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MUS. COMP. ZOOL.
LIBRARY.
BULLETINS 2° 873
HARVARD
OF UNIVERSIIM
AMERICAN
PALEONTOLOGY
VOL, LAI
1972-1973
IN MEMORIAM
Don L. Frizzell
1906-1972
Thomas G. Perry
1919-1972
CONTENTS OF VOLUME LXII
Bulletin No. Pages
271. The Trace Fossil Zoophycos as an Indicator of
Water Depth.
By Richard G. Osgood, Jr. and
Eugene J. Szmuc 1-22
272. Mature Modification and Dimorphism in Select-
ed Late Paleozoic Ammonoids.
By Richard Arnold Davis 23-130
273. Upper Cretaceous and Cenozoic Silicofiagel-
lates and Ebridians.
By Hsin Yi Ling 131-229
274. Siluro-Devonian Microfaunal Biostratigraphy in
Nevada.
By W. A. McClellan ........ 230-375
Plates
1-2
3-22
23-32
33-44
INDEX
No separate index is included in the volume. Each number is
indexed separately. Contents of the volume are listed in the begin-
ning of the volume.
ine bien iN
OF
AMERICAN
PALEONTOLOGY
(Founded 1895)
Vol. 62
No. Mis. comp. 290,
LIBRARY
Ake a
JUN 2 1979
; 1 FAA
HARVARD
LINN Crpy
1972
Paleontological Research Institution
Ithaca, New York 14850, U.S.A.
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BUREE TINS
OF
AMERICAN
PALEONTOLOGY
(Founded 1895)
Vol. 62
No. 271
THE TRACE FOSSIL ZOOPHYCOS AS AN
INDICATOR OF WATER DEPTH
By
RICHARD G. OsGoopn, JR. AND EUGENE J: SZMUC
May 31, 1972
Paleontological Research Institution
Ithaca, New York 14850 U. S. A.
Library of Congress Card Number: 72-79600
Printed in the United States of America
Arnold Printing Corporation
CONTENTS
Page
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Morphology of Zoophycos ............---.---00-------- ee eal iets eee Ss ie 6
racesrosstlsmaspindicesnot: vwatentc ep theese ee een nee nee 8
The Lower Mississippian section in northeastern Ohio ~............0.............. eR Dd 9
Occurrence of Zoophycos in the Mississippian of northeastern Ohio ~................. 11
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TBS Oy Cay 2a OF) 0) 0h 6 1 een Ie Pewee re D2 ta nae ee aR eR ce eek a ee tea 16
Appendix—detarled) localityaidataeses. eee. eee eee ee eee 18
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THE TRACE FOSSIL ZOOPHYCOS AS AN INDICATOR
OF WATER DEPTH
RicHARD G. OsGoop, JR.! AND EUGENE J. Szmuc?
ABSTRACT
A geographically widespread occurrence of the trace fossil Zoophycos is
described from the Lower Mississippian Cuyahoga Formation of northeastern
Ohio. Although Seilacher (1955, 1964, 1967a, 1967b) stated that Zoophycos is
most common in “intermediate water depths,’ numerous environmental indi-
cators (oscillation ripple marks, large scale cross lamination, and Lingula)
demonstrate that in northeastern Ohio Zoophycos flourished in shallow water
which was above wave base. At present the genus encompasses too wide a
range of morphologic variables. Detailed studies are needed in order to deter-
mine the limiting factor of the environment. Water depth does not appear to
provide the answer.
INTRODUCTION
As Seilacher demonstrated, one of the major values of trace
fossils is that they can be used to indicate water depth in cases
where other evidence is absent. In a series of papers, Seilacher
(1955, 1963, 1964, 1967b) established several “ichnofacies’ each
with its suite of trace fossils and each indicative of a certain en-
vironment. The Zoophycos Facies, characterized by the roostertail-
like trace fossil of the same name was proposed for an environment
which was “sublittoral to bathyl, below wave base, and without
turbidite sedimentation” (Seilacher 1964, p. 311). It should be
noted that Seilacher (1964, fig. 7) listed four deep-water deposits
containing Zoophycos, and that in 1967 he reported the first oc-
currence of Zoophycos in “deep-sea”? cores of Recent sediments
take by Lamont-Dougherty Geological Observatory. At the same
time he conceded that Zoophycos may occasionally occur in shal-
lower water “. . . due to local channeling or restriction” (Seilacher,
1967b, p. 418). In spite of these exceptions the authors conclude,
upon analysis of Seilacher’s works, that Seilacher believed that
the majority of occurrences of Zoophycos represent intermediate
water depth. Further justification for this belief will be given
below.
The purpose of this paper is to discuss a major widespread
occurrence of Zoophycos from the Lower Mississippian rocks of
northeastern Ohio. As will be demonstrated, the body fossils, pri-
mary sedimentary structures, and the regional geologic setting, all
indicate that large numbers of Zoophycos occur here in rocks
that were deposited in shallow water that was above wave base.
The validity of Zoophycos as a reliable depth indicator is thereby
somewhat in doubt.
1 Department of Geology, College of Wooster, Wooster, Ohio 44691
2 Department of Geology, Kent State University, Kent, Ohio 44242
6 BULLETIN 271
MORPHOLOGY OF ZOOPHYCOS
Diagnosis — slightly helicoid trace fossils possessing a marginal
tube; the Sprezte radiating from the center frequently give the
genus a “rooster-tail” like appearance.
Discussion — Zoophycos Massalongo, 1855 is a common form
which has been reported from nearly every continent in rocks rang-
ing from Ordovician through Tertiary in age. As Hantzschel (1962,
1965) and Simpson (1970) pointed out the genus is extremely
variable in form. It is badly in need of a monographic study, and
no attempt will be made here to discuss the genus in depth. For a
more detailed analysis of Zoophycos and its taxonomic problems the
reader is referred to Pliéka (1968, 1970), Lessertisseur (1955),
Hantzschel (1965), and Simpson (1970).
Perhaps the wide range of morphology exhibited by Zoophycos
can best be demonstrated by considering three forms figured
in the literature. Sarle (1906, fig. 1) illustrated a specimen from
the Silurian of New York State which resembles a “‘rooster-tail.”
The marginal tube is clearly visible as are the arcuate Spreite. A
more typical occurrence of Zoophycos is shown by Hantzschel
(1962, figs. 137-la, 2a). It is a flat helix with the Spreite radiating
from the apex of the cone. No marginal tube is present but this
may be due to imperfect preservation. According to Seilacher
(1967a) this form, which he had found in the Alpine Tertiary
section, is a variant of a more complex Cretaceous form, which
takes on a flat antler-like appearance (Seilacher, 1967a, p. 79 top).
Compounding the problem is the fact that many times Zoophycos
has a distinct three dimensional aspect. Moreover, as Sarle (1906,
fig. 2) pointed out several helices can be interlaced, making study
even more difficult. In its most extreme three dimensional form
the Zoophycos structure can resemble a corkscrew with Spreite,
where the vertical dimensions exceed the diameter of the helix.
These forms are assigned by most authors to Daedalus Rouault,
1850.3
3Simpson (1970) was well aware of the morphological variation included
in Zoophycos and made some attempt to limit it by restricting the circular forms
to Spirophyton Hall, 1963. However, as one can see, and as Simpson himself
admitted, the Zoophycos still contains a wide variety of morphologic expres-
sions (see Simpson, 1970, fig. 1).
‘TRACE Fossi1Ls: OsGoop & SZMUC 7
The Mississippian forms from Ohio contribute little to the
general knowledge of the morphology or ethology exhibited by
Zoophycos. They closely resemble those specimens illustrated by
Hantzschel (1962, fig. 137-la) and Simpson (1970, fig. Ib).
They occur in gray, micaceous siltstones and their overall pat-
tern is somewhat masked by their three-dimensional aspect. One
specimen (PI. 1, fig. 2) retains a small fragment of the marginal
tube. The tube is ovoid in cross-section and measures 5 mm by
3 mm. Because the species of Zoophycos are so poorly defined,
no attempt at specific identification will be made here.
Interpretation — The interpretations given to Zoophycos are
as varied as the morphology of the genus itself. Several earlier
authors (e.g. Rouault, 7m Lebesconte, 1884) believed that it was the
impression of marine algae. It has also been assigned to the Porifera
by Lebesconte (1887) and to the Alcyonaria by Lucas (1938).
Other workers regarded it as inorganic, caused by either current
activity (Nathorst, 1886), or gas bubbles (Korn, 1929).
Today it is generally agreed that Zoophycos represents the
feeding structure of an unknown soft-bodied wormlike organism
(Hantzschel 1962, 1965; Seilacher 1967a). A notable excepton is
Pli¢ka (1968, 1970) who, working with Tertiary material from
the Carpathians, described the arcuate Spreite as the impression of
the prostomia of sabellid annelids.
The present authors accept the trace fossil assignment although
they are unable to comment on Pli¢ka’s material. The three di-
mensional preservation of Zoophycos, as well as the lack of any
microstructure, rules out a vegetable origin. Moreover, the arcuate
Spreite are a variant of a feeding method frequently observed in
trace fossils. Phycodes and Rhizocorallium could be cited as ex-
amples. Basically each arc represents the course of the organism as
it moved along, ingesting the sediment. When the elastic limits of
the body were reached the animal contracted and proceeded to
excavate a new burrow directly adjacent to the old one. Seilacher
(1967a) postulated that the antler-shaped Zoophycos from the
Cretaceous are actually a less efficient variant of the more compact
Tertiary forms. In the former the area between the horns is left
unexploited whereas this does not occur in the compact forms.
Because the burrows of Zoophycos are normally three dimensional
8 BULLETIN 271
it is probable that they were excavated below the depositional inter-
face.
The above description of the origin of the burrow is greatly
simplified. The only detailed attempts to describe the habits of
the organism are those of Bischoff (1968) and Simpson (1970).
In summary, Zoophycos is variable in overall appearance but
all forms assigned to the genus possess the arcuate Spreite. Because
of its variability it must be regarded as a form genus. It represents
a feeding burrow although the actual mechanics of formations of
the burrow are still not clear for all members of the genus.
TRACE FOSSILS AS INDICES OF WATER DEPTH
Seilacher (1955) compared the trace fossils from five sections
of diverse geologic ages. He found some sections (e.g. Lower Cam-
brian of the Salt Range of Pakistan) were characterized by shallow
resting traces (Cubichnia of Seilacher, 1953a) which probably
served as places of temporary concealment. He interpreted these as
indicative of a shallow-water environment well within the euphotic
zone, an area where predators would be most active. Conversely
other sections such as the Tertiary Alpine Flysch contained no
Cubichnia but numerous grazing traces (Pascichnia of Seilacher
1953a) . Seilacher interpreted Pascichnia as two-dimensional feeding
trace analogous to strip mining, where the organism attempts to
cover a large area with a minimum amount of effort. This can lead
to unusual geometric patterns such as the tight spiral of Helmin-
thoida.
Observations of Recent sediments have served largely to con-
firm Seilacher’s hypotheses. It is difficult to observe Cubichnia in
the process of formation since the organism quickly covers itself
with sediment. However, Recent stelleroids and ophiuroids are
known to burrow for protection (Seilacher, 1953b), and it is not
unreasonable to assume that Rusophycus, which is abundant in
lower Paleozoic sediments represents a burrow of trilobites. While
Pascichnia remain unreported in photographs taken within the
4For a more detailed discussion of Seilacher’s ichnofacies see Osgood (1970,
pp. 399-404).
TRACE FossiLs: OsGoop & SZMUC 9
euphotic zone their presence has been confirmed by photographs
taken at great depth (see Seilacher 1967b, pl. II).
In 1964 Seilacher expanded his analysis to include 42 sections
of varying geologic age and locality. He formally named the suite
of trace fossils dominated by Pascichnia as the “Nereites Facies”
while the Cubichnia suite was entitled the “Cruziana Facies.” In
addition he proposed the “Zoophycos Facies’ at this time. Unlike
the Nereztes Facies and Cruziana Facies which normally contained
several different trace fossils, Zoophycos was commonly the sole
trace fossil. Seilacher (1964, 1967b) cited several examples from
the geologic record to support his views. Moreover, in the Ordo-
vician of Iraq there is a vertical gradation through the Nereites-
Zoophycos-Cruziana Facies.
In subsequent papers Seilacher (1963, 1967b) enlarged the
number of facies to six. These additional facies, which need not
concern us here, are indicative of shallow water or special environ-
mental conditions.
The authors of this paper take no issue with Seilacher’s inter-
pretation of the environmental significance of the Cruziana and
Nereites Facies. They are supported by both well-reasoned argu-
ments and Recent observations. The Zoophycos Facies rests on
weaker evidence. Why should this particular three-dimensional
feeding burrow be confined to “intermediate” depth water and
why are there virtually no other trace fossils occurring with it?
THE LOWER MISSISSIPPIAN SECTION IN
NORTHEASTERN OHIO
The Mississippian stratigraphic units in northern Ohio com-
prise a composite thickness of more than 300 meters of fine-to-
coarse-grained clastics that were deposited in the northwestern part
of the Appalachian marginal basin. They include shallow-water
marine, transitional, and to a lesser degree nonmarine sediments.
The stratigraphic sequence shown in Text-figure | is conform-
able except for an erosional break of considerable relief at the base
of the Berea Formation and a minor unconformity or reworked
surface at the Cuyahoga-Logan contact.
Four major types of lithologic groups are shown in Text-figure
1: 1. Conglomerate, 2. Sandstone and pebbly sandstone, 3. Gray
shale and siltstone, 4. Black and gray shale and siltstone.
10 BULLETIN 271
FORMATION MEMBER DEPOSITIONAL ENVIRONMENT
REDISTRIBUTED DELTAIC SAND-SHEET
BLACK HAND DELTAIC BAR-FINGER
= WOOSTER PRODELTAIC
ARMSTRONG PRODELTAIC AND SHALLOW OFFSHORE MARINE
RITTMAN DELTAIC BAR-FINGER
MEADVILLE PRODELTAIC AND SHALLOW OFFSHORE MARINE
—<
co
=>)
Se
—<
S=
=>
<5)
"STRONGSVILLE STAGNANT SHALLOW SEA
BEDS"
SHARPSVILLE SHALLOW OFFSHORE MARINE
ORANGEVILLE SHALLOW OFFSHORE MARINE
SUNBURY STAGNANT SHALLOW SEA
Eeearr| DELTAIC AND NON-MARINE CHANNEL
SILTSTONE AND
GRAY SHALE
@ ZooPHYCoS IN SILTSTONE
SANDSTONE BLACK SHALE
V VERTICAL LINGULA
Text-figure 1. Composite section of Lower Mississippian stratigraphic
units in northern Ohio and their deduced depositional environments. Vertical
dimensions of the units are not drawn to scale.
CONGLOMERATE = GRAY SHALE
TRACE FossiLts: OsGoop & SZMUC a
The Berne Member of the Logan Formation is a thin, wide-
spread fine-pebble conglomerate which contains a moderately abun-
dant marine invertebrate fauna. Hyde (1915) interpreted the con-
glomerate as a wave-reworked “lag concentrate” of pebbles derived
from the underlying pebbily sandstones of the Black Hand Member.
The sandstones are predominately deltaic in origin. The Berea
Sandstone is a channel-deltaic unit composed of nonmarine, transi-
tional, and marine strata (Pepper, et al., 1954). The Rittman
Member and Black Hand Member are regarded as delta-complex
deposits composed primarily of bar-finger sands which grade later-
ally into finer clastics of the Wooster Member and Meadville Mem-
ber (Barclay, 1968; Szmuc, 1970) .
The gray shale beds and the flaggy siltstone-shale beds of the
Orangeville Member, Sharpsville Member, and Wooster Member,
and the upper portion of the Meadville Member are composed of
prodeltaic and shallow-water offshore marine sediments (Barclay,
1968; Szmuc, 1970). A sparse to abundant marine fauna composed
of brachiopods, bryozoans, corals, and many other invertebrates
is dispersed irregularly throughout these units.
The black shales of the Sunbury Member and the siltstones
and black and gray shales of the basal portion of the Meadville
Member (here designated informally as the “Strongsville Beds’’)
contain a depauperate biota composed of conodonts, plant debris,
and inarticulate brachiopods. These strata were deposited in re-
stricted, stagnant, and deoxygenated shallow seas.
The Strongsville Beds are noteworthy because they typically
display one to six meters of gray and black shales intercalated be-
tween two areally extensive siltstone beds which at many expos-
ures contain profuse numbers of Zoophycos on their upper bedding
surfaces. One or both of the Zoophycos-bearing siltstones are prom-
inently displayed at Brandywine Creek, Cuyahoga Falls, Strongs-
ville, and Valley City (Sections 1, 2, 3, and 4 of this report).
OCCURRENCE OF ZOOPHYCOS IN THE MISSISSIPPIAN
OF NORTHEASTERN OHIO
The main objective of this study was to locate Zoophycos in the
field and to search for sedimentary structures, body fossils and
12 BULLETIN 271
trace fossils which would provide some evidence of water depth.
Numerous exposures were sampled and described in detail in a
five-county area, including the seven representative localities shown
in Text-figure 2. The results of the study are presented below.
N
Lake Erie
Yi
93 CUYAHOGA |
0 10 20 ?
[fs biaouea ee cient [ee a A sl]
miles A o}
e SUMMIT
MEDINA hee
plese
Q7
ASHLAND
Text-figure 2. Map of five counties in northeastern Ohio showing out-
crops of Lower Mississippian strata at selected localites. 1. Brandywine Creek.
2. Gorge Park, Cuyahoga Falls. 3. Strongsville. 4. Valley City. 5. Killbuck
Creek. 6. Wooster Quarry. 7. Buck Creek. See Appendix at end of report for
detailed locality data.
TRACE Fossi_s: OsGoop & SzMuUC 13
A) Stratigraphic distribution of Zoophycos
Zoophycos is confined to certain siltstones of the Meadville
Member and Armstrong Member. The sandstones, conglomerates,
and the gray and black shales are devoid apparently of Zoophycos.
B) Abundance of Zoophycos
The density of distribution of Zoophycos in the Strongsville
Beds is comparable to certain beds in the Mississippian Borden
Group of Kentucky, where their intertwined patterns locally cover
entire bedding planes. Although Zoophycos is more sparsely repre-
sented on bedding planes in the Armstrong and upper part of the
Meadville, its common presence clearly demonstrates that it is not
a chance occurrence. It was usually possible to locate three to four
specimens on a surface of four square meters.
The authors were able to confirm Seilacher’s (1964) observa-
tion that only rarely does Zoophycos occur with other trace fossils.
The only associated form is Palaeophycus sp. (as defined by Osgood,
1970). This is a common trace fossil and not diagnostic of any
particular environment.
C) Associated invertebrate and plant fossils
Few body fossils were found in association with Zoophycos.
This is not too surprising because the section is predominantly
clastic, and fossil occurrences within this part of the Mississippian
tend to be localized. Fragmented pelecypod, sponge, crinoid, and
articulate brachiopod remains were found from a few centimeters
to 20 meters (vertical distance) of Zoophycos, but not within the
same or adjacent beds. There are, however, two significant excep-
tions: 1) At the Strongsville locality (Section 3) specimens of
Lingula cf. melie Hall were found in situ within the Zoophycos
beds (PI. 2, fig. 1). As most of them assume the life position it is
unlikely that they were washed into the area. It is well known that
Recent Lingula prefers a shallow-water environment and is the
only living brachiopod which can sustain itself for short periods of
time in brackish waters (Rudwick, 1965). According to Hatai
(1940) Recent Lingula extends from the low-water line to about
20 or 23 fathoms, although one specimen of Lingula unguis was
recorded from a depth of about 50 fathoms near Seto, Japan.
14 BULLETIN 271
2) At the Valley City locality (Section 4) a black shale two to
three meters thick is in contact with the upper Zoophycos siltstone
of the Strongsville beds. Within the shale are a few thin (2 to 3 mm
thick) beds of cross-laminated siltstone. This shale is also present
at the Strongsville locality and apparently represents a local basin.
Its significance for this study is that it contains abundant carbon-
ized cordaite fragments, some several centimeters long. While this
is not indicative of any particular water depth, the abundance of
the plant fragments would seem to demonstrate a near-shore occur-
rence.
D) Sedimentary features associated with Zoophycos
Several lines of evidence point to abundant current or wave
activity.
1) Many of the siltstone beds in the Meadville Member and
Armstrong Member pinch and swell and some are abruptly lenticu-
lar. Cross-laminations are common and many are large-scale sets
which range from 3 to 5 meters in lateral extent and display dips of
5 to 15 degrees. While we are unable to assign the large-scale struc-
tures to a particular depositional environment (e.g. point bar) it
seems unlikely that they originated below wave-base.
2) Oscillation ripple marks are abundant at the Killbuck
Creek locality (Section 5) and occur within 5 cm (vertically) of
Zoophycos. ‘They have an amplitude of 1 cm and a wave length of
7 to 10 cm., and many grade laterally into interference ripple
marks. It is not uncommon to find three or four ripple-marked
beds within a thickness of 20 cm. While the authors realize that
such ripple marks may be produced by various mechanisms (see
McKee, 1965) it is difficult to visualize the origin of such large
numbers below wave-base. It is worthy of note that Seilacher
(1967b, fig. 2) placed Zoophycos below the occurrence of oscillation
ripple marks.
E) Regional geological setting
It is instructive to look at the environmental setting of the
formations both underlying and overlying the Cuyahoga. In north-
eastern Ohio the Berea Sandstone occurs some 45 meters below the
Zoophycos beds (Szmuc, 1970, p. 43). Pepper, et al. (1954) demon-
strated convincingly that the Berea and the subjacent Bedford
TRACE FossiLs: OsGoop & SZMUC 15
Formation represent a deltaic, prodeltaic, and channel-fill environ-
ment. The reader is referred to their paper for a more complete
discussion.
Although the Logan Formation, which lies some 35 meters
above Zoophycos, has not been thoroughly studied by the writers,
environmental indicators are abundant. The Quarry of the Wooster
Medal Brick and Tile Company at Wooster (Section 6) contains
one of the best exposures of the upper Cuyahoga Formation and
lower Logan Formation in northeastern Ohio. In this exposure
interference ripplemarks, ripple drift, and small scale channel
cross-lamination are common. Rare casts of mud cracks have also
been found. The Berne Conglomerate in the basal portion of the
Logan consists of quartz pebbles in a sandy matrix containing
chonetid brachiopods.
The stratigraphic position of the Cuyahoga beds between del-
taic and delta-derived sediments is a compelling argument for a
shallow-water habitat of the Ohio Zoophycos. The argument is
strongly enhanced by the environmental setting of the Meadville,
Armstrong, Rittman, and Black Hand Members. The deduced depo-
sitional histories (see Text-fig. 1) of these units are based not only
on the criteria presented in this paper, but also on their body-fossil
content, geometry, and stratigraphic relations with contiguous units
such as the largely nonmarine Pocono Sandstone of Pennsylvania
(Hyde, 1915; Barclay, 1968; Szmuc, 1970). Environmental interpre-
tations on these bases indicate that the Zoophycos in the Armstrong
Member is sandwiched between the Rittman and Black Hand bar-
finger sands, the stratigraphic separation of which is about 35
meters. Moreover, the Zoophycos-bearing beds in the Armstrong
and in the upper part of the Meadville are intimately associated
with off-shore marine and prodeltaic deposits.
There is nothing in the Ohio Mississippian section which sug-
gests any significant deepening of the epeiric sea during Cuyahoga
sedimentation. On the contrary, all available evidence indicates
that Zoophycos lived in shallow-water, offshore environments.
CONCLUSIONS AND PROBLEMS
1) In Ohio Zoophycos is closely associated with shallow-water
environments, as is amply attested by our interpretation of the
Strongsville Beds.
16 BULLETIN 27]
2) Seilacher’s Zoophycos Facies apparently must be broadened
or expanded from an equivocal assignment of “intermediate depth”
to include deposits which form well above wave base and proximate
to the outer margins of bar-finger sands. This opinion is strongly
indicated by our interpretation of the Armstrong beds and upper
Meadville beds and related units of the Cuyahoga Formation.
3) It seems likely that water depth, which in turn reflects many
physical factors (temperature, light), is not the sole factor in deter-
mining the distribution of Zoophycos. This conclusion is buttressed
by the presence of this genus in two different shallow-water eco-
logical niches, the Strongsville and the Armstrong-Meadville, as
well as by Seilacher’s accounts of Zoophycos in relatively deep-water
deposits.
4) ‘Trace-fossil workers are now faced with the task of con-
ducting a closer examination of this genus. Several questions re-
main unanswered, e.g., why is Zoophycos usually not associated with
other trace fossils, how much morphologic variation is to be per-
mitted to exist within the genus, and finally what are the specific
physical, chemical, and biotic factors which control its distribution?
BIBLIOGRAPHY
Barclay, C. C.
1968. Sedimentary structures and depositional history of the coarse-
clastic rocks of the Cuyahoga Formation in northern Ohio. Kent
State Univ. unpubl. master’s thesis, 108 pp.
Bischoff, B.
1968. Zoophycos, a polychaete annelid, Eocene of Greece. Jour. Paleont.,
vol. 42, pp. 1439-1443, 2 pls.
Fale J.
1863. Observations upon some spiralgrowing fucoidal remains of the
Paleozoic rocks of New York. 16th Ann. Rept. New York State
Cabinet, pp. 76-83.
Hantzschel, W.
1962. Trace-fossils and Problematica. In Treatise on Invertebrate Pale-
ont., Moore, R. C. (ed)., pt. W., Miscellanea, Lawrence, Kansas,
University of Kansas Press, pp. 177-245, figs. 109-149.
1965. Vestigia invertebratorum et Problematica. Fossilium Catalogus, 1.
Animalia, s’Gravenhage, Junk, 142 pp.
Hatai, K.
1940. The Cenozoic Brachiopoda of Japan. Sci. Rep. Tohuku Imp. Univ.,
ser. 2, vol. 20, 413 pp.
Hyde, J. E.
1915. Stratigraphy of the Waverly formations of central and southern
Ohio. Jour. Geol., vol. 23, pp. 655-682; 757-779.
TRACE Fossits: OsGoop & SZMUC 17
Korn, H.
1929. Fossile Gasblasenbahnen aus dem Thiiringer Palaeozokum. Eine
neue Deutung von Dictyodora, Z. Naturwiss., Bd. 89, pp. 25-46,
3 figs.
Lebesconte, P.
1883. Presentation a la société des oeuvres posthumes de Marie Rouault
par P. Lebesconte, suivies d’une note sur les Cruziana et Rhyso-
phycus. Soc. Geol. France, Bull., ser. 3, vol. 11, pp. 466-472.
1887. Constitution générale du Massif breton comparée a celle du
Finisterre. Soc. Geol. France, Bull., ser. 3, vol. 14, pp. 776-820, 2 pls.
Lessertisseur, J.
1955. Traces fossiles d’activité animale et leur significance paléobio-
logique. Soc. Geol. France, Mem. 74, 150 pp., 68 figs., 11 pls.
Lucas, G.
1938. Les Cancellophycus du Jurassique sont des Alcyonaires. Compte
Rendues Acad. Sci., vol. 206, p. 1914.
Massalongo, A.
1855. Zoophycos, novum genus plantorum fossilium. Pp, 52, 3 pls. Verona.
McKee, E. D.
1965. Experiments on ripple lamination. In Middleton, G. V., Primary
sedimentary structures and their Hydrodynamic Interpretation.
Soc. Econ. Paleont. and Minerol., Spec. Pub., No. 12, pp. 66-84,
6 figs.
Nathorst, A. G.
1886. Nouvelles observations sur les traces d’animaux et autres phéno-
menes d’origine purement mécanique décrits comme “Algues fos-
siles.’ Kgl. Svenska Vetensk. Akad. Handl., vol. 21, No. 14, 58
pp., 14 figs. 5 pls.
Osgood, R. G. Jr.
1970. Trace fossils of the Cincinnati area. Palaeont. Americana, vol. 6,
No. 41, pp. 281-444, 29 figs., 27 pls.
Pepper, J. F., Dewitt, W., and Demarest, D. F.
1954. Geology of the Bedford Shale and Berea Sandstone in the Ap-
palachian Basin. United States Geol. Sur., Prof. Paper 259, 109
pp., 61 figs., 13 pls.
Plicka, M.
1968. Zoophycos and a proposed classification of sabellid worms. Jour.
Paleont., vol 42, pp. 836-849, 2 pls.
1970. Zoophycos and similar fossils. In Crimes, T. P. and Harper, J. C.,
Trace Fossils. Geological Journal (Liverpool), Special Issue, No.
3, pp. 361-71, 4 figs., 2 pls.
Rouvault, M.
1883. See Lebesconte, P.
Rudwick, M. J. S.
1965. Ecology and paleoecology. In Treatise on Invertebrate Paleon-
tology. Moore, R. C. (ed.), pt. H, vol. 1, Brachiopoda, Lawrence,
Kansas, University of Kansas Press, pp. 199-214.
Sarle, C. J.
1906. Arthrophycus and Daedalus of burrow origin. Rochester Acad.
Sci., Proc., vol. 4, pp. 203-210, 4 figs.
Seilacher, A.
1953a. Uber die Methoden der Palichnologie, Studien zur Palichnologie,
No. 1. Neus Jahrb. Geologie u. Palaont., Abh., Bd. 96, pp. 421-
452, 10 figs., 1 pl.
18 BULLETIN 271
1953b. Uber die Methoden der Palichnologie, Die fossilien Ruhespuren
(Cubichnia). Neues Jahrb. Geologie u. Palaont., Abh., Bd. 98, pp.
87-124, 5 figs., 5 pls.
1955. Spuren und Lebensweise der Trilobiten; Spuren und Fazies im
Unterkambrium. In Schindewolf, O. H., and Seilacher, A., Beitrdge
zur Kenntnis des Kambriums in der Salt Range (Pakistan). Akad.
Wiss. Lit. Mainz. math.-nat. KI., Abh., No. 10, pp. 86-143, figs.
1-11, pls. 16-27.
1963. Lebenssuren und Salinitatsfazies. Fortschr. Geol. Rheinland und
Westfalia, Bd. 10, pp. 81-94, 7 figs.
1964. Biogenic sedimentary structures. In Imbrie, J., and Newell, N.
Approaches to Paleoecology, New York, John Wiley and Sons,
pp. 296-316, 7 figs.
1967a. Fossil behavior. Scientific American, vol. 217, pp. 72-80.
1967b. Bathymetry of trace fossils. Marine Geology, vol. 5, pp. 413-429, 4
figs., 2 pls.
Simpson, S.
1970. Notes on Zoophycos and Spirophyton. In Crimes, T. P. and Harper,
J. C., Trace Fossils. Geological Journal (Liverpool), Special
Issue, No. 3, pp. 505-515, 4 figs.
Szmuc, E.
1970. The Mississippian System. In Banks, P., and Feldman, R., Guide
to the Geology of Northeastern Ohio. Northern Ohio Geol. Soc.,
pp. 23-67, 31 figs., 3 pls.
APPENDIX — DETAILED LOCALITY DATA
1. Brandywine Creck locality. Top of section is at former State Route 8 bridge
at Little York, 4.2 miles south of Northfield Center; Northfield Township,
Summit County, Northfield 7.5 minute Quadrangle.
2. Gorge Park locality. City Park in Cuyahoga Falls, outcrop is in a tributary
valley on north bank of the Cuyahoga River 0.8 miles east of State Route
8 bridge; Summit County, Akron East and Akron West 7.5 minute Quad-
rangles.
3. Strongsville locality. Baker Creek, 0.1 mile west of junction of State Routes
82 and 237 and 1.25 miles west of Strongsville. Outcrop is some 300 meters
south of the bridge over route 82; Strongsville Township, Cuyahoga County,
Berea 7.5 minute Quadrangle.
4. Valley City locality. West branch of Rocky River where crossed by State
Route 252 less than one mile south of intersection of State Routes 252 and
303; Medina County, West View 7.5 minute Quadrangle.
5. Killbuck Creek locality. Outcrop begins some 200 meters downstream from
where an unnamed tributary of Killbuck Creek is crossed by Wayne County
Route 36, 2.5 miles north of village of Congress; Wayne County, West
Salem 7.5 minute Quadrangle.
6. Wooster Quarry locality. Shale pit (abandoned 1970) of Wooster Medal
Brick Company on north side of Wayne County Route 22, 1.7 miles south
of intersection of County Routes 22 and 106, Wooster: Wayne County,
Wooster 7.5 minute Quadrangle.
Buck Creek locality. Buck Creek where crossed by State Route 60 at Here-
ford; Ruggles Township, Ashland County, New London 7.5 minute Quad-
rangle.
“I
PLATES
20 BULLETIN 271
EXPLANATION OF PLATE 1
Figure Page
1. Zoophycos spr*.2...24 5 2 eee ee ee eee 6
A large incomplete specimen from the Brandywine Creek locality
(#1), UCGM 40644, x 0.4. Respository of figured specimens,
University of Cincinnati Geology Museum, Cincinnati, Ohio,
45221.
2° (LOOBNY COS SDi oy ee cece ost ee Re ea 6
A more typical “rooster-tail’” form with a small portion of the
marginal tube preserved. Buck Creek locality (#7), UCGM
40645, X 0.5.
BULL. AMER. PALEONT., VOL. 62 PrAte; 1
BULL. AMER. PALEONT., VOL. 62 PLATE 2
TRACE FossiLs: OsGoop & SZMUC Dal
EXPLANATION OF PLATE 2
Figure Page
ie Eingularct meliear Hallgren et cette tacts tinct nw cette ae 13
Preserved in situ. Although the bedding planes are not visible
the specimen was taken directly from the outcrop thus assur-
ing correct orientation. Strongsville locality (#3), UCGM
40646, X 1.2.
2. Large scale cross stratification with a dip of 5°-15° 14
The lateral distance shown in the photograph is approximately
10 meters. Killbuck Creek locality (#5).
INDEX
Note: Light face figures refer to pages. Bold face figures refer to plates.
Armstrong Member ....
Berea Formation
Berne Member
Bischoti baie.
Black Hand Member ..
Borden Group
Brandywine Creek,
section #1
Buck Creek,
Section 357)
Cruziana Facies
Cubichnia
Cuyahoga Falls,
section #2
Cuyahoga Formation ..
Daedalus
Hantzschel, W. ............
ivi Cay irae ear 2 ee
‘Ichnofacies”
Killbuck Creek,
SeChlOn7s5iee ee
PAN GULA ees ;
Logan Formation
11, 18, 20
18, 20
11s
5, 9, 14-16
14, 18, 21
13, 21
OAS
Meadville Member ...... 11, 13-16
cf. melie, Lingula ....2 13
Nereites Facies ............ 9
Orangeville Member .. alg
Pascichnia® 22.0. 8
PlickageMisy wees ee. 6,7
Rittman Member ......... Hale 15)
Sarle..\C. fet.) tne taken 6
Sellacher= Ay oe). 5-9, 13, 16
Sharpsville Member .... 11
SIMPSONS eee ee 6-8
Spirophyton- 2. 6
“Strongsville Beds” .... 11, 14, 15
Strongsville,
Secuiony Grant IG, 216}, alts}, 2A
SZMUCH tein Wy a5
Valley City,
section #4 .............. 11, 14,18
Wooster Member ...... Wal
Wooster Quarry,
Sections 09 ee 15, 18
Zoophycos Facies ....... 5, 9, 16
Zoophycos Sp ............. 1 6
ig
MUS. COMP. ZOOL.
BULIERING «|
OF SEP 23 1972
RVARD
AMERIG AN Haven Ge
PALEONTOLOGY
(Founded 1895)
Vol. 62
No. 272
MATURE MODIFICATION AND DIMORPHISM
IN SELECTED LATE PALEOZOIC AMMONOIDS
By
RicHarp ARNOLD Davis
1972
Paleontological Research Institution
Ithaca, New York 14850 U. S. A.
