
43
Min.
37
52
66
37
72
140
154
73
146
161
70
132
150
Etymology: Angustus-a-um (Latin, adj.) = narrow.
Type locality: Sample OR-589D, Warm Springs member,
Snowshoe Formation, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, Oregon;
Ghost Creek Formation, Queen Charlotte Islands; San
Hipólito Formation, Baja California Sur; Liminangcong
Chert, Philippines; Dürrnberg Formation, Austria.
Plate KAT07. Katroma angusta Yeh. Magnification x200. Fig. 1(H). Yeh 1987b, pl. 23, fig. 8. Fig. 2. QCI, GSC loc. C304281, GSC 128817. Fig. 3. QCI, GSC loc. C-080612, GSC 128818. Fig. 4. QCI, GSC loc. C-140418, GSC 128819.
Fig. 5. Whalen & Carter 2002, pl. 14, fig. 2. Fig. 6. Whalen & Carter 2002, pl. 14, fig. 1. Fig. 7. AT, BMW21-37.
217
Katroma aurita Whalen & Carter 2002
Species code: KAT08
Synonymy:
1984 Katroma spp. – Whalen & Pessagno, pl. 1, figs. 1, 7.
2002 Katroma aurita n. sp. – Whalen & Carter, p. 134, pl. 13,
figs. 4, 8, 9; pl. 18, figs. 1, 2, 5.
Original description: Test multicyrtid with cephalis, thorax, abdomen and post-abdominal chamber. Small cephalis with massive horn, usually double-pronged. Prongs of
horn at an angle of approximately 80° to each other, tapering distally, flattened, with small irregularly spaced spines
on margins; horn sometimes more irregular, without clear
development of two prongs. Cephalis and thorax hemispherical, perforate but often covered by a thin layer of
microgranular silica. Abdomen trapezoidal in outline, perforate, with pores sometimes marked by thin layer of microgranular silica. Large inflated post-abdominal chamber
sub-spherical in shape, widest at central point and ending
in a closed, tapering cylindrical tube. Meshwork on postabdominal chamber composed predominantly of pentagonal pore frames, larger medially and decreasing in size
towards the abdomen and closed terminal tube. Numerous strong circumferential spines located at widest part of
post-abdominal chamber.
Original remarks: Katroma aurita n. sp. is distinguished
from K. bicornus De Wever 1982, by the spines on the
double-pronged horn, as well as the more numerous,
circumferential spines positioned at right angles to the
post-abdominal chamber.
Measurements (µm):
(n) = number of specimens measured
Length (14) (excludes horn)
315
345
248
290
Width (max.) (15)
135
150
105
129
HT
Max.
Min.
Mean
Etymology: Auritus, a, um (Latin, adj.) = long eared lepus.
Type locality: Sample BPW80-30, San Hipólito Formation,
Vizcaino Peninsula, Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur.
Katroma bicornus De Wever 1982a
Species code: KAT09
Synonymy:
1982a Katroma bicornus n. sp. – De Wever, p. 193, pl. 3, figs. 1-4.
1982b Katroma bicornus De Wever – De Wever, p. 304, pl. 46,
figs. 1-4.
1982 Syringocapsa sp. A – Imoto et al., pl. 1, fig. 7.
1982 Katroma bicornus De Wever - De Wever & Origlia-Devos,
pl. 1, figs. H, I.
1984 Katroma aff. neagui Pessagno & Poisson – Murchey, pl. 1,
fig. 28.
1987b Katroma bifurca n. sp. – Yeh, p. 79, pl. 3, figs. 3-4, 24.
1989 Katroma sp. B – Hattori, pl. 3, fig. A.
1990 Katroma cf. bicornus De Wever – Hori, Fig. 8.19.
1996 Katroma bicornus De Wever – Tumanda et al., p. 181,
Fig. 5.1.
1997 Katroma cf. bicornus De Wever – Hori, pl. 1, fig. 21.
1997 Katroma bicornus De Wever – Yao, pl. 11, fig. 544.
1998 Katroma bicornus De Wever – Yeh & Cheng, p. 28, pl. 7,
fig. 22.
2004 Katroma bicornus De Wever – Matsuoka, fig. 121.
Original description: Katroma bearing two stout spines
disposed at 120° from one another, on each side of test
axis. Porous cephalis in shape of truncated cone. Test bears
thin, long spines probably related to cephalic spines. Porous
thorax in shape of a truncated cone. There is almost no
change in contour between cephalis and thorax. Abdomen
inflated, much larger than cephalis and thorax, wider than
long. Its median part bears about ten radial spines, rounded
in cross-section. Postabdominal tube is subcylindrical,
distally closed, with pores sometimes elongated in axial
direction. Pores are large all over test with a maximum size
on abdomen.
218
Original remarks: Pores sometimes filled by a thin network
as a spider’s web. This form differs from K. neagui Pessagno
and Poisson by its two stout horns, long cephalic spines,
larger pores and a wider, distinct abdomen.
Measurements (µm):
Based on 4 specimens.
Length of horns
Length of cephalis
Length of thorax
Length of abdomen
Length of postabdominal tube
Width of cephalis in the middle
Width of thorax in the middle
Width of abdomen
Mean
65
53
51
119
Min.
60
50
45
114
43
62
152
42
50
140
Max.
71
60
59
125
200
45
71
165
HT
60
60
59
125
42
65
165
Etymology: From latin bi-, two and cornu, -us, horn; form
with two horns.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Nicely Formation, Oregon; Franciscan Complex, California; Drimos
Formation, Greece; Tawi Sadh Member of the Guwayza
Formation, Oman; Liminangcong Chert, Philippines;
Japan.
Plate KAT08. Katroma aurita Whalen & Carter. Magnification Fig. 1a(H) x200, Fig. 1b(H) x300. Fig. 1a(H). Whalen &
Carter 2002, pl. 13, fig. 4. Fig. 1b(H). Whalen & Carter 2002, pl. 13, fig. 8.
Plate KAT09. Katroma bicornus De Wever. Magnification x200. Fig. 1(H). De Wever 1982a, pl. 3, fig. 2.
Fig. 2. OM, BR474-R17-09. Fig. 3. JP, IYII24-15. Fig. 4. Hori 1990, Fig. 8.19.
219
Katroma brevitubus Dumitrica & Goričan n. sp.
Species code: KAT12
Synonymy:
1982 Syringocapsa sp. B – Yao, pl. 4, figs. 14-15.
1982 Syringocapsa sp. B – Yao et al., pl. 2, fig. 15.
1982 Syringocapsa sp. B – Imoto et al., pl. 1, fig. 8.
1990 Syringocapsa sp. B – Hori, Fig. 8.11.
1997 Syringocapsa sp. B of Yao 1982 – Hori, pl. 1, fig. 18.
1997 Syringocapsa sp. D – Yao, pl. 11, fig. 545.
1998 Katroma megasphaera n. sp. – Yeh & Cheng, p. 28, pl. 7,
figs. 9, 20 only.
1998 Katroma sp. B – Yeh & Cheng, p. 30, pl. 9, fig. 23.
2001 Syringocapsa inflata (Yeh) – Gawlick et al., pl. 6, fig. 6.
2004 Syringocapsa sp. – Hori, pl. 5, fig. 61.
Type designation: Specimen SG 022014 from sample
OM-00-252, Musallah Formation, Jabal Buwaydah center
east.
Description: Test composed of three segments and a tubular
extension. Cephalis and thorax short, broadly conical,
collar stricture not well pronounced externally. Apical horn
indistinct, short, circular in cross-section. Cephalothorax
bears small circular pores. Abdomen inflated, spherical,
much larger than cephalothorax, covered with mediumsized hexagonal and pentagonal pore frames. Abdomen
without circumferential spines. Tubular extension short,
inverted conical, porous. Pores similar in size to those of
abdomen. Tubular extension closed; in well-preserved
220
specimens it ends with a short spine, circular in crosssection.
Remarks: Katroma brevitubus n. sp. differs from K. angusta
Yeh by having a much shorter terminal tube. It differs from
Katroma ninstintsi Carter by having a shorter cephalothorax, more expanded abdomen, and shorter terminal tube.
Measurements (µm):
Based on 11 specimens.
Length of cephalis and thorax
Length of abdomen
Length of terminal tube
Maximum width of abdomen
HT
42
145
64
159
Min.
33
104
36
124
Max.
55
156
79
183
Mean
43
125
55
141
Etymology: From Latin: brevis,-e (short) and tubus,-i
(tube); noun.
Type locality: Sample OM-00-252 from Musallah Formation, Jabal Buwaydah center east.
Occurrence: Musallah Formation, Haliw (Aqil) Formation
and Tawi Sadh Member of the Guwayza Formation, Oman;
Dürrnberg Formation, Austria; Liminangcong Chert, Philippines; Japan.
Plate KAT12. Katroma brevitubus Dumitrica & Goričan n. sp. Magnification x200. Fig. 1(H). OM-00-252-022014.
Fig. 2. OM, BR706-R14-02. Fig. 3. OM-00-252-021819. Fig. 4. OM, Haliw-039-R06-22. Fig. 5. OM, Haliw-038-R09-02.
Fig. 6. OM-00-251-021422. Fig. 7. OM-00-252-021814. Fig. 8. AT, BMW21-18. Fig. 9. JP, MNA-10, MA118883.
221
Katroma clara Yeh 1987b
Species code: KAT10
Synonymy:
1984 Katroma sp. – Whalen & Pessagno, pl. 1, fig. 2.
1987b Katroma clara n. sp. – Yeh, p. 80, pl. 3, figs. 6-7.
1987 Katroma neagui Pessagno & Poisson, emend. De Wever
– Goričan, p. 184, pl. 1, fig. 2.
1988 Katroma kurusuensis n. sp. – Hori, p. 553, fig. 6.1a-5.
1989 Katroma sp. A – Hattori, pl. 2, fig. K.
1990 Katroma sp. N – Hori, fig. 8.20.
1990 Katroma kurusuensis Hori – Hori, fig. 8.21.
1997 Katroma kurusuensis Hori – Hori, pl. 1, fig. 16.
? 1998 Katroma sp. – Cordey, p. 110, pl. 22, figs. 7, 10.
2002 Katroma clara Yeh – Whalen & Carter, p. 134, pl. 14,
figs. 4, 5, 11, 12, 15; pl. 18, figs. 12, 13.
2004 Katroma clara Yeh – Matsuoka, fig. 112.
2004 Katroma sp. – Matsuoka, fig. 113.
2005 Katroma kurusuensis Hori – Hori, pl. 9, fig. 13.
Original description: Cephalis dome-shaped, with crownlike horn having four to five short branches, each branch
circular in cross-section. Thorax and abdomen trapezoidal
in outline. Three earlier chambers narrow, long, subcylindrical in shape, sparsely perforate, covered by layer of microgranular silica. First post-abdominal chamber inflated,
subspherical in outline, terminating in narrow subcylindrical tubular extension. Test mainly comprised of pentagonal
and hexagonal pore frames. Pore frames small on earlier
chambers, medium-sized on postabdominal chamber with
thick rims and thin sides. One row of ten to twelve short
circumferential spines on equatorial surface of first postabdominal chamber. Tubular extension perforate with
small pores in spiral rows of pore frames.
222
Original remarks: Katroma clara, n. sp., differs from
Katroma bifurca, n. sp., by having a horn with four to
five branches rather than with two branches, by having
circumferential spines on inflated post-abdominal chamber,
and by possessing a test with earlier chamber subcylindrical
in shape rather than conical in shape.
Further remarks: By Whalen & Carter (2002): Katroma
clara Yeh differs from K. neagui Pessagno and Poisson 1981,
by the more globular shape of the post-abdominal chamber
and from all other species of Katroma by the distinctive,
branching horn.
Measurements (µm):
Ten specimens measured.
HT Mean Max.
Length of apical horn
22
28
40
Length of proximal conical
108 109
110
portion
Width of conical portion at base 86
83
86
162 162
162
Length of inflated segment
151 151
200
Width of inflated segment
Min.
22
108
80
150
151
Etymology: Clarus-a-um (Latin, adj.) = clear.
Type locality: Sample OR-536J, Nicely Formation, southeast side of Morgan Mountain, east-central Oregon.
Occurrence: Nicely Formation, Oregon; San Hipólito
Formation, Baja California Sur; Budva Zone, Montenegro;
Gümüslü Allochthon, Turkey; Tawi Sadh Member of the
Guwayza Formation, Oman; Japan.
Plate KAT10. Katroma clara Yeh. Magnification x200, except Fig. 7 x300. Fig. 1(H). Yeh 1987b, pl. 3, fig. 6. Fig. 2. TR,
1662D-R02-01. Fig. 3. Whalen & Carter 2002, pl. 14, fig. 4. Fig. 4. Whalen & Carter 2002, pl. 14, fig. 5. Fig. 5. JP, MNA10, MA11880. Fig. 6. JP, Ku-2-23. Fig. 7. Hori 1988, Fig. 6-1b. Fig. 8. Hori 1990, Fig. 8-21. Fig. 9. OM, BR524-R05-23.
Fig. 10. OM, BR1121-R07-25. Fig. 11. OM, BR477-R19-03.
223
Katroma elongata Carter n. sp.
Species code: KAT17
Synonymy:
1998 Katroma sp. aff. K. irvingi n. sp. – Whalen & Carter, p. 70,
pl. 19, figs. 11, 19.
1998 Katroma coliforme Hori – Yeh & Cheng, p. 28, pl. 7, fig. 6.
2001 Syringocapsa coliformis Hori – Gawlick et al., pl. 5, fig. 10;
pl. 6, fig. 7.
2001 Syringocapsa angusta (Yeh) – Gawlick et al., pl. 5, fig. 11.
2001 Gigi aff. fustis De Wever – Gawlick et al., pl. 5, fig. 12.
does not range above the Sinemurian, but the successor
species, Katroma elongata n. sp., does and is abundant
through most of the Pliensbachian. For this reason, it has
been described as a separate species.
Syringocapsa coliforme Hori (1988) differs from Katroma
elongata n. sp. in having a more massive apical horn and a
narrower abdominal profile.
Type designation: Holotype GSC 111721 from GSC loc.
C- 080612; Ghost Creek Formation (lower Pliensbachian).
Measurements (µm):
Based on 9 specimens.
Description: Cephalis small, dome-shaped, with small apical horn, circular in axial section. Thorax, abdomen and
first abdominal chamber trapezoidal in outline with small
polygonal pore frames. Final post-abdominal chamber
marginally inflated, composed of medium-sized polygonal
pore frames. No significant break between first and final
post-abdominal chambers. Terminal tube narrow, imperforate or with very small pore frames randomly distributed.
Remarks: Whalen & Carter (1998) originally included
this species with Katroma irvingi Whalen and Carter,
a species with widely ranging variation mainly in the size
of the final post abdominal chamber. Recent studies of the
Pliensbachian fauna, however, indicate that K. irvingi s. s.
224
Length (excl. horn)
Maximum width
HT
376
138
Max.
376
138
Min.
277
84
Mean
329
110
Etymology: Latin (adj.) elongatus-a-um = elongate
Type locality: Sample CAA-T-80-7 (GSC loc. C-080612),
Ghost Creek Formation, Rennell Junction, central Graham
Island, Queen Charlotte Islands, British Columbia.
Occurrence: Ghost Creek Formation and Rennell Junction
member of the Fannin Formation, Queen Charlotte Islands;
Dürrnberg Formation, Austria; Liminangcong Chert,
Philippines.
Plate KAT17. Katroma elongata Carter n. sp. Magnification x250. Fig. 1(H). QCI, GSC loc. C-080612, GSC 111721.
Fig. 2. QCI, GSC loc. C-127868, GSC 128814. Fig. 3. Carter et al. 1998, pl. 19, fig. 11.
225
Katroma neagui Pessagno & Poisson 1981, emend. De Wever 1982a
Species code: KAT13
Synonymy:
1981 Katroma neagui n. sp. – Pessagno & Poisson, p. 62, pl. 12,
figs. 1-5; pl. 15, fig. 3.
1982a Katroma neagui Pessagno & Poisson, emend. – De Wever,
p. 193, pl. 3, figs. 5-8.
1982b Katroma neagui Pessagno & Poisson, emend. De Wever
– De Wever, p. 305, pl. 45, figs. 8, 9, 11, 12.
1992 Katroma neagui Pessagno & Poisson – Pessagno &
Mizutani, pl. 99, figs. 7, 12, 16, 17, 20, 21.
Original description: Test as with genus. Meshwork consisting of massive tetragonal to pentagonal pore frames
(predominantly pentagonal); pores becoming larger on
tubular extension of first post-abdominal chamber. Cephalis with crown-like horn with four branches; branching components of horn circular in axial section. Row of
short spines (approximately 12 in number) occurring circumferentially around medial portion of post-abdominal
chamber; spines circular in axial section. Length of tubular
extension on first post-abdominal chamber more than half
of total length of test.
Original remarks: Katroma neagui n. sp., differs from Late
Jurassic and Early Cretaceous species of Podobursa (e.g.,
P. berggreni Pessagno) by having twelve rather than three
circumferentially arranged spines around the medial portion of the final post-abdominal chamber and by having
an open, tubular extension on its final post-abdominal
chamber.
Further remarks: By De Wever (1982a): The morphological characters described by E. A. Pessagno Jr. and
226
A. Poisson remain valid, except for the following. The apical horn has 3 to 5 branches and not always 4. Cephalis
bears at least one very small lateral spine similar to that of
Gigi fustis n. sp. This species has only 3 segments, not 4;
the third one is inflated and continued by a distally closed,
and not open, tube.
The occurrence of cephalic spines is paramount for the
generic assignment. As a matter of fact, E. A. Pessagno
et A. Poisson based the difference between Katroma and
Podobursa on a respectively open and closed abdominal
tube; now, fig. 6 clearly shows a closed tube.
Measurements (µm):
Based on 8 specimens.
Width of abdomen
Length of cephalis-abdomen
Length of first post-abdominal chamber
Width of first post-abdominal chamber
Length of tube on post-abdominal chamber
HT Max. Min.
70
70
50
65
110 115
100
50
120 120
310 310 235
Etymology: This species is named for Dr. Teodor Neagu,
University of Bucharest (Romania) in honor of his contributions to Mesozoic stratigraphy and micropaleontology.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey.
Plate KAT13. Katroma neagui Pessagno & Poisson. Magnification x200x. Fig. 1(H). Pessagno & Poisson 1981,
pl. 12, fig. 1. Fig. 2. TR, 1662D-R01-09. Fig. 3. TR, 1662D-R02-03.
227
Katroma ninstintsi Carter 1988
Species code: KAT14
Synonymy:
1985 Katroma sp. – Igo et al., pl. 15, fig. 14.
1987b Katroma sp. A – Yeh, p. 81, pl. 3, fig.1, pl. 6, figs. 4, 14.
1988 Katroma ninstintsi Carter n. sp. – Carter et al., p. 60, pl. 2,
figs. 4, 9.
1992 Katroma sp. – Pessagno & Mizutani, pl. 99, figs. 6, 10,
11, 15.
1996 Katroma sp. A. – Tumanda et al., p. 181, Fig. 4.15.
1998 Katroma sp. A – Yeh & Cheng, p. 30, pl. 7, figs. 7, 10.
2001 Syringocapsa inflata (Yeh) – Gawlick et al., pl. 5, fig. 9.
Original diagnosis: Tricyrtid test; abdomen expanded and
globose with long, open tubular extension. Apical horn
small and asymmetrical.
Original description: Tricyrtid test: cephalis small and
hemispherical; thorax trapezoidal, expanding more rapidly
in height than width; abdomen enlarged and globose,
terminating in an open tubular extension. Irregularly
shaped pore frames small on initial chambers, becoming
larger on expanded portion of abdomen and decreasing in
size on tubular extension. Apical horn small and more or
less asymmetrical. On some specimens this horn appears
to divide in the manner of Katroma neagui Pessagno and
Poisson (see Pl. 2, fig. 9, this report). Lateral spine (V-spine)
present, but very short and usually broken in type material.
Very small, radial spines (positioned at maximum extension
of abdomen) are more often than not eroded.
Original remarks: A number of forms are tentatively
grouped together as Katroma ninstintsi; they are thought to
represent variants of one species. Collectively they differ in
exhibiting a more or less abrupt transition from expanding
228
thorax to inflated abdomen. This change is sometimes quite
distinct (see Pl. 2, fig. 9) in other cases not (holotype, Pl. 2,
fig. 4).
Katroma ninstintsi normally differs from K. bicornus De
Wever in having only one very small, asymmetrical apical
horn and the radial abdominal spines are much smaller.
It differs from K. neagui Pessagno and Poisson in having
a more expanded abdominal chamber and, whereas the
apical horn on some specimens appears to divide, it is
much shorter and usually has only two, rather than four,
radial branches.
Measurements (µm):
Based on 13 specimens.
Length of apical horn
Length of cephalis and thorax
Length of abdomen
Length of terminal tube
Maximum width of abdomen
HT
15
46
96
106
107
Av.
22.3
85
120
144
155
Max.
61
105
150
170
200
Min.
14
46
92
120
107
Etymology: Named for Haida Indian Chief Ninstints, of
Anthony Island.
Type locality: GSC locality C-080577, Fannin Formation,
Creek locality, Maude Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Fernie Formation, northeastern British
Columbia; Nicely Formation, Oregon; Dürrnberg Formation, Austria; Liminangcong Chert, Philippines; Japan.
Plate KAT14. Katroma ninstintsi Carter. Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 2, fig. 4. Fig. 2. Carter et
al. 1988, pl. 2, fig. 9. Fig. 3. QCI, GSC loc. C-140495, GSC 128820. Fig. 4. NBC, GSC loc. C-305208, GSC 128821.
Fig. 5. NBC, GSC loc. C-305208, GSC 128822. Fig. 6. QCI, GSC loc. C-080612, GSC 128823. Fig. 7. QCI, GSC loc.
C-140495, GSC 128915. Fig. 8. NBC, GSC loc. C-305208, GSC 128916. Fig. 9. OM, Haliw-039-R07-03.
Fig. 10. OM, Haliw-039-R03-09.
229
Katroma? sinetubus Carter n. sp.
Species code: KAT16
Type designation: Holotype GSC 111722 and paratype GSC
111723 from GSC loc. C-080611; Ghost Creek Formation
(lower Pliensbachian).
Description: Test with three or four chambers and a strong
apical and antapical spine. Cephalis small, mostly imperforate with a strong triradiate horn; horn single on holotype
(pl. KAT16, fig. 1) but bifurcating on paratype (pl. KAT16,
fig. 4). Thorax and abdomen trapezoidal in outline, usually with small polygonal pore frames, but pore frames
sometimes larger. Final post-abdominal chamber strongly
inflated composed of large pentagonal and hexagonal pore
frames; pore frames reduced in size distally. Final post-abdominal chamber terminating in a short triradiate antapical spine; spine with sharp narrow ridges and wide, deep
grooves.
Remarks: Genus queried because Katroma, as described by
Pessagno & Poisson (1981), has an inflated final chamber that
terminates in a long open or closed tube. Katroma? sinetubus n. sp. differs from all other described species of Katroma
Katroma sp. 4
Species code: KAT18
Synonymy:
2002 Katroma sp. A – Whalen & Carter, p. 136, pl. 14, fig. 8.
Remarks: This species is very similar to Katroma clara Yeh,
from which it differs in having thorns along the entire apical
horn and in lacking the terminal branching. The horn can
be circular (pl. KAT18, fig. 1) or triradiate in cross section
(pl. KAT18, figs. 2, 3).
Occurrence: San Hipólito Formation, Baja California Sur;
Haliw (Aqil) Formation, Oman.
230
in that the distalmost abdominal chamber terminates in a
strong antapical spine rather than a terminal tube.
Measurements (µm):
Based on 7 specimens.
Length of test
(excl. horn and distal spine)
Maximum width
Length of distal spine
HT
Max.
Min.
Mean
233
233
188
212
171
broken
171
42
141
22
157
36 (4)
Etymology: Latin, sine + tubus = without tube; noun.
Type locality: Sample CAA-79-Ren-Phant, lms 1 (GSC
loc. C-080611), Ghost Creek Formation, Rennell Junction
section, central Graham Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Ghost Creek Formation and Rennell Junction member of the Fannin Formation, Queen Charlotte
Islands.
Plate KAT16. Katroma? sinetubus Carter n. sp. Magnification x200. Fig. 1(H). QCI, GSC loc. C-080611, GSC 111722.
Fig. 2. QCI, GSC loc. C-080611, GSC 111723. Fig. 3. QCI, GSC loc. C-304281, GSC 128824. Fig. 4. QCI, GSC loc. C304281, GSC 128825.
Plate KAT18. Katroma sp. 4. Magnification x200. Fig. 1. BCS, loc. SH-412-14. Fig. 2. OM, Haliw-038-R08-18.
Fig. 3. OM, Haliw-039-R02-10.
231
Genus: Lantus Yeh 1987b
Type species: Lantus sixi Yeh 1987b
Synonymy:
1984b Milax n. gen. – Blome, p.372.
1987b Lantus n. gen. – Yeh, p. 90.
? 1988 Hemicryptocephalis n. gen – Li, p. 329.
Original description: Test multicyrtid, conical, with horn,
usually with strictures at joints, final post-abdominal
chamber closing, with large subspherical, latticed expansion. Cephalis conical or dome-shaped. Cephalis, thorax
and abdomen sparsely perforate, covered by layer of microgranular silica. Post-abdominal chambers consisting
of single layer of dense, small, tetragonal, pentagonal, or
hexagonal pore frames. Pore frames regular to irregular in
shape and size.
Original remarks: Lantus n. gen., differs from Pseudoristola n. gen., by having a test with well-developed horn,
strictures at joints, and smaller, less regular polygonal pore
frames.
Further remarks: Lantus differs from Minocapsa Matsuoka in having a conical rather than ovoid shape and
more postabdominal chambers. It differs from Stichocapsa
Haeckel by lacking an aperture.
Milax Blome is synonymized with Lantus, because
the name Milax is occupied for a gastropod genus (Gray
1855).
Hemicryptocephalis Li is questionably synonymized with
Lantus Yeh, because the description of the former is unclear
for the following reasons: (1) Li mentions that a constricted
aperture is absent but does not indicate whether the
genus is totally closed distally or not, and no illustrations
are provided to show this characteristic; (2) the genus
supposedly has “two descending spines to form the second
post-cephalis cavity” but again this distinguishing feature
is not illustrated and it is certainly not present in the genus
Lantus.
Etymology: Lantus is a name formed by an arbitary
combination of letters (ICZN, 1985, Appendix D, pt. VI,
Recommendation 40, p.201).
Included species:
LAN05 Lantus intermedius Carter n. sp.
LAN01 Lantus obesus (Yeh) 1987b
LAN04 Lantus praeobesus Carter n. sp.
LAN02 Lantus sixi Yeh 1987b
LAN03 Lantus sp. A sensu Whalen & Carter 2002
Lantus intermedius Carter n. sp.
Species code: LAN05
Synonymy:
1988 Hemicryptocephalis dengqensis n. sp. - Li, p. 330, pl. 1,
figs. 5, 6; not fig. 4, ? fig. 10.
1989 Pseudoristola ? spp. – Hattori, pl. 15, fig. D.
1997 Parahsuum sp. NB – Yao, pl. 13, fig. 642.
Type designation: Holotype GSC 111724 from GSC loc. C304566, Rennell Junction member of the Fannin Formation
(upper lower Pliensbachian).
Description: Test conical, usually with four or five postabdominal chambers. Cephalis small, conical, apparently
lacking horn. Cephalis imperforate, thorax and abdomen
sparsely perforate covered with a layer of microgranular
silica. Post-abdominal chambers trapezoidal, variable in
width, increasing more in height than width as added, final
chamber closed with an ellipsoidal cap. Pore frames on post
abdominal chambers polygonal very gradually increasing
in size distally. Slightly raised transverse ridges and/or
slight alignment of pores on distal chambers observed on
some specimens.
Remarks: This species differs from Lantus praeobesus n. sp.
in being less inflated distally and by sometimes possessing
a few very rudimentary circumferential ridges between
232
chambers. L. intermedius differs from L. sixi Yeh in lacking
constrictions between post abdominal chambers. L. intermedius appears to be intermediate between L. praeobesus
which first appears in the earliest Pliensbachian and L. sixi
Yeh which appears later in the late Pliensbachian and Toarcian.
Measurements (µm):
Based on 12 specimens.
Max. length
Max. width
HT
258
126
Max.
340
158
Min.
226
116
Mean
273
140
Etymology: From Latin: intermedius, -a, -um = intermediate; adjective.
Type locality: Sample 99-CNA-MI-9 (GSC loc. C-304566),
Rennell Junction member of the Fannin Formation, Maude
Island, west of Ells Bay, Skidegate Inlet, Queen Charlotte
Islands, British Columbia.
Occurrence: Uppermost Ghost Creek Formation and Fannin Formation, Queen Charlotte Islands; Fernie Formation, NE British Columbia; Haliw (Aqil) Formation, Oman;
China; Japan.
Plate LAN05. Lantus intermedius Carter n. sp. Magnification x200. Fig. 1(H). QCI, GSC loc. C-304566, GSC 111724.
Fig. 2. QCI, GSC loc. C-304567, GSC 128829. Fig. 3. QCI, GSC loc. C-140495, GSC 128830. Fig. 4. QCI, GSC loc. C140495, GSC 128831. Fig. 5. QCI, GSC loc. C-140495, GSC 128832. Fig. 6. NBC, GSC loc. C-305208, GSC 128833.
Fig. 7. OM, Haliw-038-R09-25.
233
Lantus obesus (Yeh) 1987b
Species code: LAN01
Synonymy:
1987b Pseudoristola obesa n. sp. – Yeh, p. 96, pl. 14, figs. 11-12.
2001 Stichocapsa obesa (Yeh) – Gawlick et al., pl. 5, fig. 6.
2003 Stichocapsa convexa Yao – Kashiwagi & Kurimoto, pl. 4,
figs. 1, 2.
1997 Pseudoristola obesa Yeh – Yao, pl. 15, fig. 724.
2005 Sethocapsa sp. – Hori, pl. 8, figs. 29-30, 50.
Original description: Test conical, short, with two postabdominal chambers, without circumferential ridges.
Cephalis small, dome-shaped, without horn. Cephalis,
thorax, and abdomen closely spaced, sparsely perforate,
covered with layer of microgranular silica. Post-abdominal
chamber consisting of single layer of pentagonal and
hexagonal pore frames. Pore frames increasing in size
distally. Bulbous expansion large, subspherical in outline,
comprised of nearly two thirds of whole test.
Original remarks: Pseudoristola obesa differs from other
Pseudoristola spp. in this report by having an extremely
short test with a very large bulbous spherical final postabdominal chamber.
234
Further remarks: Generic name changed to Lantus because
this species lacks the typical parvicingulid arrangement of
pores.
Measurements (µm):
Ten specimens measured.
Length of proximal Width at
Length
conical part (= last
base of
of last
segment excluded) conical part segment
102
120
HT
114
100
128
Mean
111
102
138
Max.
114
Min.
107
96
120
Width
of last
segment
156
162
171
156
Etymology: Obesus-a-um (Latin, adj.) = fat.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Hyde Formation and Warm Springs member
of the Snowshoe Formation, Oregon; Fannin Formation,
Queen Charlotte Islands; Dürrnberg Formation, Austria;
Haliw (Aqil) Formation, Musallah Formation and Tawi
Sadh Member of the Guwayza Formation, Oman.
Plate LAN01. Lantus obesus (Yeh). Magnification x200. Fig. 1(H). Yeh 1987b, pl. 14, fig. 12. Fig. 2. AT, BMW21-52.
Fig. 3. QCI, GSC loc. C-304567, GSC 128826. Fig. 4. QCI, GSC loc. C305417, GSC 128917. Fig. 5. OM, BR1121-R10-12.
Fig. 6. OM, BR1121-R06-18. Fig. 7a, b. OM-00-252-021533, 021534. Fig. 8. OM, BR485-R20-06. Fig. 9. OM, Haliw-038R09-03. Fig. 10. OM, BR1122-R04-02.
235
Lantus praeobesus Carter n. sp.
Species code: LAN04
Synonymy:
1988 Hemicryptocephalis dengqensis n. sp. – Li, p. 330, pl. 1,
figs. ? 4, ? 10; not figs. 5, 6.
1993 Stichocapsa sp. – Kashiwagi & Yao, pl. 1, fig. 5.
1998 Lantus sp. A – Yeh & Cheng, p. 34, pl. 12, fig. 9.
? 2001 Stichocapsa sp. – Kashiwagi, Fig. 6.5.
Type designation: Holotype GSC 111725 from GSC loc. C304566; Rennell Junction member of the Fannin Formation
(upper lower Pliensbachian).
Description: Test broadly conical, usually with four to five
post-abdominal chambers. Cephalis small, conical, usually
without a rudimentary horn. Cephalis imperforate, thorax
and abdomen sparsely perforate covered with a layer of
microgranular silica. Post-abdominal chambers trapezoidal,
rapidly increasing in width as added, final chamber closed
with a large ellipsoidal cap. Pore frames on post-abdominal
chambers polygonal increasing in size distally.
Remarks: This species differs from Lantus obesus (Yeh)
1987 in having a shorter, less inflated final post-abdominal
chamber with smaller pore frames, and in lacking a constriction between the final two chambers. L. praeobesus
n. sp. is the oldest species of Lantus included in this catalogue; it first appears in the basal Pliensbachian and is
abundant throughout the stage.
236
Hemicryptocephalis dengquensis Li is questionably synonymized with L. praeobesus n. sp. because the description of the former is unclear (see remarks under genus
Lantus).
Measurements (µm):
Based on 6 specimens.
Max. length
Max. width
HT
255
145
Max.
289
167
Min.
203
134
Mean
239
152
Etymology: From the Latin, prae (prefix) and obesus (adj.)
= before obesus.
Type locality: Sample 99-CNA-MI-9 (GSC loc. C-304566);
Rennell Junction member of the Fannin Formation, Maude
Island, west of Ells Bay, Skidegate Inlet, Queen Charlotte
Islands, British Columbia.
Occurrence: Ghost Creek Formation and Rennell Junction member of the Fannin Formation, Queen Charlotte
Islands; Fernie Formation, NE British Columbia; Haliw
(Aqil) Formation, Musallah Formation, and Tawi Sadh
Member of the Guwayza Formation, Oman; Japan; Liminangcong Chert, Philippines.
Plate LAN04. Lantus praeobesus Carter n. sp. Magnification Figs. 1-8 x200 (scale bar A), Figs. 9-13 x 300 (scale bar B).
Fig. 1(H). QCI, GSC loc. C-304566, GSC 111725. Fig. 2. QCI, GSC loc. C-304281, GSC 128834. Fig. 3. QCI, GSC loc.
C-304281, GSC 128835. Fig. 4. QCI, GSC loc. C-080612, GSC 128836. Fig. 5. NBC, GSC loc. C-305208, GSC 128837.
Fig. 6. OM, Haliw-038-R08-31. Fig. 7. OM, Haliw-038-R09-12. Fig. 8. OM, Haliw-038-R09-17. Fig. 9. OM-00-252021918. Fig. 10. OM, BR1122-R02-11. Fig. 11. OM, BR1121-R09-20. Fig. 12. OM, BR1121-R08-02.
Fig. 13. OM, BR1121-R06-15.
237
Lantus sixi Yeh 1987b
Species code: LAN02
Synonymy:
1987b Lantus sixi n. sp. – Yeh, p. 90, pl. 4, fig. 16; pl. 17, figs. 9,
13, 17, 24.
1987b Lantus sp. cf. L. sixi n. sp. – Yeh, p. 90, pl. 4, fig. 17; pl. 5,
fig. 18; pl. 17, fig. 14.
1997 Lantus sixi Yeh – Yao, pl. 15, fig. 725.
2004 Lantus sp. – Matsuoka, figs. 96, 97.
Original description: Test small, conical in shape, usually
four to five post-abdominal chambers. Cephalis small,
conical, with short, massive horn. Earlier chambers sparsely
perforate, covered by layer of microgranular silica. Postabdominal chambers having small polygonal pore frames.
Pore frames increasing in size distally, irregular on apical
portion but tending to be aligned in a regular fashion on
final one or two post-abdominal chambers. Final postabdominal chamber closed by large, latticed, subellipsoidal
cap.
Further remarks: Differs from all other species included
here in having very slight strictures between post-abdominal chambers. Apical horn not always distinct.
Lantus sp. A sensu Whalen & Carter 2002
Species code: LAN03
Synonymy:
2002 Lantus sp. A – Whalen & Carter, p. 142, pl. 16, fig. 10.
Original remarks: This species differs from Lantus sixi Yeh
1987 in lacking constrictions and by having larger pore
frames on distal chambers.
Occurrence: San Hipólito Formation, Baja California Sur;
Rennell Junction member of the Fannin Formation, Queen
Charlotte Islands; Tawi Sadh Member of the Guwayza
Formation, Oman.
238
Measurements (µm):
Ten specimens measured.
Length of apical horn
Length of proximal conical part
(= last segment excluded)
Width at base of conical part
Length of last segment
Width of last segment
HT
Mean
13
10
Max. Min.
13
8
117
123
130
115
65
52
95
75
58
102
86
65
108
65
52
95
Etymology: This species is named for Mr. Walter M. Six, Jr.
for his help on this project.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, Oregon;
Fannin member of the Fannin Formation, Queen Charlotte
Islands; Japan.
Plate LAN02. Lantus sixi Yeh. Magnification x300. Fig. 1(H). Yeh 1987b, pl. 17, fig. 9. Fig. 2. QCI, GSC loc. C-304568,
GSC 128827. Fig. 3. QCI, GSC loc. C-175306, GSC 128828.
Plate LAN03. Lantus sp. A sensu Whalen & Carter. Magnification x300. Fig. 1. QCI, GSC loc. C-304568, GSC 128918.
Fig. 2. Whalen & Carter 2002, pl. 16, fig. 10. Fig. 3. OM, BR523-R01-13.
239
Genus: Laxtorum Blome 1984a, emend. Carter 1993
Type species: Laxtorum hindei Blome 1984a
Synonymy:
1984a Laxtorum n. gen. – Blome, p. 56.
1993 Laxtorum Blome emend. – Carter, p. 112.
Original description: Test multicyrtid, consisting of four or
more post-abdominal chambers (segments). Cephalis conical, imperforate, with a large, well-developed horn. Thorax
trapezoidal in outline, perforate, in some specimens buried by microgranular silica. Abdomen and post-abdominal
chambers trapezoidal in outline. Test wall consisting of two
layers: inner layer comprised of triangular to pentagonal
pore frames that lack nodes; outer layer comprised of triangular to hexagonal pore frames with massive, polygonal
nodes at the pore frame vertices, nodes low in relief; pores
of both layers of pore frames large, subcircular to polygonal in outline; pore frames of the outer layer generally restricted to the circumferential ridges, with the exception of
the final post-abdominal chambers. Post-abdominal chambers commonly increasing more rapidly in width than in
height.
Emended definition: Carter (1993): Genus emended to
include the following: (1) test may possess up to twelve
postabdominal chambers, (2) test may have slender, lateral spines that extend radially from medial and/or distal
post-abdominal chambers, and are distributed equally to
subequally around perimeter of test, and (3) test may terminate in an open, flaring terminal tube that lacks septal
partitions.
Original remarks: Laxtorum new genus differs from
Canoptum Pessagno, 1979, by having a test in which
the pores are not buried by an outer layer of accreted
microgranular silica.
Etymology: Laxtorum is a name formed by an arbitrary
combination of letters (ICZN, 1964, p. 113, Appendix D,
Pt. IV, Recommendation 40).
Included species:
LAX06 Laxtorum hemingense Whalen & Carter 1998
Laxtorum hemingense Whalen & Carter 1998
Species code: LAX06
Synonymy:
absence of a prominent horn. It differs from all Rhaetian
species of Laxtorum in having larger, open, more regularly
aligned pore frames and in lacking a terminal tube.
Original description: Test conical with approximately eight
to nine postabdominal chambers. Cephalis small, steeply
dome-shaped with or without a small horn; horn circular
in axial section; cephalis imperforate, covered with a thick
layer of microgranular silica. Thorax, abdomen and all
postabdominal chambers gradually increasing in width
as added. Thorax trapezoidal in outline with mediumsized pores mostly obscured by layer of microgranular
silica. Mostly tetragonal pore frames on abdomen and
postabdominal chambers subaligned in transverse rows.
Circumferential ridges composed of thicker, more irregular
pore frame bars (zig-zag structure) on proximal parts of test
becoming thinner and straighter distally; circumferential
ridges with irregular longitudinal extensions superimposed
on adjacent pore frames.
Measurements (µm):
Based on 5 specimens.
1998 Laxtorum hemingense n. sp. – Whalen & Carter, p. 80,
pl. 25, figs. 6-8, 13-14, 24-25; pl. 27, figs. 5, 6, 16, 20.
Original remarks: Laxtorum hemingense n. sp. is distinguished from Laxtorum sp. A and Laxtorum sp. B by the
240
Length (excluding horn)
240
240
218
229
Max. width
105
109
98
104
HT
Max.
Min.
Mean
Etymology: This species is named for Heming Head, located on the east side of Talunkwan Island, northwest of
the type locality.
Type locality: Sample QC-549, Sandilands Formation,
Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands.
Plate LAX06. Laxtorum hemingense Whalen & Carter. Magnification x250. Fig. 1(H). Carter et al. 1998, pl. 25, fig. 6.
Fig. 2. QCI, GSC loc. C-080612, GSC 128919. Fig. 3. QCI, GSC loc. C-175311, GSC 128920.
241
Genus: Minocapsa Matsuoka 1991
Type species: Minocapsa cylindrica Matsuoka 1991
Synonymy:
1991 Minocapsa n. gen. – Matsuoka, p. 734.
Original description: Closed cyrtid. Shell consisting of
four or more segments, pyriform to ovoidal. Cephalis hemispherical without apical horn. Thorax and abdomen truncate conical. Final segment large, hemispherical without
aperture. Pores, circular to subcircular and closely spaced.
Original remarks: Minocapsa, n. gen. is distinguished from
Stichocapsa Haeckel by lacking aperture. It also differs
from Zhamoidellum Dumitrica and Cryptamphorella
Dumitrica by not being cryptothoracic and by consisting
of four or more segments rather than three.
Further remarks: Minocapsa differs from Lantus Yeh by
having fewer segments and the height of the last segment
is greater than the remainder of the test. Both genera lack
an aperture.
Etymology: The generic name is named for the Mino Terrane which includes the type area, Nanjo Massif.
Included species:
MCP01 Minocapsa cylindrica Matsuoka 1991
MCP02 Minocapsa globosa Matsuoka 1991
TPS02 Minocapsa? megaglobosa (Matsuoka) 1991
Minocapsa cylindrica Matsuoka 1991
Species code: MCP01
Synonymy:
1987 Bagotum sp. E – Hattori, pl. 15, fig. 4.
1989 Bagotum sp. aff. B. modestum Pessagno & Whalen
– Hattori & Sakamoto, pl. 13, fig. K.
1990 Stichocapsa (?) sp. – Nagai, pl. 4, fig. 6.
1991 Minocapsa cylindrica n. sp. – Matsuoka, p. 735, figs. 10.1a-5b.
1997 Minocapsa cylindrica Matsuoka – Yao, pl. 10, fig. 450.
? 2003 Stichocapsa sp. B – Kashiwagi & Kurimoto, pl. 4, fig. 4.
2004 Minocapsa cylindrica Matsuoka – Hori, pl. 2, fig. 14.
2004 Minocapsa cylindrica Matsuoka – Matsuoka, fig. 91.
Original description: Shell of five segments, ovoidal.
Cephalis spherical; thorax and abdomen truncate conical.
The proximal three segments form a conical proximal part.
The distal two segments form a cylindrical to ovoidal distal
part. The last segment hemispherical without aperture.
Segmental joints generally indistinct externally; joint
between abdomen and fourth segment faintly marked
by change in contour from the conical proximal part
to the cylindrical to ovoidal distal part. Pores circular to
subcircular, rather densely spaced and set in tetragonal to
hexagonal pore frames. Pores on the distal part tend to be
arranged longitudinally. Pores around distal end slightly
larger than those on the rest of the shell; these pores
242
surrounded by circular rims. Small nodes or spines situated
at the pore frame vertices.
Original remarks: This species differs from Minocapsa
globosa, n. sp. by consisting of five segments rather than four,
by possessing a more slender distal part and by the pores of
the distal part tending to be longitudinally arranged.
Measurements (µm):
Six specimens measured.
Total height of shell
Max. width of shell
HT
193
120
Max.
193
135
Min.
165
115
Mean
181
125
Etymology: The specific name comes from the Latin
cylindricus-a-um (=cylindrical).
Type locality: MNA-10, Nanjo Massif, Mino Terrane,
central Japan.
Occurrence: Mino Terrane, Japan; Musallah Formation,
Oman; Skrile Formation, Slovenia; Hyde Formation,
Oregon.
Plate MCP01. Minocapsa cylindrica Matsuoka. Magnification x400. Fig. 1(H). Matsuoka 1991, Fig. 10-1a.
Fig. 2. SI, MM 5.00-010122. Fig. 3. OM-00-252-021927.
243
Minocapsa globosa Matsuoka 1991
Species code: MCP02
Synonymy:
1991 Minocapsa globosa n. sp. – Matsuoka, p. 736, figs. 11.1a-4b.
1997 Minocapsa globosa Matsuoka – Yao, pl. 10, fig. 451.
2004 Minocapsa globosa Matsuoka – Matsuoka, fig. 90.
Original description: Shell of four segments, pyriform.
Cephalis spherical; thorax and abdomen truncate conical.
The proximal three segments form a conical proximal part.
The fourth segment large, subspherical without aperture.
Collar and lumber strictures indistinct externally. Joint
between abdomen and fourth segment marked by rapid
change in contour from the conical proximal part to
spherical distal part. Pores circular to subcircular, densely
spaced and set in polygonal (largely hexagonal) pore
frames. Pores around distal end slightly larger than those
on the rest of shell and surrounded by rims. Small nodes
situated at the pore frame vertices in some specimens.
Original remarks: This species is compared to Minocapsa
cylindrica, n. sp. under the latter species.
Measurements (µm):
Six specimens measured.
Total height of shell
Max. width of shell
HT
173
140
Max.
185
149
Min.
170
135
Mean
177
142
Etymology: The specific name is derived from the Latin
globosus-a-um (=spherical).
Type locality: MNA-10, Nanjo Massif, Mino Terrane, central Japan.
Occurrence: Mino Terrane, Japan.
Minocapsa? megaglobosa (Matsuoka) 1991
Species code: TPS02
Synonymy:
1991 Tricolocapsa (?) megaglobosa n. sp. – Matsuoka, p. 724,
figs. 3. 1a-5b.
1997 Tricolocapsa megaglobosa Matsuoka – Yao, pl. 9, fig. 423.
2003 Tricolocapsa ? megaglobosa Matsuoka – Goričan et al.,
p. 297, pl. 4, fig. 12.
2004 Tricolocapsa (?) megaglobosa Matsuoka – Matsuoka,
fig. 82.
Original description: Shell of three segments with a dishlike basal appendage. Cephalis hemispherical, poreless.
Thorax truncate-conical with circular, densely spaced
pores. Abdomen large, inflated, barrel-shaped with circular,
densely spaced pores larger than those in thorax. Collar
stricture distinct externally. Lumber stricture slightly
recognizable externally. Joint between abdomen and basal
appendage marked by a row of pores slightly larger than
those in abdomen. Basal appendage half to a third of
abdomen in width, with circular to subcircular densely
spaced pores smaller than those in remaining part of shell
surface.
segments, and the presence of an inflated segment.
However, according to the generic diagnosis of Minocapsa,
the last segment is large whereas in this species the last
segment is very small and the third segment is the largest.
Regardless, even if assignation to Minocapsa is questionable,
the assignation of this species to Tricolocapsa Haeckel is
completely wrong because this Cenozoic genus has three
segments and a cephalic tube. It is possible that Minocapsa?
megaglobosa has amphipyndacid affinities because the
paratype illustrated in transmitted light by Matsuoka (1991,
fig. 3.5) seems to show a two-segmented cephalis. The
globular, imperforate cephalis of this species also suggests
amphipyndacid affinities.
Measurements (µm):
Numbers of specimens measured are in parentheses.
Total height of shell
Maximum width of shell
Width of basal appendage
HT
153
105
55
Max. Min. Mean
160 120
145
88
101
117
61
47
55
(12)
(12)
(11)
Original remarks: This species is questionably assigned
to Tricolocapsa, because it possesses a dish-like basal
appendage. This species is distinguished from Tricolocapsa
(?) fusiformis Yao by having a large, inflated abdomen,
densely spaced pores and a distinct collar stricture.
Etymology: This specific name is derived from the Latin
mega (=large) and globosus-a-um (=spherical).
Further remarks: Tricolocapsa? megaglobosa is herein
tentatively assigned to the genus Minocapsa Matsuoka
because, in common with this genus, it has the following
elements: the absence of an aperture, more than three
Occurrence: Mino Terrane, Japan; Skrile Formation, Slovenia; Tawi Sadh Member of the Guwayza Formation and
Musallah Formation, Oman.
244
Type locality: Sample MNA-10, Nanjo Massif, Mino
Terrane, central Japan.
Plate MCP02. Minocapsa globosa Matsuoka. Magnification x300. Fig. 1(H). Matsuoka 1991, Fig. 11-1a.
Fig. 2. Matsuoka 1991, Fig. 11-2. Fig. 3. Matsuoka 1991, Fig. 11-3.
Plate TPS02. Minocapsa? megaglobosa (Matsuoka). Magnification x400. Fig. 1(H). Matsuoka 1991, Fig. 3.1a.
Fig. 2. OM, BR485-R21-15. Fig. 3. OM-00-251-021611. Fig. 4. OM-00-117-021213.
245
Genus: Napora Pessagno 1977a
Type species: Napora bukryi Pessagno 1977a
Synonymy:
1977a Napora n. gen – Pessagno, p. 94.
1977b Ultranapora n. gen. – Pessagno, p. 38.
1982a Jacus n. gen. – De Wever, p. 204.
1986 Napora Pessagno emend. – Pessagno et al., p. 34.
1986 Napora Pessagno emend. – Takemura, p. 43.
2003 Napora Pessagno – Dumitrica & Zügel, p. 57.
Original description: Test dicyrtid with a large conical cephalis and a large subglobular thorax. Cephalis with massive horn bearing longitudinal ridges and grooves and often
having subsidiary spines. Thorax with coarse, equal size,
polygonal (usually hexagonal) pore frames and circular
pores and with a large circular aperture (mouth) at base;
three slightly curved feet with longitudinally developed
ridges and grooves occurring at base of thorax.
Original remarks: Napora n. gen., differs from Tripilidium
in possessing a dicyrtid test with a well-developed apical
horn.
Further remarks: By Dumitrica & Zügel (2003): The study
of the initial spicule of the species Napora modesta n. sp. in
transmitted light showed that the cephalic initial skeleton
consists of MB, V, A, two L, two l, D, and the arches VL,
Ll, lD, Al. The arch AV is absent, but an arch AD seems to
exist because one of the three blades of the apical horn has
a dorsal direction. Genus Jacus De Wever, 1982 is herein
considered a junior synonym of Napora. The differences
between these two genera concern only the superficial
structure of thorax, a character that can be considered of
specific level. The absence of the cephalocone, considered
by De Wever (1982) an additional distinctive character
from Napora, has no value because in both genera it is
nothing else than the short ventral spine. Moreover, both
genera have the apical horn with a verticil of three spines
or spinules as extensions of the three blades; exceptions are
very rare. This is also a character that differentiates Napora
and Jacus from Anaticapitula n. gen.
Etymology: Napora is an anagram for C. F. Parona, one of
the early students of Jurassic Radiolaria.
Included species:
NAP09 Napora blechschmidti Dumitrica n. sp.
NAP08 Napora bona Pessagno, Whalen & Yeh 1986
NAP02 Napora cerromesaensis Pessagno, Whalen & Yeh
1986
NAP06 Napora conothorax Carter & Dumitrica n. sp.
NAP01 Napora graybayensis Pessagno, Whalen & Yeh 1986
3410 Napora nipponica Takemura 1986
NAP03 Napora reiferensis (Pessagno, Whalen & Yeh) 1986
NAP04 Napora relica Yeh 1987b
JAC01 Napora sandspitensis (Pessagno, Whalen & Yeh)
1986
Napora blechschmidti Dumitrica n. sp.
Species code: NAP09
Type designation: Holotype pl. NAP09, fig. 3, sample
BR871-R03-08, all paratypes also from BR871, chert of
Tawi Sadh Member reworked in the Guwayza Formation,
Al Khashbah Mountains, Oman.
Description: Test high pyramidal with a thick apical horn.
Apical horn long, multi-bladed proximally, three-bladed
distally, pointed, with a crown of three small thorns between
the two portions. Blades of proximal part cover the cephalis
that is practically invisible outside. Cephalis indistinct
externally, screened by the blades of the apical horn. Thorax
pyramidal with convex sides and 2-4 transversal ribs. One
rib may be higher making a shoulder on the thorax. Pore
frames mostly quadrangular, usually aranged in transverse
rows between ribs. One row between each intercostal
area. Rows of pores toward cephalis may be more or less
disturbed. Feet straight to slightly curved, gently pointed,
three-bladed, slightly divergent; external blades rather high
along thorax where they make the edges of the pyramid.
Remarks: Napora blechschmidti resembles N. bona Pessagno, Whalen & Yeh and N. hasta Yeh & Cheng but differs
especially by having a many-bladed apical horn.
246
Measurements (µm):
Based on 4 specimens.
Total height of shell with horn and feet
Height of apical horn and cephalis
Height of thorax
Breadth of thorax
Length of feet
Min.
230
100
43
100
73
Max.
260
140
60
115
97
Etymology: The species is named for Ingo Blechschmidt as
a sign of friendship and to honour his contribution to the
geology of the Hamrat Duru Basin, Oman.
Type locality: Sample BR871, chert of Tawi Sadh Member
reworked in the Guwayza Formation, Al Khashbah Mountains, Oman.
Occurrence: Upper part of the Tawi Sadh Member of the
Guwayza Formation, Oman.
Plate NAP09. Napora blechschmidti Dumitrica n. sp. Magnification x250. Fig. 1. OM, BR871-R08-11.
Fig. 2. OM, BR871-R07-19. Fig. 3(H). OM; BR871-R06-10. Fig. 4. OM, BR871-R06-10. Fig. 5. OM, BR871-R08-14.
247
Napora bona Pessagno, Whalen & Yeh 1986
Species code: NAP08
Synonymy:
1986 Napora bona n. sp. – Pessagno et al., p. 36, pl. 6, figs. 4, 5.
1986 Napora sp. A – Pessagno et al., p. 46, pl. 7, fig. 14.
1988 Napora sp. aff. N. cosmica Pessagno, Whalen & Yeh
– Carter et al., p. 58, pl. 14, fig. 2.
1989 Napora triangularis Takemura – Hattori, pl. 20, fig. C.
1991 Napora bona Pessagno, Whalen & Yeh – Carter & Jakobs,
p. 343, pl. 3, fig. 4.
2004 Napora bona Pessagno, Whalen & Yeh – Matsuoka,
fig. 138.
Original diagnosis: Cephalis large, hemispherical, with well
developed large cephalocone and a medium-length horn.
Horn triradiate in axial section with three medium-width
longitudinal ridges alternating with three somewhat wider
longitudinal grooves; ridges wider on proximal two thirds
of horn, flaring outwards to give rise to three short spines.
Thorax pyramidal with linear, circumferentially-arranged
tetragonal pore frames; faces of distal one or two rows of
pore frames sloping inwards. Feet incurved, triradiate in
axial section with three sharply-bladed longitudinal ridges
alternating with three wide longitudinal grooves; ridges
high in relief.
Original remarks: Napora bona appears closely related to
Napora sp. A (Pl. 7, Fig. 14). It differs from the latter form
by possessing larger, less numerous, and more irregularly
shaped thoracic pore frames. Both forms display a row of
inwardly-sloping pore frames at the base of the thorax.
N. bona differs from N. cosmica, n. sp., by having a larger
cephalis with a well-developed cephalocone; shorter, less
curved feet; larger, less numerous, more irregular thoracic
pore frames; a shorter, more massive horn; and an inturned
row of pore frames at the base of the thorax.
Measurements (µm):
Number of specimens measured are in parentheses.
Length of cephalis
Length of thorax
Width of thorax at top
Width of thorax at base
Length of horn
Width of horn at base
Length of foot (maximum)
HT
20
37.5
37.5
70
32.5
20
62.5
Mean
Max.
Min.
22.5 (7) 25 (7) 12.5 (7)
32.8 (7) 37.5 (7) 25 (7)
36.4 (7) 37.5 (7) 30 (7)
77.1 (7) 95 (7) 62.5 (7)
45.6 (4) 50 (4) 32.5 (4)
21.4 (7) 25 (7) 17.5(7)
67.5 (4) 82.5 (4) 50 (4)
Etymology: (Latin) bonus = good, useful.
Type locality: Sample OR-580, Warm Springs member,
Snowshoe Formation, near bridge over South Fork of John
Day River, east-central Oregon.
Occurrence: Warm Springs member of the Snowshoe Formation, Oregon; Phantom Creek Formation, Queen Charlotte Islands; Japan.
Napora cerromesaensis Pessagno, Whalen & Yeh 1986
Species code: NAP02
Synonymy:
1986 Napora cerromesaensis n. sp. – Pessagno, Whalen & Yeh,
p. 38, pl. 4, figs. 2-4, 10, 15, 16.
2002 Napora cerromesaensis Pessagno, Whalen & Yeh – Whalen
& Carter, p. 140, pl. 15, fig. 10.
2004 Napora sp. cf. N. cerromesaensis Pessagno, Whalen & Yeh
– Ziabrev et al., Fig. 5-8.
Original diagnosis: Cephalis relatively small, hemispherical, with massive apical horn; cephalis may be partially
obscured by a thin layer of microgranular silica. Horn approximately same length as test, triradiate in axial section
with narrow, rounded, longitudinal ridges alternating with
broad, deep grooves; ridges extended into broad spines
midway from base to terminus of horn; narrow ridges of
apical horn may extend down over cephalis, although not
as pronounced as on horn; horn tapers distally. Thorax subpyramidal in shape with large, irregularly shaped tetragonal and pentagonal pore frames arranged in poorly-defined
transverse rows. Massive triradiate feet attached to base of
thorax, curved slightly inward. Mouth subtriangular in
outline, surrounded by imperforate rim.
Original remarks: Napora cerromesaensis, n. sp., is distinguished from Napora (?) graybayensis, n. sp., by the more
248
regular arrangement of the pore frames and the thin layer
of microgranular silica on the cephalis. In addition, N. cerromesaensis possesses a more massive horn and feet that are
not as curved as those of N.(?) graybayensis, n. sp.
Further remarks: This species differs from Napora bona
Pessagno, Whalen & Yeh in that the feet are more massive
and less outwardly directed.
Measurements (µm):
Numbers of specimens measured are in parentheses.
Length of cephalis
Length of thorax
Width of thorax at top
Width of thorax at base
Length of horn
Width of horn at base
Length of foot (maximum)
HT
20
80
40
90
70
20
80
Mean
16.8 (11)
70.5 (10)
38.1 (11)
99.5 (11)
86.3 (11)
22.7 (11)
95.9 (11)
Max.
20 (11)
80 (10)
45 (11)
110 (11)
100 (11)
25 (11)
120 (11)
Min.
15 (11)
60 (10)
30 (11)
90 (11)
70 (11)
20 (11)
70 (11)
Etymology: The species is named for Cerro Mesa, located
to the northeast of its type locality.
Type locality: Sample BPW-30, San Hipólito Formation,
Vizcaino Peninsula, Baja California Sur, Mexico.
Plate NAP08. Napora bona Pessagno, Whalen & Yeh. Magnification x250. Fig. 1(H). Pessagno, Whalen & Yeh 1986,
pl. 6, fig. 4. Fig. 2. JP, MNA-10, MA12702. Fig. 3. Carter & Jakobs 1991, pl. 3, fig. 4.
Plate NAP02. Napora cerromesaensis Pessagno, Whalen & Yeh. Magnification x250. Fig. 1(H). Pessagno, Whalen &
Yeh 1986, pl. 4, fig. 2. Fig. 2. Pessagno, Whalen & Yeh 1986, pl. 4, fig. 3. Fig. 3. QCI, GSC loc. C-304567, GSC 128838.
Fig. 4. QCI, GSC loc. C-304566, GSC 128839. Fig. 5. QCI, GSC loc. C-304567, GSC 128840. Fig. 6. QCI, GSC loc. C304567, GSC 128841. Fig. 7. NBC, GSC loc. C-305813, GSC 111726.
249
Occurrence: San Hipólito Formation, Baja California Sur;
Nicely Formation, Oregon; Rennell Junction member of
the Fannin Formation, Queen Charlotte Islands; Fernie
Formation, NE British Columbia; Bainang Terrane, Tibet.
Napora conothorax Carter & Dumitrica n. sp.
Species code: NAP06
Type designation: Holotype pl. NAP06, fig. 1; paratype 1,
fig. 2, both from Fernie Formation, Williston Lake, British
Columbia; paratype 2, fig. 3, Haliw Formation, Humadiyin,
Oman.
Description: Cephalis indistinct externally and imperforate,
included at the upper part of thorax. Apical horn threebladed, relatively thin and short, bearing a verticil of three
spinules and terminating in a short conical spine. Vertical
spine not visible outside. Thorax and cephalis forming
a wide, short cone with longitudinal ridges on the upper
part, especially on cephalis, and 3-4 prominent transversal
ridges. Ridges interconnected by vertical crests forming
rectangular depressions. Pores in single transverse rows
between ridges. Feet strongly triradiate and recurved, long,
and pointed, with thin ridges and deep grooves. Outer
ridge of each foot extends outward from the area of the
raised ridge.
Remarks: The apical horn and the feet of this species resemble those of Napora latissima Takemura with differing
only in that the feet are less divergent and less curved. The
paratype from Oman is rather similar to the holotype and
paratype from British Columbia but has less pronounced
transversal ribs and longitudinal ribs are more visible. Although partly broken, the only preserved foot of this specimen seems to show characters similar to the feet of the
holotype and paratype.
Measurements (µm):
Based on 3 specimens.
HT Paratype 1 Paratype 2
Length of apical horn
37
47
Height of cephalothorax 83
87
90
Diameter of thorax
100
103
100
Length of feet
157
-
Etymology: From the conical shape of the thorax; noun.
Type locality: GSC loc. C-305208, Fernie Formation,
Williston Lake, northeastern British Columbia.
Occurrence: Fernie Formation, Williston Lake, northeastern British Columbia; Haliw (Aqil) Formation, Oman.
Napora graybayensis Pessagno, Whalen & Yeh 1986
Species code: NAP01
Synonymy:
1986 Napora (?) graybayensis n. sp. – Pessagno, Whalen & Yeh,
p. 39, pl. 2, figs. 1-3, 10, 11, 14; pl. 11, figs. 3, 4.
1998 Napora? graybayensis Pessagno, Whalen & Yeh – Whalen &
Carter, p. 75, pl. 21, fig. 4.
Original diagnosis: Cephalis relatively small, hemispherical, with massive horn; pore frames of cephalis commonly
not covered by layer of microgranular silica. Horn triradiate in axial section throughout most of its length, with
three narrow, rounded, longitudinal ridges alternating
with three broad, shallow grooves; horn tapering sharply
distally; length of horn one-half to equal to length of test;
small spines commonly located on ridges approximately at
midpoint of horn. Thorax subpyramidal in shape with medium- to very large-sized, irregularly-shaped, tetragonal to
pentagonal pore frames; pore frames arranged in poorly-defined transverse rows with slight development of transverse
ridges separating the rows. Feet of moderate length, curved
inward, triradiate in cross-section, consisting of three very
narrow, longitudinal ridges alternating with broad shallow
grooves. Mouth triangular in outline, bounded by narrow,
imperforate rim.
Original remarks: This species of Napora is distinguished
from others by the irregular shape and distribution of
250
the pore frames. It is questionably assigned to the genus
Napora because of its peculiar, irregular pore frames, poor
definition externally between the cephalis and thorax, and
the lack of a microgranular layer proximally.
Measurements (µm):
Numbers of specimens measured are in parentheses.
HT
Length of cephalis
10
Length of thorax
50
30
Width of thorax at top
50
Width of thorax at base
Length of horn
50
12.5
Width of horn at base
Length of foot (maximum) 30
Mean
13.3 (9)
56.6 (9)
36.6 (9)
69.4 (9)
42.2 (7)
14.1 (9)
65 (8)
Max.
20 (9)
75 (9)
45 (9)
80 (9)
60 (7)
20 (9)
80 (8)
Min.
10 (9)
45 (9)
30 (9)
50 (9)
30 (7)
10 (9)
45 (8)
Etymology: This species is named for Gray Bay, which is
located north of its type locality.
Type locality: Sample QC-675, Sandilands Formation
(Kunga Formation of Pessagno, Whalen & Yeh, 1986)
Kunga Island - north side, Queen Charlotte Islands, British
Columbia, Canada.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands.
Plate NAP06. Napora conothorax Carter & Dumitrica n. sp. Magnification x300. Fig. 1. NBC, GSC loc. C-305208,
GSC 128843. Fig. 2. NBC, GSC loc. C-305208, GSC 128844. Fig. 3. OM, Haliw-038-R08-10.
Plate NAP01. Napora graybayensis Pessagno, Whalen & Yeh. Magnification x400. Fig. 1(H). Pessagno, Whalen & Yeh
1986, pl. 2, fig. 1. Fig. 2. Pessagno, Whalen & Yeh 1986, pl. 2, fig. 2. Fig. 3. Pessagno, Whalen & Yeh 1986, pl. 2, fig. 3.
Fig. 4. QCI, GSC loc. C-175311, GSC 128842.
251
Napora nipponica Takemura 1986
Species code: 3410
Synonymy:
1986 Napora nipponica n. sp. – Takemura, p. 44, pl. 2, figs. 16-21.
1989 Napora nipponica Takemura – Hattori & Sakamoto, pl. 2,
fig. M, not fig. L.
1991 Napora nipponica Takemura – Carter & Jakobs, p. 343,
pl. 3, fig. 1.
1993 Napora nipponica Takemura – Pessagno et al., p. 158,
pl. 8, fig. 10.
1995a Napora nipponica Takemura – Baumgartner et al., p. 330,
pl. 3410, figs. 1-2.
1997 Napora nipponica Takemura – Yao, pl. 8, fig. 372.
cal horn and feet, and a transverse arrangement of thoracic
pores. N. nipponica is also distinguishable from other species of Napora by its shape of apical horn and three feet.
Further remarks: Differs from Napora bona Pessagno,
Whalen & Yeh by having smaller, more numerous pores on
thorax and the feet are more incurved.
Measurements (µm):
Based on 15 specimens.
Length of the shell including horn and feet
Height of cephalo-thorax
Maximum width of shell including feet
Width of thorax
Min.
200
60
115
75
Max.
270
85
160
110
Original description: Cephalis small and subspherical with
straight and triradiate apical horn, and with or without
cephalocone. Ridges of apical horn may or may not originate
at the base of the cephalis. A node located at the position
of about half way along each ridge. Thorax subspherical
or hemispherical to trigonally pyramidal, with usually
transversely arranged circular pores. Three feet triradiate
and curved convexly. Aperture subtriangular to circular
with remarkable circular or subtriangular apertural ring
around it.
Etymology: The trivial name is derived from Nippon, Japan
in Japanese.
Original remarks: Although Napora nipponica n. sp. resembles in its shape to N. bukryi Pessagno, N. nipponica differs from N. bukryi in possessing a considerably long api-
Occurrence: Mino Terrane, Japan; Phantom Creek Formation, Queen Charlotte Islands; Josephine Ophiolite, California.
Type locality: Maganese carbonate ore deposit, sample
TKN-105. Gujo-Hachiman area, Mino Terrane, central
Japan.
Napora reiferensis (Pessagno, Whalen & Yeh) 1986
Species code: NAP03
Synonymy:
Original remarks: Jacus reiferensis, n. sp., is distinguished
from Jacus coronatus De Wever, 1982, by the nature of
the meshwork on the thorax and velum(?) as well as the
structure of the horn.
Original diagnosis: Cephalis medium-sized, hemispherical,
commonly covered with a thin layer of microgranular
silica; small spine may extend from base of cephalis. Apical
horn massive, more than one-half length of test, distally
trifurcating. Proximal two-thirds of horn triradiate in axial
section with narrow, rounded ridges and broad grooves;
distal portion of horn separated into three tapering lobes;
lobes elliptical in cross-section, approximately at right
angles to long axis of test and in some specimens curving
slightly downward; lobes of horn ranging from onehalf to equal the length of the triradiate portion of horn.
Thorax subpyramidal in outline with small- to mediumsized, slightly nodose, irregularly shaped, elliptical and
tetragonal pore frames; pore frames arranged in poorlydefined transverse rows separated by transverse ridges.
Massive triradiate feet, attached to base of thorax, curved
slightly inward. Subsidiary meshwork (velum?), with
very irregularly-shaped and –spaced pore frames and
imperforate rim, some attached to base of thorax. Mouth
subtriangular in outline.
Measurements (µm):
Numbers of specimens measured are in parentheses.
1986 Jacus reiferensis n. sp. – Pessagno, Whalen & Yeh, p. 32,
pl. 4, figs. 7-8, 11; pl. 5, figs. 6, 14.
2002 Napora reiferensis (Pessagno, Whalen & Yeh) – Whalen &
Carter, p. 140, pl. 15, figs. 7, 11.
252
HT
Mean
Max.
Length of cephalis
20 20.9 (11) 25 (11)
Length of thorax
50 63.1 (11) 80 (11)
Width of thorax at top
45 43.1 (11) 50 (11)
Width of thorax at base
110 100 (11) 120 (11)
Length of horn
65 63.6 (11) 85 (11)
Width of horn at base
25 20.4 (11) 25 (11)
Length of foot (maximum) 100 93.5 (10) 120 (10)
Min.
15 (11)
50 (11)
40 (11)
90 (11)
55 (11)
15 (11)
70 (10)
Etymology: This species is named for Pico Reifer, which is
located east of its type area.
Type locality: Sample BPW-30, San Hipólito Formation,
Vizcaino Peninsula, Baja California Sur, Mexico.
Occurrence: San Hipólito Formation, Baja California Sur.
Plate 3410. Napora nipponica Takemura. Magnification Fig. 1(H) x300, Fig. 2. x200. Fig. 1(H). Takemura 1986, pl. 2,
fig. 20. Fig. 2. Carter & Jakobs 1991, pl. 3, fig. 1.
Plate NAP03. Napora reiferensis (Pessagno, Whalen & Yeh). Magnification x 250. Fig. 1(H). Pessagno, Whalen & Yeh
1986, pl. 4, fig. 7. Fig. 2. Pessagno, Whalen & Yeh 1986, pl. 5, fig. 6.
253
Napora relica Yeh 1987b
Species code: NAP04
Synonymy:
1986 Jacus (?) species B – Pessagno et al., p. 34, Pl. 5, Fig. 9.
1987b Napora relica n. sp. – Yeh, p. 85, Pl. 10, Figs. 4, 17.
1987b Napora sp. aff N. relica Yeh – Yeh, p. 85, Pl. 24, Figs. 8, 21, 23.
1989 Napora sp. A – Hattori, pl. 5, fig. L.
1989 Napora sp. B – Hattori, pl. 5, fig. M.
2002 Napora sp. B of Hattori – Hori & Wakita, pl. 3, fig. 5.
2003 Napora relica Yeh – Goričan et al., p. 296, pl. 4, fig. 4.
2004 Napora relica Yeh – Matsuoka, fig. 139.
Original description: Cephalis relatively large, hemispherical, imperforate, covered with layer of microgranular
silica. Horn moderately long, triradiate with three narrow
ridges alternating with three narrow grooves; each ridge
terminating in long pointed spine. Axial part of horn also
terminating in short spine and surrounded by long curved
spines extending out from ridges. Thorax subhemispherical in outline, with mixture of tetragonal, pentagonal, and
hexagonal pore frames arranged in five to six transverse
rows; larger pore frames on middle portion of thorax. Aperture large, subcircular in outline. Feet long, tapering distally, triradiate with three narrow ridges alternating with
three grooves, grooves wider proximally and narrow distally.
Further remarks: This species is easily recognized by its
distinctive horn with three, long, curved pointed spines.
Measurements (µm):
Ten specimens measured.
Length of cephalis
Length of thorax
Width of thorax at top
Width of thorax at base
Length of horn
Length of foot (maximum)
HT
25
63
30
94
63
113
Mean
25
61
30
95
62
115
Max.
27
63
32
97
65
118
Min.
23
59
29
93
61
110
Etymology: Relicus-a-um (Latin, adj.) = outstanding.
Type locality: Sample OR-600M, Hyde Formation at IzeePaulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, Oregon;
Skrile Formation, Slovenia; Tawi Sadh Member of the Guwayza Formation, Oman; Japan.
Napora sandspitensis (Pessagno, Whalen & Yeh) 1986
Species code: JAC01
Synonymy:
1986 Jacus (?) sandspitensis n. sp. – Pessagno, Whalen & Yeh,
p. 33, pl. 2, figs. 5, 9, 13, 16, 17; pl. 11, fig. 13.
1998 Jacus(?) sandspitensis Pessagno, Whalen & Yeh – Whalen
& Carter, p. 75.
Original diagnosis: Cephalis small, hemispherical, with
small cephalocone and relatively short horn. Proximal
two-thirds of horn triradiate in axial section with three
narrow grooves alternating with three rounded ridges
of approximately the same width. Grooves often deeplyincised proximally, becoming shallower distally; ridges each
bearing short spines before their termination. Distal onethird of horn circular in axial section; lacking ridges and
grooves. Thorax with very coarse, nodose polygonal pore
frames; distal portion of thorax with mixture of tetragonal
and pentagonal pore frames arranged in rows between
angled ridges. Feet long, triradiate in axial section with
three narrow ridges that alternate with three wide grooves;
short spines occur along ridges on proximal half of each
foot, suggesting velum attachement. Mouth subcircular in
outline, surrounded by an imperforate rim.
Original remarks: This species is questionably assigned to
Jacus because it possesses a true cephalocone. It is conceivable that the spine extending from the »hole« (cephalopyle?)
of De Wever’s (1982) specimens of Jacus merely represents
a remnant of a fragile cephalocone not preserved in his
254
badly-etched material. The presence of a velum in J. (?)
sandspitensis is probably indicated by the short spines that
occur on the proximal half of the feet. It should be noted
that some specimens of Napora, such as N. praespinifera
(Pessagno, 1977b) and N. spinifera (Pessagno, 1977b) display velum-like structure below the thorax and between
the proximal halves of the feet; however, we have never observed the basal closure of this velum-like structure. Jacus
(?) sandspitensis differs from J. coronatus De Wever, 1982,
by possessing a structurally simpler horn lacking a crownlike mass at its tip, and by having considerably longer, slender feet. Both species possess ridges at the base of the thorax, which separate rows of pore frames.
Measurements (µm):
Numbers of specimens measured are in parentheses.
HT
Mean
Length of cephalis
25
23.8 (9)
Length of thorax
100 76.1 (9)
50
48.0 (9)
Width of thorax at top
112.5 105.5 (9)
Width of thorax at base
Length of horn
55
81.7 (7)
47.5 35.8 (9)
Width of horn at base
Length of foot (maximum) 170 147.5 (5)
Max.
25 (9)
100 (9)
62.5 (9)
125 (9)
92.5 (7)
47.5 (9)
175 (5)
Min.
15 (9)
55 (9)
40 (9)
95 (9)
50 (7)
25 (9)
95 (5)
Etymology: This species is named for the town of Sandspit
in the Queen Charlotte Islands, British Columbia.
Plate NAP04. Napora relica Yeh. Magnification x250x. Fig. 1(H). Yeh 1987b, pl. 10, fig. 4. Fig. 2. JP, IYII-20.
Fig. 3. Goričan et al. 2003, pl. 4, fig. 4. Fig. 4. OM, BR1121-R09-06. Fig. 5. OM, BR524-R05-24. Fig. 6. OM, BR525-R08-01.
Plate JAC01. Napora sandspitensis (Pessagno, Whalen & Yeh). Magnification x200. Fig. 1(H). Pessagno, Whalen & Yeh
1986, pl. 2, fig. 9. Fig. 2. QCI, GSC loc. C-080613, GSC 128845. Fig. 3. QCI, GSC loc. C-140495, GSC 128846.
Fig. 4. QCI, GSC loc. C-175311, GSC 111727.
255
Type locality: Sample QC-534, Rennell Junction member
of the Fannin Formation (Maude Formation in Pessagno et
al., 1986), Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands and Ghost Creek formations, and
Rennell Junction member of the Fannin Formation, Queen
Charlotte Islands; Tawi Sadh Member of the Guwayza
Formation, Oman.
Genus: Naropa Dumitrica n. gen.
Type species: Naropa vi Hori, Whalen & Dumitrica n. sp.
Description: Test conical, dicyrtid or eventually tricyrtid,
with non-bladed apical and ventral horns and three triradiate feet having two blades on the external side and one on
the internal side. Thorax latticed, wide open. Initial spicule
unknown but supposed to be similar to that of Napora.
Remarks: Superficially this new genus resembles Napora
and other ultranaporids but differs in having both apical
and ventral horns non-bladed and the feet with one blade
directed towards the axis of shell and two directed laterally.
Unlike this new genus, the apical horn of Napora is always
three-bladed and commonly has a crown of spinules, the
ventral horn is short and bladed, and one blade is always
centrifugally directed and represents the prolongation of
the dorsal and primary lateral spines of the initial spicule.
Etymology: The name is an anagram of Napora, which, in
turn, is an anagram of Parona, one of the pioneers of the
study of fossil radiolarians. Feminine gender.
Included species:
UTD01 Naropa vi Hori, Whalen & Dumitrica n. sp.
Naropa vi Hori, Whalen & Dumitrica n. sp.
Species code: UTD01
Synonymy:
1990 Dumitricaella (?) sp. A – Hori, p. 581, fig. 8.24.
2002 Ultranaporid gen. et sp. indet. – Whalen & Carter, p. 140,
pl. 16, fig. 14.
Type designation: Holotype specimen pl. UTD01, fig. 1
from sample BPW80-14 Baja California Sur.
Description: Test conical, dicyrtid. Cephalis small, with
polygonal pore frames. Apical and ventral horns long,
cylindrical and fused at the base; horns disposed in a
V shape. Thorax conical with hexagonal pore frames
arranged in staggered horizontal rows and, in some
specimens, with a circumferential ridge at middle part.
Distal opening of thorax triangular, surrounded by a wide
imperforate rim. Feet straight, divergent, triradiate, pointed
distally. Two outer blades of each foot interconnected with
the corresponding blade of neighbouring feet at the base
of the preforate part of the thorax.
Remarks: This species is rare but morphologically distinctive. It differs from typical Napora in having both apical
256
and ventral horns circular in cross-section. Moreover, the
tribladed feet display two outer blades with a wide groove
in between.
Measurements (µm):
Number of specimens measured in parentheses.
Length of cephalis
Length of thorax
Maximum width of thorax
Length of apical horn
Length of ventral horn
Maximum length of feet
HT
21
90
80
64
49
98
Mean
25.3 (23)
79.8 (23)
90.8 (23)
70.7 (22)
35.9 (13)
90.8 (23)
Max.
33 (23)
100 (23)
107 (23)
90 (22)
53 (13)
107 (23)
Min.
20 (23)
57 (23)
77 (23)
60 (22)
18 (13)
77 (23)
Etymology: From the two cephalic horns that outline the
V letter; noun.
Type locality: Sample BPW80-14 Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Inuyama, Mino terrane and Kaiji, Kuma, Chichibu terrane,
SW Japan.
Plate UTD01. Naropa vi Hori, Whalen & Dumitrica n. sp. Magnification x250. Fig. 1(H). Whalen & Carter 2002,
pl. 16, fig. 14. Fig. 2. Hori 1990, fig. 8.24. Fig. 3. JP, Ku(b)-5-16, RH(1)1026. Fig. 4. JP, UC2-22-1, RH(1)75. Fig. 5. JP,
KG9-52, RH(1)2803.
257
Genus: Noritus Pessagno & Whalen 1982
Type species: Noritus lillihornensis Pessagno & Whalen 1982
Synonymy:
1982 Noritus n. gen. – Pessagno & Whalen, p. 123.
Original description: Test conical to subconical. Cephalis
with well-developed horn. Pore frames polygonal, regular
to irregular, aligned to nonaligned; thickened by the
insertion of longitudinal and lateral ridges between pore
frames; ridges not inserted between all pore frames. Wellpreserved specimens with tubular extension on final postabdominal chamber; pore frames of tubular extension
thinner than those on previous chambers and with
rudimentary development of ridges.
Original remarks: Noritus, n. gen., differs from Droltus
n. gen., by possessing a test wall on its post-abdominal
chambers with a partially developed outer layer. Whereas
the pore frames of Droltus are thickened along bars in all
directions, those of Noritus are not; many pore frames, in
fact, may display no thickening at all (cf. pl. 4, figs. 1, 6, 10
and pl. 5, figs. 3, 4, 10, 15, 19). Because of this difference in
the mode of test construction, Noritus is tentatively placed
in the Bagotidae.
Etymology: Noritus is a name formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, Pt. VI,
Recommendation 40, p. 113).
Included species:
NTS01 Noritus lillihornensis Pessagno & Whalen 1982
Noritus lillihornensis Pessagno & Whalen 1982
Species code: NTS01
Synonymy:
1982 Noritus lillihornensis n. sp. – Pessagno & Whalen, p. 123, pl.
5, figs. 3, 4, 10, 15, 19; pl. 12, fig. 1.
1992 Noritus lillihornensis n. sp. Pessagno & Whalen – Pessagno
& Mizutani, pl. 99, figs. 1, 2, 9.
2004 Noritus sp. – Matsuoka, fig. 229.
with a longer, more massive horn. It differs from all species
of Droltus by the characters cited under the genus.
Measurements (µm):
Based on 10 specimens.
Length excluding horn
287.5
287.5
175.0
212.7
Width (max.)
100.0
110.0
62.5
91.7
HT
Max.
Min.
Mean
Original description: Test elongate with long massive horn
on hemispherical cephalis and with five post-abdominal
chambers. Cephalis and thorax with small polygonal pore
frames; thorax and subsequent chambers trapezoidal
in cross section. Post-abdominal chambers increasing
gradually in length and somewhat more rapidly in width
as added; final post-abdominal chamber with long, tubular
extension which may be 1/3 length of remainder of test.
Pore frames tetragonal to pentagonal (predominantly
tetragonal). Ridges not developed on all pore frames.
Etymology: This species is named for Lillihorn Island in
Skidegate Channel.
Original remarks: Noritus lillihornensis, n. sp., differs from
N. sp. A by having a much slenderer, more elongate test
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Japan.
258
Type locality: Sample QC 534, Fannin Formation (Maude
Formation in Pessagno & Whalen, 1982), Queen Charlotte
Islands, Maude Island, Skidegate Inlet, British Columbia.
Plate NTS01. Noritus lillihornensis Pessagno & Whalen. Magnification x250. Fig. 1(H). Pessagno & Whalen 1982,
pl. 5, fig. 3. Fig. 2. QCI, GSC loc. C-080612, GSC 128847. Fig. 3. QCI, GSC loc. C-080612, GSC 111728.
259
Genus: Orbiculiformella Kozur & Mostler 1978
Type species: Orbiculiforma railensis Pessagno 1977b
Synonymy:
1978 Orbiculiformella n. gen. – Kozur & Mostler, p. 162.
Original description: Circular spongy skeleton. Marginal
part clearly inflated, with large pores. Depressed inner
part rather thick, with very small pores. Six to twelve main
spines extending from test.
Original remarks: According to usual practice in all
other radiolarian families, subquadratic forms with four
main spines and circular forms with 6-12 main spines
will be herein considered as two separate genera. The
genus Orbiculiforma Pessagno 1973 will be restricted to
subquadratic forms with four main spines, the circular
forms with 6-12 main spines will be assigned to the new
genus Orbiculiformella.
Further remarks: The genus should be emended to comprise species with more than six spines, without limiting
the higher number. The genus is probably polyphyletic but
because of the imperfect preservation of the microsphere
and of the absence of studies of the microsphere of all species, we include here all species having the external morphologic diagnostic characters described by the authors of
the genus.
Etymology: Arbitrary word formation. This paper: the
name is a diminutive of Orbiculiforma.
Included species and subspecies:
ORB05 Orbiculiformella callosa (Yeh) 1987b
ORB06 Orbiculiformella incognita (Blome) 1984b
ORB03 Orbiculiformella lomgonensis (Whalen & Carter)
1998
ORB11 Orbiculiformella mediocircus Dumitrica n. sp.
ORB02 Orbiculiformella? robusta (Whalen & Carter) 1998
ORB08 Orbiculiformella teres (Hull) 1997
ORB13 Orbiculiformella? trispina s.l. (Yeh) 1987b
ORB09 Orbiculiformella? trispina trispina (Yeh) 1987b
ORB10 Orbiculiformella? trispina trispinula (Carter) 1988
Orbiculiformella callosa (Yeh) 1987b
Species code: ORB05
Synonymy:
1987b Orbiculiforma callosa n. sp. – Yeh, p. 41, pl. 2, fig. 25; pl. 5,
fig. 19; pl. 11, fig. 11, pl. 22, figs. 10, 12.
1988 Orbiculiforma kwunaensis Carter n. sp. – Carter et al.,
p. 44, pl. 1, figs. 8, 11.
1996 Orbiculiforma sp. A – Pujana, p. 135, pl. 1, fig. 12.
1998 Orbiculiforma callosa Yeh – Cordey, p. 93, pl. 21, figs. 2, 4, 10.
2002 Orbiculiformella kwunaensis Carter – Whalen & Carter,
p. 109, pl. 1, fig. 3.
2003 Orbiculiforma ? callosa Yeh – Goričan et al., p. 295, pl. 3,
figs. 1-4.
2004 Orbiculiformella sp. – Matsuoka, fig. 53.
Original description: Test thick, circular in outline, with
large, deep central cavity. Test consisting of concentric layers of small irregular polygonal pore frames with circular to
elliptical pores. Pore frames slightly larger on rims, smaller in central cavity and median band of margin. Margin
slightly concaved with several small peripheral spines. Peripheral spines usually thin, short, circular in cross-section.
Diameter of central cavity about half the diameter of test.
Original remarks: Orbiculiforma callosa, n. sp., differs from
O. (?) trispina, n. sp., by having a test with larger central
cavity and by having several small peripheral spines rather
than three massive, elongate spines.
Further remarks: Orbiculiformella kwunaensis Carter is
synonymized with O. callosa Yeh because the larger pore
frames in central area are now judged to be species variability.
260
Note that some Pliensbachian specimens (pl. ORB05,
fig. 8) can be more than twice as large as the holotype.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Diameter
of shell
160
170
187
158
Diameter of
central cavity
80
82
87
75
Length
of spines
38
45
30
Etymology: Callosus-a-um (Latin, adj.) = hard.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, Oregon;
Fannin Formation, Queen Charlotte Islands; Bridge River
Complex, British Columbia; San Hipólito Formation, Baja
California Sur; Sierra Chacaicó Formation, Argentina;
Skrile Formation, Slovenia; Dürrnberg Formation, Austria;
Gümüslü Allochthon; Turkey; Tawi Sadh Member of the
Guwayza Formation, Oman; Mino Terrane, Japan.
Plate ORB05. Orbiculiformella callosa (Yeh). Magnification Figs. 1-5 x200 (scale bar A), Figs. 6-11 x 150 (scale bar B).
Fig. 1a(H). Yeh 1987b, pl. 22, fig. 12. Fig. 1b(H). Yeh 1987b, pl. 22, fig. 10. Fig. 2. OM, BR871-R02-14.
Fig. 3. Goričan et al. 2003, pl. 3, fig. 1. Fig. 4. OM, BR871-R02-13. Figs. 5a, b. OM, BR1122-R04-13a, b.
Fig. 6. Carter et al. 1988, pl. 1, fig. 11. Fig. 7. AT, BMW21-21. Fig. 8. TR, 1662D-R06-16. Fig. 9a. BCS, Loc. SH-412-14.
Fig. 9b. Whalen & Carter 2002, pl. 1, fig. 3. Figs. 10a, b. OM, BR706-R06-22a, b. Fig. 11. OM, BR1121-R07-19.
261
Orbiculiformella incognita (Blome) 1984b
Species code: ORB06
Synonymy:
1984b Orbiculiforma (?) incognita n. sp. – Blome, p. 353, pl. 5,
figs. 1, 2, 8, 9, 12, 13.
1988 Spongotrochus (Stylospongidium) sp. aff. S. (S.) echinodiscus
Clark and Campbell – Carter et al., p. 46, pl. 10, figs. 7, 10.
2003 Orbiculiforma ? incognita Blome – Goričan et al., p. 296,
pl. 3, figs. 5-7.
Original description: Test thin, circular in outline. Meshwork consisting of irregular, polygonal (tetragonal to hexagonal) pore frames, becoming slightly smaller toward
central area; pores circular to elliptical in outline. Central
cavity extremely shallow, narrow; width approximately
one-third that of test diameter. Sides of test gently rounded.
Periphery of test possessing numerous, slender, rodlike peripheral spines; spines circular in axial section.
Original remarks: Orbiculiforma (?) incognita, n. sp. differs
from O. monticelloensis Pessagno 1973 as well as other species of Orbiculiforma described in this report by having an
extremely shallow central area and by having longer, rodlike peripheral spines. This form differs from other taxa
belonging to the genus Orbiculiforma by not possessing a
well-developed central cavity and is therefore questionably
assigned to this genus.
Further remarks: We include also forms with upper and
lower planar surfaces of the shell and uniformly sized pores.
Our specimens have a rather well-pronounced groove running along the test margin, whereas in the type material the
test margins are gently rounded.
Measurements (µm):
Based on 6 specimens.
Test diam. (max.)
179
181
162
172
Spine length
26-36
65
22
36
HT
Max.
Min.
Av.
Etymology: Incognitus-a-um (Latin, adj., f.) = unexamined,
unknown, unrecognized.
Type locality: Sample 80AJM 8A, Shelikof Formation,
Puale Bay, southern Alaska.
Occurrence: Shelikof Formation, southern Alaska; Phantom Creek and Graham Island formations, Queen Charlotte Island, British Columbia; Skrile Formation, Slovenia.
Orbiculiformella lomgonensis (Whalen & Carter) 1998
Species code: ORB03
Synonymy:
1998 Praeorbiculiformella? lomgonensis n. sp. – Whalen &
Carter, p. 58, pl. 9, fig. 8.
Original description: Test large, circular in outline with a
straight-sided pariphery; test relatively thick in proportion
to diameter. Seven relatively broad grooves radiating from
centre of test to margin. Meshwork composed primarily of
small, irregularly shaped polygonal pore frames; meshwork
generally uniform in size over surface of test.
Original remarks: Although the overall characteristics of
Praeorbiculliformella? lomgonensis n. sp. suggest inclusion
with Praeorbiculliformella Kozur and Mostler, the unusual
radiating grooves have not been observed on any other
species of this genus.
262
Measurements (µm):
Based on 17 specimens.
Maximum diameter of cortical shell
340
395
281
347
HT
Max.
Min.
Mean
Etymology: This species is named for Lomgon Bay located
on the north side of Tasu Sound.
Type locality: Sample 86-CAA-T-2/3, Sandilands Formation, Graham Island, Yakoun River area, Queen Charlotte
Islands, British Columbia.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands.
Plate ORB06. Orbiculiformella incognita (Blome). Magnification x200. Fig. 1(H). Blome 1984b, pl. 5, fig. 1.
Fig. 2. Carter et al. 1988, pl. 10. fig. 10. Fig. 3. Carter et al. 1988, pl. 10, fig. 7. Fig. 4. Goričan et al. 2003, pl. 3, fig. 7.
Fig. 5. SI, MM 21.70-000333. Figs. 6a, b. Goričan et al. 2003, pl. 3, figs. 5a, b.
Plate ORB03. Orbiculiformella lomgonensis (Whalen & Carter). Magnification x150. Fig. 1(H). Carter et al. 1998,
pl. 9, fig. 8. Fig. 2. QCI, GSC loc. C-305386, GSC 128848. Fig. 3. QCI, GSC loc. C-080612, GSC 128849.
263
Orbiculiformella mediocircus Dumitrica n. sp.
Species code: ORB11
Type designation: Holotype specimen 1662D-R09-05 from
sample 1662D, Gümüslü Allochthon, Taurus Mts., Turkey.
a thick circular ring between the central depression and
sloping periphery.
Diagnosis: Test spongy, circular with a large central
depression, a rounded sloping periphery and a thick spongy
circular ridge between the two.
Measurements (µm):
Based on 4 specimens.
Diameter of central cavity
External diameter of thick ring
Diameter of entire skeleton
HT
360
490
605
Min.
220
345
420
Max.
360
490
605
Description: Test spongy, circular with a wide central
depression bordered by a thick, high, spongy circular ring
which is hemispherical in transverse section. Periphery of
disc beyond the ring sloping resulting in a round-angled
circular band without solid spines but having fine spinules.
Meshwork of test with rounded polygonal meshes which
are larger on the thick ring decreasing in size on the
periphery.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Remarks: This species is well distinguished from other
species of the genus by its larger size and the presence of
Occurrence: Gümüslü Allochthon, Taurus Mts., Turkey;
Tawi Sadh Member of the Guwayza Formation, Oman.
Etymology: From the Latin medius – in the middle and
circus – circle; noun.
Orbiculiformella? robusta (Whalen & Carter) 1998
Species code: ORB02
Synonymy:
1998 Praeorbiculiformella robusta n. sp. – Whalen & Carter,
p. 58, pl. 9, figs. 2, 3, 4, 19.
Original description: Test large, nearly circular in outline
with straight-sided periphery, very thin in proportion
relative to diameter. Very broad, shallow depression in
centre of test usually destroyed. Three strong, principal
spines intersect in centre of test; point of intersection
surrounded by dense spongy meshwork with small pores;
principal spines usually triradiate in axial section and
extend from periphery of feet. Subsidiary spine located
between each principal peripheral spine; subsidiary spines
extend from margins of test only, not penetrating to centre
of test. Meshwork primarily composed of small, irregularly
shaped pentagonal and tetragonal pore frames; meshwork
generally uniform in size over entire test.
Original remarks: The large test and strong spines distinguish Praeorbiculiformella robusta n. sp. from P. yanensis
n. sp.
264
Measurements (µm):
Based on 13 specimens.
Diameter of
cortical shell
325
354
225
295
Width of
central area
195
228
195
181
Length of
longest spine
118
125
52
92
HT
Max.
Min.
Mean
Etymology: Robustus, a, um (Latin; adj.) = hard and strong
like oak.
Type locality: Sample 89-CNA-KUG-1A, Sandilands Formation, north Kunga Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte
Islands.
Plate ORB11. Orbiculiformella mediocircus Dumitrica n. sp. Magnification x150. Figs. 1(H)a-c. TR, 1662D-R09-05a-c.
Fig. 2. OM, BR476-R18-01.
Plate ORB02. Orbiculiformella? robusta (Whalen & Carter). Magnification x150. Fig. 1(H). Carter et al. 1998,
pl. 9, fig. 2.
265
Orbiculiformella teres (Hull) 1997
Species code: ORB08
1997 Orbiculiforma teres n. sp. – Hull, p. 16, pl. 1, figs. 10, 11, 15, 19.
1999 Orbiculiforma (?) teres Hull – Kiessling, p. 41, pl. 8, fig. 6.
2003 Orbiculiforma? teres Hull – Goričan et al., p. 296, pl. 3,
figs. 8-13.
differs from Orbiculiforma multifora Pessagno and Poisson
(1981) by having a very indistinct or no central depression.
De Wever (1981b, p. 46, plate 7, figs. 6-8) illustrated
a closely related spongodiscid which, in addition to
peripheral spines, possesses two polar spines.
Original description: Test large, relatively thin, subcircular
in outline. Central cavity very shallow. Twelve to eighteen
short spines along periphery; base of spines usually weakly
triradiate, becoming circular in axial section distally.
Meshwork fine, composed of tetragonal and pentagonal
pore frames.
Measurements (µm):
Based on 7 specimens. AA’ and BB’ represent the diameter
of the spongy test in two planes oriented perpendicular to
one another.
Synonymy:
Original remarks: Orbiculiforma teres, n. sp., is distinguished by its large size, very shallow central cavity, and
numerous short peripheral spines.
Further remarks: By Goričan et al. (2003): Included are
lenticular forms with angled sides of test and numerous
peripheral spines. Spines can vary from short and thin to
rather thick, sometimes bladed or flattened at the base.
Some specimens have a small shallow central depression
(Plate III, fig. 11 a, b) whereas others show a perfectly
biconvex shell (Plate III, figs. 9, 13). Orbiculiforma? teres
266
AA’
BB’
HT
225.0
75.0
Min.
215.0
50.0
Max.
252.0
80.0
Mean
235.4
68.2
Etymology: Teres (Latin, adj.) = smooth, polished.
Type locality: Sample SM-50, Volcanopelagic strata overlying the Coast Range Ophiolite, Stanley Mountain, California Coast ranges.
Occurrence: Volcanopelagic strata at Stanley Mountain,
California; Ameghino Formation, Antarctic Peninsula;
Skrile Formation, Slovenia.
Plate ORB08. Orbiculiformella teres (Hull). Magnification x250. Fig. 1(H). Hull 1997, pl. 1, fig. 10. Figs. 2a, b. Goričan
et al. 2003, pl. 3, figs. 12a, b. Fig. 3. Goričan et al. 2003, pl. 3, fig. 9. Figs. 4a, b. Goričan et al. 2003, pl. 3, figs. 8a, b.
Fig. 5. Goričan et al. 2003, pl. 3, fig. 13. Fig. 6. Goričan et al. 2003, pl. 3, fig. 10. Fig. 7. Goričan et al. 2003, pl. 3, fig. 11b.
Fig. 8. SI, MM 21.70, 010224. Figs. 9a, b. SI, MM 11.76, 000226, 000227.
267
Orbiculiformella? trispina s.l. (Yeh) 1987b
Species code: ORB13
Synonymy:
1987b Orbiculiforma (?) trispina n. sp. – Yeh, p. 42, pl. 9,
figs. 2, 10 (O. (?) trispinosa in plate caption).
See also subspecies.
Included subspecies:
ORB09 Orbiculiformella? trispina trispina (Yeh) 1987b
ORB10 Orbiculiformella? trispina trispinula (Carter) 1988
Orbiculiformella? trispina trispina (Yeh) 1987b
Species code: ORB09
Synonymy:
1987b Orbiculiforma (?) trispina n. sp. – Yeh, p. 42, pl. 9,
figs. 2, 10 (O. (?) trispinosa in plate caption).
1998 Orbiculiforma silicatilis n. sp. – Cordey, p. 93, pl. 21,
figs. 5, ? 8, not fig. 7.
Original description: Test thick, circular in outline, with
small, deep central cavity and three periphery spines.
Periphery spines elongate, relatively massive, circular in
cross-section with most portion circular in cross-section
with only proximal end triradiate with three deep grooves
alternating with three rounded ridges. Three spines nearly
equally spaced. Meshwork of test predominantly of relatively
uniform size of small irregular pore frames. Margin of test
slightly concaved.
Original remarks: Orbiculiforma (?) trispina, n. sp., can be
easily distinguished from other Orbiculiforma spp. in this
report by having three long massive periphery spines. It is
possible that this form should be assigned to a new genus.
Measurements (µm):
Ten specimens measured.
Diameter of shell
158
170
187
158
Diameter of
central cavity
53
82
88
75
Etymology: Tri = three, spina = spine.
Length of
spines
168
38
45
30
HT
Mean
Max.
Min.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, Oregon; Fannin
Formation, Queen Charlotte Islands; Bridge River and
Hozameen complexes, British Columbia.
Orbiculiformella? trispina trispinula (Carter) 1988
Species code: ORB10
Synonymy:
1988 Orbiculiforma trispinula Carter n. sp. – Carter et al., p. 44,
pl. 1, figs. 7, 10.
Original diagnosis: Test discoidal with large central
depression and three short radial spines. Meshwork spongy
and coarse, pore frames larger in central area.
Original description: Test a circular disc with large central
cavity and three radial spines. Surface of test planiform,
sides vertical. Central cavity large (approximately half test
diameter) and deeply depressed. Pore frames polygonal and
concentrically arranged; very small and delicate in the middle of the central area, but immediately surrounding ones
are somewhat larger; composed of thin fragile bars. Pore
frames on rim-like upper surfaces small, bars are coarser
and small nodes occur at vertices of bars. Spines short and
circular in section.
Original remarks: This form differs from Orbiculiforma(?)
trispina Yeh by having a larger overall diameter and thick268
ness, a wider central cavity (and narrower rim) and much
smaller radial spines.
Further remarks: Orbiculiformella? trispina trispinula is
now considered a subspecies of Orbiculiformella? trispina
(Yeh).
Measurements (µm):
Based on 7 specimens.
HT Av. Max. Min.
Maximum diameter of test
206 218 230 200
Maximum diameter of central cavity 134 130 155 110
Length of spines
46 43
46
40
Etymology: Latin, trispinula (adj.), three small spines.
Type locality: GCS locality C-080577, Fannin Formation,
Creek locality, Maude Island.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Fernie Formation, NE British Columbia.
Plate ORB09. Orbiculiformella? trispina trispina (Yeh). Magnification x 200. Fig. 1(H). Yeh 1987b, pl. 9, fig. 2.
Fig. 2. QCI, GSC loc. C-304566, GSC 128851.
Plate ORB10. Orbiculiformella? trispina trispinula (Carter). Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 1,
fig. 7. Fig. 2. NBC, GSC loc. C-305208, GSC 128852. Fig. 3. QCI, GSC loc. C-080612, GSC 128853. Fig. 4. QCI, GSC loc.
C-080612, GSC 128854 .
269
Genus: Palaeosaturnalis Donofrio & Mostler 1978,
emend. Kozur & Mostler 1981
Type species: Spongosaturnalis triassicus Kozur & Mostler 1972
Synonymy:
1978 Palaeosaturnalis n. gen. – Donofrio & Mostler, p. 33.
1981 Palaeosaturnalis Donofrio & Mostler emend. – Kozur &
Mostler, p. 55.
1983 Palaeosaturnalis Donofrio & Mostler emend. Kozur &
Mostler – Kozur & Mostler, p. 19.
1984 Palaeosaturnalis Donofrio & Mostler emend. – De Wever,
p. 15.
Original diagnosis: Form with a smooth, flat, simple ring
of very variable width, bearing spines all around; thorns
on the external side missing. In most cases, auxiliary or
sustaining bars are developed near polar bars. Cortical and
medullary shells as with family.
Emended description: Kozur & Mostler (1981): Forms
with a smooth, flattened, simple ring of variable width and
a periphery with spines. The two polar spines are always
opposite to the spines of outer margin. Sustaining spines of
second order absent. Cortical shell mostly spongy, widely
separated from the ring. One medullary shell usually
present.
Original remarks: The name Palaeosaturnalis was chosen
because Triassic representatives are all characterized by a
ring with a circular cross-section. Unlike Jurassic-Cretaceous representatives, the polar sustaining bars [of the Triassic forms] are, except some specimens, in direct continuation of the peripheral spines (see text-fig. 8). Moreover,
the stratigraphically younger forms mostly lack sustaining
bars, whereas they are very frequent in Triassic forms.
Further remarks: The genus Palaeosaturnalis differs from
Pseudoheliodiscus in not having auxiliary or/and subsidiary
rays (De Wever, 1984, p. 15-16).
Included species:
SAT13 Palaeosaturnalis aff. liassicus Kozur & Mostler 1990
SAT12 Palaeosaturnalis subovalis Kozur & Mostler 1990
SAT14 Palaeosaturnalis sp. B sensu Whalen & Carter 2002
Palaeosaturnalis aff. liassicus Kozur & Mostler 1990
Species code: SAT13
Synonymy:
aff. 1990 Palaeosaturnalis liassicus n. sp. – Kozur & Mostler, p.
192, pl. 1, figs. 2, 3; pl. 12, figs. 1, 3, 4, 6, 8-10; pl. 13,
figs. 1, 2, 6, 7.
2002 Paleosaturnalis sp. A – Whalen & Carter, p. 108, pl. 5, fig. 5.
Remarks: This species differs from typical Palaeosaturnalis
liassicus by having fewer peripolar spines.
Occurrence: San Hipólito Formation, Baja California Sur.
Palaeosaturnalis subovalis Kozur & Mostler 1990
Species code: SAT12
Synonymy:
1972 Spongosaturnalis ? sp. c – Yao, p. 35, pl. 8, fig. 3.
1987b Acanthocircus sp. B – Yeh, p. 49, pl. 5, fig. 13.
1990 Palaeosaturnalis subovalis n. sp. – Kozur & Mostler, p. 193,
pl. 1, fig. 7; pl. 13, figs. 4, 9.
1991 Palaeosaturnalis sp. aff. P. liassicus Kozur & Mostler – Yang
& Mizutani, p. 65, pl. 2, figs. 4, 11, 13; not pl. 3, figs. 2, 12, 13.
2002 Palaeosaturnalis lenggriesensis Kozur & Mostler – Tekin,
p. 182, pl. 2, fig. 2.
Not 2002 Palaeosaturnalis subovalis Kozur & Mostler – Tekin,
p. 182, pl. 2, fig. 5.
Original description: Shell slightly ellipsoidal, spongy,
consisting of several concentric layers. Microsphere latticed. Ring suboval to oval, narrow, flat, undifferentiated,
with 9-11 slender, relatively short spines. One axial spine
as long as the circumaxial spines, the other one is considerably longer and somewhat more slender. A larger smooth
segment is present to both sides of the axial spines. Polar
spines very robust.
270
Original remarks: In Palaeosaturnalis haeckeli n. sp. the
circumaxial spines are more variable in their length. On
both sides of the axial spines 1-2 large, widely spaced spines
are present. The remaining circumaxial spines are smaller
and closely spaced.
Further remarks: The very few specimens we included in
this species resemble very well the holotype and especially
one paratype (Kozur & Mostler, 1990, pl. 1, fig. 7).
Measurements (µm):
Diameter of shell
(parallel to polar axis)
Diameter of shell
(perpendicularly to polar axis)
Diameter of ring
(parallel to polar axis)
Diameter of ring
(perpendicularly to polar axis)
Min.
Max.
76
90
60
77
192
215
190
240
Plate SAT13. Palaeosaturnalis aff. liassicus Kozur & Mostler. Magnification x150. Fig. 1. Whalen & Carter 2002,
pl. 5, fig. 5.
Plate SAT12. Palaeosaturnalis subovalis Kozur & Mostler. Magnification x200. Fig. 1(H). Kozur & Mostler 1990,
pl. 13, fig. 4. Fig. 2. OM, BR706-R13-13. Fig. 3. JP, IYII17-63, RH(1)1660. Fig. 4. NBC, GSC loc. C-305208, GSC 111729.
271
Etymology: According to the suboval ring outline.
Type locality: Sample L1, Kirchstein Limestone at Mt.
Kirchstein, 6.5 km WSW of Lenggries/Isar, Bavaria,
Germany.
Occurrence: Kirchstein Limestone, Germany; Varhegy
Cherty Limestone Formation, Hungary; Hocaköy Radiolarite, Turkey; Tawi Sadh Member of the Guwayza Formation, Oman; Japan; Nadanhada Terrane, China; Nicely Formation, Oregon; Fernie Formation, NE British Columbia.
Palaeosaturnalis sp. B sensu Whalen & Carter 2002
Species code: SAT14
Synonymy:
2002 Paleosaturnalis sp. B – Whalen & Carter, p. 108, pl. 5,
figs. 6, 10.
Original remarks: This species is distinguished by having
bifurcating polar spines.
Occurrence: San Hipólito Formation, Baja California Sur.
Genus: Pantanellium Pessagno 1977a
Type species: Pantanellium riedeli Pessagno 1977a
Synonymy:
is under study from the Upper Triassic and another from
the Lower Cretaceous.
Original description: Test divided into ellipsoidal to subspherical cortical shell and spherical first medullary shell,
both with massive polygonal pore frames having nodes at
vertices. Cortical shell with bipolar primary spines possessing well-developed alternating, longitudinally arranged
ridges and grooves. One spine often somewhat shorter
than other. Primary spines interconnected and occurring
along same axes as primary beams which connect cortical shell to first medullary shell; diameter of two primary
beams about half that of primary spines. Secondary radial
beams also connecting cortical shell (pl. 6, fig. 6); extending from nodal points of pore frame vertices of both cortical and first medullary shells.
Further remarks: Kozur & Mostler (1990, p. 214) argued
that Pantanellium Pessagno is a younger synonym of
Ellipsoxiphus Dunikowski 1882 with Xiphosphaera
(Ellipsoxiphus) suessi Dunikowski 1882 as type species. In
modern radiolarian literature the name Ellipsoxiphus has
been used for Cenozoic species only. On the other hand,
Mesozoic species have been assigned to Pantanellium
by a great majority of authors and this usage has largely
prevailed even since 1990. Pantanellium is a thoroughly
studied genus with a well-established stratigraphic range
from the Late Triassic to the Early Cretaceous (Aptian).
In spite of the Principle of Priority we continue to use the
generic name Pantanellium, because its replacement with
Ellipsoxiphus would certainly cause confusion (see ICZN
1999, art. 23.2 and Preamble p. XX).
1977a Pantanellium n. gen. – Pessagno, p. 78.
Original remarks: Pantanellium n. gen., differs from Protoxiphotractus Pessagno in having bipolar spines with longitudinally arranged, alternating grooves and ridges. Many
workers, for example Foreman (1973, p. 258), have included species assignable to this genus under Sphaerostylus
Haeckel. Unfortunately, the single illustration and the description of the type species of Sphaerostylus (i.e., S. zitteli
Rüst) are exceedingly poor and of virtually no use to any
worker hoping to make a definitive identification. The resurrection of the name Sphaerostylus can serve no purpose.
It is suggested, therefore, that the name Sphaerostylus be
considered a nomen dubium. Pantanellium is known to include several species. At present, only the type species and
P. fischeri (Pessagno) have formal names. One new species
272
Etymology: This genus takes its name from Dante Pantanelli, one of the early students of Mesozoic Radiolaria.
Included species:
PAN20 Pantanellium brevispinum Carter n. sp.
PAN14 Pantanellium carlense Whalen & Carter 1998
PAN18 Pantanellium cumshewaense Pessagno & Blome
1980
PAN11 Pantanellium danaense Pessagno & Blome 1980
PAN19 Pantanellium inornatum Pessagno & Poisson 1981
PAN16 Pantanellium skedansense Pessagno & Blome 1980
Plate SAT14. Paleosaturnalis sp. B sensu Whalen & Carter. Magnification x 150. Fig. 1. Whalen & Carter 2002, pl. 5, fig. 6.
273
Pantanellium brevispinum Carter n. sp.
Species code: PAN20
Synonymy:
1980 Pantanellium sp. G – Pessagno & Blome, p. 248, pl. 6, fig. 9.
1997 Pantanellium sp. C – Yao, pl. 3, fig. 139.
Type designation: Holotype GSC 111730 from GSC loc. C080611; Ghost Creek Formation (lower Pliensbachian).
Description: Cortical shell relatively large, spherical to subspherical with medium-sized pores and very low rounded
nodes at pore frame vertices. Pore frames pentagonal and
hexagonal, thicker in Z direction than in Y direction; five
pore frames visible along AA’, six visible along BB’. Polar
spines very short, less than one-quarter diameter of test;
spines strongly tapering, one slightly longer than the other.
Spines usually triradiate in axial section with three, narrow
ridges and three relatively wide grooves.
Remarks: Pantanellium brevispinum n. sp. differs from
P. haidaense Pessagno & Blome by having much shorter
polar spines, and lower nodes at pore frame vertices.
Measurements (µm):
Based on 6 specimens. System of measurement shown in
text-figure 5 of Pessagno & Blome (1980).
AA’
A’S’
AS
BB’
cc’
dd’
HT
105
37
32
100
39
32
Max.
111
37
37
111
39
32
Min.
94
37
32
93
23
23
Mean
102
43
34
100
32
27
Etymology: From the Latin brevis + spinus, -a, -um (with
short spines).
Type locality: Sample CAA-79-Ren-Phant, lms 1 (GSC loc.
C-080611), Ghost Creek Formation, Rennell Junction section, central Graham Island, Queen Charlotte Islands, British Columbia.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Japan.
Pantanellium carlense Whalen & Carter 1998
Species code: PAN14
Synonymy:
1984 Pantanellium sp. – Whalen & Pessagno, pl. 1, fig. 17.
1998 Pantanellium carlense n. sp. – Whalen & Carter, p. 47, pl. 2,
figs. 1, 2, 13, 14, 17, 18.
2002 Pantanellium carlense Whalen & Carter – Whalen & Carter,
p. 105, pl. 9, figs. 1, 2, 10, 11.
Original description: Cortical shell subspherical in shape,
slightly elongated in plane of polar spines, with mediumto large- sized pentagonal and hexagonal pore frames. Bars
of pore frames thin along Y, thicker along Z (refer to Fig.
23). All pore frames with irregularly shaped, spinose nodes
at vertices. Five pore frames visible along AA’; five to six
pore frames visible along BB’. Polar spines triradiate in
axial section with narrow, rounded longitudinal ridges and
grooves; one polar spine approximately one third shorter
than the other; base of shorter polar spine slightly wider at
cc’ than the base of other spine at dd’. First medullary shell
with thin, fragile pentagonal and hexagonal pore frames.
Original remarks: The elongated cortical shell with spiny
nodes at pore frame vertices distinguishes Pantanellium
carlense n. sp. from all other species of Pantanellium.
274
Measurements (µm):
(n) = number of specimens measured. System of measurement shown in text-figure 5 of Pessagno & Blome (1980).
AA’
(7)
60
68
60
65
A’S’
(6)
120
120
105
116
AS
(6)
86
90
79
83
BB’
(7)
71
86
71
79
cc’
(7)
34
45
34
40
dd’
(7)
45
53
38
46
HT
Max.
Min.
Mean
Etymology: This species is named for Mount Carl, located
to the north of the type locality.
Type locality: Sample QC-675, southeast side of Kunga
Island, Queen Charlotte Islands.
Occurrence: Sandilands Formation, Queen Charlotte Islands; San Hipólito Formation, Baja California Sur.
Plate PAN20. Pantanellium brevispinum Carter n. sp. Magnification x300. Fig. 1(H). QCI, GSC loc. C-080611,
GSC 111730. Fig. 2. QCI, GSC loc. C-080611, GSC 128856.
Plate PAN14. Pantanellium carlense Whalen & Carter. Magnification x250, except Fig. 1(H)b x 500. Fig. 1(H) a, b.
Carter et al. 1998, pl. 2, figs. 1, 17. Fig. 2. Whalen & Carter 2002, pl. 9, fig. 2.
275
Pantanellium cumshewaense Pessagno & Blome 1980
Species code: PAN18
Synonymy:
1980 Pantanellium cumshewaense n. sp. – Pessagno & Blome,
p. 240, pl. 6, figs. 1, 2, 15, 17, 21, 22.
Original description: Cortical shell subspherical with
large, predominantly hexagonal pore frames with poorly
developed nodes of low relief at vertices. Bars of pore
frames moderately thick; thickness about equal along Y
and Z (text-fig. 5). Four to 5 pore frames visible along AA’;
5 pore frames visile along BB’. Polar spines triradiate in
axial section; longitudinally comprised of 3 broad grooves
alternating with 3 narrow ridges. One polar spine somewhat
shorter than other; longer spine somewhat rounded
distally, not coming to sharp point; ridges of longer spine
disappearing on distal third of spine. First medullary shell
with small, fragile pore frames.
Original remarks: P. cumshewaense, n. sp., differs from
P. baileyi, n. sp., in having better defined pore frames which
lack spines.
Measurements (µm):
Based on 11 specimens. System of measurement shown in
text-figure 5 of Pessagno & Blome (1980).
AA’
110
101
120
90
A’S’
88
97
115
80
AS
65
63
85
40
BB’
95
97
110
90
cc’
20
23
35
15
dd’
35
29
40
20
HT
Av.
Max.
Min.
Etymology: This species is named for Cumshewa Inlet
south of Skidegate Bay in the Queen Charlotte Islands.
Type locality: Sample QC 534, Fannin Formation (Maude
Formation in Pessagno & Blome, 1980), Queen Charlotte
Islands.
Occurrence: Fannin Formation, Queen Charlotte Islands.
Pantanellium danaense Pessagno & Blome 1980
Species code: PAN11
Synonymy:
1980 Pantanellium danaense n. sp. – Pessagno & Blome, p. 241,
pl. 4, figs. 9-11, 15.
1990 Ellipsoxiphus cf. danaensis (Pessagno & Blome) – Kozur &
Mostler, p. 215, pl. 14, fig. 13; ? pl. 15, fig. 15.
1998 Pantanellium danaense Pessagno & Blome – Whalen &
Carter, p. 47, pl. 2, figs. 4, 5.
2004 Pantanellium danaense Pessagno & Blome – Hori et al.,
pl. 5, fig. 4; pl. 6, fig. 16.
Original description: Cortical shell spherical with large
(occasional small) pentagonal and hexagonal pore frames;
pentagonal pore frames about equal in number to hexagonal pore frames. Bars of pore frames thin along Y; thick
along Z (text-fig. 5). All pore frames with spinose nodes at
vertices. Six pore frames visible along AA’; 5 pore frames
visible along BB’. Polar spines triradiate in axial section,
1 polar spine shorter than the other; shorter spine with 3
moderately wide longitudinal ridges alternating with 3 longitudinal grooves of about the same width. Longer spine
with 3 thin longitudinal ridges alternating with 3 wide longitudinal grooves; grooves about 3 times wider than ridges.
First medullary shell with thin, fragile, hexagonal and pentagonal pore frames.
276
Original remarks: P. danaense, n. sp., differs from P. riedeli
Pessagno in having: (1) less massive pore frames; (2) spinose
nodes at the pore frame vertices; and (3) less massive, thinner polar spines.
Measurements (µm):
Based on 6 specimens. System of measurement shown in
text-figure 5 of Pessagno & Blome (1980).
AA’
85
80
85
75
A’S’
90
79
90
65
AS
45
46
60
40
BB’
100
93
100
90
cc’
30
23
30
20
dd’
20
23
25
20
HT
Av.
Max.
Min.
Etymology: This species is named for Dana Inlet in its type
area.
Type locality: Sample QC 550, Sandilands Formation
(Kunga Formation in Pessagno & Blome, 1980), Queen
Charlotte Islands, British Columbia.
Occurrence: Sandilands, Ghost Creek and Fannin formations, Queen Charlotte Islands; Kirchstein Limestone,
Germany; Tawi Sadh Member of the Guwayza Formation,
Oman; Japan.
Plate PAN18. Pantanellium cumshewaense Pessagno & Blome. Magnification x250. Fig. 1(H). Pessagno & Blome 1980,
pl. 6, fig. 1. Fig. 2. QCI, GSC loc. C-304566, GSC 128857. Fig. 3. QCI, GSC loc. C-305417, GSC 111731.
Plate PAN11. Pantanellium danaense Pessagno & Blome. Magnification x300. Fig. 1(H). Pessagno & Blome 1980,
pl. 4, fig. 9. Fig. 2. OM, BR871-R09-01. Fig. 3. QCI, GSC loc. C-140495, GSC 128858. Fig. 4. QCI, GSC loc. C-305417,
GSC 128859. Fig. 5. QCI, GSC loc. C-140495, GSC 128860. Fig. 6. QCI, GSC loc. C-140495, GSC 111732.
277
Pantanellium inornatum Pessagno & Poisson 1981
Species code: PAN19
Synonymy:
1981 Pantanellium inornatum n. sp. – Pessagno & Poisson,
p. 56, pl. 6, figs. 1-9.
1981c Pantanellium inornatum Pessagno & Poisson –
De Wever, p. 144, pl. 5, fig. 2.
1982b Pantanellium inornatum Pessagno & Poisson –
De Wever, p. 128, pl. 1, fig. 8, 9.
1982 Pantanellium inornatum Pessagno & Poisson – De Wever
& Origlia-Devos, pl. 1, fig. N.
1982 Pantanellium sp. – Imoto et al., pl. 1, fig. 11.
1988 Pantanellium cf. browni Pessagno & Blome – Li, pl. 1, fig. 13.
1988 Pantanellium aff. inornatum Pessagno & Poisson – Li,
pl. 1, fig. 14.
1993 Pantanellium cf. kluense Pessagno & Blome – Kashiwagi &
Yao, pl. 1, fig. 13.
1995 Sphaerostylus inornatum (Pessagno & Poisson) – Suzuki,
pl. 8, fig. 7.
1996 Pantanellium sp. A – Pujana, p. 136, pl. 1, fig. 8.
1998 Pantanellium inornatum Pessagno & Poisson – Kashiwagi,
pl. 2, figs. 16, 17, 21.
1998 Pantanellium sp. aff. P. kungaense Pessagno & Blome –
Yeh & Cheng, p. 13, pl. 1, fig. 4.
1998 Pantanellium sp. cf. P. inornatum Pessagno & Poisson
– Yeh & Cheng, p. 13, pl. 5, fig. 5, not fig. 3.
1998 Pantanellium skedansense Pessagno & Blome – Yeh &
Cheng, p. 13, pl. 1, fig. 5.
Original description: Cortical shell, thin, spherical with
relatively slender triradiate bipolar spines; triradiate bipolar
spines with three rounded, narrow ridges alternating
with three narrow grooves. Meshwork of cortical shell
comprised of equal number of hexagonal and pentagonal
pore frames. Pentagonal pore frames slightly smaller than
hexagonal pore frames. Meshwork of first medullary shell
278
thick likewise comprised of hexagonal pentagonal pore
frames. Secondary radial beams between cortical shell and
first medullary shell circular in axial section.
Original remarks: Pantanellium inornatum Pessagno,
n. sp., differs from P. riedeli Pessagno (1977a) (1) by having
longer, slender polar spines with narrow ridges separated
by narrow grooves; (2) by having smaller, narrower ridges
separated by narrow grooves; (3) by having smaller, more
numerous pore frames; and (4) by having a thinner walled
cortical shell and a thicker walled first medullary shell.
Measurements (µm):
Based on 9 specimens. System of measurements after
Pessagno (1973).
A’S
110
110
85
AS
85
85
55
cc’
25
30
25
dd’
20
25
20
AA’
85
90
85
BB’
80
85
75
HT
Max.
Min.
Etymology: Inornatus-a-um (Latin, adj.): unadorned.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Ghost Creek
and Fannin formations, Queen Charlotte Islands; Sierra
Chacaicó Formation, Argentina; Dürrnberg Formation,
Austria; Haliw (Aqil) and Musallah formations, Oman;
Dengqen area, Tibet; Liminangcong Chert, Philippines;
Japan.
Plate PAN19. Pantanellium inornatum Pessagno & Poisson. Magnification x250. Fig. 1(H). Pessagno & Poisson 1981,
pl. 6, fig. 1. Fig. 2. QCI, GSC loc. C-304566, GSC 111733. Fig. 3. QCI, GSC loc. C-080611, GSC 128861. Fig. 4. QCI,
GSC loc. C-080611, GSC 128862. Fig. 5. QCI, GSC loc. C-080611, GSC 128863. Fig. 6. OM-00-115-023029. Fig. 7. OM,
Haliw-039-R03-06. Fig. 8. OM, Haliw-039-R06-21. Fig. 9. OM, Haliw-039-R02-17. Fig. 10. OM, Haliw-038-R08-11.
Fig. 11. AT, BMW 21-8. Fig. 12. JP, Ku(b)-11-24.
279
Pantanellium skedansense Pessagno & Blome 1980
Species code: PAN16
Synonymy:
1980 Pantanellium skedansense n. sp. – Pessagno & Blome,
p. 246, pl. 5, figs. 8, 9, 15, 20, 23.
1998 Pantanellium skedansense Pessagno & Blome – Whalen &
Carter, p. 49, pl. 1, fig. 12.
2002 Pantanellium skedansense Pessagno & Blome – Whalen &
Carter, p. 105, pl. 6, figs. 7, 8, 13, 14.
Original description: Cortical shell spherical to subspherical
with hexagonal and occasional pentagonal pore frames
having poorly developed nodes at their vertices. Bars of
pore frames thin in both Y and Z directions (text-fig. 5).
Five to 6 pore frames visible along AA’ and 6 along BB’. Polar
spines long, triradiate in axial section, one spine somewhat
longer than the other; some specimens with slightly curved
spinal tips. Both spines comprised of 3 broad longitudinal
grooves alternating with 3 ridges; grooves about 2 times as
wide as ridges. Grooves and ridges maintaining about same
width throughout.
Original remarks: Pantanellium skedansense, n. sp., appears
closely related to P. inornatum Pessagno and Poisson
(in press). It differs from the latter species primarily
in possessing longer polar spines with much broader
grooves. Pantanellium skedansense also closely resembles P.
sanrafaelense, n. sp., in terms of the distribution and size of
its pore frames and the structure of its primary spines; the
longer polar spine of both species usually possesses a curved
tip as well as wide grooves. Pantanellium skedansense differs
from P. sanrafaelense by lacking prominent, massive nodes
at pore frame vertices. It is suggested that P. skedansense
may be ancestral to P. sanrafaelense.
Measurements (µm):
Based on 5 specimens. System of measurement shown in
text-figure 5 of Pessagno & Blome (1980).
AA’
80
72
80
70
A’S’
115
104
115
85
AS
90
80
90
70
BB’
75
68
75
60
cc’
31
25
31
20
dd’
18
18
20
15
HT
Av.
Max.
Min.
Etymology: This species is named for Skedans Point north
of Kunga Island.
Genus: Parahsuum Yao 1982
Type species: Parahsuum simplum Yao 1982
Synonymy:
1982 Parahsuum n. gen. – Yao, p. 61.
1982 Lupherium n. gen. – Pessagno & Whalen, 1982, p. 135.
1986 Parahsuum Yao – Takemura, p. 47.
1987b Drulanta n. gen. – Yeh, p. 71.
1987b Fantus n. gen. – Yeh, p. 61.
1988 Parahsuum Yao – Hori & Yao, p. 49.
1990 Parahsuum Yao – Kozur & Mostler, p. 222.
Original description: Shell multisegmented, conical to
spindle-shaped lacking well-developed strictures. Cephalis
conical to dome-shaped, poreless with or without apical
horn. Thorax trapezoidal in outline with sparse irregulary
displaced pores. Abdomen and post-abdominal segments
with continuous edged costae. Single row of square pore
frames with circular, primary pores between costae.
Original remarks: Parahsuum differs from Hsuum Pessagno (1977a, p. 81) in having single row of pores between
costae, from Archaeodictyomitra Pessagno (1976, p. 49) in
having primary pores, and from Mita Pessagno (1977b,
p. 44) in having edged costae.
Further remarks: For the distinction between Parahsuum
Yao and Canutus Pessagno & Whalen we follow Kozur &
Mostler (1990) who wrote: “Some of the Canutus s.l. species, but not the type species Canutus tipperi Pessagno and
Whalen, 1982, belong also to Parahsuum Yao, 1982. There280
fore Canutus Pessagno and Whalen, 1982 is not synonymous with Parahsuum Yao, 1982. Canutus can be distinguished from Parahsuum by an outer layer that covers parts
or all of the inner pore frames with very large rectangular
pores. Moreover, typical Canutus have a spindle-shaped
test. More elongate subconical species of Canutus s.l., in
which the outer layer is reduced to nodes on the pore frame
vertices or directly superimposed on the inner pore frames
(no bars of the outer pore frame between the bars of the inner pore frames) as Canutus giganteus Pessagno and Whalen, 1982, C. indomitus Pessagno and Whalen, 1982 and C.
izeensis Pessagno and Whalen, 1982 are here placed into
Parahsuum Yao, 1982.”
Etymology: This genus is named according to the similarity
of the external shape with Hsuum Pessagno.
Included species:
PHS02 Parahsuum edenshawi (Carter) 1988
DRO05 Parahsuum fondrenense (Whalen & Carter) 1998
PHS09 Parahsuum formosum (Yeh) 1987b
2012 Parahsuum izeense (Pessagno & Whalen) 1982
PHS03 Parahsuum longiconicum Sashida 1988
PHS04 Parahsuum mostleri (Yeh) 1987b
PHS05 Parahsuum ovale Hori & Yao 1988
PHS01 Parahsuum simplum Yao 1982
PHS06 Parahsuum vizcainoense Whalen & Carter 2002
PHS07 Parahsuum? sp. A sensu Whalen & Carter 2002
Type locality: Sample QC 550, Sandilands Formation
(Kunga Formation in Pessagno & Blome, 1980), Queen
Charlotte Islands.
Occurrence: Sandilands and Ghost Creek formations,
Queen Charlotte Islands; Fernie Formation, Williston Lake,
northeastern British Columbia; San Hipólito Formation,
Baja California Sur.
Plate PAN16. Pantanellium skedansense Pessagno & Blome. Magnification x250. Fig. 1(H). Pessagno & Blome 1980,
pl. 5, fig. 8. Fig. 2. Whalen & Carter 2002, pl. 6, fig. 7. Fig. 3. NBC, GSC loc. C-305813, GSC 111734.
281
Parahsuum edenshawi (Carter) 1988
Species code: PHS02
1988 Drulanta edenshawi Carter n. sp. – Carter et al., p. 53,
pl. 2, fig. 5 only.
1990 Drulanta sp. cf. mostleri Yeh – De Wever et al., pl. 4, fig. 4.
? 1998 Parahsuum sp. – Kashiwagi, pl. 1, fig. 8.
2004 Parahsuum sp. – Matsuoka, fig. 210.
Original remarks: Drulanta edenshawi has a more variable
morphology than D. mostleri Yeh. It can have a shorter,
stouter test that is very rounded apically as illustrated by
the holotype (Pl. 2, fig. 5) as well as a more elongate test
illustrated by the paratype (Pl. 2, fig. 6). It is generally larger
than D. mostleri and has much coarser costae.
Original diagnosis: Test conical, rounded apically without
horn. 9 to 12 strong, rounded longitudinal costae visible
laterally between single rows of large circular to rectangular
pores. Test has a »welded« appearance.
Measurements (µm):
Based on 17 specimens.
Synonymy:
Original description: Test elongate, conical, rounded
apically without horn and usually with six or seven postabdominal chambers. Cephalis hemispherical, all other
chambers trapezoidal in outline. All but final one or
two chambers increasing gradually in height and width;
distal chambers on complete specimens slightly reduced
in width. Some tests very slightly constricted at joints.