PALEONTOLOGICAL RESEARCH INSTITUTION
1971 - 72
PRESIDENT (ince ee eos ees aa es a OA a SCF Sos Be ee Ee a RR DANIEL B. SAss
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SRERE TARY. recs ts eas 5 ye OE IE Sieh 2S bel CEN REBECCA §S. HARRIS
DIRECTOR: WEREASURER( (cnt ost se a ctoeeee e eee ene eeees KATHERINE V. W. PALMER
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BULLETINS
OF
AMERICAN
PAL EOnrOlLoG y
(Founded 1895)
Vol. 62
No. 272
MATURE MODIFICATION AND DIMORPHISM
IN SELECTED LATE PALEOZOIC AMMONOIDS
By
RicHarp ARNOLD Davis
September 13, 1972
Paleontological Research Institution
Ithaca, New York 14850 U. S. A.
Library of Congress Card Number: 72-83384
Printed in the United States of America
Arnold Printing Corporation
CONTENTS
Page
PANS trial Cty coscceet eae Sess EE Ee rele prec oe he, MER ene core Ee see tia hohe eiten ta scasseeedepees 27
Tru OC UGE Omnis ose cee 2 cee cc oP cc SE LE TA aN ci otek ae 27
ates Ontorencticy shel liVloditicatio myyse eset see see aa ee 30
MD TTAO Tp AAS ITN eee ere. et eer, ee 2 SA Eee eee ee eee Ee 36
IDKsevereyfoxahi ai = 1 Dee ey = pment Oe i ee ene ee ee es ee ere 43
Superfamily ADRIANITACEAE ................. aiter Sey vane iste) ene ri lass 43
Biarrially ge A RD AINE DAU ea 2 ee ee en eee 43
SuperfamilyseA\G Av nl ChRAGDACEA Tat 61
amily eG Adri BRAC TID IA Bian eee tet Ree ree eee eee 61
Supertamily@Y CLOLOBACE Ale 65
Beer ena ivan © VAC Ve OL © BD AE eo acs es ne te eee rere ees 66
Rarmilyss Valo RIO CE RAG AI ee ee 76
Subfamily, VND RTO CE RAGIINAR ooo. oct eke ee re ee aay 76
Supertaniilya ViEAReACISET @ NileneAl© By ANB eee eee eee 81
1p ered KoPee ING (Ped Se BG 8 ICG) Hef BD D5 pe ae ek a este ee eae Te 81
Subfamily. MAR AEIONITINAB Ss ses 6 og ee Sirs est se estes 81
Subfamily KeA GAM 1 DIN By oe oe: oot eee ee ee 88
INGRETENCES ee 2eeeese FIORE Be Geel Ae oom I eg he eh see es 2h asec He eee ee 90
LPAI ing SPN Ow AONE ee ie NUN a2 Oe Seen ae ts De ee 101
Index
i ' i aly
) ery
Ce
PN wag cult i i
bie
AE UA chi i if . CoMabikol 1 Tacs
m Agusta |
iJ LA Time Ant a at
, an
‘ it ae +f rehe # ; oon +, e
MATURE MODIFICATION AND
DIMORPHISM IN SELECTED
LATE PALEOZOIC AMMONOIDS!
Ricuarp ArNoLp Davis,
University of Cincinnati
ABSTRACT
Many ammonoid cephalopod taxa display late ontogenetic modifications
of the shell. These modifications include any number of the following: change
in coiling, modification of whorl section, change in ornament, development of
an apertural constriction, development of an apertural shell thickening, forma-
tion of a characteristic ultimate peristome (with lappets or a rostrum, for ex-
ample), changes in shell deposition, progressive approximation of the ultimate
septa, progressive simplification of the ultimate few sutures, thickening of the
last septum, and development of muscle scars. Such modifications are related to
the attainment of sexual maturity. The ecologic and physiologic implications of
these mature modifications are uncertain.
Long ago, sexual dimorphism was suggested to account for the widespread
existence of pairs of morphologically distinct mature ammonoids which are
virtually identical as juveniles. This explanation is most probably valid.
The following genera of the family Adrianitidae display mature modifi-
cations: Adrianites, Crimites, Epadrianites, Hoffmannia, Neocrimites, Paler-
mites, Pseudagathiceras, Sizilites, and Texoceras. In specimens of all of these,
changes in coiling and an apertural constriction are present. Changes in orna-
ment are present in Neocrimites, Pseudagathiceras, Sizilites, and possibly Hoff-
mannia. Lappets are known in Adrianites, Neocrimites, and Palermites. Changes
in whorl cross-section occurred in Adrianites, Crimites, Hoffmannia, Palermites,
and Sizilites. Possible changes in shel] secretion are known in Adrianites and
Texoceras.
Agathiceras, of the family Agathiceratidae, displays changes in coiling,
whorl cross-section, and ornament and has an apertural construction and lappets.
Dimorphism is present in the genus; the two forms appear to differ only in
mature diameter.
Three genera of the Cyclolobidae are known to bear mature modifications:
Mexicoceras, Waagenoceras, and Cyclolobus. All three have an apertural con-
striction and changes in coiling and whorl cross-section. Mexicoceras has a
mid-ventral salient in the peristome, and Cyclolobus has lappets.
Four genera of the Marathonitidae display mature modifications: Hyatto-
ceras, Pseudovidrioceras, Marathonites, and Peritrochia. All four have an
apertural constriction. Hyattoceras and some Marathonites display changes in
coiling. Hyattoceras, Pseudovidrioceras, and some Marathonites show changes
in whorl cross-section. Marathonites and Hyattoceras each appear to include
dimorphs; in both genera, the two members of a pair differ in the nature of
the mature modifications rather than just in size.
The genera Waagenina and Stacheoceras of the Vidrioceratidae show
mature modifications. Both have an apertural constriction, lappets, and changes
in whorl shape. Stacheoceras shows changes in coiling. Waagenina appears to
include dimorphs; the two members of a pair differ in the nature of the
mature modifications.
INTRODUCTION
An externally shelled cephalopod, at death, has left an auto-
1Contribution No. 106, Department of Geology, University of Cincinnati.
28 BULLETIN 272
biography of at least part of its life cycle in the form of the shell.
Its growth lines, the increments between the growth lines, and, in-
deed, the shape of the conch, as a whole, record a life history of the
anterior margin of the body; the septa and, hence, the sutures give
witness to the ontogeny of the septal mantle.
Were the conch complete it would give much of the story of the
architectural career of its author, but Nature has seen fit to
obliterate the last chapters of most of these autobiographies, and
many men have followed her lead by inadvertently destroying por-
tions of the body chamber during preparation of the shell. The rela-
tively few well-preserved shells that remain tell an interesting tale.
Many shells indicate that toward the end of the recorded span
of life, changes in the animal occurred — the last-formed part of
the shell has been modified with respect to the earlier-formed por-
tion. These changes in some taxa apparently were minor, but in
others, profound. The modifications may be present in both the
septa and in the conch wall or only in the septa. Moreover, in some
taxa, two conchs which were identical in early growth stages may
be strikingly different after modification, revealing dimorphism.
The purpose of this work is to examine these shell modifications
and any dimorphism thereby indicated.
Five ammonoid families have been studied in detail: the
Adrianitidae, Agathiceratidae, Cyclolobidae, Marathonitidae, and
Vidrioceratidae. At least some taxa of all of these families are known
to exhibit modification in the last-formed part of the shell. That
suitable material of these taxa is available for the present study is
due predominately to the long-term contessorative efforts of the late
A. K. Miller, of W. M. Furnish, and of Brian F. Glenister, all of
the University of Iowa.
All known genera of the five families enumerated have been
studied, and those with modifications are described.
This work should not be construed as an authority on tax-
onomy. I have not made the thorough sutural studies upon which to
found a taxonomic revision. The taxonomy used in this work is based
mostly on the synthesis of Bogoslovsky, Librovitch, and Ruzhencev
(1962), because theirs is the most recent study of most of the taxa
concerned. Newer works, when available, were used. Where the
Soviet Treatise is internally inconsistent or apparently incorrect,
PaLeozoic AMMONOIDs: DAvIs 29
its taxonomy has been modified. The diagnoses given are intended
solely to convey my understanding (or lack thereof) of the taxa
being discussed.
Some of the taxa discussed in this work originally were not
designated with names conforming to the present International
Code of Zoological Nomenclature (for example, Stacheoceras (Mara-
thonites) J. P. Smithi Bose, 1919, and P. tridens form B Haniel,
1915). In order to simplify the tracing of taxonomic history, how-
ever, I have given, at least once, each taxonomic name as it was
originally proposed.
This paper is, in the main, the dissertation I presented to the
University of Iowa. Professor Brian F. Glenister and Professor W. M.
Furnish advised me in the preparation of that thesis. I am greatly
indebted for their gifts of time, experience, and enthusiasm. In addi-
tion notes and photographs they made relating to their comprehen-
sive study of Permian ammonoid type specimens were invaluable.
Revisions were done during my current employment at the Uni-
versity of Cincinnati.
During most of the time I worked on the thesis I held a
National Science Foundation Graduate Fellowship. For that I am
most grateful.
The various scientific institutions listed below graciously lent,
traded, or gave specimens to the University of Iowa (SUI) or al-
lowed them to be studied, as did Francis G. Stehli of Case-Western
Reserve University and, likewise, M. Collignon of Académie Moirans
(the abbreviations are those used in this work): The American
Museum of Natural History (AMNH); British Museum (Natural
History) (BMNH); Geologisch Instituut der Universiteit van
Amsterdam (GIUA); The University of Copenhagen; Institut und
Museum fiir Geologie und Paliontologie der Universitat Tiibingen
(GPIT); Istituto di Geologia dell’Universita Palermo (IGUP); The
Geological Survey of Pakistan; Institut Teknologi Bandung (ITB);
Istituto di Geologia e Paleontologia dell’Universita di Pisa
(MGPU); Muséum national d’Histoire naturelle, Paris (MNHN);
Institut fiir Palaontologie der Rhein. Friedrich-Wilhelm Universitat,
Bonn (PIUB); University of Paris, Sorbonne (UPSM); The Uni-
versity of Texas (Texas); Texas Tech University; Mineralogisch
Geologisch Museum Technische Hogeschool Delft (THD); The
United States National Museum (USNM); Yale Peabody Museum
(YPM).
30 BULLETIN 272
Dr. Claude Spinosa (Boise State College, Boise, Idaho, USA)
and Dr. Eric J. Denton (The Laboratory, Citadel Hill, Plymouth,
England) both have generously provided me with unpublished in-
formation on particular aspects of their work with cephalopods.
I am grateful to them.
LATE ONTOGENETIC SHELL MODIFICATION
In the externally shelled cephalopods as a whole, the changes in
morphology which occurred toward the end of that span of life
recorded in the shell included any number of the following:
. changes in coiling,
. changes in whorl section,
. changes in ornament (including coloring),
. development of an apertural constriction,
. development of an apertural shell thickening,
. formation of a characteristic ultimate peristome (with lap-
pets or a rostrum, for example),
7. changes in shell deposition (such as, production of a black
border at the terminal peristome in Nautilus),
8. progressive reduction of interseptal interval (approximation),
9. progressive simplification of the ultimate few sutures,
10. thickening of the last septum,
11. development of muscle scars.
Some taxa are not known to have developed modifications.
This may reflect an actual lack of such modifications (Ruzhencev,
1962, p. 255) or be due to either unavailability of suitably pre-
served material or to inadequate study.
A modified individual generally has at least several aspects of
its conch affected. Various modifications occurred at diverse times in
the ontogeny of the individual; for example, a change in coiling was
wrought before the formation of an apertural constriction,
Within any genus there seems to have been but one or two
schemes of overall modification. Within any one such scheme, of
course, the development of a particular aspect of modification varied
from individual to individual.
In years past, most authors (for example, Hyatt, 1874) at-
tributed late ontogenetic shell modifications to senility — the
gerontic individual just couldn’t maintain discipline in its shell-
ON Mm RW Ne
PaLeozoic AMMonoIDs: DAvis 31
forming faculties, so chaos resulted. Hyatt (1874) also noted that
what he interpreted as pathologic dwarfs commonly are modified
very similarly to those he regarded to be large gerontic forms.
Coémme (1917) considered at least some modifications as indica-
tive that their bearer was a “small, degenerate hybrid”. Apparently
struck by the similarity of some modifications to varices in snails, to
the parabolic nodes of certain Mesozoic ammonoids, and to phrag-
mocone constrictions, certain authors (for example, Nikitin, 1884;
Siemiradzki, 1925; Davitashvili and Khimshiashvili, 1954; Sazonov,
1957; fide Makowski, 1962) have suggested that the modifications
are not late ontogenetic features at all, but were produced early
or periodically and eroded or resorbed to allow further growth. Kerr
(1931) was so impressed by the apparent external uselessness of
most shell modifications that he concluded ammonoid shells were
internal. Only one genus of externally shelled cephalopods is ex-
tant — Nautilus. In this form, shell modifications occur at and are
directly related to the onset of sexual maturity (Willey, 1902;
Stenzel, 1964).
Interpretation of late ontogenetic shell modifications in terms
of senility, pathology, and genetic problems appears to be unlikely
because of the great overall uniformity of modifications within each
taxon or subdivision of that taxon. Resorption and erosion should
have left partially resorbed and partially eroded individuals — none
are known (Makowski, 1962); and to remove all modifications
would, in some cases, necessitate erosion or resorption of more than
one and one-half whorls of the shell. These facts, coupled with the
knowledge that the conch changes in Nautilus are associated with
maturity, strongly indicate that the late ontogenetic shell modifica-
tions in the externally shelled cephalopods are terminal and truly
are mature modifications.
(“Mature” here is used as meaning sexually mature. This word
has been used previously, for example, by Miller and Furnish, 1940a,
apparently to indicate that an individual had attained what were
considered to be its taxonomically diagnostic characteristics; in this
work the word is placed in quotes when it is used in the sense of
Miller and Furnish. The state of sexual maturity has been denoted
previously as “fully mature”. )
The various mature modifications must have served some
32 BULLETIN 272
useful purpose (or, at the very least, have been neutral in effect),
otherwise these modications would not have persisted.
One could reasonably assume that mature modification pre-
pared the animal for a mode of life different from that of its youth;
this may well not have been entirely the case. It is known that at
least some of the modern dibranchiate cephalopods (for example,
Loligo and Sepia), reach maturity only to quickly breed and die
(Lane, 1960, and personal communication, E. J. Denton). Lane
(1960) and Wells (1962) pointed out that captive Octopus always
die after brooding their eggs, and Wells noted that it is possible
that cephalopods, in general, spawn only once and then die. It is
known that periodically great numbers of mature Nautilus shells
are piled onto the beaches of the southwestern Pacific Ocean, sug-
gesting mass death, perhaps after mating. At least the possibility
exists that the ammonoid had to face its environment as a mature
individual for only a brief period of time; hence mature modifica-
tions may have been related mainly to sex and have had little other
ecologic significance.
If the two sexes of a taxon were morphologically different and
spent much of their lifespan as adults, it is difficult to envision how
they could have occupied the same ecologic niche and still be of
such different morphologies. But, then, perhaps the two sexes lived
in different environments (Westermann, 1964, p. 39 et seq.; 1969).
The two sexes, after all, would have needed to have been in the
same place for only a short time.
Each mature modification must be examined both to suggest
what sexual significance the given modification might have had and
to indicate its possible nonsexual ecological implications. In order to
keep such speculations from becoming completely idle, the biology
of extant organisms must be borne in mind — particularly the
biology of the surviving cephalopods and, of these, especially
Nautilus.
Modifications relating to sex might include mechanisms:
1. for sex recognition,
2. for mating battles, either actual or ritual, between conspecific
individuals of the same sex,
. for the enclosure of sex organs,
4. for the accommodation of sex organs,
OW
PatEozoic AMMONOIDs: Davis 33
5. for the support of sex organs,
6. to aid copulation,
7. to aid oviposition,
8. for brooding eggs, young, or both.
(Because almost any feature could be claimed to have been used for
sex recognition, this aspect will not be mentioned in the following
discussion. )
Modifications (both those related to sex and those not pertain-
ing to sex) might affect the animal’s:
. hydrodynamic and hydrostatic characteristics,
. mobility and maneuverability,
. sensory ability,
. security from external organic and inorganic dangers,
. feeding,
. disposal of waste.
Ganev in coiling most probably would have had hydrodynamic
effects. Likewise the orientation of the animal would have changed;
this would have altered the position and orientation of the peristome
and, hence, the potential orientation of the hyponome.
Changes in whorl section also would have altered the hydro-
dynamic characteristics of the conch. Development of an angular
venter (for example, in Hyattoceras and in Cyclolobus) or of a
ventral keel would seem to be a modification for a more mobile
existence. A broadening of the venter in transverse section, particu-
larly if coupled with a change in coiling to render the venter flat
as viewed laterally, would seem admirably suited for bottom dwelling
(Ruzhencev, 1962). In Nautilus the conch of the male is broader
than that of the female in order to accommodate the spadix.
Changes in ornament, like the loss of color banding ventrally
and addition of coloring dorsolaterally in Nautilus, could have been
used for camouflage (see Cowen, et al., 1971). Miller (1944) specu-
lated that the spines on the penultimate one-half whorl of Pseud-
agathiceras might have served to protect the bottom-dwelling animal
from attack from above. The loss of longitudinal ornament and
the appearance of transverse plications in the ultimate one-
quarter volution of Sizilites, as well as the spines in Pseudagathiceras
would certainly have affected the hydrodynamics. The purpose of —
the deep depression in each flank of the ultimate living chamber
of Agathiceras swessi is unclear.
Nn PWN
34 BULLETIN 272
Development of an apertural constriction, an apertural shell
thickening, or both would have strengthened the ultimate shell mar-
gin; this would have been of particular importance inasmuch as shell
repair apparently ceased after mature modifications were completed.
Perna (1915) pointed out that constriction of the aperture would
have afforded the animal protection from outside influences, not
the least of which would have been the attempt by a potential preda-
tor to wrench the cephalopod from its conch (Berry, 1928). The
addition of shell materia] at the aperture would have affected the
hydrostatic characteristics of the conch, thereby altering the orienta-
tion of the hyponome, but an even greater effect on the hyponome
would have been the barricading of the aperture (Perna, 1915) —
the hyponome would have needed to shoot over the constriction
(or shell thickening or both) or to extend beyond it, as in Nautilus.
The thickening of the shell material might have been simply a
device to rid the body of excess calcium carbonate. A highly con-
stricted aperture would have imposed limitations on the size of
food that could have been consumed, unless the cephalopod could
extrude its body as do the modern cowries. Perhaps young ammon-
oids were protected by living behind the apertural constriction of a
parent (Schmidt, fide Berry, 1928). The purpose of the two pro-
nounced depressions within the apertural constriction of Cyclolobus
is unclear. Likewise, the purpose of the slight apertural shell thick-
ening in Nautilus has not been established.
Lappets might have served in mating (or other) battles; they
don’t appear strong enough to have functioned as real weapons,
but they might have done service in bluffing such as occurs in the
extant cephalopods Loligo and Sepia. They could have been utilized
as supports for sex organs or perhaps for protection of eggs or young.
Lappets might also have been used as supports of nonsexual organs;
Gillet (1937) suggested that these structures might have supported
arms. Mikesh and Glenister (1968) contemplated that hollow lateral
shell extensions in Solenochilus, a Carboniferous nautilid, might have
enclosed extensions of the mantle which served a sensory function.
The development of lappets may have been to eliminate part of
the weight of the shell, while still retaining some of its supporting
and protective attributes. On the other hand the lappets may
have added weight anteriorly to alter the orientation of the conch.
Lappets might, in addition, have served a hydrodynamic function.
PaLteozoic AMMoNoOIDs: Davis 35
Ruzhencev (1962) suggested that they supported the animal when
it advanced from the shell and perhaps allowed the creature to
move into and out of the conch more rapidly; to him this indicated
a more predational way of life.
The rostrum has been taken by many authors to indicate loss
of swimming ability, but some creatures with rostra seem from other
features to have been nektonic (Dunbar, 1924). As pointed out by
Perna (1915) a rostrum might have served as a funnel support; in
fact, the hyponome may have been bifid, one half lying on either
side of the rostrum (Trueman, 1941). The rostrum could have sup-
ported the animal when it advanced from its shell. This structure
may have been used as a weapon, either real or apparent, or as a
support for sensory or other organs. It is likely that the rostrum
would have had a hydrostatic effect, bringing the aperture down, and
might also have resulted in streamlining. Dunbar (1924) suggested
that a hyponomic sinus would be needless for an animal with a long
muscular body which could be protruded from the shell; in fact,
he said, this protrusion would displace more water so that the animal
would have become more buoyant the instant it started to swim.
Perhaps the rostrum served for brooding. Jeannet (1948) regarded
all ammonoids bearing rostra as dibranchiates but stated no reasons.
In Nautilus there apparently is a change in shell deposition
about the time the final aperture is formed — the “black layer”
which is restricted to the dorsum in juveniles is extended to ring
the aperture. Zittel (1895) equated this “black layer” with the
runzelschicht of ammonoids. The raised points on the venter of the
interior of the modified portion of the shell of, for example, some of
the adrianitids may be a ventral extension of runzelschicht and, if
so, could be homologous to the black ring of the mature Nautilus.
Pruvot-Fol (1937) suggested that the black matter in Nautilus con-
sists of metabolic wastes.
Willey (1902) noted that usually, but not always, the last few
septa of Nautilus are progressively more crowded. Such approxima-
tion also occurred in at least some ammonoids (and fossil nautiloids,
too, for that matter). Makowski (1962) reasoned (for Quenstedto-
ceras, a Mesozoic ammonoid ) that lengthening of the shell stopped
before septal secretion ceased; this resulted in a shortening of the
body chamber and a progressive approximation of the ultimate few
septa. Whether this actually occurs in Nautilus is unknown. Shorten-
36 BuLLETIN 272
ing of the living chamber might be linked with an enlargement in
whorl cross-section. In any case there probably would have been an
effect on the center of gravity and, hence, on orientation of the live
animal.
In ammonoids other than those with simple sutures septal
crowding was accompanied by a sutural simplification. Makowski
(1962) indicated that further growth of such specimens and eventual
return of the sutures to the normal shape have never been ob-
served. He stated further that the septal crowding present at places
in the phragmocones of some ammonoids is never accompanied by
sutural simplification and that the phragmocone constrictions in
goniatites and other forms are never accompanied by densely spaced
septa and simplified sutures. (Suture simplification followed by
further complication are, however, known in the early ontogenetic
stages of Agathiceras [fide Miller and Furnish, 1939] and Paracel-
tites [personal communication, Claude Spinosa] ). Sutural simplifica-
tion accompanying septal approximation would seem to be merely a
mechanical adjustment to septal crowding.
The ultimate septum of externally shelled cephalopods common-
ly is greatly thickened. This thickening, surely of hydrostatic im-
port, also may have served to provide a stronger base for muscle
support. Perhaps the animal disposed of excess calcium carbonate
in this manner. The purpose and cause of the thickening of the ulti-
mate septum in Nautilus is unknown.
Makowski (1962) reported that in certain (but not all) taxa
supplementary prismatic shell material was added at the base of
the ultimate body chamber for muscle attachment.
At present the physiology and ecology of Nautilus are virtually
unknown. Until the basic research on these subjects is done, evalua-
tion of the hypotheses concerning the physiologic and ecologic im-
plications of ammonoid mature modifications is premature. More-
over the possibility that the ammonoid was not closely similar to
any extant animal cannot be precluded completely. The various
mature modifications might have been suited for anatomies not now
within our sphere of knowledge.
DIMORPHISM
Since before the middle of the Nineteenth Century paleon-
tologists working with the ammonoid cephalopods have recognized
PaLeEozoic AMMONOIDs: Davis 37)
that mature individuals commonly occur in paired groups. One of
each pair is larger and generally has a simple peristome; the smaller
form commonly has a more ornate aperture. Particularly impressive
are the facts that the pairing is widespread in time and space and
that the two groups of a pair are closely similar except near and at
maturity. This pairing has been attributed to sexual dimorphism.
Phenetic differences between two groups of individual am-
monoids collected from the same horizon and locality could be due
to any of a number of factors.
The two groups might represent two separate taxa (of any
level); in other words, phyletic splitting had occurred. If the taxa
are conspecific, the presence of both in one place simultaneously
could be due, for example, to a recent breakdown of the barrier
which led to the phyletic split, to ocean water depth stratification
of the taxa, to activity at different times of the day, or to post-
mortem transportation.
The two groups might indicate dimorphism (that is, the
existence in one taxon of two distinct phenetic groups).
Dimorphism could seem present in the local collection because
of:
a high proportion of pathologic individuals,
a high proportion of mutant individuals,
post-mortem deformation,
post-mortem sorting,
preferential destruction, or
faulty sampling.
In one area pathologic conditions could result in two groups,
but such conditions would be unlikely to be widespread and long-
lived. Likewise, in a small area, taxonomically unimportant but
phenetically striking mutation (for example, a high proportion of six-
legged frogs in a single pond) could produce apparent dimorphism,
but, again, over a broad area and extended period of time such
mutations would be lost or become of sufficient frequency to be
taxonomically important. Post-mortem deformation could produce
apparent dimorphism (for example, Davis and Campbell, 1968, dis-
cussed specimens of the longiconic nautiloid Pseudorthoceras
knoxense from Carboniferous rock near Knoxville, Iowa, — those
specimens lying horizontally are flattened, whereas those standing
Dk ON
38 BULLETIN 272
vertically are round in cross-section), but deformation is almost
always recognizable by actual breaks in the specimen, loss of bi-
lateral symmetry, or both, and, moreover, identical conditions of
deformation would be confined to a small geographic area. Sorting or
preferential destruction might, for example, remove middle-sized
specimens, leaving apparent dimorphism — again studies of collec-
tions from other strata and localities would reveal the local nature
of such conditions of sorting or preservation. A non-representative
sample of the local fauna could give a distorted picture of even
that fauna. Actual dimorphism should be recognizable throughout
the whole taxon.
Dimorphism could be the result of:
1. alternation of morphologically distinct generations,
2. polymorphism, such as exhibited by certain hymenopteran
insects, for example,
3. morphological differences between different ontogenetic
stages (including neoteny, such as exhibited by the axolotyl,
and instars), or
4. sexual dimorphism.
Of these four, only the last has been documented in the Cephalopoda.
Whether two groups are representatives of separate taxa or are
sexual antidimorphs (opposite members of a dimorphic pair) is a
basic question. The criteria for the identification of sexual dimorph-
ism in ammonoids are well known (Makowski, 1962; Callomon, 1963;
Westermann, 1964):
1. There should be distinct dimorphism between sexual anti-
dimorphs.
2. Sexual antidimorphs should have identical phylogenies.
Sexual antidimorphs should have identical early ontogenies.
4. Sexual antidimorphs should have the same stratigraphic
range.
5. The sex ratio should be consistent both with that in modern
related organisms and within the total geographic and strati-
graphic range of the taxon.
If sexual dimorphism is to be recognized there must be some ob-
servable difference between the two sexes. Between two sexually
antidimorphic groups intermediate forms might exist either because
the ranges of variability of the two groups overlap, because of the
Ww
PaLeozoic AMMONOIDs: DAvIs 39
existence of intermediate mutant or pathologic individuals, or be-
cause of the presence of intersexes in the taxon.
Theoretically, at least, the two sexes of a species need not
evolve in the same way or at the same rate, particularly with respect
to phenetic characters directly involved with sex. Hence, sexual anti-
dimorphs might well not have identical phylogenies and ranges.
McCaleb’s (1968, p. 29) conclusion that “dimorphism is a_pre-
dominant feature at the inception of an evolution lineage and de-
creases throughout phylogeny”, if generally applicable to am-
monoids, may be important.
The respective ontogenies of the two sexes of a species need not
be identical. Sexual characters can appear early in ontogeny. Mc-
Caleb’s (1968) conclusions regarding dimorphism in Syngastrioceras
oblatum (from the Upper Carboniferous of Arkansas), perhaps
cast doubt on the almost universal application of the assumption
that sexually dimorphic characters are most pronounced at maturity.
What he considered to be antidimorphs differ most as juveniles, and
“the conch differences decrease in the more mature forms” (p. 51).
Even if the actual sex ratios in now extinct forms were con-
sistent with such ratios in related extant groups, there are many
possible sources of deviation. Differential mortality could produce
local concentrations of one or the other sex, and, likewise, different
geographic distributions in life, differential preservation, and sorting
could result in local deviations. In all these cases study of the taxon
throughout its whole range would disclose an approximation of the
actual ratio. The general collector tends to gather the more spec-
tacular fossils, so might gather the larger or more striking of the
antidimorphs or might consider poorly preserved specimens of the
smaller antidimorph to be juveniles of the larger and count them
as such. Moreover, the sex ratios reported for the extant cephalopods
(Pelseneer, 1926; Fretter and Graham, 1966; Mangold-Wirz, 1963 )
vary so markedly that virtually no antidimorph ratio could be re-
jected as inconsistent.
In short, each criterion for the identification of sexual dimorph-
ism can be open to serious doubt. It does, however, seem highly
unlikely that paired groups of ammonoids with both members having
closely similar ranges, phylogenies, and ontogenies and in a con-
40 BULLETIN 272
sistent ratio throughout their geographic and stratigraphic ranges
could be anything but sexual antidimorphs.
The conclusion that two forms are sexual antidimorphs presup-
poses that the two forms are conspecific. Hence, the person making
the conclusion has no alternative but to refer to both forms by the
same binomen.
Makowski (1962) recognized two sorts of dimorphism in am-
monoids, based primarily on number of whorls:
In Type A — one antidimorph generally has five to six whorls
but may have as few as four, and the other antidimorph generally
has seven to nine whorls but may have as few as six. The larger
form has a relatively greater whorl height adapertural of whorl five
to five and one-half.
In Type B — one antidimorph generally has seven to nine
whorls but may have as few as six, and the other antidimorph has
at least one whorl more than its opposite. (Guex, 1968, added a third
type, ‘O’, in which the smaller antidimorph has but three to four
whorls. )
Westermann (1964) also recognized two kinds of dimorphism,
based on conch morphology. In the simpler and rarer type the anti-
dimorphs differ only in size but not in other morphologic characters.
In the more abundant type one antidimorph is smaller and has a
more highly modified peristome than the other; the two antidimorphs
may also differ in other mature modifications. Westermann recog-
nized dimorphism, intermediate between the two main types, in
which the antidimorphs differ both in size and in minor morphology,
for example, in the transverse ribs on the mature conch.
Zeiss (1969) recognized three categories of dimorphism. In
Group I, one antidimorph is large and variocostate whereas the
other is small and equicostate. Both antidimorphs of Group II are
ornamented similarly, but there are differences in size and in the
shape of the apertural region. Dimorphism in Group III is suspected
but not clearly recognizable.
The various systems do not correspond. For example, each of
Makowski’s dimorphism types includes representatives of both of
Westermann’s categories.
Beginning with De Blainville (1840), who was the first person
to suggest dimorphism in ammonoids (fide Makowski, 1962), most
PaLeozoic AMMONOIDs: DAvIs 4]
workers have considered the smaller of each dimorphic pair to be
the male. Makowski (1962, p. 58) stated that in all known inverte-
brate groups in which sexual dimorphism is expressed by notable
differences in size, it is the male which is smaller: “Not a single
case of opposite morphological conditions has so far been reported
in this respect.” Wells (1962, p. 33) in discussing extant forms
noted that: “In cephalopods the males are usually larger than their
mates.” Being more specific, Wells (1966, p. 581) stated: “In
Octopus, Sepia, Sepiola, and Loligo, at least, the occurrence of
mating appears to be determined by the male, which is generally
larger than the female. . .”.
Mangold-Wirz (1963) gave length and age measurements for
sixteen extant cephalopod species. Although in some cases the maxi-
mum size of males exceeds that of females, in every example given
save one the length of the female upon reaching maturity is greater
than that of the male at maturity. In that one case, Sepia officinalis,
the female is just at the upper end of the size range of the males.
(Of interest is the fact that, except in Sepia officinalis, the female
takes longer to reach maturity. Moreover, in those cases in which
the maximum length of one sex exceeds that of the other, it appears
that the maximum age of the one also exceeds that of the other.)
Westermann (1969) pointed out that size relationships of the
sexes of extant cephalopods are extremely variable even at the genus
level. Although he gave no measurements with his examples, he
concluded that “wherever marked sexual dimorphism in_ size
(> 1.5 in diameter) is present in the cephalopoda, the female is
larger than the male.”
Mangold, Lu, and Aldrich (1969) discussed dimorphism in the
extant squids J/lex illecebrosus illecebrosus, I. illecebrosus argentinus,
and J. illecebrosus coindetu, in all of which it is generally accepted
that females are larger than males. In all three subspecies the mean
mantle length of females exceeds that of males. However, in J.
dlecebrosus coindetu, and apparently also in J. illecebrosus argen-
tinus, the males and females differ in proportions; for example, in
I, illecebrosus coindetui the arms of the male are longer, the head
of the male is broader, and the diameter of the largest suckers is
larger in males than in females.
In Nautilus the male is slightly broader than the female (Willey,
1902). Stenzel (1964) illustrated (after Willey, 1902) the maximum
42 BuLLeETIN 272
and minimum whorl width differences between the sexes of Nautilus.
The male is shown to be as much as 25% wider than the female.
(Westermann’s 1969 statement about this illustration that the fe-
male is 50% wider is apparently a typographic error.)
Coémme (1917, fide Makowski, 1962) stated that in ammonoids
the smaller antidimorph of a pair is the less abundant, as are
the males of modern cephalopods. Pelseneer (1926, and extracted
in Fretter and Graham, 1966) collated reports of sex ratios for
modern cephalopods varying from 3:1 in favor of males to nearly
7:1 against; most of his examples give females the majority. Man-
gold-Wirz (1963) gave sex ratios for 16 extant cephalopod species
which range from 143 males for every 100 females to 39 males for
every 100 females; she discussed that season of sampling, place of
sampling, and relative susceptibility of the sexes to capture may be
significant in her statistics, Westermann (1969) also discussed sex
ratios in extant cephalopods, paying particular attention to migra-
tion of populations and the commonly concomitant segregation of
sexes. Mangold, Lu, and Aldrich (1969) studied 382 males and 351
females of Illex illecebrosus coindetii as well as 452 males and 457
females of J. dlecebrosus illecebrosus. Willey (1902) listed 150
male: 66 female Nautilus pompilius, but 10 male: 16 female Nautilus
macromphalus.
One is forced to wonder whether size differences and (especially)
sex ratios in modern forms are sufficiently well understood to extra-
polate to fossil cephalopods.
Within the mature living chamber of an Eleganticeras elegantu-
lum macroconch (the larger antidimorph of a pair) Lehmann (1966)
found a cluster of spherical structures of the same size as the pro-
toconch of the species. He cautiously interpreted these structures as
egg coatings. This, if the brooding of eggs by the female in Argo-
nauta and Octopus proves to be the general case in the extant
cephalopods, might be taken to indicate that the macroconch is the
female.
As related in the excellent historical surveys of Makowski
(1962) and Callomon (1963), most students of ammonoid dimorph-
ism have concentrated their attentions on Mesozoic fossils. However
Foord and Crick (1897), Haug (1897), Perna (1914), Demanet
(1943), McCaleb and Furnish (1964), Furnish and Knapp (1966),
PaLreozoic AMMONOIDs: DAVIs 43
McCaleb (1968), and Davis, Furnish, and Glenister (1969) have
all at least suggested the existence of dimorphism in ammonoids
from the Paleozoic.
The greatest obstacle to study of dimorphism in Paleozoic am-
monoids is the marked lack of suitably preserved faunas. In order to
establish that dimorphism exists, it is necessary to study large num-
bers of fossils which retain mature modifications. In addition, the
internal whorls must be well preserved, if ontogeny is to be deci-
phered and whorls counted. In none of the five families studied 1s
any fauna known to me which fully satisfies the requirements of
abundance and suitable preservation. The Agathiceras fauna from
the Road Canyon Formation of Texas discussed by Davis, Furnish,
and Glenister (1969, pp. 103, 105-107) lacks preservation of internal
whorls.
Of the taxa discussed in the descriptive portion of this work
only four, Agathiceras, Hyattoceras, Marathomtes, and Waagenina,
exhibit two distinct mature forms. Unfortunately, paucity of avail-
able mature individuals of the last three forces me to regard the
dimorphism apparent in these three genera as only theoretical. On
the other hand, dimorphism in A gathiceras seems to be statistically
justifiable (Davis, Furnish, and Glenister, 1969). But even this
case is established only on the basis of a single collection from one
horizon and locality and, because the ontogeny and phylogeny of
the studied specimens is unknown, any too positive attribution of
the dimorphism apparent in Agathiceras to sex seems to me to be
premature.