Cephalis and thorax perforate, covered with a smooth
layer of microgranular silica. Two rows of thin, circular
to rectangular pore frames per chamber; pores circular
to subcircular. Continuous costae strong, and rounded;
superimposed on test between single longitudinal rows
of coarse pores; 9 to 12 costae visible laterally. Test has a
»welded« appearance.
Length
Maximum width
HT
296
148
Av.
290
149
Max.
350
170
Min.
210
112
Etymology: Named for Albert Edward Edenshaw, a prominent chief of the Haida Indians whose head village was at
Kiusta, later at Kung (both are located on the north coast of
Graham Island).
Type locality: GCS locality C-080577, Fannin Formation,
Creek locality, Maude Island.
Occurrence: Fannin Formation, Queen Charlotte Islands;
Haliw Formation, Tawi Sadh Member of the Guwayza
Formation, Musallah Formation, Oman; Japan.
Parahsuum fondrenense (Whalen & Carter) 1998
Species code: DRO05
Synonymy:
Further remarks: This species seems to be closely related to
Parahsuum formosum (Yeh) from which it differs, because
its vertical costae are not continuous throughout the test.
Original description: Test conical, finely costate, with approximately four to six postabdominal chambers. Cephalis
medium sized, dome-shaped, sparsely perforate, mostly
covered by layer of microgranular silica; cephalis with
short, delicate horn, circular in axial section. Thorax, abdomen, and postabdominal chambers trapezoidal in outline,
increasing gradually in width and height as added. Pore
frames of outer latticed layer on abdomen and postabdominal chambers square to rectangular, aligned in distinct
transverse and vertical rows, becoming larger as added.
First few postabdominal chambers with about 8 costae visible laterally; number of costae increasing to about 12 on
final postabdominal chamber.
Measurements (µm):
Based on 6 specimens.
1998 Droltus fondrenensis n. sp. – Whalen & Carter, p. 63, pl. 15,
figs. 9, 18.
Original remarks: The much more regularly aligned pore
frames of the outer latticed layer, distinguish Droltus
fondrenensis n. sp. from D. firmus n. sp.
282
Length (excluding horn)
195
225
150
191
Max. width
105
128
94
112
HT
Max.
Min.
Mean
Etymology: This species is named for Fondren Science
Building at the University of Texas at Dallas where much of
the systematic research for this paper was carried out.
Type locality: Sample QC-677, Sandilands Formation,
Kunga Island, Queen Charlotte Islands.
Occurrence: Sandilands Formation, Queen Charlotte
Islands.
Plate PHS02. Parahsuum edenshawi (Carter). Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 2, fig. 5. Fig. 2. QCI,
GSC loc. C-080611, GSC 128879. Fig. 3. QCI, GSC loc. C-304566, GSC 128880. Fig. 4. QCI, GSC loc. C-080612, GSC
111735. Fig. 5. OM, Haliw-039-R02-13. Fig. 6. OM, BR524-R05-15. Fig. 7. OM-00-251-021433.
Plate DRO05. Parahsuum fondrenense (Whalen & Carter). Magnification x250. Fig. 1(H). Carter et al. 1998, pl. 15,
fig. 9. Fig. 2. QCI, GSC loc. C-304281, GSC 128797. Fig. 3. QCI, GSC loc. C-175311, GSC 128798. Fig. 4. QCI, GSC loc.
C-175310, GSC 128799. Fig. 5. QCI, GSC loc. C-080611, GSC 128800.
283
Parahsuum formosum (Yeh) 1987b
Species code: PHS09
Synonymy:
1987b Drulanta formosa n. sp. – Yeh, p. 72, pl. 19, figs. 13-14.
2004 Droltus sp. – Matsuoka, fig. 204.
Measurements (µm):
Based on ten specimens.
HT
Mean
Max.
Min.
Max. length
200
205
210
194
Max. width
140
142
146
137
Original description: Test as with genus, costate throughout the test except cephalis and thorax. Costae massive,
about ten visible laterally. Cephalis relativelly small, conical.
Cephalis and thorax covered with layer of microgranular
silica. Meshwork of inner latticed layer consisting of square
to rectangular pore frames and increasing in size distally.
Chambers increasing in width rapidly and in length gradually as added.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Further remarks: See remarks under Parahsuum fondrenense (Whalen & Carter).
Occurrence: Hyde Formation, Oregon; Ghost Creek and
Fannin formations, Queen Charlotte Islands; Japan.
Etymology: Formosus-a-um (Latin, adj.) = beautiful.
Parahsuum izeense (Pessagno & Whalen) 1982
Species code: 2012
Synonymy:
1982 Canutus izeensis n. sp. – Pessagno & Whalen, p. 129, pl. 6,
figs. 8, 10, 15.
1982 Canutus giganteus n. sp. – Pessagno & Whalen, p. 127,
pl. 4, figs. 5, 13.
1987b Broctus izeensis (Pessagno & Whalen) – Yeh, pl. 4, fig. 29.
? 1987b Broctus (?) sp. A – Yeh, p. 54.
1988 Canutus giganteus Pessagno & Whalen – Carter et al.,
p. 50, pl. 3, fig. 1.
1988 Canutus izeensis Pessagno & Whalen – Carter et al., p. 51,
pl. 3, fig. 2.
1995a Parahsuum izeense (Pessagno & Whalen) – Baumgartner
et al., p. 378, pl. 2012, figs. 1-2.
1996 Canutus izeensis (Pessagno & Whalen) – Pujana, p. 138,
pl. 1, fig. 15.
1998 Canutus izeensis Pessagno & Whalen – Cordey, p. 104,
pl. 25, figs. 5, 10.
2003 Parahsuum izeense (Pessagno & Whalen) – Goričan et al.,
p. 296, pl. 5, figs. 18-19.
Original description: Test short, inflated, spindle-shaped,
usually with six post-abdominal chambers. Cephalis hemispherical, knoblike; remaining chambers trapezoidal in
cross section; cephalis and thorax usually imperforate. Abdomen and all but last two or three post-abdominal chambers increasing rapidly in width and gradually in length
as added; last two or three post-abdominal chambers decreasing somewhat in width. Inner latticed layer of postabdominal chambers consisting of moderately sized square
to rectangular pore frames with nodes at vertices; 15 rows
of pore frames visible laterally; three pore frames per row
occurring between two longitudinal ridges and joints of
chamber. Outer (second) latticed layer consisting of fragile,
284
irregular, polygonal pore frames. Outer latticed layer best
developed on earlier post-abdominal chambers.
Original remarks: Canutus izeensis, n. sp. differs from
C. tipperi, n. sp. by having a less inflated test, a hemispherical, knoblike cephalis, and considerably smaller pore frames
in the inner layer.
Further remarks: Parahsuum giganteum and P. izeense are
synonymized because they represent a continuum of test
shape from slender to broad.
Measurements (µm):
Based on 8 specimens.
Length
350.0
350.0
250.0
303.1
Width (max.)
200.0
200.0
150.0
175.0
HT
Max.
Min.
Mean
Etymology: This species is named for the village of Izee
near its type locality.
Type locality: OR-536, Nicely Formation, southeast side of
Morgan Mountain, east-central Oregon.
Occurrence: Nicely Formation, Oregon; Fannin Formation,
Queen Charlotte Islands; Bridge River Complex, British
Columbia; Franciscan Complex, California; Sierra Chacaicó
Formation, Argentina; Apennines, Italy; Skrile Formation,
Slovenia; Musallah Formation, Oman.
Plate PHS09. Parahsuum formosum (Yeh). Magnification x200. Fig. 1(H). Yeh 1987b, pl. 19, fig. 13. Fig. 2. JP, MNA-10,
MA13299. Fig. 3. QCI, GSC loc. C-175306, GSC 128881. Fig. 4. QCI, GSC loc. C-127898, GSC 111736.
Plate 2012. Parahsuum izeense (Pessagno & Whalen). Magnification x200. Fig. 1(H). Pessagno & Whalen 1982, pl. 6,
fig. 8. Fig. 2. Pessagno & Whalen 1982, pl. 4, fig. 5. Fig. 3. QCI, GSC loc. C-080577, GSC 128703. Fig. 4. Goričan et al.
2003, pl. 5, fig. 18. Fig. 5. OM-00-254-022034.
285
Parahsuum longiconicum Sashida 1988
Species code: PHS03
Synonymy:
1988 Parahsuum longiconicum n. sp. – Sashida, p. 20, pl. 2,
figs. 1-4, 16, 17.
1988 Parahsuum kanyoense n. sp. – Sashida, p. 21, pl. 1, figs. 14,
15, 20-24;
1990 Parahsuum aff. longiconicum Sashida – Hori, Fig. 8.14
1996 Parahsuum longiconicum Sashida – Tumanda et al., p. 178,
Fig. 4.2.
1997 Parahsuum sp. aff. P. longiconicum Sashida – Hori, pl. 1, fig.
24.
1998 Parahsuum longiconicum Sashida – Kashiwagi, pl. 1, fig. 4.
? 2001 Parahsuum cf. longiconicum Sashida – Kashiwagi, Fig. 6.3.
2003 Parahsuum longiconicum Sashida – Goričan et al., p. 296,
pl. 5, fig. 16.
2003 Parahsuum cf. longiconicum Sashida – Kashiwagi &
Kurimoto, pl. 3, figs. 7, 10.
2004 Parahsuum longiconicum Sashida – Hori, pl. 8,
figs. 49-50, 54.
2004 Parahsuum sp. – Hori, pl. 8, figs. 51-52, 55-57, ? fig. 59.
2004 Parahsuum kanyoense Sashida – Hori, pl. 8, fig. 53.
2004 Parahsuum longiconicum Sashida – Ishida et al., pl. 5,
figs. 3, 4.
Original diagnosis: Parahsuum of cone-shaped test with
long conical horn.
Original description: Conical test of medium width with a
massive conical horn on hemispherical cephalis. Cephalis
usually has wide and deep grooves at the base of conical
horn. Thorax and subsequent chambers trapezoidal in
outline, increasing gradually in width and length as added
except for final post-abdominal chamber. Complete
specimens with six to seven post-abdominal chambers.
Outer layer comprised continuous edged costae except for
apical region. Single row of square pore frames between
286
costae. Weak circumferential ridges present at joint part
of the first and second post-abdominal chambers. Ten to
eleven costae usually visible on side view of final postabdominal chamber.
Original remarks: This species closely resembles Parahsuum kanyoense, n. sp. in general shell shape. However, the
present new species has a longer conical horn and more
uniform frame work on the outer surface of shell.
Further remarks: Parahsuum longiconicum Sashida differs
from P. formosum (Yeh) by having linearly arranged costae
only on the distal half to two thirds of the test and by having
a much stronger horn.
Measurements (µm):
Based on 15 specimens.
Length
230
205
220
Max.
width
130
110
120
No. of postabdominal Length
chambers
of horn
7
50
Max.
6
25
Min.
6
40
Mean
Etymology: Latin, longus means long and conica, conical.
Type locality: Sample TAK-5, Takarazawa Valley, Itsukaichi area, Tokyo Prefecture, central Japan.
Occurrence: Japan; Liminangcong Chert, Philippines;
Haliw (Aqil) Formation and Tawi Sadh Member of the
Guwayza Formation, Oman; Skrile Formation, Slovenia;
Dürrnberg Formation, Austria; Fannin Formation, Queen
Charlotte Islands.
Plate PHS03. Parahsuum longiconicum Sashida. Magnification Figs. 1-5, 9, 11 x200 (scale bar A), Figs. 6-8, 10, 12-15
x300 (scale bar B). Fig. 1(H). Sashida 1988, pl. 2, fig. 1. Fig. 2. GSC loc. C-304568, GSC 128883. Fig. 3. GSC loc.
C-304568, GSC 128884. Fig. 4. GSC loc. C-080613, GSC 128885. Fig. 5. GSC loc. C-304568, GSC 111803. Fig. 6. JP,
MNA-10, MA12861. Fig. 7. JP, MNA-10, MA13012. Fig. 8. JP, IYII-11-69. Fig. 9. Hori 1990, Fig. 8-14. Fig. 10. Goričan
et al. 2003, pl. 5, fig. 16. Fig. 11. AT, BMW21-55. Fig. 12. OM, BR1121-R09-26. Fig. 13. OM, BR1122-R02-03.
Fig. 14. OM, BR1121-R08-01. Fig. 15. OM, Haliw-038-R08-32.
287
Parahsuum mostleri (Yeh) 1987b
Species code: PHS04
Synonymy:
1987b Drulanta mostleri n. sp. – Yeh, p. 72, pl. 18, Figs. 3-4, 21.
1987b Drulanta sp. cf. D. mostleri n. sp. – Yeh, p. 73, pl. 18, fig. 1.
1987b Drulanta mirifica n. sp. – Yeh, p. 72, pl. 4, fig. 8; pl. 18,
figs. 5, 7-8, 23.
1987b Drulanta sp. aff. D. mirifica n. sp. – Yeh, p. 72, pl. 3, fig. 17.
1987 Canutus aff. C. hainaensis – Hattori, pl. 15, fig. 11.
1988 Drulanta edenshawi Carter n. sp. – Carter et al., p. 53, pl. 2,
fig. 6 only.
1989 Drulanta sp. aff. D. pulchra Yeh – Hattori, pl. 12, fig. K.
1989 Drulanta sp. aff. D. mirifica Yeh – Hattori, pl. 12, fig. L.
1997 Parahsuum mirifica (Yeh) – Yao, pl. 14, fig. 648.
1998 Drulanta mirifica Yeh – Yeh & Cheng, p. 23, pl. 8, fig. 17,
pl. 9, fig. 19.
2002 Parahsuum mostleri (Yeh) – Whalen & Carter, p. 126,
pl. 15, figs. 4, 14.
2003 Parahsuum mostleri (Yeh) – Goričan et al., p. 296, pl. 5,
fig. 20.
Original description: Cephalis small, conical, usually with
seven to nine post-abdominal chambers. Cephalis domeshaped without horn. Thorax and subsequent chambers
trapezoidal in outline. Cephalis and thorax sparsely perforate, covered with layer of microgranular silica. Abdomen
and all post-abdominal chambers comprised of ten, longitudinal costae superimposed on each row of pore frames.
About nine to ten costae visible laterally. Pore frames medium in size. Chambers increasing gradually in width as
added with final post-abdominal chamber remaining in the
same width or decreasing slightly in width.
Original remarks: Drulanta mostleri, n. sp., differs from
D. mirifica, n. sp., by having a shorter test with smaller
cephalis.
Further remarks: Drulanta mostleri and Drulanta mirifica
are herein assigned to Parahsuum and synonymized because they represent variability amongst the population.
P. mostleri is more elongated and more pointed apically
than P. edenshawi (Carter).
Measurements (µm):
Ten specimens measured.
Max. length
221
218
222
216
Max. width
117
119
123
115
HT
Mean
Max.
Min.
Etymology: This species is named after Dr. H. Mostler in
honor of his studies on the Mesozoic Radiolaria.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Hyde Formation, Oregon; Fannin Formation,
Queen Charlotte Islands; San Hipólito Formation, Baja
Californian Sur; Skrile Formation, Slovenia; Dürrnberg
Formation, Austria; Liminangcong Chert, Philippines;
Japan.
Parahsuum ovale Hori & Yao 1988
Species code: PHS05
Synonymy:
1982 Parahsuum (?) sp. C – Yao, pl. 4, figs. 9-11.
1982 Parahsuum (?) sp. C – Yao et al., pl. 2, fig. 10.
1986 Parahsuum (?) sp. C – Hori, fig. 6.3.
1986 Parahsuum sp. C – Matsuoka, pl. 1, fig. 2.
1986 Parahsuum sp. C – Matsuoka & Yao, pl. 1, fig. 3.
1986 Parahsuum directiporata (Rüst) – Sato et al., fig. 17.11.
1986 Bagotum sp. A – Sashida et al., fig. 5.18.
1988 Parahsuum ovale n. sp. – Hori & Yao, p. 51, pl. 1, figs. 3a-e,
4a-c, 6-8, 9a, b.
1988 Parahsuum takarazawaense n. sp. – Sashida, p. 19, pl. 1,
figs. 6-13, 18, 19.
1990 Parahsuum ovale Hori & Yao – Hori, fig. 8.16.
1990 Parahsuum ovale Hori & Yao – Yao, pl. 2, fig. 2.
1992 Parahsuum takarazawaense Sashida – Sashida, pl. 1, fig. 7.
1993 Parahsuum ovale Hori & Yao – Kashiwagi & Yao, pl. 1, fig. 1.
1994 Parahsuum ovale Hori & Yao – Goričan, p. 79, pl. 17, fig. 13.
1997 Parahsuum ovale Hori & Yao – Hori, pl. 1, fig. 26.
1997 Parahsuum ovale Hori & Yao – Yao, pl. 14, fig. 654.
1998 Parahsuum ovale Hori & Yao – Kashiwagi, pl. 1, fig. 3;
pl. 2, fig. 20.
2003 Parahsuum takarazawaense Sashida – Kashiwagi &
Kurimoto, pl. 3, fig. 4.
2004 Parahsuum ovale Hori & Otsuka – Hori, pl. 2, figs. 1, 2;
?pl. 2, fig. 25.
288
2004 Parahsuum ovale Hori & Yao – Ziabrev et al., Fig. 5-5.
2005 Parahsuum ovale Hori & Otsuka – Hori, pl. 8, fig. 4.
Original description: Shell of 6 or more segments, oval,
without stricture. Cephalis poreless apically, flattened conical, without apical horn. Internal cephalic structure quite
indistinct. Apical surface of shell smooth, partially with
shallow irregular depressions and sparsely arranged pores.
Thorax and abdomen with small, circular pores arranged
irregularly and with polygonal pore frames. In some specimens, thorax and abdomen with regularly arranged pores
and with tetragonal pore frames. Post-abdominal segments
with 22-28 continuous longitudinal costae. Each of postabdominal segments with 2 or 3 transverse rows of pores
arranged tetragonally. Edged small nodes occurring at vertices of pore frames.
Original remarks: Parahsuum ovale sp. nov. differs from
other species of Parahsuum in having an oval shell and
being fairly flat in apical part. P. ovale is possibly co-specific with Stichocapsa directiporata Rüst. S. directiporata is
distinguished from P. ovale by having a small number (1618) of longitudinal costae.
Plate PHS04. Parahsuum mostleri (Yeh). Magnification x250. Fig. 1(H). Yeh 1987b, pl. 18, fig. 3. Fig. 2. AT, BMW21-16.
Fig. 3. Goričan et al. 2003, pl. 5, fig. 20. Fig. 4. Carter et al. 1988, pl. 2, fig. 6. Fig. 5. QCI, GSC loc. C-305388, GSC
128882. Fig. 6. Whalen & Carter 2002, pl. 15, fig. 4.
Plate PHS05. Parahsuum ovale Hori & Yao. Magnification x250. Fig. 1(H). Hori & Yao 1988, pl. 1, fig. 3a. Fig. 2. JP,
Ku(b)-11. Fig. 3. Hori 1990, Fig. 8-16. Fig. 4. OM, Haliw-038-R09-19. Fig. 5. OM, BR1121-R06-12. Fig. 6. OM-00-252021728.
289
Measurements (µm):
Based on 13 specimens.
Hight
211
225
246
211
Width
120
124
141
109
H/W
1.76
1.82
2.05
1.50
Type locality: Sample 38, Inuyama Section, Kiso River,
1.km NE of Unuma, Central Japan.
HT
Av.
Max.
Min.
Occurrence: Japan; Budva Zone, Montenegro; Haliw
(Aqil) Formation, Tawi Sadh Member of the Guwayza
Formation, Musallah Formation, Oman; Bainang Terrane,
Tibet.
Etymology: The name is derived from the Latin adjective
ovalis, meaning oval (egg-shaped).
Parahsuum simplum Yao 1982
Species code: PHS01
Synonymy:
1982 Parahsuum simplum n. sp. – Yao, p. 61, pl. 4, figs. 1-8.
1982 Parahsuum simplum Yao – Yao et al., pl. 2, fig. 9.
1982 Parahsuum simplum Yao – Imoto et al., pl. 1, figs. 1, 2.
1983 Parahsuum simplum Yao – Ishida, pl. 2, figs. 1-2.
1984 Lupherium? spp. – Whalen & Pessagno, pl. 4, figs. 8, 10, 11.
1986 Parahsuum simplum Yao – Matsuoka & Yao, pl. 1, fig. 2.
1987 Parahsuum simplum Yao – Goričan, p. 185, pl. 1, fig. 3.
1988 Parahsuum simplum Yao – Hori & Yao, p. 51, pl. 1, figs.1a-d.
1988 Parahsuum simplum Yao – Sashida, p. 19, pl. 1, figs. 1-5,
16, 17.
1990 Parahsuum simplum Yao – De Wever et al., pl. 4, fig. 1, not
fig. 9.
1990 Parahsuum simplum Yao – Hori, Fig. 8.15.
1990 Parahsuum simplum Yao – Kozur & Mostler, p. 222, pl. 17,
fig. 2.
1990 Parahsuum simplum Yao – Yao, pl. 2, fig. 1.
1992 Parahsuum simplum Yao – Sashida, pl. 1, figs, 1, 2, 5, 6, not
figs. 3, 4.
? 1992 Parahsuum simplum Yao – Sano et al., pl. 2, fig. A.
1994 Parahsuum simplum Yao – Goričan, p. 79, pl. 17,
figs. 9, 10, 12.
1996 Canutus sp. aff. C. hainaensis Pessagno & Whalen
– Pujana, p. 138, pl. 1, fig. 5.
1997 Parahsuum simplum Yao – Hori, pl. 1, fig. 25.
1997 Parahsuum simplum Yao – Sugiyama, p. 184, Fig. 28-8.
1998 Parahsuum simplum Yao – Whalen & Carter, 67, pl. 16,
ig. 6.
1998 Parahsuum simplum Yao – Kashiwagi, pl. 1, figs. 1, 2; pl. 2,
fig. 1.
1998 Parahsuum simplum Yao – Yeh & Cheng, p. 26, pl. 4,
fig. 14.
2002 Parahsuum simplum Yao – Whalen & Carter, p. 126, pl. 12,
figs. 3, 4, 12, 13; pl. 17, figs. 14, 15.
2002 Parahsuum sp. aff. P. simplum Yao – Whalen & Carter,
p. 126, pl. 12, fig. 5; pl. 17, figs. 12, 13.
2002 Parahsuum simplum Yao – Tekin, p. 189, pl. 4, fig. 3.
2003 Parahsuum simplum Yao – Kashiwagi & Kurimoto, pl. 3,
figs. 1-3.
290
2004 Parahsuum simplum Yao – Hori, pl. 1, fig. 51, pl. 2, fig. 24.
2004 Parahsuum simplum Yao – Hori et al., pl. 6, fig. 3.
2004 Parahsuum simplum Yao – Ishida et al., pl. 5, figs. 1, 2.
2005 Parahsuum simplum Yao – Hori, pl. 8, figs. 1-2.
Original description: Shell of 6 or more segments, elongate,
conical, becoming somewhat spindle-shaped in unbroken
or mature forms. Cephalis poreless, conical with short
apical horn. Internal cephalic structure quite indistinct.
Post-thoracic segments with continuous 24-32 costae.
Each of post-thoracic segments has 3 or 4 transverse rows
of pores arranged tetragonally. In some specimens, weak
circumferencial ridges present at joint part of segments.
Original remarks: Parahsuum simplum differs from Parahsuum (?) sp. A in lacking a long apical horn, and from
Parahsuum (?) sp. C in having a conical shell.
Further remarks: The considerable range in variation of
this species in overall size and width of distal chambers has
already been recognized by Hori & Yao (1988).
Measurements (µm):
Based on 27 specimens.
Height overall
Max. height of segment
Max. width of shell
Min.
161
28
85
Max.
293
39
129
Av.
216
33
104
Etymology: The name is derived from the Latin adjective
simplus, meaning simple.
Type locality: Sample 38, Inuyama Section, Kiso River,
1 km NE of Unuma, Central Japan.
Occurrence: Worldwide.
Plate PHS01. Parahsuum simplum Yao. Magnification x250. Fig. 1(H). Yao 1982, pl. 4, fig. 1. Fig. 2-3. Whalen & Carter
2002, pl. 12, figs. 3-4. Fig. 4. QCI, GSC loc. C-080613, GSC 111737. Fig. 5. QCI, GSC loc. C-080611 GSC 111738.
Fig. 6. OM, Haliw-039-R04-05. Fig. 7. OM, Haliw-039-R03-18. Fig. 8. OM, Haliw-039-R01-02. Fig. 9. OM, BR1121R08-09. Fig. 10. AT, BMW21-50.
291
Parahsuum vizcainoense Whalen & Carter 2002
Species code: PHS06
Synonymy:
1982 Lupherium sp. A – Pessagno & Whalen, p. 136, pl. 6, fig. 4.
1984 Lupherium sp. – Whalen & Pessagno, pl. 4, figs. 5-7.
1987b Lupherium sp. B – Yeh, p. 68, pl. 17, figs. 1, 4.
1987b Lupherium sp. C [Lupherium (?) sp. E in fig. captions]
– Yeh, p. 68, pl. 17, figs. 2, 3, 8.
1998 Drulanta sp. cf. D. mirifica Yeh – Yeh & Cheng, p. 23, pl. 4,
figs. 15, 16.
1998 Parahsuum sp. cf. P. officerense (Pessagno & Whalen)
– Yeh & Cheng, p. 26, pl. 8, fig. 6.
2002 Parahsuum vizcainoense n. sp. – Whalen & Carter, p. 126,
pl. 12, figs. 6, 10, 14.
Original description: Test elongate, conical to spindle
shaped, with six to seven post-abdominal chambers.
Large dome-shaped cephalis and thorax with delicate
horn, circular in cross section and often broken. Cephalis
and usually thorax and abdomen covered by a layer of
microgranular silica providing either a complete, smooth
coating or incomplete, irregular coating. Thorax, abdomen
and most post-abdominal chambers trapezoidal in outline.
Post-abdominal chambers gradually increasing in width
distally; final few chambers gradually decreasing in width;
all post-abdominal chambers gradually increasing in height.
Costae fine, closely spaced, not always perfectly aligned or
continuous. Pores circular to elliptical in outline.
Parahsuum? sp. A sensu Whalen & Carter 2002
Species code: PHS07
Synonymy:
? 1998 Drulanta sp. A – Yeh & Cheng, p. 23, pl. 8, fig. 11.
2002 Parahsuum? sp. A – Whalen & Carter, p. 128, pl. 16,
figs. 12, 13, 17.
Original remarks: The chambered test of this species is
similar to the genus but it has three, short bladed wing-like
extensions on distal post-abdominal chambers.
Occurrence: San Hipólito Formation, Baja California Sur.
292
Original remarks: The more delicate costae and strong
horn distinguish this species from Parahsuum simplum Yao.
In addition it differs from Parahsuum officerense (Pessagno
and Whalen 1982) in having a shorter test (fewer chambers)
and more irregular costae and meshwork. It is possible this
new species is the ancestor of P. officerense (Pessagno and
Whalen).
Measurements (µm):
Based on 8 specimens.
Length (excludes horn)
225
285
180
214
Width (Max.)
105
120
98
109
HT
Max.
Min.
Mean
Etymology: This species is named for the Vizcaino Peninsula, Baja California Sur.
Type locality: Sample BPW80-30, San Hipólito Formation,
Vizcaino Peninsula, Baja California Sur, Mexico.
Occurrence: San Hipólito Formation, Baja California
Sur; Franciscan Complex, California; Hyde Formation
and Warm Springs member of the Snowshoe Formation,
Oregon; Liminangcong Chert, Philippines; Japan; Skrile
Formation, Slovenia.
Plate PHS06. Parahsuum vizcainoense Whalen & Carter. Magnification x250, except Fig. 1b(H) x400.
Fig. 1(H).Whalen & Carter 2002, pl. 12, figs. 6, 10. Fig. 2. JP, IYII-3. Fig. 3. JP, MNA-10, MA12853. Fig. 4. SI, MM 6.76,
010302.
Plate PHS07. Parahsuum? sp. A sensu Whalen & Carter. Magnification x250, except Fig. 2b x400. Fig. 1. Whalen &
Carter 2002, pl. 16, fig. 12. Fig. 2. Whalen & Carter 2002, pl. 16, figs. 13, 17.
293
Genus: Parasaturnalis Kozur & Mostler 1972
Type species: Spongosaturnalis? diplocyclis Yao 1972
Synonymy:
1972 Parasaturnalis n. gen. – Kozur & Mostler, p. 43.
Original description: The double to triple ring encloses a
single row of pores and has two polar, or four to five radial
rods that connect to a spongy central shell. Moderately long
spines occur on the outer rim of the secondary ring.
Original remarks: In Heliosaturnalis n. gen. the inner ring
is firmly attached to the spongy shell. In Pseudosaturnalis
the secondary ring shows numerous pores.
Etymology: Composed of the prefix Para- and saturnalis.
Included species:
2013 Parasaturnalis diplocyclis (Yao) 1972
SAT15 Parasaturnalis yehae Dumitrica & Hori n. sp.
Parasaturnalis diplocyclis (Yao) 1972
Species code: 2013
Synonymy:
1972 Spongosaturnalis? diplocyclis n. sp. – Yao, p. 33, pl. 7,
figs. 6-10; pl. 8, figs. 1-2.
Not 1981c Japonisaturnalis diplocyclis (Yao) – De Wever, p. 141,
pl. 1, figs. 5, 7, 8.
Not 1982b Japonisaturnalis diplocyclis (Yao) – De Wever, p. 212213, pl. 13, fig. 9; pl. 14, figs. 1, 2.
1982 Parasaturnalis sp. – Wakita, pl. 4, fig. 13.
1987b Parasaturnalis vigrassi n. sp. – Yeh, p. 49, pl. 5, fig. 14;
pl. 23, fig. 11.
1987b Parasaturnalis sp. B – Yeh, p. 50, pl. 9, fig. 8.
1987 Parasaturnalis diplocyclis Yao group – Hattori, pl. 1,
figs. 7, 8, 9.
1989 Parasaturnalis diplocyclis Yao – Hattori, pl. 1, fig. 5; pl. 18,
fig. A.
1995a Parasaturnalis diplocyclis (Yao) – Baumgartner et al.,
p. 388, pl. 2013, figs. 1-3.
1996 Parasaturnalis sp. A – Yeh and Cheng, p. 108, pl. 2, fig. 6,
not pl. 7, fig. 8.
1997 Parasaturnalis diplocyclis (Yao) – Yao, pl. 5, fig. 206.
1997 Parasaturnalis sp. A – Yao, pl. 5, fig. 207.
2003 Parasaturnalis cf. diplocyclis (Yao) – Goričan et al., p. 291,
pl. 1, fig. 15.
2003 Spongosaturnalis (?) diplocyclis Yao – Kashiwagi &
Kurimoto, pl. 5, figs. 8, 9.
2004 Parasaturnalis sp. B sensu Yeh – Hori, pl. 2, fig. 20.
2004 Parasaturnalis diplocyclis (Yao) – Hori, pl. 4, fig. 40.
2004 Parasaturnalis diplocyclis (Yao) – Matsuoka, fig. 4.
Original description: Spongosaturnalid with double ring,
and with second spines on second ring. Shell not preserved, but fragmentary thorns on sturdy spines probably indicate that shell may be spongy. Polar spines a little
long or short, somewhat thin, with no ridge. Ring double,
first (inner) ring and second (outer) ring, joined by bars
(called as first spines), bilaterally symmetrical, circular to
subcircular, with smooth surface, and no ridge. First ring
curves smoothly, with no auxiliary spine on inner margin.
Second ring slightly waves with short wavelength, but in
some specimens curves smoothly. Thirteen or more first
spines (bars) on first ring, constant in size and shape, joining with second ring. Thirteen or more second spines on
second ring, situated respectively at middle point of part
joined with first spines, short, thornlike or low protrusive,
with rounded or somewhat sharp ends. Spaces enclosed by
294
both rings and first spines, elliptical or subrectangular. One
space at end of each polar spine generally larger than others, and in some specimens divided in two parts by transversal bar.
Original remarks: Although only eight specimens, which
are represented by fragmentary ring, were found, this
species is established because of the morphological feature
lacking auxiliary spines. ?S. catadelos, having a more
complicated ring, is described by Foreman (1968, p. 1112, pl. 1, figs. la-f; Latest Cretaceous, Moreno formation,
California). The ring of ?S. catadelos is broad and flat, and
perforated by numerous pores arranging in some measure
of regularity. It probably indicates that the complicated
ring is a combination of the fundamental rings (first ring,
second ring etc.) and spines (first spines, second spines
etc.). There is considerable variation in the number of
spines among specimens.
Measurements (µm):
Based on 6 specimens.
D. of 1 ring; (polar spines)
D. of 1 ring; (transversely)
D. of 2 ring; (polar spines)
D. of 2 ring; (transversely)
Diameter of shell
Length of polar spine
Length of first spine
Length of sec. spine
Breath of 1 ring
Breath of 2 ring
HT
203
260
313
325
126
23
18-25
23
15
8
Av.
180
250
285
302
113
21
15-25
11
9
7
Min.
150
190
230
270
100
13
10
3
3
3
Max.
203
315
330
350
126
30
35
25
15
13
Type locality: Sample IN-3, manganese carbonate ore,
Mino Belt, river side of the Kiso, east of Unuma, Kagamihara City, Gifu Prefecture, central Japan.
Occurrence: Japan; Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, Oregon;
Apennines, Italy; Skrile Formation, Slovenia; Tawi Sadh
Member of the Guwayza Formation, Oman; Liminangcong
Chert, Philippines.
Plate 2013. Parasaturnalis diplocyclis (Yao). Magnification x150. Fig. 1. Yao 1972, pl. 8, fig. 1. Fig. 2(H). Yao 1972,
pl. 8, fig. 2. Fig. 3. OM, BR528-R10-16. Fig. 4. Matsuoka 2004, fig. 4. Fig. 5. OM, BR871-R02-03. Fig. 6. OM, BR528R10-21. Fig. 7. OM, BR528-R10-15. Fig. 8. BR706-R13-11. Fig. 9. OM, BR706-R13-10. Fig. 10. OM, BR871-R02-04.
295
Parasaturnalis yehae Dumitrica & Hori n. sp.
Species code: SAT15
Synonymy:
1981c Japonisaturnalis diplocyclis (Yao) – De Wever, p. 141, pl. 1,
fig. 5, ? figs. 7, 8.
1982b Japonisaturnalis diplocyclis (Yao) – De Wever, p. 212-213,
pl. 13, fig. 9; pl. 14, ?figs. 1, 2.
1987b Parasaturnalis sp. A – Yeh, p. 49, pl. 3, fig. 16.
1992 Parasaturnalis spp. – Pessagno & Mizutani, pl. 99,
figs. 10, 15.
1998 Heliosaturnalis sp. A – Yeh & Cheng, p. 16, pl. 2, fig. 20;
pl. 8, fig. 9; pl. 11, fig. 11.
Type designation: The holotype was illustrated by Yeh
(1987b, pl. 3, fig. 16) from the Nicely Formation, Oregon.
Diagnosis: A species of Parasaturnalis having the spines of
secondary ring aligned with the radial bars connecting the
two rings.
Description: Ring double, subcircular to slightly elliptical
in outline, thin, smooth. Ring commonly with 12-14 radial
bars between the two rings and a similar number of short
spines on the secondary ring aligned with the radial bars
connecting the two rings. Shell spongy, many-layered,
when larger it may be connected to primary ring also by
auxiliary and/or subsidiary rays.
Remarks: This new species is clearly different from
Parasaturnalis diplocyclis (Yao) by the position of the
spines on the secondary ring relative to the radial bars
connecting the two rings. Whereas in P. diplocyclis the bars
have an alternate position, in P. yehae they are aligned with
these bars. The ring of P. yehae is comparable to ring of the
Carnian species Heliosaturnalis magnus Kozur & Mostler.
In the present state of knowledge we include in this species
specimens with and without auxiliary or subsidiary rays.
296
The fossil record seems to prove that the latter are stratigraphically younger (late Pliensbachian - Toarcian) (Yeh,
1987b) than the former (Sinemurian - early Pliensbachian)
(De Wever, 1981c, 1982b; Yeh & Cheng, 1998) and that the
disappearance of these centripetally directed radial bars,
is an evolutionary process inherited from the Late Triassic and earliest Jurassic ancestors. If so, these species could
be separated into two successive subspecies of evolutionary and stratigraphic value i.e., Parasaturnalis yehae ssp. A
and Parasaturnalis yehae yehae. In the present paper the
existance of these two morphotypes is just suggested.
Measurements (µm):
Based on 6 specimens.
Diameter of central shell
Width of primary ring at the end of polar rays
Width of secondary ring
Length of primary ring
Length of secondary ring
HT
227
327
289
360
Min. Max.
108 120
239 305
365 407
-
Etymology: The species is named after Kuei-Yu Yeh, Tainan
National College of the Arts, Tainan, Taiwan, who illustrated several specimens of this species.
Type locality: Sample OR-536J, Nicely Formation, Morgan
Mountain, east-central Oregon (Yeh, 1987b).
Occurrence: Nicely Formation, Oregon; Inuyama and Nanjo areas, Japan; Busuanga Island, Philippines; Fernie Formation, northeastern British Columbia; Gümüslü Allochthon,
Turkey; Tawi Sadh Member of the Guwayza Formation and
Haliw (Aqil) Formation, Oman.
Plate SAT15. Parasaturnalis yehae Dumitrica & Hori n. sp. Magnification x150. Fig. 1(H). Yeh 1987b, pl. 3, fig. 16.
Fig. 2. OM, BR528-R10-17. Fig. 3. NBC, GSC loc. C-305208, GSC 111739. Fig. 4. OM, BR706-R13-12. Fig. 5. JP, Nanjo
chert, NAI-132, RH(1)468.
297
Genus: Paronaella Pessagno 1971
Type species: Paronaella solanoensis Pessagno 1971
Synonymy:
1971 Paronaella n. gen. – Pessagno, p. 46.
1971 Patulibracchium n. gen. – Pessagno, p. 26.
1980 Paronaella Pessagno emend. – Baumgartner, p. 300.
1981b Paronaella Pessagno emend. – De Wever, p. 33.
1987b Sontonaella n. gen. – Yeh, p. 44.
1993 Paronaella Pessagno emend. Baumgartner – Pessagno et
al., p. 121.
1994 Fluegelium n. gen. – Steiger & Steiger, p. 457.
1999 Paronaella Pessagno, emend. Baumgartner, emend. De
Wever – Kiessling, p. 38.
Original description: Test lacks rays with bracchiopyle.
Rays always nearly equal in length; expanded or thickened
ray tips lacking. Meshwork linear to sublinear; comprised
of irregular polygonal pore frames. Pore frames comprised
of bars connected to weakly developed nodes.
Original remarks: Paronaella n. gen. differs from Patulibracchium n. gen. and Halesium n. gen. by lacking a bracchiopyle and expanded ray tips and by always having rays
which are nearly equal in length.
Further remarks: We include all forms with or without a
bracchiopyle and with or without bulbous tips.
Etymology: This genus is named for C. F. Parona, one of the
early students of Mesozoic Radiolaria.
Included species and subspecies:
PAR13 Paronaella corpulenta De Wever 1981b
PAR22 Paronaella curticrassa Carter & Dumitrica n. sp.
PAR24 Paronaella fera s.l. (Yeh) 1987b
PAR15 Paronaella fera fera (Yeh) 1987b
PAR10 Paronaella fera jamesi Whalen & Carter 1998
PAR16 Paronaella grahamensis Carter 1988
PAR17 Paronaella notabilis Whalen & Carter 2002
2005 Paronaella skowkonaensis Carter 1988
PAR19 Paronaella snowshoensis (Yeh) 1987b
PAR20 Paronaella tripla De Wever 1981b
PAR21 Paronaella variabilis Carter 1988
Paronaella corpulenta De Wever 1981b
Species code: PAR13
Synonymy:
1981b Paronaella corpulenta n. sp. – De Wever, p. 33, pl. 2,
figs. 7-9.
1982b Paronaella corpulenta De Wever – De Wever, p. 245,
pl. 22, fig. 7; pl. 23, figs. 1-3.
1988 Paronaella sp. C – Carter et al., p. 42, pl. 11, fig. 7.
2002 Paronaella corpulenta De Wever – Whalen & Carter,
p. 107, pl. 2, figs. 6, 12.
2003 Paronaella spp. – Goričan et al., p. 295, pl. 2, fig. 4 only.
? 2004 Paronaella corpulenta De Wever s.l. – Matsuoka, fig. 32.
Original description: Massive form with three very wide,
club-shaped arms terminating in a short, robust triradiate
spine.
Arms, almost as wide as long, composed of a spongy
network which is embedded within other more delicate
spongy material that usually remains only as fragments.
This secondary spongy material is responsible for the plump
shape of this species. Primary spines at distal ends of arms
seem to be deep-set as in a cushion. Some forms possess
secondary spines between or on arms; these spines are
always thinner than primary spines, although sometimes
longer.
Original remarks: This species differs from Paronaella
obesa (Pessagno) by absence of bracchiopyle, more massive
shape, finer less frequent secondary spines and presence
298
of a patagium. Its plump shape, very rounded outline and
primary spine (which appears to be deep-set as in a cushion)
easily distinguishes this species from others. It differs from
Paronaella tripla n. sp. by the presence of secondary spines
and, mainly by its different network.
Measurements (µm):
Based on 10 specimens.
Length of ray
HT
185
Min.
167
Max.
200
Av.
177
Length of primary spines is difficult to measure since they
arise from the depressed part of a “cushion”; they are generally about 50µm long.
Etymology: From Latin corpulentus, -a, -um, adj. = corpulent, by analogy with the rotund shape of this species.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Ghost Creek,
Fannin, Whiteaves and Phantom Creek formations, Queen
Charlotte Islands; San Hipólito Formation, Baja California
Sur; Skrile Formation, Slovenia; Tawi Sadh Member of the
Guwayza Formation, Oman.
Plate PAR13. Paronaella corpulenta De Wever. Magnification x150. Fig. 1(H). De Wever 1981b, pl. 2, fig. 7.
Fig. 2. QCI, GSC loc. C-175311, GSC 128864. Fig. 3. QCI, GSC loc. C-175311, GSC 128865. Fig. 4. QCI, GSC loc.
C-080612, GSC 128866. Fig. 5. QCI, GSC loc. C-080612, GSC 128867. Fig. 6. QCI, GSC loc. C-080612, GSC 128868.
Fig. 7. OM, BR1123-R05-01. Fig. 8. OM, BR706-R03-21a. Fig. 9. Goričan et al. 2003, pl. 2, fig. 4.
299
Paronaella curticrassa Carter & Dumitrica n. sp.
Species code: PAR22
Synonymy:
1987b Sontonella sp. B – Yeh, p. 47, pl. 11, fig. 10.
1988 Paronaella sp. A – Carter et al., p. 42, pl. 4, fig. 10.
Type designation: Holotype, pl. PAR22, fig. 2, Carter et al.
1988, pl. 4, fig. 10 (GSC 80574); paratype, fig. 1, QCI, GSC
loc. C-177371, GSC 128875.
Description: Test small with a large slightly raised central
area and three short stubby rays widely expanded and clubshaped at tips; each ray with a single short spine. Meshwork
of test usually massive on central area and proximal part of
arms, pore frames triangular to quadrangular or irregularly
polygonal with nodes at vertices; meshwork slightly smaller
towards ray tips. Lateral sides of rays vertical. Spines on ray
tips triradiate at base then circular in axial section.
Remarks: Paronaella curticrassa n. sp. differs from P. grahamensis Carter in having a larger central area and shorter
rays with more widely expanded ray tips. The holotype
300
appears to be morphologically intermediate between the
two other specimens illustrated.
Measurements (µm):
Based on 4 specimens.
HT
Min. Max. Av.
Length of ray from shell centre 135-145 73
160 117
Minimum width of ray
50
36
85
58
120-130 65
170 111
Width of expanded tip
Etymology: From the Latin curtus – short and crassus – fat,
plump; adjective.
Type locality: GSC loc. C-080577, Maude Island, Queen
Charlotte Islands, British Columbia.
Occurrence: Fannin Formation, Queen Charlotte Islands;
Hyde Formation and Warm Springs member of the
Snowshoe Formation, Oregon; Tawi Sadh Member of the
Guwayza Formation, Oman.
Plate PAR22. Paronaella curticrassa Carter & Dumitrica n. sp. Magnification x200. Fig. 1. QCI, GSC loc. C-177371,
GSC 128875. Fig. 2(H). Carter et al. 1988, pl. 4, fig. 10. Fig. 3. OR600A-R03-13.
301
Paronaella fera s.l. (Yeh) 1987b
Species code: PAR24
Synonymy:
1987b Sontonella fera n. sp. – Yeh, p. 46, pl. 1, fig. 3; pl. 3, fig. 20.
See also subspecies.
Included subspecies:
PAR15 Paronaella fera fera (Yeh) 1987b
PAR10 Paronaella fera jamesi Whalen & Carter 1998
Paronaella fera fera (Yeh) 1987b
Species code: PAR15
Synonymy:
1987b Sontonaella fera n. sp. – Yeh, p. 46, pl. 1, fig. 3; pl. 3, fig. 20.
1987b Sontonaella sp. aff. S. fera n. sp. – Yeh, p. 46, pl. 1, fig. 4.
Original description: Rays nearly equal in length and
width, with large expanded tips. One tip larger than other
two and flanking with two short lateral spines at distal surface. The other two tips with long, massive central spines.
All spines circular in cross section. Test mainly with triangular and tetragonal pore frames having prominent nodes
at vertices. Pore frames irregularly arranged, medium in
size, with smaller pore frames on tips.
Original remarks: Sontonaella fera Yeh, n. sp., can be easily
distinguished from other Sontonaella spp. in this report by
having one large tip with two small spines and two smaller
tips with extremely long and massive spines.
302
Measurements (µm):
Ten specimens measured. System of measurement shown
in text-figure 7 of Yeh (1987b).
HT
Mean
Max.
Min.
LA
186
177
186
160
LB
186
177
186
160
LC
186
177
186
160
WR
50
56
60
50
LT
86
77
86
70
WT
143
140
143
135
Etymology: Ferus-a-um (Latin, adj.) = wild.
Type locality: Sample OR-536D, Nicely Formation, southeast side of the Morgan Mountain, east-central Oregon.
Occurrence: Nicely Formation, Oregon; Ghost Creek and
Fannin formations, Queen Charlotte Islands and Fernie
Formation, Williston Lake, British Columbia; Tawi Sadh
Member of the Guwayza Formation.
Plate PAR15. Paronaella fera fera (Yeh). Magnification x150. Fig. 1(H). Yeh 1987b, pl. 1, fig. 3. Fig. 2. QCI, GSC loc. C304568, GSC 128869. Fig. 3. NBC, GSC loc. C-305208, GSC 128870. Fig. 4. NBC, GSC loc. C-305208, GSC 128871.
Fig. 5. QCI, GSC loc. C-080612, GSC 128872. Fig. 6. OM, BR528-R10-04.
303
Paronaella fera jamesi Whalen & Carter 1998
Species code: PAR10
Synonymy:
section and spines are shorter and slightly flat with single
central spine on ray tips. Several forms originally assigned
to P. jamesi Whalen & Carter have now been assigned to
P. grahamensis Carter.
Original description: Test with short equally spaced rays;
rays approximately equal in length, narrow proximally,
gradually expanding in width distally. Ray tips moderately
expanded and bulbous. Meshwork composed mostly of
strong, triangular and tetragonal pore frames with small
nodes at pore frame vertices; slight lineation of pore frames
developed externally on proximal portion of rays. Each ray
usually with short, circular, flat-bladed to slightly triradiate
central spine; some rays with several smaller spines rather
than one primary spine.
Measurements (µm):
Based on 8 specimens.
1998 Paronaella jamesi n. sp. – Whalen & Carter, p. 51, pl. 13,
figs. 19, 23, not figs. 18, 22, 24.
2002 Paronaella jamesi Whalen & Carter – Tekin, p. 181, pl. 1,
fig. 11.
Original remarks: See remarks under Paronaella skenaensis
n. sp.
Remarks under P. skenaensis Whalen & Carter n. sp. (p.
52): Paronaella skenaensis n. sp. differs from P. jamesi n. sp.
in having much finer meshwork with smaller nodes and
more delicate spines.
Further remarks: This species differs from Paronaella fera
fera (Yeh) because the rays are not rectangular in cross-
Length
of longest ray
174
249
146
184
Average width
of rays at base
53
66
50
59
Max. width
of ray tips
121
184
113
139
HT
Max.
Min.
Mean
Etymology: This species is named for James Helwig, Dallas, Texas who assisted with fieldwork and construction of
plates.
Type locality: Sample 89-CNA-KUH-8, Sandilands Formation, north side of Kunga Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands; Hocaköy Radiolarite, Turkey.
Paronaella grahamensis Carter 1988
Species code: PAR16
Synonymy:
1987 Paronaella (?) sp. V – Hattori, pl. 4, fig. 12.
1988 Paronaella grahamensis Carter n. sp. – Carter et al., p. 40,
pl. 11, figs. 11, 12; not pl. 11, fig. 10.
1998 Paronaella jamesi n. sp. – Whalen & Carter, p. 51, pl. 13,
figs. 18, 22, 24, not figs. 19, 23.
2002 Paronaella grahamensis Carter – Whalen & Carter, p. 107,
pl. 2, figs. 3, 4, 9, 11, 13.
2001 Paronaella grahamensis Carter – Gawlick et al., pl. 2,
fig. 17.
2004 Paronaella sp. – Matsuoka, fig. 30.
Original diagnosis: Moderate in size with short rays,
expanded tips and slender, almost cylindrical central spine.
Rays subrectangular in cross-section.
Original description: Three-rayed patulibracchiid of
moderate size with slender central spine. Rays short,
approximately equal in length. Tips enlarged and rounded;
expansion may occur gradually throughout ray length or
more abruptly in distal portion only. Pore frames irregular
to sublinearly aligned, uniform in size, mostly tetragonal.
Central spines on ray tips are slender, variable in length,
circular in section. Rays subrectangular in cross-section.
304
Original remarks: This abundant form is similar to, but
much smaller than, Paronaella bona Yeh. Strong central
spines are evident on almost all specimens.
Measurements (µm):
Based on 7 specimens.
Length of longest ray
Width of ray
Width of ray tip
Length of longest spine
HT
171
47
118
65
Av.
167
60
122
58
Max.
200
70
140
130
Min.
125
47
102
35
Etymology: Named for Graham Island; type locality in central portion of island.
Type locality: GSC locality C-080583, Phantom Creek Formation, Graham Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Fannin, Whiteaves and Phantom Creek formations, Queen Charlotte Islands; Fernie Formation, NE
British Columbia; San Hipólito Formation, Baja California
Sur; Skrile Formation, Slovenia; Dürrnberg Formation,
Austria; Japan.
Plate PAR10. Paronaella fera jamesi Whalen & Carter. Magnification x150. Fig. 1(H). Carter et al. 1998, pl. 13, fig. 23.
Fig. 2. Carter et al. 1998, pl. 13, fig. 19. Fig. 3. QCI, GSC loc. C-305417, GSC 128873.
Plate PAR16. Paronaella grahamensis Carter. Magnification x150. Fig. 1(H). Carter et al. 1988, pl. 11, fig. 11.
Fig. 2. QCI, GSC loc. C-304568, GSC 128874. Fig. 3. JP, MNA-10, MA10845. Fig. 4. NBC, GSC loc. 305208, GSC 111740.
Fig. 5. Whalen & Carter 2002, pl. 2, fig. 4. Fig. 6. Whalen & Carter 2002, pl. 2, fig. 3. Fig. 7. SI, MM 5.00, 010106.
305
Paronaella notabilis Whalen & Carter 2002
Species code: PAR17
Synonymy:
1988 Paronaella sp. D – Carter et al., p. 42, pl. 11, fig. 9.
2002 Paronaella notabilis n. sp. – Whalen & Carter, p. 107, pl. 2,
figs. 7, 10, 14; pl. 3, figs. 1, 8, 11.
2002 Paronaella notabilis Whalen & Carter – Tekin, p. 181, pl. 1,
figs. 12, 13.
Original description: Test with three stout rays; ray length
slightly greater than ray width; distal part of ray somewhat
inflated to bulbous. Each ray with small, tapering principal spine, circular in axial section. Small and mediumsized subsidiary spines sometimes present on ray tips.
Meshwork composed of irregularly shaped tetragonal and
pentagonal pore frames with slight development of nodes
at pore frame vertices; no apparent development of pore
frame lineation.
Original remarks: The slightly more elongate, less bulbous
rays, along with the presence of subsidiary spines on
the ray tips, distinguish Paronaella notabilis n. sp. from
P. corpulenta De Wever 1981b.
Measurements (µm):
Based on 11 specimens.
Length of ray
90
105
83
93
Length of spine
53
75
45
55
HT
Max.
Min.
Mean
Etymology: Notabilis (Latin, adj.) = remarkable, striking,
noteworthy.
Type locality: Sample BPW80-30, San Hipólito Formation,
Vizcaino Peninsula, Baja California Sur, Mexico.
Occurrence: Phantom Creek and Graham Island formations, Queen Charlotte Islands; San Hipólito Formation,
Baja California Sur; Hocaköy Radiolarite, Turkey.
Paronaella skowkonaensis Carter 1988
Species code: 2005
Synonymy:
1987 Paronaella sp. O – Hattori, pl. 4, fig. 14.
1988 Paronaella sp. F – Hattori, pl. 6, fig. D.
1989 Homoeoparonaella sp. – Hattori & Sakamoto, pl. 6, fig. J.
1988 Paronaella skowkonaensis Carter n. sp. – Carter et al.,
p. 40, pl. 11, figs. 4-5.
1989 Tritrabs (?) spp. – Hori & Otsuka, pl. 4, fig. 7, not fig. 6.
1989 Paronaella (?) sp. – Hori & Otsuka, pl. 4, fig. 8.
1995a Paronaella skowkonaensis Carter – Baumgartner et al.,
p. 398, pl. 2005, figs. 1-2.
2004 Homoeoparonaella sp. – Matsuoka, fig. 35.
Original diagnosis: Three-rayed patulibracchiid having
long, slender rays with clavate to wedge-shaped tips. Meshwork fine and irregular. Ray tips have numerous short fine
spines.
Original description: Test large with three long slender rays
expanding at tips. Rays subequal in length at approximately
120°. Tips rounded to wedge-shaped. External pore frames
small, sublinearly arranged; tetragonal to pentagonal
with weak nodes at vertices. Numerous short, fine spines
extend from ray tips of well preserved specimens. Internal
meshwork layered and spongy.
306
Original remarks: This form strongly resembles Rhopalastrum trixiphus Rüst 1898, but differs in having several
short spines rather than a single central one on each ray
tip. It has been assigned to Paronaella because of its layered spongy meshwork.
Measurements (µm):
Based on 10 specimens.
Length of ray AX:
Length of ray BX:
Length of ray CX:
Width of ray:
Width of tip:
HT
196
188
182
50
149
Av.
197
81
146
Min.
230
70
205
Max.
150
50
80
Etymology: Named for Skowkona Mountain, southeast of
the type locality.
Type locality: GSC locality C-080584, Phantom Creek
Formation, Yakoun River, Graham Island, Queen Charlotte
Island, British Columbia.
Occurrence: Fannin, Whiteaves and Phantom Creek
formations, Queen Charlotte Islands; Apennines, Italy;
Japan.
Plate PAR17. Paronaella notabilis Whalen & Carter. Magnification x200. Fig. 1(H). Whalen & Carter 2002, pl. 3,
fig. 1. Fig. 2. Whalen & Carter 2002, pl. 2, fig. 7.
Plate 2005. Paronaella skowkonaensis Carter. Magnification x150. Fig. 1(H). Carter et al. 1988, pl. 11, fig. 4.
Fig. 2. Matsuoka 2004, fig. 35. Fig. 3. Hori & Otsuka 1989, pl. 4, fig. 8.
307
Paronaella snowshoensis (Yeh) 1987b
Species code: PAR19
Synonymy:
1987b Sontonaella snowshoensis n. sp. – Yeh, p. 46, pl. 21, fig. 3;
pl. 22, figs. 9, 14.
2002 Paronaella snowshoensis (Yeh) – Whalen & Carter, p. 107,
pl. 3, figs. 2, 9, 12, 14.
Original description: Test large with three rays nearly
equal in length. Rays long, moderate in width, subcircular
in outline, terminating in large rhombohedral tips with a
medium length triradiate spines originating from central
portion of tips. Test comprised of nearly uniformly sized
sublinearly arranged triangular pore frames on ray shafts
and more irregularly arranged polygonal pore frames on
tips. Pore frames larger on ray shafts and smaller in central
area, all pore frames with nodes at vertices.
Original remarks: Sontonaella snowshoensis, n. sp., differs
from S. bona, n. sp., by having rays consisting chiefly
of triangular pore frames rather than tetragonal pore
frames, and by having three ray shafts with three large
308
rhombohedral tips rather than with one rhombohedral tip
and two ellipsoidal tips.
Measurements (µm):
Ten specimens measured. System of measurement
shown in text-figure 7 of Yeh (1987b).
HT
Mean
Max.
Min.
LA
315
320
355
300
LB
315
320
355
300
LC
315
320
355
300
WR
63
70
80
63
LT
125
115
125
100
WT
200
210
220
200
LSP
150
152
155
150
Etymology: This species is named for the Snowshoe Creek
near its type locality.
Type locality: OR-589D, Warm Springs member, Snowshoe
Formation, east-central Oregon.
Occurrence: Hyde Formation and Warm Springs member
of the Snowshoe Formation, Oregon; San Hipólito Formation, Baja California Sur; Japan.
Plate PAR19. Paronaella snowshoensis (Yeh). Magnification x 150. Fig. 1(H). Yeh 1987b, pl. 22, fig. 9.
Fig. 2. JP, MNA-10, MA10797. Fig. 3. BCS, Loc. BPW80-30. Fig. 4. Whalen & Carter 2002, pl. 3, fig. 2.
309
Paronaella tripla De Wever 1981b
Species code: PAR20
Synonymy:
1981b Paronaella tripla n. sp. – De Wever, p. 34, pl. 3, figs. 5, 6.
1982b Paronaella tripla De Wever – De Wever, p. 248, pl. 25,
figs. 3-4.
1988 Paronaella sp. B – Carter et al., p. 42, pl. 11, fig. 6.
Original description: Form with three massive arms each
terminating in a short primary spine. Arms wide at base
increasing more in width distally. Five to six longitudinal
beams clearly visible on arms, connected to oblique
bars framing triangular pores. Nodes, sometimes well
developed, present at beam-bar intersections. Fine, loose
spongy material visible on well preserved specimens on all
test surfaces, especially on general plane of flattening.
Original remarks: This species differs from Paronaella
elegans by its much more massive shape, wider arms,
thinner beams and bars and more abrupt brachial end. It
is distinguished from Paronaella petroleumensis (Pessagno)
1971 by the absence of a bracchiopyle and its less inflated
central part, and from P. californiensis (Pessagno) 1971
310
by the shape of its arms and absence of a bracchiopyle;
Paronaella kotura Baumgartner (1980) is thinner with
slimmer arms.
Measurements (µm):
Based on 6 specimens.
Length of arm, from center
to base of terminal spine
Width of arm
(before inflated part)
Mean
Min.
Max.
HT
196
157
220
220
107
100
120
120
Etymology: From latin triplus, -a, -um, adj. = triple. This
species has three arms, three primary spines, and pores
which are triangular most of the time.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Dürrnberg
Formation, Austria; Phantom Creek Formation, Queen
Charlotte Islands.
Plate PAR20. Paronaella tripla De Wever. Magnification x150. Fig. 1(H). De Wever 1981b, pl. 3, fig. 5.
Fig. 2. TR, 1662D-R08-05. Fig. 3. Carter et al. 1988, pl. 11, fig. 6. Fig. 4. AT, BMW21-29.
311
Paronaella variabilis Carter 1988
Species code: PAR21
Synonymy:
1988 Paronaella variabilis Carter n. sp. – Carter et al., p. 41,
pl. 11, figs. 1-3.
2002 Paronaella variabilis Carter – Whalen & Carter, p. 107,
pl. 3, figs. 3, 4, 7, 10.
2004 Paronaella sp. cf. P. bona (Yeh) – Ziabrev et al., fig. 5-7.
Original diagnosis: Three-rayed patulibracchiid of variable
morphology. Rays generally slender with greatly expanded
elliptical, wedge or club-shaped tips. Mesh size irregular,
coarse to medium, finer on ray tips. Rays terminate in numerous small spines.
Original description: Three-rayed form variable in many
respects. Rays moderately slender, expanding to large elliptical, club, or wedge-shaped tips. The largest interradial
angle varies continuously from 120º˙to almost 150º (n=81).
Pore frames irregularly arranged on most specimens, sublinear on others; always smaller on ray tips. Pore frames
triangular and tetragonal, nodes highly developed. Rays
cylindrical to subrectangular in cross-section.
n=81
Maximum angle
Minimum angle
Mean
128.41º
111.99º
Standard deviation
5.88
5.93
Original remarks: A variable form extremely abundant
in all upper Toarcian samples. Resembles Paronaella sp.
cf. P. kotura figured by Baumgartner (1980, p. 304, Pl. 9,
fig. 14), but lacks fine meshwork in central area and has
numerous fine spines on ray tips. It differs from P. kotura
by having highly variable interradial angles, shorter, wider
rays with more expanded tips, and by lacking small pores
in the central area.
Measurements (µm):
Based on 10 specimens.
Length of ray AX
BX
CX
Width of ray
Width of tip
HT
196
188
182
50
149
Av.
Max.
Min.
197
230
150
81
146
70
205
50
80
Etymology: Latin, variabilis (adj.), changeable.
Type locality: GSC locality C-080584, Phantom Creek Formation, Yakoun River, Graham Island, Queen Charlotte
Island, British Columbia.
Occurrence: Ghost Creek, Fannin, Whiteaves and Phantom
Creek formations, Queen Charlotte Islands; San Hipólito
Formation, Baja California Sur; Haliw (Aquil) Formation,
Oman; Bainang Terrane, Tibet.
Genus: Perispyridium Dumitrica 1978
Type species: Trilonche? ordinaria Pessagno 1977a
Synonymy:
1978 Perispyridium n. gen. – Dumitrica, p. 35.
1987b Protoperispyridium n. gen – Yeh, p. 91.
Original description: Flat eptingiids with cephalis small,
surrounded in frontal plane by a triangular or subcircular
peripheral latticed shell; sagittal ring inserted in the cephalic
wall; arches more or less distinct.
312
Original remarks: Perispyridium seems to be the last survivor of the family. It bears the most advanced spumellarian
morphology among the eptingiids, the cephalis being able
to be easily confused with the microsphere and the peripheral latticed shell with the cortical shell.
Included species:
PSP03 Perispyridium hippaense (Carter) 1988
PSP01 Perispyridium oregonense (Yeh) 1987b
Plate PAR21. Paronaella variabilis Carter. Magnification x 150. Fig. 1(H). Carter et al. 1988, pl. 11, fig. 1.
Fig. 2. Carter et al. 1988, pl. 11, fig. 3. Fig. 3. QCI, GSC C-080611, GSC 128876. Fig. 4. Whalen & Carter 2002, pl. 3,
fig. 3. Fig. 5. OM, Haliw-039-R03-10. Fig. 6. OM, Haliw-039-R03-11.
313
Perispyridium hippaense (Carter) 1988
Species code: PSP03
Synonymy:
1987b Protoperispyridium sp. A – Yeh, p. 93, pl. 13, figs. 13, 23.
1988 Protoperispyridium hippaensis Carter n. sp. – Carter et al.,
p. 59, pl. 6, figs. 1-2.
2004 Perisypridium sp. – Matsuoka, fig. 183.
Original diagnosis: Strongly triangular, thickened peripheral shell with concave sides, heavy nodes and prominent
sleeve-like extensions. Three spines are massive and triradiate with crown-like extensions (produced by subsidiary
spines on ridge tips).
Original description: Cephalis indistinct, peripheral shell
triangular in outline, sides concave; shell thickened at
right angles to frontal plane. Apical and two primary lateral spines often not at 120º. Spines massive, triradiate
with alternating ridges and grooves. Ridges wide and deep,
grooves relatively shallow, merging to a point. Outer tip of
each ridge is widened and blunt. The subsidiary spines together produce a crown-like structure.
Original remarks: Differs from Protoperispyridium sp. B,
in having a thicker shell with concave sides, more massive
pore frames and heavier nodes.
Further remarks: Perispyridium hippaense differs from
P. oregonense (Yeh) in having large nodes on the peripheral
shell and spines terminating in a crown-like structure.
Measurements (µm):
Based on 10 specimens.
Width of test
(along spine axis)
Length of longest spine
HT
Av.
Max.
Min.
120
130
150
112
93
96
110
82
Etymology: Named for Hippa Island, on the west coast of
Graham Island.
Type locality: GSC locality C-080588,Graham Island Formation, Rennell Junction, Graham Island, Queen Charlotte
Islands, British Columbia.
Occurrence: Whiteaves, Phantom Creek, and Graham Island formations, Queen Charlotte Islands; Hyde Formation, Oregon; Japan.
Perispyridium oregonense (Yeh) 1987b
Species code: PSP01
Synonymy:
1987b Protoperispyridium oregonense n. sp. – Yeh, p. 91, pl. 13,
figs. 14, 16, 18, 20, 22, 24.
1987b Perispyridium (?) sp. A – Yeh, p. 91, pl. 3, fig. 8, pl. 24,
fig. 4, 13, 22, 24.
1987b Protoperispyridium sp. D – Yeh, p. 93, pl. 24, fig. 14.
1989 Protoperispyridium sp. cf. P. oregonense Yeh – Hattori,
pl. 3, fig. J.
1997 Perispyridium sp. E0 – Yao, pl. 15, fig. 708.
2003 Perispyridium cf. oregonense (Yeh) – Goričan et al., p. 296,
pl. 4, fig. 1.
Original description: Test convex at right angle to frontal
plane. Cephalis large, with irregular large polygonal pore
frames forming raised cephalic wall. Sides of peripheral
shell concave inwards. Pore frames with thin rims and thick
sides. Peripheral shell connected to three spines with truncated, sleeve-like extensions. Apical spines and two primary lateral spines displaced equally, about equal in length,
and triradiate with three wide ridges alternating with three
wide grooves, grooves about as wide as ridges. Narrow subsidiary grooves occurring on ridges. Peripheral shell consisting of large irregular pore frames with thickest portion
on sleeve-like extension and connected to cephalis by two
or three butresses which separate one or two small rounded
pericephalic pores.
314
Measurements (µm):
Ten specimens measured. LT = length of test along the axis
of apical spine, WT = maximum width of test normal to the
axis of apical spine.
HT
Mean
Max.
Min.
LT (max.)
123
125
141
123
WT (max.)
135
131
135
128
Etymology: This species is named for the state of Oregon.
Type locality: OR-600M, Hyde Formation at Izee-Paulina
road, east-central Oregon.
Occurrence: Hyde Formation, Oregon; Ghost Creek and
Fannin formations, Queen Charlotte Islands; Skrile Formation, Slovenia; Japan.
Plate PSP03. Perispyridium hippaense (Carter). Magnification x250. Fig. 1(H). Carter et al. 1988, pl. 6, fig. 1.
Fig. 2. Carter et al. 1988, pl. 6, fig. 2. Fig. 3. JP, MNA-10, MA11437.
Plate PSP01. Perispyridium oregonense (Yeh). Magnification x250. Fig. 1(H). Yeh 1987b, pl. 13, fig. 14. Fig. 2. QCI,
GSC loc. C-080611, GSC 111741. Fig. 3. QCI, GSC loc. C-175309, GSC 111742.
315
Genus: Pleesus Yeh 1987b
Type species: Pleesus aptus Yeh 1987b
Synonymy:
1987b Pleesus n. gen. – Yeh, p. 82.
Original description: Test multicyrtid, spindle-shaped
(conical when broken), with or without constrictions between joints. Cephalis dome-shaped with horn. Earlier
chambers covered with layer of microgranular silica, remaining chambers consisting of single layer of regular to
subregular polygonal pore frames. Final post-abdominal
chambers slightly inflated and terminating in open, narrow,
elongate, tubular extension.
Original remarks: Pleesus n. gen., differs from Katroma
Pessagno and Poisson (1981) by lacking a horn on
cephalis, and by having a less inflated final post-abdominal
chamber.
Etymology: Pleesus is a name formed by an arbitrary
combination of letters (ICZN, 1985, Appendix D, pt.Vl,
Recommendation 40, p. 201).
Included species:
PLE01 Pleesus aptus Yeh 1987b
Pleesus aptus Yeh 1987b
Species code: PLE01
Synonymy:
1987b Pleesus aptus n. sp. – Yeh, p. 82, pl. 10, figs. 9, 18, 23;
pl. 23, fig. 7.
1996 Pleesus sp. – Tumanda et al., p. 181, Fig. 5.9.
1998 Pleesus sp. aff. P. aptus Yeh – Yeh & Cheng, p. 34, pl. 4,
fig. 17.
2004 Pleesus aptus Yeh – Matsuoka, fig. 122.
Original description: Test as with genus, elongate, spindleshaped, with slight constrictions between joints, with as
many as nine post-abdominal chambers. Cephalis domeshaped, medium in size, without horn. Thorax and subsequent chambers trapezoidal in outline, gradually increasing in width as added, with final post-abdominal chamber
slightly inflated and terminating in narrow, latticed, tubular extension. Cephalis imperforate, thorax and abdomen
sparsely perforate, covered by layer of microgranular silica.
Post-abdominal chambers consisting of single layer of subregular tetragonal, pentagonal, and hexagonal pore frames.
Pore frames gradually increasing in size distally.
316
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Length
Width of Length of
Max.
of conical conical part inflated
width
part
at base
part
187
90
75
115
73
118
192
110
209
128
75
126
185
90
71
112
Etymology: Aptus-a-um (Latin, adj.) = useful.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Hyde Formation and Warm Springs member
of the Snowshoe Formation, Oregon; Liminangcong Chert,
Philippines; Tawi Sadh Member of the Guwayza Formation,
Musallah Formation, Oman; Japan.
Plate PLE01. Pleesus aptus Yeh. Magnification x200. Fig. 1(H). Yeh 1987b, pl. 10, fig. 9. Fig. 2. Matsuoka 2004, fig. 122.
Fig. 3. OM, BR706-R12-08. Fig. 4. BR706-R12-04. Fig. 5. OM-00-252-021817. Fig. 6. OM-00-251-021417.
317
Genus: Plicaforacapsa O’Dogherty, Goričan & Dumitrica 2006
Type species: Stylocapsa catenarum Matsuoka 1982a
Synonymy:
2006 Plicaforacapsa O’Dogherty, Goričan & Dumitrica n. gen.
- O’Dogherty et al., p. 443.
Original diagnosis: Test elongated fusiform, composed
of two or more segments. Outer surface of shell with
longitudinal plicae bearing one row of small circular pores.
Cephalis small, hemispherical, last segment inflated with a
constricted aperture at the base.
Original remarks: Plicaforacapsa differs from other genera
in having pores on the longitudinal plicae. At present two
species with this ornamentation have been found in the
Jurassic: Stichocapsa elegans Matsuoka from the Toarcian
(Matsuoka 1991a) and Stylocapsa catenarum Matsuoka
from the middle to late Bathonian (Baumgartner et al.,
1995a). A direct phylogenetic relationship between them
has not been demonstrated yet.
Etymology: Referring to perforated plicae, feminine gender.
Included species:
SCP02 Plicaforacapsa? elegans (Matsuoka) 1991
Plicaforacapsa? elegans (Matsuoka) 1991
Species code: SCP02
Synonymy:
1991 Stichocapsa elegans n. sp. – Matsuoka, p. 731, Fig. 7. 1a – 5b.
2003 Stichocapsa elegans Matsuoka – Goričan et al., p. 297,
pl. 4, fig. 11.
2004 Stichocapsa elegans Matsuoka – Matsuoka, fig. 86.
Original description: Shell of four to five segments,
elongate ovoidal. Cephalis hemispherical, poreless. Thorax,
abdomen and the fourth segment in the case of fivesegmented specimens truncate conical. All segments but
the last one form conical proximal part; the last segment
large, hemispherical with a strongly constricted aperture.
Strictures between segments indistinct externally. Ten to
12 longitudinal plicae observed on the conical proximal
portion of shell in lateral view. One row of pores runs on the
plicae. The distal portion of shell with smooth, perforated
surface. Pores small and circular.
Original remarks: Stichocapsa elegans, n. sp. differs from
S. plicata, n. sp. by its slender form and by possessing
318
perforated longitudinal plicae on the conical proximal
portion of the shell.
Measurements (µm):
Numbers of specimens measured are in parentheses.
Total height of shell
Max. width of shell
Diameter of aperture
HT
170
89
8
Max.
179
105
8
Min.
152
81
6
Mean
165
89
7
(15)
(15)
(2)
Etymology: The specific name comes from the Latin elegans
(=elegant).
Type locality: MNA-10, Nanjo Massif, Mino Terrane,
central Japan.
Occurrence: Mino Terrane, Japan; Skrile Formation,
Slovenia; Tawi Sadh Member of the Guwayza Formation,
Oman.
Plate SCP02. Plicaforacapsa? elegans (Matsuoka). Magnification x400. Fig. 1(H) a-b. Matsuoka 1991, fig. 7.1a-b.
Fig. 2. Goričan et al. 2003, pl. 4, fig. 11. Fig. 3. OM, BR1121-R09-17.
319
Genus: Podocapsa Rüst 1885, emend. Foreman 1973
Type species: Podocapsa guembeli Rüst 1885 (subsequent designation by Campbell, 1954)
Synonymy:
1885 Podocapsa n. gen. – Rüst, p. 304.
1973 Podocapsa Rüst emend. – Foreman, p. 267.
Original description: The three following species required
the definition of a new genus. A diagnosis would be: Monocyrtida clausa eradiata, testa subsphaerica, appendicibus
tribus vel pluribus ubique clathratis, and it would have its
analogue in the genera Haeckel’s dicyrtid genus Sethrochytris and Ehrenberg’s Lithochytris. Of the two latticed extensions the two opposite ones, which could be named basal
extensions, are always equal, while the third one, the apical
extension, is developed different.
Further remarks: By Foreman (1973): When Campbell
(1954, p. D122) subsequently designated Podocapsa
guembeli Rüst, 1885 as the type species of Podocapsa, he did
not, in the absence of a type designation by Rüst, indicate
which of the two entirely different specimens illustrated by
Rüst was to be considered as the lectotype of P. guembeli.
He did reproduce one of Rüst’s illustrations, fig. 5 on pl. 36.
However, since he very frequently selected a specimen other
than the one or more which were eligible to be the type
of the species designated as type species of the genera he
treated, this illustration by Campbell is not considered to be
a designation. We therefore designate Rüst’s specimen (pl.
36, fig. 6) as the lectotype of Podocapsa guembeli. Although
Rüst considered this specimen to be a monocyrtid with two
porous wings and a porous apical extension, it is apparent
that the latter is actually the terminal tube of the distalmost
segment and that the proximal segments have been broken
off. The generic definition of Podocapsa is thus emended as
follows: Shell of at least three segments, the proximal part
small, made up of all but the distalmost segment which is
large, globose, and bears three porous wings and a porous
terminal tube.
By Baumgartner et al. (1995a): This genus can be distinguished from the genus Podobursa by the nature of the
laterally directed porous wings as opposed to laterally directed spines on Podobursa.
Included species:
POD01 Podocapsa abreojosensis Whalen & Carter 2002
Podocapsa abreojosensis Whalen & Carter 2002
Species code: POD01
Synonymy:
1984 unidentified Radiolaria – Whalen & Pessagno, pl. 1, fig. 8.
1998 Podocapsa sp. A – Yeh & Cheng, p. 30, pl. 6, figs. 12, 16.
2002 Podocapsa abreojosensis n. sp. – Whalen & Carter, p. 136,
pl. 14, figs. 6, 7, 13, 14; pl. 18, figs. 10, 11.
Original description: Test composed of cephalis, thorax
and abdomen. Cephalis hemispherical, with small horn,
small pores masked by a layer of microgranular silica. Thorax trapezoidal in outline with small, irregularly sized and
shaped pores covered by layer of microgranular silica. Abdominal chamber spherical, inflated, much larger than first
two chambers, terminating in porous, cylindrical terminal
extension. Pore frames on abdomen pentagonal, hexagonal,
as well as irregular in construction, much larger than on
first two segments; abdominal pore frames larger in medial
position becoming smaller towards thorax and terminal
tube. Three, large, porous arms circumferentially arranged
along widest part of abdomen, situated approximately 120°
from each other. Both arms and terminal tube gently tapering distally. Arms and terminal tube usually with broken
tips but some specimens show closure, particularly on the
arms (holotype). Pores on arms and terminal tube slightly
smaller than on abdomen, sometimes showing slight trochospiral arrangement.
Original remarks: Podocapsa abreojosensis n. sp., is
distinguished from P. amphitreptera Foreman 1973, by the
320
less globose shape of the cephalis and thorax and longer
arms. It is possible Podocapsa? abreojosensis n. sp., may be
a heterochronous homeomorph as it is so disconnected in
time from all other species of this genus; for this reason,
the genus is queried. However, at this time we are unable to
recognize characteristics that would suggest establishing a
new genus.
Measurements (µm):
(n) = number of specimens measured
Length (10)
(maximum)
278
278
173
213
Length of arms (11)
135
150
60
96
HT
Max.
Min.
Mean
Etymology: This species is named for Punta Abreojos
located to the south of the type area.
Type locality: Sample BPW80-30, San Hipólito Formation,
Punta San Hipólito, Vizcaino Peninsula, Baja California,
Mexico.
Occurrence: San Hipólito Formation, Baja California Sur;
Liminangcong Chert, Philippines; Haliw (Aquil) Formation, Oman.
Plate POD01. Podocapsa abreojosensis Whalen & Carter. Magnification x200 except Fig. 1b(H) x300. Fig. 1(H).
Whalen & Carter 2002, pl. 14, figs. 6, 13. Fig. 2. Whalen & Carter 2002, pl. 14, fig. 7. Fig. 3. OM, BR682-R09-13. Fig. 4.
OM, Haliw-039-R03-04.
321
Genus: Praeconocaryomma Pessagno 1976
Type species: Praeconocaryomma universa Pessagno 1976
Synonymy:
1976 Praeconocaryomma n. gen. – Pessagno, p. 40.
Original description: Cortical shell invariably with radial
spines protruding from mammae. Pore frames differing
in size, shape, and distribution between cortical shell and
each medullary shell, tending to be proportionately and
progressively larger on each medullary shell. Radial beams
connecting medullary shells about one-third as thick as
those connecting cortical shell to first medullary shell.
Original remarks: Praeconocaryomma n. gen., differs
from Conocaryomma Lipman by invariably having three
rather than four or five medullary shells. It differs from
Phaenicosphaera Haeckel by possessing three medullary
shells rather than one.
“Acanthosphaera” magnimamma Rüst and “Heliosphaera”
mammillaria Rüst from the Kieselkalk of Cittiglio, Italy,
appear to be early representatives of Praeconocaryomma.
These same species are now known from Tithonian strata
in the California Coast Ranges. Forms figured by Rüst
(1885, pl.28, figs.12-13) from the Koprolithen of Ilsede,
northwestern Germany, as “Carposphaera” circumplicata
Rüst and “Carposphaera” affinis Rüst are probably
assignable at least to the Praeconocaryommidae. Likewise,
“Conosphaera” sphaeroconus Rüst (1898, pl. 4, fig. 8)
from the Kieselkalk of Cittiglio may be assignable to the
Praeconocaryommidae. All three of the last-named species
have cortical shells with mammae lacking radial spines;
they should probably be assigned to a new genus.
Included species:
PRY05 Praeconocaryomma bajaensis Whalen n. sp.
PRY01 Praeconocaryomma decora gr. Yeh 1987b
PRY02 Praeconocaryomma immodica Pessagno & Poisson
1981
PRY03 Praeconocaryomma parvimamma Pessagno &
Poisson 1981
PRY07 Praeconocaryomma sarahae Carter n. sp.
PRY04 Praeconocaryomma whiteavesi Carter 1988
PRY06 Praeconocaryomma? yakounensis Carter n. sp.
Praeconocaryomma bajaensis Whalen n. sp.
Species code: PRY05
Synonymy:
1989 Praeconocaryomma spp. – Hattori, pl. 9, fig. M.
1996 Praeconosphaera sphaeroconus (Rüst) Yang – Pujana,
p. 136, pl. 1, fig. 21.
1997 Praeconocaryomma sp. A – Yao, pl. 1, fig. 32.
2002 Praeconocaryomma sp. A – Whalen & Carter, p. 108, pl. 8,
fig. 8.
2003 Praeconocaryomma spp. – Goričan et al., p. 291, pl. 1,
fig. 14 only.
Type designation: Holotype USNM 401912 (pl. PRY05,
fig. 1) from sample BPW80-30. Paratype, GSC 111744 from
GSC loc. C-140418 (pl. PRY05, fig. 2).
Description: Test spherical with large, closely spaced porous mammae that comprise a large part of the total surface
area. Surfaces of mammae rounded, penetrated by many
small circular pores centered around a small rod-like spine.
Pores in intermammary area generally small, very irregular
in shape. Outermost medullary shell with variably-sized
pentagonal pore frames. Thin triradiate beams connect
medullary shell with mammae on cortical shell.
Remarks: This form is similar to Praeconocaryomma
whiteavesi Carter but differs in having less numerous
but much larger mammae and correspondingly smaller
intermammary areas with smaller pores.
322
Measurements (µm):
Based on 6 specimens
Diameter of cortical shell
Diameter of mammae
Height of mammae
HT
224
32
20
Max.
308
53
31
Min.
207
25
14
Mean
247
37
21
Etymology: Named for Baja California Sur, Mexico.
Type locality: Loc. BPW80-30, sandstone member, San
Hipólito Formation, Punta San Hipólito, Vizcaino Peninsula, Baja California Sur, Mexico. Paratype from Louise Island, Queen Charlotte Islands.
Occurrence: San Hipólito Formation, Baja California Sur;
Ghost Creek and Fannin formations, Queen Charlotte
Islands; Sierra Chacaicó Formation, Argentina; Skrile
Formation, Slovenia; Tawi Sadh Member of the Guwayza
Formation, Oman; Japan.
Plate PRY05. Praeconocaryomma bajaensis Whalen n. sp. Magnification Figs. 1-4, 6 x200 (scale bar A), figs. 5, 7-9 x150
(scale bar B). Fig. 1(H). Whalen & Carter 2002, pl. 8, fig. 8. Fig. 2. QCI, GSC loc. C-140418, GSC 111744.
Fig. 3. QCI, GSC loc. C-304566, GSC 111745. Fig. 4. Goričan et al. 2003, pl. 1, fig. 14. Fig. 5. SI, MM 5.00, 010114.
Fig. 6. OM, BR1121-R06-02. Fig. 7. QCI, GSC loc. C-305417, GSC 111746. Fig. 8. OM, BR1121-R06-06.
Fig. 9. OM, BR1121-R07-24.
323
Praeconocaryomma decora gr. Yeh 1987b
Species code: PRY01
Synonymy:
1987b Praeconocaryomma decora n. sp. – Yeh, p. 39, pl. 6, fig. 15;
pl. 20, figs. 1-2, 9, 16, 19.
1987b Praeconocaryomma sp. A – Yeh, p. 40, pl. 2, figs. 17, 22;
pl. 20, fig. 4.
1990 Praeconocaryomma decora Yeh – Nagai, pl. 6, fig. 6.
1998 Praeconocaryomma decora Yeh – Yeh & Cheng, p. 15,
pl. 11, figs. 1, 5.
2002 Praeconocaryomma sp. A Yeh – Whalen & Carter, p. 108,
pl. 8, fig. 5.
2003 Praeconocaryomma spp. – Goričan et al., p. 291, pl. 1,
fig. 10 only.
Original description: Cortical shell with closely spaced
mammae. Mammae uniform in size, moderately large, high
in relief, distal surface rounded, hexagonal or subcircular
in outline, with long primary spine originating from
center of each mamma. Primary spines slender, circular in
axial section. Each mamma with five to seven short rays
connecting directly to adjacent mammae or linking with
rays from surrounding mammae. Rays single, bifurcate,
or trifurcate. Mammary pore frames poorly developed,
with thin bars connecting massive rays and formnig small
subtriangles beneath mammae. Intermammary pore frames
large, regular in size, mostly triangular and tetragonal in
outline.
Original remarks: Praeconocaryomma decora, n. sp., is
very similar to P. immodica Pessagno & Poisson (1981) by
having a cortical shell with large rounded, closely spaced
mammae and by having primary spines circular in axial
section. These two species can be distinguished by different
patterns of intermammary pore frames.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Diameter of
cortical shell
180
180
184
174
Diameter of
medullary shell
90
95
87
Length of spines
48
70
92
45
Etymology: Decorus-a-um (latin, adj.) = graceful.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, Oregon; San
Hipólito Formation, Baja California Sur; Skrile Formation,
Slovenia; Liminangcong Chert, Philippines; Tawi Sadh
Member of the Guwayza Formation, Oman.
Praeconocaryomma immodica Pessagno & Poisson 1981
Species code: PRY02
Synonymy:
1977a Praeconocaryomma magnimamma (Rüst) – Pessagno,
p. 77, pl. 5, figs. 14-16; pl. 6, fig. 1.
1981 Praeconocaryomma immodica n. sp. – Pessagno & Poisson,
p. 57, pl. 7, figs. 2-9.
1984 Praeconocaryomma immodica Pessagno & Poisson
– Pessagno et al., p. 24, pl. 1, figs. 22-24.
1987 Praeconocaryomma aff. P. immodica Pessagno & Poisson
– Hattori, pl. 21, fig. 1.
1988 Praeconocaryomma immodica Pessagno & Poisson
– Carter et al., p. 31, pl. 1, fig. 1.
1996 Praeconocaryomma immodica Pessagno & Poisson
– Tumanda et al., p. 173, Fig. 4.7.
1996 Praeconocaryomma immodica Pessagno & Poisson – Yeh
& Cheng, p. 100, pl. 2, fig. 12.
1998 Praeconocaryomma immodica Pessagno & Poisson
– Cordey, p. 89, pl. 22, figs. 1, 2.
1998 Praeconocaryomma immodica Pessagno & Poisson – Yeh
& Cheng, p. 15, pl. 1, fig. 11, 14; pl. 11, fig. 15.
2004 Praeconocaryomma immodica Pessagno & Poisson – Hori,
pl. 5, fig. 9.
2004 Praeconocaryomma immodica Pessagno & Poisson
– Ziabrev et al., Fig. 5-6.
2005 Praeconocaryomma immodica Pessagno & Poisson – Hori,
pl. 8, fig. 53.
Original description: Cortical shell with prominent
mammae which tend to be exceedingly high in relief.
Distal surfaces (tops) of mammae imperforate, somewhat
324
flattened, pentagonal in outline; mammae with radially
arranged primary spines that are circular in axial section.
Each face of pentagonal mammae with large pores; pores
separated by stout rays which project into intermammary
areas; individual rays bifurcate or trifurcate linking up
with rays of adjoining mammae and forming triangular
intermammary pore frames. Massive nodes present
at point of bifurcation or trifurcation. Well preserved
specimens with thinner rays projecting from bottom
side of rays at nodal points forming subsidiary triangular
pore frames. Primary radial beams (circular in axial
section) continuous with radial beams connecting cortical
shell with first medullary shell and first medullary shell
with second medullary shell. First medullary shell with
triangular meshwork comprised of equilateral triangular
pore frames; second medullary shell with polygonal pore
frames.
Original remarks: P. immodica, n. sp., differs from P. media, n. sp., (1) by having mammae which are pentagonal
rather than hexagonal in ouline and which are considerably higher in relief; (2) by having mammae which are
more closely spaced; and (3) by having more complex
intermammary areas. P. media has triangular mammary
pore frames closed by a bar at their base; however, the
mammary pore frames of P. immodica lack the basal bar
and are open basally.
Plate PRY01. Praeconocaryomma decora gr. Yeh. Magnification x250. Fig. 1(H). Yeh 1987b, pl. 19, fig. 2.
Fig. 2. Whalen & Carter 2002, pl. 8, fig. 5. Fig. 3. QCI, GSC loc. C-304568, GSC 111804. Fig. 4. OM, BR1121-R09-13.
Fig. 5. Goričan et al. 2003, pl. 1, fig. 10. Fig. 6. OM, BR1121-R06-01. Fig. 7. OM, BR1121-R08-29.
Fig. 8. OM, BR1121-R06-03.
Plate PRY02. Praeconocaryomma immodica Pessagno & Poisson. Magnification x250. Fig. 1(H). Pessagno & Poisson
1981, pl. 7, fig. 2. Fig. 2. QCI, GSC loc. C-080613, GSC 111743. Fig. 3. OM, BR1121-15929.
325
This species seems to be the most advanced form in the
P. parvimamma, n. sp., lineage group. At present it is not
possible to link it directly to earlier and simpler forms
such as P. media. It would, however, appear that the basal
bar of the P. media mammary pore frames has been lost in
the course of the evolution of this lineage.
Measurements (µm):
Based on 10 specimens.
HT
Max.
Min.
Diameter of cortical shell
206
225
193
Height of mammae
44
44
25
Etymology: Immodicus-a-um (Latin, adj.): immoderate,
excessive.
Type locality: Sample BK 605, red radiolarian chert in mélange, Franciscan Complex, California.
Occurrence: Franciscan Complex, California; Fannin,
Whiteaves and Phantom Creek formations, Queen Charlotte Islands; Bridge River Complex, British Columbia; Liminangcong Chert, Philippines; Japan; Bainang Terrane,
Tibet; Tawi Sadh Member of the Guwayza Formation,
Oman.
Praeconocaryomma parvimamma Pessagno & Poisson 1981
Species code: PRY03
Synonymy:
1981 Praeconocaryomma parvimamma n. sp. – Pessagno &
Poisson, p. 58, pl. 8, figs. 5-8, pl. 9, fig. 2.
1998 Praeconocaryomma parvimamma Pessagno & Poisson
– Cordey, p. 89, pl. 22, figs. 3, 6.
Original description: Cortical shell with mammae having
radially arranged relatively long primary spines originating
from the center of their flat distal (top) surfaces. Primary
spines relatively long, circular in cross-section. Distal
flattened surfaces of mammae hexagonal in outline; six
sides of each mamma with massive triangular mammary
pore frames at their base; mammary pore frames with
massive nodes at their base only; pore frames and sides
of mammae sloping gently outward. Six rays originating
from position of nodes at base of mammary pore frames,
aligned with legs of each mammary pore frame and
interconnecting with rays of adjoining mammae. Large
subelliptical pores occurring between rays. Cortical shell
and two medullary shells connected by radial beams
which are circular in axial section. First medullary shell
with triangular pore frames having nodes at their vertices;
second medullary shell with polygonal (pentagonal?) pore
frames.
Original remarks: P. parvimamma, n. sp., differs from
P. media, n. sp., (1) by having much smaller, less inclined
mammary pore frames and (2) by having mammae which
are smaller with flattened distal (top) surfaces.
P. parvimamma appears to be the earliest and simplest
form of a lineage group (termed the P. parvimamma lineage group here) which includes at least four morphotypes.
The data at hand indicate the the P. parvimamma lineage
group makes its first appearance in the Lower Pliensbachian (?Upper Sinemurian) and its final appearance in
the Lower Tithonian. During the period from Early
326
Pliensbachian to Early Tithonian times this lineage tends
to change through an increase in the width and height of
mammae and by developing more complex structure in
the intermammary areas. All members of this group display a first medullary shell with equilateral triangular pore
frames.
It should be noted that the form figured by Pessagno
(1977) as P. magnimamma (Rüst) is assigned to P. immodica, n. sp., herein. Rüst’s (1898, Pl. IV, fig. 1) illustration of A. magnimamma shows a form with mammae and
intermammary areas perforated by numerous small pores.
Pessagno originally assumed that the small pores were a
figment of Rüst’s imagination and that the extremely large
mammae with long smooth (circular in axial section) primary spines were the distinguishing feature of P. magnimamma. Unfortunately, however, a form quite similar to
Rüst’s form occurs in Pliensbachian cherts from the Franciscan Complex. This form is referred to P. sp. aff. P. magnimamma (Rüst) herein.
Measurements (µm):
Based on 9 specimens.
Diameter of cortical shell
Height of mammae
HT
235
20
Min.
200
12
Max.
260
20
Etymology: Parvus-a-um (Latin, adj.): small + mamma
(-ae, F.) = breast.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Franciscan
Complex, California; Bridge River Complex, British Columbia; Tawi Sadh Member of the Guwayza Formation,
Oman.
Plate PRY03. Praeconocaryomma parvimamma Pessagno & Poisson. Magnification x200. Fig. 1(H). Pessagno &
Poisson 1981, pl. 8, fig. 5. Fig. 2. TR, 1662D-R04-09. Fig. 3. TR, 1662D-R04-02. Fig. 4. TR, 1662D-R04-07.
Fig. 5. OM, BR1121-R07-20.
327
Praeconocaryomma sarahae Carter n. sp.
Species code: PRY07
Synonymy:
? 1987 Praeconocaryomma sp. B – Hattori, pl. 20, fig. 18.
? 1997 Praeconocaryomma ? sp. D0 – Yao, pl. 1, fig. 36.
2001 Praeconocaryomma media Pessagno & Poisson – Gawlick
et al., pl. 6, fig. 2.
2002 Praeconocaryomma media Pessagno & Poisson – Suzuki et
al., p. 172, fig. 4-A.
2002 Praeconocaryomma parvimamma Pessagno & Poisson
– Suzuki et al., p. 172, fig. 4-B.
Type designation: Holotype GSC 111747 from GSC loc. C304568 (pl. PRY07 fig. 1), Rennell Junction member of the
Fannin Formation (upper lower Pliensbachian). Paratype
GSC 111751 from GSC loc. C-305417 (pl. PRY07, fig. 5),
Sandilands Formation (basal Pliensbachian).
Description: Multi-layered cortical shell with mammae,
some having remnants of short, circular primary spines.
Distal surfaces of mammae low to moderately raised with
relatively flat surfaces; usually hexagonal (occasionally
pentagonal or septagonal) in shape. Each mamma usually
surrounded by six massive triangular mammary pore
frames whose outer vertices link with other mammae
forming an interlocking meshwork. Inner layer(s) of shell
with triangular pore frames that apparently are connected
to the inner surface of external mammae directly or are
connected by some type of pillar-like structures. Inner layer
of pore frames slightly rotated, but usually aligned. Size of
pore frames gradually decreasing towards center of test.
Central structure of test unknown.
It is interesting to note that the outer shell structure
of P. sarahae n. sp. closely parallels that of the genus
Pseudopantanellium Yeh, and species P. floridum Yeh. Both
328
species are approximately the same age which suggests
the possibility that some type of horizontal gene transfer
(Dumitrica & Guex, 2003) may have been operational
during this time.
Remarks: Praeconocaryomma sarahae n. sp. differs from P.
parvimamma Pessagno & Poisson in lacking the six small
pores that surround the base of each mamma. Pessagno &
Poisson (1991) suggest that P. parvimamma is probably the
earliest and simplest form of the genus Praeconocaryomma.
P. sarahae n. sp. is even simpler morphologically and may
be the older of the two.
Measurements (µm):
Based on 6 specimens.
Diameter of cortical shell
Height of mammae
HT
207
10
Max.
232
11
Min.
197
7
Mean
210
10
Etymology: Named for Sarah K. Carter, a mammal biologist and ecologist for her help with the author’s research
and her constant support.
Type locality: Sample 99-CNA-MI-11 (GSC loc. C-304568),
Rennell Juction member of the Fannin Formation, Maude
Island, Skidegate Inlet, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands and Ghost Creek formations and
Rennell Junction member of the Fannin Formation, Queen
Charlotte Islands; Pucara Group, Peru; Dürrnberg Formation, Austria.
Plate PRY07. Praeconocaryomma sarahae Carter n. sp. Magnification x200. Fig. 1(H). GSC loc. C-304568, GSC
111747. Fig. 2. QCI, GSC loc. C-140495, GSC 111748. Fig. 3. QCI, GSC loc. C-080612, GSC 111749. Fig. 4. QCI, GSC
loc. C-140495, GSC 111750. Fig. 5. QCI, GSC loc. C-305417, GSC 111751. Fig. 6. QCI, GSC loc. C-305417, GSC 111752.
Fig. 7. QCI, GSC loc. C-305417, GSC 111753. Fig. 8. QCI, GSC loc. C-305417, GSC 111754. Fig. 9. QCI, GSC loc. C305417, GSC 111755. Fig. 10. AT, BMW21-01. Fig. 11. AT, BMW21-07.
329
Praeconocaryomma whiteavesi Carter 1988
Species code: PRY04
Synonymy:
1981 Praeconocaryomma sp. aff. P. magnimamma (Rüst)
– Pessagno & Poisson, p. 59, pl. 9, figs. 3-5.
1987b Praeconocaryomma sp. C – Yeh, p. 40, pl. 2, fig. 28; pl. 20,
fig. 5.
1988 Praeconocaryomma whiteavesi Carter n. sp. – Carter et al.,
p. 31, pl. 1, figs. 3, 6.
1989 Praeconocaryomma whiteavesi Carter – Hattori, pl. 18, fig. F.
1989 Praeconocaryomma spp. – Hattori, pl. 19, fig. B.
1998 Praeconocaryomma whiteavesi Carter – Cordey, p. 90,
pl. 22, fig. 8.
Not 1998 Praeconocaryomma whiteavesi Carter – Yeh & Cheng,
p. 15, pl. 1, figs. 9, 12.
1998 Praeconocaryomma sp. A – Yeh & Cheng, p. 15, pl. 1, fig. 10.
2005 Praeconocaryomma sp. aff. P. magnimamma (Rüst) sensu
Pessagno & Poisson – Kashiwagi et al., pl. 6, fig. 12.
Original diagnosis: Spherical test with small, closely spaced
porous mammae. Pores in intermammary areas normally
much larger; elliptical and subtriangular in shape.
Original description: Test spherical with small, closely
spaced, porous mammae. Surfaces of mammae penetrated
by a number of small circular pores centred around a small
spine, which is circular in section. Pores in intermammary
area irregularly sized; larger pores subtriangular in shape,
smaller pores more elliptical. Occasional nodes arise near
centres of intermammary areas where a number of pores
converge. First medullary shell has pentagonal pore frames
of varying size with weakly developed nodes at bar vertices.
Sturdy triradiate beams connect medullary shell with
mammae on cortical shell.
Original remarks: This form is somewhat similar to Praeconocaryomma sp. aff. P. magnimamma (Rüst) illustrated
by Pessagno and Poisson (1981, p. 59, pl. 9, figs. 3-5). It differs by having smaller mammae and correspondingly larger intermammary areas with larger pores. These differences
become even more apparent when this form is compared
with P. magnimamma (Rüst) 1898 and P. sp. aff. P. magnimamma figured by Feary and Pessagno (1980, figs. 3, 4),
and are considered significant enough to warrant designating this form a new species.
Further remarks: Praeconocaryomma sp. aff. P. magnimamma (Rüst) figured by Pessagno and Poisson (see above) is
now considered variability of P. whiteavesi.
Measurements (µm):
Based on 7 specimens.
Diameter of cortical shell
Height of mammae
HT
196
15
Av.
189
14
Max.
200
20
Min.
180
11
Etymology: Named in honour of J. F. Whiteaves, who
studied the early paleontological collection from Maude
Island.
Type locality: GCS locality C-080577, Fannin member of
the Fannin Formation, Creek locality, Maude Island, Queen
Charlotte Islands, British Columbia.
Occurrence: Fannin Formation, Queen Charlotte Islands
and Bridge River Complex, British Columbia; Warm
Springs member of the Snowshoe Formation, Oregon;
Gümüslü Allochthon, Turkey; Tawi Sadh Member of the
Guwayza Formation and Musallah Formation, Oman;
Liminangcong Chert, Philippines; Japan.
Praeconocaryomma? yakounensis Carter n. sp.
Species code: PRY06
Type designation: Holotype GSC 111794 (pl. PRY06, fig. 1),
and paratype GSC 111795 (pl. PRY06, fig. 2) from GSC loc.
C-140441; Sandilands Formation (upper Sinemurian).
Description: Large multi-layered cortical shell with prominent mammae. Distal surfaces of mammae small to
medium in size, low to moderately raised, pentagonal to
hexagonal in shape, with slightly rounded surfaces. Each
mamma surrounded by six strong triangular pore frames
whose vertices link with other mammae. Inner layer(s)
of shell comprised of smaller irregular to subtriangular
pore frames connected to the outer triangular pore frames
forming a slightly depressed interconnecting meshwork.
Short peripheral spines visible on some specimens. Central
structure of test unknown.
Remarks: Praeconocaryomma? yakounensis n. sp. differs
from P. sarahae n. sp. in possessing a larger test. Furthermore, the inner layer of pore frames connects to the outer
triangular pore frames forming an interlocking meshwork
330
rather than telescoping inward as in P. sarahae n. sp. In
Queen Charlotte Islands, this is the oldest form (late Sinemurian) that can even questionably be assigned to the genus Praeconocaryomma Pessagno.
Measurements (µm):
Based on 5 specimens.
Diameter of cortical shell
Height of mammae
HT
279
21
Max.
302
21
Min.
279
15.6
Mean
292
17.5
Etymology: Named for the type locality on the western
bank of the Yakoun River, central Graham Island.
Type locality: Sample CAA-86-T-2/3 (GSC loc. C-140441),
Sandilands Formation, Yakoun River area, central Graham
Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands, British Columbia.
Plate PRY04. Praeconocaryomma whiteavesi Carter. Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 1, fig. 3.
Fig. 2. OM-00-251, 021521. Fig. 3. OM, BR1121-R06-04. Fig. 4. OM, BR1121-R10-07. Fig. 5. OM, BR1121-R08-16.
Plate PRY06. Praeconocaryomma? yakounensis Carter n. sp. Magnification x200. Fig. 1(H). GSC loc. C-140441, GSC
111794. Fig. 2. QCI, GSC loc. C-140441, GSC 111795. Fig. 3. QCI, GSC loc. C-140441, GSC 111796. Fig. 4. QCI, GSC
loc. C-140441, GSC 111797. Fig. 5. QCI, GSC loc. C-140441, GSC 111798.
331
Genus: Praehexasaturnalis Kozur & Mostler 1983,
emend. Kozur & Mostler 1990
Type species: Palaeosaturnalis tenuispinosus Donofrio & Mostler 1978
Synonymy:
1983 Praehexasaturnalis n. gen. – Kozur & Mostler, p. 30.
1990 Praehexasaturnalis Kozur & Mostler, emend. – Kozur &
Mostler, p. 194.
Original diagnosis: Ring narrow but not differentiated
yet. Its cross section is flat. Outline of ring hexagonal to
octagonal. 8-6 very strong marginal spines. 2 polar spines
opposite to marginal spines. No auxiliary spines. Cortical
shells spongy. Medullary shells latticed.
Emended description: By Kozur & Mostler (1990): Spongy
shell globular, consisting of several concentric layers.
Microsphere latticed. Ring mostly narrow, rarely moderately
broad, in most primitive forms with 8 spines and with
rounded octagonal outline to rounded hexagonal outline,
later invariably with rounded hexagonal to hexagonal
outline and 6 spines, occasionally with 1-2 further, mostly
distinctly smaller spines. Most primitive forms only with
polar spines, higher evolved forms additionally with 2-12
auxiliary spines.
Original remarks: In the ring outline this new genus is
quite identical with Hexasaturnalis n.gen. but the ring is
still flat to shallow oval in cross section and has no ridges.
Moreover, the polar spines are situated opposite to the
marginal spines. Palaeosaturnalis Donofrio and Mostler,
1978, is distinguished by its circular outline. Moreover, in
most species the ring is broader. Praehexasaturnalis n.gen.
is a perfect transitional form between Palaeosaturnalis
Donofrio and Mostler, 1978, and Hexasaturnalis n.gen. As
for the first time in this genus the typical ring outline of the
Hexasaturnalinae n.subfam. appears, it is already placed in
this subfamily.
Further remarks: By Kozur & Mostler (1990): Kozur and
Mostler (1983) assumed that Praehexasaturnalis Kozur and
Mostler, 1983 represents the forerunner of Hexasaturnalis
Kozur and Mostler, 1983, because the striking hexagonal
ring outline is in both genera the same and Hexasaturnalis
began later than Praehexasaturnalis. As already pointed
out under the remarks to the Saturnaliacea Deflandre,
1953 and Parasaturnalidae Kozur and Mostler, 1972,
Praehexasaturnalis is a dead-ending sidebranch of the
Parasaturnalidae, in which the same hexagonal ring outline
as in primitive Acanthocircinae Pessagno, 1977b emend.
(Hexasaturnalis Kozur and Mostler, 1983, Yaosaturnalis
Kozur and Mostler, 1983) evolved, but the other
characteristics of the Acanthocircinae (peripolar spines,
ridges on the outer margin of the ring) never evolved.
Within the genus Praehexasaturnalis for the first time the
development of taxa with auxiliary spines from taxa without
auxiliary spines have been observed. Both morphologically
and phylogenetically near related forms, connected by
transitional forms, are here united into Praehexasaturnalis
that is therefore here used in a broader sense as by Kozur
and Mostler (1983). The forms with auxiliary spines
continued seemingly until the Upper Cretaceous without
larger morphological changes. These Upper Cretaceous
forms have the highest number of auxiliary spines (about
12). However, the inner structure of the shell of these Upper
Cretaceous forms is unknown. Therefore they can be only
tentatively assigned to Praehexasaturnalis.
Etymology: According to the supposed phylogenetic line
Praehexasaturnalis n. gen. – Hexasaturnalis n. gen.
Included species:
SAT01 Praehexasaturnalis tetraradiatus Kozur & Mostler
1990
Praehexasaturnalis tetraradiatus Kozur & Mostler 1990
Species code: SAT01
Synonymy:
1984 Pseudoheliodiscus (?) spp. – Whalen & Pessagno, pl. 3,
fig. 12, 13.
1990 Praehexasaturnalis tetraradiatus n. sp. – Kozur & Mostler,
p. 195, pl. 6, figs. 8, 9, 11, 12.
1994 Praehexasaturnalis tetraradiatus Kozur & Mostler – Carter,
pl. 1, fig. 19.
1998 Praehexasaturnalis tetraradiatus Kozur & Mostler
– Whalen & Carter, p. 54, pl. 14, figs. 1, 2, 5, 6, 9, 10.
2002 Praehexasaturnalis tetraradiatus Kozur & Mostler
– Whalen & Carter, p. 108, pl. 5, figs. 7, 11, 12.
2002 Praehexasaturnalis tetraradiatus Kozur & Mostler – Tekin,
p. 184, pl. 2, fig. 10.
332
Original description: Shell large, globular, spongy, consisting of several concentric layers. Microsphere latticed. Shell
surface with delicate, short, needle-like spines. Ring narrow, flat, undifferentiated, ouline hexagonal. 6 very large,
slender spines in the 6 corners of the ring. Axial spines a
little larger and more robust than the 4 circumaxial spines.
Polar spines robust. Two auxiliary spines cross-like arranged with the polar spines, mostly short, elongated triangular.
Original remarks: The phylomorphogenetic development
within the genus Praehexasaturnalis Kozur & Mostler, 1983
emend. is now well known. The oldest forms (Lower to
Middle Norian) have 8 needle-like spines, all of about the
same length (the axial spines may be somewhat larger than
the circumaxial spines, like in the stratigraphically younger
forms). The main stock (with narrow ring) of these oldest
Praehexasaturnalis is represented by Praehexasaturnalis
burnensis (Blome, 1984a), synonym, see under the genus.
The outline of this species is still variable (roundish
octagonal, roundish suboval, subquadratic).
The next younger form is Praehexasaturnalis elegans
(Kozur & Mostler, 1972) from the Middle and Upper Norian. In this species the 2 spines perpendicularly to the polar spines are already distinctly shorter than the remaining spines, the outline of the ring is rounded hexagonal to
rounded subquadratic.
In the Upper Norian (? to Rhaetian) Praehexasaturnalis tennuispinosus (Donofrio & Mostler, 1978) occurs, in
which the 2 spines perpendicular to the polar spines are
not more present. The ring outline is hexagonal.
In the (highest Upper Norian?) Rhaetian and Hettangian Praehexasaturnalis tetraradiatus n. sp. occur, that coincides morphologically with P. tenuispinosus, but has
2 auxiliary spines additional to the polar spines. These
4 inner spines are cross-like arranged.
In the Hettangian P. kirchsteinensis evolved then remained morphologically unchanged, but displays 5-10
auxiliary spines.
Maybe Upper Cretaceous forms with the same morphological character, but with peripolar spines, are the last representatives of this line that yields important guide forms
for the Norian to Hettangian time-interval. 5 distinct
zones can be discriminated within this interval by evaluation of the phylomorphogenetic development within the
genus Praehexasaturnalis: P. burnensis without P. elegans
(Lower Norian), P. elegans without P. tenuispinosus (Middle Norian), P. tenuispinosus without P. tetraradiatus (Upper Norian, higher Upper Norian radiolarian faunas not
yet well investigated) P. tetraradiatus without P. kirchsteinensis (?uppermost Norian, Rhaetian), P. tetraradiatus and
P. kirchsteinensis (Hettangian).
Taxonomically this development is interesting, because
taxa with auxiliary spines evolved from taxa without auxiliary spines. Moreover, in P. tetraradiatus the 2 polar spines
and the 2 auxiliary spines are cross-like arranged, like in the
genus Stauracanthocircus Kozur & Mostler, 1983 emend.
This is surely a homoeomorphy (see under this genus).
Measurements (µm):
Diamater of shell
Diameter of ring (in polar axis)
Diamater of ring (perpendicular to polar axis)
Width of ring
Length of spines
Min.
110
144
144
13
90
Max.
120
170
170
21
125
Etymology: According to the 4 rays at the inner margin of
the ring (2 polar spines, 2 auxiliary spines).
Type locality: Kirchstein Limestone, 6.5 km WSW of
Lenggries/Isar, Bavaria, Germany.
Occurrence: Kirchstein Limestone, Bavaria; Csövár Limestone and Várhegy Cherty Limestone formations, Hungary; Sandilands Formation, Queen Charlotte Islands; San
Hipólito Formation, Baja California Sur; Hocaköy Radiolarite, Turkey.
Plate SAT01. Praehexasaturnalis tetraradiatus Kozur & Mostler. Magnification x150. Fig. 1(H). Kozur & Mostler 1990,
pl. 6, fig. 11. Fig. 2. Whalen & Carter 2002, pl. 5, fig. 7. Fig. 3. Whalen & Carter 2002, pl. 5, fig. 11.
333
Genus: Praeparvicingula Pessagno, Blome & Hull 1993
Type species: Parvicingula profunda Pessagno & Whalen 1982
Synonymy:
Further remarks: In this catalogue we include Praecaneta
with Praeparvicingula, because on many morphotypes
the »H-linked« circumferential ridges are not clearly
distinctive.
Original description: Test conical to subcylindrical (never
spindle-shaped) with horn. Final postabdominal chamber(s) either increasing in width or maintaining same
width. Final chamber lacking narrow terminal tube as with
Parvicingula s. s.
Etymology: From the Latin prae prefix = before, and Parvicingula Pessagno.
1993 Praeparvicingula Pessagno, Blome & Hull n. gen.
– Pessagno et al., p. 144.
1993 Praecaneta Pessagno, Blome & Hull n. gen. – Pessagno et
al., p. 142.
Original remarks: Praeparvicingula was originally referred
to by Pessagno et al. (1987a, 1987b, 1989) as Parvicingula
s. l. Praeparvicingula differs from Parvicingula s. s. by lacking a narrow tube on the final postabdominal chamber.
Moreover, the final postabdominal chamber/chambers of
Praeparvicingula continue to increase in width, either rapidly or slowly, as added.
Included species:
PVG01 Praeparvicingula aculeata (Carter) 1988
PVG02 Praeparvicingula elementaria (Carter) 1988
PVG03 Praeparvicingula gigantocornis (Kishida & Hisada)
1985
PVG04 Praeparvicingula nanoconica (Hori & Otsuka) 1989
TVS01 Praeparvicingula? spinifera (Takemura) 1986
PCA02 Praeparvicingula tlellensis Carter n. sp.
Praeparvicingula aculeata (Carter )1988
Species code: PVG01
Synonymy:
1982 Parvicingula sp. A – Wakita, pl. 1, fig. 7.
1988 Parvicingula aculeata Carter n. sp. – Carter et al., p. 54,
pl. 18, figs. 1, 2, 7.
1988a Parvicingula sp. aff. P. schoolhousensis Pessagno &Whalen
– Hattori, pl. 10, fig. M.
1997 Parvicingula dhimenaensis dhimenaensis Baumgartner
– Yao, pl. 13, fig. 625.
Original diagnosis: Test subcylindrical with 10 or more
post-bdominal chambers and very short, slender horn. Test
has rows of pointed nodes in place of circumferential ridges
between abdomen and first few post-abdominal chambers.
Original description: Test elongate, subcylindrical with 10
or more post-abdominal chambers when well preserved.
Cephalis small, conical with very short, slender horn
(not visible on all specimens). Thorax, abdomen and first
few post-abdominal chambers trapezoidal, remaining
chambers subrectangular in outline. Cephalis and thorax
sparsely perforate. Post-abdominal chambers have three
lateral rows of symmetrical (predominantly pentagonal)
pore frames between ridges. Pore frames in rows flanking
circumferential ridges slope steeply away from ridges. Those
in central row depressed, smaller and staggered; pores
elliptical. Test has rows of sharp pointed nodes, rather than
discrete circumferential ridges, between abdomen and first
three or four post-abdominal chambers. More distal ridges
are narrow with small rounded nodes.
Original remarks: Differs from all other species of Parvicingula by having a very short, almost non-existent, horn
and sharp pointed nodes separating the abdomen and first
few post-abdominal chambers.
Praeparvicingula elementaria (Carter) 1988
Species code: PVG02
Synonymy:
1988 Eucyrtidium elementarius Carter n. sp. – Carter et al.,
p. 60, pl. 17, fig. 13.
1988 Parvicingula sp. B – Carter at al., p. 56, pl. 18, figs. 3, 4.
2001 Eucyrtidium ex gr. elementarius Carter – Vishnevskaya,
p. 162, pl. 60, fig. 9; pl. 69, figs. 6, 8.
2001 Laxtorum ? jurassicum Isozaki & Matsuda – Vishnevskaya,
p. 166, pl. 69, fig. 4 only.
Original diagnosis: Ovate, smooth multicyrtid with thickwalled test (at least two layers), and medium-sized symmetrical horn.
334
Original description: Test ovate, usually with six to eight
post-abdominal chambers, and a medium-sized symmetrical horn. Boundaries between initial chambers indistinct;
distal chambers separated by slightly thickened ridges and/
or lateral rows of small, poorly developed nodes. All chambers separated internally by planiform partitions with circular apertures. Apical chambers increase rapidly in width;
intermediate chambers are cylindrical; final chamber always slightly constricted. Height of all chambers appears to
be constant. Test walls thick, composed of at least two (and
likely more) layers of pores set in hexagonal pore frames.
Measurements (µm):
Based on 7 specimens.
Length (excluding horn)
Maximum width
HT
321
126
Av.
308
116
Max.
346
130
Etymology: Latin, aculeatus (adj.), prickly.
Min.
260
109
Type locality: GSC locality C-080595, Graham Island Formation, Graham Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Graham Island Formation, Queen Charlotte
Islands; Tawi Sadh Member of the Guwayza Formation,
Oman; Japan.
Plate PVG01. Praeparvicingula aculeata (Carter). Magnification x300. Fig. 1(H). Carter et al. 1988, pl. 18, fig. 1.
Fig. 2. Carter et al. 1988, pl. 18, fig. 2. Fig. 3. OM, BR871-R07-27.
Plate PVG02. Praeparvicingula elementaria (Carter). Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 17,
fig. 13. Fig. 2. Carter et al. 1988, pl. 18, fig. 4.
335
Pores circular on outer layer; small on proximal chambers,
becoming larger and more uniform in size on distal chambers.
Original remarks: This form bears no resemblance to any
known species of Eucyrtidium. Specimens (not illustrated)
of middle Toarcian age are very similar (possibly ancestral ?) to this species but differ in having a greater number
of postabdominal chambers and coarser meshwork with
smaller, more irregularly arranged pore frames.
Further remarks: This species is now assigned to Praeparvicingula because we consider Eucyrtidium elementarius and
Parvicingula sp. B of Carter (in Carter et al. 1988) conspecific and Parvicingula sp. B has slight ridges.
Measurements (µm):
Based on 17 specimens.
HT
Maximum length (excluding horn) 244
Maximum width
144
Length of horn
38
Av.
287
153
28
Max.
350
170
40
Min.
230
135
20
Etymology: Latin, elementarius (adj.), pertaining to rudiments of first principles.
Type locality: Locality GSC C-080595, Graham Island Formation, Graham Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Whiteaves, Phantom Creek and Graham Island formations, Queen Charlotte Islands; Koryak, Far East
Russia.
Praeparvicingula gigantocornis (Kishida & Hisada) 1985
Species code: PVG03
Synonymy:
1982 Parvicingula? sp. A – Kishida & Sugano, pl. 7, fig. 8.
? 1982 Parvicingula sp. – Matsuda & Isozaki, pl. 1, figs. 14, 17.
1982 Parvicingula sp. – Kido, pl. 4, figs. 11, 12.
1985 Parvicingula gigantocornis n. sp. – Kishida & Hisada,
p. 118, pl. 4, figs. 1-5.
1988 Parvicingula gigantocornis Kishida & Hisada – Sashida,
p. 22, pl. 2, figs. 5, 10-12, 20, 21; pl. 3, figs. 4, 5.
1988 Parvicingula sp. aff. P. media Pessagno &Whalen – Carter
et al., p. 55, pl. 18, figs. 9, 11.
1990 Parvicingula cf. gigantocornis Kishida & Hisada – Hori, Fig.
9.31.
1991 Parvicingula sp. B – Carter & Jakobs, p. 343, pl. 3, fig. 20.
1992 Parvicingula aff. gigantocornis Kishida & Hisada – Sashida,
pl. 2, fig. 7.
1993 Parvicingula sp. – Fujii et al., pl. 1, fig. 2.
2001 Praeparvicingula gigantocornis (Kishida & Hisada)
– Kashiwagi, Fig. 6.6.
2003 Parvicingula gigantocornis Kishida & Hisada – Kashiwagi
& Kurimoto, pl. 3, fig. 21.
2004 Parvicingula gigantocornis Kishida & Hisada – Hori, pl. 13,
figs. 49-52.
2004 Parvicingula gigantocornis Kishida & Hisada – Ishida et al.,
pl. 5, fig. 14.
Original diagnosis: Test multicyrtid, with long, massive
horn. Each circumferentail ridge widely spaced, separated
by two or sometimes three rows of small circular pores.
Original description: Test conical, with six to eight preserved chambers, terminating in long, thick, massive apical
horn. Cephalis small, hemispherical; subsequent chambers
trapezoidal in outline. Cephalis and thorax sparsely pored.
Each circumferential ridge widely spaced, separated by two
336
or sometimes three rows of small circular pores; center row
of pores poorly developed. Abdomen and post-abdominal
chambers increasing gradually in height and increasing
rapidly in width as added.
Original remarks: Parvicingula gigantocornis, n. sp.,
appears to be closely related to such species as Parvicingula
matura, P. grantensis, P. vera and others described by
Pessagno & Whalen (1982) from Middle Jurassic strata
in North America. However, P. gigantocornis differs from
these species, in having smaller pores and poorly developed
center row of pores. It is likely that P. gigantocornis is a more
primitive species of the genus Parvicingula.
Measurements (µm):
Based on 7 specimens.
Length
128
160
120
138
Width
88
128
80
90
HT
Max.
Min.
Av.
Etymology: The name is derived from the Latin noun
gigantocornis, meaning giant horn.
Type locality: Locality 253, black bedded chert, Ueno-mura
area, Kanto Mountains, Central Japan.
Occurrence: Kanto Mountains, Japan; Tawi Sadh Member
of the Guwayza Formation and Musallah Formation,
Oman; Phantom Creek and Graham Island formations,
Queen Charlotte Islands.
Plate PVG03. Praeparvicingula gigantocornis (Kishida & Hisada). Magnification x300. Fig. 1(H). Kishida &
Hisada 1985, pl. 4, fig. 1. Fig. 2. OM-00-256-022434. Fig. 3. OM, BR871-R07-06. Fig. 4. BR871-R09-02.
Fig. 5. Carter & Jakobs 1991, pl. 3, fig. 20. Fig. 6. Carter et al. 1988, pl. 18, fig. 11. Fig. 7. OM, BR871-R06-20.
Fig. 8. OM, BR871-R06-21.
337
Praeparvicingula nanoconica (Hori & Otsuka) 1989
Species code: PVG04
Synonymy:
1985 but the former is distinguished from the latter by
having clearly meshwork of outer layer and rather longer
test. The present authors doubted whether this form was
one of poor preserved specimens of P. gigantocornis at the
first. Resulting from comparison between the two from
siliceous mudstones, this form was considered the different
species from P. gigantocornis. Both species, P. nanoconica
and P. gigantocornis, differ from all other species of
Parvicingula by possessing a very small test and a long
horn. P. nanoconica also morphologically resembles P. vera
Pessagno and Whalen, 1982 and P. profunda Pessagno and
Whalen, 1982. The latter two species are distinguished
from the former in these following features additionally by
lacking of a small test and a very long horn; by having more
weakly developed circumferential ridges and by possessing
chamber comprised of a central row of smaller pores than
that of other two rows respectively. Almost all pores in each
chamber of P. nanoconica are about equal in size.
Original description: Test consisting of 5 to 7 chambers,
possibly more, conical with developed circumferential
ridges. Cephalis hemispherical with long apical horn;
horn, solid, elongated cone. Thorax and subsequent
chambers, truncated cone, increasing in width except
distalmost chamber. Surface of cephalis having irregularly
arranged pores. Meshwork of outer layer clearly, hexagonal
symmetrically constructed by 3 rows of pore frames between
two circumferential ridges; in some species, 4 rows of pore
frames are served on distal part. Pores above and below
adjoining circumferential ridges lined up in radius distance
lag. Pore frames slope steeply away from ridges and formed
wavy or nodose structure of circumferential ridges. In some
specimens, proximal portion of test possessing nodose or
spiny circumferential ridges.
Measurements (µm):
Based on 17 specimens.
1982 Parvicingula sp. – Matsuda & Isozaki, pl. 1, figs. 13, 16, 17.
1984 Parvicingula sp. A – Murchey, pl. 1, fig. 22.
1989 Parvicingula nanoconica n. sp. – Hori & Otsuka, p. 183,
pl. 2, figs. 1-6.
1990 Parvicingula nanoconica Hori & Otsuka – Hori, Fig. 9.40.
1993 Parvicingula gigantocornis Kishida & Hisada – Fujii et al.,
pl. 1, fig. 1.
1996 Praeparvicingula nanoconica (Hori & Otsuka) – Yeh &
Cheng, p. 116, pl. 6, fig. 5; pl. 9, figs. 1, 2, 7.
1997 Parvicingula sp. D2 – Yao, pl. 13, fig. 624.
1999 Praeparvicingula nanoconica (Hori & Otsuka) – Hori,
pl. 1, fig. 9.
2004 Parvicingula nanoconica Hori & Otsuka – Hori, pl. 3,
ig. 32, pl. 4, figs. 8-12; pl. 23. figs. 8, 9.
2004 Parvicingula sp. – Hori, pl. 23, fig. 14.
2005 Parvicingula nanoconica Hori & Otsuka – Hori, pl. 12,
fig. 42.
2005 Parvicingula nanoconica Hori & Otsuka – Kashiwagi et al.,
pl. 6, fig. 5.
HT
Av.
Max.
Min.
Height
198+
184
233
161+
Width
98
103
134
80
H/W
2.0+
1.8
2.4+
1.3+
Length of apical horn
55+
55
64
37
Etymology: The name is derived from the Latin adjective
nano-conicus, meaning small coned.
Type locality: The Mt. Norikuradake area, Azumi village,
Azumi-gun, Nagano Prefecture, central Japan.
Occurrence: Japan; Franciscan Complex, California; Musallah Formation, Oman; Liminangcong Chert, Philippines.
Original remarks: Parvicingula nanoconica sp. nov. is
apparently similar to P. gigantocornis Kishida & Hisada,
Praeparvicingula? spinifera (Takemura) 1986
Species code: TVS01
Synonymy:
1986 Triversus spinifer n. sp. – Takemura, p. 63, pl. 10,
figs. 21-23; pl. 11, figs. 1-2.
1987 Ristola sp. E – Hattori, pl. 19, fig. 6.
1989 Ristola spp. – Hattori, pl. 14, fig. H.
1997 Parvicingula aff. spinifer (Takemura) – Yao, pl. 13, fig. 609.
2003 Parvicingula spinifer (Takemura) – Goričan et al., p. 297,
pl. 5, fig. 5.
2004 Triversus spinifer Takemura – Matsuoka, fig. 239.
Original description: Conical to spindle-shaped shell of
seven to nine segments with many spines or nodes on its
surface. Cephalis small, spherical and poreless, with or
without an apical horn. Thorax truncated-conical, poreless
or with sparsely distributed small pores. Abdomen truncated-conical, with irregularly or transversely distributed
338
small pores. Post-abdominal segments truncated-conical
proximally, cylindrical in subsequent segments and deflated in distal segments, with circular pores arranged hexagonally and in transverse three rows. Spines conical or triradiate, situated at joints of the segments.
Original remarks: Triversus spinifer n. sp. is distinguished
from T. japonicus in possessing many spines or nodes on
its surface.
Further remarks: Triversus is not a valid name because this
name had been used a few years previously for a nematod
(Sher, 1974). This species is questionably assigned to
Praeparvicingula because of its Amphipyndax-like cephalic
skeletal structure as shown by Takemura (1986).
Plate PVG04. Praeparvicingula nanoconica (Hori & Otsuka). Magnification x300. Fig. 1(H). Hori & Otsuka 1989,
pl. 2, fig. 1a. Fig. 2. Hori & Otsuka 1989, pl. 2, fig. 5. Fig. 3. OM-99-137-000816.
Type locality: Sample TKN-105, manganese carbonate ore,
Gujo-Hachiman area, Mino terrane, central Japan.
Measurements (µm):
Based on 10 specimens.
Length of shell
Max. width of shell including spines
Min.
135
80
Max.
210
110
Occurrence: Japan; Tawi Sadh Member of the Guwayza
Formation, Oman; Skrile Formation, Slovenia.
Etymology: The name, spinifer, means thorny.
Plate TVS01. Praeparvicingula? spinifera (Takemura). Magnification x400. Fig. 1(H). Takemura 1986, pl. 10, fig. 21.
Fig. 2. JP, HM1-22, RH627. Fig. 3. JP, MNA-10, MA12396. Fig. 4. OM, BR871-R09-15. Fig. 5. Goričan et al. 2003,
pl. 5, fig. 5.
339
Praeparvicingula tlellensis Carter n. sp.
Species code: PCA02
Synonymy:
1984 Ristola sp. B – Murchey, pl. 1, fig. 10.
1987 Ristola sp. D – Hattori, pl. 19, fig. 5.
1987 Ristola sp. N – Hattori, pl. 19, fig. 9.
1987b Pseudoristola sp. B – Yeh, p. 97, pl. 14, fig. 21.
1988 Parvicingula sp. E – Carter et al., p. 56, pl. 5, fig. 13.
1989 Ristola spp. – Hattori, pl. 14, fig. I.
1990 Pseudoristola sp. B of Yeh – Nagai, pl. 2, fig. 9.
1997 Parvicingula sp. B – Yao, pl. 13, fig. 607.
2003 Parvicingula aff. decora (Pessagno & Whalen) – Goričan et
al., p. 297, pl. 5, figs. 6-9.
Type designation: Holotype GSC 80699 (Carter in Carter
et al. 1988, pl. 15, figs. 1-2) from GSC loc. C-080586; Phantom Creek Formation (Aalenian).
Description: Test conical, almost rounded apically with a
very rudimentary horn. Cephalis hemispherical, imperforate; thorax slightly trapezoidal, sparsely perforate. Abdomen and five to seven post- abdominal chambers slightly
widening distally as added; all with three rows of hexagonal pore frames. Outer rows of pore frames staggered with
respect to central row; circumferential ridges separating
chambers smooth, nodes low and rounded.
Remarks: Praeparvicingula tlellensis n. sp. differs from
P. decora (Pessagno & Whalen) in having a more gradually
340
conical outline, in lacking H-linked circumferential ridges,
and in having much smaller nodes on ridges. P. tlellensis
n. sp. differs from Parvicingula (?) spinata (Vinassa) in
having a more rounded cephalis, smaller pores on postabdominal chambers, and rounded rather than sharp
circumferential ridges.
Measurements (µm):
Based on 10 specimens.
Length (excl. horn)
Maximum width
HT
192
100
Max.
234
109
Min.
184
90
Mean
210
100
Etymology: Named for the Tlell River northeast of the type
locality, name of Haida origin meaning place-of-big-surf
and alternately, land-of-berries.
Type locality: Sample GSC loc. C-080586, Phantom Creek
Formation, waterfall locality on the east side of Branch
Road 59, central Graham Island, Queen Charlotte Islands,
British Columbia.
Occurrence: Whiteaves, Phantom Creek and Graham Island formations, Queen Charlotte Islands; Hyde Formation, Oregon; Franciscan Complex, California; Skrile Formation, Slovenia; Japan.
Plate PCA02. Praeparvicingula tlellensis Carter n. sp. Magnification x300. Fig. 1(H). Carter et al. 1988, pl. 5, fig. 13.
Figs. 2-5. Goričan et al. 2003, pl. 5, figs. 6-9. Fig. 6. SI, MM 21.70, 010221.
341
Genus: Protopsium Pessagno & Poisson 1981
Type species: Protopsium ehrenbergi Pessagno & Poisson 1981
Synonymy:
1981 Protopsium n. gen. – Pessagno & Poisson, p. 53.
Original description: Primary test ellipsoidal (sometimes
somewhat flattened) with two polar spines. Patagium-like
mass of irregularly shaped and distributed pore frames
occurring in one plane. Secondary spines of variable size
radiating out from primary test into patagium-like mass
seemingly offering support for the irregular meshwork.
Polar spines with or without alternating grooves and ridges,
occasionally bifurcating.
Original remarks: Protopsium n. gen., differs from Archaeospongoprunum Pessagno (1973): (1) by possessing a
patagium-like mass supported by secondary spines; (2) by
having polar spines which may or may not have alternating
ridges and grooves and which sometimes bifurcate; and
(3) by sometimes displaying a somewhat compressed test.
Protopsium like Archaeospongoprunum possesses meshwork
arranged in concentric layers.