DESCRIPTIVE PART
Superfamily ADRIANITACEAE Schindewolf, 1931
Family ADRIANITIDAE Schindewolf, 1931
Diagnosis. — The Adrianitidae includes ammonoids in which
the suture underwent goniatitic ontogenetic development; sub-
sequent new elements were then added in the saddle positioned on
the early adolescent umbilical seam, and these elements migrated
toward the dorsum and venter respectively.
Genus ADRIANITES Gemmellaro, 1887
Diagnosis. — Adrianites includes adrianitids in which the
44 BULLETIN 272
“mature” suture on each side has six or seven lobes between the
bifid ventral lobe and the umbilical shoulder.
Synonymy. — Glenister and Furnish (1961, p. 726) suggested
that Neocrimites (Sosiocrimites) Ruzhencev, 1950, and Aricoceras
(Neoaricoceras) Ruzhencev, 1950, are synonyms of Adrianites.
Specific composition. — Adrianites elegans Gemmellaro, 1887,
is the type species of Adrianites by original designation. Adrianites
kingi Gemmellaro, 1887, is the only species which was referred to
Aricoceras (Neoaricoceras) by Ruzhencev (1950). Adrianites instg-
nis Gemmellaro, 1887, and A. insignis var. biassalensis Toumanskaya,
1931, are the only two taxa which were referred to Neocrimites
(Sosiocrimites) by Ruzhencev (1950); Furnish and Glenister (in
Davis, Furnish, and Glenister, 1969, explanation to plate 3) consid-
ered A. insignis to belong in Neocrimites. In addition to the pre-
viously listed taxa, A. isomorphus Gemmellaro, 1888, is referred to
Adrianites, and the following inadequately described taxa may be-
long in the genus: A. bobkovae Toumanskaya, 1949, A. permicus
Toumanskaya, 1935, A. planus Toumanskaya, 1935, and A. vino-
gradovi Toumanskaya, 1949. A. schmidti Toumanskaya, 1937a, and
A. bobkowi Toumanskaya, 1937a, are apparently nomina nuda,
Immature shell. — The conch of Adrianites has been described
as discoidal to subspherical, involute, and with a small umbilicus
(Ruzhencev, 1950, p. 204).
Constrictions are present in most, if not all, individuals of
Adrianities. In the type species A. elegans they are prominent on
the shell exterior, producing a slight but conspicuous decrease in
whorl height immediately adoral of each constriction. Similar de-
creases in whorl height occur in representatives of A. insignis and
A. isomorphus, even though the constrictions in these two taxa are
less prominent than in the type species. In A. kimgi, on the other
hand, the only evidence of constrictions on the shell exterior is the
periodic closer spacing of the transverse lirae. The maximum num-
ber of constrictions per whorl apparently varies from two (in A.
tsomorphus) through three (in A. elegans and A. kingi) to four
(in A. insignis).
In the type species Adrianites elegans the longitudinal and
transverse lirae are equally conspicuous. In the genus as a whole
the relative prominence of these two features varies from that dis-
played by A. tsomorphus, in which the transverse components are
PaLeozoic AMMONOIDs: Davis 45
much the less striking, to that shown by A. king, in which the longi-
tudinal ornament is restricted to the flanks in the form of scallops
in the transverse lirae. The longitudinal lirae are weak or absent
in the constrictions. In some individuals the longitudinal ornament
affects both the shell exterior and interior.
The overall trend of each half of the external suture is arcuate.
Mature shell. — After about two-thirds of the ultimate whorl
of Adrianites had been formed, there occurred a slight decrease in
the radius of curvature of coiling, followed by a slightly greater in-
crease; this resulted in the venter being somewhat flattened in
longitudinal section. This change in coiling is particularly evident
in the umbilici of specimens of A. tsomorphus and A. kingit where
it resulted in an actual decrease in the size of the umbilicus. In A.
elegans and in at least some representatives of A. imsigmis the
umbilicus seems little affected (but in these taxa, the flattening
of the venter is, itself, slight). In at least some individuals, for
example of A. tsomorphus and A. elegans, the change in coiling was
accompanied by a slight but distinct increase in relative whorl
width; this bulging appears greatest in the dorsolateral portions of
the flanks.
Near the adapertural end of the mature body chamber of
Adrianites is a constriction whose trend is somewhat different from
those adapical of it. Shortly before the apertural constriction was
formed, a decrease in the radius of curvature of coiling occurred,
leading directly into the constriction.
As in the immature constrictions of Adrianites, longitudinal
lirae are not present in the ultimate constriction or are less prom-
inent there than adapical of it. The growth lines are more closely
spaced and are commonly less prominent in the terminal constriction
than adapical of it. Beginning in the adoral portion of the apertural
constriction each successive growth line shows a more accentuated
ventrolateral salient on each side, forming the lappets of the ulti-
mate peristome. The lappet figured by Gemmellaro (1888, pl. D,
fig. 10) is straight. Each lappet bears an adaperturally directed
branch of the ultimate constriction.
The leading edge of the dorsal shell of Adrianites has a distinct
lip (for example, USNM-a and SUI 32442). The surface of the dorsal
shell is ornamented with irregularly spaced ridges and raised points
46 BuLLeETIN 272
arranged in a generally radial pattern; this is interpreted as runzel-
schicht.
The ventral surface of the internal mold of the mature body
chamber in some individuals (for example, USNM-b) bears minute,
irregularly spaced pits which correspond to raised points on the in-
ward-facing surface of the shell. In one place on USNM unnum-
bered specimen b the outermost portion of the shell has broken
away, revealing that the raised points on the inward-facing surface
of the shell correspond to globular- or rod-shaped bodies within the
shell itself. The presence of these bodies in some and their absence
in other individuals even in the same species remains unexplained.
Gemmellaro reported the following living chamber lengths for
members of Adrianites: one and one-third whorls in A. elegans (1887,
p. 44), more than one volution in A. imsignis (1887, p. 45), and a
whorl and a half in A. king (1887, p. 47) and in A. tsomorphus
(1888, p. 14). The ultimate constriction and final septum were ob-
served by the present author in only one individual (A. tsomorphus
— SUI 32442); they lie about one and one-third whorls apart.
The maximum diameter of specimens of Adrianites known to
the author are: A. elegans — 20% mm and 21 mn, A. kingi — 22
mm, A. itsomorphus — 25 mm, 26 mm, 27 mm, 27 mm, and A.
insignis — 27 mm and 29 mm.
Material. — The preceding observations are based on study of:
Gemmellaro (1887, 1888), Toumanskaya (1931, 1935, 1937a, 1949),
Miller and Furnish (1940a), Ruzhencev (1950), Glenister and
Furnish (1961), Bogoslovsky, Librovitch, and Ruzhencev (1962),
Davis, Furnish, and Glenister (1969), three mature and one nearly
mature specimens of A. tsomorphus (USNM unnumbered, SUI
32442), two mature examples of A, cf. A. insignis (GPIT unnum-
bered, BMNH C 37654), and photographs and notes taken by
W. M. Furnish and Brian F. Glenister in the summer of 1965.
Illustrations. — The following mature specimens of Adrianites
are figured: two A. elegans — IGUP 85 (PI. 3, figs. 1-3), (Pl. 3,
figs. 5-7), two A. insignis — MGPU unnumbered (PI. 5, figs. 4, 5),
IGUP 84 (PI. 6, figs. 7, 8), one A. cf. A. insignis — BMNH C37654
(Pl. 5, figs. 1-3), four A. tsomorphus — SUI 32442 (PI. 3, fig. 4),
IGUP 86 (PI. 4, figs. 1-3), USNM unnumbered a (PI. 4, figs. 7, 8),
USNM unnumbered b (PI. 4, fig. 9), and one A. kingi — IGUP 81
(Pl. 5, figs. 6-8).
PaLEozoic AMMONOIDs: DAvIs 47
Genus CRIMITES Toumanskaya, 1937b
Diagnosis. — Crimites includes narrowly umbilicate adrianitids
in which the “mature” suture on each side has three complete lobes
between the bifid ventral lobe and the umbilical shoulder.
Specific composition. — The type species of Crimites is Agathi-
ceras krotowi Karpinsky, 1889, by subsequent designation of Glenis-
ter and Furnish (1961, p. 726). In addition to the type species the
following are referred to Crimites: C. elkoensis Miller, Furnish, and
Clark, 1957, C. glomulus Ruzhencev, 1952, C. pamuricus var. darensis
Toumanskaya, 1949, C. singularis Bogoslovskaya, 1962, and C.
subkrotowit Ruzhencev, 1938. Adrianites globosus Chernov, 1939,
is a synonym of Crimites krotowi (fide Ruzhencev, 1956a, p. 249).
Crimites busterensis Toumanskaya, 1937b, C. pamiricus Toumans-
kaya, 1937b, C. smithi Toumanskaya, 1937b, C. spathi Toumans-
kaya, 1937b, and C. undosus Ruzhencev, 1952, are apparently all
nomina nuda. Adrianites gemmellarot Toumanskaya, 1931, A. hamiel
Toumanskaya, 1931, Agathiceras oyensi Haniel, 1915, A. rothpletz
Haniel, 1915, and Crimites subkrotovi var. murgabensis Toumans-
kaya, 1949, may belong in Crimites.
Immature shell. — The conch form of Crimites has been
described as spherical to strongly globose (Ruzhencev, 1950, p. 202).
The shell is involute.
Only one individual of Crimites known to the author (Ruzhen-
cev, 1956a, pl. 39, fig. 3) has immature constrictions; three shallow
ones are visible in its outermost preserved whorl.
The relative prominence of the longitudinal and transverse lirae
of Crimites is variable; the spiral ornament may be much the more
striking, may be just as conspicuous as the transverse lirae, or may
be represented only in the form of scallops in the growth lines. In
those individuals with prominent transverse lirae, each lira is at
the crest of a transverse shell plication. On the internal mold the
peak of each plication is, itself, scalloped. Along the umbilical
shoulder of some representatives of the genus there runs a thicken-
ing of the shell layer (Ruzhencev, 1941, p. 884).
In addition to the three lobes between the bifid ventral lobe
and the umbilical shoulder of Crimites, on each side there are three
lobes between the umbilical seam and the dorsal lobe, and there are
a variable number of lobes on the umbilical wall. Each “internal
48 BuLLETIN 272
lateral lobe” is tied to its respective “external lateral lobe”. The
lobe just ventrad of the shoulder may be bifid.
Mature shell. — Only comparatively slight changes in coiling
occurred in the final stages of the ontogeny of Crimites. In some
individuals, after al] but about the last quarter of the ultimate volu-
tion was formed there was a slight decrease in the radius of curvature
of coiling, followed by a like increase; this resulted in a slight flat-
tening of the venter as viewed in longitudinal section. This change
in coiling is accompanied by a gentle increase in the ratio W/D.
The umbilicus appears to have been little affected.
Near the adapertural end of the mature living chamber of
Crimites is a broad constriction.
The plications in the shell of Crimites do not seem to occur in
the ultimate constriction, although growth lirae are present. Longi-
tudinal ornament extends into the adapertural constriction; at least
the transverse lirae there are scalloped.
No specimen of Crimites known to the author retains the ulti-
mate peristome.
The ultimate constriction of Crimites varies from three-quarters
to just less than one whorl adapertural of the ultimate septum.
All three mature specimens of Crimites known to the author
have a diameter between 22 and 23 mm.
Material. — The preceding observations are based on study of:
Haniel (1915), Toumanskaya (1931, 1937a, 1937b, 1949), Ruzhen-
cev (1941, 1950, 1951, 1952, 1956a), Miller, Furnish, and Clark
(1957), Glenister and Furnish (1961), Bogoslovskaya (1962), and
two mature and one nearly mature specimen of Crimites krotowti
(SUT 5981).
Illustrations. — Two mature specimens of Crimites krotowt are
figured: SUI 5981a (PI. 6, fig. 6) and SUI 5981b (PI. 6, figs. 3-5).
Genus EPADRIANITES Schindewolf, 1931
Diagnosis. — Epadriamtes includes globose, fairly broadly
umbilicate adrianitids in which the “mature” suture on each side
has four or five lobes between the bifid ventral lobe and the umbilical
shoulder, four or five lobes between the umbilical seam and the
dorsal lobe, and two or three lobes on the umbilical wall.
Synonymy.— Those specimens which have been referred to
PaLEozoic AMMONOIDs: DAvis 49
Basleoceras Ruzhencev, 1950, are regarded as congeneric with in-
dividuals of Epadriamtes.
Specific composition. — The type species of Epadrianites is
Agathiceras timorense Boehm, 1907, by original designation.
Agathiceras beyrichi Haniel, 1915, is the type species, and Adriamites
haueri Gemmellaro, 1888, is the only other species referred to
Basleoceras by Ruzhencev (1950, p. 203). In addition, Agathaceras
timorense var. involuta Haniel, 1915, is referred to Epadrianites.
Immature shell. — The conch of Epadrianites has been described
as moderately involute and spherical or with width greater than the
diameter (Ruzhencev, 1950, p. 203).
The majority of specimens of Epadrianites bear no evidence of
constrictions. One specimen of E. timorense (SUI 32439) has six
gentle transverse ridges in its outermost preserved whorl on which
the longitudinal lirae are less distinct than elsewhere; another
individual of the same species (PIUB unnumbered) is similarly
ornamented. The ridges of both of the examples are apparently
constrictions.
The most striking ornamental feature of Epadrianites is the
presence of prominent longitudinal lirae with faint transverse growth
lines. Basleoceras and Epadrianites were distinguished on the basis
of the difference in the spacing of the longitudinal lirae (Ruzhencev,
1950), but Haniel (1915, pl. 50, fig. 14) illustrated what is an ap-
parently transitional form. Ruzhencev (1950, p. 203) stated that
Basleoceras has growth lines with a weak ventral sinus, whereas in
Epadrianites the growth lines each form a ventral salient. As far as
the present author is able to determine, there is little if any dif-
ference in the trend of the growth lines of the two “genera”.
Mature shelil.— Only one mature specimen of Epadrianites is
known to the author (IGUP 75). After about three-quarters of the
ultimate whorl was formed, there was a slight decrease in the
radius of curvature of coiling followed immediately by a slightly
greater increase. Just before the apertural constriction was formed,
there was a marked decrease in the radius of curvature leading
directly into the constriction. These changes in coiling result in a
flattening of the longitudinal section of the venter with a terminal
upturning of the venter. The longitudinal ornament continues on into
the ultimate constriction. The final peristome of Epadrianites is
50 BuLueTIN 272
unknown; the growth lines seem to retain their immature trend as
far as preserved.
The exact length of the mature living chamber of Epadriamtes
is not known, although Gemmellaro (1888, p. 17) stated that it is
much greater than one whorl. The diameter of the one known
mature individual is 66 mm.
Material. — The preceding observations are based on study of:
Gemmellaro (1888), Haniel (1915), Miller and Furnish (1939),
Ruzhencev (1950), Glenister and Furnish (1961), one immature
individual of Epadrianites beyricht (SUI 32440), three immature
specimens of FE. timorense (SUI 32439, 12321), and photographs
and notes taken by W. M. Furnish and Brian F, Glenister in the
summer of 1965.
Illustrations. —One mature specimen of Epadrianites haueri
is figured — IGUP 75 (PI. 8, figs. 1-4). One immature example of
E. tumorense is also illustrated — PIUB 21b (PI. 9, figs. 8, 9).
Genus HOFFMANNIA Gemmellaro, 1887
Diagnosis. — Hoffmannia includes strongly evolute, transversely
plicate adrianitids in which the “mature” suture on each side has
five lobes between the bifid ventral lobe and the umbilical seam.
Specific composition.—The type species of Hoffmannia is
Adrianites (Hoffmannia) hoffmanni Gemmellaro, 1887, by mono-
typy. In addition to the type species, Adrianites (Hoffmannia)
burgensis Gemmellaro, 1888, is referred to Hoffmannia. H. fisher
Plummer and Scott, 1937, is probably not closely related to the
genus (fide Miller and Furnish, 1940a, p. 110).
Immature shell. — The conch of Hoffmannia is serpentine, evo-
lute, with narrow whorls and a broad umbilicus (Bogoslovsky,
Librovitch, and Ruzhencev, 1962, p. 394).
All specimens of Hoffmannia known to the author have con-
strictions which extend from seam to seam. There are two or three
of these constrictions per whorl.
The flanks of specimens of Hoffmannia are marked by con-
spicuous simple or dichotomizing transverse plications. These plica-
tions are restricted to the flanks, but on the venter there are deli-
cate transverse ribs which appear to be growth lirae.
Mature shell.—Only one individual known to the author
(IGUP 78) may be mature. Its venter has been eroded, and ap-
PaLeozoic AMMONOIDs: Davis 51
parently the creature had been seriously injured in life, for the plica-
tions about one-eighth whorl adapical of the terminal constriction
are markedly disturbed.
The apparent flattening of the venter of the last quarter volu-
tion as viewed in longitudinal section may be a mature modification
or might be the result of erosion. The apparent rapid increase in
whorl height in the region just adapical of the terminal constriction
may be a modification, may have resulted from the animal’s injury,
or may be the product of erosion of the shell apical of the region.
Nonetheless, there does seem to be a gradual increase in the coarse-
ness of the ventral ribbing in an adapertural direction throughout
the ultimate whorl.
~The living chamber of Hoffmanma is one and one-half whorls
long (Gemmellaro, 1887, p. 50).
The diameter of the one possibly mature individual is 24 mm.
Material. — The preceding observations are based on study of:
Gemmellaro (1887, 1888), Plummer and Scott (1937), Miller and
Furnish (1940a), Glenister and Furnish (1961), Bogoslovsky,
Librovitch, and Ruzhencey (1962), two immature specimens of
Hoffmannia hoffmanni (USNM unnumbered, SUI 32434), and
notes and photographs taken by W. M. Furnish and Brian F.
Glenister in the summer of 1965.
Illustrations. — One specimen of Hoffmannia is figured, a
mature individual of H. hoffmanni — IGUP 78 (PI. 10, figs. 1-3).
Genus NEOCRIMITES Ruzhencev, 1940a
Diagnosis. — Neocrimites includes narrowly umbilicate adriani-
tids in which the “mature” suture on each side has four or five lobes
between the bifid ventral lobe and the umbilical shoulder, three to
five lobes between the dorsal lobe and the umbilical seam, and one
or two lobes on the umbilical wall.
Synonymy. — Neocrimites (Metacrimites) Ruzhencev, 1950,
and Neocrimites (Neocrimites) were considered consubgeneric by
Glenister and Furnish (1961, p. 726, 727); Aricoceras (Aricoceras)
Ruzhencev, 1950, and A. (Metaricoceras) Ruzhencev, 1950, were
referred to Neocrimites by the same authors (ibid.).
Specific composition.—The type species of Neocrimites is
Adriamites fredericksi Emeliancev, 1929, by original designation.
52 BuLLeTIN 272
Adrianites newelli Miller and Furnish, 1940a, the type species, and
Adrianites adamsi Miller and Furnish, 1940a, were the only taxa
referred to Neocrimites (Metacrimites) by Ruzhencev (1950, p.
202). In addition to the type species, Ruzhencev (1950, p. 202;
1956a, p. 255) included the following in Neocrimites (Neocrimites):
Adrianites defordi Miller and Furnish, 1940a, A. dunbari Miller and
Furnish, 1940a, Waagenia koswae Krotow, 1885, Adrianites mara-
thonensis Bose, 1919, A. nalivkini Toumanskaya, 1949, A. plummeri
Miller, 1944, Agathiceras stuckenbergi Karpinsky, 1889, and Adria-
mites warrem Miller and Crockford, 1936. Miller and Furnish (1940a,
p. 116), however, believed that the type of A. marathonensis might
be only an immature Waagenoceras dienert. Ruzhencev (1950, p.
203) included the following in Aricoceras (Aricoceras): Adriamtes
ensifer Gemmellaro, 1887, the type species, Agathiceras cancellatum
Haniel 1915 (sensu Haniel, p. 75), and Adrianites (Neocrimites)
meridionalis Teichert and Fletcher, 1943. Agathiceras martini var.
globosa Haniel, 1915, is apparently a synonym of Neocrimites can-
cellatum (Furnish — personal communication, 1965 notes). The
type species and only taxon which has been referred to Aricoceras
(Metaricoceras) is Agathiceras cancellatum form. discoidalis Haniel,
1915, although the suture of Agathiceras cancellatum form. globosa
Haniel, 1915 (pl. 50, fig. Ic) agrees with Ruzhencev’s (1950, p.
203) diagnosis of the subgenus. Furnish and Glenister (in Davis,
Furnish, and Glenister, 1969, explanation to plate 3) included
Adrianites insignis Gemmellaro, 1887, in Neocrimites.
Immature shell. —The conch form of Neocrimites has been
described as spherical to subdiscoidal (Glenister and Furnish, 1961,
p. 728). The conch is involute (zbid.) to comparatively involute
(Ruzhencev, 1950, p. 202).
At least some individuals of Neocrimites bear transverse con-
strictions. These constrictions are irregularly spaced, which may
account for their apparent lack in some individuals. The amplitudes
of these features are not uniform, nor do the variations seem to
follow any regular pattern. The outermost preserved whorl of one
specimen (one of IGUP 82) has nine transverse bands in which
the growth lines are disturbed; these apparently reflect constric-
tions and, if so, constitute the largest number observed by the
present author in one whorl. Other than in this specimen, the
PaLEozoic AMMONOIDs: DAvis 53
maximum number of constrictions observed per volution is four. The
constrictions run from shoulder to shoulder.
The shell of Neocrimites bears both longitudinal and transverse
lirae, both of which in some individuals are so prominent as to be
plications. The relative prominence of the transverse and longi-
tudinal ornament varies. The spiral ornament of some specimens
tends to fade at constrictions, and the growth lines are more closely
spaced there. Along the umbilical shoulder of some representatives
of the genus there runs a thickening of the shell layer (Ruzhencev,
1941, p. 884).
Mature shell — After nearly three-quarters of the ultimate whorl
of Neocrimites had been formed there was a slight decrease in the
radius of curvature of coiling, followed by a slightly greater increase.
Just prior to the formation of the ultimate constriction there was
another decrease in the radius of curvature which led directly into
the constriction. In the majority of known specimens all of these
changes in coiling are slight.
The trend of the ultimate constriction of Neocrimites differs
from that of those adapical of it. This terminal constriction is devoid
of longitudinal ornament.
After almost all of the ultimate constriction of Neocrimites had
been formed the ventrolateral portions of each successive growth
line formed a larger salient, so that the ultimate peristome has one
prominent ventrolateral lappet on each side, each bearing a branch
of the terminal constriction. These lappets curve toward the dorsum.
Neocrimites sp. of Glenister and Furnish (1961, pp. 729-731)
was said to have a terminal flare adapertural of the ultimate con-
striction. The internal mold of the living chamber of Neocrimites
dunbari was said to bear a small, low, rounded ridge or raised line
along the venter of the mature living chamber (Miller, 1944, p. 99).
Gemmellaro (1887, p. 46) stated that the living chamber of
Neocrimites ensifer is one and one-half whorls long. Neocrimites sp.
of Glenister and Furnish (1961, pp. 729-731) was said to have a
body chamber only one whorl long. In SUI 32443 the living chamber
is no longer than one volution, and in SUI 32605 it is no more
than one and one-quarter whorls long.
The smallest mature individual of Neocrimites known to the
author is 27 mm in diameter (viz. N. sp. of Glenister and Furnish,
54 BULLETIN 272
1961). The largest (SUI 32443) is not yet mature at a diameter of
40 mm.
Material. — The preceding observations are based on study of:
Gemmellaro (1887), Haniel (1915), Bose (1919), Wanner (1932),
Miller and Crockford (1936), Miller and Furnish (1940a), Ruzhen-
cev (1940a, 1941, 1950, 1956a), Miller (1944), Toumanskaya
(1949), Glenister and Furnish (1961), Bogoslovskaya (1962), Arm-
strong, Dear, and Runnegar (1967), Davis, Furnish, and Glenister
(1969), about one hundred immature Neocrimites (N. sp. — GPIT
unnumbered, SUI 32444-32448, 32221, 32601-32603, defordi — SUI
1398, discoidalis — SUI 32450, dunbari — SUI 1360, 1763-1765,
13595, 32443, fredericksi — SUI 32449, 32452, 32543, globosus —
SUI 32451, newelli — SUI 1361, 1478, 13596), eight mature indi-
viduals (SUI 32443, 1360, 32600, 32604-32606), and photographs
and notes taken by W. M. Furnish and Brian F. Glenister in the
summer of 1965.
Illustrations. — The following mature specimens of Neocrimites
are figured: N. cancellatus (fide Furnish and Glenister in Davis,
Furnish, and Glenister, 1969) — PIUB unnumbered (PI. 10, figs.
4,5), GIUA T328 (PI. 3, figs. 8, 9), N. ensifer — IGUP 82 (PI. 7,
figs. 1-3), and (PI. 7, figs. 4, 5). In addition, one immature specimen
is illustrated — N. ensifer — IGUP 82 (PI. 6, figs. 1, 2).
Genus PALERMITES Toumanskaya, 1937a
Diagnosis. — Palermites includes moderately evolute adrianitids
in which the “mature” suture on each side has five lobes between
the bifid ventral lobe and the umbilical shoulder, five lobes between
the dorsal lobe and the umbilical seam, and one or two lobes on the
umbilical wall.
Specific composition. — The type species of Palermites is
Adriamtes distefanot Gemmellaro, 1887, by original designation. In
addition, Adrianites distefanoi var. minor Toumanskaya, 1931, is
referred to Palermites.
Immature shell.—The conch of Palermites is not broad, is
moderately involute, and has a large umbilicus (Ruzhencev, 1950, p.
203):
Every specimen of Palermites known to the author bears trans-
verse constrictions, concave outward on both the shell exterior and
the steinkern. Most individuals have three constrictions per whorl.
PaLeozoic AMMONOIDS: DAVIs 55
Each constriction continues from umbilical seam to umbilical seam.
Longitudinal lirae are restricted to the umbilical region but are
absent from the umbilical wall. The lira which is most prominent
is situated on the shoulder. On the more ventral portions of the
conch, longitudinal ornament is present only as scallops in the trans-
verse lirae. Transverse lirae are crowded slightly adoral of each con-
striction.
Mature shell. — After slightly more than half of the ultimate
whorl of Palermites was formed there was a slight decrease in the
radius of curvature of coiling followed by a slightly greater increase;
this change in coiling is evident in the course of the umbilical seam
but does not appear to have affected the venter. After about three-
quarters of the last volution was formed, there was another pair of
changes in coiling similar to the previous but affecting both the
seam and the venter. Coupled with the latter changes in coiling was
a gradual increase in the ratio W/H, so that the venter appears
somewhat flattened both in longitudinal and transverse section.
Close to the ultimate constriction the height increased relative to the
width, so that the venter again took on a smaller radius of curvature
in transverse section; there is also a tightening of coiling leading
directly into the ultimate constriction.
The terminal constriction of Palermites is marked by a gentle
ridge which parallels the trend of the constriction. From its broadest
point on the venter the constriction becomes narrower and shallower
toward the umbilical shoulders.
Ventrolateral lappets in Palermites were commenced after al-
most all of the ultimate constriction was formed. These lappets are
shorter and more delicate and appear to be more strongly oblique
than in other genera of the family.
The living chamber of Palermites distefanoi is considerably
longer than the ultimate whorl (Gemmellaro, 1887, p. 48).
Two mature individuals of Palermites are known to the author,
both from Sicily. Their diameters are 30% mm and 45 mm.
Material. — The preceding observations are based on study of:
Gemmellaro (1887), Toumanskaya (1931, 1937a), Ruzhencev
(1950), Glenister and Furnish (1961), about a dozen immature
individuals (USNM unnumbered), three other immature speci-
mens (SUI 32455), one mature example (SUI 32456), and notes
56 BULLETIN 272
and photographs taken by W. M. Furnish and Brian F. Glenister
in the summer of 1965.
Illustrations. —Two mature specimens of Palermites distefanot
are figured: SUI 32456 (PI. 9, figs. 3-5) and IGUP 76 (PI. 9, figs.
Sa fy
Genus PSEUDAGATHICERAS Schindewolf, 1931
Diagnosis. — Pseudagathiceras includes large-umbilicate adriani-
tids in which the “mature” suture is straight in overall trend and
on each side has three or four lobes between the bifid ventral lobe
and the umbilical shoulder, three lobes between the dorsal lobe
and the umbilical seam, and one or two lobes on the umbilical wall.
Specific composition. — The type species of Pseudagathiceras is
Agathiceras (Doryceras?) wichmanni Haniel, 1915, by original des-
ignation. In addition, P. difuntense Miller, 1944, and P. spinosum
Miller, 1944, are referred to Pseudagathiceras.
Immature shell. — The shell of Pseudagathiceras has been de-
scribed as very broad and evolute or moderately involute, with a
fairly large umbilicus (Ruzhencev, 1950, p. 203). The umbilical
shoulders are rounded.
Every specimen of Pseudagathiceras known to the author with
the possible exception of the holotype of the type species bears
transverse constrictions. These extend from umbilical seam to um-
bilical seam. The maximum number of constrictions observed in
any one whorl is five; the majority have but three per volution,
but these are smaller specimens.
Pseudagathiceras, with the possible exception of the holotype
of the type species, bear irregularly spaced transverse plications.
The plications are stronger near the umbilicus, so much so that
some individuals appear nodose.
All specimens of Pseudagathiceras in which shell is preserved
bear longitudinal lirae. These lirae seem to fade out on the umbilical
walls. In most specimens the transverse lirae are much less promin-
ent than the longitudinal ornament, but in one specimen available
to the author the lirae in the two directions are equally distinct.
Mature shell. — After about two-thirds of the ultimate whorl
was formed there was a slight decrease in the radius of curvature
of coiling followed by a slightly greater increase in the radius of
coiling. Just prior to the formation of what may be an apertural
PaLeozoic AMMONOIDs: DAvis Ly
constriction there was a decrease in the radius of curvature of coil-
ing. The umbilicus displays only a gradual increase in the radius of
curvature followed by decrease in the vicinity of the apertural (?)
constriction, There seems to be a constriction or particularly prom-
inent plication just where the first change in coiling occurs.
The transverse plications are more prominent on the ultimate
living chamber of Pseudagathiceras than on the phragmocone. In
the adapical half of the living chamber certain of the plications
bear a ventrolateral spine on each side. Not every plication is so
favored, and the spines do not seem to be regularly positioned on
the circumference. In the adapertural half of the living chamber
no spines were formed, but the plications are prominent.
Miller (1944, pp. 101, 103) reported that the living chamber
of Pseudagathiceras is almost one whorl long.
The maximum diameter (exclusive of spines) of the Pseudaga-
thiceras figured by Miller (1944, pl. 29, figs. 1, 2) is 21 mm,
Material. — The preceding observations are based on study of:
Haniel (1915), Miller (1944), Ruzhencev (1950), Glenister and
Furnish (1961), about one and one-half dozen immature specimens
of Pseudagathiceras difuntense (SUI 32457, 1363, 1362, 1770, 13621),
and photographs and notes taken by W. M, Furnish and Brian F.
Glenister in the summer of 1965.
Illustrations. — One mature specimen of Pseudagathiceras spin-
osum is figured — YPM 16310 (PI. 9, figs. 1, 2). One immature ex-
ample of P. wichmanni is also illustrated —THD 12752 (PI. 10,
figs. 6-8).
Genus SIZILITES Toumanskaya, 1937a
Diagnosis. — Sizilites includes large-umbilicate adrianitids in
which the “mature” suture is arcuate in overall trend and on each
side has four lobes between the bifid ventral lobe and the umbilical
seam and has four lobes between the dorsal lobe and the seam.
Specific composition. — The type species of Sizilites is Adrianites
affinis Gemmellaro, 1888, by original designation. In addition, A.
craticulatus Gemmellaro, 1888, is referred to the genus.
Immature shell.—The conch of Sizilites is described as not
wide, moderately involute, and with a large umbilicus (Ruzhencevy,
1950, p. 204). The umbilical shoulder has a large radius or is absent.
Every specimen of Sizilites known to the author bears two to
58 BULLETIN 272
four constrictions in the outermost preserved whorl. These constric-
tions tend to fade out on the venter and on the umbilical wall por-
tions of the conch, but they are prominent elsewhere.
In some individuals of Sizilites longitudinal lirae occur on all
but the umbilical wall portions of the shell, but in other specimens
these lirae fade on the venter to become merely scallops in the trans-
verse lirae. The longitudinal ornament fades at constrictions.
Mature shell. — After about one-half of the ultimate whorl of
Sizilites was formed there was a slight decrease in the radius of
curvature of coiling followed by a slightly greater increase. After
about two-thirds of the whorl was formed there occurred another
pair of changes in coiling similar to the first pair. These latter
changes were apparently followed by a gradual relative increase in
width, Just prior to the formation of the apertural constriction a
decrease in the radius of curvature of the coiling occurred, leading
directly into the constriction.
The apertural constriction of Szzilites is much more prominent
and of a somewhat different trend from those adapical of it.
The longitudinal ornament ceases about one-quarter whorl
adapical of the apertural constriction. The growth lines, on the
other hand, become more prominent, to the extent of becoming
plications in the last quarter volution.
The mature living chamber of Stzilites craticulatus was stated
by Gemmellaro (1888, p. 15) to be one and one-half whorls long.
The two mature specimens of Sizilites known to the author are
each 154 mm in diameter.
Material. — The preceding observations are based on study of:
Gemmellaro (1888), Toumanskaya (1937a), Ruzhencey (1950),
Glenister and Furnish (1961), seventeen immature specimens (six
USNM unnumbered, five SUI 32454, and six GPIT unnumbered),
one mature individual (GPIT unnumbered) and photographs and
notes taken by W. M. Furnish and Brian F. Glenister in the sum-
mer of 1965.
Illustrations. —One mature specimen of Sizilites craticulatus is
figured: IGUP 87 (Pl. 11, figs. 1, 2). Two immature examples of
Sizilites are also illustrated: S. craticulatus — MGPU unnumbered
(Pl. 11, figs. 8-10) and S. affims — IGUP 88 (PI. 11, figs. 3-5).
PaLEozoic AMMONOIDs: DAvIs 59
Genus TEXOCERAS Miller and Furnish, 1940a
Diagnosis. — Texoceras includes adrianitids in which the “ma-
ture” suture on each side has three complete lobes between the bifid
ventral lobe and the umbilical seam, two complete lobes between
the dorsal lobe and the seam, and one lobe on the seam.
Specific composition. — The only species which has been re-
ferred to Texoceras is Agathoceras [sic] texanuwm Girty, 1908. This
species was made the type species of the genus by original designa-
tion. Marathonites delawareanus Plummer and Scott, 1937, is a
synonym (fide Miller and Furnish, 1940a, p. 111).
Immature shell.— The conch of T'exoceras is sublenticular to
thickly subdiscoidal and is deeply involute, with an open, moderately
small umbilicus (Miller and Furnish, 1940a, p. 111).
Most individuals of Texoceras examined display no constric-
tions. Two specimens (SUI 32462, 32461) have four constrictions
per whorl at a diameter of about 6 mm; but another example (SUI
32437) displays no constrictions on a whorl of the same diameter.
Available larger specimens lack constrictions except at the ultimate
peristome.
The growth lines of T’exoceras are prominently scalloped; the
points of these scallops extend orad to form discontinuous longi-
tudinal lirae. This spiral ornament is absent from the umbilical walls.
Each growth lira lies at the crest of a gentle plication; these plica-
tions, as displayed on the internal mold, are crenulate. Each side
of the conch bears about ten coarse transverse plications per whorl;
these are restricted to the dorsolateral portions of the conch and are
quite distinct from the smaller plications which bear growth lines.
The larger plications are not everywhere evenly spaced.
The internal molds of all Texoceras living chambers known to
the author display a gentle longitudinal ridge in mid-ventral posi-
tion. In some individuals (for example, SUI 32461 R, Z, and AE)
this ridge extends the full length of the living chamber; in others
(such as SUI 32461 N and AF, and 13609) it is discontinuous. Only
a few suitably preserved phragmocones were examined by the
author; all of them seem to bear the ventral longitudinal ridge on
the internal mold. Miller and Furnish (1940a, p. 111) stated that
this ridge is present only on some specimens of Texoceras.