Etymology: Protopsium is a name formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, Pt. IV,
Recommendation 40, p. 113).
Included species:
PTP01 Protopsium gesponsa De Wever 1981c
Protopsium gesponsa De Wever 1981c
Species code: PTP01
Synonymy:
1981c Protopsium gesponsa n. sp. – De Wever, p. 145, pl. 5,
fig. 9-11.
1981c Protopsium sp. aff. P. gesponsa – De Wever, p. 148, pl. 5,
fig. 19.
1981 Protopsium sp. A – Pessagno and Poisson, p. 54, pl. 4,
figs. 1, 4.
1981 Protopsium sp. C – Pessagno and Poisson, p. 54, pl. 4,
figs. 3, 5-8.
1982b Protopsium gesponsa De Wever – De Wever, p. 185, pl. 10,
figs. 11-13; pl. 15, figs. 3-6.
2003 Protopsium gesponsa De Wever – Goričan et al., p. 295,
pl. 2, figs. 11, 12.
2004 Protopsium gesponsa De Wever – Matsuoka, fig. 15.
Original description: Protopsium with an ovoid shell
bearing two stout spines triradiate in cross-section over
half their length. Spines frequently asymmetric, conical in
general shape.
Original remarks: This species differs from P. ispartaensis
by its less massive spines, triradiate in cross-section
over half their length, and a shell with a finer network
often prolongated by long spurs subparallel to spines
(pl. 5, fig. 9).
342
This form is distinguished from P. ehrenbergi by its very
elongate central shell and conical spines that are partially
rounded in cross-section. It differs from P. libidonosum and
P. posinos by the presence of two spines rather than three
or more.
Measurements (µm):
Based on 4 specimens.
Length of central shell
Width of shell
HT
130
82
Av.
123
90
Min.
110
82
Max.
130
100
Spines can reach 110 µm in length.
Etymology: Anagram of E. A. Pessagno Jr. who illustrated
this form.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Skrile Formation, Slovenia; Tawi Sadh Member of the Guwayza Formation, Oman; Mino Terrane, Japan.
Plate PTP01. Protopsium gesponsa De Wever. Magnification x250. Fig. 1(H). De Wever 1981c, pl. 5, fig. 11.
Fig. 2. Matsuoka 2004, fig. 15. Fig. 3. OM, BR706-R05-20. Figs. 4, 5. Goričan et al. 2003, pl. 2, figs. 11-12.
343
Genus: Protunuma Ichikawa & Yao 1976
Type species: Protunuma fusiformis Ichikawa & Yao 1976
Synonymy:
1976 Protunuma n. gen. – Ichikawa & Yao, p. 114.
Original description: Spindle shaped, multisegmented
form with inversely subconical last segment which has
a small aperture at its base. No indentation at surface
junction of segments. Numerous small circular pores on
surface aligned in longitudinal rows and in diagonal aspect.
Numerous longitudinal plicae on surface generally running
continuously through segments. Apical horn not present
or, if present, insignificant.
Original remarks: This genus differs from Unuma in the
last segment, which has no basal appendage with large
pores but has a constricted, small, terminal aperture. At
present, no spiny form like Unuma (Spinunuma) has been
observed in the genus Protunuma.
Included species:
PRU01 Protunuma paulsmithi Carter 1988
Protunuma paulsmithi Carter 1988
Species code: PRU01
Synonymy:
1988 Protounuma paulsmithi Carter n. sp. – Carter et al., p. 54,
pl. 6, figs. 9, 12.
1991 Protounuma paulsmithi Carter – Carter & Jakobs, p. 344,
pl. 3, fig. 17.
Original diagnosis: Spindle-shaped, inflated, lacking horn.
Base partially constricted; aperture half of maximum
diameter of test. Test surface has ten to fourteen longitudinal
plicae, with two to three longitudinal rows of circular pores
between flanking plicae.
Original description: Test spindle-shaped, multisegmented,
final chamber partially constricted at base, aperture half
of maxmum diameter of test. Apical horn lacking on all
specimens examined. Cephalis small and imperforate. All
chambers, except for the final one or two, increase rapidly
in width; final chamber(s) slightly constricted. Ten to
fourteen longitudinal plicae superimposed on surface of
test; plicae mostly continuous from thorax to aperture.
Two to three longitudinal rows of circular pores between
adjacent plicae. Pores usually arranged diagonally, but
sometimes horizontally. Pore size increases very slightly
from apex to base.
Original remarks: This species is larger than Protunuma
fusiformis Ichikawa & Yao, has a larger aperture, fewer
plicae and larger pores; but like P. fusiformis it too is quite
344
variable, having both short broad forms and more elongate
‘slender’ ones. Differs from P. costata (Heitzer) in having
a larger terminal aperture, fewer rows of pores between
adjacent plicae, and these pores are larger.
Further remarks: Protunuma paulsmithi differs from typical Middle and Late Jurassic Protunuma species in having a
widely open instead of a strongly constricted aperture. This
species, however, lacks a perforate basal appendage, characteristic of Unuma Ichikawa & Yao. Protunuma paulsmithi
seems to be the oldest representative of the genus.
Measurements (µm):
Based on 19 specimens.
Maximum length
Maximum width
HT
216
121
Av.
179
107
Max.
216
121
Min.
130
98
Etymology: Named in honour of Dr. P. L. Smith, of the
University of British Columbia, for his contributions to the
study of Jurassic ammonite biostratigraphy.
Type locality: GSC locality C-080579, Whiteaves Formation, Creek locality, Maude Island, Queen Charlotte Islands, British Columbia.
Occurrence: Whiteaves and Phantom Creek formations,
Queen Charlotte Islands.
Plate PRU01. Protunuma paulsmithi Carter. Magnification x300, except Fig. 1b(H) x500. Fig. 1(H)a, b. Carter et al.
1988, pl. 6, figs. 9, 12. Fig. 2. Carter & Jakobs 1991, pl. 3, fig. 17.
345
Genus: Pseudocrucella Baumgartner 1980
Type species: Crucella sanfilippoae Pessagno 1977a
Synonymy:
medullary shell is on one side axially attached to the cortical
shell. On the other side it is surrounded by cortical space.
Primary canals are large, with a vertical axis of symmetry,
surrounded by small, less regularly distributed canals which
connect with the cortical space (see text-fig. 4K).
Original description: Test as with subfamily composed of
4 rays at right angles, usually with tapering tips and long
triradiate central spines. Cortical shell composed of 2 lateral
and 1 to 3, sometimes merging, median external beams on
each side connected by transverse bars with more or less
developed nodes at intersections. Pores circular, rectangular
or parallelogram-shaped in 2 or more partly continuous
rows. Central area with irregular meshwork, nodose with
smaller pores, or with a depression exposing the medullary
shell. Lateral sides exposing the medullary rays with 2 or 3
paired or alternating rows of circular or rectangular pores.
Cross section of rays rectangular or square. The discoidal
Original remarks: Pseudocrucella n. gen. differs from
other four-rayed hagiastrids by its inner structure, by a
rectangular cross section of rays and less regularly arranged pore rows on top and bottom sides. The described
internal structure has been reconfirmed in topotypes of
P. sanfilippoae from Point Sal (NSF 907, Pessagno collection) (pl. 8, figs. 23-24).
1980 Pseudocrucella n. gen. – Baumgartner, p. 291.
1982b Pseudocrucella Baumgartner – De Wever, p. 239.
1984b Pseudocrucella Baumgartner – Blome, p. 351.
1995a Pseudocrucella Baumgartner – Baumgartner et al., p. 442.
Included species:
PDC03 Pseudocrucella ornata De Wever 1981b
3126 Pseudocrucella sanfilippoae (Pessagno) 1977a
PDC04 Pseudocrucella sp. C sensu Carter 1988
Pseudocrucella ornata De Wever 1981b
Species code: PDC03
Synonymy:
1981b Pseudocrucella (?) ornata n. sp. – De Wever, p. 32, pl. 2,
figs. 1-6.
1982b Pseudocrucella (?) ornata De Wever – De Wever, p. 240,
pl. 22, figs. 1-6.
1987b Pseudocrucella jurassica n. sp. – Yeh, p. 29, pl. 2, figs. 5, 19.
1988 Pseudocrucella sp. A – Carter et al., p. 29, pl. 7, figs. 8-9.
1991 Pseudocrucella sp. A, n. sp. – Carter & Jakobs, p. 344, pl. 2,
fig. 13.
Original description: Form with four orthogonal arms
and patagium (sometimes poorly developed or preserved).
Subcylindrical arms ending in spines, triradiate in crosssection at base and distally rounded. In cross-section,
arms show three primary canals that are subtriangular in
shape, surrounding the primary beam (pl. 2, fig. 6). Three
small secondary canals prolongate the primary blades. This
structure resembles that described by P. O. Baumgartner
(1980, text-fig. 4, C) for Tetratrabs gratiosa. Arms, on
surface, show three (or four) longitudinal beams, connected
by transverse bars, aligned from one beam to the other thus
delimiting all orthogonal network. Well-developed node
present where beam and bar intersect. Central part of shell
inflated with few nodes on most specimens.
Original remarks: This species differs from “Pseudocrucella
sp. C” cited by Baumgartner (1980, pl. 8, fig. 11) by its more
regularly arranged pores, more massive nodes and the
presence of a patagium.
Histiastrum elizabethae Rüst (1898, p. 30) has more spindle-shaped, slim arms that do not have aligned pores on
346
their central part. Spines extend from arms without abrupt
change in outline. And finally, one does not know the inner
structure.
Histiastrum valanginica Aliev (1965, p. 33) has a central
lacuna while P. (?) ornata n. sp. has an inflated center.
This species is tentatively assigned to Pseudocrucella
because it does seem to have all the criteria of the genus. It
has the proper external architecture, but the primary and
secondary canals with their respective proportions evoke
the Tritrabinae structure.
Measurements (µm):
Based on 8 specimens.
Av. Min. Max. HT
Total length (of two rays without spines) 217 220 310 264
Length of spines of rays
55 54
57
54
52 40
62
55
Width of rays
Etymology: From Latin ornatus, -a, -um (adj.) = adorned,
decorated.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Nicely Formation, Oregon; Phantom Creek Formation, Queen Charlotte
Islands, British Columbia.
Plate PDC03. Pseudocrucella ornata De Wever. Magnification x200. Fig. 1(H). De Wever 1981b, pl. 2, fig. 1.
Fig. 2. Carter & Jakobs 1991, pl. 2, fig. 13. Fig. 3. Carter et al. 1988, pl. 7, fig. 8.
347
Pseudocrucella sanfilippoae (Pessagno) 1977a
Species code: 3126
Synonymy:
1977a Crucella sanfilippoae n. sp. – Pessagno, p. 72, pl. 2,
figs. 15-16.
1982 Crucella sanfilippoae Pessagno – Aita, pl. 3, fig. 9.
1980 Pseudocrucella sanfilippoae (Pessagno) – Baumgartner,
p. 291, pl. 8, figs. 1, 23-24.
1981 Pseudocrucella sanfilippoae (Pessagno) – Kocher, p. 88,
pl. 16, fig. 1.
1984 Pseudocrucella sanfilippoae (Pessagno) – Baumgartner,
p. 781, pl. 7, fig. 17.
1988 Pseudocrucella sanfilippoae (Pessagno) – Carter et al.,
p. 29, pl. 7, figs. 1, 4.
1989 Pseudocrucella (?) sp. A – Hori & Otsuka, pl. 4, fig. 5.
1995a Pseudocrucella sanfilippoae (Pessagno) – Baumgartner et
al., p. 444, pl. 3126, figs. 1-3.
Original description: Meshwork with linearly arranged
square pore frames having massive nodes at their corners.
Spines triradiate in axial section proximally and circular in
axial section distally.
Original remarks: Crucella sanfilippoae n.sp. differs from
C. messinae by virtue of its linearly arranged square pore
frames and the structure of its spines.
Measurements (µm):
Based on 10 specimens.
Length of ray
Width of ray
Length of spines
Min.
100
50
55
Max.
170
60
120
Etymology: This species is named for Annika Sanfilippo
(Scripps Institution of Oceanography) in honor of her
contributions to the study of Jurassic Radiolaria
Type locality: Sample NSF 907, Point Sal, Santa Barbara
County, California.
Occurrence: Worldwide.
Pseudocrucella sp. C sensu Carter 1988
Species code: PDC04
Synonymy:
1988 Pseudocrucella sp. C – Carter et al. p. 30, pl. 7, fig. 7.
2003 Pseudocrucella sp. C sensu Carter – Goričan et al., p. 293,
pl. 1, fig. 20.
Remarks: This species is characterized by long slender rays
and a rather irregular arrangement of pores. In Carter et
al. (1988) it was considered identical with Pseudocrucella
sp. C of Baumgartner (1980, p. 292, pl. 8, figs. 10, 11) but it
differs from the latter by having rounded and not vertical
sides of rays.
Occurrence: Whiteaves and Phantom Creek formations,
Queen Charlotte Islands, British Columbia; Skrile Formation, Slovenia; Tawi Sadh Member of the Guwayza Formation and Musallah Formation, Oman.
Plate PDC04. Pseudocrucella sp. C sensu Carter. Magnification x250. Fig. 1. Carter et al. 1988, pl. 7, fig. 7. Fig. 2. OM,
BR825-3-R09-07. Fig. 3. OM-00-114, 023132. Fig. 4. Goričan et al. 2003, pl. 1, fig. 20. Fig. 5. SI, MM 6.76, 010330.
Fig. 6. SI, MM 6.76, 000518.
348
Plate 3126. Pseudocrucella sanfilippoae (Pessagno). Magnification x200. Fig. 1(H). Pessagno 1977a, pl. 2, fig. 15.
Fig. 2. Carter et al. 1988, pl. 7, fig. 1. Fig. 3. Carter et al. 1988, pl. 7, fig. 4. Fig. 4. JP, NK9-62.
349
Genus: Pseudoeucyrtis Pessagno 1977b
Type species: Eucyrtis (?) zhamoidai Foreman 1973
Synonymy:
Original remarks: Pseudoeucyrtis n. gen. differs from Eucyrtis
Haeckel (type species = E. conoidea Rüst, 1885; see Foreman, 1973, p. 264) by lacking strictures, by having a more
coarsely and densely perforate test, and by being spindle
shaped.
Original description: Test elongate, spindle shaped, multisegmented termination in a closed (?) tube. Cephalis imperforate with short, often massive horn. Remaining chambers coarsely perforate with polygonal pore frames; pore
frames often spinose. Post-cephalic chambers (exclusive
of terminal tube) increasing gradually in height, but somewhat more rapidly in width to middle of test where they
begin to decrease in width. Test devoid of strictures.
Further remarks: Some species may possess weakly developed strictures. Some species lack apical horn (e.g.,
Pseudoeucyrtis safraensis Dumitrica & Goričan n. sp.).
1977b Pseudoeucyrtis n. gen. – Pessagno, p. 58.
1990 Pseudoeucyrtis Pessagno – Yang & Wang, p. 213.
1994 Pseudoeucyrtis Pessagno – O’Dogherty, p. 179.
1997 Pseudoeucyrtis Pessagno – Hull, p. 158.
Included species:
PSE02 Pseudoeucyrtis angusta Whalen & Carter 1998
PSE04 Pseudoeucyrtis busuangaensis (Yeh & Cheng) 1998
PSE03 Pseudoeucyrtis safraensis Dumitrica & Goričan n. sp.
Pseudoeucyrtis angusta Whalen & Carter 1998
Species code: PSE02
Synonymy:
1998 Pseudoeucyrtis angusta n. sp. – Whalen & Carter, p. 73,
pl. 18, figs. 9-12, 16.
? 1998 Pseudoeucyrtis angusta Whalen & Carter – Yeh & Cheng,
p. 31, pl. 9, figs. 3, 11.
Not 2002 Pseudoeucyrtis angusta Whalen & Carter – Whalen &
Carter, p. 138, pl. 16, fig. 3.
Not 2004 Pseudoeucyrtis angusta Whalen & Carter – Matsuoka,
fig. 123.
Original description: Test multicyrtid, cylindrical, with
narrow terminal tube. Cephalis very small, hemispherical
with short horn; horn circular in axial section, sometimes
slightly bifurcate; cephalis with small, polygonal pore
frames usually covered by layer of microgranular silica.
Thorax and abdomen roughly trapezoidal in outline with
small irregularly shaped pore frames usually covered by
a layer of microgranular silica. Seven to ten postabdominal chambers, rectangular, with medium-sized, irregularly
shaped pore frames; chambers very gradually increasing in size to central widest part of test, then gradually
decreasing in width; chambers in central part of test just
slightly wider than first and last postabdominal chambers;
strictures usually weakly developed between postabdomi-
nal chambers in central part of test. Terminal tube usually
open, with numerous circular pores.
Original remarks: Pseudoeucyrtis angusta n. sp. differs
from P. sp. A by possessing weakly developed strictures.
The very narrow elongate test of P. angusta n. sp. distinguishes it from Protokatroma aquila n. sp.
Measurements (µm):
Based on 5 specimens.
HT
Max.
Min.
Mean
Length (excluding horn)
592
592
526
551
Max. width
99
99
86
92
Etymology: Angustus, a, um (Latin; adj.) = narrow, tight.
Type locality: Sample 89-CNA-KUD-16, Sandilands Formation, Kunga Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands.
Pseudoeucyrtis busuangaensis (Yeh & Cheng) 1998
Species code: PSE04
Synonymy:
1998 Protokatroma busuangaensis n. sp. – Yeh & Cheng, p. 30,
pl. 7, figs. 2, 3; pl. 9, figs. 4, 5, 24.
1998 Protokatroma sp. aff. P. aquila Whalen & Carter – Yeh &
Cheng, p. 30, pl. 7, fig. 1; pl. 9, figs. 6, 7.
1998 Protokatroma sp. B. – Yeh & Cheng, p. 31, pl. 10, figs. 2, 3.
2004 Pseudoeucyrtis sp. – Hori, pl. 2, figs. 49-51.
Original description: Test slender, slightly swollen at median portion, with short tubular horn. Final post-abdomi350
nal chamber terminating with moderately long cylindrical
closed tubular extension which gently decreases in width
towards its distal end.
Original remarks: This form is characterized by having a
slender test which is slightly inflated at its median portion.
Further remarks: The specimens from the Haliw Formation of Oman (pl. PSE04, figs. 2-7) resemble the holotype
Plate PSE02. Pseudoeucyrtis angusta Whalen & Carter. Magnification x200. Fig. 1(H). Carter et al. 1998, pl. 18,
fig. 10. Fig. 2. Carter et al. 1998, pl. 18, fig. 9.
Plate PSE04. Pseudoeucyrtis busuangaensis (Yeh & Cheng). Magnification x200. Fig. 1(H). Yeh & Cheng 1998,
pl. 9. fig. 4. Fig. 2. OM, Haliw-039-R01-05a. Fig. 3. OM, Haliw-039-R02-07. Fig. 4. OM, Haliw-039-R01-07.
Fig. 5. OM, Haliw-039-R01-06. Fig. 6. OM, Haliw-038-R08-23. Fig. 7. OM, Haliw-039-R01-05.
351
but differ only in being more inflated and the maximum
diameter is in the proximal third of shell rather than in the
middle.
Measurements (µm):
Based on 7 specimens.
HT
Max.
Min.
Mean
Length
(excluding horn)
380
397
380
387
Width
(maximum)
74
77
66
72
Etymology: This form is named after its type locality, Busuanga Island, Philippines.
Type locality: Sample CR91-49E, Liminangcong Chert, a
reddish bedded chert sequence exposed at a road side of
the Coron Highway, about 400 m to the west of milestone
KM35.
Occurrence: Liminangcong Chert, Busuanga Island, Philippines; Haliw (Aqil) Formation, Oman; Japan.
Pseudoeucyrtis safraensis Dumitrica & Goričan n. sp.
Species code: PSE03
Type designation: Holotype specimen R20-04 from sample
BR 485, Tawi Sadh Member, Guwayza Formation, Jabal
Safra; paratype specimen 021416, sample OM-00-251,
Musallah Formation, Al Aridh Group, Jabal Buwaydah,
Oman.
Description: Test multicyrtid, spindle-shaped, elongate.
Cephalis small, hemispherical; lacking apical horn.
Following five to seven segments roughly trapezoidal in
outline, increasing slowly in width as added; segments with
polygonal pore frames arranged diagonally. Remainder
of test slowly decreasing in width, mostly comprising a
long tube; pore frames of tube larger, mostly rectangular,
arranged linearly. Intersegmental constrictions indistinct,
usually weakly developed on proximal part but always
lacking on the terminal portion with linearly arranged
pores.
Remarks: Pseudoeucyrtis safraensis n. sp. differs from
P. angusta Whalen & Carter 1998 and P. busuangaensis
Yeh & Cheng 1998 in having linearly arranged pores. In
P. safraensis the linear pore arrangement characterizes at
least one third of the test. P. safraensis is similar to Foremania
352
sandilandsensis Whalen & Carter 1998 in the alignment of
pores, but differs in being proportionally narrower, distally
more constricted, and in lacking a large, branched apical
horn. Also, the pores of F. sandilandensis are arranged in
both longitudinal and transversal rows (square pattern),
whereas in P. safraensis they are arranged in longitudinal
rows only.
Measurements (µm):
Based on 5 specimens.
HT
Max.
Min.
Mean
Length
670
670
408
--
Max. width
81
100
81
90
Etymology: Named after type locality.
Type locality: Sample BR 485, Tawi Sadh Member, Guwayza Formation, Jabal Safra, Oman.
Occurrence: Tawi Sadh Member of the Guwayza Formation, Sabt and Musallah formations, Oman.
Plate PSE03. Pseudoeucyrtis safraensis Dumitrica & Goričan n. sp. Magnification x200. Fig. 1(H). OM, BR485-R20-04.
Fig. 2. OM, BR706-R12-18. Fig. 3. OM-01-21, 010830. Fig. 4. OM-01-21, 010829. Fig. 5. OM-00-251, 021416.
Fig. 6. OM-00-252, 021822.
353
Genus: Pseudogodia Tekin 1999, emend. Carter herein
Type species: Pseudogodia sonmezi Tekin 1999
Synonymy:
1999 Pseudogodia n. gen. – Tekin, p. 120.
Original description: Test thick, roughly hexagonal in outline composing of big nodes. Each side of the test slightly
convex. Rim of the test mainly includes big nodes (six to
seven) in different size mainly situated in every corner but
sometimes irregular. Nodes separating from each other and
from central nodes by shallow depressions. Central nodes
have same feature those of nodes situated at the side of the
test. Meshwork consisting of irregular polygonal (mainly
trigonal and hexagonal) pore frames and mainly circular
pores in different size. Test possesses two peripheral spines
tapering distally and circular in cross section, usually one
of them robust and ticker than the other could be the polar
spine.
Emended description: Test disc-shaped, thick, subhexagonal in outline. Planar surfaces of test slightly convex,
composed of large raised central area surrounded by one or
more rings of large irregularly-sized tubercules. Tubercules
separated from each other and from central area by
shallow depressions. Meshwork of tubercules consisting of
irregular polygonal (mainly trigonal and hexagonal) pore
frames with irregular circular pores. Test with two or more
peripheral spines.
Original remarks: This genus could be distinguished from
the genus Orbiculiforma Pessagno by having polygonal
outline instead of circular, strong nodes both at rim and
center of the test. It differs also from Cretaceous genus Godia
Wu by possessing polygonal outline instead of circular and
whole test of former composing mainly large nodes, latter
has small pores on the surface of the test.
Emended remarks: This genus is distinguished from Godia
Wu in possessing a polygonal outline. By its polygonal
outline it also differs from Orbiculiforma Pessagno, which
is now restricted to subquadratic forms with four main
spines (see original remarks under Orbiculiformella Kozur
& Mostler). Pseudogodia further differs from Godia Wu
and Orbiculiformella Kozur & Mostler in possessing a
large central area and large raised tubercules around the
periphery.
Etymology: For the similarity to Cretaceous genus Godia
Wu.
Included species:
ORB12 Pseudogodia deweveri Carter n. sp.
Pseudogodia deweveri Carter n. sp.
Species code: ORB12
Synonymy:
1981c G. sp. indet. – De Wever, p. 150, pl. 5, figs. 28, 29.
Type designation: Holotype GSC 111756 from GSC loc.
C 3045686 and paratype GSC 128855 from GSC loc. C080613; Rennell Junction member of the Fannin Formation
(upper lower Pliensbachian).
Description: Test disc-shaped, large and thick, subcircular
to slightly scalloped in outline with short spines radiating
from the periphery in different planes. Upper and lower
surfaces of test with well defined raised central area, width
greater than one-half diameter of test. Central area with
numerous large, slight- to moderately-raised tubercules
(Holotype, pl. ORB12, fig. 1). Outer rim of test covered
with fine spongy pore frames; pore frames in central part of
central area sometimes larger and well defined (Paratype,
pl. ORB12, fig. 2), but pore frames on tubercules always
much smaller. Peripheral spines short, irregularly spaced,
variable in width, circular in axial section.
Remarks: This distinctive species was first recognized in the
Pliensbachian of Turkey (De Wever, 1981c) and mentioned
354
as having a pseudoaulophacid outline similar to Pseudoaulophacus lenticulatus Pessagno, a central thickening, and
peripheral spines.
There are pronounced similarities between Pseudogodia
deweveri n. sp. and Orbiculiformella lomgonensis Whalen
& Carter 1998, both similar in age. The former possesses
large porous tubercules on the raised central area,
whereas the latter has a depressed central area and raised
tubercules around the periphery of the test. Until more
is known, O. lomgonensis is still included with the genus
Orbiculiformella.
Measurements (µm):
Based on 8 specimens.
Diameter of cortical shell
Diameter of central area
HT
363
199
Max.
363
206
Min.
281
160
Mean
322
184
Etymology: Named for Patrick De Wever, Muséum National d’Histoire Naturelle, Paris, to honour his pioneering contribution to the knowledge of Pliensbachian radiolarians
and for his helpful guidance in the author’s early studies of
the late Early Jurassic of Queen Charlotte Islands.
Type locality: Sample 99-CNA-MI-11 (GSC loc. C-304568),
Rennell Junction member of the Fannin Formation, Maude
Island, east of Ells Bay, Skidegate Inlet, Queen Charlotte
Islands, British Columbia.
Occurrence: Fannin Formation, Queen Charlotte Islands;
Gümüslü Allochthon, Turkey.
Plate ORB12. Pseudogodia deweveri Carter n. sp. Magnification x150. Fig. 1(H). QCI, GSC loc. C-304566, GSC 111756.
Fig. 2. QCI, GSC loc. C-080613, GSC 128855.
355
Genus: Pseudoheliodiscus Kozur & Mostler 1972, emend. De Wever 1984
Type species: Pseudoheliodiscus riedeli Kozur & Mostler 1972
Synonymy:
1972 Pseudoheliodiscus n. gen. – Kozur & Mostler, p. 24.
1979 Pseudoheliodiscus Kozur & Mostler, emend. Pessagno
– Pessagno et al., p. 169.
1984 Pseudoheliodiscus Kozur & Mostler emend. – De Wever,
p. 15.
1990 Pseudoheliodiscus Kozur & Mostler, emend. Pessagno
– Kozur & Mostler, p. 189.
Original diagnosis: Shell spongy to delicately latticed, with
two prominent needle-shaped polar spines on the inner
side. Usually with one or two medullary shells which are
not always present. Shell connected directly with a girdle
bearing very long radial spines.
Emended diagnosis: De Wever (1984): Palaeosaturnalinae
with simple ring, with auxiliary and/or subsidiary rays.
Original remarks: This genus is transitional between
Heliodiscus Haeckel 1862, where the spines are not fused to
a ring, and Saturnalinae, where the ring is clearly detached
from the shell.
Further remarks: We use here the emendation of the genus
by De Wever (1984) that takes into account the presence
or absence of polar spines. For Pessagno (in Pessagno et
al., 1979) and Kozur & Mostler (1990) this character has
no value.
Etymology: By similarity with Heliodiscus Haeckel, 1862.
Included species:
SAT16 Pseudoheliodiscus aff. alpinus Kozur & Mostler 1990
sensu Whalen & Carter 2002
SAT07 Pseudoheliodiscus yaoi gr. Pessagno 1981
Pseudoheliodiscus aff. alpinus Kozur & Mostler 1990 sensu Whalen & Carter 2002
Species code: SAT16
Synonymy:
1984 Pseudoheliodiscus (?) spp. – Whalen & Pessagno, pl. 3,
fig. 8, 9.
aff. 1990 Pseudoheliodiscus alpinus n. sp. – Kozur & Mostler,
p. 189, pl. 5, fig. 1, 3, 5-9, 11, 12.
2002 Pseudoheliodiscus sp. aff. P. alpinus Kozur & Mostler
– Whalen & Carter, p. 108, pl. 5, figs. 8, 14.
Original remarks: This species differs from P. alpinus Kozur
& Mostler by having fewer rays between the polar spines
that are shorter and less robust.
Occurrence: San Hipólito Formation, Baja California Sur.
Pseudoheliodiscus yaoi gr. Pessagno 1981
Species code: SAT07
Synonymy:
1981 Pseudoheliodiscus yaoi Pessagno n. sp. [Pseudoheliodiscus
yaoi Pessagno & Poisson n. sp. in fig. captions] – Pessagno
& Poisson, p. 55, pl. 4, fig. 9; pl. 5, figs. 1, 4, 7-9; pl. 13, fig. 2.
1981c Pseudoheliodiscus yaoi Pessagno – De Wever, p. 144,
pl. 5, fig. 1.
1981c Pseudoheliodiscus yaoi Pessagno ? – De Wever, p. 144,
pl. 4, figs. 8-10.
1982b Pseudoheliodiscus yaoi Pessagno – De Wever, p. 224,
pl. 20, fig. 6.
1982b Pseudoheliodiscus yaoi Pessagno ? – De Wever, p. 225,
pl. 20, figs. 1-3.
1982 Pseudoheliodiscus yaoi Pessagno – De Wever & OrigliaDevos, pl. 1, fig. T.
Original description: Test with extremely broad, flat ring
having thirteen to fourteen peripheral spines and about
twelve auxiliary spines. Central spongy cortical shell
occupying most of ring on most specimens; cortical shell
comprised of concentric layers of irregular polygonal
(triangular, tetragonal, pentagonal) pore frames.
Original remarks: Pseudoheliodiscus yaoi, n. sp., differs
from P. riedeli Kozur & Mostler (1972) by having a much
broader ring with shorter peripheral spines. It differs from
356
P. finchi Pessagno (1979) by having a somewhat wider
ring and thirteen to fourteen as opposed to ten or eleven
peripheral spines.
Measurements (µm):
Based on 8 specimens.
HT Max. Min.
Diameter of spongy cortical shell
140 150 135
Diameter of test including cortical shell and
230 260 230
ring, excluding peripheral spines on ring
Width of ring, exclusive of peripheral spines 30
45
30
Etymology: This species is named for Dr. Akira Yao (Osaka
City University) in honor of his contributions to the study
of Parasaturnalidae.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus
Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Ghost Creek
Formation, Queen Charlotte Islands; Drimos Formation,
Greece; Tawi Sadh Member fo the Guwayza Formation,
Oman.
Plate SAT16. Pseudoheliodiscus aff. alpinus Kozur & Mostler sensu Whalen & Carter. Magnification x200.
Fig. 1. Whalen & Carter 2002, pl. 5, fig. 8.
Plate SAT07. Pseudoheliodiscus yaoi gr. Pessagno. Magnification x200. Fig. 1(H). Pessagno & Poisson1981, pl. 4, fig. 9.
Fig. 2. QCI, GSC loc. C-305417, GSC 111757. Fig. 3. QCI, GSC loc. C-080612, GSC 111758.
Fig. 4. OM, BR485-R20-13. Fig. 5. OM, BR485-R20-16.
357
Genus: Pseudopantanellium Yeh 1987b
Type species: Pseudopantanellium floridum Yeh 1987b
Synonymy:
1987b Pseudopantanellium n. gen. – Yeh, p. 50.
Original description: Test spherical to ellipsoidal with
two triradiate polar spines. Meshwork of test comprised of
numerous concentric subspherical rings of pentagonal and
hexagonal pore frames anchoring to pillar-like structure
at pore frame vertices. Size of pore frames decreasing
gradually toward center of spherical test.
Original remarks: Pseudopantanellium n. gen., has superficial resemblance to Pantanellium Pessagno (1977b).
However, Pseudopantanellium can be distinguished from
Pantanellium by having a series of concentric layers of
pentagonal or hexagonal pore frames rather than consisting only of a cortical shell and a medullary shell.
Etymology: From the Latin pseudo = false, and Pantanellium.
Included species:
PPN01 Pseudopantanellium floridum Yeh 1987b
Pseudopantanellium floridum Yeh 1987b
Species code: PPN01
Synonymy:
1987b Pseudopantanellium floridum n. sp. – Yeh, p. 50, pl. 10,
figs. 3, 21; pl. 20, fig. 10.
1987b Pseudopantanellium sp. A – Yeh, p. 51, pl. 2, figs. 7, 13, 24.
1987b Pseudopantanellium sp. B – Yeh, p. 51, pl. 2, fig. 14.
1987b Pseudopantanellium sp. C – Yeh, p. 51, pl. 10, fig. 1, 15.
1987b Pseudopantanellium sp. D – Yeh, p. 51, pl. 10, fig. 2; pl. 23,
figs. 4, 15.
1987 Gn. Sp. indet. – Hattori, pl. 22, fig. 17.
1996 Pseudopantanellium sp. A – Pujana, p. 137, pl. 1, fig. 2.
Original description: Test as with genus, large, spherical
in shape, polar spines relatively thin, medium in length,
triradiate with three ridges alternating with three narrow
grooves, extremely narrow subsidiary grooves occurring
on ridges. Concentric rings of meshwork usually of large
pentagonal and hexagonal pore frames anchoring to pillarlike structure at vertices. Pore frames of outermost layer
large, with prominent thin in [sic] rims and moderately
thick in [sic] sides, with prominent nodes at vertices. Five
to six pore frames visible.
358
Original remarks: Pseudopantanellium floridum n. sp.,
can be distinguished from other Pseudopantanellium spp.
in this report by having a large spherical test with very
symmetrical pore frames.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Maximum
diameter
of test
200
175
200
143
Maximum
width
of spine
30
25
30
17
Maximum
length
of polar spines
150
156
165
145
Etymology: From the Latin floridus = flowery.
Type locality: Sample OR-600M, Hyde Formation at IzeePaulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, Oregon; Ghost
Creek and Fannin formations, Queen Charlotte Islands;
Sierra Chacaicó Formation, Argentina; Japan.
Plate PPN01. Pseudopantanellium floridum Yeh. Magnification x150. Fig. 1(H). Yeh 1987b, pl. 10, fig. 3. Fig. 2. QCI,
GSC loc. C-140495, GSC 111759. Fig. 3. QCI, GSC loc. C-140495, GSC 111760. Fig. 4. QCI, GSC loc. C-304566, GSC
111761.
359
Genus: Pseudopoulpus Takemura 1986
Type species: Pseudopoulpus yamatoensis Takemura 1986
Synonymy:
1986 Pseudopoulpus n. gen. – Takemura, p. 39.
1987b Pseudopoulpus Takemura emend. – Yeh, p. 86.
2003 Pseudopoulpus Takemura – Dumitrica & Zügel, p. 32.
Original description: Shell of one segment, cephalis with
apical horn and three feet. Cephalis subspherical, large and
perforated by many irregularly or hexagonally arranged
pores. Cephalis subdivided usually slightly into two parts
by longitudinal grooves on cephalic surface, which accords
with arches Al. Apical horn, which is a prolongation of A,
usually triradiate and thinner than the three feet. Three
feet, prolongations of two L and D, triradiate and strong.
MB, A, V, D, two L and two l as cephalic skeletal elements
and two arches Al existing. VL, Ll, and lD at collar portion.
Arch AV not existing. V not on the same plane defined by
MB and two L.
Original remarks: Pseudopoulpus, n. gen. differs from the
genera Saitoum Pessagno and Poulpus De Wever, which
belong to the subfamily Poulpinae De Wever, in lack of arch
AV, which is the sagittal ring. Although Pseudopoulpus, n.
gen. has a thick and latticed cephalic shell, this new genus
is tentatively assigned to the family Plagoniidae Haeckel,
emend. Riedel in the present paper; because of its tripod
skeleton and considerably large cephalis. Cephalic skeletal
structures of Cenozoic Plagoniids, however, have not yet
been clarified sufficiently.
Etymology: The genus name, is derived from “pseudo” and
the genus name Poulpus De Wever.
Included species:
2007 Pseudopoulpus acutipodium Takemura 1986
POU01 Pseudopoulpus sp. A sensu Whalen & Carter 2002
Pseudopoulpus acutipodium Takemura 1986
Species code: 2007
Synonymy:
1986 Pseudopoulpus acutipodium n. sp. – Takemura, p. 40, pl. 1,
figs. 5-8.
1987b Pseudopoulpus pessagnoi n. sp. – Yeh, p. 89, pl. 12,
figs. 7-9, 12, 13, 22, 26.
1987b Pseudopoulpus sp. A – Yeh, p. 89, pl. 26, fig. 18.
1987b Pseudopoulpus sp. B – Yeh, p. 89, pl. 26, fig. 14.
1987b Pseudopoulpus sp. D – Yeh, p. 90, pl. 12, fig. 1.
1991 Pseudopoulpus acutipodium Takemura – Carter & Jakobs,
p. 344, pl. 3, fig. 10.
2004 Pseudopoulpus acutipodium Takemura – Matsuoka,
fig. 162.
Original description: Cephalis large and subspherical, with
irregularly or hexagonally arranged usually circular pores
and pore frames. Apical horn thin, short and triradiate
proximally. Three feet strong, straight, triradiate and
sharply pointed.
Measurements (µm):
Based on 5 specimens.
Length of shell including horn and feet
Height of cephalis
Maximum width of shell including feet
Width of cephalis
Min.
175
85
180
95
Max.
240
100
215
120
Etymology: Acutipodium, means sharpened foot.
Type locality: Sample TKN-105, Gujo-Hachiman area in
the Mino terrane, central Japan.
Occurrence: Mino Terrane, Japan; Phantom Creek Formation, Queen Charlotte Islands; Hyde Formation and Warm
Springs member of the Snowshoe Formation, Oregon.
Original remarks: P. acutipodium n. sp. is distinguished from
P. yamatoensis by its sharply pointed feet.
Pseudopoulpus sp. A sensu Whalen & Carter 2002
Species code: POU01
Synonymy:
1984 unidentified Radiolaria – Whalen & Pessagno, pl. 1, fig. 4.
2002 Pseudopoulpus sp. A – Whalen & Carter, p. 130, pl. 13,
figs. 1, 2, 14.
Original remarks: This species is similar to Pseudopoulpus
pessagnoi Yeh 1987, but differs in having a more massive
apical horn.
360
Further remarks: Pseudopoulpus pessagnoi Yeh is now
assigned to Pseudopoulpus acutipodium Takemura.
Occurrence: San Hipólito Formation, Baja California Sur.
Plate 2007. Pseudopoulpus acutipodium Takemura. Magnification x250, except Fig. 5c x500. Fig. 1(H). Takemura 1986,
pl. 1, fig. 5. Fig. 2. Matsuoka 2004, fig. 162. Fig. 3. Carter & Jakobs 1991, pl. 3, fig. 10. Fig. 4. OR600A, 13165.
Figs. 5a-c. OR600A-R03-08a-c.
Plate POU01. Pseudopoulpus sp. A sensu Whalen & Carter. Magnification x250. Fig. 1. Whalen & Carter 2002,
pl. 13, fig. 2.
361
Genus: Pseudoristola Yeh 1987b
Type species: Pseudoristola faceta Yeh 1987b
Synonymy:
1987b Pseudoristola n. gen. – Yeh, p. 95.
Original description: Test multicyrtid, conical to subconical,
with two to six post-abdominal chambers. Cephalis conical
to dome-shaped without horn. Thorax and subsequent
chambers trapezoidal in outline. Earlier chambers covered
with layer of microgranular silica, remaining chambers
consisting of single layer of regular to subregular pentagonal
and hexagonal pore frames. Each post-abdominal chamber
with two to three transverse rows of pore frames. Pore
frames slightly variable in size, usually with largest pores
along partitions. Test with or without poorly developed
circumferential ridges. Final post-abdominal chamber
closing and with latticed bulbous expansion.
Original remarks: Pseudoristola n. gen., differs from Ristola
Pessagno and Whalen (1982) by having a test with pore
frames more irregular in shape, by lacking well-developed
circumferential ridges on post-abdominal chambers, and
by having its final post-abdominal chamber closing with a
large latticed bulbous expansion rather than terminating in
an open tubular extension.
Further remarks: See original remarks under Lantus Yeh.
Etymology: From the Latin pseudo = false, and Ristola.
Included species:
PRL01 Pseudoristola megaglobosa Yeh 1987b
Pseudoristola megaglobosa Yeh 1987b
Species code: PRL01
Synonymy:
1987b Pseudoristola megaglobosa n. sp. – Yeh, p. 96, pl. 14,
fig. 13; pl. 23, figs. 17, 22.
1987b Pseudoristola sp. cf. P. megaglobosa n. sp. – Yeh, p. 96,
pl. 14, fig. 15.
2004 Pseudoristola megaglobosa Yeh – Matsuoka, fig. 93.
Original description: Test conical, without horn, with five to
six post-abdominal chambers. Cephalis conical, remaining
chambers trapezoidal in outline and gradually increasing
in width as added. Earlier chambers closely spaced,
sparsely perforate, covered with layer of microgranular
silica. Post-abdominal chamber consisting of two to three
rows of polygonal pore frames in each chamber. Final postabdominal chamber terminating in large subspherical
expansion. Pore frames subregular, slightly variable in
size, with largest pore frames on bulbous expansion. Test
without prominent circumferential ridges, with H-linked
pattern along joints.
362
Measurements (µm):
Ten specimens measured.
Length of proximal conical
part (= last segment excluded)
Width at base of conical part
Length of last segment
Width of last segment
HT
Mean
Max.
Min.
166
180
198
165
105
133
177
107
145
190
110
160
220
105
132
177
Etymology: Megaglobosa: mega (=large) + globosa-a-um
(=spherical).
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Hyde Formation and Warm Springs member
of the Snowshoe Formation, Oregon; Ghost Creek and
Fannin formations, Queen Charlotte Islands; Musallah
Formation, Oman; Japan.
Plate PRL01. Pseudoristola megaglobosa Yeh. Magnification x200. Fig. 1(H). Yeh 1987b, pl. 23, fig. 22. Fig. 2. Matsuoka
2004, fig. 93. Fig. 3. JP, MNA-10, MA13457. Fig. 4a, b. OM-00-118, 000618, 000619. Fig. 5. QCI, GSC loc. C-304566,
GSC 111762. Fig. 6. QCI, GSC loc. C-080611, GSC 111763. Fig. 7. QCI, GSC loc. C-175309, GSC 111764.
Fig. 8. QCI, GSC loc. C-080613. GSC 111765.
363
Genus: Religa Whalen & Carter 2002
Type species: Religa globosa Whalen & Carter 2002
Synonymy:
2002 Religa n. gen – Whalen & Carter, p. 142.
Original description: Test multicyrtid with cephalis, thorax,
abdomen and usually two post-abdominal chambers.
Cephalis with strong horn. Final post-abdominal chamber
spherical, very inflated, significantly larger than other
chambers, with circumferentially arranged, solid spines at
widest part.
Original remarks: The absence of a tubular extension on the
terminal chamber distinguishes this genus from Podobursa
Wisniowski 1889; emend. Foreman 1973, from Podocapsa
Rüst, 1885; emend. Foreman 1973, and from Katroma
Pessagno and Poisson 1981; emend. De Wever 1982, while
the presence of strong spines circumferentially arranged
along the widest part of the final post-abdominal chamber
distinguishes it from Sethocapsa Haeckel 1881. The very
inflated nature of the final chamber of Religa distinguishes
it from Lantus Yeh 1987b. Arcanicapsa Takemura 1986 is
distinguished from Religa n. gen. by having an inflated
abdominal chamber and irregularly distributed spines.
Etymology: The genus Religa is named from an arbitrary
combination of letters (ICZN, 1985, Appendix. D, pt. VI,
Recommendation 40, p. 201).
Included species:
REG01 Religa globosa Whalen & Carter 2002
REG02 Religa sp. A
Religa globosa Whalen & Carter 2002
Species code: REG01
Synonymy:
1998 Katroma sp. aff. K. pinquitudo Whalen & Carter – Yeh &
Cheng, p. 29, pl. 9, fig. 20.
2002 Religa globosa n. sp. – Whalen & Carter, p. 144, pl. 15,
figs. 2, 3, 8, 9; pl. 18, figs. 14, 15.
2004 Religa globosa Whalen & Carter – Matsuoka, fig. 103.
Original description: Test with small, dome-shaped
cephalis with small pores masked by a thin layer of
microgranular silica. Cephalis with strong horn, subcircular
in axial-section with two short branches irregular in
length and shape. Dome-shaped thorax much larger than
cephalis, covered by a thin layer of microgranular silica.
Abdomen and first post-abdominal chamber trapezoidal
in outline; abdominal and post-abdominal pores more
exposed, surrounded by raised areas of microgranular
silica; thorax, abdomen and first post-abdominal chamber
sometimes defined by transversely-aligned discontinuous
ridges produced by buildup of microgranular silica. Postabdominal chamber spherical, inflated, much larger than
other chambers, closed distally. Pores on globular postabdominal chamber larger medially becoming smaller
towards first post-abdominal chamber and distal part
of test. Circumferential spines usually at widest part of
post-abdominal chamber but sometimes in a more distal
position; spines triradiate in axial section at base becoming
circular in axial section distally.
Original remarks: Religa globosa n. sp. is an abundant
and distinctive species in the upper Sinemurian and
Pliensbachian and as yet, has no comparatives.
Measurements (µm):
Based on 12 specimens
HT
Max.
Min.
Mean
Length (excludes horn)
195
195
154
174
Width (max.)
143
150
120
132
Etymology: Globosus, a, um (Latin: adj.) = globular,
spherical. This species is named for the globular shape of its
post-abdominal chamber.
Type locality: Sample BPW80-30, San Hipólito Formation,
Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Ghost Creek and Fannin formations, Queen Charlotte
Islands; Liminangcong Chert, Philippines; Japan.
Religa sp. A
Species code: REG02
Remarks: This species has a much smaller apical horn than
Religa globosa Whalen and Carter and lacks circumferential
spines.
364
Occurrence: Ghost Creek Formation, Queen Charlotte Islands; Skrile Formation, Slovenia.
Plate REG01. Religa globosa Whalen & Carter. Magnification x200. Fig. 1(H). Whalen & Carter 2002, pl. 15, fig. 2.
Fig. 2. Whalen & Carter 2002, pl. 15, fig. 3. Fig. 3. Matsuoka 2004, fig. 103. Fig. 4. QCI, GSC loc. C-304281, GSC 111766.
Fig. 5. QCI, GSC loc. C-080610, GSC 111767. Fig. 6. QCI, GSC loc. C-304281, GSC 111768.
Plate REG02. Religa sp. A. Magnification x250. Fig. 1. QCI, GSC loc. C-080612, GSC 111769. Fig. 2. QCI, GSC loc.
C-080612, GSC 111770. Fig. 3. SI, MM 5.00, 992106.
365
Genus: Rolumbus Pessagno, Whalen & Yeh 1986
Type species: Rolumbus mirus Pessagno, Whalen & Yeh 1986
Synonymy:
1986 Rolumbus n. gen. – Pessagno, Whalen & Yeh, p. 26.
Original diagnosis: Test as with family but possessing
two massive horns: a vertical horn attached to the vertical
bar and an apical horn attached to the apical bar. Tubular,
velum-like structure extending from base of thorax on
well-preserved specimens (Pl. 1, fig. 2).
Description of family Farcidae Pessagno, Whalen &
Yeh 1986 (p. 22): Test dicyrtid with single layer of latticed
meshwork on both cephalis and thorax. Latticed layer
of cephalis and occasionally proximal portion of thorax
covered by thin outer layer of microgranular silica. Cephalis
large, hemispherical with one horn (e.g., Farcus n. gen.), or
two horns (e.g., Rolumbus n. gen.), which are triradiate in
axial section. Cephalic skeletal elements cyrtoid, including
vertical bar, primary left lateral bar, primary right lateral
bar, median bar, secondary left lateral bar, secondary right
lateral bar, and apical bar (dorsal bar absent). Thorax large,
inflated, with four (rarely five) feet that are triradiate in
axial section. Four feet opposed to two primary lateral and
two secondary lateral bars; fifth foot, if present, opposed to
vertical bar. Base of thorax hemispherical with centrallyplaced circular aperture (mouth) that has an imperforate
rim. Thorax with (e.g., Rolumbus n. gen.) or without
(e.g., Farcus n. gen.) fragile tubular, velum-like structure
extending distally from aperture (mouth) of well-preserved
specimens.
Original remarks: Rolumbus n. gen., differs from Hilarisirex
Takemura and Nakaseko, 1982, by being dicyrtid rather
than tricyrtid, by having only a single layer of latticed
meshwork, and by lacking A-frames. It differs from Farcus
n. gen., by having two rather than one horn and by having
a tubular, velum-like structure extending from the base of
the thorax (see Rolumbus sp. Pl. 1, Fig. 2).
Etymology: Rolumbus (masc.) is a name formed by an
arbitrary combination of letters (ICZN, 1964, Appendix D,
Pt. VI, Recommendation 4, p.113).
Included species:
RBS01 Rolumbus gastili Pessagno, Whalen & Yeh 1986
RBS02 Rolumbus halseyensis Pessagno, Whalen & Yeh 1986
Rolumbus gastili Pessagno, Whalen & Yeh 1986
Species code: RBS01
Synonymy:
1986 Rolumbus gastili n. sp. – Pessagno, Whalen & Yeh, p. 26,
pl. 4, figs. 1, 5, 6, 9, 12-14..
2002 Rolumbus gastili Pessagno, Whalen & Yeh – Whalen &
Carter, p. 124, pl. 11, figs. 4, 9, 13, 16, 17.
Original diagnosis: Cephalis large, dome-shaped with an
irregular layer of microgranular silica; pore frames may
be exposed at base of cephalis. Horns straight; apical horn
longer than vertical horn; horns triradiate in axial section
with narrow, rounded, longitudinal ridges and narrow
grooves; discontinuous, narrow ridges may lie between
three main ridges, extending part way up horn. Thorax
with very irregularly-sized and -shaped tetragonal and
pentagonal pore frames; pore frames arranged in poorlydefined transverse rows separated by ridges. Four feet of
medium length, triradiate in axial section with narrow,
rounded ridges and broad grooves. Mouth circular in
outline. Tubular, velum-like structure may be preserved at
base of thorax.
Original remarks: The moderately-developed transverse
ridges of Rolumbus gastili, n. sp., distinguish it from
Rolumbus halseyensis, n. sp.
366
Measurements (µm):
Numbers of specimens measured are in parentheses, X =
broken.
HT
Mean
Max.
Min.
Length of cephalis
30 25.3 (9) 30 (9) 20 (9)
Length of thorax
85 76.8 (8) 90 (8) 65 (8)
55 49.8 (9) 58 (9) 40 (9)
Width of thorax at top
100 97.3 (9) 110 (9) 75 (9)
Width of thorax at base
Length of apical horn
90 88.3 (9) 108 (9) 60 (9)
Length of vertical horn
20X 52.6 (3) 60 (3) 48 (3)
Distance between horn tips 150 126.6 (3) 140 (3) 120 (3)
Length of foot (maximum) 80 69.2 (7) 80 (7) 50 (7)
Etymology: This species is named for R. Gordon Gastil
(San Diego State University, San Diego, CA) in honor of his
immense contribution to our understanding of the geology
of Baja California.
Type locality: Sample BPW-30, San Hipólito Formation,
Baja California Sur.
Occurrence: San Hipólito Formation, Baja California Sur;
Tawi Sadh Member of the Guwayza Formation, Oman.
Plate RBS01. Rolumbus gastili Pessagno, Whalen & Yeh. Magnification x300. Fig. 1(H). Pessagno, Whalen & Yeh 1986,
pl. 4, fig. 5. Fig. 2. Pessagno, Whalen & Yeh 1986, pl. 4, fig. 1. Fig. 3. OM, BR485-R20-07.
367
Rolumbus halseyensis Pessagno, Whalen & Yeh 1986
Species code: RBS02
Synonymy:
1986 Rolumbus halseyensis n. sp. – Pessagno, Whalen & Yeh,
p. 28, pl. 3, figs. 1, 6, 18, 19.
2002 Rolumbus halseyensis Pessagno, Whalen & Yeh – Whalen
& Carter, p. 124, pl. 11, figs. 5, 8, 12, 14.
Measurements (µm):
Numbers of specimens measured are in parentheses, X =
broken.
HT
Mean
Length of cephalis
35 27.1 (6)
Length of thorax
55 59.6 (6)
60 49.8 (6)
Width of thorax at top
90 75.8 (6)
Width of thorax at base
Length of apical horn
82
79 (4)
Length of vertical horn
44 43.6 (5)
Distance between horn tips 155 131.2 (4)
Length of foot (maximum) 50X 55 (4)
Max.
Min.
35 (6) 15 (6)
80 (6) 50 (6)
55 (6) 44 (6)
90 (6) 55 (6)
100 (4) 54 (4)
58 (5) 28 (5)
152 (4) 98 (4)
90 (4) 30X (4)
Original diagnosis: Cephalis large, hemispherical, with
layer of microgranular silica and small nodes. Horns straight
to slightly curved downward, apical horn much larger than
vertical horn; horns triradiate in axial section with narrow,
rounded, longitudinal ridges and broad grooves; small pores
may be open at base of horns. Thorax with small tetragonal
and pentagonal pore frames, commonly obscured by layer
of microgranular silica?; pore frames arranged in poorlydefined transverse rows separated by transverse ridges.
Four feet of moderate length, triradiate in axial section
with narrow, rounded ridges and broad grooves. Mouth
circular in outline. Tubular velum-like structure commonly
extending from base of thorax.
Type locality: Sample SH-412-14, San Hipólito Formation,
Baja California Sur.
Original remarks: This species is compared to Rolumbus
gastili, n. sp., under the latter species.
Occurrence: San Hipólito Formation, Baja California Sur;
Fannin Formation, Queen Charlotte Islands.
368
Etymology: This species is named for Monte Halsey, which
is located southeast of its type area.
Plate RBS02. Rolumbus halseyensis Pessagno, Whalen & Yeh. Magnification x300. Fig. 1(H). Pessagno, Whalen & Yeh
1986, pl. 3, fig. 1. Fig. 2. Pessagno, Whalen & Yeh 1986, pl. 3, fig. 6. Fig. 3. QCI, GSC loc. C-140495, GSC 111771.
369
Genus: Spongosaturninus Campbell & Clark 1944
Type species: Spongosaturninus ellipticus Campbell & Clark 1944
Synonymy:
1944 Spongosaturninus n. gen. – Campbell & Clark, 497.
1990 Spongosaturninus Campbell & Clark – Kozur & Mostler,
p. 210.
Original description: Spongostylinae with 3 concentric
shells, outer shell spongy, inner a double lattice shell, and
having 2 equal polar spines, distal ends of which are connected by an elliptical ring.
Revised description: By Kozur & Mostler (1990): The shell
consists of a tiny latticed microsphere, a second latticed
medullary shell and a rather large third latticed medullary
shell (or cortical shell) covered by a thick layer of spongy
meshwork that reaches along the peripolar spines on the
ring or even beyond the ring. Ring transversally strongly
elongated elliptical, with distinct ridge on the inner margin
of the ring. Cross-section of the ring therefore triangular
with broad base inside. The ring has mostly 1-3 spines in
each polar region of the long axis, but may be additionally
spined around the whole outer ring margin.
Original remarks: Possibly this new genus was derived
from Saturninus by the development of a spongy, instead
of latticed cortical shell, from Saturnalis by the addition of
a spongy shell to the 2 concentric shells, or from our other
new genus, Spongosaturnalis, by the addition of inner shells
to the spongy shell.
Further remarks: By Kozur & Mostler (1990): Spongosaturninus Campbell and Clark, 1944 is a transitional group
to the Saturnalidae Deflandre, 1953. The third medullary
shell is so large that it can be also regarded as cortical shell
(see Dumitrica, 1985). However, the spongy layer on this
shell is always thicker than the distance between the second and third (medullary) shells. In Saturnalis Haeckel,
1881 disappeared this thick outer spongy layer. By this the
outer latticed medullary shell was transformed into a latticed cortical shell.
Included species:
SAT18 Spongosaturninus bispinus (Yao) 1972
Spongosaturninus bispinus (Yao) 1972
Species code: SAT18
Synonymy:
1972 Spongosaturnalis bispinus n. sp. – Yao, p. 28, pl. 2, figs. 1-9.
1996 Acanthocircus bispinus (Yao) – Yeh & Cheng, p. 106, pl. 2,
figs. 4, 5; pl. 7, fig. 3.
1997 Acanthocircus bispinus (Yao) – Yao, pl. 5, fig. 210.
2004 Spongosaturninus bispinus (Yao) – Matsuoka, fig. 8.
Original diagnosis: Spongosaturnalis with fundamentally
one sharp spine at each opposite narrow end of ring.
Original description: Shell spherical, loosely spongy, composed of irregular meshes. Ring narrow, subovoidal, with
one spine (in some specimens three spines) at each opposite narrow end. Spines short or in some specimens long,
smooth, with sharp tip. Ring generally smooth, in some
specimens with weak ridges on both edges near narrow
ends. Ridge on outer edge disappears at spines and not
on them, and ridge on inner edge becomes obsolete near
proximal part of ring. Polar spines short, smooth, their
extensions into shell change to conical sturdy spines with
fragmentary thorns joining to spongy shell.
Original remarks: This species differs from S. protoformis
in having two (or more) spines on the ring, and in lacking
the clear ridge on the ring. S. bispinus appears to be simlar
to Saturnalis minimus Squinabol (1914, p. 287-288, pl. 22,
fig. 1, and pl. 23, figs. 6a-b; Jurassic, Cittiglio (Laveno),
370
and Middle Cretaceous, Novale (Vicentino), Italy) and
Spongosaturninus parvulus Campbell and Clark (1944b, p. 9,
pl. 3, figs. 1, 3 and 5; Late Cretaceous, Middle California)
in the shape of the saturnalin ring, but differs from them in
having a completely spongy shell.
There is a considerable variation in length of spines
and presence of ridges among specimens. According to
Foreman (1968), the ridge on the saturnalin ring is an
important element in specific classification of Saturnalin
radiolaria. In this species, however, importance of the ridge
is not quite definitive.
Measurements (µm):
Based on 6 specimens.
D. of ring along polar spines
D. of ring transversely
D. of shell
L. of polar spine
L. of spine
B. of ring
HT
158
337
(86)
23
30
8-22
Min.
115
212
80
10
18
8-11
Max.
210
348
185
25
85
13-21
Mean
160
302
102
17.6
36.7
Type locality: Sample IN 11, Kiso River, Inuyama area,
central Japan.
Occurrence: Mino Belt, Japan; Liminangcong Chert, Busuanga Island, Philippines; Tawi Sadh Member of the Guwayza Formation, Oman.
Plate SAT18. Spongosaturninus bispinus (Yao). Magnification x200. Fig. 1(H). Yao 1972, pl. 2, fig. 1. Fig. 2. JP, IYII-52.
Fig. 3. Matsuoka 2004, fig. 8. Fig. 4. OM, BR871-R01-03. Fig. 5. OM, BR871-R01-06. Fig. 6. OM, BR871-R01-09.
Fig. 7. OM, BR871-R02-01. Fig. 8. OM, BR871-R01-04.
371
Genus: Stauromesosaturnalis Kozur & Mostler 1990
Type species: Stauromesosaturnalis schizospinosus Kozur & Mostler 1990
Synonymy:
1990 Stauromesosaturnalis n. gen. – Kozur & Mostler, p. 201.
Original diagnosis: Shell spherical or subspherical with
rounded subquadratic equatorial outline, spongy, consisting
of several concentric layers and a tiny latticed microsphere.
Ring narrow, flat, undifferentiated, with subelliptical to
rounded quadratic outline and numerous, relatively short
peripheral spines. 2 peripolar and 2 auxiliary spines (also
opposite to interspine spaces) are cross-like arranged. In
early forms the peripolar spines are somewhat broader than
the auxiliary spines. In highly evolved forms all 4 spines
have the same size. At least opposite to the peripolar spines,
in higher evolved forms opposite to all 4 inner spines the
ring is concavely incised.
Original remarks: Stauracanthocircus Kozur and Mostler,
1983 emend. has polar spines. Praemesosaturnalis Kozur
and Mostler, 1983 has several small auxiliary spines, not
cross-like arranged with the peripolar spines. This genus
is probably the forerunner of Stauromesosaturnalis n. gen.
However, also a direct derivation from Stauracanthocircus
Kozur and Mostler, 1983 emend. by transformation of the
polar spines into peripolar spines cannot be excluded.
Japonisaturnalis Kozur and Mostler, 1972 has evolved from
Stauromesosaturnalis species with bifurcated peripheral
spines. If the terminal branches of adjacent spines grow
together, a second ring evolved, separated from the primary
ring by a pore ring. The first true Japonisaturnalis Kozur
and Mostler, 1972 is known from the Pliensbachian.
Etymology: According to the peripolar spines (as in Mesosaturnalis Kozur and Mostler, 1983), cross-like arranged
with 2 auxiliary spines.
Included species:
SAT19 Stauromesosaturnalis deweveri Kozur & Mostler 1990
Stauromesosaturnalis deweveri Kozur & Mostler 1990
Species code: SAT19
Synonymy:
1981c Pseudoheliodiscus ? sp. aff. P. concordis – De Wever, p. 142,
pl. 2, fig. 4.
1984 Pseudoheliodiscus(?) sp. – Whalen & Pessagno, pl. 3,
figs. 10, 11.
1990 Stauromesosaturnalis deweveri n. sp. – Kozur & Mostler,
p. 202.
1997 Kozurastrum sp. A – Yao, pl. 5, fig. 205.
2002 Stauracanthocircus sanrafaelensis n. sp. – Whalen & Carter,
p. 108, pl. 6, figs. 1, 2; pl. 17, fig. 3.
2004 Stauromesosaturnalis deweveri Kozur & Mostler
– Matsuoka, fig. 11.
Original diagnosis: Shell globular, spongy. Ring narrow,
flat, undifferentiated, outline rounded subquadratic, opposite to the 4 first order spines concavely incised. 21 short,
triangular peripheral spines. 4 first order inner spines of
equal size in cross-like arrangement, all opposite to interspine spaces. Peripolar and auxiliary spines cannot be separated.
Original remarks: See under Stauromesosaturnalis schizospinosus n. gen. n. sp.
Original remarks under S. schizospinosus: Stauromesosaturnalis schizospinosus n. gen. n. sp. is the forerunner of the
Pliensbachian Japonisaturnalis n. sp. A (= Japonisaturnalis
japonicus sensu De Wever, 1981c, pl. 1, fig. 6). By fusion of
the terminal branches of adjacent peripheral spines a sec-
372
ond ring evolved that encloses together with the primary
ring and the primary peripheral spines of the pore ring. The
Pliensbachian Stauromesosaturnalis deweveri n. sp. displays
terminally unbranched spines, the ring is concavely incised
opposite to all 4 first order spines that have all the same
size.
Measurements (µm):
Holotype from De Wever (1981c), other values from Whalen
& Carter (2002), (n) = number of specimens measured.
Diameter of ring Width of ring
(max.) (6)
(max.) (7)
HT
250 and 260
15-18
Max.
300
23
Min.
255
15
Mean
269
17
Length of peripheral
spine (max.) (7)
14-25
15
4
10
Etymology: In honor of Dr. P. De Wever, Paris, who figured
this species for the first time.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; Tawi Sadh
Member of the Guwayza Formation, Oman; San Hipólito
Formation, Baja California Sur; MinoTerrane, Japan.
Plate SAT19. Stauromesosaturnalis deweveri Kozur & Mostler. Magnification x150. Fig. 1(H). De Wever 1981c, pl. 2,
fig. 4. Fig. 2. OM, BR485-R21-02. Fig. 3. Whalen & Carter 2002, pl. 6, fig. 2. Fig. 4. Whalen & Carter 2002, pl. 6, fig. 1.
373
Genus: Stichocapsa Haeckel 1881
Type species: Stichocapsa jaspidea Rüst 1885
Synonymy:
1881 Stichocapsa n. gen. – Haeckel, p. 439.
Original description:
5d-Tribe: Stichocapsida
Closed, eradiate Stichocyrtida.
A. Obtuse (with cephalis smooth, not spiny).
Further remarks: This genus, as used in modern radiolarian
literature, is polyphyletic. Although it was originally defined
as having a distally closed test, most species still assigned
to Stichocapsa have a constricted last segment with an
aperture.
Etymology: Greek Stichocapsa, jointed capsule.
Included species:
SCP01 Stichocapsa biconica Matsuoka 1991
Stichocapsa biconica Matsuoka 1991
Species code: SCP01
Synonymy:
1991 Stichocapsa biconica n. sp. – Matsuoka, p. 733, Fig. 8. 1a – 5b.
1997 Stichocapsa biconica Matsuoka – Yao, pl. 9, fig. 439.
2004 Stichocapsa biconica Matsuoka – Matsuoka, fig. 87.
conical with a constricted aperture. Collar stricture rather
distinct. Strictures between segments, except for the collar
one, indistinct. Pores small to moderate and circular to
subcircular. Aperture moderate in size, circular.
Original description: Shell of five segments, spindle
shaped. Cephalis hemispherical, poreless, occasionally
with rough surface. Thorax and abdomen truncate conical.
Fourth segment barrel-shaped. The last segment inverted
Original remarks: This species differs from Cyrtocapsa (?)
kisoensis Yao by consisting of five segments rather than four
and by lacking an apical horn.
Genus: Tetraditryma Baumgartner 1980
Type species: Tetraditryma pseudoplena Baumgartner 1980
Synonymy:
1980 Tetraditryma n. gen. – Baumgartner, p. 296.
1993 Saldorfus Pessagno, Blome & Hull n. gen. – Pessagno et al.,
p. 126.
Original description: Test as with subfamily, composed of
4 rays of equal length. Cortical shell composed of 2 strong
lateral and 1 weak median external beams, connected by
short, thin bars branching at right angles to beams, forming
2 rows of paired circular pores. Lateral sides concave, with
3 to 4 alternating horizontal rows of uniform circular to
rhombic pores. Centrally placed discoidal medullary shell
connected by subsidiary beams to cortical shell. Medullary
rays composed of 3 primary canals lie on each top or
bottom side of the medullary shell; they connect with the
cortical space and are confined by rows of subsidiary beams
linking medullary and external beams. Ray tips inflated or
tapered.
Original remarks: Tetraditryma differs from Pseudocrucella
n. gen. and all other four-rayed hagiastrids by the paired
rows of pores on top and bottom surfaces and by the
horizontal symmetry axis of the arrangement of primary
canals. The cortical wall of some species in this genus
seems to be a relict of an additional lateral external beam
on each side which can be observed on early forms of this
subfamily.
Etymology: Greek: tetra, four-, di-, two-, tryma (feminine),
hole - 4 rays with 2 rows of pores.
Included species:
3407 Tetraditryma cf. praeplena Baumgartner sensu Carter
& Jakobs 1991
Tetraditryma cf. praeplena Baumgartner sensu Carter & Jakobs 1991
Species code: 3407
Synonymy:
1991 Tetraditryma cf. praeplena Baumgartner – Carter &
Jakobs, p. 344, pl. 2, fig. 1.
1995a Tetraditryma sp. cf. T. praeplena Baumgartner –
Baumgartner et al., p. 558, pl. 3407, fig. 1.
1997 Tetraditryma cf. praeplena Baumgartner – Yao, pl. 7, fig. 328.
374
Original remarks: Lacks slender triradiate lateral spines
that extend from the ray tips at a 60-70° angle to the ray
axis, but otherwise is very similar to P. praeplena and may
be its immediate ancestor.
Occurrence: Phantom Creek Formation, Queen Charlotte
Islands; Japan.
Etymology: The specific name is derived from the Latin bi
(=two) and conicus-a-um (=conical).
Measurements (µm):
Numbers of specimens measured are in parentheses.
Total height of shell
Maximum width of shell
Diameter of aperture
HT
153
77
9
Max.
153
77
9
Min.
110
57
9
Mean
130
68
9
(15)
(15)
(2)
Type locality: MNA-10, Nanjo Massif, Mino Terrane, central Japan.
Occurrence: Mino Terrane, Japan.
Plate SCP01. Stichocapsa biconica Matsuoka. Magnification x400. Fig. 1(H). Matsuoka 1991, Fig. 8.1a.
Figs. 2-3. Matsuoka 1991, Fig. 8.3-4. Figs. 4a, b. Matsuoka 1991, Fig. 8.2a-b.
Plate 3407. Tetraditryma cf. praeplena Baumgartner sensu Carter & Jakobs. Magnification x150.
Fig. 1. Carter & Jakobs 1991, pl. 2, fig. 1.
375
Genus: Thetis De Wever 1982a, emend. Dumitrica herein
Type species: Thetis oblonga De Wever 1982a
Synonymy:
1982 a Thetis n. gen. – De Wever, p. 195.
? 1984 Eucyrtidiellum n. gen. – Baumgartner, p. 764.
? 1986 Monosera n. gen. – Takemura & Nakaseko, p. 1021.
? 1986 Eucyrtidiellum Baumgartner – Takemura, p. 66.
? 1990 Eucyrtidiellum Baumgartner – Nagai & Mizutani, p. 593.
Original description: Multicyrtid with a stout apical horn
and three thoracic spines. Last segment is prolonged by a
velum.
Emended description: Test composed of four segments: the
first three are strong and rapidly increase in size as added,
the fourth is usually cylindrical, thinner-walled, open or
closed. Cephalis small, poreless with a conical or cylindrical
(never bladed), apical horn and ventral pore in the
prolongation of ventral spine. Initial spicule with MB and
apical, dorsal, and ventral spines. Secondary lateral spines
absent. On outer test wall, L and D expressed as thin spines
descending along thorax. Thorax porous; abdomen much
larger, porous. First postabdominal segment cylindrical,
thinner-walled, straight or undulate in outline; wide open
or tending to close distally.
Original remarks: This genus differs from Ectonocorys by
its hemispherical cephalis.
Further remarks: The original definition of this genus is so
imprecise that it can be applied to many nassellarian genera.
In the emended definition, the genus includes only two of
the three species originally (De Wever, 1982a) included:
T. oblonga De Wever and T. undulata De Wever. T. (?) stolata
De Wever undoubtedly belongs to another genus because it
has a three-bladed apical horn (a characteristic never found
in either Thetis or Eucyrtidiellum), robust bladed spines L
and D, and a dicyrtid test. Such characteristics indicate a
closer affinity to Jacus? anatiformis De Wever and the genus
Anaticapitula Dumitrica & Zügel.
The genus Thetis was not defined as having a ventral pore
on cephalis but this feature is present on the type species
(see further remarks under this species), and De Wever
(1982a) mentioned it with T. undulata.
Thetis is structurally very close to Eucyrtidiellum Baumgartner. The only characteristic that could differentiate
these two genera is the presence of the L and D spines on
thorax. Unfortunately we do not know the initial spicule of
the genus Thetis: it could be similar to Eucyrtidiellum (see
Takemura, 1986) i.e., with L expressed only as short thorns,
or it could have very thin L bars that tend to disappear. The
latter possibility is supported by the presence of the spines
L and D outside test wall. Regardless, the two genera are so
close that Eucyrtidiellum could be a junior synonym of the
genus Thetis, or a subgenus.
Etymology: Dedicated to Thetis, goddess of the sea, granddaughter of Tethys and mother of Achilles.
Included species:
THT01 Thetis oblonga De Wever 1982a
Thetis oblonga De Wever 1982a
Species code: THT01
Synonymy:
1982a Thetis oblonga n. sp. – De Wever, p. 195, pl. 4, figs. 10-14.
1982a Thetis oblonga ? n. sp. – De Wever, p. 196, pl. 4, figs. 15-16.
1982b Thetis oblonga De Wever – De Wever, p. 312, pl. 42,
figs. 1-5.
1982b ? Thetis oblonga De Wever – De Wever, p. 313, pl. 42,
figs. 7, 8.
? 1982 Thetis oblonga De Wever – De Wever & Origlia-Devos,
pl. 1, fig. A.
2002 Thetis oblonga De Wever – Whalen & Carter, p. 138, pl. 16,
fig. 4.
Original description: Three-segmented form with a subcylindrical velum. Cephalis usually imperforate. Cephalis,
thorax, and abdomen with a relatively thick wall; external
surface frequently covered with lumps that may represent
an external silica layer. Apical horn long, circular in cross
section. From the collar section, or the upper thorax, three
376
spines are developed that are extensions of A, Ll and Lr spines
and are linked to the thorax and/or the abdomen by small
bars. The three spines are sub-parallel to the test outline;
being fragile, they are often broken. Pores distributed in
approximately transversal rows.
The velum prolonging abdomen has a thin wall that corresponds to the internal layer of the abdominal wall. It is
cylindrical or has a slight distal constriction.
Original remarks: This species differs from Ectonocorys
spinosa Yao (1979, p. 44, pl. 11, figs. 10-17) by having
a circular apical horn, a smaller cephalis, more slender
thoracic spines closer to the test, and by the absence of
abdominal spines. It differs from Ectonocorys (?) furcillata
n. sp. by lacking a forked apical horn and having spines on
thorax. It differs from Thetis undulata n. sp. by having only
three segments and by the outline of the distal part.
Further remarks: The original description is incomplete
concerning both cephalis and abdomen. The description
does not mention, for instance, the presence of a ventral
pore on the cephalis that is aligned with the ventral spine.
This pore with a protruding rim exists and is visible on the
right side of the cephalis on the paratype illustrated in De
Wever, 1982a, pl. 4, fig. 13 and pl. 4, fig. 16 (pl. THT01,
figs. 2, 3 in this catalogue), directed towards the reader. An
additional topotype specimen, illustrated from the same
position as the former (pl. THT01, fig. 4), clearly shows
this pore. The three thoracic spines representing external
prolongations of spines D, Lr, Ll of the initial spicule are
not always all present. The shape of the abdomen is very
characteristic in this species; it is truncated conical in the
proximal third and cylindrical with straight or concave
sides on the remaining part; the boundary between the two
parts forms a rather characteristic shoulder.
Measurements (µm):
Based on 10 specimens.
Length of apical horn
Width of cephalis
Width of thorax
Width of abdomen
Length of cephalis+thorax
+abdomen (without apical horn)
Total length (including
apical horn and velum)
HT Min. Max. Mean
56
40
64
50
21
16
26
20
37
25
38
34
53
40
64
53
56
50
78
60
144
135
182
157
Etymology: From the Latin oblongus, -a, -um, adj., oblong.
Type locality: Sample 1662D, Gümüslü Allochthon, Taurus Mts., Turkey.
Occurrence: Gümüslü Allochthon, Turkey; San Hipólito
Formation, Baja California Sur.
Plate THT01. Thetis oblonga De Wever. Magnification x400. Fig. 1(H). De Wever 1982a, pl. 4, fig. 10.
Figs. 2, 3. De Wever 1982a, pl. 4, figs. 13, 16. Fig. 4. TR, 1662D-R02-06. Fig. 5. Whalen & Carter 2002, pl. 16, fig. 4.
377
Genus: Thurstonia Whalen & Carter 1998
Type species: Thurstonia minutaglobus Whalen & Carter 1998
Synonymy:
1998 Thurstonia n. gen. – Whalen & Carter, p. 42.
Original description: Test with spherical shell and six
prominent tapering spines; two spines bipolar, at right angles to four equally spaced spines, in radial plane. Cortical shell with triangular and tetragonal pore frames with
rounded nodes at pore frame vertices; large pores sometimes apparent at base of spines. Spines circular or triradiate in axial section.
Original remarks: The internal structure of Thurstonia
n. gen. is not completely understood but because of the suggestion of an internal spicular network (see Pl. 8, Fig. 13),
we tentatively assign this genus to the Subfamily Charlottinae. Thurstonia n. gen., is distinguished from all the included genera of the Subfamily Charlotteinae by having six
prominent spines, four of which are in the same plane.
Etymology: Thurstonia n. gen. is named for the “Nellie
G. Thurston”, a schooner owned by the Pacific Fish and
Cold Storage Company and a regular visitor to the fishing
grounds of the Queen Charlotte Islands in the early 1900s.
Included species:
THU01 Thurstonia gibsoni Whalen & Carter 1998
THU04 Thurstonia timberensis Whalen & Carter 1998
Thurstonia gibsoni Whalen & Carter 1998
Species code: THU01
Synonymy:
1998 Thurstonia gibsoni n. sp. – Whalen & Carter, p. 42, pl. 6,
figs. 1, 2.
2002 Thurstonia sp. aff. T. timberensis Whalen & Carter – Tekin,
p. 188, pl. 3, fig. 20.
2004 Thurstonia gibsoni Whalen & Carter – Hori et al., pl. 5, fig. 6.
Original description: Test with small, spherical cortical
shell and six narrow spines. Cortical shell composed of
small, irregularly shaped and distributed triangular and
tetragonal pore frames with prominent rounded nodes at
pore frame vertices. Spines approximately equal in length,
circular in axial section with length of each spine slightly
less than diameter of cortical shell.
Original remarks: The small cortical shell and very delicate
spines (circular in axial section) of Thurstonia gibsoni n. sp.
distinguish it from T. timberensis n. sp.
378
Measurements (µm):
Based on 15 specimens.
Maximum diameter
of cortical shell
135
135
96
115
Length
of longest spine
85
131
66
80
HT
Max.
Min.
Mean
Etymology: This species is named in honor of Gibson (Gib)
Carter, Vernonia, Oregon, who assisted the authors in many
aspects of this research.
Type locality: Sample 89-CNA-SKUD-27, Sandilands Formation, southeast side of Kunga Island, Queen Charlotte
Islands, British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands; Hocaköy Radiolarite, Turkey; Tawi Sadh Member of
the Guwayza Formation, Oman; Japan.
Plate THU01. Thurstonia gibsoni Whalen & Carter. Magnification x200. Fig. 1(H). Carter et al. 1998, pl. 6, fig. 1.
Fig. 2. QCI, GSC loc. C-080611, GSC 111772. Fig. 3. OM, BR682-R10-20.
379
Thurstonia timberensis Whalen & Carter 1998
Species code: THU04
Synonymy:
1989 Genus 4 spp. – Hattori, pl. 17, figs. B, C.
1990 Beturiella ? sp. – Nagai, pl. 6, figs. 1, 2.
1998 Thurstonia timberensis n. sp. – Whalen & Carter, p. 43,
pl. 6, figs. 3, 4, 5, 10.
1998 Thurstonia sp. B – Yeh & Cheng, p. 11, pl. 8, fig. 8.
Original description: Test with medium-sized spherical
cortical shell with six strongly tapering spines. Cortical
shell composed of small- to medium-sized, irregularly
shaped and distributed triangular and tetragonal pore
frames with prominent, rounded nodes at pore frame
vertices; pore frame bars very thin in Y direction and very
thick in Z direction; very large pores present at contact of
spine with cortical shell. All spines approximately equal
in length, usually longer than diameter of cortical shell.
Spines triradiate in axial section proximally with narrow,
rounded longitudinal ridges and broad, rounded, tapering
longitudinal grooves; spines becoming circular in axial
section distally.
Original remarks: The broad, strongly tapering spines and
smaller, more delicate pore frames of Thurstonia timberensis n. sp. distinguish it from Thurstonia minutaglobus n. sp.
380
Measurements (µm):
Based on 11 specimens.
Maximum diameter
of cortical shell
169
169
131
152
Length of longest spine
156
188
122
151
HT
Max.
Min.
Mean
Etymology: This species is named for Timber Road, the
road leading to Beth Carter’s ranch in Vernonia, Oregon
where much of the biostratigraphic work for this paper was
completed.
Type locality: Sample 87-CNA-KUD-14, Sandilands Formation, Kunga Island, Queen Charlotte Islands, British
Columbia.
Occurrence: Sandilands, Ghost Creek and Fannin formations, Queen Charlotte Islands; Fernie Formation, Williston Lake, NE British Columbia; Hyde Formation, Oregon;
Tawi Sadh Member of the Guwayza Formation, Oman; Liminangcong Chert, Philippines; Japan.
Plate THU04. Thurstonia timberensis Whalen & Carter. Magnification Figs. 1-4 x150 (scale bar A) Figs. 5-8 x200 (scale
bar B). Fig. 1(H). Carter et al. 1998, pl. 6, fig. 3. Fig. 2. QCI, GSC loc. C-304281, GSC 111773. Fig. 3. NBC, GSC loc. C305208, GSC 111810. Fig. 4. QCI, GSC loc. C-304566, GSC 111774. Fig. 5. JP, MNA-10, MA11342.
Fig. 6. OR600A-R02-02. Fig. 7. OR600A-R03-02. Fig. 8. BR871-R02-17.
381
Genus: Trexus Whalen & Carter 1998
Type species: Trexus dodgensis Whalen & Carter 1998
Synonymy:
1998 Trexus n. gen. – Whalen & Carter, p. 81.
pore frames rather than square to rectangular, linearly
arranged pore frames on both the inner and outer layers
(see Pl. 24, fig. 22).
Canutus? beehivensis and C.? ingrahamensis described by
Carter from the Rhaetian part of the Sandilands Formation
in Queen Charlotte Islands, are herein reassigned to the
genus Trexus Whalen and Carter n. gen.
Original description: Robust, inflated, dome-shaped
multicyrtid, with small horn. Test lacks strictures. Thick
multilayered wall: inner layer composed of large, fragile,
irregularly polygonal pore frames; outer layers more massive,
composed of irregularly sized and shaped polygonal pore
frames; outer layers connected by irregularly sized and
shaped pillars; small nodes at pore frame vertices on outer
latticed layer.
Etymology: Trexus n. gen. is a name formed by an arbitrary combination of letters (ICZN, 1985, Appendix D,
pt. VI, Recommendation 40, p. 201).
Original remarks: Trexus n. gen. differs from Canutus
Pessagno and Whalen by possessing irregularly polygonal
Included species:
TRX01 Trexus dodgensis Whalen & Carter 1998
Trexus dodgensis Whalen & Carter 1998
Species code: TRX01
Synonymy:
Original remarks: The irregularly sized and shaped pore
frames of the outer layers of T. dodgensis n. sp. distinguish
it from all other Jurassic multicyrtid Nassellariina with
multiwalled construction. T. dodgensis n. sp. differs from
T. beehivensis (Carter) and T. ingrahamensis (Carter) by
having a more dome-shaped test and it lacks a terminal
tube.
Original description: Test large, dome-shaped, inflated
with three to four postabdominal chambers. Cephalis small,
hemispherical to dome-shaped, with short horn. Thorax and
abdomen trapezoidal in outline. Postabdominal chambers
subrectangular in outline, increasing very gradually in
width, more rapidly in height as added. Outer latticed
layer consisting of irregularly sized and shaped polygonal
pore frames (mostly pentagonal and hexagonal) with small
nodes at pore frame vertices. Inner latticed layer composed
of delicate, polygonal pore frames.
Measurements (µm):
Based on 7 specimens.
1998 Trexus dodgensis n. sp. – Whalen & Carter, p. 82, pl. 24,
figs. 11, 12, 16, 22, 23.
2001 Trexus dodgensis Whalen & Carter – Gawlick et al., pl. 2,
fig. 26; pl. 6, fig. 5.
2002 Trexus dodgensis Whalen & Carter – Suzuki et al., p. 184,
fig. 9 D.
2004 Canutus sp. – Matsuoka, fig. 213.
Length (excluding horn)
150
173
150
155
Maximum width
120
143
120
132
HT
Max.
Min.
Mean
Etymology: This species is named for Dodge Point, Queen
Charlotte Islands, British Columbia, located to the southeast of the type locality.
Genus: Triactoma Rüst 1885
Type species: Triactoma tithonianum RÜST 1885 (subsequent designation by Campbell, 1954).
Synonymy:
Further remarks: Herein we follow Baumgartner et al.
(1995a) who considered Tripocyclia Haeckel (as emended
by Pessagno & Yang) to be a synonym of Triactoma.
Original description: Spherical latticed test with three long,
slender spines arranged in one plane. 10 rows of round
pores, 10 per row. Not frequent. (Haeckel, 1881, p. 457)
Spined Phacodiscida, with marginal spines situated in the
equatorial plane of the lens; with three equidistant spines
with the medullary shell single, and without a spiny zone.
Included species:
3409 Triactoma jakobsae Carter 1995
TCA01 Triactoma rosespitensis (Carter) 1988
1885 Triactoma n. gen. – Rüst, p. 289.
1989 Tripocyclia Haeckel, emend. Pessagno & Yang – Pessagno et
al., p. 212.
382
Type locality: Sample QC-574, Sandilands Formation,
north side of Kunga Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands; Pucara Group, Peru; Dürrnberg Formation, Austria;
Japan.
Plate TRX01. Trexus dodgensis Whalen & Carter. Magnification Fig. 1 x300, Fig. 2. x150. Fig. 1(H). Carter et al. 1998,
pl. 24, fig. 16. Fig. 2. QCI, GSC loc. C-304567, 111790.
383
Triactoma jakobsae Carter 1995
Species code: 3409
Synonymy:
1988 Tripocyclia sp. B – Carter et al., p. 27, pl. 10, figs. 2-3.
1989 Triactoma sp. – Hattori & Sakamoto, pl. 7, figs. J, K.
1989 Tripocyclia ? sp. – Hattori, pl. 45, fig. L.
1989 Tripocyclia sp. A – Pessagno & Yang, p. 229, pl. 2, fig. 10.
1995a Triactoma jakobsae Carter n. sp. – Baumgartner et al.,
p. 588, pl. 3409, figs. 1-3.
1996 Tripocyclia yaoi n. sp. – Yeh & Cheng, p. 102, pl. 2, fig. 3,
pl. 5, figs. 1, 2, 4, 5, 11.
1996 Tripocyclia sp. aff. T. yaoi n. sp. – Yeh & Cheng, p. 102,
pl. 5, figs. 7, 8, 10.
1997 Triactoma jakobsae Carter – Yao, pl. 1, fig. 25.
2004 Triactoma jakobsae Carter – Suzuki & Ogane, pl. 4, fig 1.
Original description: Cortical shell spherical to subspherical. Outer latticed layer thick, composed of small, mostly
hexagonal pore frames with small, rather sharp nodes at
vertices of pore frames. Spines short to moderate in length
but length never exceeds diameter of cortical shell. Spines
robust, triradiate, composed of longitudinal ridges and
grooves. Ridges rounded with small to medium-sized tear
drop-shaped subsidiary grooves tapering towards distal
part of spine. Longitudinal grooves narrow and deep, tapering distally but open to spine tips. Spine tips bluntly terminating with the three longitudinal ridges turned outward to
form crown-like structures. On well preserved specimens a
short, robust, central spine extends beyond ridge terminations. Cortical buttresses weakly developed.
Original remarks: Triactoma jacobsae n.sp. is larger than
Tripocyclia wickiupensis Pessagno & Yang in all respects
and further differs from that species in having a spherical to
subspherical cortical shell composed of larger pore frames,
and in having spine tips with better developed triradiate
structures.
Further remarks: Forms with less massive spines (see synonymy) are now included in Triactoma jakobsae Carter.
Measurements (µm):
Based on 13 specimens.
Diameter of cortical shell
Length secondary spines
Width of spine base
HT
155
127
59
Av.
154
120
53
Max.
169
135
60
Min.
150
99
43
Etymology: Named for Dr. Giselle K. Jakobs for her contribution to the biostratigraphy of the Toarcian and Aalenian
of Western North America.
Type locality: GSC locality C-176579- Section 12, Phantom
Creek Formation, Yakoun River, central Graham Island,
Queen Charlotte Islands, British Columbia.
Occurrence: Phantom Creek Formation, Queen Charlotte
Islands; Warm Springs member of the Snowshoe Formation, Oregon; Liminangcong Chert, Philippines; Japan.
Triactoma rosespitensis (Carter) 1988
Species code: TCA01
Synonymy:
1987b Tripocyclia sp. A – Yeh, p. 52, pl. 3, fig. 9; pl. 26, fig. 8.
1988 Tripocyclia rosespitense Carter n. sp. – Carter et al., p. 27,
pl. 10, fig. 1.
? 1990 Tripocyclia sp. – Hori, Fig. 9.41.
1996 Tripocyclia sp. cf. T. yaoi n. sp. – Yeh & Cheng, p. 102, pl. 5,
figs. 3, 6, 9, 12.
2002 Triactoma rosespitensis (Carter) – Whalen & Carter,
p. 112, pl. 8, fig. 9.
Original diagnosis: Test small, with uniform, mostly hexagonal, pore frames and three slender tribladed spines.
Original description: Test small, spherical, and slightly
flattened in plane of spines. Three triradiate spines are
slender with alternating ridges and grooves. Ridges narrow
and rounded; grooves about twice width of ridges. Terminal
portion of spines normally pointed, but occasionally ridges
widen at tip to produce a crown-like extension. Pore frames
small, most are hexagonal, a few pentagonal.
Original remarks: This species is similar in pore frame
size and shape, and spine structure to Tripocyclia trigonium (Rüst), (Rüst, 1885, p. 23, Pl. 30(5), fig. 3; Parona,
384
1890, p. 155, Pl. 2, fig. 15; and Pessagno, 1977a, p. 80, Pl. 7,
figs. 6, 7), but differs in having a distinctly spherical rather
than subtriangular test shape.
Measurements (µm):
Based on 7 specimens.
Diameter of test
Length of longest spine
(on 4 complete specimens)
HT
141
Av.
143
Max.
150
Min.
140
136
126
148
110
Etymology: Named for Rose Spit, on the northeastern tip
of Graham Island.
Type locality: GSC locality C-080597, Phantom Creek Formation, Yakoun River, Graham Island, Queen Charlotte
Islands, British Columbia.
Occurrence: Phantom Creek Formation, Queen Charlotte
Islands, British Columbia; Nicely Formation and Warm
Springs member of the Snowshoe Formation, Oregon; San
Hipólito Formation, Baja California Sur; Liminangcong
Chert, Philippines; Japan.
Plate 3409. Triactoma jakobsae Carter. Magnification x150. Fig. 1(H). Baumgartner et al. 1995a. pl. 3409, fig. 1.
Fig. 2. Baumgartner et al. 1995a. pl. 3409, fig. 3. Fig. 3. Baumgartner et al. 1995a. pl. 3409, fig. 2.
Plate TCA01. Triactoma rosespitensis (Carter). Magnification x200. Fig. 1(H). Carter et al. 1988, pl. 10, fig. 1.
Fig. 2. QCI, GSC loc. C-304567, GSC 11181.Fig. 3. Matsuoka 2004, fig. 73. Fig. 4. Whalen & Carter 2002, pl. 8, fig. 9.
385
Genus: Trillus Pessagno & Blome 1980
Type species: Trillus siedersi Pessagno & Blome 1980
Synonymy:
1980 Trillus n. gen – Pessagno & Blome, p. 248.
possessing a well-developed median band. The phylogenetic
relationship of Trillus to other genera of Pantanellinae is
discussed elsewhere in this report.
Original description: Cortical shell with well developed
raised median band comprised of pore frames which are
greatly thickened in Z direction (text-fig. 5). Pore frames of
raised median band lacking massive secondary spines.
Etymology: Trillus is a name formed by an arbitrary combination of letters (ICZN, 1964, Appendix D, pt. 6, recommendation 40, p. 113).
Original remarks: Trillus n.gen. differs from Zartus n.gen.
in possessing a raised median band without large, massive
secondary spines. It differs from Pantanellium Pessagno in
Included species:
TRL01 Trillus elkhornensis Pessagno & Blome 1980
TRL02 Trillus seidersi Pessagno & Blome 1980
Trillus elkhornensis Pessagno & Blome 1980
Species code: TRL01
Synonymy:
1980 Trillus elkhornensis n. sp. – Pessagno & Blome, p. 249,
pl. 6, figs. 11, 12, 16, 20, 25; pl. 9, fig. 11.
1982 Trillus elkhornensis Pessagno & Blome – Nishizono et al.,
pl. 1, fig. 10.
1987b Trillus elkhornensis Pessagno & Blome – Yeh, p. 37, pl. 5,
fig. 5.
1987b Trillus sp. aff. T. elkhornensis Pessagno & Blome – Yeh,
p. 37, pl. 5, fig. 7.
1987b Trillus sp. A – Yeh, p. 37, pl. 5, figs. 5, 25.
1989 Trillus elkhornensis Pessagno & Blome – Hattori &
Sakamoto, pl. 4, fig. B.
1989 Trillus sp. C – Hattori & Sakamoto, pl. 4, fig. G.
1989 Trillus sp. I – Hattori & Sakamoto, pl. 4, fig. N.
1989 Trillus elkhornensis Pessagno & Blome – Hattori, pl. 8,
figs. E, F.
1989 Trillus elkhornensis Pessagno & Blome – Hori & Otsuka,
pl. 4, fig. 15.
1990 Trillus elkhornensis Pessagno & Blome – De Wever et al.,
pl. 3, fig. 12.
1990 Trillus elkhornensis Pessagno & Blome – Hori, Fig. 8.30.
1992 Trillus elkhornensis Pessagno & Blome – Sashida, pl. 2,
figs. 21, 22.
1993 Trillus sp. – Fujii et al., pl. 1, fig. 6.
1996 Trillus elkhornensis Pessagno & Blome – Yeh & Cheng,
p. 98, pl. 8, fig. 2.
1996 Trillus sp. cf. T. elkhornensis Pessagno & Blome – Yeh &
Cheng, p. 98, pl. 6, figs. 8, 12.
1996 Trillus sp. B – Yeh & Cheng, p. 98, pl. 8, fig. 1.
1996 Trillus elkhornensis Pessagno & Blome – Pujana, p. 137,
pl. 1, fig. 1.
1997 Trillus elkhornensis Pessagno & Blome – Hori, pl. 1, fig. 14.
1997 Trillus elkhornensis Pessagno & Blome – Yao, pl. 3, fig. 147.
2003 Trillus elkhornensis Pessagno & Blome – Goričan et al.,
p. 291, pl. 1, fig. 2.
2003 Trillus elkhornensis Pessagno & Blome – Kashiwagi &
Kurimoto, pl. 4, figs. 10, 12, not fig. 11.
2004 Trillus elkhornensis Pessagno & Blome – Hori, pl. 4, fig. 41,
pl. 13, fig. 55; pl. 23, ? fig. 25.
2004 Trillus elkhornensis Pessagno & Blome – Matsuoka, fig. 16.
2005 Trillus elkhornensis Pessagno & Blome – Kashiwagi et al.,
pl. 6, fig. 8.
386
Original description: Cortical shell subspherical with
well-developed median band; pore frames pentagonal and
hexagonal, about equal in number, with poorly developed
nodes of low relief at vertices. Bars of pore frames of equal
thickness along Y and Z (text-fig. 5). Five to 6 pore frames
visible along AA’; 6 along BB’. Both polar spines triradiate in
axial section, shorter spine about 3/4 length of longer spine;
spines comprised of massive, rather wide ridges alternating
longitudinally with somewhat wider grooves.
Original remarks: Trillus elkhornensis, n. sp., is compared
with T. seidersi, n. sp., under the latter species.
Measurements (µm):
Based on 9 specimens. System of measurement shown in
text-figure 5 of Pessagno & Blome (1980).
AA’
113
99
113
88
A’S’
125
113
125
88
AS
94
88
94
81
BB’
100
97
106
88
cc’
25
29
31
25
dd’
31
28
31
25
HT
Av.
Max.
Min.
Etymology: T. elkhornensis is named for Elkhorn Creek
near its type locality.
Type locality: Sample OR 536, Nicely Formation, northeast
side of Morgan Mountain, eastern Oregon.
Occurrence: Worldwide.
Plate TRL01. Trillus elkhornensis Pessagno & Blome. Magnification x250. Fig. 1(H). Pessagno & Blome 1980,
pl. 6, fig. 12. Fig. 2. GSC loc. C-080611, GSC 111775. Fig. 3. GSC loc. C-080611, GSC 111776. Fig. 4. NBC, GSC
loc. C-305208, GSC 111777. Fig. 5. Hori 1990, Fig. 8.30. Fig. 6. Goričan et al. 2003, pl. 1, fig. 2. Fig. 7. OM, BR706-R14-05.
Fig. 8. OM, BR1121-R09-08.
387
Trillus seidersi Pessagno & Blome 1980
Species code: TRL02
Synonymy:
1980 Trillus seidersi n. sp. – Pessagno & Blome, p. 249, pl. 9,
figs. 2-4, 9, 19.
1987b Trillus sp. cf. T. seidersi – Yeh, p. 37, pl. 7, figs, 10, 11, 16, 25.
1988 Trillus sp. cf. T. seidersi Pessagno & Blome – Carter et al.,
p. 38, pl. 16, fig. 1.
1989 Trillus sp. aff. T. elkhornensis – Hattori, pl. 8, fig. H.
1996 Trillus sp. A – Yeh & Cheng, p. 98, pl. 2, figs. 1, 2.
1997 Trillus sp. D – Yao, pl. 3, fig. 146.
Original description: Cortical shell small, subspherical
with about equal number of pentagonal and hexagonal
pore frames which lack nodes at vertices. Surface of test
sloping steeply from median band toward polar spines.
Bars of pore frames of medium thickness along Y and 2 to 3
times thicker along Z; tending to be thicker along Z in area
of median band. Five pore frames visible along AA’ and
BB’. Polar spines quite long, triradiate in axial section; one
spine somewhat longer than the other and usually showing
curvature of its tip. Both polar spines having 3 very broad
longitudinal grooves alternating with 3 relatively narrow
ridges. Ridges and grooves maintaining same width for
most of their length.
Original remarks: Trillus seidersi, n. sp., differs from Trillus
elkhornensis, n. sp., in having larger and fewer pore frames
and polar spines with much wider grooves. Both species
have long polar spines. It is conceivable that T. elkhornensis
gave rise to T. seidersi through a reduction in the number
of pore frames and an increase in the width of the grooves
on its polar spines.
Further remarks: Forms referred to Trillus cf. seidersi in
Carter et al. (1988) are herein considered T. seidersi s.s.
Measurements (µm):
Based on 9 specimens. System of measurement shown in
text-figure 5 of Pessagno & Blome (1980).
AA’
90
81
90
60
A’S’
130
122
140
105
AS
90
89
100
80
BB’
90
85
92
80
cc’
25
24
30
20
dd’
25
26
42
20
HT
Av.
Max.
Min.
Etymology: This species is named for Dr. Victor M. Seiders
in honor of his significant contribution to the study of California Coast Ranges.
Type locality: Sample OR 513, Snowshoe Formation, along
Izee-John Day Road, eastern Oregon.
Occurrence: Snowshoe Formation, Oregon; Rennell Junction member of the Fannin Formation and Graham Island
Formation, Queen Charlotte Islands; Tawi Sadh Member
of the Guwayza Formation, Oman; Liminangcong Chert,
Philippines; Japan.
Genus: Tripocyclia Haeckel 1881, emend. Kiessling 1999
Type species: Tripocyclia trigonum Rüst, subsequent designation by Campbell (1954)
Synonymy:
1881 Tripocyclia n. gen. – Haeckel, p. 458.
1999 Tripocyclia Haeckel 1881, emend. – Kiessling, p. 39.
Original description:
2b. Spiny Coccodiscida with marginal spines lying in equatorial plane and radiating from the margin of the lensshaped test (without chambered rays).
B. With three equidistant spines.
Emended description: By Kiessling (1999): Spongy lenticular disc, triangular in axial view. Three strong triradiate
primary spines, symmetrically arranged, but sometimes of
slightly different size. Spines originating from a pseudoaulophacid microsphere as thin cylindrical beams, triradiate
development only on the surface or slightly inside the test.
Spongy meshwork internally arranged in indistinct concentric layers. Externally the disc commonly shows very
small pores. Secondary spines rarely present, very small.
No tholi and no distinct nodes developed.
Remarks: By Kiessling (1999): The emendation of this
genus is based on material from the Southern Alps (Italy
and Switzerland), where common specimens of the
type species were found in Bathonian to Kimmeridgian
388
sediments (Kiessling 1995b). Well-preserved specimens
of Tripocyclia trigonum were also figured by De Wever
et al. (1986, pl. 16, figs. 17-19). The forms fit very well
to the original description, figure and measurements of
Tripocyclia trigonum provided by Rüst (1885), but are
definitely not synonymous with Tripocyclia trigonum as
described by Pessagno (1977a). T. trigonum also shows no
similarity with the species described under Tripocyclia by
Pessagno and Yang in Pessagno et al. (1989), Yang (1993),
and Hull (1997). All those species are Xiphostylidae, whereas
Tripocyclia has to be assigned to the Pseudoaulophacidae.
Thus the emendation of Tripocyclia by Pessagno & Yang
(1989) cannot be considered valid. Much of the taxonomic
confusion about Tripocyclia stems from the poor original
description of the genus (Haeckel 1881) and the type
species (Rüst 1885). Although the subsequent assignment
of T. trigonum as the type species of Tripocyclia by Campbell
(1954) may have altered the original concept of Haeckel
(1881) it has to be accepted according to the IRZN (1985).
Tripocyclia differs from Tripodictya Haeckel by lacking
concentric chambered rings internally. The differences to
Spongotripus Haeckel are less clear. The emended definition
of Tripocyclia actually fits better to the original definition
of Spongotripus than to that of Tripocyclia. However, the
subsequent designation of Spongotripus regularis Haeckel
Plate TRL02. Trillus seidersi Pessagno & Blome. Magnification x250. Fig. 1(H). Pessagno & Blome 1980, pl. 9, fig. 2.
Fig. 2. Carter et al. 1988, pl. 16, fig. 1. Fig 3. QCI, GSC loc. C-304 566, GSC 111778. Fig. 4. OM, BR871-R07-07.
Fig. 5. OM, BR706-R06-26. Fig. 6. OM, BR1121-R09-22.
389
as the type species of Spongotripus (Campbell 1954) is
not considered valid, since this species was not figured by
Haeckel (1887) and hence represents a nomen dubium.
A possibility to define a valid type species could be
Spongechinus neumayri Dunikowski which was originally
included into the type-series by Haeckel (1887). However,
this species cannot serve as type species, owing to its
spherical test. Spongotripus pauper Rüst is amongst the
first species of this genus that was sufficiently described
and figured and fits to the original definition of the genus
and should be assigned as type species. This species differs
externally from Tripocyclia by its test being flat rather than
lenticular and its much shorter spines. More work needs to
be done on the status of both Tripocyclia and Spongotripus
to clarify the taxonomic problems.
Further remarks: We have strong doubts that the microsphere of this genus is of pseudoaulophacid structure as
mentioned in the emended description. Sections to microsphere made by one of us (PD) in several species assignable to Tripocyclia Haeckel as emended by Kiessling,
species coming from the very well preserved fauna of the
Bajocian sample IN7 of Yao (1997), show a central microsphere similar to that of Paronaella rather than to the
Pseudoaulophacidae (see Dumitrica & Zügel (2002) and
Dumitrica (1997) for the morphology of the two types of
microsphere).
Regarding the type species of the genus Spongotripus
Haeckel, we must use the species designated by Campbell (1954) and not Spongotripus pauper Rüst 1888 as redesignated by Kiessling (1999) because the redesignation
of a type species is against the ICZN, art. 61. The reason
Kiessling designated another type species for Spongotripus
is because he considered that the designation of S. regularis
by Campbell is not valid since this species was not illustrated by Haeckel. However, the ICZN art. 69A only specifies that “In designating a type-species for a genus, a zoologist should give preference to a species that is adequately
figured”. There is no mention that the zoologist must designate an illustrated species. In fact neither Spongotripus
regularis as type species nor S. pauper solve the taxonomical problems of Mesozoic species with spongy disk-shaped
test and three three-bladed equatorial spines. Spongotripus
comprises probably only Cenozoic Spongodiscidae because
its description was based on Cenozoic material and because
its type species has conical rather than three-bladed spines,
whereas Jurassic and Cretaceous species have bladed spines
and may belong to differing genera of several families such
as: Angulobracchiidae, Pseudoaulophacidae, Tritrabidae,
etc. Moreover, Spongotripus pauper as illustrated by Rüst
(1888) does not seem to be an equatorial section because
it shows no regular structure in the center and the three
spines do not reach the central part, thus giving no information on the family group to which it may belong. In
summary, Mesozoic species answering the original generic
diagnosis of Spongotripus should always be questionably
assigned to this genus.
Included species:
SPT01 Tripocyclia? tortuosa Dumitrica, Goričan &
Whalen n. sp.
Tripocyclia? tortuosa Dumitrica, Goričan & Whalen n. sp.
Species code: SPT01
Synonymy:
1990 Triactoma (?) sp. – Nagai, pl. 5, fig. 1.
1997 Spongotripus sp. B0 – Yao, pl. 5, fig. 235.
2003 Spongotripus sp. B0 sensu Yao – Goričan et al., p. 296,
pl. 2, fig. 6.
Type designation: Holotype specimen BR523-R02-04
(pl. SPT01, fig. 1), paratypes specimens BR524-R04-10 and
BR 825-3-R09-06 (pl. SPT01, figs. 2, 4) from the Guwayza
Formation, Tawi Sadh Member, Jabal Safra, Oman.
Description: Cortical shell subspherical with three triradiate spines at approximately 120º to each other.
Meshwork of cortical shell delicate, composed of thin bars
forming a dense spongy pattern arranged in concentric
layers. Superficial layer of mature specimens with small
but distinct nodes. Spines long, dextrally torsioned, with
pointed tip; degree of torsion and length of torsioned
portion variable.
Remarks: Tripocyclia? tortuosa n. sp. differs from other related species by having torsioned spines. From Spongotripus incomptus Carter 1988 it further differs by having much
longer spines and a less dense and triangular test. In the
structure of the nodose shell surface, Tripocyclia? tortuosa
390
n. sp. is the most similar to Alievium? juskatlaensis Carter
1988 but the latter species has straight spines. Our specimens are not well enough preserved to make equatorial sections in order to know the structure of the microsphere, but
specimens with a superficially similar test and very slightly
torsioned spines from the Bajocian of Japan (sample IN 7
of Yao, 1997) show in their center a microsphere similar to
that of Paronaella.
Measurements (µm):
Based on 5 specimens.
Diameter of shell
Preserved length of spines
HT
83
95
Min.
83
-
Max.
113
100
Etymology: From Latin: tortuosus, -a, -um = tortuous,
referring to torsioned spines.
Type locality: Sample BR523, Guwayza Formation, Tawi
Sadh Member, Jabal Safra, Oman.
Occurrence: Tawi Sadh Member of the Guwayza Formation,
Oman; Skrile Formation, Slovenia; Japan; San Hipólito
Formation, Baja California Sur; Hyde Formation, Oregon.
Plate SPT01. Tripocyclia? tortuosa Dumitrica, Goričan & Whalen n. sp. Magnification x250. Fig. 1(H). OM, BR523R02-04. Fig. 2. OM, BR524-R04-10. Fig. 3. Goričan et al. 2003, pl. 2, fig. 6. Fig. 4. OM, BR825-3-R09-06.
Fig. 5. BCS, SH-412-14.
391
Genus: Turanta Pessagno & Blome 1982, emend. Takemura 1986
Type species: Turanta capsensis Pessagno & Blome 1982
Synonymy:
be easily distinguished externally by the asymmetrical
placement of the horn and feet of Turanta as well as the
flattened area between the horn and vertical foot.
Original description: Test dicyrtid. Cephalic wall
incompletely formed, mostly open, partially lacking sides
and roof, but possessing well-developed cyrtoid collar plate
(pl. 3, figs. 2-3, 11-12); collar plate usually visible externally.
Prominently curved to slightly curved horn attached to
apical bar; base of horn covering collar plate on dorsal side
(in position of dorsal bar) (pl. 3, figs. 2, 11; 3, 12). Thorax
with two feet connected with and in line with vertical and
dorsal bars (pl. 4, fig. 17). Horns and feet varying in length
and curvature with species, typically triradiate in axial
section with alternating ridges and grooves. Horn and feet
usually in same plane; distance between horn and vertical
foot always differing from distance between horn and dorsal
foot. Thorax subspherical, often somewhat compressed at
right angles to the plane of the horn and feet; flattened area
occurring between horn and vertical foot; thorax lacking
mouth and with symmetrical pentagonal and hexagonal
pore frames.
Emended diagnosis: By Takemura (1986): Shell of one
segment, cephalis, large, subspherical and latticed, with
three usually triradiate spines, which are prolongations of
A, V, and D. Three spines located in a same plane, which
is perpendicular to the collar plate. Dorsal spine usually
curved slightly to the side of the apical spine. Cephalic
pores large and circular, regularly or irregularly distributed.
Cephalic skeletal elements, MB, V, D, two L and two l at the
base of the cephalis, and A inside the cephalis.
1982 Turanta n. gen. – Pessagno & Blome, p. 296.
1986 Turanta Pessagno & Blome emend. – Takemura, p. 64.
Original remarks: Turanta, n.gen., differs from all other
Mesozoic dicyrtid Nassellariina by virtue of the partially
formed, naked nature of its cephalis. It may well be that
the missing cephalic walls were extremely thin and fragile
in character and were not capable of being fossilized; they
are for the most part missing on all specimens of Turanta
observed during this study.
The spumellarian-like test of Turanta at first suggests a
kinship to Tripocyclia Rüst. However, the two genera can
Further remarks: By Takemura (1986): Pessagno & Blome
(1982) described Turanta as dicyrtid Nassellaria, of which
cephalis was naked. However, Turanta possesses all the
cephalic skeletal elements and specially A, originated at the
point where MB, D and two l join, prolonging into an apical
spine (pl. 11, 17-18), penetrating inside the shell which is
described as “thorax” by Pessagno & Blome (1982). This fact
clearly indicates that the large subspherical latticed shell of
Turanta is the cephalis. Therefore, the horn described by
Pessagno & Blome (1982) is the dorsal spine and the two
feet are the apical and vertical spines.
Etymology: Turanta (f.) is a name formed by an arbitrary
combination of letters (ICZN, 1964, Appendix D, pt. VI,
Recommendation 40, p. 113).
Included species:
3247 Turanta morinae gr. Pessagno & Blome 1982
Turanta morinae gr. Pessagno & Blome 1982
Species code: 3247
Synonymy:
1982 Turanta morinae n. sp. – Pessagno & Blome, p. 300, pl. 1,
figs. 3-4, 8, 11, 16.
1982 Turanta officerensis n. sp. – Pessagno & Blome, p. 301,
pl. 2, figs. 2-3, pl. 8, fig. 1.
1988 Turanta morinae Pessagno & Blome – Carter et al., p. 62,
pl. 14, fig. 8.
1991 Turanta sp. A, n. sp. – Carter & Jakobs, p. 344, pl. 3, fig. 13.
1995a Turanta morinae gr. Pessagno & Blome – Baumgartner et
al., p. 616, pl. 3247, figs. 1-3.
1996 Turanta sp. A – Yeh & Cheng, p. 122, pl. 8, fig. 10.
1997 Turanta morinae gr. Pessagno & Blome – Yao, pl. 8,
fig. 356.
Original description: Cephalis as with genus. Thorax subspherical with equal number of pentagonal and hexagonal
pore frames having weakly developed nodes at vertices;
hexagonal pore frames somewhat larger than pentagonal
pore frames. Horn and feet relatively long, triradiate in axial section with grooves and ridges of equal width; grooves
392
and ridges gradually decreasing in width distally. Proximal
1/2 of horn curved; distal 1/2 straight. Feet straight, widely
separated, nearly at right angles to horn.
Original remarks: Turanta morinae, n. sp., differs from
Turanta silviensis n. sp. by having a longer horn, longer feet
and fewer pore frames. Furthermore, whereas T. morinae
tends to have about the same number of hexagonal and
pentagonal pore frames, T. silviensis has predominantly
hexagonal pore frames.
Measurements (µm):
Based on 8 specimens.
Length of thorax
Width of thorax
Length of foot
Length of horn
HT
162.5
137.5
150+
150
Mean
154.68
146.25
164.58
122.5
Min.
125
125
100
87.5
Max.
175
175
225
150
Etymology: This species is named for Karen E. Morin in
honor of her recent contributions to the study of Upper
Cretaceous Radiolaria.
Occurrence: Snowshoe Formation, Oregon; Phantom
Creek Formation, Queen Charlotte Islands; Sogno Formation, Italy; Liminangcong Chert, Philippines; Japan.
Type locality: Sample OR-580, Warm Springs member,
Snowshoe Formation, near bridge over South Fork of John
Day River, east-central Oregon.
Plate 3247. Turanta morinae Pessagno & Blome. Magnification x200. Fig. 1(H). Pessagno & Blome 1982, pl. 1, fig. 3.
Fig. 2 Carter et al. 1988, pl. 14, fig. 8.
393
Genus: Tympaneides Carter 1988
Type species: Tympaneides charlottensis Carter 1988
Synonymy:
1988 Tympaneides Carter n. gen. – Carter et al., p. 37.
Original description: Test is a flattened sphere (drumshaped) with four spines extending from sides to form a
cross in one plane. Top and bottom surfaces planiform,
sides vertical to slightly concave. Latticed cortical shell
composed of two layers of pore frames on planar surfaces
and a single layer on the sides. Nodes on outer layer interconnected by fragile bars to form triangular or tetragonal
pore frames.
Original remarks: Tympaneides n. gen. is assigned to the
Staurolonchidae Haeckel because of its shape, mode of shell
construction and spine structure. It differs from Emiluvia
Foreman in having a test that is circular and drum-shaped
rather than rectangular, and from Staurolonche Haeckel
in having a double-, rather than a single-layered cortical
shell.
Etymology: From the Greek tympanon = drum.
Included species:
3408 Tympaneides charlottensis Carter 1988
Tympaneides charlottensis Carter 1988
Species code: 3408
Synonymy:
1988 Tympaneides charlottensis Carter n. sp. – Carter et al.,
p. 37, pl. 9, figs. 4, 5.
1989 Tympaneides sp. cf. T. charlottensis Carter – Hori & Otsuka,
pl. 4, fig. 10.
1991 Tympaneides charlottensis Carter – Carter & Jakobs,
p. 344, pl. 2, fig. 2.
1991 Tympaneides charlottensis Carter – Tipper et al., pl. 9,
fig. 10.
1995a Tympaneides charlottensis Carter – Baumgartner et al.,
p. 618, pl. 3408, figs. 1,2.
1998 Tympaneides charlottensis Carter – Cordey, p. 95, pl. 20,
fig. 12.
with alternating ridges and grooves. Ridges rounded and
approximately twice as wide as grooves, which are narrow
and deep.
Original remarks: This species is very abundant in middle
to upper Toarcian samples.
Measurements (µm):
Based on 14 specimens.
Diameter of test
Length of longest spine
HT
129
162
Av.
118
170
Max.
150
238
Min.
80
123
Original diagnosis: Test circular, drum-shaped. Meshwork
on planar surfaces very fine, pore frames triangular, nodes
minute. Equatorial spines long, slender and triradiate.
Etymology: This species is named for Queen Charlotte (wife
of George III of England) after whom the Queen Charlotte
Islands were named.
Original description: Test circular (drum-shaped) with
four long spines extending from sides of test at 90° to
one another. Outer layer of cortical shell covered with
very small triangular pore frames composed of thin bars
with fine nodes at their vertices. Spines long (one to three
times test diameter), slender and of uniform width. Spines
Type locality: GSC locality C-080583, Phantom Creek
Formation, Yakoun River, Graham Island, Queen Charlotte
Islands, British Columbia.
394
Occurrence: Phantom Creek Formation, Queen Charlotte
Islands; Fernie Formation, Williston Lake; Bridge River
Complex, British Columbia; Japan.
Plate 3408. Tympaneides charlottensis Carter. Magnification Fig. 1 x150 (scale bar A), Figs. 2-9 x200 (scale bar B).
Fig. 1. Carter & Jakobs 1991, pl. 2, fig. 2. Fig. 2. Carter et al. 1988, pl. 9, fig. 5. Fig. 3(H). Carter et al. 1988, pl. 9, fig. 4.
Fig. 4. QCI, GSC loc. C-080612, GSC 128705. Fig. 5. QCI, GSC loc. C-304281, GSC 111779. Fig. 6. QCI, GSC loc. C304281, GSC 111780. Fig. 7. NBC, GSC loc. C-205813, GSC 111781. Fig. 8. QCI, GSC loc. C-080611, GSC 111782.
Fig. 9. Hori & Otsuka 1989, pl. 4, fig. 10.
395
Genus: Udalia Whalen & Carter 1998
Type species: Udalia dennisoni Whalen & Carter 1998
Synonymy:
1998 Udalia n. gen. – Whalen & Carter, 1998, p. 59.
Original description: Test square or rectangular in outline,
usually quite thick with four prominent spines in the same
plane, one at each corner. Upper and lower surfaces of test
planiform, sides vertical. Test composed of multiple layers of
fine meshwork; meshwork of cortical shell with numerous,
irregularly shaped (mostly triangular and tetragonal)
pore frames with small nodes at pore frame vertices. Pore
frames of inner layers much thinner and lack nodes. Inner
structure unknown as center of test hollow in all specimens
observed. Spines circular or triradiate, tapering distally.
Original remarks: Udalia n. gen. differs from Sophia n. gen.
by lacking an entactiniid-like inner structure, having a
thinner shell, more pore frames, and much smaller nodes
at pore frame vertices. We note that Udalia n. gen. has
ferresiid-like meshwork. It is possible this genus could be
assigned to the Ferresiidae Carter if that family were revised
to include forms with four spines in the cruciform position.
Comparison with Emiluvia Foreman is difficult because of
the absence of information on the inner structure of Udalia
n. gen.
Etymology: This genus named for the “Udal”, a mission
ship built in Sandspit, Queen Charlotte Islands in the early
1900s.
Included species:
UDA05 Udalia plana Whalen & Carter 1998
Udalia plana Whalen & Carter 1998
Species code: UDA05
Synonymy:
1998 Udalia plana n. sp. – Whalen & Carter, p. 60, pl. 5, figs. 7, 8,
12, 13; pl. 7, figs. 13, 14, 15, 18.
2002 Udalia plana Whalen & Carter – Suzuki et al., p. 178, fig. 7 J.
2002 Udalia plana Whalen & Carter – Whalen & Carter, p. 112,
pl. 7, figs. 9-10.
2002 Udalia plana Whalen & Carter – Tekin, p. 185, pl. 3, fig. 3.
Original description: Cortical shell diamond-shaped in
outline, very thick, with planiform upper and lower surfaces
and vertical sides. Meshwork composed of irregularly
shaped triangular and tetragonal pore frames with large,
rounded nodes at pore frame vertices; pore frame bars
thinner in Y direction than in Z direction (refer to Pl. 4,
fig. 11 for measurement system). Test with four long spines,
all equal in length and width and tapering distally; spines
triradiate in axial section with narrow, rounded longitudinal
ridges and grooves.
Original remarks: See remarks under Udalia parvacapsa
n. sp.
Remarks under Udalia parvacapsa n. sp.: Udalia parvacapsa n. sp. differs from U. plana n. sp. in having a smaller
test and much longer spines.
396
Measurements (µm):
(n) = number of specimens measured.
Maximum diameter
of cortical shell (7)
113
143
105
122
Maximum length
of primary spines (6)
131
131
79
93
HT
Max.
Min.
Mean
Etymology: Planus, a, um (latin; adj.) = even, level, flat.
Type locality: Sample QC-676, Sandilands Formation,
Kunga Island, Queen Charlotte Islands, British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands; Fernie Formation, NE British Columbia; San Hipólito Formation, Baja California Sur; Pucara Group, Peru;
Hocaköy Radiolarite, Turkey.
Plate UDA05. Udalia plana Whalen & Carter. Magnification x150. Fig. 1(H)a-b. Carter et al. 1998, pl. 5, figs. 8, 7.
Fig. 2. Whalen & Carter 2002, pl. 7, fig. 10. Fig. 3. NBC, GSC loc. C-305208, GSC 111783. Fig. 4. NBC, GSC loc. C305208, GSC 111784.
397
Genus: Unuma Ichikawa & Yao 1976
Type species: Unuma (Unuma) typicus Ichikawa & Yao 1976
Synonymy:
1976 Unuma n. gen. – Ichikawa & Yao, p. 111.
1976 Unuma sensu stricto – Ichikawa & Yao, p. 112.
1976 Spinunuma n. subgen. – Ichikawa & Yao, p. 112.
1986 Unuma Ichikawa & Yao – Takemura, p. 58.
Original description: Spindle-shaped multisegmented
form with inversely subconical basal appendage which
has pores much larger than those of the preceding main
segments. Junction of segments not externally expressed
as an indentation. Numerous small circular pores on
the surface, aligned in longitudinal and diagonal rows.
Numerous longitudinal plicae on the surface generally
running continuously through segments. Apical horn may
be minute or large; radial spines and basal spine may or
may not be present.
Original remarks: A spindle-shaped form with small
pores, the absence of an externally expressed stricture, the
presence of large pores on the inversely subconical basal
appendage and of longitudinal plicae are stable diagnostic
features of the genus Unuma.
The last segment is represented by the distal part of the
main spindle-shaped shell with small pores. The basal
portion with large pores may be considered as a lid-like
appendage of the last segment rather than as the last
segment itself.
Two subgenera, Unuma (Unuma) and Unuma (Spinunuma), are distinguished on the basis of the presence or
absence of a distinct apical horn, radial spines, and basal spine. The morphological difference between the type
species of these subgenera may appear to be significantly
great at first glance, but there exist some forms transitional
with respect to the degree of development of radial spines,
so that a separation at subgeneric level is applied here.
Etymology: From the locality of the type specimens of the
type species. Unuma (regarded as masculine).
Included species:
UNM01 Unuma unicus (Yeh) 1987b
Unuma unicus (Yeh) 1987b
Species code: UNM01
Synonymy:
Further remarks: This species is herein assigned to Unuma
Ichikawa & Yao, because the complete specimens (including
the holotype) have a perforate basal appendage. Unuma
unicus seems to be the oldest representative of the genus.
Original description: Test very small, spindle-shaped, with
four to five post-abdominal chambers. Cephalis domeshaped without horn. Test comprised of inner latticed layer
of tetragonal pore frames with circular to elliptical pores;
outer layer with long massive costae usually extending
continuously throughout test. One to three rows of pore
frames between two adjacent costae. Seven to eight costae
visible laterally. Final post-abdominal chamber terminating
in latticed, hemispherical cap.
Measurements (µm):
Ten specimens measured.
Original remarks: This form is questionably assigned to
Hsuum Pessagno because it consists of a latticed, hemispherical closing cap at final post-abdominal chamber and
also because it lacks a horn.
Type locality: OR-600M, Hyde Formation at Izee-Paulina
road, east-central Oregon.
1987b Hsuum (?) unicum n. sp. – Yeh, p. 66, pl. 17, figs. 15-16, 21.
1997 Unuma unicum (Yeh) – Yao, pl. 10, fig. 476.
2004 Hsuum (?) unicum Yeh – Matsuoka, fig. 190.
398
HT
Mean
Max.
Min.
Total length
160
153
160
147
Max. width
120
118
120
115
Etymology: Unicus-a-um (Latin, adj.) = unique.
Occurrence: Hyde Formation, Oregon; Japan.
Plate UNM01. Unuma unicus (Yeh). Magnification x300. Fig. 1(H). Yeh 1987b, pl. 17, fig. 21.
Fig. 2. Matsuoka 2004, fig. 190.
399
Genus: Williriedellum Dumitrica 1970
Type species: Williriedellum crystallinum Dumitrica 1970
Synonymy:
1970 Williriedellum n. gen. – Dumitrica, p. 69.
Original diagnosis: Cryptothoracic tricyrtids with large
inflated abdomen having a constricted aperture and a
complex sutural pore; cephalis free, poreless, with four
collar pores, with or without a short apical horn; thorax
porous, campanulate, small, without descending spines and
partly depressed into abdomen.
Original remarks: Williriedellum is morphologically rather
similar to Zhamoidellum n. gen. Cryptamphorella n.gen.
and Hemicryptocapsa Tan. It differs from the first two
ones particularly in having a constricted aperture, which
constitutes the first external distinctive character. From
the last one, with which it seems to be closely related by
their constricted aperture, it differs in lacking the three
descending thoracic spines and in the complex structure of
its sutural pore.
Etymology: This genus is dedicated to William R. Riedel
(Scripps Institution of Oceanography) as a homage for his
sustained and indefatigable work in the study of Radiolaria.
Neuter gender.
Included species:
TPS01 Williriedellum? ferum (Matsuoka) 1991
Williriedellum? ferum (Matsuoka) 1991
Species code: TPS01
Synonymy:
1991 Tricolocapsa (?) fera n. sp. – Matsuoka, p. 726,
Fig. 4. 1a-3b.
2004 Tricolocapsa(?) fera Matsuoka – Matsuoka, fig. 83.
Original description: Shell of three segments, pyriform.
Cephalis hemispherical, poreless with a small apical horn.
Thorax truncate conical or cylindrical with small pores.
Abdomen large, subspherical with a constricted aperture.
Collar stricture slightly recognizable externally. Lumber
stricture pronounced. Pores moderate in size, circular to
subcircular, densely spaced and set in polygonal (largely
hexagonal) pore frames. Abdomen possessing small
pointed spines situated at the pore frame vertices. Aperture
moderate in size, circular with a short protruding rim.
Original remarks: This species is questionably assigned
to Tricolocapsa because it possesses a small apical horn.
However, it is uncertain whether the presence of an apical
horn is an important criterion for classification at generic
level of three-segmented nassellaria. The horn seems to be
an extension of apical spine. Tricolocapsa (?) fera, n. sp., is
similar to T. tetragona Matsuoka and T. sp. cf. T. ruesti Tan
Sin Hok (in Yao, 1979) in general outline but differs from
both by having an apical horn and by densely spaced pores
on the abdomen.
400
Further remarks: In the present state of knowledge
there is no Mesozoic genus to which this species can be
confidently assigned. Since assignation to Tricolocapsa
cannot be sustained any longer we assign it provisionally
to Williriedellum because of the general shape, the number
of segments and the presence of an aperture. However,
the cephalis is divided into two chambers by an oblique
septum as in the Amphipyndacidae and particularly in the
Cretaceous genus Squinabollum Dumitrica. Squinabollum
is similar in the general shape of shell, it has an apical horn,
but lacks an aperture.
Measurements (µm):
Numbers of specimens measured are in parentheses.
Total height of shell
Max. width of shell
Diameter of aperture
HT
139
100
11
Max.
146
125
14
Min.
125
96
8
Mean
133
107
11
(6)
(6)
(5)
Etymology: This specific name is derived from the Latin
ferus-a-um (= wild).
Type locality: Sample MNA-10, Nanjo Massif, Mino Terrane, central Japan.
Occurrence: Mino Terrane, central Japan.
Plate TPS01. Williriedellum? ferum (Matsuoka). Magnification x400. Fig. 1(H). Matsuoka 1991, Fig. 4.1a.
Fig. 2. Matsuoka 1991, Fig. 4.2a.
401
Genus: Wrangellium Pessagno & Whalen 1982, emend. Yeh 1987a
Type species: Wrangellium thurstonense Pessagno & Whalen 1982
Synonymy:
1982 Wrangellium n. gen. – Pessagno & Whalen, p. 126.
1987a Wrangellium Pessagno & Whalen, emend. – Yeh, p. 67.
Original description: Test conical, multicyrtid, large, lobulate in outline with numerous closely spaced post-abdominal chambers separated by nodose circumferential ridges
with H-linked structure. Longitudinally aligned, paired
circular to elliptical primary pores situated in symmetrical,
polygonal (mostly tetragonal) pore frames sloping steeply
to either side of circumferential ridges; ridges continuous with platform-like septal partitions possessing large,
circular apertures. Post-abdominal chambers with medially situated constrictions in areas between ridges. Single
transverse row of large polygonal pore frames situated in
constrictions between ridges, often completely obscured
by veneer of micro-granular silica (pl. 3, fig. 10). Cephalis
and thorax imperforate, covered by veneer of microgranular silica. Cephalis lacking horn. Test terminating in a large
(approximately 1/3 length of test) flaring, tubular structure
with large irregular pores and longitudinal ridges (pl. 3, fig.
18); tubular structure lacking septal partitions.
Original remarks: Wrangellium n. gen., differs from
Canoptum Pessagno by having large primary pores on its
circumferential ridges which remain open during ontogeny and by having a single row of large symmetrical pore
frames in the constrictions between ridges. It is likely that
Wrangellium was derived from a Canoptum stock with
H-linked circumferential ridges.
Emended definition: By Yeh (1987a): As with that of
Pessagno & Whalen (1982, p. 126), but including forms with
three pores aligned perpendicular to each circumferential
ridge, and also including forms with spine on the cephalis.
Etymology: Wrangellium is named for the Mesozoic terrane
of Wrangellia (Jones, Silberling and Hillhouse, 1977).
Included species:
WNG03 Wrangellium oregonense Yeh 1987a
WNG01 Wrangellium thurstonense Pessagno & Whalen
1982
WNG04 Wrangellium sp. A sensu Pessagno & Whalen 1982
Wrangellium oregonense Yeh 1987a
Species code: WNG03
Synonymy:
1987a Wrangellium oregonense n. sp. – Yeh, p. 69, pl. 2, figs. 1-8,
14-15, 19-20.
1987b Wrangellium oregonense Yeh – Yeh, p. 58, pl. 15, figs. 5-7,
14, 18-19, 24; pl. 27, figs. 3, 22.
1987b Crubus sp. B – Yeh, p. 71, pl. 23, fig. 6.
1988 Wrangellium oregonense Yeh – Carter et al., p. 50, pl. 6,
figs. 8, 11.
Original description: Test conical, lobate, with as many
as eight post-abdominal chambers. Cephalis conical,
moderately large, with small delicate horn when wellpreserved. Thorax and subsequent chambers rapidly
increasing in width and gradually increasing in length as
added. Cephalis, thorax, and abdomen imperforate, covered
with thick layer of microgranular silica and separated from
each other by one row of small elliptical pores. Each postabominal chamber with central row of pore frames covered
with microgranular silica, flanking rows of pore frames
remaining open. Pores rapidly increasing in size distally.
About eight to ten pores visible laterally in each row.
402
Original remarks: Wrangellium oregonense, n. sp., differs
from W. izeense, n. sp., by having a smaller, more conical
cephalis with small spine, by having a test more conical
in shape, and by possessing larger, paired pores aligned
perpendicular to the ridges.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Max. length
258
274
288
258
Max. width
123
123
135
118
Etymology: This species is named for the state of Oregon.
Type locality: Sample OR-600A, Hyde Formation along
Izee-Paulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Snowshoe Formation, Oregon; Fannin and Whiteaves formations, Queen Charlotte Islands.