Mature shell.— After about three-quarters of the ultimate
60 BULLETIN 272
whorl of Texoceras had been formed there occurred a slight decrease
in the radius of curvature of coiling, followed by a slightly greater
increase; this resulted in a flattening of the venter of the last quarter-
volution as viewed laterally. Just before the ultimate peristome was
reached another decrease in the radius of curvature of coiling oc-
curred, leading directly into an apertural constriction. These changes
in coiling are also visible in the umbilical region.
The ornament of the immature portion of the shell continues to
the adapertural end of the mature living chamber. In most indi-
viduals the apertural constriction is relatively deep and distinct.
In SUI 32461 M and AG, however, it is relatively shallow, but the
peristome is flared and thus apparently mature.
As pointed out by Miller and Furnish (1940a, p. 113), the
inner surface of the test of Texoceras is pustulose; this has produced
pits in the surface of the steinkern. The pits (and, hence, the
pustules) appear to have no regular arrangement and, indeed, vary
as to size and spacing from individual to individual, Every known
specimen with a distinct apertural constriction bears these features;
they seem to be most distinct and closely spaced in the ultimate
constriction, fading apically so that they are apparently restricted
to the ultimate one-quarter whorl. In the two specimens with in-
distinct constrictions (SUI 32461 M and AG), the pits in the in-
ternal mold appear to be present only in patches in the vicinity of
the aperture (2.e. for about % whorl).
The ultimate peristome of Texoceras is immediately adjacent
and parallel to the ultimate constriction. In SUI 32461 P and Z the
peristome is apparently scalloped and is situated approximately %
mm from the constriction.
The last few septa of Texoceras are approximated, and the
characteristically rounded lobes become shallower and more pointed.
The ultimate living chamber of Texoceras varies from seven-
eighths of one whorl to just under one volution in length.
In 15 mature specimens of Texoceras the diameter varies from
a minimum of 22 mm to a maximum of 29 mm, with the majority
clustering toward the low end of the range,
Material. — The preceding observations are based on study of:
Girty (1908), Plummer and Scott (1937), Miller and Furnish
(1940a), Ruzhencev (1950), Glenister and Furnish (1961), Bogo-
PaLteozoic AMMONOIDs: Davis 61
slovsky, Librovitch and Ruzhencev (1962), about 40 immature in-
dividuals, and about 40 mature specimens (SUI 13609, 32435-
32438, 32461, and 32462).
Illustrations. — Eight mature specimens of Texoceras texanwm
are figured: SUI 13609 (PI. 11, fig. 15), 32461 I (PI. 11, fig. 12),
32461 M (PI. 11, fig. 7), 32461 P (PI. 11, figs. 13, 14), 32461 Z (PI.
11, fig. 6), 32461 AA (PI. 10, fig. 9), 32461 AG (PI. 11, fig. 11), and
32461 AJ (PI. 10, fig. 10).
Superfamily AGATHICERATACEAE Arthaber, 1911
Family AGATHICERATIDAE Arthaber, 1911
Diagnosis. —The Agathiceratidae comprises ammonoids_ in
which the suture underwent goniatitic ontogenetic development;
following this, the adolescent lateral lobe became trifid, but the in-
ternal lobe remained undivided.
Genus AGATHICERAS Gemmellaro, 1887
Diagnosis. — Agathiceras comprises agathiceratids with “ma-
ture” sutures in which the three subdivisions of the primary lateral
lobe are discrete and subequal in size.
Synonymy. — Agaticeras Gregorio, 1930, and Agathoceras
Girty, 1908, are apparently merely typographic errors.
Taxonomic composition. — Ruzhencev (1950) recognized three
subgenera: Agathiceras (Agathiceras), A. (Gaetanoceras) Ruzhen-
cev, 1938, and A. (Paragathiceras) Ruzhencev, 1950. Glenister and
Furnish (1961) questioned the validity of the latter two but appar-
ently revised their opinion in Davis, Furnish, and Glenister (1969).
The type species of Agathiceras is A. swessi Gemmellaro, 1887,
by original designation. In addition, the following are referred to
Agathiceras: A. anceps Gemmellaro, 1887, A. anceps var, kitschiensts
Toumanskaya, 1931, A. anceps var. sarabansis Toumanskaya, 1931,
A. applanatum Teichert, 1944, A. applini Plummer and Scott, 1937,
A. asiaticum Toumanskaya, 1935, A. asselicum Ruzhencev, 1938
(junior synonym of A. uralicum - fide Ruzhencev, 1950), A. beschui
Toumanskaya, 1931, A. bodraki Toumanskaya, 1931, A. brouweri
Smith, 1927, A. ciscoense Smith, 1903, A. contractwm Plummer and
Scott, 1937, A. frechi Bise, 1919, A. frechi var. ircutensis Toumans-
kaya, 1949. A. girtyi Bose, 1919, A. katschae Toumanskaya, 1931,
A. martini Haniel, 1915, A. planum Toumanskaya, 1931, A. sundat-
62 BULLETIN 272
cum Haniel, 1915, A. tornatwm Gemmellaro, 1887, and Gontatites
uralicus Karpinsky, 1874. A number of other taxa, inadequately de-
scribed, may also belong in Agathiceras: Gomiatites fultonensis Mil-
ler and Gurley, 1896, Ammonites hildrethi Morton, 1836, Agathiceras
kinshanensis Grabau, 1924, Agathiceras mediterraneum ‘Youmans-
kaya, 1949, and Agathiceras tegengrem Grabau, 1924.
Furnish (personal communication, 1965) concluded that the
holotype of Agathiceras martini var. globosa Haniel, 1915, is an
adrianitid, as are the specimens described and figured by Wanner
(1932
Immature shell. —Glenister and Furnish (1961, p. 695)
described the conch form of Agathiceras as generally ellipsoidal in
cross-section but noted that some individuals are parabolic or
circular. W/D decreased in ontogeny, and Dixon (1960, p. 54) con-
cluded that the seven species he studied (A. applint, A. frechi, A.
girtyi, A. suesst, A. sundaicum, A. tornatum, and A. uralicum) can
best be differentiated on the basis of differences in the rates of onto-
genetic change of W/D.
The constrictions of the phragmocone of Agathiceras are pre-
dominantly internal thickenings of the shell but are expressed as
depressions on the shell exterior of some individuals. As many as
six constrictions have been observed in a single whorl, but some in-
dividuals (for example, A. applanatum of Glenister and Furnish,
1961, p. 696, and “A. asselicum” of Ruzhencev, 1938, p. 263) ap-
parently bear no constrictions. There does seem to have been an
ontogenetic increase in the number of constrictions per whorl.
Except for Agathiceras martini, the genus is characterized by
longitudinal lirae which are present on the ventral and lateral por-
tions of the conch and even on the umbilical wall; at least some
individuals bear traces of these “lirae” on the internal mold. The
number of such lirae per individual seems to have remained essen-
tially constant throughout ontogeny, and the lirae are evenly spaced,
This longitudinal ornament is present as far back as the second
whorl.
The siphuncle of Agathiceras was not ventral in the earliest
growth stages, so that the earliest formed sutures lack the bifurca-
tion of the ventral lobe. The umbilical lobe of the “mature” sutures
resulted from an ontogenetic coalescence of the two lobes adjacent
PaLtEozoic AMMONoOIDs: Davis 63
to the early adolescent umbilical seam. At “maturity” the prongs
of the ventral lobe are about the same size as the lateral lobes and
are straight.
Mature shell. — After two-thirds to three-quarters of the length
of the mature living chamber of Agathiceras had been formed, a
slight decrease in the radius of curvature of coiling occurred, followed
by a slightly greater increase. This resulted in a flattening of the
venter as viewed in longitudinal section. In at least some forms
(for example, A. swesst) the changes in coiling were accompanied
by a distinct increase in the whorl width; this bulging is greatest
on the dorsolateral portions of the flanks but is discernible across
the venter where it resulted in a slight increase in the radius of
curvature of the venter as viewed in transverse section. In at least
some individuals (for example, many SUI 32460) the umbilical
wall portion of the shell just adapical of the mature peristome flares
dorsad, resulting in a decrease in the size of the umbilicus.
In Agathiceras suessi each flank bears a deep depression about
one-quarter whorl adapical of the adaperturalmost constriction.
The position of the deep depression corresponds to the change in
spiral and the initiation of the lateral bulge. These depressions affect
both the shell exterior and the internal mold more profoundly than
any ordinary constriction, are elongate in a radial direction, and oc-
cupy about the middle third of the flanks. Other species of the genus
apparently lack these depressions: one specimen of A. sp. (SUI
12297) has an ordinary constriction in a position corresponding to
that of the depressions of A. swessi, but over one hundred individuals
of A. uralicum (fide Furnish and Glenister in Davis, Furnish, and
Glenister, 1969) have neither constrictions nor depressions between
the apertural constriction and slightly less than one-half whorl
adapical of it (SUI 32460),
The adapertural constriction of Agathiceras has a trend some-
what different from that of all the constrictions preceding it; more-
over, the adapertural constriction has a greater amplitude, strongly
affecting both the shell exterior and the internal mold, In Agathi-
ceras suesst there is a distinct ventrolateral bulge on each side im-
mediately adapical of the ultimate constriction and corresponding in
position to each ventrolateral Jappet of the terminal peristome; in
64 BULLETIN 272
the constriction itself there is one pit on each side just ventral of
the ventrolateral bulge.
At least some specimens possess small raised points on the shell
interior. These points seem to lie in definite radial rows near the
umbilicus, whereas their arrangement is random on the flanks and
venter, The significance of these features is not apparent at present.
The mature living chamber of Agathiceras is seven-eighths of a
whorl to one whorl in length.
The mature diameters of seven Agathiceras sp. are 16, 21, 23,
25, 33, 55, and about 55 mm. The maximum diameters of seven
Agathiceras suessi are 20, 35, 36, 37, 37, about 40, and 42 mm. One
mature A. wralicum is 33 mm in diameter.
A sample (SUI 32460) consisting of 110 mature specimens of
Agathiceras uralicum (fide Furnish and Glenister in Davis, Furnish,
and Glenister, 1969) from one horizon in the Road Canyon Forma-
tion at one location in the Glass Mountains region of Texas has been
studied statistically. Within this population of 110 specimens two
distinct size groupings are recognizable (see Davis, Furnish, and
Glenister, 1969):
1. one group of 28 individuals with an average diameter of
20.7 mm (as calculated from measurement of 19 specimens),
and
2. another group of 82 individuals with a mean diameter of
30.0 mm (as calculated from measurement of 38 specimens).
Conch proportions, ornament, and mature modifications in both
groups appear the same, Poor internal preservation precludes onto-
genetic study and even the determination of the number of whorls.
Material. — The preceding observations are based on study of:
Morton (1836), Karpinsky (1874, 1889), Krotow (1885), Gem-
mellaro (1887, 1888), Miller and Gurley (1896), Haug (1898),
Smith (1903, 1927), Girty (1908), Haniel (1915), Bose (1919),
Fredericks (1921), Grabau (1924), Gregorio (1930), Toumanskaya
(1931, 1935, 1939b, 1939c, 1949), Plummer and Scott (1937),
Ruzhencev (1938, 1950, 1951, 1962), Miller and Furnish (1939,
1940a, 1957b), Miller (1944, 1945a), Teichert (1944), Miller and
Youngquist (1947), Gerth (1950), Miller and Downs (1950),
Glenister and Furnish (1961), Unklesbay (1962), Hayasaka (1963,
1965), Chao (1965), Davis, Furnish, and Glenister (1969), about
PaLeozoic AMMONOIDs: Davis 65
1500 immature specimens (Agathiceras sp. — SUI unnumbered,
applini — SUI 1386, 1766, 1768, 13612, browwers — SUI 32429,
ciscoense — SUI 13764, 17025, frechi — SUI 13765-13767, 32420,
32421, girtyi — SUI 1400, 1966, 2126, 13623, 32418, 32419, swesst —
GPIT unnumbered, SUI 8297, 12613, 32424, 32426-32428, 32432,
32433, USNM unnumbered, F. Stehli loan, swndaicwm — SUI 13615,
tornatum — SUI 32423, USNM unnumbered, wralicum — SUI
1387, 1767, 10567-10570, 13611, 32413-32415, 32417), about 250
mature Agathiceras (A. sp. — BMNH C73338, GPIT 24543, GPIT
PV24409, SUI 12297, 32432, 32458, F. Stehli loan, swesst — GPIT
24543, SUI 32433, USNM unnumbered, F. Stehli loan, wralicum —
SUI 10569, 32460), and photographs and notes taken by W. M.
Furnish and Brian F. Glenister in the summer of 1965.
Illustrations. — Eight mature specimens of Agathiceras are
figured: two A. sp. — SUI 12297 (PI. 14, fig. 9), SUI 32458 (PI.
13, figs. 8, 9), three A. swesst — IGUP 119 (PI. 12, figs. 1-4, Pl. 12,
figs. 7, 8), and MGPU unnumbered (PI. 12, figs. 5, 6), three A.
uralicum (fide Furnish and Glenister in Davis, Furnish, and Glenis-
ter, 1969), SUI 32460 BA (PI. 13, figs. 1,2), SUI 32460 BE (PI. 13,
figs. 5-7), SUI 32460 BN (PI. 13, figs. 3, 4). One immature speci-
men of A. martint is also illustrated — THD 12735 (Pl. 14, fig. 8).
Superfamily CYCLOLOBACEAE Zittel, 1903
Diagnosis. — The Cyclolobaceae includes those ammonoids with
sutures that underwent goniatitic development, following which the
umbilical, internal lateral, and dorsal lobes each split into three, and
the lateral lobe split into three or two resulting in the formulas:
CVU PAL hi Uy Unt, te Ti DiDyD,.)
pl aL Pipe) eR eee ge eg ey et Fe Py al
or
Can We Illa (DD LD )
sae | ee Dae Die ee ee Ze le &2
In the first case, subsequent new elements were added by successive
bifurcation of that subdivision of the lateral lobe nearest the um-
bilicus, according to the scheme:
a Ginn Gale oh Snes Ele EL (LE 18 ) ete.
re: hey en marie? 1
-1 2-1-1 2-1-
In the second case the subdivision of the lateral lobe nearest to the
66 BuLLeETIN 272
ventral lobe bifurcated once; other subsequent new elements were
added by successive bifurcation of the subdivision of the lateral lobe
nearest the umbilicus.
Family CYCLOLOBIDAE Zittel, 1903
Diagnosis. — The Cyclolobidae includes cyclolobaceans with a
“mature” suture in which the lateral and internal lobes are denticu-
late both at their bases and also on the sides.
Genus MEXICOCERAS Ruzhencev, 1955
Diagnosis. — Mexicoceras includes cyclolobids in which initial
subdivision of the primary lateral lobe was through bifurcation; the
“mature” suture includes five or six pairs of digitate external lateral
lobes.
Specific composition.— The type species of Mextcoceras is
Waagenoceras cummingsi var. guadalupense Girty, 1908, by original
designation (W. clavatum Plummer and Scott, 1937, is a synonym).
This is the only species which has been referred to the genus aside
from MV. globosum Chao, 1965, a nomen nudum.
Miller and Furnish (1940a) recognized three sympatric sub-
species of Mexicoceras guadalupense: W. guadalupense guadalupense
Girty, 1908, W. guadalupense smithi Miller and Furnish, 1940a, and
W. guadalupense thompson Miller and Furnish, 1940a.
Immature shell.—The conch of Mexicoceras is subspherical
(Bogoslovsky, Librovitch, and Ruzhencev, 1962). The three sub-
species of the type species were defined mainly on differences in
conch proportions (Miller and Furnish, 1940a): Mexicoceras guada-
lupense smitht has W greater than D in “moderate sized specimens”;
M. g. thompson has W about two-thirds of D in “moderate sized
specimens”; in the typical subspecies W/D is between the ratios of
the other two in “moderate sized specimens”. Miller and Furnish
(1940a) stated that the three subspecies are completely inter-
gradational and that “large individuals” have a greater uniformity
in conch proportions than “moderate sized individuals”.
The majority of individuals of Mewxicoceras examined by the
author have at least one constriction visible in the outermost pre-
served whorl. There does not seem to be a regular pattern for the
spacing of constrictions. In SUI 1782, for example, the first con-
striction occurs one whorl beyond the protoconch, at a diameter of
PaLeozoic AMMoNoIDs: Davis 67
0.9 mm; the next constriction is about 255° further, at a diameter
of 1.4 mm; the third is about 780° (2 whorls + 60°) further, at a
diameter of 3.8 mm; and the fourth is about 175° further, at a
diameter of 4.8 mm. Apparently the lack of constrictions in the
outermost preserved whorl of some individuals may be merely a
function of irregular spacing of constrictions, rather than an actual
lack of constrictions,
The shell is thicker at a constriction than between constrictions,
but constrictions as observed in median longitudinal sections of in-
dividuals of Mexicoceras vary considerably. In some instances (for
example, the first constriction in SUI 1782) both the shell exterior
and the internal mold are concave outward at the venter. At other
constrictions (such as the second constriction of SUI 1782) the
shell exterior is convex outward, and the internal mold seems affected
little if any. At the third constriction of SUI 1782 the shell exterior
is virtually unaffected, but the internal mold is concave outward.
And at the fourth constriction of SUI 1782 the internal mold is con-
cave outward at the venter, but the shell exterior appears to be
slightly convex outward.
In the majority of examined specimens which seem to fall with-
in Mexicoceras guadalupense smithi (for example, most of the SUI
12612 series) immediately adapical of each definite constriction are
first a ridge and then a groove, both of which parallel the constric-
tion. The groove is more distinct near the umbilicus than on the
venter, where it may disappear entirely; the groove is everywhere
more shallow than the immediately adjacent constriction.
In general a constriction is shallower and broader at the venter
than at the umbilicus.
Most well-preserved individuals of Mexicoceras display trans-
verse undulations in the internal mold. Some of these undulations
are due to shell plication, but some are also due to thickenings in
the shell. The undulations (hence, the plications, the shell thicken-
ings or both) are irregularly spaced and of different amplitudes,
but there does not seem to be a regular pattern for these differences
in amplitude. As with constrictions, these undulations are commonly
less prominent at the venter than near the umbilicus.
Those ammonoids which Miller and Furnish (1940a) referred
to Mextcoceras guadalupense smithi are characterized by nodes on
68 BULLETIN 272
the umbilical shoulders, which are longitudinally elongate. In plicate
individuals not every plication has a node. In some individuals (for
example, SUI 12612AE) only the inner whorls bear nodes, and no
individual of the species of diameter larger than 25 mm has been
observed to bear nodes. In SUI 12612AB the nodes lie slightly ven-
tral of the shoulder so that this individual seems to have two
shoulders on each side.
Growth lines in Mexicoceras parallel] the trend of the closest
constriction.
Mature shell.— The approach to maturity in Mewicoceras is
heralded by a slight decrease in the radius of curvature of coiling fol-
lowed by a slightly greater increase; this occurs about two-thirds
of the way through the ultimate whorl. These changes in coiling
appear to coincide with a change in conch proportions: the whorl
height decreases in the last third of the living chamber, whereas the
width continues to increase.
Miller and Furnish (1940a, p. 163) reported that mature
Mexicoceras guadalupense has a constriction just adapical of the
ultimate peristome. Seven specimens available to the author display
a body chamber which terminates in a constriction. Miller and
Furnish further stated (ibid., p. 161) that specimens of the species
rarely if ever have constrictions on the fully mature portion of the
conch except at the aperture. Of the seven specimens only one
(Texas-k) has a constriction in the living chamber other than that
at the apertural end; in this individual the second constriction is
about 230° adapical of the first. Texas-k’s adapertural constriction
apparently lacks the ventral salient present in the apertural constric-
tions of the other six individuals.
No individual examined possesses a recognizable ultimate
peristome. Growth lines and plications in the shell of the living
chamber of SUI 12614 indicate that the ventral salient developed
late in the ontogeny.
There is a decrease in the septal interval towards the mature
body chamber. The ultimate septum of a mature individual is
distinctly thicker than those adapical of it.
Six of the seven individuals with the apertural constriction
preserved have living chambers slightly less than one whorl] in
length; the body chamber of Texas-k is about one and one-quarter
volutions in length. Plummer and Scott (1937, p. 161) reported
PaLeozoic AMMONOIDs: DAvIs 69
specimens of Waagenoceras clavatum having living chambers longer
than one volution and with the last three or four septa crowded.
The significance of Plummer and Scott’s specimens and of Texas-k
is unclear.
The maximum diameter of mature individuals ranges from 48
mm (USNM 144423) to at least 200 mm (unidentified individual
mentioned by Miller and Furnish, 1940a, p. 163).
Material. — The preceding observations are based on study of:
Girty (1908), Bése (1919), Diener (1921), Plummer and Scott
(1937), Miller and Furnish (1940a), Miller and Unklesbay (1943),
Miller (1944), Ruzhencev (1955, 1956b), Bogoslovsky, Librovitch,
and Ruzhencev (1962), Chao (1965), Furnish (1966), Davis,
Furnish, and Glenister (1969), Furnish and Glenister (1970), about
70 immature individuals (AMNH unnumbered, SUI 1781, 1782,
12609, 12610, 12612, 12616-12619, 13598, Univ. Texas unnumbered,
USNM 144419, 144422, 144437) and about ten mature specimens
(AMNH unnumbered, SUI 12614, 12615, Univ. Texas unnumbered,
USNM 144423).
Illustrations. —One mature specimen of Mextcoceras guadalu-
pense guadalupense is figured —USNM 144423 (PI. 15, figs. 1-4).
Two immature individuals of Mexicoceras are illustrated: M. guada-
lupense guadalupense —USNM 144422 (PI. 15, fig. 7; Pl. 17, figs.
1,2), and M. guadalupense thompsoni—USNM 144419 (PI. 15,
fics. 5,6).
Genus WAAGENOCERAS Gemmellaro, 1887
Diagnosis. —Waagenoceras includes globose cyclolobids in
which initial subdivision of the primary lateral lobe produced three
lobes; the “mature” suture includes seven or eight external lateral
lobes.
Synonymy. — Wanneroceras Toumanskaya, 1937c, a nomen nu-
dwm based on W. gemmellarot Toumanskaya, 1937c, another nomen
nudum, is listed as a synonym of Waagenoceras by Bogoslovsky,
Librovitch, and Ruzhencev (1962) and of Timorites by Miller, Fur-
nish, and Schindewolf (1957).
Specific composition.— The type species of Waagenoceras is
W. mojsisovicsi Gemmellaro, 1887, by subsequent designation ( Die-
ner, 1921, p. 25). In addition to the type species the following taxa
70 BULLETIN 272
are referred to Waagenoceras: W. dieneri Bose, 1919, W. dienert
girtyi Miller and Furnish, 1940a, W. dieneri karpinskyi Miller, 1944,
W. lidacense Roever, 1940, W. nikitini Gemmellaro, 1888, and W.
stachei Gemmellaro, 1887. W. adrianense Gregorio 1930, W. minutum
Gregorio, 1930, and W. nikitini var, pergloboswm Gregorio, 1930,
may belong in Waagenoceras. W. coahwila Toumanskaya, 1937c,
is apparently a nomen nudum. W. gemmellaroi Haniel, 1915, was
retained in Waagenoceras by Ruzhencev (1955); other authors (for
example, Miller and Furnish, 1940a) have referred this species to
Timorites. W. richardson’ Plummer and Scott, 1937 (W. dieneri
richardsom of Miller and Furnish, 1940a) was recognized as being
transitional between Waagenoceras and Timorites by the latter two
authors; Glenister and Furnish (in Davis, Furnish, and Glenister,
1969) based a new genus, Newellites, on W. richardsont.
Immature shell. —The conch of Waagenoceras is globose (Gem-
mellaro, 1887, p. 11) to subspherical (Bogoslovsky, Librovitch, and
Ruzhencev, 1962, p. 396), is involute, and has a small and deep
umbilicus (Gemmellaro, 1887, p. 11).
Most individuals of Waagenoceras examined display two to
four constrictions in the outermost preserved whorl. The constric-
tions seem to be variably spaced; for example, in SUI 1784 the first
constriction is 346° from the protoconch, the second is 514° fur-
ther, and the third is 280° further.
The shell in a constriction is thicker than between constrictions,
but constrictions, as observed in median longitudinal sections of in-
dividuals of Waagenoceras, are variable. In some instances (for ex-
ample, the first constriction of SUI 1784) the shel] exterior is un-
affected, and the internal mold is concave outward at a constric-
tion; in other cases (such as the second and third constrictions of
SUI 1784) both the shell exterior and the internal mold are concave
outward at a constriction.
Most individuals of Waagenoceras examined by the author bear
transverse undulations in the surface of the internal mold. Although
many of these are due to plications in the shell, at least some are
the result of thickenings in the shell. Generally these undulations
are of low amplitude, but their amplitude and spacing are variable.
In SUI 1402 the outermost preserved whorl has a non-undulate sur-
face, but the next whorl within bears strong transverse plications.
PaLeozoic AMMonoIDs: Davis 7a
Well-preserved individuals of Waagenoceras bear transverse
growth lines. These lines are crowded at constrictions; in one un-
numbered individual from AMHN, for example, just adapical of a
constriction at a diameter of about 40 mm there are 34 growth
increments per millimeter of circumference, but about 1 mm after
the constriction there are 20/mm. As noted by Gemmellaro (1888,
p. 4) in some individuals clusters of growth lines occur in various
places not corresponding to constrictions.
Waagenoceras dieneri karpinskyi was defined on the basis of the
presence of reticulate ornament (in three individuals).
Gemmellaro (1887, p. 11) described the runzelschicht of
Waagenoceras as consisting of fine “dotting” connected to one an-
other by subtle undulating striations.
Mature shell. —In at least some individuals of Waagenoceras,
after formation of two-thirds of the ultimate whorl there occurred a
slight decrease in the radius of curvature of coiling followed by a
slightly greater increase, so that the venter appears slightly flattened
in longitudinal section in the final one-third volution; these changes
in coiling are likewise apparent in the course of the umbilical wall.
Gemmellaro (1887, p. 12, 13; 1888, p. 4) reported that the ven-
tral portion of at least the first two-thirds of the ultimate whorl of
Waagenoceras is narrow, whereas in the ultimate one-third whorl
the conch is wide and depressed.
In specimens SUI 32317 and 32318 the mature shell is genicu-
late and has a narrow, almost angular venter. Although sutural de-
tails are not preserved in either of the specimens at hand nor in
similar specimens from the same locality (Texas Tech unnumbered),
on the basis of associated fossils and sutural generalities, Furnish
(1967, personal communication) referred these specimens to W. rich-
ardsom. The conch form of these two individuals may be due, at
least in part, to postmortem deformation; the conch form certainly
is unlike that of any other known cyclolobid. The two examples and
others from the same locality may well not be congeneric with
Waagenoceras at all (indeed, Furnish and Glenister, in a footnote
in Davis, Furnish, and Glenister, 1969, erected the genus Newellites
for “Waagenoceras” richardsoni); on the other hand, SUI 32317
and 32318 may possibly represent the antidimorph of “ordinary”
Waagenoceras, or, perhaps, they are referable to Timorites. (A
72 BULLETIN 272
thorough study of the Cyclolobidae is necessary before this can be
resolved. )
No known individual of Waagenoceras bears any constrictions
in the wall of the ultimate body chamber except for that at the
adapertural end. Striations and undulations in the shell are present
in the living chamber, however.
The internal mold of most mature Waagenoceras living cham-
bers bears numerous, irregularly arranged and spaced, minute pits,
but these are absent in some individuals. In those individuals pos-
sessing these pits, they mark an area extending a variable distance
adapical from the ultimate constriction (1/4 whorl in SUI 32405;
between 1/3 and 1/2 whorl in SUI 32410; nearly one full whorl
in an unnumbered AMNH specimen).
As in many other taxa, the interseptal distance of Waagenoceras
decreases in the space occupied by the last several septa. Likewise,
the ultimate septum is thickened.
The ultimate peristome of Waagenoceras lacks either lappets
or a ventral prolongation.
The ultimate living chamber of Waagenoceras was reported by
Gemmellaro (1887, p. 11) to be one and one-half volutions long,
and that of W. dieneri was reported by Miller and Furnish (1940a,
p. 171) to be about one whorl long. In the four specimens avail-
able to the author in which both the apertural constriction and the
ultimate septum are visible, the living chamber is about one and
one-quarter whorls long.
The maximum diameter of mature individuals of Waagenoceras
ranges from about 90 mm (AMNH unnumbered) to 160 mm
(IGUP 32). The diameters of SUI 32317 and 32318 are 190 mm
and 170 mm respectively.
Material. — The preceding observations are based on study of:
Gemmellaro (1887, 1888), Mojsisovics (1888), White (1891), Smith
(1903), Bose (1919), Diener (1921, 1927), Berry (1928), Gregorio
(1930), Miller (1933, 1944, 1945a, 1945b, 1953), Greco (1935),
Toumanskaya (1937c), Plummer and Scott (1937), Marez Oyens
(1938), Roever (1940), Miller and Furnish (1940a, 1957a), Miller
and Unklesbay (1943), Hayasaka (1947, 1965), Ruzhencev (1955,
1956b), Miller, Furnish, and Schindewolf (1957), Bogoslovsky,
Librovitch, and Ruzhencev (1962), Furnish (1966), Davis, Furnish,
PaLEozoic AMMONOIDs: Davis 73
and Glenister (1969), Furnish and Glenister (1970), about 25 im-
mature undifferentiated Waagenoceras dienert (AMNH unnum-
bered, SUI 1401, 12492, 13601, 13602, 16690, 32402, 32406-32410),
about 10 mature undifferentiated W. dieneri (AMNH unnumbered,
SUI 32405, 32410, 32411), 1 immature W. dieneri dieneri (SUI
16690), about 20 immature W. dieneri girtyi (SUI 1366-1370, 1779,
1780, 1783, 1784, 12611, 13599, 13600), 1 mature W. dieneri girtyi
(SUI 12613), 1 immature W. lidacense (GIUA drawer E229 un-
catalogued), 1 immature and 2 mature specimens of W. richardsoni
(SUI 1402, 32317, 32318 respectively), and notes and photographs
taken by W. M. Furnish and Brian F. Glenister in the summer of
1965.
Illustrations. — Three mature specimens of Waagenoceras are
figured: W. mojstsovicst —IGUP 32 (PI. 16, fig. 1), IGUP 34 (PI.
16, figs. 2,3), and W. richardsont — SUI 32318 (PI. 16, figs. 6, 7).
One immature specimen of W. stachez is also illustrated — IGUP 37
GPT. 6, figs. 4;.5).
Genus CYCLOLOBUS Waagen, 1879
Diagnosis. — Cyclolobus includes those narrow, ellipsoidal to
lenticular cyclolobids in whose “mature” suture nine to fourteen
pairs of external lateral lobes are present. L! divides the first lateral
saddle medially, and U is relatively inconspicuous and is not sub-
divided.
Synonymy. — Bogoslovsky, Librovitch, and Ruzhencev (1962)
listed Godthaabites Frebold, 1932, and Procyclolobus Toumanskaya,
1939a, as synonyms of Cyclolobus. Nassichuk, Furnish, and Glen-
ster (1965) considered Cyclolobus (Cyclolobus) and Cyclolobus
(Krafftoceras) Diener, 1903, to be consubgeneric.
Specific composttion.— The type species of Cyclolobus is
Phylloceras oldhami Waagen, 1872, by original designation and by
monotypy; Furnish and Glenister (1970) considered C. insignis
Diener, 1903, to be a junior synonym. In addition, the following
taxa are referred to Cyclolobus: Godthaabites kullingi Frebold, 1932,
Arcestes (Cyclolobus Waag.) persulcatus Rothpletz, 1892, C. teich-
ertt Furnish and Glenister, 1970, and C. walkeri Diener, 1903. Fur-
nish and Glenister (1970) suppress all the following as junior syn-
onyms of C. walkeri: C. astrei Besairie, 1936, C. (Krafftoceras)
haydeni Diener, 1903, C. (Krafftoceras) kraffti Diener, 1903, and
74 BULLETIN 272
C. walkeri madagascariensis Besairie, 1936. The relationships of
Cyclolobus hoshanensis Sun, 1939, are unclear. Cyclolobus subcum-
minst Haniel, 1915, is a perrinitid.
Immature shell. — The conch of Cyclolobus is thinly ellipsoidal
to lenticular (Bogoslovsky, Librovitch, and Ruzhencev, 1962).
Most individuals of Cyclolobus formed constrictions at least
sometime in their ontogeny, but the spacing and, even, shape of
these structures seem to be quite variable. In C. walkeri (sensu Vail-
lant-Couturier-Treat, 1933, and Besairie, 1936) there are up to
four constrictions per whorl, generally restricted to diameters of less
than 25 mm, but constrictions may occur up to a diameter of 80
mm, albeit very slightly prominent; near the ultimate septum of
nearly all these specimens is a depression which Vaillant-Couturier-
Treat (1933, p. 25) thought could be considered a constriction. In
“C. krafftc” constrictions are restricted to the early whorls. In “C.
haydenw” there are four to five constrictions per whorl persisting
in the adult. The holotype of the type species bears six constric-
tions in a whorl of about 105 mm diameter. In C. walkert (sensu
Vaillant-Couturier-Treat, 1933) and C. cf, C. kullingi the constric-
tions in the earliest whorls are falciform, but in later whorls these
structures became biconvex. The constrictions of the holotype of the
type species are falciform at a diameter of about 105 mm.
At least some specimens of Cyclolobus have prominent trans-
verse plications in their early volutions; these fade out by a diameter
of 10 mm in C. oldhamt, by 30 mm in C. kullingi, and by 5 mm in
“C. krafftv’ according to Nassichuk, Furnish, and Glenister (1965).
In C. walkeri (sensu Vaillant-Couturier-Treat, 1933) barely percep-
tible ribs may be present even at maturity.
The transverse growth lines of specimens of Cyclolobus parallel
the constrictions, and at least in “C. insignis” they are bunched in
places (Diener, 1903, p. 165).
The suture of Cyclolobus is strongly arched at “maturity”.
Mature shell. — After about two-thirds of the ultimate whorl
of C'yclolobus was formed, there occurred a slight decrease in the
radius of curvature of coiling followed by a slightly greater in-
crease. The changes in coiling are much more prominently displayed
in the outline of the umbilicus than in the whorl periphery. In at
least C. walkert (sensu Vaillant-Couturier-Treat, 1933) these changes
PaLeozoic AMMOoNOIDs: Davis 75
in coiling were accompanied by an accentuation of the angularity
of the shoulder and by development of a dorsal bulge which covers
the preceding whorl and masks part of the umbilicus; in addition
the venter became keeled in the last three-fourths to two-thirds
of the ultimate volution (Vaillant-Couturier-Treat, 1933, p. 25).
Some individuals of Cyclolobus display constrictions in the ulti-
mate living chamber in addition to the terminal constriction. In
“C. insignis” these features extend across the venter (5/whorl);
in C. oldhami they are restricted to the umbilical region; constric-
tions are obsolete in C. walkeri (sensu Vaillant-Couturier-Treat,
1933). In “‘C. haydeni” there are four to five constrictions per whorl
persisting in the adult (Besairie 1936, p. 105).
The mature’ living chamber of Cyclolobus terminates in a bi-
convex constriction. The two convexities closer to the venter form
a pair of lappets. Within the constriction, just ventrad of each
lappet, is a deep pit, apparently affecting both the shell exterior
and the internal mold.
At least some Cyclolobus walkeri (sensu Vaillant-Couturier-
Treat, 1933) bear gentle biconvex ribs in the living chamber.
The length of the ultimate living chamber of Cyclolobus varies
from eight-ninths of a whorl to one full whorl,
The mature diameter of Cyclolobus is variable: “C. haydent”
— 57 mm (Diener, 1903, p. 168), “C. insignis” —66 mm (1bid., p.
165), C. walkeri (sensu Vaillant-Couturier-Treat, 1933) —90 mm
(UPSM #28) to nearly mature at 160 mm (MNHN Vaillant-Cou-
turier-Treat, 1933, pl. 2, fig. 1).