Plate WNG03. Wrangellium oregonense Yeh. Magnification x250. Fig. 1(H). Yeh 1987b, pl. 15, fig. 5. Fig. 2. Carter et al.
1988, pl. 6, fig. 8. Fig. 3. QCI, GSC loc. C-304566, GSC 111785. Fig. 4. QCI, GSC loc. C-080613, GSC 111786.
403
Wrangellium thurstonense Pessagno & Whalen 1982
Species code: WNG01
Synonymy:
Original remarks: Wrangellium thurstonense, n. sp. differs
from Wrangellium sp. A by having much less pronounced
circumferential ridges. In addition, Wrangellium sp. A
possesses larger pores within the constrictions between
ridges.
Original description: Test with as many as 13 post-abdominal chambers with rare spines attached to circumferential
ridges (many more spines probably broken off). Majority
of large primary pores circular, set in square pore frames
along circumferential ridges. When visible, pores within
constrictions large, circular. Cephalis and thorax conical,
imperforate, completely covered by microgranular silica.
Proximal post-abdominal chambers very gradually increasing in width with distal post-abdominal chambers remaining the same width. Flaring, tubular structure rapidly
increasing in width, about 50% wider than test at its termination. Irregularly sized pores randomly distributed over
tubular structure with two sharp ridges extending along its
length.
Measurements (µm):
Based on 7 specimens.
1982 Wrangellium thurstonense n. sp. – Pessagno & Whalen,
p. 126, pl. 2, figs. 7, 13; pl. 3, figs. 1, 3, 10, 18; pl. 12, fig. 13.
1998 Wrangellium thurstonense Pessagno & Whalen – Whalen
& Carter, p. 65, pl. 17, figs. 5, 6; pl. 26, fig. 7.
Length
325.0
382.0
245.0
303.1
Width (max.)
100.0
160.0
100.0
112.4
HT
Max.
Min.
Mean
Etymology: This species is named for Turston Harbor,
northwest of its type locality.
Type locality: Sample QC 590A, Sandilands Formation
(Kunga Formation in Pessagno & Whalen, 1982), Queen
Charlotte Islands, British Columbia.
Occurrence: Sandilands Formation, Queen Charlotte Islands.
Wrangellium sp. A sensu Pessagno & Whalen 1982
Species code: WNG04
Synonymy:
1982 Wrangellium sp. A – Pessagno & Whalen, p. 126, pl. 3, figs.
2, 8, 9.
Remarks: Differs from Wrangellium thurstonense Pessagno
& Whalen 1982 in having fewer postabdominal chambers
and a more lobate outline. It further differs from W. izeense
Yeh 1987a in having a conical rather than rounded cephalis
and two rows of pores rather than three per chamber.
Occurrence: Franciscan Complex, California; Sandilands
and Ghost Creek formations, Queen Charlotte Islands.
404
Plate WNG01. Wrangellium thurstonense Pessagno & Whalen. Magnification x250. Fig. 1(H). Pessagno & Whalen
1982, pl. 2, fig. 7. Fig. 2. Carter et al. 1998, pl. 17, fig. 5.
Plate WNG04. Wrangellium sp. A sensu Pessagno & Whalen. Magnification x250. Fig. 1. Pessagno & Whalen 1982,
pl. 3, fig. 2. Fig. 2. QCI, GSC loc. C-080612, GSC 111787.
405
Genus: Xiphostylus Haeckel 1881, emend. Pessagno & Yang 1989
Type species: Xiphostylus attenuatus Rüst 1885 (subsequent designation by Campbell, 1954)
Synonymy:
referred to herein as cortical buttresses (pl. 1, figs. 3-4).
Outer latticed layer of cortical shell usually not as thick
as that of Tripocyclia Hackel or Triactoma Rüst (cf. pl. 1,
figs. 2, 5-6, 8, 10-11, 13).
Emended description: By Pessagno & Yang in Pessagno
et al. (1989): Test with subspherical to ellipsoidal cortical
shell with opposed secondary spines. Secondary spines
subequal in length, predominantly triradiate in axial section with three longitudinal grooves alternating with three
longitudinal ridges. Shorter spine often more massive and
wider than longer spine. Spines attached to latticed cortical shell by means of latticed protrusions of cortical shell
Emended remarks: By Pessagno & Yang in Pessagno et al.
(1989): Xiphostylus Haeckel differs from Triactoma Rüst
by possessing two opposed secondary spines with cortical
buttresses, and a less spherical cortical shell.
1881 Xiphostylus n. gen. – Haeckel, p. 449.
1989 Xiphostylus Haeckel emend. Pessagno & Yang – Pessagno
et al., p. 232.
Included species:
XTL02 Xiphostylus duvalensis Carter n. sp.
XTL01 Xiphostylus simplus Yeh 1987b
Xiphostylus duvalensis Carter n. sp.
Species code: XTL02
Type designation: Holotype GSC 111788 from GSC loc. C080610; Fannin member of the Fannin Formation (upper
Pliensbachian).
Description: Cortical shell large, spherical in outline,
with two polar spines approximately equal in length. Shell
comprised of small pentagonal and hexagonal pore frames
with thin rims and deep walls; small pointed nodes at pore
frame vertices. Spines relatively short (less than diameter of
shell), and fairly constant in width. Spines triradiate with
wide rounded ridges and narrow grooves; ridges terminate
with small thorns, raised at right angles to axis of spine.
Spine tips circular in axial section.
Remarks: Xiphostylus duvalensis n. sp. differs from X. simplus Yeh in having spines whose ridges terminate in small
thorns.
Measurements (µm):
Based on 2 specimens.
Diameter of cortical shell
Length of longer polar spine
Length of shorter polar spine
HT
136
93
82
2nd spec.
107
97
83
Etymology: Species named for Duval Rocks, west of type
locality on north shore of Cumshewa Inlet, Queen Charlotte Islands.
Type locality: Sample CAA-85-SP-27, lms. 2 (GSC loc.
C-080610), Fannin member of the Fannin Formation, north
shore Cumshewa Inlet, Moresby Island, Queen Charlotte
Islands, British Columbia.
Occurrence: Fannin member of the Fannin Formation,
Queen Charlotte Islands; Tawi Sadh Member of the Guwayza Formation, Oman.
Xiphostylus simplus Yeh 1987b
Species code: XTL01
Synonymy:
1987b Xiphostylus simplus n. sp. – Yeh, p. 52, pl. 10, fig. 7; pl. 22,
fig. 4.
1987b Xiphostylus sp. A – Yeh, p. 53, pl. 3, fig. 15; pl. 10, fig. 10.
1987b Xiphostylus sp. B – Yeh, p. 53, pl. 26, figs. 7, 11.
1987 Xiphosphaera spp. – Hattori, pl. 22, figs. 9-14, not fig. 15.
1989 Xiphostylus sp. – Hattori & Sakamoto, pl. 1, fig. K.
1989 Xiphostylus spp. – Hattori, pl. 4, fig. B, C, D.
1990 Xiphostylus sp. – Nagai, pl. 5, fig. 5.
1997 Xiphostylus simplus Yeh – Yao, pl. 1, fig. 15.
1997 Xiphostylus sp. P2 – Yao, pl. 1, fig. 16.
2003 Xiphostylus spp. – Goričan et al., p. 291, pl. 1, fig. 1.
Original description: Test large, subspherical in outline,
with two polar spines unequal in length. One spine long
406
and massive, triradiate with three rounded ridges alternating with three wide grooves; ridges and grooves displaying
slight torsion. One polar spine extremely short and pointed in nature. Test comprised of regularly sized pentagonal
and hexagonal pore frames. Pore frames thin in rims and
thick in sides with numerous laminations superimposed
on each pore frame (plate 22, figure 4).
Original remarks: This form is characterized by having a
large spherical test with one extremely short polar spine
and one long, massive polar spine.
Further remarks: Included are specimens with simple polar
spines regardless of their length.
Plate XTL02. Xiphostylus duvalensis Carter n. sp. Magnification x250. Fig. 1(H). QCI, GSC loc. C-080610,
GSC 111788. Fig. 2. OM, BR473-R16-02.
Measurements (µm):
Ten specimens measured.
HT
Mean
Max.
Min.
Diameter
of test
110
111
115
108
Length of
longer polar spine
110
108
110
102
Etymology: Simplus-a-um (Latin, adj.) = simple.
Length of
shorter polar spine
25
27
30
23
Type locality: Sample OR-600M, Hyde Formation at IzeePaulina road, east-central Oregon.
Occurrence: Nicely and Hyde formations, and Warm
Springs member of the Sowshoe Formation, Oregon; Skrile
Formation, Slovenia; Tawi Sadh Member of the Guwayza
Formation, Oman; Japan.
Plate XTL01. Xiphostylus simplus Yeh. Magnification x200. Fig. 1(H). Yeh 1987b, pl. 10, fig. 7. Fig. 2. OM, BR706-R13-01.
Fig. 3. Goričan et al. 2003, pl. 1, fig. 1.
407
Genus: Zartus Pessagno & Blome 1980
Type species: Zartus jonesi Pessagno & Blome 1980
Synonymy:
1980 Zartus n. gen. – Pessagno & Blome, p. 249.
Original description: Cortical shell spherical to ellipsoidal
with well developed raised median band. Pore frames on
median band thicker in Z direction (text-fig. 5) than those
of remainder of test. Raised median band with short, broad,
often massive, triradiate secondary spines; secondary
spines centered on pore frame vertices with ridges of spines
extending onto 3 bars of adjacent pore frames. Test with
2 polar spines of different length; polar spines usually
triradiate but sometimes partially circular in axial section.
First medullary shell with thin, fragile pore frames.
Original remarks: The triradiate secondary spines of
Zartus n. gen. are centered on the pore frame vertices
along the center of the median band. Their ridges extend
distally onto the bars of 3 adjacent pore frames (pl. 7, figs.
6, 12). The pore frames of Zartus, which are normally quite
thick in the Z direction (text-fig. 5), are even thicker in
the Z direction along the median band. Such an increase
in thickness along the median band may offer stouter
support for the massive secondary spines. Zartus n. gen.
differs from Pantanellium Pessagno in possessing a welldeveloped, raised median band with triradiate secondary
spines. The phylogenetic relationship of Zartus to other
genera of the Pantanellinae is discussed elsewhere in this
report.
Etymology: Zartus is a name formed by an arbitrary combination of letters (ICZN, 1964, Appendix D, pt. IV, Recommendation 40, p. 113).
Included species:
ZRT01 Zartus mostleri Pessago & Blome 1980
ZRT03 Zartus stellatus Goričan & Matsuoka n. sp.
Zartus mostleri Pessagno & Blome 1980
Species code: ZRT01
Synonymy:
1980 Zartus mostleri n. sp. – Pessagno & Blome, p. 252, pl. 6,
figs. 3-5, 13.
1989 Zartus sp. A – Hattori, pl. 9, fig. G.
1989 Zartus sp. B – Hattori, pl. 9, fig. H.
1989 Zartus sp. C – Hattori, pl. 9, fig. I.
1989 Zartus spp. – Hattori, pl. 9, fig. K.
1997 Zartus sp. B0 – Yao, pl. 4, fig. 153.
1997 Zartus dicksoni Pessagno & Blome – Yao, pl. 4, fig. 154.
2003 Zartus aff. mostleri Pessagno & Blome – Goričan et al.,
p. 291, pl. 1, figs. 4-6.
Original description: Cortical shell ellipsoidal with large,
hexagonal to pentagonal pore frames with spinose nodes
at vertices. Bars of pore frames thin in Y direction; thick
in Z direction (text-fig. 5). Four to 5 pore frames visible
along AA’; 5 along BB’. Median band weakly developed,
having 4 wide, long (16 x 25µm on holotype) relatively
massive triradiate secondary spines. Polar spines triradiate
in axial section; shorter spine with 3 massive, wide ridges
alternating with 3 moderately narrow grooves; grooves
about equal in width to ridges. Shorter polar spine about
2/3 length of longer polar spine. Longer polar spine with
3 wide grooves alternating with 3 relatively narrow ridges.
First medullary shell with fragile hexagonal and pentagonal
pore frames.
Original remarks: Zartus mostleri n. sp., differs from
Z. jurassicus, n. sp., in having much longer polar spines and
a much narrower median band. It appears to have been the
408
first species of Zartus to appear and may have arisen from
Pantanellium danaense, n. sp., ancestor.
Further remarks: In this species we also include forms
with many short triradiate secondary spines, extending
from most of the pore-frame vertices on the median band
(pl. ZRT01, figs. 3, 4). These forms differ from Zartus
mostleri s. s. by having more numerous (more than four)
and shorter secondary spines.
Measurements (µm):
Based on 6 specimens. System of measurement shown in
text-figure 5 of Pessagno & Blome (1980).
AA’
95
98
110
90
A’S’
125
117
125
100
AS
115
83
115
65
BB’
100
101
115
90
cc’
35
30
35
25
dd’
25
28
40
25
HT
Av.
Max.
Min.
Etymology: This species is named for Dr. Helfried Mostler
in honor of his pioneering contribution to the study of
Triassic Radiolaria.
Type locality: Sample QC 534, Fannin Formation (Maude
Formation in Pessagno Blome, 1980), Queen Charlotte
Islands.
Occurrence: Ghost Creek and Fannin formations, Queen
Charlotte Islands; Skrile Formation, Slovenia; Japan.
Plate ZRT01. Zartus mostleri Pessagno & Blome. Magnification x250. Fig. 1(H). Pessagno & Blome 1980, pl. 6, fig. 3.
Fig. 2. QCI, GSC loc. C-080612, GSC 111789. Fig. 3. Goričan et al. 2003, pl. 1, fig. 4. Fig. 4. SI, MM 6.76, 000407.
Fig. 5. Goričan et al. 2003, pl. 1, fig. 6.
409
Zartus stellatus Goričan & Matsuoka n. sp.
Species code: ZRT03
Synonymy:
? 1989 Zartus spp. – Hattori, pl. 9, fig. L.
1997 Zartus sp. A0 – Yao, pl. 4, fig. 152.
2003 Zartus sp. A0 sensu Yao – Goričan et al., p. 291, pl. 1, fig. 3.
2004 Zartus (?) sp. – Matsuoka, fig. 17.
Type designation: Holotype specimen MA 11322 from
sample MNA-10, Nanjo Massif, Mino Terrane, Japan.
Description: Cortical shell subspherical with relatively
small hexagonal and pentagonal pore frames. Five to six
pore frames visible along AA’. Median band bears several
(more than four, usually seven) long massive triradiate
secondary spines, pyramidal in shape. Polar spines long
and robust, approximately equal in length; width constant
through most of the length, then decreasing more rapidly towards spine tips. Polar spines triradiate with three
rounded ridges alternating with three deep grooves.
Remarks: Zartus stellatus n. sp. differs from all other Zartus
species by having larger polar spines and numerous long
secondary spines on median band.
410
Measurements (µm):
Based on 5 specimens.
HT
Min. Max.
Width of shell along BB’
76
75
90
Length of polar spines
128, 145 72
145
30
40
Width of polar spines at base 40, 34
Maximum length
46
26
46
of secondary spines
Mean
80
108
35
36
Etymology: Stellatus-a-um (Latin, adj.) = stellate, arranged
like a star, radiating.
Type locality: Sample MNA-10, Nanjo Massif, Mino
Terrane, Japan.
Occurrence: Mino Terrane, Japan; Skrile Formation,
Slovenia.
Plate ZRT03. Zartus stellatus Goričan & Matsuoka n. sp. Magnification x250. Fig. 1(H). Matsuoka 2004, fig. 17.
Fig. 2. JP, MNA-10, MA11505. Fig. 3. Goričan et al. 2003, pl. 1, fig. 3. Fig. 4. SI, MM 6.76, 000525. Fig. 5. SI, MM 6.76,
000406.
411
Genus: Zhamoidellum Dumitrica 1970
Type species: Zhamoidellum ventricosum Dumitrica 1970
Synonymy:
1970 Zhamoidellum n. gen. – Dumitrica, p. 79.
1992 Complexapora n. gen. – Kiessling & Zeiss, p. 190.
Original description: Cryptothoracic tricyrtids with large
inflated abdomen without aperture or sutural pore. Cephalis poreless, with four collar pores, with or without a short
apical horn; thorax campanulate, porous, partly depressed
into the abdominal cavity, its opening without descending
spines.
Original remarks: This new genus is very similar to
Cryptamphorella n. gen. from which it differs, firstly, by the
porous structure of its thorax and, secondly, by having no
sutural pore. In fact, Cryptamphorella sometimes does not
possess a sutural pore. The members of this genus are very
frequent in the Upper Jurassic. We described herein only
two better preserved species.
Further remarks: It is possible that this genus is a junior
synonym of Trisyringium Vinassa, 1901 (type species
Trisyringium capellinii Vinassa) from which it only differs
in lacking spines on the abdominal segment. Complexapora
Kiessling & Zeiss 1992 (type species Complexapora tirolica
Kiessling) should be considered a junior synomym of
Zhamoidellum. The latter genus was originally considered
to differ from Zhamoidellum only in having a sutural pore;
however, this pore is only a depression of the abdominal
wall. Such a depression is present but very weakly developed
in the type species.
Etymology: The genus is dedicated to Dr. A. I. Zhamoida as a
homage to his activity for disentangling the biostratigraphy
of the Mesozoic radiolaritic series. Neuter gender.
Included species:
COM01 Zhamoidellum yehae Dumitrica n. sp.
Zhamoidellum yehae Dumitrica n. sp.
Species code: COM01
Synonymy:
1988 Dicolocapsa aff. verbeeki Tan – Li, pl. 1, fig. 23.
1998 Complexapora sp. A – Yeh & Cheng, p. 33, pl. 4, fig. 13;
pl. 9, fig. 18, 22.
2005 Tricolocapsa sp. – Kashiwagi et al., pl. 6, fig. 18.
Remarks: Zhamoidellum yehae n. sp. differs from Z. ventricosum Dumitrica in being smaller, in having a spherical
abdomen and a small sutural depression.
Type designation: Holotype specimen BR1121-R06-22
from sample BR 1121, Guwayza Formation, Tawi Sadh
Member, Wadi Mu’aydin, Oman.
Measurements (µm): Based on 6 specimens. Diameter
of abdomen 123-132 (holotype 132). The two specimens
illustrated by Yeh & Cheng (1998) are larger; the diameter
of their abdomen is 157 and 182 respectively.
Diagnosis: A small species of Zhamoidellum with cephalothorax half or more depressed in the cavity of a spherical
abdomen.
Etymology: The species is named for Kuei-Yu Yeh, Taiwan,
honoring her contribution to the knowledge of Mesozoic
Radiolaria.
Description: Cephalothorax with apically rounded, imperforate cephalis and with thorax depressed in the abdominal
cavity. Abdomen spherical with numerous small pores of
various size and irregular arrangement. Pore frames circular or polygonal, usually hexagonal, with or without tiny
thorns at vertices. Sutural pore represented by a small, irregular depression in the vicinity of cephalothorax.
Type locality: Sample BR1121, Tawi Sadh Member, Guwayza Formation, Wadi Mu’aydin, Oman.
412
Occurrence: Tawi Sadh Member of the Guwayza Formation
and Haliw (Aqil) Formation, Oman; Liminangcong Chert,
Philippines; Dengqen area, Tibet; Japan; Ghost Creek
Formation, Queen Charlotte Islands.
Plate COM01. Zhamoidellum yehae Dumitrica n. sp. Magnification x300. Fig. 1(H)a-b. OM, BR1121-R06-22a, b.
Fig. 2. OM, Haliw-038-R08-20. Fig. 3. JP, Ku(b)-11-77. Fig. 4. QCI, GSC loc. C-304281, GSC 128751.
Fig. 5. OM, BR1121-R05-05b. Fig. 6. JP, MNA-10, MA12277.
413
414
3. DESCRIPTION OF
LOCALITIES
Locality data are provided only for specimens that are
illustrated in the catalogue. The exact location, a short
description of lithology and the overall stratigraphic range
of the studied successions are given. Stratigraphically
important co-occurring fossils are also indicated. The
number preceding each area corresponds to the locality
number on the world map (Fig. 1.2, p. 11). Stratigraphic
ranges of all studied formations are summarized in Fig.
3.1, p. 416.
1. NORTHEASTERN BRITISH COLUMBIA
Williston Lake
NTS 94B/3 Mt. Brewster; Peach Reach, UTM 497385N,
6215650W. Fernie Formation at Black Bear Ridge.
GSC loc. C-305208. Parts of a calcareous concretion collected in thinly laminated black, siliceous shales and
siltstones from the lower part of the formation at Pardonet Creek, by R. Hall (University of Alberta, Calgary). Faunas present include a “bedding plane exposure of crinoids, which I assume to be Seirocrinus ….”
and a few “scrappy ammonite impressions” (R.H. Hall,
pers. comm. 2000). Radiolarians are probably equivalent to faunas from the Whiteavesi Ammonite Zone in
Queen Charlotte Islands (ESC).
NTS 94B/3 Mt. Brewster. UTM 497670N, 6215500W. 4 km
northeast of Nabesche River. Fernie Formation. Sample
collected 3.5 m west (?upsection) from crinoid bed at
Black Bear Ridge section.
GSC loc. C-305813 (01-OF-BBR 6A), Pliensbachian.
Sample collected by R. Hall.
2. QUEEN CHARLOTTE ISLANDS
Sinemurian
Graham Island, Yakoun River area
NTS 103 F/8 Graham Island. UTM 681500m E; 5921800m
N (53° 24’ 57.3” N; 132° 16’ 8.3” W). Sandilands Formation, Kunga Group. Sample collected by B.E.B. Cameron
in transit along west side of Yakoun River, south of junction with Ghost Creek, central Graham Island. Eroded riverside sequence of sandstone, siltstone and shale with rare
limestone concretions. Ammonites collected in association
with limestone sample are upper Sinemurian and belong
to the Harbledownense Assemblage of Pálfy (1991) (H.W.
Tipper, personal communication, 1994).
415
416
Fig. 3.1. Stratigraphy of included sites.
CAA- 86-T-2/3 GSC loc. C-140441. Carbonate concretion
collected 76 m above base. Upper Sinemurian. Sample
collected by B.E.B. Cameron.
concretions and lenses in siltstone and fine sandstone.
GSC loc. C-140418). Limy concretionary lens. ?Kunae/
Carlottense Zones, upper Pliensbachian.
Pliensbachian
Maude Island, below ammonite type section
NTS 103 F/1 Skidegate Channel. Lat. 53° 11’ 06”N, Long.
132° 04’02”W. Sandilands Formation, Kunga Group.
Shoreline exposure ~ 0.5 km west of type section. Sequence composed mainly of siltstone and shale with minor
sandstone and limy lenses.
GSC loc. C-304428. Sample from discontinuous limestone bed 6 cm thick. ?Tetraspidoceras Ammonite Assemblage; uppermost Sinemurian/basal Pliensbachian
(J. Pálfy, pers. comm. 2001).
Pliensbachian locality data from Queen Charlotte Islands
(excluding Section 4 from Carter et al., 1988) are from an
unpublished report to the Geological Survey of Canada by
E. S. Carter (March 2002). Nearly all radiolarian samples
co-occur with ammonites identified by H.W. Tipper (Geological Survey of Canada, deceased 2005) who also provided the ammonite zonal assignment.
Kunga Island
NTS 103 b/13 & 103 B/14 Louise Island. Lat. 52° 45’ 53”N,
Long. 131° 33’ 08”W. Sandilands Formation, Kunga Group.
Stratigraphic sequence along the southeast shore of Kunga
Island. Note: the Sandilands Formation extends upward
to the lowermost Pliensbachian at this locality whereas in
nearly all others it extends only to the upper Sinemurian.
Sequence composed mainly of interbedded siltstone and
shale with minor sandstone, limy lenses and concretions.
GSC loc. C-305417. Limestone concretion collected
10.94 m above base of section. Tetraspidoceras
Ammonite Assemblage, basal Pliensbachian.
Central Graham Island
NTS 103 F/8, Yakoun Lake. Lat. 53° 22’ 30”N, Long. 132°
16’ 00”W. Rennell Junction member of the Fannin Formation, Maude Group. Quarry above Rennell Junction,
Rd. 19. Section consists of thick sandstone beds alternating with sequences of limy concretions and lenses in siltstone and fine sandstone.
GSC loc. C-177371. Limestone sample; base of Whiteavesi
Zone, upper lower Pliensbachian.
Cumshewa Inlet, Moresby Island
NTS 103 G/4 Cumshewa Inlet. Lat. 53° 02’ 53”N, Long.
131° 56’ 05”W. Basal two-thirds of section composed of
shale with very minor limestone concretions (Ghost Creek
Formation); upper part composed of limy concretions
and lenses in siltstone and minor shale (Rennell Junction
member of Fannin Formation); uppermost beds more
sandy with fewer concretionary lenses.
GSC loc. C-080610. Fannin Formation; spot sample
collected in upper part of section by B.E.B. Cameron
in 1986. ?Kunae Zone; upper Pliensbachian.
GSC loc. C-140413. Rennell Junction member of the Fannin formation. Collected 54 m below top of Fannin
Formation. Whiteavesi Zone, upper lower Pliensbachian.
Louise Island
NTS 103 G/4 Cumshewa Inlet. Lat. 53° 02’ 06”N, Long.
131° 52’ 47”W. Fannin Formation. North shore of Louise
Island approximately 2 km northeast along shore from
Beattie Anchorage. Upper Pliensbachian stratigraphic
sequence consists of sandstone alternating with limy
Maude Island, Tipper Creek
NTS 103 F/1 Skidegate Channel. Lat. 53° 11.82’N, Long.
132° 3.63’W. Fannin Formation, Maude Group. Tipper
Creek flows onto beach approximately 100 m west of Fannin Bay (southwest coast of Maude Island) about midway
through the type section of the Ghost Creek and Fannin
formations (Cameron & Tipper, 1985). Stratigraphic sequence composed of interbedded medium grey sandstone,
dark grey siltstone with rare shale and sandy limestone.
GSC loc. C-080577. Limestone sample collected 54.3 m
stratigraphically below top of Fannin Formation. Carlottense Zone, upper Pliensbachian (see section 4, Bulletin 386, Carter et al., 1988).
Maude Island, west of Ells Bay
NTS 103 F/1 Skidegate Channel. Lat. 53° 11’ 57”N, Long.
132° 2’ 56”W. West side of Ells Bay. Type section of the
Maude Formation of Sutherland Brown (1968). Total section extends over 112 m stratigraphically. The lower Ghost
Creek Formation (46 m) is thick dark grey shale with
rare limestone concretions and lenses; the overlying Rennell Junction member of the Fannin Formation (~37m)
is comprised mainly of limy concretions and lenses interbedded in dark grey siltstone, minor shale and sandstone.
Note: the upper beds of the Fannin Formation (~30 m) are
much more sandy and contain only rare limy concretionary beds. The upper part of the Rennell Junction member
yields the best preserved and most abundant radiolarian
faunas in the entire sequence.
GSC loc. C-305386. Ghost Creek Formation, collected
1.0 m above base of section. Imlayi Ammonite Zone,
lower Pliensbachian.
GSC loc. C-305388. Ghost Creek Formation, collected
3.0 m above base of section. Imlayi Ammonite Zone,
lower Pliensbachian.
GSC loc. C-175310. Rennell Junction member of the
Fannin Formation, collected 62 m above base of
section. Whiteavesi Ammonite Zone, upper lower
Pliensbachian.
GSC loc. C-304565. Rennell Junction member of the
Fannin Formation, collected 68.9 m above base of
section. Whiteavesi Ammonite Zone, upper lower
Pliensbachian.
GSC loc. C-304566. Rennell Junction member of the
417
Fannin Formation, collected 70.7 m above base of
section. Whiteavesi Ammonite Zone, upper lower
Pliensbachian.
GSC loc. C-304567. Rennell Junction member of the
Fannin Formation, collected 71.8 m above base of
section. Whiteavesi Ammonite Zone, upper lower
Pliensbachian.
GSC loc. C-304568. Rennell Junction member of the Fannin Formation, collected 75.4 m above base of section.
Whiteavesi/Freboldi Ammonite Zones, upper lower
Pliensbachian.
GSC loc. C-175306. Fannin Formation, collected 85 m
above base of section. Basal Kunae Ammonite Zone;
upper Pliensbachian.
Graham Island, Ghost Creek
NTS 103 F/8, Yakoun Lake. Lat. 53° 25’ 46”N, Long. 132°
17’ 16”W. Fannin Formation; Quarry on road north of
Ghost Creek.
GSC loc. C-175309. Limy concretion in sandy beds of
the Fannin Formation collected for radiolarians in
1990, by H.W. Tipper. Kunae Ammonite Zone, upper
Pliensbachian.
Rennell Junction, Graham Island
NTS 103 F/8, Yakoun Lake. Lat. 53° 24’ 26”N, Long. 132°
18’ 13”W. High waterfall section east of ‘Queen Charlotte
Main’, 0.25 km north of junction of ‘Queen Charlotte
Main’ with road to Rennell Sound. Lower half of sequence
mainly shale with rare limy concretions and lenses (Ghost
Creek Formation); upper part of the sequence comprised
of interbedded siltstone, sandstone, limestone and shale
with rare concretionary lenses. This locality was collected
in 1979-82 soon after the area was logged. It is now totally
overgrown with a tall, dense forest and is, for all but the
most doggedly-determined, totally inaccessible.
GSC loc. C-175311. Ghost Creek Formation. Collected
0.9 m above base of section. Imlayi Ammonite Zone,
basal Pliensbachian.
GSC loc. C-080612. Ghost Creek Formation. Collected
2.8 m above base of section. Imlayi Ammonite Zone,
lowermost Pliensbachian.
GSC loc. C-127868. Ghost Creek Formation. Collected
~6 m above base of section. Imlayi Ammonite Zone,
lower Pliensbachian.
GSC loc. C-127867. Ghost Creek Formation. Collected
~6 m above base of section. Imlayi Ammonite Zone,
lower Pliensbachian.
GSC loc. C-080611. Ghost Creek Formation. Collected
58.1 m above base of section. Whiteavesi Ammonite
Zone, lower Pliensbachian.
GSC loc. C-080613. Rennell Junction member of the
Fannin Formation. Collected 94.2 m above base
of section. Freboldi Ammonite Zone, upper lower
Pliensbachian.
GSC loc. C-140495. Rennell Junction member of the
Fannin Formation. Collected 110 m above base of
section. Basal Freboldi Ammonite Zone, upper lower
Pliensbachian.
418
Toarcian, Aalenian and Bajocian localities
For further information see Appendix 1 in Carter et al.
(1988).
Skidegate Inlet, Maude Island, south side
NTS 103 F/1, Skidegate Channel. Lat. 53° 11.94’N,
Long. 132° 3.25’W. Whiteaves Formation, Maude
Group. Creek flows onto beach just east of Fannin Bay,
above a prominent sandstone with nodular coquinoid
beds containing ammonites, bivalves, brachiopods and
nautiloids (see Section 6 in Carter et al., 1988). Sequence
comprised of pale grey-green weathering shale with
limestone nodules and septarian nodules.
GSC loc. C-080579. Dark grey limestone nodule collected
in creek bed 20.5 m stratigraphically below top of
Whiteaves Formation. Occurs with Phymatoceras sp.;
middle Toarcian.
Central Graham Island
Yakoun River
NTS 103 F/8, Yakoun Lake. Phantom Creek Formation,
Maude Group. Yakoun River, Graham Island, approximately 2 km south of Ghost Creek; east side of river (see
Section 12 in Carter et al., 1988). Grey-green weathering
shale overlain by pale brown sandstone with minor shale
interbeds and common buff-weathering sandy limestone
lenses.
GSC loc. C-080583 and GSC loc. C-080584. Lat. 53°
25.19’N, Long. 132° 16.64’W. Light grey to brownishgrey sandy limestone collected 10.5 m and 14.5 m
respectively, above top of Whiteaves Formation.
Samples occur with ammonites and belemnites of late
middle or early late Toarcian age.
GSC loc. C-080597. Lat. 53° 25.22’N, Long. 132° 15.73’W.
Light grey limestone sample collected by H.W. Tipper,
1.8 km south of Ghost Creek. Occurs with late Toarcian
ammonites.
GSC loc. C-156399. Lat. 53° 25’ 20”N, Long. 132° 15’
45”W. Sample from large ellipsoidal buff-weathering
carbonate concretion (60 cm diameter) collected
5 m above base of belemnite sandstone member of
Phantom Creek Formation. Associated ammonites
include Erycitoides howelli (White) which occurs 1.0 m
above sample, and Bredia sp., which occur both 0.7 m
below and 4.3 m above sample. Sample is probably
early late Aalenian in age. For further information see
Carter & Jakobs (1991).
Branch Road 59
NTS 103 F/8, Yakoun Lake. Lat. 53° 23.19’N, Long. 132°
16.23’W. Phantom Creek Formation, Maude Group. Small
waterfall locality on east side of Branch Road 59, 0.5 km
from ‘Queen Charlotte Main’ about 0.75 km north of
junction with road to Rennell Sound (see section 13 in
Carter et al., 1988). Exposed is upper part of Phantom
Creek Formation and base of Graham Island Formation.
Lowest beds are grey-green weathering shale and siltstone,
equivalent to shales on Maude Island, Yakoun River and
above the waterfall at Rennell Junction. Irregularly bedded,
porly sorted sandstone overlies the shale and above the
sandstone is a thick sequence of interbedded shale and tuff
(Graham Island Formation).
GSC loc. C-080586. Medium dark grey sandy pelletal
limestone talus sample. ‘Hammatoceratid’ ammonites
below sample location; Tmetoceras sp. above; Aalenian.
Branch Road 57
NTS 103 F/8, Yakoun Lake. Branch Road 57, Graham
Island. Graham Island Formation, Yakoun Group. Branch
road 57 intersects ‘Queen Charlotte Main’ approximately
2 km north of its junction with the Rennell Sound Road
(see section 14 in Carter et al., 1988). Base of section in
fault contact with Phantom Creek Formation. Exposed are
dark grey shale and siltstone with rare sandy layers and
thin beds of buff weathering concretionary limestone.
GSC loc.C-080592. Lat. 53° 23.64’N, Long. 132° 16.21’W.
Brownish grey limestone sample collected in shale
34 m stratigraphically above base of Graham Island
Formation. Occurs with early Bajocian ammonites.
GSC loc. C-080593. Lat. 53° 23.63’N, Long. 132° 16.1’W.
Greenish grey limestone sample, collected in shale
57 m stratigraphically above base of Graham Island
Formation. Occurs with early Bajocian ammonites.
GSC loc. C-080595. Lat. 53° 23.63’N, Long. 132° 16.07’W.
Greenish brown limestone sample collected in shale
63 m stratigraphically above base of Graham Island
Formation. Occurs with early Bajocian ammonites.
3. EAST-CENTRAL OREGON
Hyde Formation
OR-600A. Massive, medium grey, volcaniclastic sandstone
(volcanic wacke) with occasional thin interbeds of
tuffaceous mudstone and siltstone. Well preserved
silicified Radiolaria occurring in small, dark grey,
micritic limestone nodules about 7.5 cm in diameter.
State highway 63 (Izee-Paulina road) along South Fork
of John Day River just west of bridge over river. 61 m
above base. Early Toarcian. Data from Yeh (1987b).
Material illustrated herein studied by P. Dumitrica.
Residue gift from K.-Y. Yeh.
Snowshoe Formation
OR 555. Warm Springs member of the Snowshoe Formation. Reddish-brown weathering, dark-gray, fissile
shales with dark-grey, micritic limestone nodules and
lenticular masses of silty limestone. Limestone nodules
commonly bear well-preserved silicified Radiolaria.
Sample collected 70 m above contact with the underlying Hyde Formation. National Forest road 16020 near
Duncan Hollow, 2.88 km west of intersection with
State Highway 63 (Izee-Paulina road). Probably lowermost Bajocian (discites Standard Zone) or uppermost
Aalenian (concavum Standard Zone). Data from Pessagno et al. (1986). Material illustrated herein studied by
P. Dumitrica. Residue gift from E.A. Pessagno.
4. BAJA CALIFORNIA SUR
Punta San Hipólito, Vizcaíno Peninsula
Lower third of sandstone member of the San Hipólito
Formation, type section, Pliensbachian (probably early
Pliensbachian). No age-diagnostic fossils, other than radiolarians are associated. Samples collected by P.A. Whalen
in 1980 and 1982. Detailed description and radiolarian inventory in Whalen & Carter (2002).
SH-412-14. Thin to medium-bedded, poorly sorted, olive
grey to light greenish brown, tuffaceous sandstone
interbedded with silty and cherty tuffs and silty,
tuffaceous, light grey limestone. Sample from a
light grey micritic calcareous concretion containing
abundant well-preserved silicified Radiolaria. Sample
collected approximately 250 m above base of sandstone
member. Sample from Dr. David Barnes, Western
Michigan University.
BPW80-14. Lithology same as for SH-412-14 (above).
Sample from a light grey micritic calcareous concretion containing abundant well-preserved silicified Radiolaria. Sample collected 261 m above base of sandstone member.
BPW80-15A. Lithology same as SH-412-14 (above).
Sample from a light grey slightly silty micritic
limestone bed 12-15 cm thick, containing wellpreserved silicified Radiolaria. Sample collected 270 m
above base of sandstone member.
BPW80-15B. Lithology same as for SH-412-14 (above).
Sample from a light grey, slightly silty, micritic
limestone bed 15 cm thick. Sample collected 270 m
above base of sandstone member.
BPW80-26. Lithology same as for SH-412-14 (above).
Sample from a light grey slightly silty micritic
calcareous concretion 45 cm in diameter, with rare
moderately well-preserved silicified Radiolaria. Sample
collected 282 m above base of sandstone member.
BPW80-16. Lithology same as for SH-412-14 (above).
Sample from a calcareous cannonball concretion 30 cm
in diameter, with abundant well-preserved silicified
Radiolaria. Sample collected 283 m above base of
sandstone member.
BPW80-27. Lithology same as for SH-412-14 (above).
Sample from a light greyish-tan, slightly silty micritic
calcareous concretion 60 cm long and 18 cm thick,
with moderately well-preserved silicified Radiolaria.
Sample collected 293 m above base of sandstone
member.
BPW80-28. Lithology same as for SH-412-14 (above).
Sample from a tan micritic limestone bed 6.3 cm thick
containing well-preserved silicified Radiolaria. Sample
collected 303 m above base of sandstone member.
BPW80-29. Lithology same as for SH-412-14 (above).
Sample from a light grey micritic limestone bed 10 cm
thick containing moderately well-preserved silicified
Radiolaria. Sample collected approximately 345 m
above base of sandstone member.
BPW80-30. Lithology same as for SH-412-14 (above).
Sample from a medium grey slightly silty micritic
419
limestone bed 5 cm thick containing abundant wellpreserved silicified Radiolaria. Sample collected 345 m
above base of sandstone member.
5. SLOVENIA
Mt. Mangart, Julian Alps
The section is exposed near the Slovenian–Italian border
(N 46026’80”, E 13039’18”, alt. 2164 m). The succession is
28 m thick and consists of organic and manganese rich
calcareous shales with interbedded dark grey siliceous
radiolarian-bearing limestone (Skrile Formation of Šmuc,
2005). No age-diagnostic fossils other than radiolarians are
associated. Detailed description and radiolarian inventory
given in Goričan et al. (2003). The lower part of the
succession (samples MM 5.00, MM 6.76 and MM 11.76)
is assigned to the early Toarcian, the upper part (samples
MM 21.70 and MM 27.20) may range to the middle
Toarcian.
MM 5.00. Dark grey siliceous limestone, 3.60 m above the
base of the Skrile Formation.
MM 6.76. Dark grey siliceous limestone.
MM 11.76. Dark grey siliceous limestone.
MM 21.70. Dark grey siliceous limestone.
MM 27.20. Siliceous limestone 2 m below the top of the
Skrile Formation.
6. AUSTRIA
Teltschengraben, Northern Calcareous Alps
Located in Teltschengraben, east of Bad Mitterndorf.
Dürrnberg Formation (Gawlick et al., 2001). Grey marl
and marly limestone, partly siliceous bedded limestone.
Liassic continuation of the Zlambach Formation but much
more cherty.
BMW-21. Siliceous limestone with radiolarians and sponge
spicules. Slide in Bathonian-Callovian radiolaritic
matrix (Hallstatt Mélange). Sample collected by H.-J.
Gawlick, studied by L. O’Dogherty.
7. TURKEY
Gümüslü Allochthon, Domüz Dag massif
Located 1 km NW of the Gümüslü village at 1400 m
altitude. Alternation of light grey radiolarian-bearing
limestone and marl. The overlying limestone (Ammonitico
rosso facies) contains late Pliensbachian ammonites of
the Margaritatus Zone. Detailed locality description and
stratigraphic column given in De Wever (1982b, p. 93).
1662D. Light grey bedded limestone. Sample collected
by A. Poisson. Radiolarians previously described by
Pessagno & Poisson (1981) and De Wever (1981b, c;
1982a, b). Sample restudied for this catalogue by P. De
Wever and P. Dumitrica.
420
8. OMAN
Hamrat Duru Group
In the Hamrat Duru Group of the Hawasina Nappes
the late Pliensbachian to early Bajocian radiolarian
assemblages occur in the Tawi Sadh Member of the
Guwayza Formation (Blechschmidt et al., 2004). This
member underlies the Oolitic Limestone Member of the
same formation and corresponds to the upper member
of the Matbat Formation of Béchennec (1987) and
BRGM group (1984-1993). The Tawi Sadh Member is
lithologically variable but generally consists of softer
rocks than the underlying and overlying units. For the
most part it consists of greenish to dark grey shale, greygreen bedded chert up to 10 cm thick and interbedded
shale and, in some sections and especially in the upper
part, a mixed carbonate/siliciclastic sequence consisting
of pelletal calcarenite with a variable content of sand-sized
quartz and limestone lithoclasts, sandstones and some
chert levels. The member may be very thick, up to 150 m,
or thinner. Its lower boundary may be tectonic or may lay
normally over the Al Ayn Formation, but its contact with
the overlying Oolitic Limestone Member is gradual. The
member is late Pliensbachian to early-middle Bajocian in
age on the basis of radiolarians, which are practically the
only fossils, other than sponge spicules that occur.
The Tawi Sadh Member was sampled in several sections,
those most important for the purpose of this catalogue are
described below. The samples were collected and studied
by P. Dumitrica.
Section 1 – Wadi Mu’aydin
UTM 569514/2538712 (Blechschmidt et al., 2004, fig. 8).
Several samples have been collected from this section
but only the following ones contained determinable
radiolarians.
BR1120. Grey chert at the base of the section (upper
Pliensbachian?-lower Toarcian).
BR1121. Grey chert ~4 m above the base of the section.
BR1122. Grey chert 7-8 m above base.
BR1123. Grey chert 11-12 m above base.
BR1128. Grey chert 32 m above base.
BR1129. Grey chert 34 m above base.
BR1130. Grey chert 44 m above base.
BR1131. Grey chert 46 m above base (upper Aalenianlower Bajocian).
OM-00-92. Approximately equivalent to BR1123 (studied
by Š. Goričan).
Section 2 (composite section) – Jabal Safra
UTM 582215/2513436 and UTM 586090/2512975. Three
sections were sampled in this area, two on the northern
side of the Jabal Safra ridge (sections 2A, 2B) and one on
the southern side (section 2C). The fourth section (2D)
from the northern side of Jabal Safra yielded only one
sample (BR706). Together these sections offer the most
complete radiolarian sequence of the upper Pliensbachian
to lower Bajocian interval.
On the northern side of the Jabal Safra ridge the Tawi
Sadh Member is not complete, the upper part presumably
comprising Aalenian and Bajocian strata is missing. The
missing part is exposed on the southern side of the ridge.
The two sections from the northern side of Jabal Safra
(UTM 582215/2513436) are two neighbouring parallel
sections that complement each other and partly overlap.
One section begins at the contact with the underlying Al
Ayn Formation (BR469) and ends 32 m above (BR488)
where the upper part of the succession is covered by scree
from the overlying Oolitic Limestone Member of the
Guwayza Formation. The other section (samples BR523
to BR533) continues the succession of the first section for
another 28.5 m. A portion of the lower part is probably
overlapped on a portion of the upper part of the first
section. The contact of this section with the overlying
member is covered by a scree of oolitic limestone, but
fragments of chert scattered among the scree allow the
determination of the radiolarian assemblage from beneath
the contact. Both sections consists of a monotonous
succession of green shale and grey-green, yellow
weathering chert.
The section 2C from the southern side of the Jabal Safra
ridge (UTM 586090/2512975) (samples BR824 to BR828)
differs from those exposed on the northern side. It consists
of a succession of light grey or yellowish platy limestones
with some more or less silicified levels and red radiolarites
towards the top.
Section 2A - Jabal Safra, northern side
Lower part of the Tawi Sadh Member of the Guwayza
Formation.
BR469. 1.5-3.0 m above base, green claystone and several
beds of silicified claystone and a bank of calcarenite at
the middle of the interval (Pliensbachian).
BR470. 3-4.5 m above base, intercalations of green
claystone and weakly silicified claystone.
BR471. 4.5-6 m above base, intercalations of more or less
silicified green claystone.
BR472. 6-7.5 m above base, intercalations of more or less
silicified green claystone.
BR473. 7.5-9 m above base, green claystone and more or
less silicified chert.
BR474. 9-10.5 m above base, green claystone and more or
less silicified chert.
BR475. 10.5-12 m above base, more or less silicified green
claystone.
BR476. 12-13.5 m above base, more or less silicified green
claystone.
BR477. 13.5-15 m above base, green chert and claystone.
BR478. 15-16.5 m above base, green chert and claystone.
BR479. 16.5-18 m above base, green chert and claystone.
BR480. 18-19.5 m above base, green chert and claystone.
BR481. 19.5-21 m above base, green chert and claystone.
BR482. 21-22.5 m above base, grey or green chert and
claystone.
BR483. 22.5-24 m above base, green chert and claystone.
BR484. 24-25.5 m above base, green chert.
BR485. 25.5-27 m above base, green chert.
BR486. 27-28.5 m above base, green chert.
BR487. 28.5-30 m above base, green chert and claystone.
BR488. 30-32 m above base, green chert and claystone
(Pliensbachian).
Section 2B - Jabal Safra, northern side
Tawi Sadh Member of the Guwayza Formation.
BR523. 3-4.5 m above base, green chert and claystone
(Pliensbachian).
BR524. 4.5-6 m above base, green chert and claystone.
BR525. 6-7.5 m above base, green chert and claystone.
BR526. 7.5-9 m above base, grey chert and grey marl.
BR527. 9-10.5 m above base, green chert and grey marl.
BR528. 10.5-12 m above base, grey chert and marl.
BR529. 12-13.5 m above base, grey chert and marl.
BR530. 13.5-15 m above base, grey chert, marl and 50 cm
of oolitic limestone.
BR531. 15-16.5 m above base, grey chert, marl and 20 cm
of calcarenite (upper Pliensbachian?-lower Toarcian).
BR532. 22 m above base, grey chert float.
BR533. 28.5 m above base, yellow brown chert float.
OM-99-83 and OM-99-89. Approximately equivalent to
the interval between BR529 and BR531 (studied by
Š. Goričan).
Section 2C - Jabal Safra, southern side
BR824. cca 60 m above base of Tawi Sadh Member, red
chert (upper Toarcian-lower Aalenian).
BR825/3. cca 63 m above base of Tawi Sadh Member, red
chert.
BR825. cca 74 m above base of Tawi Sadh Member, red
chert.
BR826. cca 88 m above base of Tawi Sadh Member, red
chert (Bajocian).
Section 2D - Jabal Safra, northern side
BR706. 72-76 m above base of section, which is located
about 1 km east of section 2B. The section begins in
the Al Ayn Formation and continues in the Tawi Sadh
Member of the Guwayza Formation. Sample BR706
is from the base of the Tawi Sadh Member (upper
Pliensbachian-?lower Toarcian).
BR682. Jabal Safra, coordinates not taken. Tawi Sadh
Member of the Guwayza Formation, lower Toarcian.
Section 3 - Al Sawad
UTM 578108/2544420. The section is well exposed on the
right side of the Wadi Muti upstream of the village of Al
Sawad. The succession of the Tawi Sadh Member consists
of three lithologic units. Radiolarians occur only in the
lower unit, measuring about 80 m and characterized by a
succession of dark green, more or less silicified shale with
green radiolarian chert and intercalations of yellowishbrown lithoclastic limestone. Radiolarians are frequent at
several levels but moderately preserved.
BR560. Base of section (Aalenian).
BR586. Approximately 60 m above base of section.
BR587. 2.5-3 m above BR586.
BR590. 9 m above BR587.
BR591. 11-12 m above BR587 (Aalenian-?lower Bajocian).
421
Section 4 - Wadi Saal
UTM 489029/2524022 and UTM 49001/2523209. In the
Wadi Saal, in the western part of the Hamrat Duru Range,
the Tawi Sadh Member is not well exposed mostly due to
tectonic complications, partly to its softer rocks. It is represented by white, light grey or green chert and shale and
yellowish-brown lithoclastic or oolitic limestone. Radiolarians, although visible in the chert, were generally not sufficiently preserved to be extracted. Only the following three
samples from three levels of the succession contained a determinable fauna.
BR117. cca 7-8 m above base, white chert and silicified
limestone (upper Pliensbachian?-lower Toarcian).
BR131. cca 15 m above base, white chert and silicified
limestone.
BR137. 55 m above base, with a level of chert in a
succession of yellowish-brown lithoclastic limestone
(Aalenian).
Section 5 - Al Khashbah Mt.
UTM 609565/2504490.
BR871 is a yellowish-brown chert reworked in the Oolitic
Limestone Member of the Guwayza Formation
(Aalenian).
Section 6 - FB 007
Sabt, east of Ibra (N 22°28’20’’, E 59°04’14’’). Sabt Formation according to Béchennec et al. (1993). The 60 m thick
Pliensbachian-Toarcian succession is composed predominantly of green shale and siliceous mudstone with rare
intercalations of radiolarian sandstone; chert beds occur
toward the top. This succession is overlain by calcareous
turbidites.
OM-01-21. Light green chert, collected 2.5 m below
calcareous turbidites.
Al Aridh Group
The studied samples are from the lower part of the
Musallah Formation as defined by Béchennec (1987). The
oldest age previously obtained in the Musallah Formation
was late Callovian-early Kimmeridgian (Béchennec et al.,
1993). Current radiolarian dating reveals that the base of
the formation is as old as Pliensbachian. The Pliensbachian
to upper Toarcian-lower Aalenian succession treated in
this catalogue consists of varicoloured bedded radiolarian
chert interstratified locally with resedimented limestones.
No age-diagnostic fossils other than radiolarians are
associated.
Sections measured by F. Béchennec and C. Robin,
radiolarian samples collected and studied by Š. Goričan.
Section 1 – FB 2841
Near Al Aridh north of Ibri (N 23°21’50’’, E 56°36’19’’).
Lower part of the Musallah Formation.
OM-99-131. Light reddish argillaceous chert, 1 m above
the outcropping base (lowermost part of the Musallah
Formation not exposed).
OM-99-133. Light greenish yellow argillaceous chert,
6.5 m above OM-99-131.
422
OM-99-137. (Bajocian). Red nodular argillaceous chert,
23 m above OM-99-133.
Section 2 – FB 2895
Jabal Buwaydah East (N 22º52’06.2’’, E 57º05’40.9’’). Lower
part of the Musallah Formation.
OM-00-251. Light reddish chert, 25.5 m above the contact
with medium-grained calcareous turbidites.
OM-00-252. Light reddish chert, 3 m above OM-00-251.
OM-00-254. Beige argillaceous chert, 8 m above OM-00252.
OM-00-255. White chert, 2.5 m above OM-00-254.
OM-00-256. Light yellow chert, 12.5 m above OM-00-255,
0.5 m below a succession of coarse limestone breccias.
OM-00-258 (upper Aalenian-lower Bajocian). Red nodular
argillaceous chert, approximately 30 m above OM-00256.
Section 3 – FB 156
Jabal Buwaydah Center East (N 22°55’20’’, E 57°05’35’’).
Lower part of the Musallah Formation, a 7 m thick chert
level intercalated between resedimented oolitic limestone
below and limestone breccia above.
OM-00-117. Yellow argillaceous chert near the top of the
chert level.
OM-00-118. Light red argillaceous chert, approximate
lateral equivalent of sample OM-00-117.
Section 4 – FB 124
Jabal Buwaydah East (N 22°51’36’’, E 57°07’43’’). Lower
part of the Musallah Formation.
OM-00-263. Red chert within a 1 m thick chert level
intercalated between two limestone breccia beds.
Section 5 – FB 298
Jabal Buwaydah Center West (N 22º59’9’’, E 57º01’8’’).
Lower part of the Musallah Formation, 7 m thick chert
succession bounded by a tectonic contact below and
limestone breccias above.
OM-00-115. Vivid red chert, base of chert succession.
OM-00-114. Yellow chert with red spots, 6 m above sample
OM-00-115.
Umar Group
Humadiyin
UTM 2247563N/5747425E, Haliw (Aqil) Formation.
According to Béchennec (1987), the Umar Group is
the uppermost structural unit of the Hawasina Nappes
and comprises two formations: the Sinni Formation,
composed mainly of Triassic volcanic rocks and the Aqil
Formation, composed predominantly of Middle Triassic
to Cretaceous siliceous pelagic deposits, calcirudite and/or
megabreccias of reworked carbonates. The latter formation
was initially separated by Glennie et al. (1974) as the Haliw
Formation from the same area as the Aqil Formation
and, consequently, has priority over the latter. The upper
Norian to Lower Jurassic is represented here by red chert
and is now under study.
O38 and O39. For the moment the only information
on the Early Jurassic radiolarians from the Haliw
Formation comes from these two lower Pliensbachian
samples, collected from floated fragments under
megabreccias that cover the chert sections near the
locality Humadiyin, on the north side of the road
Nizwa-Sinew. Radiolarian samples collected by
L. Krystyn and studied by P. Dumitrica.
9. JAPAN
Mino Terrane - Inuyama area
Lower Jurassic (Hettangian-Toarcian), Lat. 35° 25±2’N;
long. 136° 58±2’E. The Kamiaso Unit (Wakita, 1988),
Mino Terrane, SW Japan, is one of the Jurassic accretionary complexes comprised of bedded cherts and clastic
sedimentary rocks. A late Bathonian – early Callovian ammonite, Choffatia sp., was reported from the clastic rocks
(Sato, 1974; Sato & Westermann, 1985). Triassic-Lower
Jurassic radiolarian and upper Lower Triassic and Middle–Upper Triassic conodont fossils were obtained from
bedded chert sequences in this area and biostratigraphic
studies were summarized (e.g. Hori, 1988; 1990; Matsuda
& Isozaki, 1991; Sugiyama, 1997).
Samples for this study were collected by R. S. Hori
(Katsuyama (UF), Iwayakannon (IY), Kurusu (KU),
UC sections) and R.S. Hori and M. Takeuchi (KA log =
Pliensbachian – Toarcian part of UF section and IW log =
those of IY section). See Hori (1990) for section locations
in the Inuyama area. All samples were obtained from
bedded chert sequences.
The outcrops of these sections are located on the right
bank of the Kisogawa River (UF, IY and UC sections)
and the left bank (KU section). Four chert sequences of
the Kamiaso Unit are exposed along the Kisogawa River,
namely CH-1, CH-2, CH-3 and CH-4 in structurally
ascending order (Yao et al., 1980).
The sedimentation rate of Lower Jurassic chert sequences
from the Inuyama area is ca. 1m/m.y. = 0.1cm/kyr (Hori et
al., 1993).
Katsuyama (UF) Section (Hori, 1990)
This section is located on the CH-3 chert-sheet near the
Katsuyama road junction, Gifu prefecture, SW Japan.
Detailed sample locations of the outcrop and vertical
distributions of representative taxa are shown in Hori
(1992). Between UFI19 and UFI21, the Toarcian Oceanic
Anoxic Event (OAE) was recorded lithologically and
geochemically (Hori, 1993).
The UF section consists of ca. 32 m thick chert sequence
which is one of the most complete sections of Upper
Triassic to Lower Jurassic strata in the Inuyama are. The
outcrop extends down to the Middle and upper Lower
Triassic (Matsuoka et al., 1994). Hori (1992) and Carter &
Hori (2005) reported detailed radiolarian and conodont
biostratigraphy at the Triassic/Jurassic boundary of this
section. The basal horizon (0 cm) is located at UFI3, which
is the dusty red chert bed (T/J boundary event level) =Kb1
(01) of Hori (1992) and Carter & Hori (2005).
UFI6. 560 cm above top of dusty red chert bed, red bedded
chert.
UFI7. 720 cm above top of dusty red chert bed, red bedded
chert.
UFI8. 860 cm above top of dusty red chert bed, red bedded
chert.
UFI9. 960 cm above top of dusty red chert bed, red bedded
chert.
UFI10. 1070 cm above top of dusty red chert bed, red
bedded chert.
UFI11. 1170 cm above top of dusty red chert bed, red
bedded chert.
UFI12. 1280 cm above top of dusty red chert bed, red
bedded chert.
UFI13. 1410 cm above top of dusty red chert bed, red
bedded chert.
UFI14. 1840 cm above top of dusty red chert bed, green
bedded chert.
UFI15. 1990 cm above top of dusty red chert bed, green
bedded chert
UFI16. 2050 cm above top of dusty red chert bed, grey
bedded chert.
UFI17. 2170 cm above top of dusty red chert bed, grey
bedded chert.
UFI18. 2240 cm above top of dusty red chert bed, grey
bedded chert.
UFI19. 2460 cm above top of dusty red chert bed, grey
bedded chert; just below the OAE.
UFI20. 2540 cm above top of dusty red chert bed; high
peak of Toarcian OAE. (70 cm below the base of white
massive chert bed). No radiolarian data.
UFI21. 2720 cm above top of dusty red chert bed, green
and red bedded chert; above the OAE.
UFI22. 2860 cm above top of dusty red chert bed, green
bedded chert.
UFI23. 3160 cm above top of dusty red chert bed, green
bedded chert.
UFI24. 3220 cm above top of dusty red chert bed, green
bedded chert.
Kb log of UF section (Hori, 1992)
This log is located around the Triassic/Jurassic boundary
of the UF section. Original data and sample locations were
published in Hori (1992). Kb01 is the same level as Kb1 in
Carter & Hori (2005). The base of this Kb log is located at
the Kb01 which is the purple red (or dusty red) chert bed
characterized by the extinction level of conodont Misikella
posthernsteini.
Kb07. 281 cm above the Kb01 bed.
Kb09. 368 cm above the Kb01 bed.
Kb10. 419 cm above the Kb01 bed.
Kb11. 493 cm above the Kb01 bed.
Kb12. 543 cm above the Kb01 bed.
Kb13. 575 cm above the Kb01 bed.
Kb14. 679 cm above the Kb01 bed.
Kb15. 724 cm above the Kb01 bed and 70 cm below the
UFI10 of the UF section.
423
KA log of UF section (Takeuchi, 2001)
The KA log is the upper Pliensbachian - Toarcian part of
the UF section, focusing on the interval of the Toarcian
OAE. The base of this section is a white massive chert bed
2790 cm above the top of the dusty red chert bed (Kb01)
of the UF section. The sketch map of this white massive
chert was shown in Fig. 3 of Hori (1992), the lithological
column of this section was illustrated in Takeuchi (2001).
KA:-170. 504 cm below base of white massive chert bed.
KA:-150. 470 cm below base of white massive chert bed.
KA:-121. 393 cm below base of white massive chert bed.
KA:-95. 311 cm below base of white massive chert bed; pre
OAE level.
KA:-5. 20 cm below base of white massive chert bed; in
OAE level.
KA:+22. 127 cm above base of white massive chert bed;
post black chert (OAE) level.
KA:+25. 131 cm above base of white massive chert bed.
KA:+40. 157 cm above base of white massive chert bed.
KA:+55. 187 cm above base of white massive chert bed.
KA:+60. 202 cm above base of white massive chert bed.
KA:+68. 214 cm above base of white massive chert bed.
Iwayakannon (IY) Section (Hori, 1988)
The sketch map and sample locations of this section are
shown in Hori (1988). This section consists of ca. 25 m
bedded chert sequence located on the CH-4 chert-sheet
of Yao et al. (1980). Detail sample locations and outline
of vertical distributions of representative taxa are shown
in Hori (1988). The top of this section is in contact with
Middle Jurassic green mudstone containing the Unuma
echinatus assemblage. The Toarcian OAE level is correlated
with a thick shale bed just below IYII 14.
IYIII1. 1940 cm below top of section.
IYII24. 1880cm below top of section.
IYII23. 1810 cm below top of section.
IYII22. 1630 cm below top of section.
IYII21. 1490 cm below top of section.
IYII18. 1140 cm below top of section.
IYII17. 960 cm below top of section.
IYI7. 810 cm below top of section.
IYII14. 580 cm below top of section.
IYI6. 510 cm below top of section.
IYII13. 490 cm below top of section.
IYII12. 440 cm below top of section.
IYII11. 380 cm below top of section.
IYII10. 310 cm below top of section.
IYII9. 200 cm below top of section.
IYI4. 180 cm below top of section.
IYII8. 120 cm below top of section.
IYII7. 90 cm below top of section.
IYII6. 80 cm below top of section.
IYII5. 65 cm below top of section.
IYII4. 40 cm below top of section.
IYII3. 20 cm below top of section.
IYII2. At the top of the section.
IW log of IY section (Takeuchi, 2001)
The standard level of this section is located at the base
424
of a thick green shale bed just below IYII 14 (6 m below
the top of the IY section). The lithological column of this
section is illustrated in Takeuchi (2001).
IW:-2. 5 cm below base of thick green shale bed.
IW:+7. 22 cm above base of thick green shale bed.
IW:+18. 45 cm above base of thick green shale bed.
IW:+24. 57 cm above base of thick green shale bed.
IW:+37. 97 cm above base of thick green shale bed.
IW:+51. 125 cm above base of thick green shale bed.
IW:+57. 137 cm above base of thick green shale bed.
Kurusu (KU) Section (Hori, 1988)
The KU section was first described by Hori (1988). This
section is located on the left bank of the Kisogawa River,
Inuyama City, Aichi Prefecture. It is one of the most
complete sequences of Triassic/Jurassic boundary bedded
cherts in the Inuyama area. The cherts are ca. 35 m thick
without remarkable lithological change such as boundary
clay.
KU(b)14. 2261 cm below KU (a)1, green chert.
KU(b)13. 2126 cm below KU (a)1, greenish grey chert.
KU(b)12. 1950 cm below KU(a)1, greenish grey chert.
KU(b)11. 1820 cm below KU(a)1, greenish grey chert.
KU(b)5. 1519 cm below KU(a)1, black chert.
KU(b) 4. 751 cm below KU(a)1, grey chert.
KU(b)3. 670 cm below KU(a)1, black chert.
KU(b)1. 420 cm below KU(a)1, black chert.
KU(a)7. 310 cm below KU(a)1, green chert; thick shale
bed intercalated which corresponds to Toarcian OAE
level.
KU(a)5. 260 cm below KU(a)1, grey chert.
KU(a)4. 220 cm below KU(a)1, greenish grey chert.
KU(a)3. 140 cm below KU(a)1, green chert.
KU(a)2. 70 cm below KU(a)1, green chert.
KU(a)1. Top 0 cm grey chert.
UC Section (Hori, 1986; also data provided by Hori, 1990,
1993)
This section consists of 11 m of bedded chert that extends
from the upper Sinemurian to Aalenian. Black cherts
including FeS2 minerals occur in the upper part of the
section (ca. 520 cm level), which suggest the Toarcian OAE
(Hori, 1993). Lithological and preliminary radiolarian data
are shown in Hori (1986, 1990).
UC1. 0 cm, greenish red chert (Pliensbachian chert).
UC2. 27 cm above UC1, greenish red chert.
UC3. 86 cm above UC1, greenish red chert.
UC4. 118 cm above UC1, red chert.
UC5. 138 cm above UC1, red chert.
UC6. 164 cm above UC1, greenish red chert.
UC7. 205 cm above UC1, greenish red chert.
UC8. 227 cm above UC1, greenish grey chert.
UC9. 280 cm above UC1, grey chert.
UC10. 341 cm above UC1, grey chert.
UC11. 373 cm above UC1, grey chert.
UC12. 423 cm above UC1, grey chert.
UC13. 472 cm above UC1, grey chert.
UC14. 515 cm above UC1, grey chert; just below the
beginning of black chert of the Toarcian OAE.
UC15. 714 cm above UC1, greenish black chert; Toarcian
OAE.
UC16. 755 cm above UC1, greenish black chert; Toarcian
OAE.
UC17. 842 cm above UC1, greenish black chert; Toarcian
OAE.
Mino Terrane - Nanjo area
This area is located in the northwestern part of the Mino
Terrane (Lat. 35˚, 42±1’N; Long.136˚, 17±1E). Lithological
data by Hori (1990 MSc thesis). Samples collected by
R. S. Hori.
Mélange rocks belonging to the Lower Jurassic accretionary complex in the Mino Terrane are exposed in this
area. A chert sample (85072502B) obtained from the same
locality as Ito & Matsuda (1980) is from one of the tectonic
blocks and the mudstone sample is from the matrix of the
mélange.
85072502B. Green chert block in Jurassic sedimentary
complex.
850725044. Mudstone, mélange matrix of Jurassic
complex.
Nanjo Massif - Imajo unit
MNA-10. Manganese band in a siliceous mudstone layer.