Material. — The preceding observations are based on study of:
Waagen (1872, 1879), Mojsisovics (1873, 1882), Rothpletz (1892),
Diener (1903, 1904, 1921, 1927); Haniel (1915), Grabaw (1924),
Vaillant-Couturier-Treat (1926, 1933), Smith (1927), Besairie
(1930, 1936), Frebold (1932), Wanner (1932), Miller (1933), Tou-
manskaya (1939a), Sun (1939), Miller and Furnish (1940b), Ruz-
hencev (1955), Jeannet (1959), Bogoslovsky, Librovitch, and Ruz-
hencey (1962), Nassichuk, Furnish, and Glenister (1965), Furnish
(1966), Davis, Furnish, and Glenister (1969), Furnish and Glen-
ister (1970), about three dozen immature examples of Cyclolobus
sp. (BMNH C34155, Collignon unnumbered, Copenhagen unnum-
bered, SUI 12313, 12343, 12344, 32397-32401, 32403, 32404), two
76 BULLETIN 272
mature Cyclolobus sp. (Collignon unnumbered, SUI 32396), about
ten immature specimens of C. oldhami (BMNH C10459, GPIT un-
numbered, Geological Survey of Pakistan unnumbered, USNM
unnumbered), three immature individuals of C. walkeri (BMNH
C36641-C36643 ), and photographs and notes taken by W. M. Fur-
nish and Brian F. Glenister in the summer of 1965.
Illustrations. —Two mature specimens of Cyclolobus walken
are figured: MNHN B7520 (PI. 17, fig. 7), and BMNH C36642 (PI.
17, fig. 8). Two immature examples of Cyclolobus are also illus-
trated: C. walkeri — MNHN B7517 (PI. 17, figs. 3-5), and “C.
astrev’? — UPSM unnumbered (PI. 17, fig. 6).
Family VIDRIOCERATIDAE Plummer and Scott, 1937
Diagnosis. — The Vidrioceratidae comprises cyclolobaceans
with “mature” sutures in which there are three to ten pairs of ex-
ternal lateral lobes which are denticulate only at the base.
Subfamily VIDRIOCERATINAE Plummer and Scott, 1937
Diagnosis. — Vidrioceratinae comprises ellipsoidal vidrioceratids.
Genus WAAGENINA Krotow, 1888
Diagnosis. — Waagenina comprises vidrioceratins in which four
or five lobes arose from each primary lateral lobe; of these four or
five lobes of the “mature” suture, the dorsal component is bifid.
Synonymy.— The following were considered synonyms of
Waagenina by Bogoslovsky, Librovitch, and Ruzhencev (1962):
Grioceras Toumanskaya, 1939a, Martoceras Toumanskaya, 1938,
Pamirites Toumanskaya, 1938, and Waagenia Krotow, 1885 (non
Kriechbaumer, 1874, fide Bogoslovsky, Librovitch, and Ruzhencev,
1962) (non Neumayr, fide Diener, 1897b).
Specific composition.— The type species of Waagenina is
Waagenma subinterrupta Krotow, 1885. (Popanoceras krasnopoly-
skyt Karpinsky, 1889, Marathonites krasnopolskyi var. asymmetrica
Ruzhencev, 1933, M. k. var. bidentata Ruzhencev, 1933, M. k. var.
quadridentata Ruzhencev, 1933, M. k. var. tridentata Ruzhencev,
1933, M. k. forms, a, B, y, and A of Voinova, 1934, and M. k. formae
a, b, c, d, e, and f of Maximova, 1935, are synonyms.) In addition
to the type species, the following are referred to Waagenina: Stacheo-
ceras benedictinum Gemmellaro, 1887, Pamirites clinei Toumans-
PaLeozoic AMMmonorpDs: Davis rad
kaya, 1938, Stacheoceras darae Gemmellaro, 1887, Marathomtes
dieneri Smith, 1927 (= Popanoceras timorense form 8 Haniel,
1915), Stacheoceras perspectivum Gemmellaro, 1887, and P. timo-
rense form y Haniel, 1915. P. tridens form 8 and P. t. form y Haniel,
1915, and Waagenia jazwae Krotow, 1885, may belong in Waagenima.,
Immature shell. —The conch of Waagenina is inflated, ellipsoi-
dal, involute, and has a narrow umbilicus (Ruzhencev, 1940d, p.
118).
In specimens of Waagenina perspectivum the constrictions are
narrow, affect both the shell exterior and the internal mold, and
extend neither across the venter nor to the umbilical shoulders.
In other species the constrictions are shallow and broad, extend
from shoulder to shoulder, but affect the shell exterior only slightly.
The occurrence and spacing of these constrictions are variable,
In some specimens of Waagenina the internal mold is marked
by gentle transverse undulations which correspond in position to
the growth lines. In some individuals (for example, SUI 32597,
32598) there are stronger undulations in the internal mold; they
are restricted to the flanks of the conch and correspond in spacing
to the interseptal intervals, Growth lines are more closely spaced
at constrictions than between.
In some individuals of Waagenina (such as, SUI 10588, 32595)
there is a definite shell thickening in the umbilicus. This thickening
is bordered by a distinct edge which approximately corresponds
to the umbilical shoulder.
In a number of examples of Waagenina (for example, SUI
10588, 32595) there are numerous minute holes on the flanks of
the phragmocone steinkern. Other individuals (such as, SUI 32596)
lack these features. In size, shape, and arrangement, these small
pits are similar to those near the ultimate constriction of many
adrianitids. The pits do not correspond in form to the runzelschicht
observed in some specimens (for example, GPIT PV24409).
Mature shell. — The only mature representatives of Waagenina
known to the author were studied by Gemmellaro and referred by
him to Stacheoceras darae. In these specimens there was little, if any,
change in coiling in the ultimate whorl. However, the venter of the
ultimate portion of the conch appears more broadly rounded than
in inner whorls.
78 BULLETIN 272
The mature living chamber of Waagenina darae terminates in
a constriction. There appear to be two distinct trends of the ultimate
constrictions in this genus: in the specimen of Gemmellaro’s 1887
plate 4, figures 2 and 3 (IGUP 68) there is a distinct ventral] re-
éntrant, whereas in that of his 1887 plate 4, figure 11 (IGUP 41)
the reéntrant is slight. The ultimate peristome is marked by a ven-
trolateral lappet on each side, flanking a ventral sinus.
The ultimate living chambers of Waagenina perspectivum and
W. darae were stated by Gemmellaro (1887, pp. 36, 39) to be longer
than the ultimate whorl and only as long as the terminal volution,
respectively.
Maximum diameters of only two individuals (of Waagenina
darae) are known to the author: 54% mm (IGUP 64) and 78 mm
(IGUP 68).
Material. — The preceding observations are based on study of:
Krotow (1885), Gemmellaro (1887), Karpinsky (1889), Diener
(1897b), Smith (1927), Schindewolf (1931), Ruzhencev (1933,
1938, 1940d, 1951, 1956a, 1962), Voinova (1934), Greco (1935),
Maximova (1935), Toumanskaya (1938, 1939a, 1949), Bogoslov-
skaya (1962), Bogoslovsky, Librovitch, and Ruzhencev (1962),
about 20 immature specimens of Waagenina (subinterrupta — SUI
10588, 32595-32598, darae — GPIT PV24397, PV24409, perspec-
tivwm —GPIT PV24409), and photographs and notes taken by
W. M. Furnish and Brian F. Glenister in the summer of 1965.
Illustrations. —Three mature specimens of Waagenina darae
are figured:,IGUP 64 (Pl;18) figs. 5,6), (PINS iiss 95910) and
IGUP 68 (Pl. 18, figs. 11-13). In addition, three immature in-
dividuals of Waagenina are illustrated: W. subinterrupta — SUI
32597 (PIN lS htigss 3.4), SUL 32597 (CPI2 19; fiets9)eandeWaene=
Spectiuum — IGUP 66 (PI. 18, figs. 1, 2).
Genus STACHEOCERAS Gemmellaro, 1887
Diagnosis. — Stacheoceras comprises vidrioceratins with “ma-
ture” sutures in which there are six to twelve subdivisions of the
primary lateral lobe on each side.
Synonymy. — Neostacheoceras Schindewolf, 1931, is a synonym
of Stacheoceras.
Specific composition. — The type species of Stacheoceras is S.
PaLreozoic AMMONOIDS: Davis 79
mediterraneum Gemmellaro, 1887, by subsequent designation
(Diener, 1921, p. 22), but Smith (1927) and Schindewolf (1931)
considered Arcestes antiquus Waagen, 1879, to be the type species
of Stacheoceras by original designation. In addition, the following
species are referred to the genus: Stacheoceras andrussow: Touman-
skaya, 1931, S. arthaberi Smith, 1927 (= Popanoceras tridens form
8 and « of Haniel, 1915), S. boeset Toumanskaya, 1931, S. borisstaki
Toumanskaya, 1931, Popanoceras collignoni Besairie, 1936, S. medt-
terraneum Gemmellaro var, crimensis Toumanskaya, 1931, S. diblasu
Gemmellaro, 1887, S. gaudryi Gemmellaro, 1888, S. gemmellarot
Miller, 1944, S. gordoni Miller, Furnish, and Clark, 1957, S. gruene-
waldti Gemmellaro, 1887 (= S. gruenwaldti of Hayasaka, 1965),
Neostacheoceras hanieli Schindewolf, 1931 (= Popanoceras timo-
rense form 9 of Haniel, 1915), Stacheoceras iwaizakiense Mabuti,
1935, S. karpinskyi Gemmellaro, 1887, S. rothi Miller and Furnish,
1940a, S. sexlobatum Miller and Cline, 1934, Popanoceras timorense
form 8, £, and » of Haniel, 1915, Stacheoceras towmanskyae Miller
and Furnish, 1940a, Arcestes tridens Rothpletz, 1892, Popanoceras
tridens form ¢ Haniel, 1915, Stacheoceras trimurti Diener, 1897a,
Popanoceras tschernyschewi Stoyanow, 1910, and Stacheoceras vogtt
Toumanskaya, 1931. The following may also belong in Stacheoceras:
S. discoidale Toumanskaya, 1935, S. globosum Gemmellaro, 1887,
S. globulinwm Gregorio, 1930, S. pelagicum Gemmellaro, 1887, S.
quadridens Hayasaka, 1947, S. sosiense Gregorio, 1930, S. tepense
Toumanskaya, 1931, and S. tietzei Gemmellaro, 1887. Stacheoceros
[sic] marcoui Gemmellaro, 1887, and Stacheoceras trimarti Diener,
1899, of Furnish (1966) are both apparently nomina nuda.
Immature shell. — The immature conch of Stacheoceras is sub-
globular to subdiscoidal, with a small umbilicus (Miller and Furnish,
1940; p.. 129).
Constrictions in the shell of Stacheoceras are predominantly in-
ternal thickenings in the shell and, therefore, affect the shell ex-
terior slightly, if at all. The number of constrictions per whorl varies
from zero to four, with no apparent orderly relation between
diameter and the number of constrictions per volution.
Growth lines in Stacheoceras parallel the constrictions.
In some individuals there is a shell thickening in the umbilicus.
This thickening is bordered by a distinct edge.
80 BuLLeTIN 272
Mature shell.—In at least some specimens of Stacheoceras,
after about three-quarters of the ultimate whorl was formed, there
was a decrease in the radius of curvature of coiling followed by a
slightly greater increase. In SUI 32610A an earlier change in
coiling occurred after slightly less than two-thirds of the ultimate
volution had been produced. There is a terminal dorsal flaring of the
shell into the umbilicus displayed by SUI 1364W and 32608D.
The ultimate living chamber of Stacheoceras terminates in a
constriction. In addition to this terminal constriction, the body
chambers of IGUP 71 and SUI 16652 each bear a constriction
about one-half whorl adapical of the peristome and another about
three-quarters adapical of the ultimate constriction.
There are gentle transverse undulations in the steinkern of the
final living chamber of Stacheoceras.
There is one short, broad, ventrolateral lappet on each side in
the ultimate peristome of Stacheoceras. Each lappet bears a branch
of the terminal constriction.
The length of the ultimate living chamber in Stacheoceras is
variable. Gemmellaro (1887, p. 29 et seq.) reported the body cham-
ber lengths in S. mediterranewm and S. diblasii as one and one-half
and one and one-third whorls, respectively. Wanner (1932, p. 272)
noted that the body chamber of S. tridens is one and two-fifths
volutions long. One specimen of S. gemmellaroi (SUI 32612) has an
ultimate living chamber of just less than one whorl in length. Two
specimens of S. towmanskyae (SUI 32607E, 16652F) have body
chambers of one and one-eighth and seven-eighths whorls, re-
spectively.
Wanner (1932, p. 272) reported the mature diameter of one
specimen of Stacheoceras tridens as 125% mm. One S. tietzet (IGUP
71) is mature at a diameter of 47% mm, One S. gemmellaroi (SUI
32612) is mature at 65 mm. One S. towmanskyae has a mature
diameter of about 85 mm. Three specimens of S. sp. (SUI 32610D,
32608D, 32610A) have maximum diameters of about 90 mm, about
90 mm, and 107 mn, respectively.
Material. — The preceding observations are based on study of:
Waagen (1879), Krotow (1885), Gemmellaro (1887, 1888), Roth-
pletz (1892), Haug (1894, 1898), Diener (1897a, 1921), Stoyanow
(1910) Haniel (1915), Smith (1927), Gregorio (1930), Schinde-
wolf (1931), Toumanskaya (1931, 1935), Wanner (1932), Miller
PaLeozoic AMMONOIDs: DAvis 81
and Cline (1934), Mabuti (1935), Greco (1935), Besairie (1936),
Plummer and Scott (1937), Hayasaka (1940, 1947, 1965), Miller
and Furnish (1940a, 1957a, 1957b), Miller and Unklesbay (1943),
Miller (1944, 19452), Miller, Furnish, and Clark (1957), Triimpy
(1960), Gordon and Merriam (1961), Bogoslovsky, Librovitch,
and Ruzhencev (1962), Shevyrev (1965), Furnish (1966), Davis,
Furnish, and Glenister (1969), about 130 immature specimens of
Stacheoceras (S. sp. — Copenhagen unnumbered, SUI 12314, 13597,
32608-32611, 32613, antiquum — GPIT unnumbered, gemmellaroi
— SUI 1365, 13610, 13656, sexlobatum — SUI 706, tridens — SUI
32615, towmanskyae — SUI 1364, 16652, 32607), about ten mature
individuals (S. sp. — SUI 32608, 32610, gemmellaroi — SUI 32612,
toumanskyae — SUI 1364, 16652, 32607), and photographs and notes
taken by W. M. Furnish and Brian F. Glenister in the summer of
1965.
Illustrations. — Four mature pugcueene of Stacheoceras are
figured: two S. sp. — SUI 32608D (PI. 19, fig. 5), SUI 32610A (PI.
19, figs. 6-8), one S. tvetzei — IGUP 71 (PI. 4, figs. 4, 5), and one
S. towmanskyae — SUI 13640 (PI. 19, figs. 3, 4). In addition, three
immature individuals of Stacheoceras are illustrated: S. gaudryi —
IGUP 59 (PI. 4, fig. 6), S. mediterranewum — IGUP 62 (PI. 19, figs.
1,2), and S. pelagicum — MGPU unnumbered (PI. 18, figs. 7, 8).
Superfamily MARATHONITACEAE Ruzhencevy, 1938
Family MARATHONITIDAE Ruzhencev, 1938
Diagnosis. — The Marathonitidae comprises ammonoids whose
suture underwent goniatitic development, following which the pri-
mary lateral, umbilical, and internal lateral lobes each split into
three, producing the basic formula:
ee ee Te Usich Dep.
Dig lee e ee eee ee Oink LS
Subsequent complication arose through denticulation of the lobes
with a bifurcation of the dorsad Ls and the ventrad /» or a trifurca-
tion of D, or with all three.
Subfamily MARATHONITINAE Ruzhencev, 1938
Diagnosis. — The Marathonitinae comprises marathonitids with
sutures in which the dorsal lobe is trifid and the three (or four)
82 BuLLeETIN 272
subdivisions of the initial lateral lobe are primarily trifid; during
phylogeny this latter trifidity developed into complex subdivision,
particularly in the ventrad two or three subdivisions of the primary
lateral lobe.
Genus HYATTOCERAS Gemmellaro, 1887
Diagnosis. — Hyattoceras comprises marathonitins with “ma-
ture” sutures in which each branch of the ventral lobe has denticles
on its dorsad side; there are four subdivisions of the primary lateral
lobe of which the ventrad three are strongly subdivided.
Synonymy. — Abichia Gemmellaro, 1887, (not Abichia as used
by Andrussow, 1907) is considered to be congeneric with Hyatto-
ceras.
Familial assignment. — Furnish and Glenister (in Davis, Fur-
nish, and Glenister, 1969) considered Hyattoceras to belong in a
family Hyattoceratidae.
Specific composition. — The type species of Hyattoceras is H.
geinitzi Gemmellaro, 1887, by subsequent designation (Diener,
1921, p. 25). In addition to the type species, the following are re-
ferred to Hyattoceras: H. (Abichta) abichi Gemmellaro, 1887, 7.
guembeli Gemmellaro, 1888, H. subgeimitzi Haniel, 1915, and H.
turgidum Gemmellaro, 1887.
Immature shell.— The immature conch of Hyattoceras is el-
lipsoidal to discoidal, involute, and has a narrow umbilicus (Bogo-
slovsky, Librovitch, and Ruzhencev, 1962).
The only ornament characteristic of immature Hyattoceras
is the transverse striae.
Mature shell. — As the middle one-third of the ultimate volu-
tion of Hyattoceras geinitzi was formed there was a progressive nar-
rowing of the venter. The narrowest place is at the end of this
middle one-third and corresponds to two changes in coiling, first a
slight decrease in the radius of curvature of coiling, followed im-
mediately by a slightly greater increase. These changes in coiling
caused the venter of the ultimate one-third whorl to appear flat-
tened in lateral view. The venter of this ultimate one-third whorl
also became progressively more flattened in transverse section, for,
after the marked narrowing of the venter in the middle one-third
of the ultimate whorl, the venter became progressively broader. Also
in the last one-third of the terminal volution there developed a
dorsad flaring of the peristome, completely closing the umbilicus.
PaLeozorc AMMONOIDs: DAvIs 83
In Hyattoceras guembeli and H. turgidwm the changes in
coiling are less striking, and the narrowing of the venter followed
by a flattening appears to be less pronounced or even absent.
The mature conch of Hyattoceras abicht is markedly geniculate,
with a narrowing of the venter followed by a broadening similar
to that of Hyattoceras gemmitzt. Haug (1897) suggested that Abichia
is the antidimorph of Hyattoceras, the former being male and the
latter, female.
The ultimate living chamber of Hyattoceras terminates in a
pronounced constriction. In the ventral region of H. geinitz this
constriction is immediately adoral of a tranverse ridge, and the
ventral part of the constriction is depressed, producing a slight mid-
ventral salient in the peristome.
The mature living chamber of Hyattoceras geinitzt was reported
by Gemmellaro (1887, p. 16) to be one and one-third whorls long,
of H. abichi to be one and one-third volutions long (1887, p. 18),
and of H. guembeli to be almost one and one-half whorls long (1888,
D7).
The maximum diameter of three mature Hyattoceras geinitzi
is about 40 mm, of one mature H. guembeli is 67 mm, of one mature
H, turgidum is 23% mm, and of five mature H. abichi is between
18 mm and 23 mm.
Material. — The preceding observations are based on study of:
Gemmellaro (1887, 1888), Haug (1897), Haniel (1915), Smith
(1927), Plummer and Scott (1937), Miller and Furnish (1940a),
Toumanskaya (1949), Bogoslovsky, Librovitch, and Ruzhencev
(1962), Davis, Furnish, and Glenister (1969), two immature speci-
mens of Hyattoceras sp. (GIUA Dr. 511 no. T581, SUI 32581),
three immature (BMNH (33681, C73335, C73336) and three
mature (GPIT unnumbered, USNM unnumbered) examples of
Hi. geinitzt, three mature individuals of H. abichi (USNM unnum-
bered), about ten immature specimens of H. subgeinitzi (GIUA
Dr. 551 no, T582, PIUB unnumbered, SUI 12322, 32463) and photo-
graphs and notes taken by W. M. Furnish and Brian F. Glenister
in the summer of 1965.
Illustrations. — Six mature specimens of Hyattoceras are fig-
ured: two H. abichi—IGUP 55 (PI. 21, figs. 1-3), USNM unnum-
bered (PI. 21, figs. 4, 5), two H. geinitzi —IGUP 52 (PI. 20, figs.
5-8), GPIT unnumbered (PI. 21, figs. 10), one H. guembeli — IGUP
84 BuLLETIN 272
unnumbered (PI. 20, figs. 3, 4), and one H. turgidum — IGUP 51
GEIN20; figs: 15 2):
Genus PSEUDOVIDRIOCERAS Ruzhencev, 1936
Diagnosis. — Pseudovidrioceras comprises those marathonitins
with “mature” sutures in which the prongs of the ventral lobe
are undenticulate, in which each of the subdivisions of the primary
lateral lobe terminates in three denticles, and in which there are
three complete lobes plus part of a fourth on each side between the
bifid ventral lobe and the umbilical shoulder. (Furnish and Glenister
[in Davis, Furnish, and Glenister, 1969] considered Pseudovidrio-
ceras to be a subgenus of Marathonites.)
Specific composition. — The type species of Pseudovidrioceras
is Vidrioceras girtyi Miller and Cline, 1934, by original designation.
The only other species which has been referred to Pseudovidrioceras
is Stacheoceras pygmeum Gemmellaro, 1887, Peritrochia novomexi-
cana Miller and Parizek, 1948, and P. dunbari Miller and Furnish,
1940a, may belong in Pseudovidrioceras.
Immature shell.—The conch of Pseudovidrioceras is sub-
discoidal (Miller and Cline, 1934, p. 290) to discoidal (Gemmellaro,
1887, p. 41). The venter is rounded, and the conch is compressed
laterally. The umbilical shoulders are rounded, and the umbilicus
is closed. In the one known specimen of P. girtyi W is greater than
H in all whorls save the last; in P. pygmeum the height is greater
than the width.
In Pseudovidrioceras pygmeum the only ornament on the im-
mature shell consists of the fine, radial striae which are sinuous
in the ventral region.
Mature shell.— There were apparently no changes in coiling
in the growth of the mature living chambers of Pseudovidrioceras
before the formation of the apertural constriction. In P. girtyi, how-
ever, there was a change of conch proportions such that H exceeds
W in the ultimate whorl. In P. pygmeum the dorsal part of the
ultimate one-third volution extends over the umbilicus, completely
covering it.
The living chamber of the type species of Pseudovidrioceras
lacks constrictions except for that at the aperture. On the other
hand, in P. pygmeum (according to Gemmellaro, 1887) the internal
mold of the ultimate whorl bears a few broad and superficial
PaLreozoic AMMONOIDs: Davis 85
“strangolamenti” (apparently minor constrictions) which extend
from the umbilicus straight across the venter without sinuations.
A transverse bulge is present in at least Pseudovidrioceras pyg-
meum just orad of the ultimate constriction. Miller and Cline (1934,
p. 290) concluded that the aperture of P. girtyi is large and flaring;
the aperture of P. pygmeum is neither.
The interseptal spacing of the last several septa of Pseudo-
vidrioceras girtyi decreases progressively.
The length of the mature living chamber of Pseudovidrioceras
is one whorl.
The maximum diameter of many specimens of P. pygmeum
doesn’t exceed 9 mm (Gemmellaro 1887, p. 41), but one specimen
is at least 11, and another, 12 mm in diameter (ibid.). The soli-
tary known specimen of P. girtyi has a mature diameter of 15% mm.
Material. — The preceding observations are based on study of:
Gemmellaro (1887), Toumanskaya (1931), Miller and Cline (1934),
Ruzhencev (1936, 1950), Miller and Furnish (1940a), Miller and
Parizek (1948), Bogoslovsky, Librovitch, and Ruzhencev (1962),
Davis, Furnish, and Glenister (1969), and photographs and notes
taken by W. M. Furnish and Brian F. Glenister in the summer of
1965.
Illustrations. — One mature specimen of Pseudovidrioceras pyg-
meum is figured — IGUP 65 (PI. 21, figs. 11-13).
Genus MARATHONITES Bose, 1919
Diagnosis. — Marathonites comprises marathonitins with “ma-
ture” sutures in which each branch of the ventral lobe is bifid and
each of the three subdivisions of the primary lateral lobe is trifid.
Familial assignment. — Furnish and Glenister (in Davis,
Furnish, and Glenister, 1969) considered Marathonites to belong
in the family Vidrioceratidae.
Taxonomic composition.—The type species of Marathonites was
Stacheoceras (Marathonites) J. P. Smithi Bése, 1919, by original
designation, Plummer and Scott (1937) and Miller and Furnish
(1940a) considered S. (M.) J. P. Smithi Bose, 1919, to be a junior
synonym of Popanoceras ganti Smith, 1903; if the two names denote
the same taxon, then the type species of the genus should be called
Marathonites ganti (Smith, 1903). In addition to the type species,
86 BULLETIN 272
the following taxa are referred to Marathonites: M. (Almites)
busterensis Toumanskaya, 1949, M. electraensis Plummer and
Scott, 1937, M. gracilis Smith, 1927, M. arathomites invariabilis
Ruzhencev, 1933, Stacheoceras kussicum Toumanskaya, 1931, S. k.
var. djidairensis Toumanskaya, 1931, M. (Almites) pamiricus
Toumanskaya, 1949, M. (A.) pressulus Ruzhencev, 1956a, Popano-
ceras romanowskyt Karpinsky, 1889, M. sellardsi’ Plummer and
Scott, 1937, Stacheoceras swevt Toumanskaya, 1931, M. sulcatus
Bose, 1919, M. (M.) wuralensis Ruzhencev, 1940d, Stacheoceras
uralicum Fredericks, 1921, and M. vidriensis Bose, 1919.
Two subgenera of Marathomites have been recognized: Mara-
thonites (Almites) Toumanskaya, 1941, and Marathonites (Mara-
thomtes) Bose, 1919. In addition, Furnish and Glenister (in Davis,
Furnish, and Glenister, 1969) considered the previously discussed
Pseudovidrioceras to be a subgenus of Marathonites.
Immature shells. — The conch of Marathonites is subglobose,
strongly involute, and narrowly umbilicate (Ruzhencev, 1940d, p.
125). The ratio W/D decreases during ontogeny.
Most immature specimens of Marathonites exhibit at least one
constriction in the outermost preserved whorl. The distance be-
tween constrictions varies (for example, in SUI 1065, a specimen of
M. ganti, there are three constrictions in one whorl, but two of the
interconstrictional distances are one-quarter whorl). Nonetheless
there seems to be a general increase in the number of constrictions
per whorl with ontogeny up to a maximum of six. The constrictions
affect the internal mold only.
Commonly individuals of Marathonites bear transverse undula-
tions in the surface of the internal mold, These undulations ap-
parently correspond in position and trend to the growth lines on
the shell exterior. These shell plications are much more prominent
immediately adapical of a constriction than adapertural of it; this
latter position corresponds to a crowding of growth lines. In the
region adapical of a constriction the plications are not absolutely
parallel to that constriction, but each successive plication in an
adapertural direction has a trend which is more closely similar to
that of the constriction.
In at least some individuals of Marathonites the shell exhibits
longitudinal lirae. These structures are present only on the flanks
PaLEozoic AMMONOIDs: Davis 87
in specimens of M. ganti and are less prominent than the growth
lines. In M. sellardsi there is one striking rib on each shoulder and
one single, less prominent lira just dorsal and another just ventral
of each main rib. Development of this rib is even more pronounced
in M. uralensis in which there are two very prominent ribs, one on
each shoulder.
Mature shell. —There are two distinct mature conch morpho-
logies displayed by representatives of Marathomites.
In at least M. ganti, M. invariabilis, and M. pressulus the
mature living chamber is distinguished only by a prominent aper-
tural constriction.
In four specimens of Marathonites sp. from Timor (SUI 12301,
12304-12306) the mature living chamber also has an apertural con-
striction, but the conch is geniculate with the venter broad and
rounded except at the more adapertural of the two changes in
coiling. Here the venter is narrow. (Furnish and Glenister [im Davis,
Furnish, and Glenister, 1969] referred these four specimens to
Pseudovidrioceras, which they considered a subgenus of MMara-
thonites.)
In both groups of mature forms the length of the living chamber
is slightly less than one whorl to approximately one whorl. A group
of three mature specimens of M. ganti varies from 28 to 33 mm in
diameter. One individual of M. invariabilis (Ruzhencev, 1956a, pl.
37, fig. 3) has a mature diameter of 50 mm; three others (SUI
10594B, 32616B, 10594A) have diameters of 39 mm, 49 mm, and
51 mm, respectively. One mature example of M. pressulus (ibid., pl.
38, fig. 2) is 34 mm in diameter. Three of the four specimens of the
other mature form of Marathonites are 28 or 29 mm in diameter.
The significance of these two mature forms of Marathonites is
unclear in the light of the material available, However, if the dif-
ferences are of subgeneric taxonomic value, they are at variance with
the subgenera of Marathonites as currently defined [viz. M. (M.)
gantt, M. (Almites) tnvariabilis and M. (A.) pressulus all have the
same type of modification]. Perhaps the two forms are antidimorphs.
Material. — The preceding observations are based on study of:
Karpinsky (1889), Smith (1903, 1927), Bose (1919), Fredericks
(1921), Schindewolf (1931), Toumanskaya (1931, 1939b, 1941,
1949), Miller and Cline (1934), Plummer and Scott (1937),
Ruzhencev (1938, 1940c, 1940d, 1950, 1956a, 1962), Miller and
88 BULLETIN 272
Furnish (1940a), Miller and Youngquist (1947), Miller and Downs
(1950), Bogoslovsky, Librovitch, and Ruzhencev (1962), Davis,
Furnish, and Glenister (1969), about 60 immature specimens of
Marathonites (M. sp. — SUI 32589, ganti— SUI 1061-1067, 1898,
10525, 11046, 13817-13819, 17033, 17034, 32588, 32590, 32594,
invariabilis — SUI 10594, 32616, 32617, J. P. Smithi — SUI
13821, sellardsi — SUI 1396, 1406, 32591, John Britts Owen Collec-
tion 695), and about ten mature examples of Marathonites (M. sp.
— SUI 12301, 12304-12306, gantz — SUI 1063, 13817, invartabilis
— SUI 10594, 32616).
Illustrations. — Four mature specimens of Marathomtes are
figured: two M. sp. — SUI 12301 (PI. 14, fig. 7, Pl. 21, figs. 7-9),
SUI 12305 (PI. 21, fig. 6), one M. ganti — SUI 1063 (PI. 22, figs.
5-7), and one M. invariabilis — SUI 10594 A (PI. 22, figs, 8-10).
In addition, one immature specimen of M. gantz is illustrated — SUI
17033 (PI. 22, figs. 1, 2).
Subfamily KARGALITINAE Ruzhencev, 1960
Diagnosis. — The Kargalitinae comprises marathonitids with
sutures in which the dorsal lobe is either bifid or trifid and the ven-
trad lobe of the three (or four) subdivisions of the primary lateral
lobe is primarily bifid.
Genus PERITROCHIA Girty, 1908
Diagnosis. — Peritrochia includes kargalitins with “mature”
sutures in which the branches of the ventral lobe are not subdivided;
the dorsad of the three subdivisions of the primary lateral lobe and
the ventrad of the three subdivisions of the primary internal lateral
lobe each bifurcated during ontogeny; the dorsal lobe is entire.
Familial assignment. — Furnish and Glenister (in Davis,
Furnish, and Glenister, 1969) considered Peritrochia to belong in
the family Vidrioceratidae.
Synonymy. — Paratrochia Diener (1921, p. 10) is apparently a
typographic error,
Specific composition.— The type species and only named
species referred to Perttrochia is P. erebus Girty, 1908, by monotypy.
Immature shell. — The conch of Peritrochia is parabolic (Bogo-
slovsky, Librovitch, and Ruzhencev, 1962, p. 392), subdiscoidal to
subglobular with a small umbilicus.
PaLeozoic AMMONOIDs: DAvis 89
There are up to five constrictions per whorl in Perttrochia.
These features affect only the internal mold. In addition to constric-
tions, most individuals display gentle transverse plications as well
as growth lines. From the Bone Spring Limestone in Texas one
specimen of Peritrochia (SUI 32584) was recovered which appar-
ently is a new species; this individual bears transverse plications
which are relatively conspicuous. The trends of the growth lines,
plications, and contrictions are parallel; early in ontogeny all three
are essentially straight, becoming more sinuous with ontogenetic
development.
In addition to the transverse ornament, well-preserved shells
and internal molds of Peritrochia exhibit faint longitudinal undula-
tions.
In the immediate vicinity of the umbilicus is an area in which
the shell appears thicker than ventrad of this area. This thickened
shell forms a distinct edge on the umbilical shoulder.
Mature shell.— There is little if any change in coiling, in
conch proportions, or in both in the ultimate living chamber of Peri-
trochta.
Constrictions are lacking on the ultimate whorl save for the
apertural modification. Transverse plications, striations, and longi-
tudinal ornament are present in the ultimate living chamber, as is
the spire of thickened shell at the umbilicus.
The ultimate body chamber of Peritrochia appears to be just
less than one whorl long.
The maximum diameter of Peritrochia varies from 18 to 24 mm.
Material. — The preceding observations are based on study of:
Gemmellaro (1887), Girty (1908), Toumanskaya (1931), Greco
(1935), Plummer and Scott (1937), Miller and Furnish (1940a,
1957b), Ruzhencev (1940c, 1940d), Mullerried, Miller, and Furnish
(1941), Miller (1945a), Miller and Youngquist (1947), Miller and
Parizek (1948), Miller and Downs (1950), Unklesbay (1962), Bogo-
slovsky, Librovitch, and Ruzhencev (1962), Davis, Furnish, and
Glenister (1969), about 20 immature (SUI 32583) and eight ma-
ture (SUI 32583, 32585, 32586) representatives of Peritrochia erebus,
and one immature specimen of a new species of Peritrochia (fide
W. M. Furnish, 1967, personal communication).
90 BULLETIN 272
Illustrations. —Two mature specimens of Peritrochia erebus
are figured: SUI 32585 (PI. 14, figs. 4-6) and SUI 32586 (PI. 14,
figs. 1, 2). In addition, two immature specimens of Peritrochia are
illustrated: P. n. sp. — SUI 32584 (PI. 14, fig. 3) and P. erebus —
SUMG2583t hr ( PlO22 ehiesh Sy 4):
REFERENCES
Andrussow, N.
1907. About the genus Arcicardium Fischer. Annual of Geology &
Mineralogy of Russia, vol. 9, pp. 105-114, pls. 3, 4 [in Russian
and German].
Armstrong, J. D., Dear, J. F., and Runnegar, B.
1967. Permian ammonoids from eastern Australia. Geol. Soc. Australia,
Jour., vol. 14, pt. 1, pp. 87-97, pls. 5, 6.
Arthaber, G.
1911. Die Trias von Albanien. Beitr. zur Palaont. u. Geol. Osterr.-Un-
garns, Bd. 24, pp. 169-277, pls 17-24.
Berry, Edward W.
1928. Cephalopod adaption—the record and its interpretation. Quar. Rev.
Biology, vol. 3, No. 1, pp. 92-108.
Besairie, Henri
1930. Recherches géologiques & Madagascar. Soc. Hist. Nat. Toulouse,
Bull., vol. 60, 2d fascicule, pp. 1-272, pls. 1-27.
1936. Recherches géologiques &@ Madagascar, Ire Suite—La géologie du
Nord-Ouest, Chapitre 3—Les fossiles. Acad. Malgache Mém., Fasc.
21, pp. 105-207, pls. 4-24.
Blainville, M. H. D. de
1840. Prodrome d’une monographie des ammonites, pp. 1-31 of Supple-
ment Du Dictionaire Des Sciences Naturelles, Bertrand, Paris.
Boehm, G.
1907. Geologische Mitteilungen aus dem Indo-Australischen Archipel.
VIb Jiingeres Paldozoicum von Timor. N. Jb. Min., Geol., Palaont.,
Beilage-Band 25, pp. 303-323, pls. 10, 11.
Bohmers, J. C. A.
1936. Bau und Struktur von Schale und Sipho bei permischen Am-
monoidea. Univ. Amsterdam, pp. 1-125, pls. 1, 2.
Bose, Emil.
1919. The Permo-Carboniferous ammonoids of the Glass Mountains,
West Texas, and their stratigraphical significance. Univ. Texas
Bull., No. 1762 (1917), pp. 1-241, pls. 1-11.
Bogoslovskaya, M. F.
1962. Artinskian ammonoids from the Central Urals. Acad. Sci. USSR,
Works of Paleont. Institute [Trudy], vol. 87, pp. 1-117, pls. 1-11
[in Russian].
Bogoslovsky, V. I., Librovitch, L. S., and Ruzhencev, V. E.
1962. Pp. 334-409 of Molluscs-Cephalopods. I. vol. (V. E. Ruzhencey,
ed.) of Principles of Paleontology [Osnovy] (Yu. A. Orlov, overall
editor), Publishing House, Acad. Sci. USSR, Moscow [in Russian].
Callomon, J. H.
1963. Sexual dimorphism in Jurassic ammonites. Leicester Lit. & Phil.
Soc., Trans., vol. 57, pp. 21-56, pl. 1.
Chao, King-Koo
1965. The Permian ammonoid-bearing formations of South China.
Scientia Sinica, vol. 14, No. 12, pp. 1813-1825, pls. 1, 2.
PaLeozoic AMMONOIDs: DAvIs 91
Chernov, A. A.
1939. Order Ammonoidea. The Ammonoids. Atlas of the principle forms
of fossils of the fauna of the USSR, vol. 6. Permian System, pp.
160-190 [in Russian].
Coémme, S.
1917. Note critique sur le genre Cadmoceras. Soc. Géol. France, Bull.,
ser. 4, vol. 17, pp. 44-54, pl. 6.
Cowen, Richard, Gertman, Richard, and Wiggett, Gail
1971. Color patterns in Nautilus, and their implications for cephalopod
paleobiology [Abstract]. Geol. Soc. America, Abstracts with pro-
grams, vol. 3, No. 2, pp. 104-105.
Davis, Richard Arnold, and Campbell, Carlyle B.
1968. An orthocerid cephalopod from the Cherokee Group of Marion
County, Iowa. Iowa Acad. Sci., Proc., vol. 73, pp. 279-284.
Davis, Richard Arnold, Furnish, W. M., and Glenister, Brian F.
1969. Mature modification and dimorphism in Late Paleozoic am-
monoids. pp. 101-110, pls. 2-5 Sexual Dimorphism in Fossil Metazoa
and Taxonomic Implications (G. E. G. Westermann, editor), In-
ternational Union of Geological Sciences, ser. A, No. 1.
Davitashvili, L. Sh., and Khimshiashvili, N. G
1954. On the question of the biological significance of the apertural
formation of ammonites. Works [Trudy] of the Paleobiology Sec-
tion of the Academy of Science of the Georgian Soviet Socialist
Republic, vol. 2, pp. 44-76 [in Russian].
Demanet, F.
1943. Les horizons marins du Westphalien de la Belgique et leurs faunes.
Mus. Roy. Hist. Nat. Belgique, Mém., No. 101, pp. 1-166, pls. 1-9.
Diener, Carl
1897a. The Permocarboniferous fauna of Chitichun, No. I, Geol. Surv.
India, Mem., Palaeontologia Indica, ser. 15, Himdlayan Fossils,
vol. 1, No. 3, pp. 1-105, pls. 1-13.
1897b. The cephalopoda of the Lower Trias. Geol. Surv. India, Mem.,
Palaeontologia Indica, ser. 15, Himdlayan Fossils, vol. 2, No. 1,
pp. 1-181, pls. 1-23.
1903. Permian fossils from the central Himalayas. Geol. Surv. India,
Mem., Palaeontologia Indica, ser. 15, Himdlayan Fossils, vol. 1
pt. 5, pp. 1-204, pls. 1-10.
1904. Note on Cyclolobus Haydeni, Diener. Geol. Sur. India, Records,
vol. 31, pt. 2, pp. 56-58.
1921. Ammonoidea permiana. pt. 14 (pp. 1-36) of Fossilium Catalogus I:
Animalia (C. Diener, ed.), W. Junk, Berlin.
1927. Leitfossilien des marinen Perm. 5th Leiferung (pp. 1-84, pls. 1-14)
Leitfossilien (Georg Giirich, ed.), Gebriider Borntraeger, Ber-
in.
Dixon, Joe Scott
1960. A statistical study of seven species of the Pennsylvanian-Permian
goniatite, Agathiceras. M. S. Thesis, University Iowa, pp. 1-58.
Dunbar, Carl Owen
1924. Phases of cephalopod adaption. Pp. 187-223 in Organic Adaption
to Environment (M. R. Thorpe, ed.), Yale Univ. Press, New
Haven, Conn.
Emeliancev, T.
1929. Artinsk cephalopods from the Polyudovsk Ridge district in the
Urals. Geol. Comm. (Leningrad). Transactions [Izvestiia], vol.
48, No. 8, pp. 145 (1229) - 152 (1236), pl. 51 [in Russian].
’
9? BuLieTIN 272
Foord, A. H., and Crick, G. C.
1897. Catalogue of the fossil Cephalopoda in the British Museum, Part
3. Bactritidae and Ammonoidea. pp. i-xxxviii + 1-303, London.
Frebold, Hans
1932. Marines Unterperm in Ostgrénland und die Frage der Grenzzie-
hung zwischen dem pelagischen Oberkarbon und Unterperm. Med-
delelser om Grgnland, vol. 84, No. 4, pp. 1-33, pl. 1.
Frech, Fritz
1915. Loses und geschlossenes Gehduse der tetrabranchiaten Cephalo-
poden. Centralblatt Min., Geol., Palaont., Jahrg. 1915, pp. 593-606.
Fredericks, George
1921. On some ammonoids of the Upper Carboniferous of the Ural
Mountains. Soc. Paleont. Russia, Ann., vol. 3, pp. 73-94, pl. 1
[in Russian].
Fretier, V., and Graham, A.
1966. Reproduction. Chapter 4 (pp. 127-164) of vol. 1, Physiology of
Mollusca (Karl M. Wilbur and C. M. Yonge, ed.), Academic
Press, New York.
Furnish, W. M.
1966. Ammonoids of the Upper Permian Cyclolobus-zone. N. Jb. Geol.
Palaont., Abh., vol. 125, pp. 265-296, pls. 23-26.
Furnish, W. M., and Glenister, Brian F.
1970. Permian ammonoid Cyclolobus from the Salt Range, West Pakistan.
pp. 153-175. Stratigraphic Boundary Problems: Permian and Trias-
sic of West Pakistan (Bernhard Kummel and Curt Teichert, ed.),
University Press of Kansas, Lawrence, Kansas.
Furnish, W. M., and Knapp, W. D.
1966. Lower Pennsylvanian fauna from eastern Kentucky; Part 1, Am-
monoids. Jour. Paleont., vol. 40, No. 2, pp. 296-308, pl. 35.
Gemmellaro, Gaetano Georgio
1887. La fauna dei calcari con Fusulina della valle del Fiume Sosio
nella provincia di Palermo, Fascio I, Cephalopoda, Ammonoidea.
Gior. Sci. Nat. Econ., vol. 19, pp. 1-106, pls. 1-10.
1888. La fauna dei calcari con Fusulina della valle del Fiume Sosio
nella provincia di Palermo, Appendice Fascio I, Cephalopoda,
Ammonoidea. Gior. Sci. Nat. Econ., vol. 20, pp. 9-36, pls. A-D.
Gerth, H.
1950. Die Ammonoideen des Perms von Timor und ihre Bedeutung fiir
die stratigraphische Gliederung der Permformation. N. Jb. Min.,
Geol., Palaont., Bd. 91, Abt. B, pp. 233-320.
Gillet, S.
1937. Les Ammonites du Bajocien d’Alsace et de Lorraine. Serv. Carte
Géol. Alsace et Lorraine, Mém., vol. 5, pp. 1-130.
Girty, G. H.
1908. The Guadalupian fauna. U. S. Geol. Sur., Prof. Paper 58, pp. 1-
651, pls. 1-31.
Glenister, B. F., and Furnish, W. M.
1961. The Permian ammonoids of Australia. Jour. Paleont., vol. 35, No.
4, pp. 673-736, pls. 78-86.
Gordon, Mackenzie, Jr., and Merriam, C. W.
1961. Late Permian ammonoids in the Inyo Range, California, and their
significance. U. S. Geol. Sur., Prof. Paper 424-D, pp. D-238-D-239.
Grabau, A. W.
1923-1924. Stratigraphy of China, Part 1, Paleozoic and Older. China
Geol. Sur., Peking, pp. 1-528, pls. 1-6.
PaLeozoic AMMONOIDs: DAvis 93
Greco, Benedetto '
1935. La Fauna Permiana del Sosio Conservata nei Musei di Pisa, di
Firenze e di Padova, Palaeontographia Italica, vol. 35, pp. 101-190,
pls. I-IV (12-15).
Gregorio, M. A. De
1930. Sul Permiano di Sicilia. Annales de géologie et de paléontologie,
No. 52, pp. 1-70, pls. 1-21.
Guex, Jean
1968. Note Préliminaire sur le dimorphisme sexuel des Hildocerataceae
du Toarcien moyen et supérieur de Aveyron (France). Bull. Soc.
Vaudoise de Sciences Naturelles, Lausanne, vol. 70, No. 327, pp.
57-84, pls. 1-3.
Haniel, C. A.
1915. Die Cephalopoden der Dyas von Timor. Abh. 6 (pp. 1-153, pls. 46-
56) of Lieferung 3 of Paldontologie von Timor (J. Wanner, ed.),
Stuttgart.
Haug, Emile
1894. Les Ammonites du Permien et du Trias. Remarques sur leur classi-
fication. Soc. Géol. France, Bull., ser. 3, vol. 22, pp. 385-412.
1897. [Observations on Glangeaud, 1897]. Soc. Géol. France, Bull.,
Ser 3.2 V0len25.9 py 107.
1898. Etudes sur les Goniatites. Soc. Géol. France, Mém., Paléont., Mém.
18, pp. 1-112, pl. I.
Hayasaka, Ichiro
1940. On two Permian ammonoids from the Kitakami Mountains, north
Japan. Geol. Soc. Japan, Jour., vol. 47, No. 565, pp. 422-427.
1947. A Permian cephalopod faunule from Chéchiang Prov., China.
Acta Geol. Taiwanica, vol. 1, No. 1, pp. 13-38, pls. 1, 2.
1963. Some Permian fossils from southern Kitakami, I1I—Ammonoidea.
Japan Acad., Proc., vol. 39, No. 8, pp. 594-599.
1965. Some cephalopods in the Permian faunule of Takokura-Yama,
Fukushima Prefecture, Japan. Palaeont. Soc. Japan, Trans. Proc.,
N.S., No. 57, pp. 8-27, pls. 2, 3.
Hyatt, Alpheus
1874. Abstract of a memoir on the “Biological relations of the Jurassic
ammonites”. Boston Soc. Nat. Hist., Proc., vol. 17, pp. 235-241.
Jeannet, Alphonse
1948. Sur la forme de l’ouverture chez certains Nautiles mésozoiques.
Schweiz. Miner. & Petrogr. Mitteil., vol. 28, fase. 1 (Festschrift
P. Niggli), pp. 178-187, figs. 1-5.
1959. Ammonites Permiennes et Faunes Triasiques de Himalaya Cen-
tral. Geol. Surv. India, Mem., Palaeontologia Indica, new ser.,
vol. 34, mem. 1, pp. 1-189, pls. 1-21.
Karpinsky, A. P.
1874. Geological investigation in the Orenburgian region. Records Min.
Soc., ser. 2, No. 9, pp. 212-310, pl. 12 [in Russian].
1889. Uber die Ammoneen der Artinsk-Stufe und einige mit denselben
verwandte carbonische Formen. Imp. Acad. Sci. St. Petersburg,
Mem., ser. 7, vol. 37, No. 2, pp. 1-104, pls. 1-5.
Kerr, J. G.
1931. Notes upon the Dana specimens of Spirula and upon certain prob-
lems of cephalopod morphology. Oceanographical Reports edited by
the “Dana” Committee, No. 8, pp. 1-34, pls. 1-20.
Krotow, P.
1885. The Artinskian Stage. A geological-paleontological monograph of
the sandstone of Artinsk. Works of the Society of Naturalists
94 BULLETIN 272
attached to the Imperial Kazan University, vol. 13, No. 5, pp.
1-312, pls. 1-4 [in Russian].
1888. Geological investigations on the western slope of the Solikamsk
and Cherdin Regions of the Urals. Works of the Geol. Comm. of
St. Petersburg, vol. 6, No. 1, pp. 1-563 [in Russian].
Kummel, Bernhard, and Lloyd, R. M.
1955. Experiments on relative streamlining of coiled cephalopod shells.
Jour. Paleont., vol. 29, No. 1, pp. 159-170.
Lane, F. W.
1960. Kingdom of the octopus. Sheridan House, New York, pp. 1-300.
Lehmann, Ulrich
1966. Dimorphismus bei Ammoniten der Ahrensburger Lias-Geschiebe.
Palaont. Zeitschr., vol. 40, No. 1/2, pp. 26-55, pis. 3, 4.
Mabuti, S.
1935. On the occurrence of Stacheoceras in the Kitakami mountainland,
etc. Saito Ho-on Kai Mus. Bull., No. 6, pp. 143-149, pl. 1.
Makowski, Henryk
1962. Problem of sexual dimorphism in ammonites. Paleont. Polonica,
12-1962, pp. 1-92, pls. 1-20.
Mangold-Wirz, Katharina
1963. Biologie des Céphalopodes benthiques et nectoniques de la mer
Catalane. Vie et Milieu, suppl. No. 13, pp. 1-285. pls. 1-4.
Mangold-Wirz, Katharina, Lu, C. C., and Aldrich, Frederick A.
1969. A reconsideration of forms of squid of the genus Illex (Illicinae,
Ommastrephidae). II. Sexual dimorphism. Canadian Jour. Zoology,
vol. 47, No. 6, pp. 1153-1156, pls. 1-4.
Marez Oyens, F. A. H. W. De
1938. Preliminary note on the occurrence of a new ammonoid fauna of
Permian age on the island of Timor. Koninklijke Nederlandsche
Akademie van Wetenschappen, Proc., vol. 41, No. 10, pp. 1122-1126.
Maximova, S. V.
1935. On some new species of artinsk ammonoids. Moscow Soc. Natural-
ists, Bull., Sec. Geology, vol. 13, No. 2, pp. 273-288, pl. 1 [in
Russian |.
McCaleb, James A.
1968. Lower Pennsylvanian ammonoids from the Bloyd Formation of
Arkansas and Oklahoma. Geol. Soc. Amer., Sp. Paper 96, pp. 1-
123eeplssul=12e
McCaleb, James A., and Furnish, W. M.
1964. The Lower Pennsylvanian ammonoid genus Axinolobus in the
southern midcontinent. Jour. Paleont., vol. 38, No. 2, pp. 249-255,
pl. 40.
Mikesh, D. L., and Glenister, B. F.
1968. Solenochilus springeri (White & St. John, 1868) from the Pennsyl-
vanian of southern Iowa. Iowa Acad. Sci., Proc., vol. 73, pp. 269-
278.
Miller, A. K.
1933. Age of the Permian limestones of Sicily. Amer. Jour. Sci., ser. 5,
vol. 26, No. 154, pp. 409-427.
1944. Permian cephalopods. pt. IV (pp. 71-127, pls. 20-45) of King, et al.
Geology and paleontology of the Permian area northwest of Las
Delicias, southwestern Coahuila, Mexico. Geol. Soc. Amer., Sp.
Paper 52.
1945a. Some exceptional Permian ammonoids from West Texas. Jour.
Paleont., vol. 19, No. 1, pp. 14-21, pls. 6-8.
PaLEozoic AMMONOIDs: DAvVIs 95
1945b. - eeniae ammonoid from Sonora. Jour. Paleont., vol. 19, No. 1,
1953. Pephalnsolte Pp. 81-82, pl. 24D, figs. 21-23 of Cooper, et al.
Permian fauna at El Antimonio, W estern Sonora, Mexico. Smith-
sonian Misc. Coll., vol. 119, No. 2.
Miller, A. K., and Cline, L. M.
1934. The cephalopods of the Phosphoria Formation of ge
United States. Jour. Paleont., vol. 8, No. 3, pp. 281-302, pl.
Miller, A. K., and Crockford, M. B.
1936. Permian cephalopods from British Columbia. RH Soc. Canada,
Trans., 3d ser., sec. IV, vol. 30, pp. 23-28, pl.
Miller, A. K., and Downs, H. R.
1950. Ammonoids of the Pennsylvanian Finis Shale of Texas. Jour.
Paleont., vol. 24, No. 2, pp. 185-218, pls. 31-35.
Miller, A. K., and Furnish, 'W. M.
1939. The late Paleozoic ammonoid families Adrianitidae and Aga-
thiceratidae. Palaont. Zeitschr., vol. 21, No. 4, pp. 297-303.
1940a. Permian ammonoids of the Guadalupe Mountain Region and ad-
jacent areas. Geol. Soc. Amer., Sp. Paper 26, pp. 1-242, pls. 1-44.
1940b. Cyclolobus from the Permian of eastern Greenland. Meddelelser
om Grénland, vol. 112, No. 5, pp. 1-10, pl. 1
1957a. Ammonoids of the basal Word Formation, Glass Mountains, West
Texas. Jour. Paleont., vol. 31, No. 6, pp. 1052-1056.
1957b. Permian ammonoids from Tunisia. Jour. Paleont., vol. 31, No. 4,
pp. 705-712, pl. 83.
Miller, A. K., Furnish, W. M., and Clark, D. L.
1957. Permian ammonoids from western United States. Jour. Paleont.,
vol. 31, No. 6, pp. 1057-1068, pls. 133, 134.
Miller, A. K., Furnish, W. M., and Schindewolf, O. H.
1957. Paleozoic ammonoids. Pp. L11-L79 of Treatise on Invertebrate
Paleontology (R. C. Moore, ed.), Geol. Soc. Amer. and Univ.
Kansas Press, Lawrence, Kansas.
Miller, A. K., and Parizek, E. J.
1948. A lower Permian ammonoid fauna from New Mevxico. Jour.
Paleont., vol. 22, No. 3, pp. 350-358, pls. 56-58.
Miller, A. K., and Unklesbay, A. G.
1943. ive siphuncle of late pal oem ammonoids. Jour. Paleont., vol.
17, No.9], pp: 1-25, spls.1-
Miller, A. K., and Youngauist, Welter,
1947. emer Permian cephalopods from the Texas Colorado River
Brey: Univ. Kansas, Paleont. Contrib., No. 2, art. 1, pp. 1-15,
pls. 1-3.
Miller, S. A., Tand Gurley, W. F. E.
1896. New species of Paleozoic invertebrates from Illinois and other
states. Ill]. State Museum, Bull. No. 11, pp. 1-50, pls. 1-5.
Mojsisovics von Mojsvar, E.
1873,1875. Die Mollusken-Faunen der Zlambach-und Hallstdatter-Schich-
ten. K.-K. Geol. Reichsanstalt, Abhand., vol. 6, pp. 1-174, pls. 1-70.
1882. Die Cephalopoden der mediterranen Triasprovinz. K.-K. Geol.
Reichsanstalt, Abhand., vol. 10, pp. 1-322, pls. 1-94.
1888. Uber einige arktische Trias- Ammoniten des Lala ag Siberien
Imp. Acad. Sci. St. Petersbourg, Mem., ser. 7, vol. 36, No. 5, pp.
1-21, pls. 1-3.
Morton, S. G.
1836. Appendix. Being a notice and description of the organic remains
96 BuLLETIN 272
embraced in the preceding paper. Amer. Jour. Sci., vol. 29, pp.
149-154, pls. 1-36.
Mullerried, F. K. G., Miller, A. K., and Furnish, W. M. i
1941. The middle Permian of Chiapas, southernmost Mexico, and its
fauna, Amer. Jour. Sci., vol. 239, No. 6, pp. 397-406, pl. 1.
Nassichuk, W. W., Furnish, W. M., and Glenister, B. F.
1965. The Permian ammonoids of Arctic Canada. Geol. Sury. Canada,
Bull. 131, pp. 1-56, pls. 1-5.
Nikitin, S.
1884. General geological map of Russia, Sheet 56. Works [Trudy]
Geol. Comm., vol. 1, No. 2, pp. 1-153, pl. 1-3 [in Russian, Ger-
man summary ].
Pelseneer, P.
1926. La proportion relative des sexes chez les animaux et particuliére-
ment chez les Mollusques. Acad. Roy. Belgique, cl. Sci., Mém.,
2deser., vol.) 8) tascy Ihe ppyal=258:
Perna, A.
1915. On life habits of the goniatites. Geological Proceedings, vol. 1, No.
1, pp. 6-14 [in Russian].
Perna, E. Ya.
1914. The ammonoids of the upper Neodevonian of the east slope of the
southern Urals. Works of the Geol. Comm. of St. Petersburg,
n.s., vol. 99, pp. 1-114, pls. 1-4 [in Russian, German summary ].
Plummer, F. B., and Scott, Gale
1937. Upper Paleozoic ammonites in Texas. The geology of Texas, vol.
III, pt. 1; Texas Univ. Bull. 3701, pp. 1-516, pls. 1-41.
Pruvot-Fol, A.
1937. Remarques sur le Nautile. Comptes Rendus du XII& Congrés
International de Zoologie — Lisbonne (1935), vol. 3, pp. 1652-1663.
Roever, W. P. De
1940. Description of some Permian ammonoids from F. Koekatoe, Lidak.
pp. 206-210, pl. V, fig. 2, of vol. I, Geological Expedition to the
Lesser Sunda Islands (H. A. Brouwer, ed.), Amsterdam.
Rothpletz, A.
1892. Die Perm-, Trias- und Jura-Formation auf Timor und Rotti im
indischen Archipel. Palaeontographica, vol. 39, pp. 57-106, pls. 9-14.
Ruzhencev, V. E.
1933. Concerning some Lower Permian ammonoids of the Aktyubinsk
Region. Moscow Soc. Naturalists, Bull., new ser. vol. 41, Sec.
Geology, vol. 11(2), pp. 164-180, pls. 4, 5 [in Russian, French
summary].
1936. Paleontological notes about Carboniferous and Permian ammonites.
Problems Soviet Geology, No. 12, pp. 1072-1088 [in Russian, English
summary ].
1937. The Carboniferous and Permian problem. Moscow Soc. Natural-
ists, Bull., new ser. vol. 45, Sec. Geology, vol. 15(5), pp. 401-411 [in
Russian, English title].
1938. Ammonoids of the Sakmarian Stage and their stratigraphic sig-
nificance. Problems Paleontology, vol. 4, pp. 187-285, pls. 1-7 [in
Russian, English introductory material].
1939. The importance of ontogeny for the natural classification of am-
monites. Acad. Sci. USSR, Transactions [Izvestiia], Ser. Biol.,
No. 1, pp. 13-32 [in Russian, English summary].
1940a. On the family Adrianitidae Schindewolf. Acad. Sci. USSR, Re-
ports [Doklady], vol. 26, No. 8, pp. 837-840.
1940b. Ad new genus Aristoceras from the Upper Carboniferous of the
Urals. Acad. Sci. USSR, Reports [Doklady], vol. 27, No. 5, pp.
524-528.
PaLteozoic AMMONOIDs: Davis 97
1940c. On the question of the taxonomic position of some of the Upper
Paleozoic ammonites. Acad. Sci. USSR, Reports [Doklady], vol.
28, No. 3, pp. 285-288.
1940d. An essay on the natural classification of some Late Paleozoic am-
monites. Acad. Sci. USSR, Works Paleont. Inst. [Trudy], vol.
11, No. 3, pp. 1-134, pls. 1-6 [in Russian, English summary].
1941. The first representative of the genus Emilites Ruzhencev in the
Upper Carboniferous of the Urals. Acad. Sci. USSR, Reports
[Doklady], vol. 30, No. 9, pp. 884-886.
1950. Upper Carboniferous ammonites of the Urals. Acad. Sci. USSR,
Works Paleont. Inst. [Trudy], vol. 29, pp. 1-223, pls. 1-15 [in
Russian ].
1951. Lower Permian ammonites of the southern Urals—I. Ammonites of
the Sakmarian Stage. Acad. Sci. USSR, Works Paleont. Inst.
[Trudy], vol. 33, pp. 1-188, pls. 1-15 [in Russian].
1952. Biostratigraphy of the Sakmarian Stage in the Aktyubinsk region
of Kazakhsh SSR, Acad. Sci. USSR, Works Paleont. Inst. [Trudy],
vol. 42, pp. 1-87, pls. 1-6 [in Russian].
1955. About the family Cyclolobidae Zittel. Acad. Sci. USSR, Reports
[Doklady], vol. 103, No. 4, pp. 701-703 [in Russian].
1956a. Lower Permian ammonites of the southern Urals—II. Ammonites
of the Artinskian Stage. Acad. Sci. USSR, Works Paleont. Inst.
[Trudy], vol. 60, pp. 1-275, pls. 1-39 [in Russian].
1956b. About some new genera of ammonoids. Acad. Sci. USSR, Reports
[Doklady], vol. 107, No. 1, pp. 158-161 [in Russian].
1960. Principles of systematics, system and phylogeny of Paleozoic am-
monoids. Acad. Sci. USSR, Works Paleont. Inst. [Trudy], vol.
83, pp. 1-331 [in Russian].
1962. Superorder Ammonoidea. The ammonoids—general part. Pp. 243-
334 of Molluscs-Cephalopods. I. Vol. (V. E. Ruzhencev, ed.)
Principles of Paleontology [Osnovy] (Yu. A. Orlov, overall editor),
Publishing House, Acad. Sci. USSR, Moscow [in Russian].
Sazonov, N. T.
1957. Jurassic deposits of the Central Region of the Russian Platform.
Leningrad, pp. 1-155 [in Russian].
Schindewolf, O. H.
1931. Uber den Ammoniten-Sipho. Preuss. Geol. Landesanst. Sitzungsber.,
Heft 6, pp. 197-209, pl. 5.
Shevyrev, A. A.
1965. Superorder Ammonoidea. Pp. 166-182, pl. 17-24 of Evolution and
change of marine organisms at the Paleozoic-Mesozoic Boundary
(V. E. Ruzhencev and T. G. Sarycheva, ed.), Acad. Sci. USSR,
Works Paleont. Inst. [Trudy], vol. 108 [in Russian].
Siemiradzki, J.
1925. Podrecznik paleontologit. C:1. Paleozoologia, Warsaw, pp. 1-389.
Smith, J. P.
1903. The Carboniferous ammonoids of America. U.S. Geol. Surv., Mon-
ogr. 42, pp. 1-211, pls. 1-29.
1927. Permian ammonoids of Timor. Jaarboek van het Mijnwezen in
Nederlandsch-Indié, Jaargang 55 (1926), Verhandelingen 1, pp.
1-90, pls. 1-16.
Stenzel, H. B.
1964. Living Nautilus. Pp. K59-K93, Treatise on Invertebrate Paleon-
tology (R. C. Moore, ed.), Geol. Soc. Amer. and Univ. Kansas
Press, Lawrence, Kansas.
Stoyanow, A. A.
1910. On the character of the boundary of Palaeozoic and Mesozoic
near Djulfa. Imp. Russ. Mineralogical Soc., Mem., vol. 47, No.
1 pps 6-139.
98 BULLETIN 272
Sun, Y. C.
1939 (1947). The uppermost Permian ammonoids from Kwangsi and
their stratigraphical significance. The 40th Anniv. Papers Nat.
Univ. Peking, pp. 35-49, pls. 1, 2, (Contr. from Geol. Inst., Nat.
Univ. Peking, No. 28, pp. 35-49, pls. 1, 2).
Teichert, Curt
1944. Two new ammonoids from the Permian of Western Australia.
Jour. Paleont., vol. 18, No. 1, pp. 83-89, pl. 17.
Teichert, Curt, and Fletcher, H. O.
1943. A Permian ammonoid from New South Wales and the correla-
tion of the Upper Marine Series. Australian Mus., Rec., vol. 21,
pp. 156-163, pl. 11.
Toumanskaya, O. G.
1931. Permo-carboniferous beds of the Crimea. Part I. Cephalopoda,
Ammonoidea. Supreme Council of National Political Economy of
the USSR, Principal Geological Prospecting Bureau, Moscow, pp.
1-117, pls. 1-8 [in Russian, English summary ].
1935. Permian ammonoids of the River Kubergandy and their strati-
graphic significance. Acad. Sci. USSR, Tajik-Pamir Expedition
1933, Works of the Expedition, Pub. 31, pp. 43-130, pls. 1-5 [in
Russian, English summary ].
1937a. About representatives of the family Adrianitidae Schindewolf.
Moscow Soc. Naturalists, Bull., Sec. Geology, vol. 15, No. 4, pp.
375-378 [in Russian].
1937b. On representatives of the new genus Crimites in Permian deposits.
Yearbook of All-Russian Paleontology. General, vol. 11, pp. 146-147
[in Russian, English abstract].
1937c. La stratigraphie du systeme pérmien d’aprés les ammonées. Inter-
national Geological Congress, 17th Session, Abstracts of papers,
pp. 93-94.
1938. On certain new genera of the family Popanoceratidae Hyatt. Sov.
Geol., 1938, No. 12, pp. 106-108 [in Russian].
1939a. On the correlation of the Permian deposits of the USSR with the
deposits of other countries. Moscow Soc. Naturalists, Bull., Sec.
Geology, vol. 17, Nos. 2, 3, pp. 17-26 [in Russian].
1939b. Permian ammonoids of the Transalai Range and their significance
for the stratigraphy of the Pamirs. Problems Paleontology, vol. 5,
pp. 93-137, pls. 1-8 [in Russian, English summary ].
1939c. Permian ammonoids of the Range of Peter I. Problems Paleon-
tology, vol. 5, pp. 139-145, pl. 1 [in Russian, English summary].
1941. On the stratigraphy of the Permian succession of the Crimea. Acad.
Sci. USSR, Reports [Doklady], vol. 32, No. 4, pp. 261-264.
1949. On the Permian ammonoids of Central Asia. Moscow Soc. Natural-
ists, Bull., Sec. Geology, vol. 24, No. 6, pp. 49-84, pls. 1-6 [in
Russian].
Trueman, A. E.
1941. The ammonite body-chamber, with special reference to the
buoyancy and mode of life of the living ammonite. Geol. Soc.
London, Quar. Jour., vol. 96, pt. 4, pp. 339-378, (379-383: dis-
cussion).
Trimpy, Rudolf
1960. Uber die Perm-Trias Grenze in Ostgrinland und iiber die Prob-
lematik stratigraphischer Grenzen. Geol. Rdsch., vol. 49, No. 1,
pp. 97-103.
PaLEeozoic AMMOoNoOIDs: DAvis 99
Unklesbay, A. G.
1962. Pennsylvanian cephalopods of Oklahoma. Oklahoma Geol. Surv.,
Bull. 96, pp. 1-150, pls. 1-19.
Vaillant-Couturier-Treat, Ida
1926. Note sur le Permienne marin de Madagascar. Comptes Rendus,
Acad. Sci. Paris, vol. 182, pp. 1092-1094.
1933. Paléontologie de Madagascar, XIX—Le Permo-Trias Marin. An-
nales de Paleontologie, vol. 22, fasc. 2, pp. 37-96, pls. 5-10.
Voinova, E.
1934. Some ammonites from the Baigendja Horizon of the Artinskian
of the Southern Urals. The United Geological and Prospecting
Service of USSR, Trans., fasc. 352, 55 numbered pages, pls. 1-5
[in Russian].
Waagen, William
1872. On the occurrence of Ammonites, associated with Ceratites, and
Gontatites in the Carboniferous deposits of the Salt Range. Geol.
Sur. India, Mem., Palaeontologia Indica, vol. 9, pp. 351-358, pl. 1.
1879. Productus Limestone fossils, 1. Pisces-Cephalopoda. Geol. Sur.
India, Mem., Palaeontologia Indica, ser. 13—Salt Range Fossils,
vol. 1, pp. 1-72, pls. 1-6.
Wanner, J.
1932. Zur Kenntnis der permischen Ammonoideen-fauna von Timor.
N. Jb. Min., Geol., Palaont., Beil.-Bd. 67, Abt. B, pp. 257-286, pls.
9-11.
Wells, M. J.
1962. Brain and behavior in cephalopods. Stanford Univ. Press, Stan-
ford, California, pp. 1-171.
1966. The brain and behavior of cephalopods. Chapter 15 (pp. 547-590),
vol. 2 of Physiology of Mollusca (Karl M. Wilbur and C. M
Yonge, ed.), Academic Press, New York.
Westermann, G. E. G.
1964. Sexual-Dimorphismus bei Ammonoideen und seine Bedeutung fiir
die Taxonomie der Otoitidae. Palaeontographica, Bd. 124, Abt. A,
Lief. 1-3, pp. 33-73, pls. 6-9.
1969. Supplement: Sexual Dimorphism, migration, and segregation in
living cephalopods. Pp. 18-20 in Sexual dimorphism in fossil
Metazoa and taxonomic implications (G. E. G. Westermann, ed.),
Int. Union Geol. Sci., ser. A, No. 1.
White, C. A.
1891. The Texas Permian and its Mesozoic type of fossils. U. S. Geol.
Sur., Bull. 77, pp. 1-51, pls. 1-4.
Willey, Arthur
1902. Contribution to the natural history of the pearly nautilus. Zoological
results based on material from New Britain, New Guinea, Loyalty
Islands, and elsewhere, collected during the years 1895, 1896, and
1897. Pt. 6, pp. 691-830, pls. 75-86, Cambridge University Press.
Zeiss, Arnold
1969. Dimorphismus bei Ammoniten des Unter-Tithon. Pp. 155-162 of
Sexual dimorphism in fossil Metazoa and taxonomic implications
(G. E. G. Westermann, ed.), Int. Union Geol. Sci., ser. A, No. 1.
Zittel, Karl Alfred Von
1895. Grundziige der Palaeontologie. Pp. 386-435. R. Oldenbourg,
Minich.
1903. Grundziige der Palaeontologic (2d edition). Abt. 1, pp. 398-468.
R. Oldenbourg, Miinich.
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PLATES
The cost of reproduction of the plates has been met by a grant from the
Graduate Division of the University of Cincinnati.
Enlargements stated to nearest 0.1.
102 BULLETIN 272
EXPLANATION OF PLATE 3
Figure Page
1-7, ‘Adrianites'Gemmellaro, 1887. 2..005.sc¢....0..6ch2 200.0004 43
1-3. A. elegans, syntype (IGUP 85, Gemmellaro, 1887, pl. 6,
figs. 14, 15), Sosio Limestone, Passo di Burgio, Pro-
vince of Palermo, Sicily, maximum diameter—20%
mm, X2.0; the specimen is mature; 5-7. A. elegans,
syntype (IGUP 85, Gemmellaro, 1887, pl. 7, fig. 23),
Sosio Limestone, Passo di Burgio, Province of Pal-
ermo, Sicily, maximum diameter—21 mm, X 2.0; the
specimen is mature; 4. A. isomorphus Gemmellaro,
(SUI 32442), Sosio Limestone, near Palazzo Adriano,
Sicily, maximum width—17 mm, x 2.0; the ultimate
peristome of the dorsal shell is shown.
3:9. (Neocrimites Ruzhencev; 1940a° 1 2...00... 3... ..2.. 6503-00: ee 51
8,9. N. cancellatus (Haniel) (fide Furnish and Glenister in
Davis, Furnish, and Glenister, 1969), hypotype (GIUA
Drawer 55, No. 1328, zbid., pl. 3, figs. 4, 5), Maoen
Mollo, Timor, xX 2.0; the specimen is mature.