This band contains manganese carbonate spherules
ranging from 0.5 to 2.0 mm in diameter. Radiolarian
fauna diverse and well preserved. One of the bestpreserved radiolarian-bearing samples of Toarcian age.
Radiolarians studied by A. Matsuoka (1991, 2004, this
catalogue).
IH84120461 and IH84120462. Two samples given by I.
Hattori to P. Dumitrica. According to Hattori (1989)
the samples are from rhodochrosite concretions in
red shale outcropping in the Nanjo Massif, SugentanMinami locality (Hattori, 1989, fig. 3). Radiolarians
from this locality have been illustrated by Hattori
(l. cit.) in plates 18-34 and the list of species occurring
in the two samples (61 and 62) are in Hattori (1989,
Table 2). The assemblage proves an Aalenian age.
Mino Terrane - Mt. Norikuradake area
Yukawa Complex by Otsuka (1988), northeastern part of
the Mino Terrane (Lat. 36˚, 8±1’N; Long. 137˚, 27±1’E),
SW Japan. Samples collected by R. S. Hori. PliensbachianToarcian bedded cherts and Toarcian-Aalenian siliceous
shale occur in this area. Lithological data and Toarcian
radiolarian fauna documented by Hori & Otsuka (1989).
Descriptions of sample locations shown in Hori (1988,
1990) and Hori & Otsuka (1989).
Norikuradake (NK) section (Hori & Otsuka, 1989)
Lat. 36˚ 7’36”N; Long. 137˚ 27’ 30”E.
NKI7. +0 m chert bed.
NKI8. +8.04 m above the NKI7 chert, chert bed.
NKI9. +17.1 m above the NKI7 chert, chert bed.
NKI10. +24 m above the NKI7 chert, chert bed. A minor
fault developed this level.
NKI11. – 21.3 m below the top of NKI17 chert bed, just
above the fault, chert bed.
NKI12. –13.2 m below the top of NKI17 chert bed.
NKII2. –9.64 m below the top of NKI17 chert bed.
NKII3. –7.07 m below the top of NKI17 chert bed.
NKII4. –3.93 m below the top of NKI17 chert bed.
NKII5. – 2.68 m below the top of NKI17 chert bed.
NKI17. +0 m just below the red shale.
NKII7. -2 +0.43 m above the NKI17, red siliceous shale.
NKII8. +1 m above the NKI17, red siliceous shale.
NKII9. +1.7 m above the NKI17, red chert.
NKII10. +2.31 m above the NKI17, red siliceous shale.
NKI20. + 2.68 m above the NKI17, red siliceous shale.
Mino Terrane - Middle Jurassic
Inuyama area
Radiolarians studied by A. Matsuoka.
MIN-1. Inuyama area, Aichi Prefecture. The locality is
on the left bank of the Kisogawa River. The sample is
a manganese carbonate band included in a siliceous
mudstone layer south of the CH-2 chert-sheet of Yao
et al. (1980). It contains 93 nassellarian species (Matsuoka, 1992) and is assignable to the Tricolocapsa plicarum Zone or JR 4 of Matsuoka (1995).
MIN-10. Inuyama area, Aichi Prefecture. Same locality as
MIN-1. The sample is assignable to the Laxtorum (?)
jurassicum Zone or JR 3 of Matsuoka (1995).
Kamiaso (Hisuikyo) area
Lat. 35˚, 32’52”N; Long.137˚, 7’47”E. Triassic and Jurassic
radiolarian bedded cherts and Middle Jurassic siliceous
mudstones well exposed along the Hida River, which
formed Hisukyo Gorge in the Kamiaso area, Gifu Prefecture of SW Japan.
Matsuda & Isozaki (1982), and Isozaki & Matsuda
(1985) documented Lower Jurassic radiolarian fossils
from this area. In particular, a black chert bed containing
manganese carbonate spherules from the Hisuikyo Gorge
contains well-preserved radiolarian fossils described by
Isozaki and Matsuda (1985); this is the type locality of the
Hsuum hisuikyoense Assemblage. Sample mentioned in
this study (only for systematic part) collected by R. S. Hori
from the black chert horizon of Isozaki & Matsuda (1985).
Gujo-Hachiman area
Lat. 35˚, 49.8’N; Long.136˚, 52’55”E. Radiolarian fossils
from this area were described by Takemura (1986), and
Yao (1997).
Samples in the systematic part of this study collected
by R. S. Hori from the same area as Takemura (1986).
Extremely well preserved radiolarian fossils from
manganese carbonate nodules in black shale (e.g. Wakita,
1982, 1984). The radiolarian faunas are correlated with
UAZ 3 of Baumgartner et al. (1995b) (late Aalenian?) as
discussed by Yao (1997).
425
MKM-1 (studied by A. Matsuoka). The locality is the same
as that of Takemura (1986). The sample contains 64
nassellarian species (Matsuoka, 1992) and is assignable
to the Laxtorum (?) jurassicum Zone or JR 3 of
Matsuoka (1995).
Chichibu Terrane
Kuma area
The Kuma area is located in western Kyushu, southwest
Japan. The biostratigraphy of Mesozoic radiolarians has
been documented mainly by Nishizono et al. (1982),
426
Nishizono & Murata (1983), Sato & Nishizono (1983) and
Nishizono (1996).
Samples mentioned in this study (only for systematic
descriptions) were collected by R. S. Hori. Precise sample
numbers and radiolarian data were shown in Hori (1990).
Kaiji (KG) and Kajiki-1 (=Kajiki: KS in Hori, 1990)
sections were investigated by Matsuoka & Yao (1986),
Hori (1990), and Matsuoka (1995). The KG section ranges
from the Parahsuum simplum subzone I to subzone
IV (Sinemurian to Pliensbachian) and the KS section
corresponds to the Mesosaturnalis hexagonus Assemblage
Zone to the Hsuum hisuikyoense Assemblage Zone
(Toarcian to Aalenian) (Hori, 1990).
KG-9. Chert. Top of Parahsuum simplum subzone II.
4. LISTING OF SPECIES
4.1. Alphabetical listing by genus
2001
4066
JAC02
JAC04
ADM01
ADM02
ADM03
3149
3271
HAG01
HAG02
ASP01
ATT01
ARS03
ARS07
ARS06
4061
3001
4032
ARS04
ARS01
ARS02
ARS08
4008
ATA02
BAG01
BAG03
BAG02
BAG04
BAG05
BAG06
BAG07
ORB04
PDC01
SPI03
ORB07
CRU18
3222
BER01
BPD13
BPD05
BPD14
Acaeniotylopsis ghostensis (Carter) 1988
Acaeniotylopsis triacanthus Kito & De Wever
1994
Anaticapitula anatiformis (De Wever) 1982a
Anaticapitula omanensis Dumitrica n. sp.
Archaeodictyomitra munda (Yeh) 1987b
Archaeodictyomitra sp. A
Archaeodictyomitra sp. B
Archaeohagiastrum longipes Baumgartner 1995
Archaeohagiastrum munitum Baumgartner 1984
Archaeohagiastrum oregonense (Yeh) 1987b
Archaeohagiastrum pobi Whalen & Carter 1998
Archaeospongoprunum coyotense Whalen &
Carter 2002
Archaeotritrabs hattorii Dumitrica n. sp.
Ares armatus De Wever 1982a
Ares avirostrum Dumitrica & Matsuoka n. sp.
Ares cuniculiformis Dumitrica & Whalen n. sp.
Ares cylindricus s.l. (Takemura) 1986
Ares cylindricus cylindricus (Takemura) 1986
Ares cylindricus flexuosus (Takemura) 1986
Ares mexicoensis Whalen & Carter 2002
Ares moresbyensis Whalen & Carter 1998
Ares sutherlandi Whalen & Carter 1998
Ares takemurai Dumitrica & Matsuoka n. sp.
Ares sp. A sensu Baumgartner et al. 1995a
Atalanta emmela Cordey & Carter 1996
Bagotum erraticum Pessagno & Whalen 1982
Bagotum funiculum Whalen & Carter 2002
Bagotum helmetense Pessagno & Whalen 1982
Bagotum kimbroughi Whalen & Carter 2002
Bagotum maudense Pessagno & Whalen 1982
Bagotum modestum Pessagno & Whalen 1982
Bagotum pseudoerraticum Kishida & Hisada
1985
Beatricea? argescens (Cordey) 1998
Beatricea? baroni Cordey 1998
Beatricea christovalensis Whalen & Carter 1998
Beatricea sanpabloensis (Whalen & Carter) 2002
Beatricea? sp. A
Bernoullius delnortensis Pessagno, Blome & Hull
1993
Bernoullius saccideon (Carter) 1988
Bipedis calvabovis De Wever 1982a
Bipedis diadema Whalen & Carter 1998
Bipedis fannini Carter 1988
427
BPD15
BPD16
BIS04
BIS02
BIS01
BIS03
BRO02
BRO03
BRO01
CAN12
CAN13
CAN08
CAN09
CAN11
CAN14
CTS06
CTS08
CTS09
CTS10
CTS15
CTS03
CTS12
CTS16
SUM03
CHA02
CHA09
CHA10
CHA03
CHA05
CHA07
CHA08
CHA11
XNM01
4033
CIT05
CRB01
CRU21
CRU11
CRU12
PDC02
CRU22
CRU10
CRU20
CRU19
PDC05
CRU13
CRU14
CRU15
CRU16
3131
CYC01
CYC02
CYC03
CYC04
DAN02
DRO07
DRO02
DRO03
428
Bipedis japonicus Hori n. sp.
Bipedis yaoi Hori n. sp.
Bistarkum mangartense Goričan, Šmuc &
Baumgartner 2003
Bistarkum phantomense (Carter) 1988
Bistarkum rigidium Yeh 1987b
Bistarkum saginatum Yeh 1987b
Broctus kuensis Pessagno & Whalen 1982
Broctus ruesti Yeh 1987b
Broctus selwynensis Pessagno & Whalen 1982
Canoptum anulatum Pessagno & Poisson 1981
Canoptum artum Yeh 1987b
Canoptum columbiaense Whalen & Carter 1998
Canoptum dixoni Pessagno & Whalen 1982
Canoptum margaritaense Whalen & Carter 1998
Canoptum rugosum Pessagno & Poisson 1981
Canutus baumgartneri Yeh 1987b
Canutus diegoi Whalen & Carter 2002
Canutus hainaensis Pessagno & Whalen 1982
Canutus nitidus Yeh 1987b
Canutus rennellensis Carter n. sp.
Canutus rockfishensis Pessagno & Whalen 1982
Canutus tipperi gr. Pessagno & Whalen 1982
Canutus sp. O
Carterwhalenia minai (Whalen & Carter) 2002
Charlottea amurensis Whalen & Carter 1998
Charlottea hotaoensis Carter n. sp.
Charlottea penderi Carter n. sp.
Charlottea proprietatis Whalen & Carter 1998
Charlottea triquetra Whalen & Carter 1998
Charlottea sp. A sensu Whalen & Carter 2002
Charlottea sp. B
Charlottea sp. C
Charlottea? sp. Y
Citriduma hexaptera (Conti & Marcucci) 1991
Citriduma radiotuba De Wever 1982a
Crubus chengi Yeh 1987b
Crucella angulosa s.l. Carter 1988
Crucella angulosa angulosa Carter 1988
Crucella angulosa longibrachiata Carter n. ssp.
Crucella beata (Yeh) 1987b
Crucella cavata s.l. Whalen & Carter 1998
Crucella cavata cavata Whalen & Carter 1998
Crucella cavata giganticava Carter n. ssp.
Crucella cavata intermedicava Carter n. ssp.
Crucella jadeae Carter & Dumitrica n. sp.
Crucella mijo De Wever 1981b
Crucella mirabunda Whalen & Carter 2002
Crucella spongase De Wever 1981b
Crucella squama (Kozlova) 1971
Crucella theokaftensis Baumgartner 1980
Cyclastrum asuncionense Whalen & Carter 2002
Cyclastrum scammonense Whalen & Carter 2002
Cyclastrum veracruzense Whalen & Carter 2002
Cyclastrum sp. A
Danubea sp. A sensu Whalen & Carter 2002
Droltus eurasiaticus Kozur & Mostler 1990
Droltus hecatensis Pessagno & Whalen 1982
Droltus laseekensis Pessagno & Whalen 1982
DRO06
DRO08
DUC01
JAC05
3411
PHS08
ELD02
ELD03
2021
EUC09
EUC10
EUC03
EUC06
EUC07
EUC04
FAR02
FAR04
FAR03
FRM01
GIG01
GOR02
GOR03
HCK05
HCK04
HAG06
HAG03
HAG04
TPS03
SCP03
SCP06
SCP04
SCP05
3502
SAT11
3089
HIG01
HIG04
HIG03
HOM01
HOM02
HSU01
HSU02
HSU03
HSU04
HSU05
3195
3278
HSU06
Droltus lyellensis Pessagno & Whalen 1982
Droltus sanignacioensis Whalen & Carter 2002
Ducatus hipolitoensis Whalen & Carter 2002
Dumitricaella trispinosa Dumitrica n. sp.
Elodium cameroni Carter 1988
Elodium? mackenziei Carter n. sp.
Elodium pessagnoi Yeh & Cheng 1996
Elodium wilsonense (Carter) 1988
Eospongosaturninus protoformis (Yao) 1972
Eucyrtidiellum disparile gr. Nagai & Mizutani
1990
Eucyrtidiellum gujoense (Takemura & Nakaseko)
1986
Eucyrtidiellum gunense gr. Cordey 1998
Eucyrtidiellum nagaiae Dumitrica, Goričan &
Matsuoka n. sp.
Eucyrtidiellum omanojaponicum Dumitrica,
Goričan & Hori n. sp.
Eucyrtidiellum ramescens Cordey 1998
Farcus asperoensis Pessagno, Whalen & Yeh 1986
Farcus graylockensis Pessagno, Whalen & Yeh
1986
Farcus kozuri Yeh 1987b
Foremania sandilandsensis gr. Whalen & Carter
1998
Gigi fustis De Wever 1982a
Gorgansium gongyloideum Kishida & Hisada
1985
Gorgansium morganense Pessagno & Blome
1980
Haeckelicyrtium crickmayi Carter n. sp.
Haeckelicyrtium sp. B sensu Whalen & Carter
2002
Hagiastrum macrum gr. De Wever 1981b
Hagiastrum majusculum Whalen & Carter 1998
Hagiastrum rudimentum Whalen & Carter 1998
Helvetocapsa minoensis (Matsuoka) 1991
Helvetocapsa nanjoensis (Matsuoka) 1991
Helvetocapsa plicata s.l. (Matsuoka) 1991
Helvetocapsa plicata plicata (Matsuoka) 1991
Helvetocapsa plicata semiplicata (Matsuoka)
1991
Hexasaturnalis hexagonus (Yao) 1972
Hexasaturnalis octopus Dumitrica & Hori n. sp.
Hexasaturnalis tetraspinus (Yao) 1972
Higumastra laxa Yeh 1987b
Higumastra lupheri Yeh 1987b
Higumastra transversa Blome 1984b
Homoeoparonaella lowryensis Whalen & Carter
2002
Homoeoparonaella reciproca Carter 1988
Hsuum altile Hori & Otsuka 1989
Hsuum arenaense Whalen & Carter 2002
Hsuum busuangaense Yeh & Cheng 1996
Hsuum exiguum Yeh & Cheng 1996
Hsuum lucidum Yeh 1987b
Hsuum matsuokai Isozaki & Matsuda 1985
Hsuum medium (Takemura) 1986
Hsuum mulleri Pessagno & Whalen 1982
Hsuum optimum Carter 1988
Hsuum philippinense Yeh & Cheng 1996
Hsuum plectocostatum Carter n. sp.
Hsuum sp. A sensu Carter 1988
Katroma angusta Yeh 1987b
Katroma aurita Whalen & Carter 2002
Katroma bicornus De Wever 1982a
Katroma brevitubus Dumitrica & Goričan n. sp.
Katroma clara Yeh 1987b
Katroma elongata Carter n. sp.
Katroma neagui Pessagno & Poisson 1981,
emend. De Wever 1982a
KAT14 Katroma ninstintsi Carter 1988
KAT16 Katroma? sinetubus Carter n. sp.
KAT18 Katroma sp. 4
LAN05 Lantus intermedius Carter n. sp.
LAN01 Lantus obesus (Yeh) 1987b
LAN04 Lantus praeobesus Carter n. sp.
LAN02 Lantus sixi Yeh 1987b
LAN03 Lantus sp. A sensu Whalen & Carter 2002
LAX06 Laxtorum hemingense Whalen & Carter 1998
MCP01 Minocapsa cylindrica Matsuoka 1991
MCP02 Minocapsa globosa Matsuoka 1991
TPS02 Minocapsa? megaglobosa (Matsuoka) 1991
NAP09 Napora blechschmidti Dumitrica n. sp.
NAP08 Napora bona Pessagno, Whalen & Yeh 1986
NAP02 Napora cerromesaensis Pessagno, Whalen & Yeh
1986
NAP06 Napora conothorax Carter & Dumitrica n. sp.
NAP01 Napora graybayensis Pessagno, Whalen & Yeh
1986
3410
Napora nipponica Takemura 1986
NAP03 Napora reiferensis (Pessagno, Whalen & Yeh) 1986
NAP04 Napora relica Yeh 1987b
JAC01 Napora sandspitensis (Pessagno, Whalen & Yeh)
1986
UTD01 Naropa vi Hori, Whalen & Dumitrica n. sp.
NTS01 Noritus lillihornensis Pessagno & Whalen 1982
ORB05 Orbiculiformella callosa (Yeh) 1987b
ORB06 Orbiculiformella incognita (Blome) 1984b
ORB03 Orbiculiformella lomgonensis (Whalen & Carter)
1998
ORB11 Orbiculiformella mediocircus Dumitrica n. sp.
ORB02 Orbiculiformella? robusta (Whalen & Carter)
1998
ORB08 Orbiculiformella teres (Hull) 1997
ORB13 Orbiculiformella? trispina s.l. (Yeh) 1987b
ORB09 Orbiculiformella? trispina trispina (Yeh) 1987b
ORB10 Orbiculiformella? trispina trispinula (Carter)
1988
SAT13 Palaeosaturnalis aff. liassicus Kozur & Mostler
1990
SAT12 Palaeosaturnalis subovalis Kozur & Mostler 1990
SAT14 Palaeosaturnalis sp. B sensu Whalen & Carter
2002
PAN20 Pantanellium brevispinum Carter n. sp.
PAN14 Pantanellium carlense Whalen & Carter 1998
PAN18 Pantanellium cumshewaense Pessagno & Blome
1980
HSU07
HSU08
HSU11
HSU10
KAT07
KAT08
KAT09
KAT12
KAT10
KAT17
KAT13
Pantanellium danaense Pessagno & Blome 1980
Pantanellium inornatum Pessagno & Poisson
1981
PAN16 Pantanellium skedansense Pessagno & Blome 1980
PHS02 Parahsuum edenshawi (Carter) 1988
DRO05 Parahsuum fondrenense (Whalen & Carter) 1998
PHS09 Parahsuum formosum (Yeh) 1987b
2012
Parahsuum izeense (Pessagno & Whalen) 1982
PHS03 Parahsuum longiconicum Sashida 1988
PHS04 Parahsuum mostleri (Yeh) 1987b
PHS05 Parahsuum ovale Hori & Yao 1988
PHS01 Parahsuum simplum Yao 1982
PHS06 Parahsuum vizcainoense Whalen & Carter 2002
PHS07 Parahsuum? sp. A sensu Whalen & Carter 2002
2013
Parasaturnalis diplocyclis (Yao) 1972
SAT15 Parasaturnalis yehae Dumitrica & Hori n. sp.
PAR13 Paronaella corpulenta De Wever 1981b
PAR22 Paronaella curticrassa Carter & Dumitrica n. sp.
PAR24 Paronaella fera s.l. (Yeh) 1987b
PAR15 Paronaella fera fera (Yeh) 1987b
PAR10 Paronaella fera jamesi Whalen & Carter 1998
PAR16 Paronaella grahamensis Carter 1988
PAR17 Paronaella notabilis Whalen & Carter 2002
2005
Paronaella skowkonaensis Carter 1988
PAR19 Paronaella snowshoensis (Yeh) 1987b
PAR20 Paronaella tripla De Wever 1981b
PAR21 Paronaella variabilis Carter 1988
PSP03 Perispyridium hippaense (Carter) 1988
PSP01 Perispyridium oregonense (Yeh) 1987b
PLE01 Pleesus aptus Yeh 1987b
SCP02 Plicaforacapsa? elegans (Matsuoka) 1991
POD01 Podocapsa abreojosensis Whalen & Carter 2002
PRY05 Praeconocaryomma bajaensis Whalen n. sp.
PRY01 Praeconocaryomma decora gr. Yeh 1987b
PRY02 Praeconocaryomma immodica Pessagno &
Poisson 1981
PRY03 Praeconocaryomma parvimamma Pessagno &
Poisson 1981
PRY07 Praeconocaryomma sarahae Carter n. sp.
PRY04 Praeconocaryomma whiteavesi Carter 1988
PRY06 Praeconocaryomma? yakounensis Carter n. sp.
SAT01 Praehexasaturnalis tetraradiatus Kozur &
Mostler 1990
PVG01 Praeparvicingula aculeata (Carter )1988
PVG02 Praeparvicingula elementaria (Carter) 1988
PVG03 Praeparvicingula gigantocornis (Kishida &
Hisada) 1985
PVG04 Praeparvicingula nanoconica (Hori & Otsuka)
1989
TVS01 Praeparvicingula? spinifera (Takemura) 1986
PCA02 Praeparvicingula tlellensis Carter n. sp.
PTP01 Protopsium gesponsa De Wever 1981c
PRU01 Protunuma paulsmithi Carter 1988
PDC03 Pseudocrucella ornata De Wever 1981b
3126
Pseudocrucella sanfilippoae (Pessagno) 1977a
PDC04 Pseudocrucella sp. C sensu Carter 1988
PSE02 Pseudoeucyrtis angusta Whalen & Carter 1998
PSE04 Pseudoeucyrtis busuangaensis (Yeh & Cheng)
1998
PAN11
PAN19
429
Pseudoeucyrtis safraensis Dumitrica & Goričan
n. sp.
ORB12 Pseudogodia deweveri Carter n. sp.
SAT16 Pseudoheliodiscus aff. alpinus Kozur & Mostler
1990 sensu Whalen & Carter 2002
SAT07 Pseudoheliodiscus yaoi gr. Pessagno 1981
PPN01 Pseudopantanellium floridum Yeh 1987b
2007
Pseudopoulpus acutipodium Takemura 1986
POU01 Pseudopoulpus sp. A sensu Whalen & Carter 2002
PRL01 Pseudoristola megaglobosa Yeh 1987b
REG01 Religa globosa Whalen & Carter 2002
REG02 Religa sp. A
RBS01 Rolumbus gastili Pessagno, Whalen & Yeh 1986
RBS02 Rolumbus halseyensis Pessagno, Whalen & Yeh
1986
SAT18 Spongosaturninus bispinus (Yao) 1972
SAT19 Stauromesosaturnalis deweveri Kozur & Mostler
1990
SCP01 Stichocapsa biconica Matsuoka 1991
3407
Tetraditryma cf. praeplena Baumgartner sensu
Carter & Jakobs 1991
THT01 Thetis oblonga De Wever 1982a
THU01 Thurstonia gibsoni Whalen & Carter 1998
PSE03
430
THU04
TRX01
3409
TCA01
TRL01
TRL02
SPT01
3247
3408
UDA05
UNM01
TPS01
WNG03
WNG01
WNG04
XTL02
XTL01
ZRT01
ZRT03
COM01
Thurstonia timberensis Whalen & Carter 1998
Trexus dodgensis Whalen & Carter 1998
Triactoma jakobsae Carter 1995
Triactoma rosespitensis (Carter) 1988
Trillus elkhornensis Pessagno & Blome 1980
Trillus seidersi Pessagno & Blome 1980
Tripocyclia? tortuosa Dumitrica, Goričan &
Whalen n. sp.
Turanta morinae gr. Pessagno & Blome 1982
Tympaneides charlottensis Carter 1988
Udalia plana Whalen & Carter 1998
Unuma unicus (Yeh) 1987b
Williriedellum? ferum (Matsuoka) 1991
Wrangellium oregonense Yeh 1987a
Wrangellium thurstonense Pessagno & Whalen
1982
Wrangellium sp. A sensu Pessagno & Whalen
1982
Xiphostylus duvalensis Carter n. sp.
Xiphostylus simplus Yeh 1987b
Zartus mostleri Pessago & Blome 1980
Zartus stellatus Goričan & Matsuoka n. sp.
Zhamoidellum yehae Dumitrica n. sp.
4.2. Alphabetical listing by species
abreojosensis
aculeata
acutipodium
alpinus
altile
amurensis
anatiformis
angulosa
angulosa
angusta
angusta
anulatum
aptus
arenaense
argescens
armatus
artum
asperoensis
asuncionense
aurita
avirostrum
bajaensis
baroni
baumgartneri
beata
biconica
bicornus
bispinus
blechschmidti
bona
brevispinum
brevitubus
busuangaense
busuangaensis
callosa
calvabovis
cameroni
carlense
cavata
cavata
cerromesaensis
charlottensis
chengi
christovalensis
clara
columbiaense
conothorax
corpulenta
coyotense
crickmayi
cumshewaense
cuniculiformis
curticrassa
cylindrica
cylindricus
cylindricus
POD01
PVG01
2007
SAT16
HSU01
CHA02
JAC02
CRU21
CRU11
KAT07
PSE02
CAN12
PLE01
HSU02
ORB04
ARS03
CAN13
FAR02
CYC01
KAT08
ARS07
PRY05
PDC01
CTS06
PDC02
SCP01
KAT09
SAT18
NAP09
NAP08
PAN20
KAT12
HSU03
PSE04
ORB05
BPD13
3411
PAN14
CRU22
CRU10
NAP02
3408
CRB01
SPI03
KAT10
CAN08
NAP06
PAR13
ASP01
HCK05
PAN18
ARS06
PAR22
MCP01
4061
3001
Podocapsa abreojosensis Whalen & Carter 2002
Praeparvicingula aculeata (Carter )1988
Pseudopoulpus acutipodium Takemura 1986
Pseudoheliodiscus aff. alpinus Kozur & Mostler 1990 sensu Whalen & Carter 2002
Hsuum altile Hori & Otsuka 1989
Charlottea amurensis Whalen & Carter 1998
Anaticapitula anatiformis (De Wever) 1982a
Crucella angulosa s.l. Carter 1988
Crucella angulosa angulosa Carter 1988
Katroma angusta Yeh 1987b
Pseudoeucyrtis angusta Whalen & Carter 1998
Canoptum anulatum Pessagno & Poisson 1981
Pleesus aptus Yeh 1987b
Hsuum arenaense Whalen & Carter 2002
Beatricea? argescens (Cordey) 1998
Ares armatus De Wever 1982a
Canoptum artum Yeh 1987b
Farcus asperoensis Pessagno, Whalen & Yeh 1986
Cyclastrum asuncionense Whalen & Carter 2002
Katroma aurita Whalen & Carter 2002
Ares avirostrum Dumitrica & Matsuoka n. sp.
Praeconocaryomma bajaensis Whalen n. sp.
Beatricea? baroni Cordey 1998
Canutus baumgartneri Yeh 1987b
Crucella beata (Yeh) 1987b
Stichocapsa biconica Matsuoka 1991
Katroma bicornus De Wever 1982a
Spongosaturninus bispinus (Yao) 1972
Napora blechschmidti Dumitrica n. sp.
Napora bona Pessagno, Whalen & Yeh 1986
Pantanellium brevispinum Carter n. sp.
Katroma brevitubus Dumitrica & Goričan n. sp.
Hsuum busuangaense Yeh & Cheng 1996
Pseudoeucyrtis busuangaensis (Yeh & Cheng) 1998
Orbiculiformella callosa (Yeh) 1987b
Bipedis calvabovis De Wever 1982a
Elodium cameroni Carter 1988
Pantanellium carlense Whalen & Carter 1998
Crucella cavata s.l. Whalen & Carter 1998
Crucella cavata cavata Whalen & Carter 1998
Napora cerromesaensis Pessagno, Whalen & Yeh 1986
Tympaneides charlottensis Carter 1988
Crubus chengi Yeh 1987b
Beatricea christovalensis Whalen & Carter 1998
Katroma clara Yeh 1987b
Canoptum columbiaense Whalen & Carter 1998
Napora conothorax Carter & Dumitrica n. sp.
Paronaella corpulenta De Wever 1981b
Archaeospongoprunum coyotense Whalen & Carter 2002
Haeckelicyrtium crickmayi Carter n. sp.
Pantanellium cumshewaense Pessagno & Blome 1980
Ares cuniculiformis Dumitrica & Whalen n. sp.
Paronaella curticrassa Carter & Dumitrica n. sp.
Minocapsa cylindrica Matsuoka 1991
Ares cylindricus s.l. (Takemura) 1986
Ares cylindricus cylindricus (Takemura) 1986
431
danaense
decora
delnortensis
deweveri
deweveri
diadema
diegoi
diplocyclis
disparile
dixoni
dodgensis
duvalensis
edenshawi
elegans
elementaria
elkhornensis
elongata
emmela
erraticum
eurasiaticus
exiguum
fannini
fera
fera
ferum
flexuosus
floridum
fondrenense
formosum
funiculum
fustis
gastili
gesponsa
ghostensis
gibsoni
giganticava
gigantocornis
globosa
globosa
gongyloideum
grahamensis
graybayensis
graylockensis
gujoense
gunense
hainaensis
halseyensis
hattorii
hecatensis
helmetense
hemingense
hexagonus
hexaptera
hipolitoensis
hippaense
hotaoensis
immodica
incognita
inornatum
432
PAN11
PRY01
3222
ORB12
SAT19
BPD05
CTS08
2013
EUC09
CAN09
TRX01
XTL02
PHS02
SCP02
PVG02
TRL01
KAT17
ATA02
BAG01
DRO07
HSU04
BPD14
PAR24
PAR15
TPS01
4032
PPN01
DRO05
PHS09
BAG03
GIG01
RBS01
PTP01
2001
THU01
CRU20
PVG03
MCP02
REG01
GOR02
PAR16
NAP01
FAR04
EUC10
EUC03
CTS09
RBS02
ATT01
DRO02
BAG02
LAX06
3502
4033
DUC01
PSP03
CHA09
PRY02
ORB06
PAN19
Pantanellium danaense Pessagno & Blome 1980
Praeconocaryomma decora gr. Yeh 1987b
Bernoullius delnortensis Pessagno, Blome & Hull 1993
Pseudogodia deweveri Carter n. sp.
Stauromesosaturnalis deweveri Kozur & Mostler 1990
Bipedis diadema Whalen & Carter 1998
Canutus diegoi Whalen & Carter 2002
Parasaturnalis diplocyclis (Yao) 1972
Eucyrtidiellum disparile gr. Nagai & Mizutani 1990
Canoptum dixoni Pessagno & Whalen 1982
Trexus dodgensis Whalen & Carter 1998
Xiphostylus duvalensis Carter n. sp.
Parahsuum edenshawi (Carter) 1988
Plicaforacapsa? elegans (Matsuoka) 1991
Praeparvicingula elementaria (Carter) 1988
Trillus elkhornensis Pessagno & Blome 1980
Katroma elongata Carter n. sp.
Atalanta emmela Cordey & Carter 1996
Bagotum erraticum Pessagno & Whalen 1982
Droltus eurasiaticus Kozur & Mostler 1990
Hsuum exiguum Yeh & Cheng 1996
Bipedis fannini Carter 1988
Paronaella fera s.l. (Yeh) 1987b
Paronaella fera fera (Yeh) 1987b
Williriedellum? ferum (Matsuoka) 1991
Ares cylindricus flexuosus (Takemura) 1986
Pseudopantanellium floridum Yeh 1987b
Parahsuum fondrenense (Whalen & Carter) 1998
Parahsuum formosum (Yeh) 1987b
Bagotum funiculum Whalen & Carter 2002
Gigi fustis De Wever 1982a
Rolumbus gastili Pessagno, Whalen & Yeh 1986
Protopsium gesponsa De Wever 1981c
Acaeniotylopsis ghostensis (Carter) 1988
Thurstonia gibsoni Whalen & Carter 1998
Crucella cavata giganticava Carter n. ssp.
Praeparvicingula gigantocornis (Kishida & Hisada) 1985
Minocapsa globosa Matsuoka 1991
Religa globosa Whalen & Carter 2002
Gorgansium gongyloideum Kishida & Hisada 1985
Paronaella grahamensis Carter 1988
Napora graybayensis Pessagno, Whalen & Yeh 1986
Farcus graylockensis Pessagno, Whalen & Yeh 1986
Eucyrtidiellum gujoense (Takemura & Nakaseko) 1986
Eucyrtidiellum gunense gr. Cordey 1998
Canutus hainaensis Pessagno & Whalen 1982
Rolumbus halseyensis Pessagno, Whalen & Yeh 1986
Archaeotritrabs hattorii Dumitrica n. sp.
Droltus hecatensis Pessagno & Whalen 1982
Bagotum helmetense Pessagno & Whalen 1982
Laxtorum hemingense Whalen & Carter 1998
Hexasaturnalis hexagonus (Yao) 1972
Citriduma hexaptera (Conti & Marcucci) 1991
Ducatus hipolitoensis Whalen & Carter 2002
Perispyridium hippaense (Carter) 1988
Charlottea hotaoensis Carter n. sp.
Praeconocaryomma immodica Pessagno & Poisson 1981
Orbiculiformella incognita (Blome) 1984b
Pantanellium inornatum Pessagno & Poisson 1981
intermedicava
intermedius
izeense
jadeae
jakobsae
jamesi
japonicus
kimbroughi
kozuri
kuensis
laseekensis
laxa
liassicus
lillihornensis
lomgonensis
longibrachiata
longiconicum
longipes
lowryensis
lucidum
lupheri
lyellensis
mackenziei
macrum
majusculum
mangartense
margaritaense
matsuokai
maudense
mediocircus
medium
megaglobosa
megaglobosa
mexicoensis
mijo
minai
minoensis
mirabunda
modestum
moresbyensis
morganense
morinae
mostleri
mostleri
mulleri
munda
munitum
nagaiae
nanjoensis
nanoconica
neagui
ninstintsi
nipponica
nitidus
notabilis
obesus
oblonga
octopus
omanensis
CRU19
LAN05
2012
PDC05
3409
PAR10
BPD15
BAG04
FAR03
BRO02
DRO03
HIG01
SAT13
NTS01
ORB03
CRU12
PHS03
3149
HOM01
HSU05
HIG04
DRO06
PHS08
HAG06
HAG03
BIS04
CAN11
3195
BAG05
ORB11
3278
TPS02
PRL01
ARS04
CRU13
SUM03
TPS03
CRU14
BAG06
ARS01
GOR03
3247
PHS04
ZRT01
HSU06
ADM01
3271
EUC06
SCP03
PVG04
KAT13
KAT14
3410
CTS10
PAR17
LAN01
THT01
SAT11
JAC04
Crucella cavata intermedicava Carter n. ssp.
Lantus intermedius Carter n. sp.
Parahsuum izeense (Pessagno & Whalen) 1982
Crucella jadeae Carter & Dumitrica n. sp.
Triactoma jakobsae Carter 1995
Paronaella fera jamesi Whalen & Carter 1998
Bipedis japonicus Hori n. sp.
Bagotum kimbroughi Whalen & Carter 2002
Farcus kozuri Yeh 1987b
Broctus kuensis Pessagno & Whalen 1982
Droltus laseekensis Pessagno & Whalen 1982
Higumastra laxa Yeh 1987b
Palaeosaturnalis aff. liassicus Kozur & Mostler 1990
Noritus lillihornensis Pessagno & Whalen 1982
Orbiculiformella lomgonensis (Whalen & Carter) 1998
Crucella angulosa longibrachiata Carter n. ssp.
Parahsuum longiconicum Sashida 1988
Archaeohagiastrum longipes Baumgartner 1995
Homoeoparonaella lowryensis Whalen & Carter 2002
Hsuum lucidum Yeh 1987b
Higumastra lupheri Yeh 1987b
Droltus lyellensis Pessagno & Whalen 1982
Elodium? mackenziei Carter n. sp.
Hagiastrum macrum gr. De Wever 1981b
Hagiastrum majusculum Whalen & Carter 1998
Bistarkum mangartense Goričan, Šmuc & Baumgartner 2003
Canoptum margaritaense Whalen & Carter 1998
Hsuum matsuokai Isozaki & Matsuda 1985
Bagotum maudense Pessagno & Whalen 1982
Orbiculiformella mediocircus Dumitrica n. sp.
Hsuum medium (Takemura) 1986
Minocapsa? megaglobosa (Matsuoka) 1991
Pseudoristola megaglobosa Yeh 1987b
Ares mexicoensis Whalen & Carter 2002
Crucella mijo De Wever 1981b
Carterwhalenia minai (Whalen & Carter) 2002
Helvetocapsa minoensis (Matsuoka) 1991
Crucella mirabunda Whalen & Carter 2002
Bagotum modestum Pessagno & Whalen 1982
Ares moresbyensis Whalen & Carter 1998
Gorgansium morganense Pessagno & Blome 1980
Turanta morinae gr. Pessagno & Blome 1982
Parahsuum mostleri (Yeh) 1987b
Zartus mostleri Pessago & Blome 1980
Hsuum mulleri Pessagno & Whalen 1982
Archaeodictyomitra munda (Yeh) 1987b
Archaeohagiastrum munitum Baumgartner 1984
Eucyrtidiellum nagaiae Dumitrica, Goričan & Matsuoka n. sp.
Helvetocapsa nanjoensis (Matsuoka) 1991
Praeparvicingula nanoconica (Hori & Otsuka) 1989
Katroma neagui Pessagno & Poisson 1981 emend. De Wever 1982a
Katroma ninstintsi Carter 1988
Napora nipponica Takemura 1986
Canutus nitidus Yeh 1987b
Paronaella notabilis Whalen & Carter 2002
Lantus obesus (Yeh) 1987b
Thetis oblonga De Wever 1982a
Hexasaturnalis octopus Dumitrica & Hori n. sp.
Anaticapitula omanensis Dumitrica n. sp.
433
omanojaponicum EUC07
optimum
HSU07
oregonense
HAG01
oregonense
PSP01
oregonense
WNG03
ornata
PDC03
ovale
PHS05
parvimamma
PRY03
paulsmithi
PRU01
penderi
CHA10
pessagnoi
ELD02
phantomense
BIS02
philippinense
HSU08
plana
UDA05
plectocostatum HSU11
plicata
SCP06
plicata
SCP04
pobi
HAG02
praeobesus
LAN04
praeplena
3407
proprietatis
CHA03
protoformis
2021
pseudoerraticum BAG07
radiotuba
CIT05
ramescens
EUC04
reciproca
HOM02
reiferensis
NAP03
relica
NAP04
rennellensis
CTS15
rigidium
BIS01
robusta
ORB02
rockfishensis
CTS03
rosespitensis
TCA01
rudimentum
HAG04
ruesti
BRO03
rugosum
CAN14
saccideon
BER01
safraensis
PSE03
saginatum
BIS03
sandilandsensis FRM01
sandspitensis
JAC01
sanfilippoae
3126
sanignacioensis DRO08
sanpabloensis
ORB07
sarahae
PRY07
scammonense
CYC02
seidersi
TRL02
selwynensis
BRO01
semiplicata
SCP05
simplum
PHS01
simplus
XTL01
sinetubus
KAT16
sixi
LAN02
skedansense
PAN16
skowkonaensis 2005
snowshoensis
PAR19
spinifera
TVS01
spongase
CRU15
squama
CRU16
434
Eucyrtidiellum omanojaponicum Dumitrica, Goričan & Hori n. sp.
Hsuum optimum Carter 1988
Archaeohagiastrum oregonense (Yeh) 1987b
Perispyridium oregonense (Yeh) 1987b
Wrangellium oregonense Yeh 1987a
Pseudocrucella ornata De Wever 1981b
Parahsuum ovale Hori & Yao 1988
Praeconocaryomma parvimamma Pessagno & Poisson 1981
Protunuma paulsmithi Carter 1988
Charlottea penderi Carter n. sp.
Elodium pessagnoi Yeh & Cheng 1996
Bistarkum phantomense (Carter) 1988
Hsuum philippinense Yeh & Cheng 1996
Udalia plana Whalen & Carter 1998
Hsuum plectocostatum Carter n. sp.
Helvetocapsa plicata s.l. (Matsuoka) 1991
Helvetocapsa plicata plicata (Matsuoka) 1991
Archaeohagiastrum pobi Whalen & Carter 1998
Lantus praeobesus Carter n. sp.
Tetraditryma cf. praeplena Baumgartner sensu Carter & Jakobs 1991
Charlottea proprietatis Whalen & Carter 1998
Eospongosaturninus protoformis (Yao) 1972
Bagotum pseudoerraticum Kishida & Hisada 1985
Citriduma radiotuba De Wever 1982a
Eucyrtidiellum ramescens Cordey 1998
Homoeoparonaella reciproca Carter 1988
Napora reiferensis (Pessagno, Whalen & Yeh) 1986
Napora relica Yeh 1987b
Canutus rennellensis Carter n. sp.
Bistarkum rigidium Yeh 1987b
Orbiculiformella? robusta (Whalen & Carter) 1998
Canutus rockfishensis Pessagno & Whalen 1982
Triactoma rosespitensis (Carter) 1988
Hagiastrum rudimentum Whalen & Carter 1998
Broctus ruesti Yeh 1987b
Canoptum rugosum Pessagno & Poisson 1981
Bernoullius saccideon (Carter) 1988
Pseudoeucyrtis safraensis Dumitrica & Goričan n. sp.
Bistarkum saginatum Yeh 1987b
Foremania sandilandsensis gr. Whalen & Carter 1998
Napora sandspitensis (Pessagno, Whalen & Yeh) 1986
Pseudocrucella sanfilippoae (Pessagno) 1977a
Droltus sanignacioensis Whalen & Carter 2002
Beatricea sanpabloensis (Whalen & Carter) 2002
Praeconocaryomma sarahae Carter n. sp.
Cyclastrum scammonense Whalen & Carter 2002
Trillus seidersi Pessagno & Blome 1980
Broctus selwynensis Pessagno & Whalen 1982
Helvetocapsa plicata semiplicata (Matsuoka) 1991
Parahsuum simplum Yao 1982
Xiphostylus simplus Yeh 1987b
Katroma? sinetubus Carter n. sp.
Lantus sixi Yeh 1987b
Pantanellium skedansense Pessagno & Blome 1980
Paronaella skowkonaensis Carter 1988
Paronaella snowshoensis (Yeh) 1987b
Praeparvicingula? spinifera (Takemura) 1986
Crucella spongase De Wever 1981b
Crucella squama (Kozlova) 1971
stellatus
subovalis
sutherlandi
takemurai
teres
tetraradiatus
tetraspinus
theokaftensis
thurstonense
timberensis
tipperi
tlellensis
tortuosa
transversa
triacanthus
tripla
triquetra
trispina
trispina
trispinosa
trispinula
unicus
variabilis
veracruzense
vi
vizcainoense
whiteavesi
wilsonense
yakounensis
yaoi
yaoi
yehae
yehae
sp. A
sp. A
sp. A
sp. A
sp. A
sp. A
sp. A
sp. A
sp. A
sp. A
sp. A
sp. A
sp. B
sp. B
sp. B
sp. B
sp. C
sp. C
sp. O
sp. Y
sp. 4
ZRT03
SAT12
ARS02
ARS08
ORB08
SAT01
3089
3131
WNG01
THU04
CTS12
PCA02
SPT01
HIG03
4066
PAR20
CHA05
ORB13
ORB09
JAC05
ORB10
UNM01
PAR21
CYC03
UTD01
PHS06
PRY04
ELD03
PRY06
BPD16
SAT07
SAT15
COM01
ADM02
4008
CRU18
CHA07
CYC04
DAN02
HSU10
LAN03
PHS07
POU01
REG02
WNG04
ADM03
CHA08
HCK04
SAT14
CHA11
PDC04
CTS16
XNM01
KAT18
Zartus stellatus Goričan & Matsuoka n. sp.
Palaeosaturnalis subovalis Kozur & Mostler 1990
Ares sutherlandi Whalen & Carter 1998
Ares takemurai Dumitrica & Matsuoka n. sp.
Orbiculiformella teres (Hull) 1997
Praehexasaturnalis tetraradiatus Kozur & Mostler 1990
Hexasaturnalis tetraspinus (Yao) 1972
Crucella theokaftensis Baumgartner 1980
Wrangellium thurstonense Pessagno & Whalen 1982
Thurstonia timberensis Whalen & Carter 1998
Canutus tipperi gr. Pessagno & Whalen 1982
Praeparvicingula tlellensis Carter n. sp.
Tripocyclia? tortuosa Dumitrica, Goričan & Whalen n. sp.
Higumastra transversa Blome 1984b
Acaeniotylopsis triacanthus Kito & De Wever 1994
Paronaella tripla De Wever 1981b
Charlottea triquetra Whalen & Carter 1998
Orbiculiformella? trispina s.l. (Yeh) 1987b
Orbiculiformella? trispina trispina (Yeh) 1987b
Dumitricaella trispinosa Dumitrica n. sp.
Orbiculiformella? trispina trispinula (Carter) 1988
Unuma unicus (Yeh) 1987b
Paronaella variabilis Carter 1988
Cyclastrum veracruzense Whalen & Carter 2002
Naropa vi Hori, Whalen & Dumitrica n. sp.
Parahsuum vizcainoense Whalen & Carter 2002
Praeconocaryomma whiteavesi Carter 1988
Elodium wilsonense (Carter) 1988
Praeconocaryomma? yakounensis Carter n. sp.
Bipedis yaoi Hori n. sp.
Pseudoheliodiscus yaoi gr. Pessagno 1981
Parasaturnalis yehae Dumitrica & Hori n. sp.
Zhamoidellum yehae Dumitrica n. sp.
Archaeodictyomitra sp. A
Ares sp. A sensu Baumgartner et al. 1995a
Beatricea? sp. A
Charlottea sp. A sensu Whalen & Carter 2002
Cyclastrum sp. A
Danubea sp. A sensu Whalen & Carter 2002
Hsuum sp. A sensu Carter 1988
Lantus sp. A sensu Whalen & Carter 2002
Parahsuum? sp. A sensu Whalen & Carter 2002
Pseudopoulpus sp. A sensu Whalen & Carter 2002
Religa sp. A
Wrangellium sp. A sensu Pessagno & Whalen 1982
Archaeodictyomitra sp. B
Charlottea sp. B
Haeckelicyrtium sp. B sensu Whalen & Carter 2002
Palaeosaturnalis sp. B sensu Whalen & Carter 2002
Charlottea sp. C
Pseudocrucella sp. C sensu Carter 1988
Canutus sp. O
Charlottea? sp. Y
Katroma sp. 4
435
4.3. Listing in ascending order of species/subspecies codes
Acaeniotylopsis ghostensis (Carter) 1988
Paronaella skowkonaensis Carter 1988
Pseudopoulpus acutipodium Takemura 1986
Parahsuum izeense (Pessagno & Whalen) 1982
Parasaturnalis diplocyclis (Yao) 1972
Eospongosaturninus protoformis (Yao) 1972
Ares cylindricus cylindricus (Takemura) 1986
Hexasaturnalis tetraspinus (Yao) 1972
Pseudocrucella sanfilippoae (Pessagno) 1977a
Crucella theokaftensis Baumgartner 1980
Archaeohagiastrum longipes Baumgartner 1995
Hsuum matsuokai Isozaki & Matsuda 1985
Bernoullius delnortensis Pessagno, Blome & Hull
1993
3247
Turanta morinae gr. Pessagno & Blome 1982
3271
Archaeohagiastrum munitum Baumgartner 1984
3278
Hsuum medium (Takemura) 1986
3407
Tetraditryma cf. praeplena Baumgartner sensu
Carter & Jakobs 1991
3408
Tympaneides charlottensis Carter 1988
3409
Triactoma jakobsae Carter 1995
3410
Napora nipponica Takemura 1986
3411
Elodium cameroni Carter 1988
3502
Hexasaturnalis hexagonus (Yao) 1972
4008
Ares sp. A sensu Baumgartner et al. 1995a
4032
Ares cylindricus flexuosus (Takemura) 1986
4033
Citriduma hexaptera (Conti & Marcucci) 1991
4061
Ares cylindricus s.l. (Takemura) 1986
4066
Acaeniotylopsis triacanthus Kito & De Wever
1994
ADM01 Archaeodictyomitra munda (Yeh) 1987b
ADM02 Archaeodictyomitra sp. A
ADM03 Archaeodictyomitra sp. B
ARS01 Ares moresbyensis Whalen & Carter 1998
ARS02 Ares sutherlandi Whalen & Carter 1998
ARS03 Ares armatus De Wever 1982a
ARS04 Ares mexicoensis Whalen & Carter 2002
ARS06 Ares cuniculiformis Dumitrica & Whalen n. sp.
ARS07 Ares avirostrum Dumitrica & Matsuoka n. sp.
ARS08 Ares takemurai Dumitrica & Matsuoka n. sp.
ASP01 Archaeospongoprunum coyotense Whalen &
Carter 2002
ATA02 Atalanta emmela Cordey & Carter 1996
ATT01 Archaeotritrabs hattorii Dumitrica n. sp.
BAG01 Bagotum erraticum Pessagno & Whalen 1982
BAG02 Bagotum helmetense Pessagno & Whalen 1982
BAG03 Bagotum funiculum Whalen & Carter 2002
BAG04 Bagotum kimbroughi Whalen & Carter 2002
BAG05 Bagotum maudense Pessagno & Whalen 1982
BAG06 Bagotum modestum Pessagno & Whalen 1982
BAG07 Bagotum pseudoerraticum Kishida & Hisada
1985
BER01 Bernoullius saccideon (Carter) 1988
BIS01
Bistarkum rigidium Yeh 1987b
BIS02
Bistarkum phantomense (Carter) 1988
BIS03
Bistarkum saginatum Yeh 1987b
2001
2005
2007
2012
2013
2021
3001
3089
3126
3131
3149
3195
3222
436
BIS04
BPD05
BPD13
BPD14
BPD15
BPD16
BRO01
BRO02
BRO03
CAN08
CAN09
CAN11
CAN12
CAN13
CAN14
CHA02
CHA03
CHA05
CHA07
CHA08
CHA09
CHA10
CHA11
CIT05
COM01
CRB01
CRU10
CRU11
CRU12
CRU13
CRU14
CRU15
CRU16
CRU18
CRU19
CRU20
CRU21
CRU22
CTS03
CTS06
CTS08
CTS09
CTS10
CTS12
CTS15
CTS16
CYC01
CYC02
CYC03
CYC04
DAN02
DRO02
DRO03
DRO05
DRO06
Bistarkum mangartense Goričan, Šmuc &
Baumgartner 2003
Bipedis diadema Whalen & Carter 1998
Bipedis calvabovis De Wever 1982a
Bipedis fannini Carter 1988
Bipedis japonicus Hori n. sp.
Bipedis yaoi Hori n. sp.
Broctus selwynensis Pessagno & Whalen 1982
Broctus kuensis Pessagno & Whalen 1982
Broctus ruesti Yeh 1987b
Canoptum columbiaense Whalen & Carter 1998
Canoptum dixoni Pessagno & Whalen 1982
Canoptum margaritaense Whalen & Carter 1998
Canoptum anulatum Pessagno & Poisson 1981
Canoptum artum Yeh 1987b
Canoptum rugosum Pessagno & Poisson 1981
Charlottea amurensis Whalen & Carter 1998
Charlottea proprietatis Whalen & Carter 1998
Charlottea triquetra Whalen & Carter 1998
Charlottea sp. A sensu Whalen & Carter 2002
Charlottea sp. B
Charlottea hotaoensis Carter n. sp.
Charlottea penderi Carter n. sp.
Charlottea sp. C
Citriduma radiotuba De Wever 1982a
Zhamoidellum yehae Dumitrica n. sp.
Crubus chengi Yeh 1987b
Crucella cavata cavata Whalen & Carter 1998
Crucella angulosa angulosa Carter 1988
Crucella angulosa longibrachiata Carter n. ssp.
Crucella mijo De Wever 1981b
Crucella mirabunda Whalen & Carter 2002
Crucella spongase De Wever 1981b
Crucella squama (Kozlova) 1971
Beatricea? sp. A
Crucella cavata intermedicava Carter n. ssp.
Crucella cavata giganticava Carter n. ssp.
Crucella angulosa s.l. Carter 1988
Crucella cavata s.l. Whalen & Carter 1998
Canutus rockfishensis Pessagno & Whalen 1982
Canutus baumgartneri Yeh 1987b
Canutus diegoi Whalen & Carter 2002
Canutus hainaensis Pessagno & Whalen 1982
Canutus nitidus Yeh 1987b
Canutus tipperi gr. Pessagno & Whalen 1982
Canutus rennellensis Carter n. sp.
Canutus sp. O
Cyclastrum asuncionense Whalen & Carter 2002
Cyclastrum scammonense Whalen & Carter 2002
Cyclastrum veracruzense Whalen & Carter 2002
Cyclastrum sp. A
Danubea sp. A sensu Whalen & Carter 2002
Droltus hecatensis Pessagno & Whalen 1982
Droltus laseekensis Pessagno & Whalen 1982
Parahsuum fondrenense (Whalen & Carter) 1998
Droltus lyellensis Pessagno & Whalen 1982
DRO07
DRO08
DUC01
ELD02
ELD03
EUC03
EUC04
EUC06
EUC07
EUC09
EUC10
FAR02
FAR03
FAR04
FRM01
GIG01
GOR02
GOR03
HAG01
HAG02
HAG03
HAG04
HAG06
HCK04
HCK05
HIG01
HIG03
HIG04
HOM01
HOM02
HSU01
HSU02
HSU03
HSU04
HSU05
HSU06
HSU07
HSU08
HSU10
HSU11
JAC01
JAC02
JAC04
JAC05
KAT07
KAT08
KAT09
KAT10
Droltus eurasiaticus Kozur & Mostler 1990
Droltus sanignacioensis Whalen & Carter 2002
Ducatus hipolitoensis Whalen & Carter 2002
Elodium pessagnoi Yeh & Cheng 1996
Elodium wilsonense (Carter) 1988
Eucyrtidiellum gunense gr. Cordey 1998
Eucyrtidiellum ramescens Cordey 1998
Eucyrtidiellum nagaiae Dumitrica, Goričan &
Matsuoka n. sp.
Eucyrtidiellum omanojaponicum Dumitrica,
Goričan & Hori n. sp.
Eucyrtidiellum disparile gr. Nagai & Mizutani
1990
Eucyrtidiellum gujoense (Takemura & Nakaseko)
1986
Farcus asperoensis Pessagno, Whalen & Yeh 1986
Farcus kozuri Yeh 1987b
Farcus graylockensis Pessagno, Whalen & Yeh
1986
Foremania sandilandsensis gr. Whalen & Carter
1998
Gigi fustis De Wever 1982a
Gorgansium gongyloideum Kishida & Hisada
1985
Gorgansium morganense Pessagno & Blome
1980
Archaeohagiastrum oregonense (Yeh) 1987b
Archaeohagiastrum pobi Whalen & Carter 1998
Hagiastrum majusculum Whalen & Carter 1998
Hagiastrum rudimentum Whalen & Carter 1998
Hagiastrum macrum gr. De Wever 1981b
Haeckelicyrtium sp. B sensu Whalen & Carter
2002
Haeckelicyrtium crickmayi Carter n. sp.
Higumastra laxa Yeh 1987b
Higumastra transversa Blome 1984b
Higumastra lupheri Yeh 1987b
Homoeoparonaella lowryensis Whalen & Carter
2002
Homoeoparonaella reciproca Carter 1988
Hsuum altile Hori & Otsuka 1989
Hsuum arenaense Whalen & Carter 2002
Hsuum busuangaense Yeh & Cheng 1996
Hsuum exiguum Yeh & Cheng 1996
Hsuum lucidum Yeh 1987b
Hsuum mulleri Pessagno & Whalen 1982
Hsuum optimum Carter 1988
Hsuum philippinense Yeh & Cheng 1996
Hsuum sp. A sensu Carter 1988
Hsuum plectocostatum Carter n. sp.
Napora sandspitensis (Pessagno, Whalen & Yeh)
1986
Anaticapitula anatiformis (De Wever) 1982a
Anaticapitula omanensis Dumitrica n. sp.
Dumitricaella trispinosa Dumitrica n. sp.
Katroma angusta Yeh 1987b
Katroma aurita Whalen & Carter 2002
Katroma bicornus De Wever 1982a
Katroma clara Yeh 1987b
Katroma brevitubus Dumitrica & Goričan n. sp.
Katroma neagui Pessagno & Poisson 1981
emend. De Wever 1982a
KAT14 Katroma ninstintsi Carter 1988
KAT16 Katroma? sinetubus Carter n. sp.
KAT17 Katroma elongata Carter n. sp.
KAT18 Katroma sp. 4
LAN01 Lantus obesus (Yeh) 1987b
LAN02 Lantus sixi Yeh 1987b
LAN03 Lantus sp. A sensu Whalen & Carter 2002
LAN04 Lantus praeobesus Carter n. sp.
LAN05 Lantus intermedius Carter n. sp.
LAX06 Laxtorum hemingense Whalen & Carter 1998
MCP01 Minocapsa cylindrica Matsuoka 1991
MCP02 Minocapsa globosa Matsuoka 1991
NAP01 Napora graybayensis Pessagno, Whalen & Yeh
1986
NAP02 Napora cerromesaensis Pessagno, Whalen & Yeh
1986
NAP03 Napora reiferensis (Pessagno, Whalen & Yeh)
1986
NAP04 Napora relica Yeh 1987b
NAP06 Napora conothorax Carter & Dumitrica n. sp.
NAP08 Napora bona Pessagno, Whalen & Yeh 1986
NAP09 Napora blechschmidti Dumitrica n. sp.
NTS01 Noritus lillihornensis Pessagno & Whalen 1982
ORB02 Orbiculiformella? robusta (Whalen & Carter)
1998
ORB03 Orbiculiformella lomgonensis (Whalen & Carter)
1998
ORB04 Beatricea? argescens (Cordey) 1998
ORB05 Orbiculiformella callosa (Yeh) 1987b
ORB06 Orbiculiformella incognita (Blome) 1984b
ORB07 Beatricea sanpabloensis (Whalen & Carter) 2002
ORB08 Orbiculiformella teres (Hull) 1997
ORB09 Orbiculiformella? trispina trispina (Yeh) 1987b
ORB10 Orbiculiformella? trispina trispinula (Carter)
1988
ORB11 Orbiculiformella mediocircus Dumitrica n. sp.
ORB12 Pseudogodia deweveri Carter n. sp.
ORB13 Orbiculiformella? trispina s.l. (Yeh) 1987b
PAN11 Pantanellium danaense Pessagno & Blome 1980
PAN14 Pantanellium carlense Whalen & Carter 1998
PAN16 Pantanellium skedansense Pessagno & Blome
1980
PAN18 Pantanellium cumshewaense Pessagno & Blome
1980
PAN19 Pantanellium inornatum Pessagno & Poisson
1981
PAN20 Pantanellium brevispinum Carter n. sp.
PAR10 Paronaella fera jamesi Whalen & Carter 1998
PAR13 Paronaella corpulenta De Wever 1981b
PAR15 Paronaella fera fera (Yeh) 1987b
PAR16 Paronaella grahamensis Carter 1988
PAR17 Paronaella notabilis Whalen & Carter 2002
PAR19 Paronaella snowshoensis (Yeh) 1987b
PAR20 Paronaella tripla De Wever 1981b
PAR21 Paronaella variabilis Carter 1988
PAR22 Paronaella curticrassa Carter & Dumitrica n. sp.
KAT12
KAT13
437
PAR24
PCA02
PDC01
PDC02
PDC03
PDC04
PDC05
PHS01
PHS02
PHS03
PHS04
PHS05
PHS06
PHS07
PHS08
PHS09
PLE01
POD01
POU01
PPN01
PRL01
PRU01
PRY01
PRY02
PRY03
PRY04
PRY05
PRY06
PRY07
PSE02
PSE03
PSE04
PSP01
PSP03
PTP01
PVG01
PVG02
PVG03
PVG04
RBS01
RBS02
REG01
438
Paronaella fera s.l. (Yeh) 1987b
Praeparvicingula tlellensis Carter n. sp.
Beatricea? baroni Cordey 1998
Crucella beata (Yeh) 1987b
Pseudocrucella ornata De Wever 1981b
Pseudocrucella sp. C sensu Carter 1988
Crucella jadeae Carter & Dumitrica n. sp.
Parahsuum simplum Yao 1982
Parahsuum edenshawi (Carter) 1988
Parahsuum longiconicum Sashida 1988
Parahsuum mostleri (Yeh) 1987b
Parahsuum ovale Hori & Yao 1988
Parahsuum vizcainoense Whalen & Carter 2002
Parahsuum? sp. A sensu Whalen & Carter 2002
Elodium? mackenziei Carter n. sp.
Parahsuum formosum (Yeh) 1987b
Pleesus aptus Yeh 1987b
Podocapsa abreojosensis Whalen & Carter 2002
Pseudopoulpus sp. A sensu Whalen & Carter
2002
Pseudopantanellium floridum Yeh 1987b
Pseudoristola megaglobosa Yeh 1987b
Protunuma paulsmithi Carter 1988
Praeconocaryomma decora gr. Yeh 1987b
Praeconocaryomma immodica Pessagno &
Poisson 1981
Praeconocaryomma parvimamma Pessagno &
Poisson 1981
Praeconocaryomma whiteavesi Carter 1988
Praeconocaryomma bajaensis Whalen n. sp.
Praeconocaryomma? yakounensis Carter n. sp.
Praeconocaryomma sarahae Carter n. sp.
Pseudoeucyrtis angusta Whalen & Carter 1998
Pseudoeucyrtis safraensis Dumitrica & Goričan
n. sp.
Pseudoeucyrtis busuangaensis (Yeh & Cheng)
1998
Perispyridium oregonense (Yeh) 1987b
Perispyridium hippaense (Carter) 1988
Protopsium gesponsa De Wever 1981c
Praeparvicingula aculeata (Carter )1988
Praeparvicingula elementaria (Carter) 1988
Praeparvicingula gigantocornis (Kishida &
Hisada) 1985
Praeparvicingula nanoconica (Hori & Otsuka)
1989
Rolumbus gastili Pessagno, Whalen & Yeh 1986
Rolumbus halseyensis Pessagno, Whalen & Yeh
1986
Religa globosa Whalen & Carter 2002
REG02
SAT01
SAT07
SAT11
SAT12
SAT13
SAT14
SAT15
SAT16
SAT18
SAT19
SCP01
SCP02
SCP03
SCP04
SCP05
SCP06
SPI03
SPT01
SUM03
TCA01
THT01
THU01
THU04
TPS01
TPS02
TPS03
TRL01
TRL02
TRX01
TVS01
UDA05
UNM01
UTD01
WNG01
WNG03
WNG04
XNM01
XTL01
XTL02
ZRT01
ZRT03
Religa sp. A
Praehexasaturnalis tetraradiatus Kozur &
Mostler 1990
Pseudoheliodiscus yaoi gr. Pessagno 1981
Hexasaturnalis octopus Dumitrica & Hori n. sp.
Palaeosaturnalis subovalis Kozur & Mostler 1990
Palaeosaturnalis aff. liassicus Kozur & Mostler
1990
Palaeosaturnalis sp. B sensu Whalen & Carter
2002
Parasaturnalis yehae Dumitrica & Hori n. sp.
Pseudoheliodiscus aff. alpinus Kozur & Mostler
1990 sensu Whalen & Carter 2002
Spongosaturninus bispinus (Yao) 1972
Stauromesosaturnalis deweveri Kozur & Mostler
1990
Stichocapsa biconica Matsuoka 1991
Plicaforacapsa? elegans (Matsuoka) 1991
Helvetocapsa nanjoensis (Matsuoka) 1991
Helvetocapsa plicata plicata (Matsuoka) 1991
Helvetocapsa plicata semiplicata (Matsuoka) 1991
Helvetocapsa plicata s.l. (Matsuoka) 1991
Beatricea christovalensis Whalen & Carter 1998
Tripocyclia? tortuosa Dumitrica, Goričan &
Whalen n. sp.
Carterwhalenia minai (Whalen & Carter) 2002
Triactoma rosespitensis (Carter) 1988
Thetis oblonga De Wever 1982a
Thurstonia gibsoni Whalen & Carter 1998
Thurstonia timberensis Whalen & Carter 1998
Williriedellum? ferum (Matsuoka) 1991
Minocapsa? megaglobosa (Matsuoka) 1991
Helvetocapsa minoensis (Matsuoka) 1991
Trillus elkhornensis Pessagno & Blome 1980
Trillus seidersi Pessagno & Blome 1980
Trexus dodgensis Whalen & Carter 1998
Praeparvicingula? spinifera (Takemura) 1986
Udalia plana Whalen & Carter 1998
Unuma unicus (Yeh) 1987b
Naropa vi Hori, Whalen & Dumitrica n. sp.
Wrangellium thurstonense Pessagno & Whalen
1982
Wrangellium oregonense Yeh 1987a
Wrangellium sp. A sensu Pessagno & Whalen
1982
Charlottea? sp. Y
Xiphostylus simplus Yeh 1987b
Xiphostylus duvalensis Carter n. sp.
Zartus mostleri Pessago & Blome 1980
Zartus stellatus Goričan & Matsuoka n. sp.
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