PLATE 3
‘BULL. AMER. PALEONT., VOL. 62
PLATE 4
BULL. AMER. PALEONT., VOL. 62
Figure
PaLeozoic AMMoNoIDs: Davis
EXPLANATION OF PLATE 4
fs G.o Adrianites GemmellarOslS87. 1s. fcccecccso8%..sobteonwdtete,.s.008hede-
1-3. A. isomorphus, syntype (IGUP 86, Gemmellaro, 1888,
pl. B, figs. 5,6), Sosio Limestone, Rupe di San Bene-
detto, Province of Palermo, Sicily, maximum diameter
—25 mm, X 2.0; the specimen is mature. (Designated
lectotype by Furnish and Glenister in the explanation
of plate 3 of Davis, Furnish, and Glenister, 1969).
7,8. A. isomorphus (USNM Unnumbered specimen a),
Sosio Limestone, Rocca di Salomone, Province of
Palermo, Sicily, & 2.5; figure 7 shows a thickening
of the dorsal shell at a constriction, and figure 8
shows the ultimate peristome of the dorsal shell. 9.
A. isomorphus (USNM unnumbered specimen b),
Sosio Limestone, Rupe di San Cologero, Province of
Palermo, Sicily, x 3.7; the figure shows the pits in
the internal mold at the ultimate constriction and the
spherical bodies in the shell which correspond to these
pits.
4-6. Stacheoceras Gemmellaro, 1887 22.00...
4,5. S. tietzei, syntype (IGUP 71, Gemmellaro, 1887, pl. 5,
figs. 1-3), Sosio Limestone, Passo di Burgio, Province
of Palermo, Sicily, maximum diameter—47%4 mm, xX
1.0; the specimen is mature. 6. S. gaudryi, syntype
(IGUP 59, Gemmellaro, 1888, pl. D, fig. 1), Sosio
Limestone, Pietra di Salamone, Province of Palermo,
Sicily, maximum diameter—34 mm, X 1.2; the speci-
men is immature.
78
104 BuLLeTINn 272
EXPLANATION OF PLATE 5
Figure Page
1-8.. Adrianites, Gemmellaro, 1887) 0220.00... eee 43
1-3. Adrianites cf. A. insignis Gemmellaro, (BMNH C37654),
Sosio Limestone, Province of Palermo, Sicily, maximum
diameter—2144 mm, X 2.0; the specimen is mature.
(This specimen is figured as pl. 3, figs. 6-8, of Davis,
Furnish, and Glenister, 1969, but was referred to
Neocrimites by Furnish and Glenister.) 4,5. 4. insignis,
hypotype (MGPU unnumbered, Greco, 1935, pl. 3,
figs. 12a,b), Sosio Limestone, Pietra di Salomone,
Province of Palermo, Sicily, maximum diameter—29
mm, X 2.0; the specimen is mature. 6-8. A. kingi,
syntype (IGUP 81, Gemmellaro, 1887, pl. 9, figs.
31,32), Sosio Limestone, Rocca di San Benedetto, Pro-
vince of Palermo, Sicily, maximum diameter—22 mm,
xX 2.0; the specimen is mature.
PLATE 5
But. AMER. PALEONT., VOL. 62
BULL. AMER. PALEONT., VOL. 62
PLATE 6
PaLEeozoic AMMONOIDs: Davis 105
EXPLANATION OF PLATE 6
Figure Page
2 Neocrimites Ruzhencey.’ 194089 eR RRR a 51
1,2. N. ensifer, syntype (IGUP 82, Gemmellaro, 1887, pl. 6,
figs. 11,12), Sosio Limestone, Passo di Burgio, Pro-
vince of Palermo, Sicily, maximum diameter—24%4
mm, X 2.0; the specimen is immature.
ao (Crimifes oe roumanskayass SOS7D) 25.3 eieshe asec sesechececesseteessnanc:
3-5. C. krotowi (Karpinsky), (SUI 5981b), Artinsk Stage,
Aktasty River, Aktyubinsk District, USSR, maximum
diameter—23 mm, x 2.0; the specimen is mature. 6.
C. krotowi (Karpinsky), (SUI 5981a), Artinsk Stage,
Aktasty River, Aktyubinsk District, USSR, maximum
diameter—244%4 mm, X 2.0; the specimen is mature.
io: Adrianites Gemmellaro, 1887 ...........:.....ccccsccecssesccessetvestoeetens
7,8. A. insignis, syntype (IGUP 84, Gemmellaro, 1887, pl.
6, figs. 8,9), Sosio Limestone, Passo di Burgio, Pro-
vince of Palermo, Sicily, maximum diameter—27 mm,
X 2.0; the specimen is mature.
43
106 BULLETIN 272
EXPLANATION OF PLATE 7
Figure Page
1-5. Neocrimites Ruzhencev, 1940a 2.0.0.0... 51
1-3. N. ensifer, syntype (IGUP 82, Gemmellaro, 1887, pl. 7,
fig. 25), Sosio Limestone, Passo di Burgio, Province
of Palermo, Sicily, maximum diameter—32 mm, X
2.0; the specimen is mature. 4,5. N. ensifer (IGUP
82), Sosio Limestone, Passo di Burgio, Province of
Palermo, Sicily, maximum diameter—31 mm, X 2.0;
the specimen is mature.
BuLL. AMER. PALEONT., VOL. 62 PLATE 7
PLATE 8
62
PALEONT., VOL.
BULL. AMER
PaLEozoic AMMONOIDs: Davis 107
EXPLANATION OF PLATE 8
Figure Page
1-4. Epadrianites Schindewolf, 1931. ................0000. en 48
1-4. E. haueri, syntype (IGUP 75, Gemmellaro, 1888, pl. C,
figs. 13,15), Sosio Limestone, Rupe de San Benedetto,
Province of Palermo, Sicily, maximum diameter—66
mm, X 1.0; the specimen is mature.
108
Figure
1,2.
3-7.
8,9.
BULLETIN 272
EXPLANATION OF PLATE 9
Pseudagathiceras Schindewolf, 1931 .......00...000e
1G 20ehe
spinosum, holotype (YPM 16310, Miller, 1944, pl.
29, figs. 1,2), zone of Waagenoceras (beds 5, 6, and
7 of the Malascachas Section), 1200 m S17°W of
Noria de Malascachas, Valle de Las Delicias, Coahuila,
Mexico, maximum diameter—21 mm, xX 2.0; the speci-
men is mature.
Palermites Toumanskaya, 1987a oo000.0000000cccceeeececceeeeeeees
3-5. P. distefanoi (Gemmellaro), hypotype (SUI 32456, Mil-
ler, Furnish, and Schindewolf, 1957, fig. 2, ill. 5a,
5b), Sosio Limestone, Palazzo Adriano, Province of
Palermo, Sicily, maximum diameter—45 mm, X 1.2;
the specimen is mature. 6,7. P. distefanoi, syntype
(IGUP 76, Gemmellaro, 1887, pl. 9, figs. 36,37),
Sosio Limestone, Rupe di San Benedetto, Province of
Palermo, Sicily, maximum diameter—301%4 mm, X 1.2;
the specimen is mature.
Epadrianites Schindewolf, 1931) .......:05..4...2.....:.s eee
8,9. E. timorense (Boehm), hypotype (PIUB 21b, Haniel,
1915, pl. 5, figs. 7a,b), Koeafeoe, Timor, maximum
diameter (fide Haniel)—30 mm, 1.2; the specimen
is immature.
54
48
|
Butt. AMER. PALEONT., VOL. 62 PLATE 9
BULL. AMER.
PALEONT., VOL. 62
PLATE 10
Figure
1-3.
4,5.
6-8.
9,10.
PaLeozoic AMMONOIDs: Davis 109
EXPLANATION OF PLATE 10
Hoffmannia Genmellaro: 1887.3 ober. oct... kone 50
1-3. H. hoffmanni, syntype (IGUP 78, Gemmellaro, 1887,
pl. 7, fig. 3), Sosio Limestone, Passo di Burgio, Pro-
vince of Palermo, Sicily, maximum diameter—27 mm,
< 2.0; the specimen apparently is mature.
Neocrimites Ruzhencev, 1940a ................000ccccccccccceccceeeeeecceees 51
4,5. N. cancellatus (Haniel) (fide Furnish and Glenister in
Davis, Furnish, and Glenister, 1969), hypotype (PIUB
unnumbered, Wanner, 1932, pl. 9, figs. 3a,b; Davis,
Furnish, and Glenister, 1969, pl. 3, fig. 3), Basleo,
Timor, maximum dimension—22 mm, xX 2.0; the
specimen is mature.
Pseudagathiceras Schindewolf, 1931 0.000000... 56
6-8. P. wichmanni, holotype (THD 12752, Haniel, 1915, pl.
5, figs. 15a-c), Basleo, Timor, maximum diameter—
16% mm, X 2.0; the specimen is immature.
Texoceras Miller and Furnish, 1940a ..0000..0.000eee. 59
9. T. texanum (Girty), (SUI 32461 AA), Bone Spring
Limestone, road cut on the west side of US 62-180,
1.8 mi. NNE of the junction of US 54 & US 62-180,
Culberson County, Texas, maximum diameter—26%
mm, X 2.0; the specimen is mature. 10. T. texanum
(Girty), (SUI 32461 AJ), Bone Spring Limestone,
road cut on the west side of US 62-180, 1.8 mi. NNE
of the junction of US 54 & US 62-180, Culberson
County, Texas, < 5.1; figure shows approximation
and simplification of the ultimate few sutures.
110
Figure
BULLETIN 272
EXPLANATION OF PLATE 11
1-5,8-10.. Sizilites ‘Toumanskaya, 1937a 20.2... 0 ee eee
1,2. S. craticulatus, syntype (IGUP 87, Gemmellaro, 1888,
pl. C, fig. 16), Sosio Limestone, Rupe di San Bene-
detto, Province of Palermo, Sicily, maximum diameter
—1514 m, X 2.0; the specimen is mature. 3-5. S. affinis
(Gemmellaro), syntype (IGUP 88, Gemmellaro, 1888,
pl. D, figs. 6,7), Sosio Limestone, Rupe di San Bene-
detto, Province of Palermo, Sicily, maximum dia-
meter—14 mm, xX 2.0; the specimen is immature.
8-10. §. craticulatus, hypotype (MGPU unnumbered,
Greco, 1935, pl. 3, figs. 13a-f), Sosio Limestone, Pietra
di Salomone, Province of Palermo, Sicily, maximum
diameter—1014 mm, X 2.0; the specimen is immature.
6,7,11-15. Texoceras Miller and Furnish, 1940a ..............0000000000000...
6. T. texanum (Girty), (SUI 32461 Z), Bone Spring Lime-
stone, road cut on the west side of US 62-180, 1.8 mi.
NNE of the junction of US 54 & US 62-180, Culberson
County, Texas, maximum diameter—24 mm, X 2.0;
the specimen is mature. 7. T. texanum (Girty), SUI
32461 M), Bone Spring Limestone, road cut on the
west side of US 62-180, 1.8 mi. NNE of the junction
of US 54 & US 62-180, Culberson County, Texas,
maximum diameter—211%4 mm, X 2.0; the specimen
apparently is mature. 11. T. texanum (Girty), (SUI
32461 AG), Bone Spring Limestone, road cut on the
west side of US 62-180, 1.8 mi. NNE of the junction
of US 54 & US 62-180, Culberson County, Texas,
2.0; the specimen apparently is mature. 12. T. texanum
(Girty), (SUI 32461 I), Bone Spring Limestone, road
cut on the west side of US 62-180, 1.8 mi. NNE of the
junction of US 54 & US 62-180, Culberson County,
Texas, X 2.0; the specimen is mature. 13,14. T. tex-
anum (Girty), (SUI 32461 P), Bone Spring Limestone,
road cut on the west side of US 62-180, 1.8 mi. NNE of
the junction of US 54 & US 62-180, Culberson County,
Texas, maximum diameter—22% mm, X 2.0; the
specimen is mature. 15. T. texanum (Girty), hypo-
type (SUI 13609, Miller and Furnish, 1940a, pl. 26,
figs. 1,2), Bone Spring Limestone, near Bone Springs,
Guadalupe Mountains, Texas, maximum diameter—
25% mm, X 2.0; the specimen is mature.
59
Buu. AMER. PALEONT., VOL. 62 PLATE 11
N
ite)
3
(eo)
>
et
Pa
(e)
Q
4
<
fae
od
je3|
=
<
=
—]
(=)
—
Figure
PaLeozoic AMMONOIDs: Davis it
EXPLANATION OF PLATE 12
Page
i-S= Agathiceras; Gemmellaro, 1887 > .2c22....250ic...2. 00. .2008. 00s ceeencaen 61
1-4. A. suessi, syntype (IGUP 119, Gemmellaro, 1887, pl. 6,
figs. 1,2), Sosio Limestone, Passo di Burgio, Province
of Palermo, Sicily, maximum diameter—35 mm, xX
1.6; the specimen is mature (designated lectotype by
Furnish and Glenister in the explanation of plate 2
of Davis, Furnish, and Glenister, 1969). 5,6. A. suessi,
hypotype (MGPU unnumbered, Greco, 1935, pl. 3,
figs. 14a,b; Davis, Furnish, and Glenister, 1969, pl.
2, fig. 8), Sosio Limestone, Rocca di San Benedetto,
Province of Palermo, Sicily, maximum diameter—
37 mm, X 1.6; the specimen is mature. 7,8. A. suessi,
syntype (IGUP 119, Gemmellaro, 1887, pl. 6, fig. 4),
Sosio Limestone, Passo di Burgio, Province of Paler-
mo, Sicily, X 1.6; the specimen is mature.
112 BULLETIN 272
EXPLANATION OF PLATE 13
Figure Page
1-9. Agathiceras Gemmellaro, 1887 2000000... cocccoccceeeeeeeeeeeeeee 61
12a
uralicum (fide Furnish and Glenister in Davis,
Furnish, and Glenister, 1969), (SUI 32460 BA), Road
Canyon Formation, saddle and slope W of Dugout
Mountain, 0.3 mi. NNW of hill 4861 (USNM loc.
732z), Glass Mountains Uplift, Brewster County,
Texas, X 1.6; the specimen is mature. 3,4. 4. uralicum
(Karpinsky) (fide Furnish and Glenister in Davis,
Furnish, and Glenister, 1969), hypotype (SUI 32460
BN, ibid., pl. 2, figs. 3,4), Road Canyon Formation,
saddle and slope W of Dugout Mountain, 0.3 mi. NNW
of hill 4861 (USNM loc. 732z), Glass Mountains Up-
lift, Brewster County, Texas, maximum diameter—
19 mm, X 1.6; the specimen is mature. 5-7. A. urali-
cum (Karpinsky) (fide Furnish and Glenister in
Davis, Furnish, and Glenister, 1969), hypotype (SUI
32460 BE, zbid., pl. 2, figs. 1,2), Road Canyon For-
mation, saddle and slope W of Dugout Mountain, 0.3
mi. NNW of hill 4861 (USNM loc. 732z), Glass Moun-
tains Uplift, Brewester County, Texas, maximum
diameter—32 mm, X 1.6; the specimen is mature.
8,9. A. sp., (SUI 32458), Waagenoceras—zone shale,
about 1 km § of Noria de las Malascachas near a
goat trail on west side of draw, Valle de Las Delicias,
Coahuila, Mexico, maximum diameter—30 mm, X 1.6;
the specimen is mature.
PLATE 13
BULL. AMER. PALEONT., VOL. 62
|
|
PLATE 14
BULL. AMER. PALEONT., VOL. 62
Figure
PaLeozoic AMMONOIDs: Davis
EXPLANATION OF PLATE 14
AEG PPP OHINFOGMEAy Gry: 19OG 92a oc st es5p eet actospuander ep conse eee eros
1,2. P. erebus Girty, (SUI 32586), upper black limestone
member of Bone Spring Limestone, about 314 mi. SSE
of El Capitan on west side of US 62 just N of the
bend in road where it starts into Guadalupe Pass,
Culberson County, Texas, maximum diameter—18%
mm, X 2.0; the specimen is mature. 3. Peritrochia
n. sp., (SUI 32584), upper black limestone member of
Bone Spring Limestone, about 3%4 mi. SSE of El Capi-
tan on west side of US 62 just N of the bend in road
where it starts into Guadalupe Pass, Culberson Coun-
ty, Texas, X 2.3; the specimen is immature. 4-6. P.
erebus, (SUI 32585), Cut-off Shale Member of the
Bone Spring Formation, pipe line road S of El Capi-
tan, Culberson County, Texas, maximum diameter—
21% mm, X 2.0; the specimen is mature.
(ae Marathonites BGSeGs 199 a. ree ee sas ee eeaasaee
7. M. sp., (SUI 12301), Bitauni horizon, Bitauni, Timor,
maximum diameter—29 mm, xX 1.6, (see other views
on plate 21); the specimen is mature.
figs. 18a-c), Bitauni, Timor, maximum diameter—
35%4 mm, X 1.6; the specimen is immature. 9. 4. sp.,
(SUI 12297), Bitauni horizon, Bitauni, Timor, maxi-
mum diameter—37%4 mm, X 1.6; the specimen is ma-
ture.
113
85
61
114 BuLLeTIN 272
EXPLANATION OF PLATE 15
Figure Page
1-7. Mexicoceras Ruzhencev, 1955 .o......cccceeceeteecseeceeeees 66
1-4.
M. guadalupense guadalupense (Girty), hypotype
(USNM 144423, Miller and Furnish, 1940a, pl. 41,
figs. 3,4, pl. 42, fig. 6; Davis, Furnish, and Glenister,
1969, pl. 5, figs. 4,5), South Wells Limestone, about
2 mi. SE of D Ranch, South Wells, Culberson
County, Texas (USGS loc. 7649), maximum diameter
—48 mm, X 1.4; the specimen is mature. 5,6. M. guada-
lupense thompsoni (Miller and Furnish), syntype
(USNM 144419, Miller and Furnish, 1940a, pl. 25,
figs. 4-6), South Wells Limestone, about 2 mi. SE of
D Ranch, South Wells, Culberson County, Texas
(USGS loc. 7649), maximum diameter—18 mm, X 1.4;
the specimen is immature. 7. M. guadalupense guada-
lupense (Girty), hypotype (USNM 144422, Miller
and Furnish, 1940a, pl. 41, figs. 1,2), South Wells
Limestone, about 2 mi. SE of D Ranch, South Wells,
Culberson County, Texas (USGS loc. 7649), maximum
diameter—30 mm, X 1.4 (see other views on Plate
17) ; the specimen is immature.
PLATE 15
BuLL. AMER. PALEONT., VOL. 62
PLATE 16
BULL. AMER. PALEONT., VOL. 62
Figure
PaLEozoic AMMONOIDs: Davis 15
EXPLANATION OF PLATE 16
Page
1-7. Waagenoceras Gemmellaro, 1887 20.00.0000... 69
1. W. mojsisovicsi, syntype (IGUP 32, Gemmellaro, 1887,
pl. 1, figs. 1-3), Sosio Limestone, Passo di Burgio,
Province of Palermo, Sicily, maximum diameter—160
mm, X 0.4; the specimen is mature (designated lecto-
paratype by Furnish and Glenister in the explanation
of plate 5 of Davis, Furnish, and Glenister, 1969).
2,3. W. mojsisovicsi, syntype (IGUP 34, Gemmellaro,
1887, pl. 2, figs. 1,2), Sosio Limestone, Passo di Burgio,
Province of Palermo, Sicily, maximum diameter—95
mm, X 0.6; the specimen is mature (designated lecto-
type by Furnish and Glenister in the explanation of
plate 5 of Davis, Furnish, and Glenister, 1969). 4,5.
W. stachei, syntype (IGUP 37, Gemmellaro, 1887, pl.
1, figs. 4-6), Sosio Limestone, Passo di Burgio, Pro-
vince of Palermo, Sicily, maximum diameter—35 mm,
< 1.2; the specimen is immature. 6,7. W. richardsoni
(Plummer and Scott), hypotype (SUI 32318, Davis,
Furnish, and Glenister, 1969, pl. 5, fig. 7), very top
of a big channel sand just below grey limestone beds,
Manzanita Member, Cherry Canyon Formation, half-
way up slope some 1500 ft. ESE of Last Chance Wells,
Sibley Ranch, Culberson County, Texas, maximum dia-
meter—170 mm, xX 0.4; the specimen is mature.
116
Figure
BULLETIN 272
EXPLANATION OF PLATE 17
1,2. iMexicoceras Ruzhencev; 1955 © ):...20..0..0....2 ee
1,2. M. guadalupense guadalupense (Girty), same specimen
as Plate 15, figure 7, * 1.4; the specimen is immature.
3-8. | ‘Cyclolobus ‘Waagen: "1879" oe eee
3-5. C. walkeri Diener, hypotype (MNHN B 7517, Vaillant-
Couturier-Treat, 1933, pl. 2, fig. 6), Ankitohazo,
Madagascar, maximum diameter—23 mm, X 1.6; the
specimen is immature. 6. “C. astrei’, holotype (UPSM
unnumbered, Besairie, 1930, pl. 5, fig. 4), Ankitohazo,
Madagascar, maximum diameter—45 mm, X 1.2; the
specimen is immature. 7. C. walkeri Diener, hypotype
(MNHN B 7520, Vaillant-Couturier-Treat, 1933, pl.
1, fig. 4; Davis, Furnish, and Glenister, 1969, pl. 5,
fig. 1; Furnish and Glenister, 1970, pl. 2, fig. A),
Ankitohazo, Madagascar, maximum diameter—93 mm,
xX 0.6; the specimen is mature. 8. C. walkeri Diener,
(BMNH C36642), Ankitohazo, Ambilobe, Madagas-
car, maximum diameter—104 mm, x 0.6; the specimen
is mature.
|
PLATE 17
BuLL. AMER. PALEONT., VOL. 62
|
|
BuLL. AMER. PALEONT., VOL. 62 PLATE 18
Figure
Pa.Leozoic AMMONOIDs: Davis
EXPLANATION OF PLATE 18
Het o-ete WaaGOMinad ISTOCOW, TOO cee ce crore cnn ese e ane vareus-ceuneveraresets
7,8. Stacheoceras Gemmellaro, 1887
7,8. S. pelagicum Gemmellaro, hypotype (MGPU_ unnum-
1,2. W. perspectivum, syntype (IGUP 66, Gemmellaro, 1887,
pl. 8, figs. 12-14), Sosio Limestone, Rocca di San
Benedetto, Province of Palermo, Sicily, maximum dia-
meter—12 mm, X 2.0; the specimen is immature. 3,4.
W. subinterrupta, (SUI 32597), upper Artinsk Stage,
western slope of the southern Urals, Kyaly-Coorty
River, Soonyduk-Say, USSR, maximum diameter—15%4
mm, X 2.0; the specimen is immature. 5,6. W. darae,
syntype (IGUP 64, Gemmellaro, 1887, pl. 4, fig. 11),
Sosio Limestone, Passo di Burgio, Province of Paler-
mo, Sicily, X 0.8; the specimen is mature. 9,10. W.
darae, syntype (IGUP 64, Gemmellaro, 1887, pl. 4,
figs. 7,8), Sosio Limestone, Passo di Burgio, Province
of Palermo, Sicily, maximum diameter—544%4 mm, X
0.8; the specimen is mature. 11-13. W. darae, syntype
(IGUP 68, Gemmellaro, 1887, pl. 4, figs. 2,3), Sosio
Limestone, Passo di Burgio, Province of Palermo,
Sicily, maximum diameter—78 mm, 0.8; the speci-
men is mature.
Pietra di Salomone, Province of Palermo, Sicily, maxi-
mum diameter—934 mm, X 2.3; the specimen is imma-
ture.
7
78
118
Figure
BULLETIN 272
EXPLANATION OF PLATE 19
1-8. Stacheoceras Gemmellaro, 1887 ........0000000ccccccecccecccceceeeereeeeee
1,2. S. mediterraneum, syntype (IGUP 62, Gemmellaro, 1887,
pl. 7, figs. 11-13), Sosio Limestone, Passo di Burgio,
Province of Palermo, Sicily, maximum diameter—
1234 mm, X 2.3; the specimen is immature. 3,4. S.
toumanskyae Miller and Furnish, (SUI 1364 O), con-
cretionary shale (beds 5 and 7 of Difunta section),
along strike between Cerro Wencelao on the south
to 300 m W of El Indio on the north, Valle de Las
Delicias, Coahuila, Mexico (King’s loc. 20), x 0.6;
the specimen is mature. 5. §. sp. (SUI 32608 D), Tim-
orites-zone shale, King’s locality 20, Las Delicias,
Coahuila, Mexico, maximum diameter—85 mm, x 0.6;
the specimen is mature. 6-8. S. sp., (SUI 32610 A),
Timorites-zone shale, 1 km NE of Cerro El Indio,
Las Delicias, Coahuila, Mexico (near King’s loc. 10),
maximum diameter—107 mm, X 0.6; the specimen
is mature.
9: “‘Waagenina Krotow, 1888 0000 eee
9. W. subinterrupta (Karpinsky) (Krotow), (SUI 32597),
upper Artinsk Stage, western slope of southern Urals,
Kyaly-Coorty River, Soonyduk-Say, USSR, maximum
diameter—17%4 mm, X 2.0; the specimen is immature.
76
PLATE 19
BuLL. AMER. PALEONT., VOL. 62
- ea
BULL. AMER. PALEONT., VOL. 62
Figure
PaLtEozoic AMMoNnoIDs: DAvVIs 119
EXPLANATION OF PLATE 20
Page
1-8. Hyattoceras Gemmellaro, 1887 ................cc ee 82
1,2. H. turgidum, holotype (IGUP 51, Gemmellaro, 1887,
pl. 2, figs. 8,9), Sosio Limestone, Passo di Burgio,
Province of Palermo, Sicily, maximum diameter—23%
mm, X 2.0; the specimen is mature. 3,4. H. guembeli
Gemmellaro, hypotype (IGUP unnumbered, Davis,
Furnish, and Glenister, pl. 4, fig. 7), Sosio Limestone,
Pietra di Salomone, Province of Palermo, Sicily, maxi-
mum diameter—67 mm, X 1.0; the specimen is ma-
ture. 5-8. H. geinitzi, syntype (IGUP 52, Gemmellaro,
1887, pl. 8, figs. 1-3), Sosio Limestone, Rocca di San
Benedetto, Province of Palermo, Sicily, maximum dia-
meter—40 m, X 1.4; the specimen is mature (desig-
nated lectotype by Furnish and Glenister in the ex-
planation of plate 4 of Davis, Furnish, and Glenister,
1969).
120
Figure
BULLETIN 272
EXPLANATION OF PLATE 21
1-5,10. Hyattoceras Gemmellaro, 1887 ..........0..0..0.0ccilieceeeeeeeeeees
6,7-9. Marathonites Bose, 1919
6. M. sp., (SUI 12305), Bitauni horizon, Bitauni, Timor,
11-13. Pseudovidrioceras Ruzhencev, 1936
11-13. P. pygmeum, syntype (IGUP 65, Gemmellaro, 1887,
1-3. H. abichi, syntype (IGUP 55, Gemmellaro, 1887, pl. 2,
figs. 10-13), Sosio Limestone, Passo di Burgio, Pro-
vince of Palermo, Sicily, maximum diameter—22 mm,
< 2.0; the specimen is mature (designated lectotype
by Furnish and Glenister in the explanation of plate 4
of Davis, Furnish, and Glenister, 1969). 4,5. H. abichi
Gemmellaro, (USNM unnumbered), Sosio Limestone,
Rocca di Salomone, Province of Palermo, Sicily, xX
2.0; the specimen is mature. 10. H. geinitzi Gemmellaro,
hypotype (GPIT unnumbered, Davis, Furnish, and
Glenister, 1969, pl. 4, fig. 3), unlabelled, but almost
certainly from Sosio, < 1.4; the specimen is mature.
X< 1.6; the specimen is mature. 7-9. M. sp., same speci-
men as Plate 14, figure 7, * 1.6; the specimen is
mature.
pl. 8, figs. 15-17), Sosio Limestone, Rocca di San Bene-
detto, Province of Palermo, Sicily, maximum dia-
meter—1114 mm, X 3.9; the specimen is mature.
85
84
BuLL. AMER. PALEONT., VOL. 62 PLATE 21
PLATE 22
PALEONT., VOL. 62
BULL. AMER
PaLEeozoic AMMONOIDs: DAvIs 121
EXPLANATION OF PLATE 22
Figure
He 5-10) Marathonifes Bose, 1919) oc h.ciceiee liv eccsceccoseseceascoensoscerstevass 85
1,2. M. ganti (Smith), (SUI 17033), Graham (Wayland)
Shale, 64% mi. NW of Jacksboro, Texas, maximum
diameter—16 mm, X 2.0; the specimen is immature.
5-7. M. ganti (Smith), (SUI 1063), Finis Shale, 3%
mi. E of Jacksboro, Texas, maximum diameter—31%4
mm, X 1.6; the specimen is mature. 8-10. M. invari-
abilis Ruzhencev, (SUI 10594 A), Artinsk Stage,
Aktyubinsk District, Aktasty River, USSR, maximum
diameter—51¥4%4 mm, X 1.2; the specimen is mature.
See Peritrocnia Girty: TOOG Ree poe vin. ee Abt issccanscscneee eee 88
3,4. P. erebus Girty, (SUI 32583 E), Bone Spring Lime-
stone, road cut on the west side of US 62-180, 1.8
mi. NNE of the junction of US 54 & US 62-180, Cul-
berson County, Texas, maximum diameter—11 mm,
xX 2.3; the specimen is immature.
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INDEX
Note: Light face figures refer to page numbers. Bold face figures refer
to the plate numbers.
A
a, Marathonites
krasnopolskyi forma 76
PAIDICHTAM erence. 82
abichi, Hyattoceras
(Aichi)! io 21824835120
Académie Moirans
(see Collignon)
adamsi, Adrianites .... 52
Metacrimites .......... 52
Neocrimites ............ 52
adrianense,
Waagenoceras ...... 70
Adrianites ...... 3,4, 5,6 43-46, 47, 49,
50, 51, 52, 54,
5, 102) 103:
104, 105
Adrianites
(otimannia) =... 50
Adrianites
(Neocrimites) .......... 52
affinis,
Adrianites! 2.9)... 57
Sizilitess ee 11 57, 58, 110
Agathiceras ..12, 13, 14 33, 36, 43, 47,
49, 52, 56, 61-
G5 ule:
113
Agathiceras
(Doryceras”)) .2.-...-. 56
Agathiceras
(Gaetanoceras) ........ 61
Agathiceras
(Paragathiceras) ...... 61
Sat NOCerAS -....++..1e2 59, 61
AIS ALICeT AS Een ee 61
Alktasty River .........% 105, 121
Aktyubinsk District... 105, 121
Aildnich, Be Ay... 4. 41, 42,94
PAIMILCS. Bens ke es. 86, 87
Ambilobe,
Madagascar .............. 116
AMNH — American
Museum of Natural
ERISUOE Vid cee ae 3: 29, 69, 71, 72,
73
Amsterdam, Geologisch
Instituut der Univer-
siteit van (see CIUA)
anceps, Adrianites .... 61
andrussowi,
Stacheoceras ............ 79
Ankitohazo,
Madagascar .............. 116
antiqua (-um, -us),
Arecestes) 3 2a ener! 79
Stacheoceras ............ 79, 81
applanatum,
Agathicerasy 2.2.25 61
applini, Agathiceras.... 61, 62, 65
(AT COSTCS 3) eth neds 73, 79
Arcestes (Cyclolobus).. 73
argentinus, Ilex
illecebrosus .............. 41, 42
Argonautal o...cteeee 42
Aricoceras .................- 44, 51, 52
Aricoceras
(Metaricoceras) ........ 51, 52
Aricoceras
(Neoaricoceras) ...... 44
ArkanSasiacensckc none 39
arthaberi,
Stacheoceras ............ 79
Artinsk Stage 0.0.0.0... 105, 117, 118,
121
asiaticum, Agathiceras 61
asselicum, Agathiceras 61. 62
astrei, Cyclolobus .17 73, 76,116
asvmmetrica,
Marathonites
krasnopolskyi var. .. 76
B
b, Marathonites
krasnopolskyi forma 76
iBasleocerasi..3 ee 49
Basleowehimor......0e 109
benedictinum,
Stacheoceras ............ 76
Waagenina .............. 76
beschui, Agathiceras .. 61
beyrichi,
Agathiceras. .............: 49
Basleocerasi...........) 49
Epadrianites ............ 49, 50
biassalensis,
Adrianites insignis
Ae ie eee 44
Neocrimites (Sosio-
crimites) insignis
if fick t 5) Soe ee ey 44
bidentata,
Marathonites
krasnopolskyi var. .. 76
Bisaunin Limon 113, 120
123
INDEX
BMNH — British
Museum (Natural
HIStOGY) eee 29, 46, 65, 75,
76, 83, 104,
116
bobkovae, Adrianites.. 44
bobkowi, Adrianites .. 44
bodraki, Agathiceras.. 61
boesei, Stacheoceras .. 79
Bone Springs
Kimestone) . ==.) 89, 109, 110,
113, 121
Bone Springs, Texas .. 110
borissiaki,
Stacheoceras ............ 79
Brewster County,
TEXAS ok i ge 112
brouweri, Agathiceras 61, 65
burgensis, Adrianites
(Hoffmannia) .......... 50
Hofimannia, 22 50
busterensis,
Crimites. 47
Marathonites
(Allmites)) -2..0.....-.- 86
Cc
c, Marathonites
krasnopolskyi forma 76
Campbell iG. By =. 37, 91
cancellata (-um, -us),
Agathiceras ............ 52
Aricoceras
(Aricoceras)) -.....--.... 52
Neocrimites ....3,10 52,54, 102,
109
Carboniferous ............ 34, 37, 39
Cerro El Indio ............ 118
Cerro Wencelao .......... 118
Cherry Canyon
Hormation... a 115
Cincinnati,
University of .2........ 27, 29, 101
ciscoense, Agathiceras 61, 65
Clark] Davidil. = 47, 48, 79, 81,
95
clavatum,
Waagenoceras ........ ;
Cline) 7 Me Sime 79, 80-81, 84,
; 85, 87, 95
clinei, Pamirites ........ 76
Waaceninagees se. 76
Coahuila 108, 112, 118
coahuila,
Waagenoceras ....... 70
coindetii, Illex
illecebrosus .............. 41, 42
Collignon; M20 es 29, 75, 76
collignoni,
Popanoceras ............ 79
Stacheoceras ............ 79
contractum,
Agathiceras .............. 61
Copenhagen,
University of .......... 29, 75, 81
craticulatus,
Adrianites ................ 57
Sizilites) .....:.. 283 1). 5.585420
Crick) (Gs Cy. .c.08 eee 42, 92
crimensis, Stacheo-
ceras mediterraneum
79
47-48, 105
Crockford, M. B. ........ 52, 54, 95
Culberson County,
TOXASH, Hee cscs 109, 110, 113,
114, 115, 121
cummingsi,
Waagenoceras ........
Cyclolobus) 222... 17 33, 34, oa
Cyclolobus
(Krafftoceras) ........ 73
D
d, Marathonites
krasnopolskyi forma 76
darae, Stacheoceras .... 77
Waagenina .......... 8 77,78 417
darensis, Crimites
pamiricus var. ........ 47
Dear Ws eee 54, 90
DeBlainville (see
Blainville)
defordi,
Adrianites) 52
Neocrimites ............ 52, 54
delawareanus,
Marathonites.......... 59
Delicias, Valle de Las 108, 112, 118
Denton, Eric J. ............ 30, 32
diblasii,
Stacheoceras ............ 79, 80
Dierners Carlie 79
dieneri,
Marathonites .......... tL
Waasenina
Waagenoceras ........ 52, 10; dlewe.
Waagenoceras
dieneris {0 see 73
Dituntae eee 118
difuntense,
Pseudagathiceras .... 51a}, BY
INDEX
discoidale (-is), fultonensis,
Agathiceras cancel- Gontatites) ic. /nes... 62
latum’ form’ .8.-.2...; 52 Surnish, W.-M: «..:.....-.. 28, 29, 31, 36,
Aricoceras (Metari- 42, 43, 44, 46,
coceras) cancellatum 47, 48, 50, 51,
fOEMG-.. ee 52 5200, 04055
Neocrimites ............ 52, 54 56, 57, 58, 59,
Stacheoceras ............ 79 60, 61, 62, 63,
distefanoi, 64, 65, 66, 67,
ACriAanites, <<... sce.s 68, 69, 70, 71,
Palermites ............ 9 54, 55, 56 72, 73, 74, 75,
76, 78, 79, 81,
djidairensis, Stacheo- 82, 83, 84, 85,
ceras kussicum var. 86 86, 87, 88, 89,
Dorycerase 56 91, 92, 94, 95,
Downs been: tks... 64, 88, 89, 95 96, 102, 103,
Dugout Mountain, 104, 108, 109,
ROXAS Hc ea. 2: IOS Labs alee
dubari, 114, 115, 116,
Adrianites 200..0c006. 52 119, 120
Neocrimites ............ 52, 53, 54
Peritrochia .............. 84 G
: Gaetanoceras .............. 61
e, Marathonites ganti,
krasnopolskyi forma 76 Marathonites ...... 22 85, 86, 87, 88,
El Capitan, Texas ...... 113 121
electraensis, Marathonites
Marathonites .......... 86 (Marathonites) ........ 87
elegans, Popanoceras ............ 85
Adrianites .....:..:.. 3 44,45,46, gaudryi,
102 Stacheoceras .......... 4 79, 81, 103
Eleganticeras .............. 42 geinitzi,
elegantulum, Hyattoceras ..20,21 82, 83,119,
Eleganticeras .......... 42 120
elkoense, Crimites ...... 47 Gemmellaro, G. G. ...... TU
ensifer, gemmellaroi,
Adrianites ................ 52 Adrianites) 22:52. 47
Aricoceras Stacheoceras .......... 79, 80, 81
(Aricoceras) ......... 52 Waagenoceras .......... 70
Neocrimites ...... 6,7 52, 53, 54, Wanneroceras ........ 69
105,106 Gertman, Richard ...... 33, 91
Epadrianites .......... 8,9 48-50,107. girtyi,
108 Agathiceras ............ 61, 62, 65
erebus, Pseudovidrioceras .. 84, 85
Peritrochia ....14, 22 88, 89, 90, Vidrioceras .......:.:.-.. 84
11313}, RAL Waagenoceras
GUGHEEI HS ® eee. 70, 73
F Glass Mountains,
f, Marathonites REX ASM hh eee 64, 112
krasnopolskyi forma 76 Glenister, Brian F. .... 28, 29, 34, 43,
Hinisi Shales... 121 44, 46, 50, 51,
fisheri, Hoffmannia .. 50 52, 54, 56, 57,
mletcher Hei: eee 52, 98 58, 61, 63. 64,
frechi, Agathiceras .... 61, 62, 65 65, 69, 70, 71,
fredericksi, TO aen sD,
Adrianites|) 51 76, 78, 81. 82,
Neocrimites............ 51, 54 83, 84, 85, 86,
125
INDEX
87, 88, 89, 91,
92, 94, 96, 102,
103, 104, 109,
111, 112, 114,
115, 116, 119,
120
globosa (-um, -us),
AGrianites) es... 47
Agathiceras can-
cellatum form ........ 52
Agathiceras
Marini Vara es2-s:..5.0: 52, 62
Mexicoceras ............ 66
Neocrimites ............ 52, 54
Stacheoceras ............ 79
globulinum,
Stacheoceras .......... 79
glomulus, Crimites .... 47
Godthaabites .............. ie
GOntatIteS ie... seen. 62
gordoni, Stacheoceras 79
gracilis,
Marathonites .......... 86
Grahams sAe @6....... asd 39, 42, 92
Graham (Wayland)
Shale ek sie eh 121
Griocerasiees....... eat 76
gruenewaldti,
Stacheoceras .......... 79
gruenwaldti,
Stacheoceras ............ 79
Guadalupe Mountains,
ROXAS HR ais 110
guedalupense,
Mexicoceras_ ..15, 17 66, 67, 68, 69,
14, 116
Mexicoceras
guadalupense 15,17 69, 114,116
Waagenoceras
cummingsi var. ........ 66
Waagenoceras .......... 66
W aagenoceras
guadalupense .......... 66
Guadalupe Pass,
TEXAS es ee ee As
guembeli,
Hyattoceras ......... 20 #82. 83.119
Gurley; W.'F. Bec. 62, 64, 95
H
hanieli,
AGrianites- 47
Neostacheoceras .... 79
Stacheoceras ............ 79
haueri,
Adrianites! 22... 49
Basleoceras .............. 49
Epadrianites .......... 8 49,50, 107
haydeni, Cyclolobus
(Krafftoceras) .......... 73, 74, 75
hildrethi, Ammonites 62
Hofimannial =. 10 50-51, 109
hoffmanni,
Adrianites
(Hoffmannia) ........ 50
Hoffmannia .......... 10 350,51, 109
hoshanensis,
Cyclolobus ..........- 74
Hyattoceras ...... 20, 21 33, 43, 82-84,
119, 120
Hyattoceras (Abichia) 82
illecebrosus, Illex ...... 41, 42
DDK Shy eee eee is 41, 42
insignis,
Adrianites ......... 5, 6 44, 45, 46, 52,
104, 105
€yclolobus) 2s 73, 74, 75
Neocrimites. ............ 44, 52
Neocrimites
(Sosiocrimites) ........ 44
invariabilis,
Marathonites ..... 22 86, 87. 88,
121
Marathonites
(Almites)) 2 87
involuta (-um, -us),
Agathiceras
timorense var. ........ 49
Epadrianites ............ 49
TOWalPee es eo ae 37
ircutensis, Agathi-
ceras frechi var. .... 61
isomorphus,
Adrianites ........ 3,4 44.45, 46,
102, 103
ITB—Institut Tekno-
logi Bandung .......... 29
iwaizakiense,
Stacheoceras .......... 79
J
Jacksboro, Texas ........ 124
jazwae, Waagenia ...... 77
J. P. Smithi,
Marathonites .......... 29, 85, 88
Stacheoceras
(Marathonites) ........ 29, 85
126
INDEX
K
Karpinskyi,
Stacheoceras ............ 79
Waagenoceras
GICNEEI et hens 70, 71
katschae, Agathiceras 61
Khimshiashvili, N. G... 31, 91
kingi,
Adrianites ........... 5 44, 45, 46,
104
Aricoceras
(Neoaricoceras) ...... 44
kinshanense,
Agathicerasi! oe. 62
kitschiensis, Agathi-
ceras anceps var. .... 61
Kinapps WhroDitet...c-.c.-- 42,92
knoxense,
Pseudorthoceras ...... 37
Knoxville, Iowa .......... 37
Koeafeoe, Timor ........ 108
koswae,
Neocrimites ............ 52
Waagenia =~. 300..4 52
kraffti, Cyclolobus
(Krafftoceras) ........ 73, 74
KeratitoO@eraS) .......6.0-.... th
krasnopolskyi,
Marathonites ............ 76
Popanoceras ............ 76
krotovi, Crimites ........ 47
krotowi, Agathiceras 47
Crimitesey, 6 47,48, 105
kullingi, Cyclolobus ... Tey te!
Godthaabites .......... 73
kussicum (-us),
Marathonites .......... 86
Stacheoceras ............ 86
Kyaly-Coorty River .... iolze, 1b ks}
L
Keibroviteh, 1. S: .2... 28, 46, 50, 51,
60, 61, 66, 69,
MO olos das
75, 16. 10,01,
82, 83, 85, 88,
89, 90
lidacense,
Waagenoceras ........ Ode
WRONG ON pete 32, 34, 41
ILLITE RC ORE Opie ih gaa sobres 41, 42, 94
M
macromphalus,
Nautilus 22. fee ee 42
Madagascar .................. 116
madagascariensis,
Cyclolobus walkeri
ia Taarceeee Laced. Ska 74
Malascachas, Noria de 108, 112
Mangold-Wirz, 1963 .... 39, 41, 42, 94
Manzanita Member .... 115
Maoen Mollo, Timor . 102
marathonensis,
Adrianitess....o.. 52
Neocrimites ............ 52
Marathonites 14, 21, 22 29, 43, 59, 76,
77, 84, 85-88,
31205121
Marathonites
CAlmaItES)e.8 en, 86, 87
Marathonites
(Pseudovidrioceras) 84, 86, 87
marcoui, Stacheoceros 79
martini,
Agathiceras ........ 14 52, 61, 62, 65,
113
Martocerasivete. 76
mediterraneum,
Agathiceras ............ 62
Stacheoceras ........ 19 79, 80, 81,
118
meridionale (-is),
Adrianites (Neo-
Crimites)) eee 52
Aricoceras (Ari-
COCEraAS) eee 52
Neocrimites ............ 52
Merriam, C. W.. ............ 81, 92
Metacrimites .............. 51, 52
Metaricoceras .............. Hie?
MexiCOlg ee eee. 108, 112, 118
Mexicoceras ...... 15, 17 66- 69, ta
MGPU—Istituto di
Geologia e Paleon-
tologia dell’Univer-
SitaudisPisa, soe 29, 46, 58, 65,
81, 104, 110,
ib tal ealilyy
Miller, ARK ®. 8s: 28, 31, 89, 96
minor,
Adrianites diste-
PAN OUR Aes 54
Palermites? <.:......0. 54
minutum,
Waagenoceras Sh ee 70
MNHN — Muséum
national d’Histoire
naturelle, Paris ...... 29, 75, 76,
116
moisisovicsi,
Waagenoceras ..16 69, 73,115
127
INDEX
murgabensis, Crimites
subkrotovi var. ........ 47
N
nalivikini,
Adrianites ................ 52
Neocrimites ............ 52
INautilUS ee eee SONGS 2ea3:
34, 35, 36, 41,
42
Neoaricoceras ............ 44
Neocrimites 3,6,7,10 44, 51-54,
102, 104, 105,
106, 109
Neocrimites
(Metacrimites) ........ HIS?
Neocrimites
(Sosiocrimites) ........ 44
Neostacheoceras ....... 78, 79
newelli,
Adrianites: =....-..... 52
Neocrimites ............ 52, 54
Neocrimites
(Metacrimites) ........ 52
Newellites ...000000...00..8 70, 71
nikitini,
Waagenoceras ........ 70
novomexicana,
Peritrochia -......0.. 84
oO
oblatum,
Syngastrioceras ...... 39
OCtOpUSH eee. 32, 41, 42
officinalis, Sepia ........
oldhami,
Cyclolobus ©..2%...... 73, 74, 75, 76
Phylloceras .............. 73
oyensi, Agathiceras .... 47
Pp
Pakistan, Geological
Sunveyioleeees 29, 76
Palazzo Adriano ........ : 102, 108
Palermitesi....-. 9 54-56, 108
105, 106, 107,
108, 109, 110,
PALE,
118, 119, 120
pamiricus,
Crimitess ee 47
Marathonites
CAlmites)) eee 86
Raminitesmee 76
Paraceltitesa 36
Paragathiceras ............ 61
Raratrochiay- =e 88
Barizek, Be Jo ccc 84, 85, 89, 95
Passo di Burgio .......... 102, 103, 105,
106, 109, 111,
115) eas:
119, 120
pelagicum,
Stacheoceras ........ 18 79,8
Peritrochia ........ 14, 22 84, 88-90,
iais} 1PAl
perglobosum,
Waagenoceras
Mi katina vate 70
permicus, Adrianites. 44
perspectivum,
Stacheoceras ............
Waagenina ......... 18.) 77; 78107
persulcatus,
Arcestes (Cyclo-
LObUS) Pe eee 73
Cyclolobus 73
Phylloceras®<...........! 73
PIUB — Institut fiir
Palaontologie der
Rhein. Friedrich-
Wilhelm Univer-
Sitat; Bonn’. 2.) ...... 29, 49, 50, 54,
83, 108, 109
plana (-um, -us),
Adrianites) =... 44
Agathiceras ............ 61
plummeri,
Adrianitess— 52
Neocrimites ........... 52
pompilius, Nautilus .... 42
Popanoceras ................ 76; 77,79, 85.
pressulus, Mara-
thonites (Almites) ... 86, 87
Procyclolobus .......... 73
Pseudagathiceras 9, 10 33. 56-57,
108, 109
Pseudorthoceras si
Pseudovidrioceras . 21 84-85, 86,
87, 120
pygmeum,
Pseudovidrio-
CeraSey er oe 21 ~=84, 85, 120
Stacheoceras ............ 84
Q
quadridens,
Stacheoceras ............ 79
quadridentata,
Marathonites
krasnopolskyi var..... 76
Quenstedtoceras ........ 35
128
INDEX
R
richardsoni,
Newellites ................ 71
Waagenoceras ....16 70, 71, 73
115
Waagenoceras
Gener ee 70
Road Canyon
HoOrmatlOonue scene 43, 64, 112
romanowskyi,
Marathonites
CAlmaIteS) ees ccna: 86
Popanoceras ............ 86
rothi, Stacheoceras .... 79
rothpletzi,
Acaunicerasmuee.s 47
Runnegar, Be ..:ccsse 54, 90
Ruzhencevy, V. E. ........ 28, 46, 50, 51,
60, 61, 66, 69,
70, 72, 73, 74,
75, 76, 78, 81,
82, 83, 85, 88,
89, 90
S
Salomone, Rocca
(Rietra) dies... 103, 104, 110,
117, 119, 120
San Benedetto, Rupe
ROCCA)EGIe ate 103, 104, 107,
108, 110, 111,
117, 119, 120
San Cologero, Rupe di 103
sarabansis, Agathi-
ceras anceps var..... 61
Schindewolf, O. H. .... 69, 72, ae
schmidti, Adrianites .. 44
Sea kik CS eee 50, 51, 59, 60,
61, 64, 66, 68,
69, 70, 72, 76,
81, 83, 85, 86,
87, 89, 96
sellardsi,
Marathonites 86, 87, 88
SISYOIIEY | ae SAN oe ope a 32, 34, 41
MEDION. tse ates 41
sexlobatum,
Stacheoceras .... 79, 81
Sipley, Ranch. .......... 115
ICTY ec oe 102, 103, 104,
105, 106, 107,
108, 109, 110,
Talk Wales alalze
118, 119, 120
singularis,
Crimites ........ 47
Sizilites 2. ee 11 33, 57-58,
110
smithi,
Crimites.- ee 47
Mexicoceras
guadalupense .......... 66, 67
Waagenoceras
guadalupense .......... 66
Solenochilust=........... 34
Soonyduk-Say, USSR... 117, 118
sosiense,
Stacheoceras .......... 79
Sosio Limestone ........ 102, 103, 104,
105, 106, 107,
108, 109, 110,
1D Liye
118, 119, 120
Sosiocrimites .............. 44
South Wells
Limestone ................ 114
South Wells, Texas .... 114
spathi, Crimites .......... 47
Spinosa, Claude .......... 30, 36
spinosum,
Pseudagathiceras .9 56,57, 108
stachei,
Waagenoceras ..16 70,73, 115
Stacheoceras 4, 18, 19 29, 76, 77, 78-
81, 84, 86,
103, 117, 118
Stacheoceras
(Marathonites) ........ 85
Stacheoceros ................. 7
Stehli, Francis G. ...... 29, 65
stuckenbergi,
Agathiceras ©...........:. 52
Neocrimites ............ 52
subcumminsi,
Cyclolobus' .2........... 74
subgeinitzi,
bvattoceras)..---.- 82, 83
subinterrupta,
Wiaacenias es 76
Waagenina 18,19 76, 78, ats
8
subkrotovi, Crimites .. 47
subkrotowi, Crimites.. 47
suessi,
Agathiceras ........ 12 33, 61. 62, 63,
64, 65, 111
suevi,
Marathonites .......... 86
Stacheoceras ............. 86
suleatus, Marathonites 86
sundaicum,
Asathieerdsi nes 61, 62, 65
Syngastrioceras ..........
129
INDEX
T
tegengreni,
Asathiceras <....—5 62
teicherti, Cyclolobus .. 73
tepense, Stacheoceras 79
texanum,
Agathiceras ............ 59
Texoceras ...... 10,11 59-61, rere
eb KR or care 43, 64, 89,
109, 110, 112,
113, 114, 115,
121
Texas Tech
Wniversitys —.2......--- 29,71
Texas, University of . 29, 68, 69
Texoceras’......... 10, 11 59-61, 109,
110
THD—Mineralogisch
Geologisch Museum
Technische Hoge-
school Delft” =.....= 29, 57, 65,
109, 113
thompsoni,
Mexicoceras guada-
LUPENSE <.:.:8%.....:-: 15 ~=66, 69, 114
Waagenoceras
guadalupense ........ 66
tietzei, Stacheoceras 4 79, 80, bc
FIN TTA OTe kee ree ese eee 102, 108, 109,
113, 120
timorense,
Agathiceras ............ 49
Epadrianites ......... 9 49,50, 108
Popanoceras ............ 77, 79
MUMOLUCSee ee ceeeeee 69, 70, 71
tornatum,
AN SaUMICeLAS a eeceeeee 62, 65
toumanskyae,
Stacheoceras ..... 19 79, 80, 81,
118
tridens,
IATCOStES eee 79
Popanoceras ............ 77, 79
Stacheoceras ............ 79, 80, 81
tridentata,
Marathonites
krasnopolskyi var. .. 76
trimarti, Stacheoceras 79
trimurti, Stacheoceras 79
tschernyschewi,
Stacheoceras ............ 79
turgidum,
Hyattoceras ....... 20 82, 83, 84,
119
U
undosus, Crimites ...... 47
Unklesbay, A. G. ........ 69, 72, 81, 95
UPSM—University of
Paris, Sorbonne .... 29, 75, 76,
116
uralensis,
Marathonites .......... 86, 87
uralica (-um, -us),
Agathiceras ........ 13 61, 62, 63, 64,
65, 112
Goniatites ................ 62
Marathonites .......... 86
Stacheoceras ............ 86
Ural Mountains .......... 117, 118
USNM—tUnited States
National Museum .. 29, 45, 46, 51,
55, 58, 65, 69,
76, 83, 103,
112, 114, 120
WSS yy weet es ee 105, 117, 118,
121
Vv
vidriensis,
Marathonites .......... 86
Widriocerasn eee 84
vinogradovi,
IAdrianitese ne 44
vogti, Stacheoceras .... 79
WwW
Wiaaseniay v...c5:.-3e oe 52,06
Waagenina ........ 18, 19 43, 76-78,
117, 118
Waagenoceras ........ 16 52, 66, 69-73,
115
walkeri,
Cyclolobus .......... 17 73, 74, 75, 76,
116
Wanneroceras ............. 69
warreni,
Adrianites) = 52
Neocrimites ............ 52
wichmanni,
Agathiceras
(Doryceras?) ........ 56
Pseudagathiceras 10 56, 57, 109
Wiggett, Gail .............. 33, 91
Y
Youngquist, W. .......... 64, 88, 89, 95
YPM—Yale Peabody
MUSCUINI eee 29, 57, 108
130
LIII.
LIV.
LV.
AYE
LVII.
LVIII.
LIX.
LX.
LXI.
LXII.
Volume lI.
II.
Il.
FY.
VI.
VII.
(Nos. 225-230). BUS UD Des 4 Gres 2e-eale, ee kN arash less aie or oie ee 18.00
Venezuela and Florida cirripeds, Antarctic forams, Lin-
naean Olives, Camerina, Ordovician conodonts, Niagaran
forams.
(NOs tes ls2S2\ enn 42 Op Diy LON DISS. ces ceeccecenerecteenstereenesemenencrecteas 18.00
Antarctic bivalves, Bivalvia catalogue.
(INGS4233-0250) ou 1S SD Die 43 eS 5) etetese weno esac tae ee eee eee 18.00
New Zealand forams, Stromatoporoidea, Indo-Pacific, Mio-
cene-Pliocene California forams.
(INiosi237-238) ie) 2 488tipp.p 45) Is. cascc28 cee. coces od cctcceconse rereeceeee te 18.00
Venezuela Bryozoa, Kinderhookian Brachiopods.
(Nos. 239-245). SIO Dts OD |Sceeeeeeesense cent eens eee ee 18.00
Dominican ostracodes, Texan pelecypods, Wisconsin mol-
lusks, Siphocypraea, Lepidocyclina, Devonian gastropods,
Miocene Pectens Guadaloupe.
(Nos. 246-247). HEA Toy oh (10h mi ONES ES 5: <2. ee ee an emS Se Be 18.00
Cenozoic corals, Trinidad Neogene mollusks.
(Nos. 248-254). SZ) RD PA ADS syee ose ote ete es ST 18.00
American Foraminifera, North Carolina fossils, coral types,
Belanski types, Venezuelan Cenozoic Echinoids, Cretaceous
Radiolaria, Cymatiid gastropods.
(Nos. 255-256). SOI p pee G2 al Sel es ce es Od ok glee re i 18.00
Jurassic ammonites.
(Nos. 257-262). EKO Y a) ohre SIL). 0) pee eco ead te ee el Sen ne 18.00
Cretaceous Radiolaria, Cretaceous Foraminifera, Pacific
Silicoflagellates, North American Cystoidea, Cincinnatian
Cyclonema, new species Vasum.
(No. 263). SITE To) 0 ORs eR ea RIA aie | aa ee Ue MA SI Ree 18.00
Bibliography of Cenozoic Echinoidea.
(Nos. 264-267). SSIS De yO Se Pl Sey, ete eee ee eee ee Pea 18.00
Jurassic-Cretaceous Radiolaria, cirripeds, Bryozoa, paly-
nology.
(Nos. 268-270). BOSD De Sleep Sse ee eee ee ee 18.00
Mollusks, Murex catalogue, Cretaceous Radiolaria.
ONTOS 2 0127.4 te ees coos See enn ae ree 18.00
Trace fossils, ammonoids, Silicoflagellates, microfauna.
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(Nos. 26-33). AD DARTS TM / OMT NS Cees co 75. plsty--.-a2e ee
Forams, ostracods, mollusks, Carriacou, fossil plants.
(Nos: 177-183) .; -448 pps, 36splss 222 a
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(No 184)... (996 spp. 1 ipl ee ee
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(Nos:0185-192)). S80 ppes S'Swiplsaet eee eet
Forams, mollusks, carpoids, Corry Sandstone.
CINog 193) 35673 pp ae 48) (plsy 5 ote ee eee eee ee
Venezuelan Cenozoic gastropods.
GNoss1942198) 99427 “pips 20M pl sa nice ce rere oe ee te ee
Ordovician stromatoporoids, Indo-Pacific camerinids, Mis-
sissippian forams, Cuban rudists.
(Nos: 199-203) 4. 365; eppsi68 ‘pls: 222 eee
Puerto Rican, Antarctic, New Zealand forams, Lepidocy-
clina, Eumalacostraca.
(No: 204) S647 spp: 6S) spss a-ha eee eons ee
Venezuela Cenozoic pelecypods.
(Nos::205=211). 1 409\pp:, 70) plss-2..22.n xe
Forams, Crustacea, brachipods, Recent mollusks.
(Nos: 212-217)... 584 pps-83) plsijn2ccs er ee ee
Forams, mollusks, polychaetes, ammonites.
(No.218).) 1058 pp:; Siplss) 022s es ee ee
Catalogue of the Paleocene and Eocene Mollusca of the
Southern and Eastern United States.
(Nos: 219-224) ;.''\.671-1pps, ‘83: pls. 34.2 ee
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No. 273 HARVARD
WNIVRRSITY,
UPPER CRETACEOUS AND CENOZOIC
SILICOFLAGELLATES AND EBRIDIANS
By
HSIN YI LING
1972
Paleontological Research Institution
Ithaca, New York 14850, U.S.A.
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BORE EINS
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PALEONTOLOGY
(Founded 1895)
Vol. 62
No. 273
UPPER CRETACEOUS AND CENOZOIC
SILICOFLAGELLATES AND EBRIDIANS
By
HSIN YI LING
October 18, 1972
Paleontological Research Institution
Ithaca, New York 14850, U.S.A.
Library of Congress Card Number: 72-86843
Printed in the United States of America
Arnold Printing Corporation
CONTENTS
Page
DANDSS E:1ck CB eee eres eee ee ee ae eee eee neeoze ee fee ee rea ceebbectoc 135
DINAN EGO ELEANOR OY ee A ee Sec ead hee ar SE SS eee ee eer 135
PN ekmonvyil CCST Sy soem ae eon enna bop Seti Aree SA en an ee ten oer 136
Samples Studied seers ccna enews eee cee ee ees ee ae 137
1Wal ara t OT yam PT OCC GUT Se eeetrs eee a ne owes wet ce cee eee er eee e ee 144
Systema tiG mill CLOpall COLO] O ay aece seesaw aera ce erence eaeaeneee 145
Briss ASH ICGpon SE Wee EC pear eRe re ae ene nee 146
bytEbridians® 245 ee, ee eee Se ee 195
ODE SC 11S S 1 Or OEP ee aE ee ee ee coe SecA es tevin Saez ape svtes deettiee 199
Eley LOH 0 Vu) meeps 2 Bee Oe ee A ee eee eee eee ee 199
pamiViedtternane anim (litaly,)me ne sito Ty eae eee ieee ne ee 202
TRAST S ESE pe A i ga ee ace ah ear te ke te hy doe oc Ree 205
Relate Siero: eee es EN 5 OE eee eye ae eS i eae 213
TEXT-FIGURE
1. Index map showing the geographic location of samples investigated .... 139
2. Distribution of silicoflagellates and ebridians in upper Cretaceous and
ea COR EME: SC CEO TN yas aes a eee ce Oe nacre eee ee a ee 140
3. Distribution of silicoflagellates and ebridians in Neogene section ............ 141
4. Distribution of silicoflagellates and ebridians in experimental Mohole
bier Mo Vee Sey eyer a yay Ue cere Re ae Sa a ae eh See a eae a ae emer 143
5. erminology, of the silicoflagellate skeleton) 22.222. 2...-esceccecceceeeeseceeeee-eeeeseee-s 147
Coe erminolocyaormtheeDridianm skeletons sess ee cene erence 196
7. Distribution of silicoflagellates from Neogene Mediterranean (Italy)
21 © Tipe ete seed met Ek Oe BN ANY Ee Seth ers ie) A oe Soon Ace eee eee 204
TABLE
NM PmPbIStMO te SAMIPLESMSEITU LES xn cce eer ce aoc cece see oan ccaceccncccece se eankes See oeoReet Se SoR cae oo 138
aTnaATHog
-T | '
A a
se) Le a pdr ging? ed pibea Jere A) eeady Beep re ie oe meen
ttt wo
atr :
+
SOF oni Se edna
ve] ; ~ Gielsieasteqarha 7
ae | pn at ae
ee ott
se
at Sin lay:
ogee OF GI01) Quads
rl
Tyg
ru axel
rit . beg sa va! Fetetig te aintemut O47 i Vag pil nivale of |
née TTT (eal ot ee Ovtl! ma test
Lake htleg et taailia lo
OH . a Se wie HTien
160 cream. Aghirer shies adihirl® La@ catellegnifvadl) Jom
Hndate (paws ies Ay tmeth eds fb “ix llooelhialae be galing
rt ae heey it,
TH a eorlath eepliogalTovliie ody te squalene
ee
: a ; ion, Cotghyte paitiety adi tee wate
Mamet), rerree eclibel# te ait rile wahelt ghana w
IE Se oS ieslt e : a NE
i a St Lat tae oe? a i
“ute ry eee
4 7
er! = — St tee © ating eeponcome == mies yar, oe
f WD
‘ 7 : !
' i eter,
es a a
od er) © ¢
; ; iad : 74 ) ——
7 i s ti ,
_ > ; = }
TUS ' i we + | mf i) wey a Y
UPPER CRETACEOUS AND CENOZOIC
SILICOFLAGELLATES AND EBRIDIANS*
Hsin Yr Linc
Department of Oceanography
University of Washington
ABSTRACT
This comprehensive examination of silicoflagellates and ebridians is based
on samples mainly from deep-sea sediments, including experimental Mohole
drilling and Legs I-III of the Deep Sea Drilling Project as well as land
outcrops, their ages ranging from the late Cretaceous (Maestrichtian) to Recent.
A detailed account of systematics is given and 56 taxa are discussed
with illustrations included from reference samples of type localities for these
forms. Stratigraphic occurrences are compared with those of other micro-
planktonic groups; planktonic Foraminifera, calcareous nannoplankton, and
Radiolaria.
Silicoflagellates and ebridians are grouped into seven assemblages and
two horizons are additionally recognized. Mediterranean samples are also
examined to evaluate the use of these microfloral assemblages in the middle
latitude region of Europe.
INTRODUCTION
The recent upsurge of micropaleontologic research has resulted
in the establishment of numerous biostratigraphic zones by means
of different microfossil groups from the late Mesozoic to Recent.
Coupled with the advancement of paleomagnetic stratigraphy and
the method of radiometric determination, these zones have been
combined (Berggren, 1969, table 2).
One group of siliceous microorganic remains, silicoflagellates
and ebridians, first described by Ehrenberg (1839), has not been
recognized for its potentiality by biostratigraphers. One of the main
reasons is that earlier investigators placed their emphasis on
describing new forms rather than on investigating their stratigraphic
occurrences.
Hanna (1928) and more recently Stradner (1961), Mandra
(1968), Bachmann and Papp (1968) suggested and attempted some
zonations using land outcrops from various parts of the world; and
Hays (et al., 1969) included one silicoflagellate species in their
study of the Plio-Pleistocene deep-sea sediments from the equatorial
Pacific which brackets the Jaramillo Event within the Pleistocene.
Jousé (1963, 1969, 1971), Muhina (1963, 1966, 1969), Ling (1970),
and Martini (1971b) subsequently recorded the importance of sili-
coflagellates in their studies of Pacific sediments, and the author
also found the same low-latitude species from mid-latitude central
north Pacific sediments.
The successful recovery of long cores by the Deep Sea Drilling
*Contribution No. 620, Department of Oceanography, University of Washington.
136 BULLETIN 273
Project (DSDP or JOIDES) provides a unique opportunity to
define the range of microfossils and to seek the phylogenic relation-
ship of various microfossils. This project also coordinates the efforts
of micropaleontologists in achieving accurate correlation. By com-
parison with the biostratigraphic framework thus refined by other
groups of planktonic microfossils, the value of silicoflagellates and
ebridians as a possible biostratigraphic index can be tested.
The present study is undertaken with the following objectives
in mind: (1) to define the range of taxa mainly from deep-sea sedi-
ments; (2) to decipher the relationship or evolutionary lineage, if
any, among silicoflagellates and ebridians; and (3) if possible, to
recognize assemblages based on the occurrence of these siliceous
microfossils.
ACKNOWLEDGMENTS
The author is indebted to William R. Riedel of Scripps Institu-
tion of Oceanography, University of California at San Diego, for
kindly providing the samples as well as the pertinent biostratigraphic
data upon which the present paper is based. Without his assistance,
the work could not have been accomplished at this time. Thanks
are also due to the Sample Distribution Panel of the Deep Sea
Drilling Project (DSDP), who kindly made samples available from
Leg I to Leg III, carried out under the financial support of the
National Science Foundation.
Sincere appreciation is due to Alfred Bachmann and Herbert
Stradner of Austria, N. de B. Hornibrook and A. R. Edwards of
New Zealand, York T. Mandra, Alfred R, Loeblich, Jr., and John
W. Ruth of the United States, for their graciously providing refer-
ence samples and unpublished data which are of great assistance in
clarifying the taxonomic problems; to Erlend Martini of West Ger-
many for his information on the unpublished data from DSDP Leg
VII; and to Thomas R. Worsley of the Department of Oceanography
at the University of Washington, for identifying some calcareous
nannofossil assemblages,
Initial phase of this study was assisted by Mrs. Carol J. Stadum,
and the completion by Mrs. Linda M. McPherson and Mrs. Shirley
A. Verzosa.
The study was supported financially by National Science
Foundation grants (GA-11308, GA-26499) and Office of Naval Re-
search contract Nonr 477 (37), Project NR 083 012.
SILICOFLAGELLATES AND EBRIDIANS: LING 137
SAMPLES STUDIED
The deep-sea sediments and land outcrop samples used for the
present study are listed in alphabetical order (Table 1) and their
geographic locations are shown in Text-figure 1.
For deep-sea samples from the Scripps Institution of Ocean-
ography collection, the abbreviations used by the Institution are
followed, and the position of the samples is the depth measured (in
centimeters) from the top of the core. For designating the Deep Sea
Drilling Project (DSDP) samples, the general format that appeared
in the Initial Reports of the Project is here adopted; namely, hole
number — core number — section number, followed by sample level
(in centimeters) from the top of the section. The samples from the
core sediments recovered during the experimental deep-sea drilling
phase of the Mohole Project at Guadalupe site are designated by
the abbreviation “EM” to conform with the original description
(Riedel, et al., 1961), followed by the number of the hole — the
number of the core run, and the depths of the sample, in centi-
meters, below the surface sediments of the particular core.
The stratigraphic positions of these samples are show in Text-
figures 2 and 3, except for some DSDP samples (marked with an
asterisk in Table 1) and a sample WR TR 39 H (Thyrsocyrtis
bromia Zone; Riedel, personal communication) which were barren
of these siliceous microfossils, to avoid an unnecessary overcrowding
in the figures. The scheme proposed by Berggren (1969, table 2)
relating radiometric time scale, European and West Coast (Cali-
fornia) marine stages, and the Cenozoic planktonic foraminiferal
zonation is adopted here, Calcareous nannoplankton zonations pro-
posed by Martini (1970) and Martini and Worsley (1970), and the
radiolarian zonation proposed by Riedel and Sanfilippo (1970,
1971) are included; the appropriate stratigraphic position of the
samples was then decided.
Despite these considerations, the sample position shown in Text-
figures 2 and 3 should still be regarded as merely an approximate
indication based on available knowledge. Inevitably a few difficulties
are encountered, According to Riedel and Sanfilippo (1970), sample
RIS 111P, 280-281 cm, belongs to Zone N.4 of planktonic Foramini-
fera but is regarded as the upper part of the Theocyrtis tuberosa
Zone of Radiolaria. The examination of calcareous nannoplankton
microflora of the sample by Thomas R. Worsley (personal com-
138 BULLETIN 273
TABLE 1. LIST OF SAMPLES STUDIED
Sample and Intervals (cm) Longitude Latitude Depth(m)
AMPH 6P 17-18 128°21’W 4°52’N 4460
50-51 128°21’W 4°52’N 4460
37P 250-251 121°05’W 18°16’S 3720
350-351 121°05’W 18°16’S 3720
CAS 1144 (California Academy
of Science Loc. 1144) 120°45’W 36°40’N
CHUB 15 78-82 125°25’W 8°31’N 4462
EM (Experimental Mohole 117°30’W 29°59’N 3566
drilling at Guadalupe site)
JYN V 16 P 231-233 149°44’W 7°44'N 5168
248-250 149°44’W 7°44’N 5168
DSDP (Deep Sea Drilling Project)
1P-1-6 40-41* 92°11’W 25°52’N 2827
2-1-3 115-116* 92°35’W 23°27'N 3572
2-2-1 130-131* 92°35’W 23°27’N 3572
2-3-1 45-46* 92°35’W 23°27'N 3572
2-3-2 100-101* 92°35’W 23°27'N 3572
2-4-2 15-16* 92°35’W 23°27'N 3572
3-5-3 62-63* 92°01’W 23°01’N 3737
3-7-1 50-51* 92°01’W 23°01’N 3737
3-10-2 45-46¢ 92°01’W 23°01’N 3737
4-4-] 10-11* 73°48’W 24°29’N 5319
5-1-1 33-34* 73°38’W 24°44’N 5361
5-1-2 73-74* 73°33’W 24°44’N 5361
6-4-1 6-8 67°39'W 30°50’N 5125
115-117 67°39'W 30°50’N 5125
6-4-2 7-10 67°39'W 30°50’N 5125
125-128 67°39'W 30°50’N 5125
6-4-3 92-94 67°39'W 30°50’N 5125
110-112 67°39'W 30°50’N 5125
6-5-1 33-35 67°39°'W 30°50’N 5125
6-6-2 1-2 67°39 W 30°50’N 5125
7-1-2 48-49* 68°18’W 30°08’N 5185
7A-3-2 51-52* 68°18’W 30°08’N 5185
9-5-1 8-9 59°12’W 32°46'N 4981
9-5-2 99-100 59°12’W 32°46’N 4981
9-5-4 117-118* 59°12’W 32°46’N 4981
10-9-2 60-61* 52°13’'W 32°52’N 4612
10-9-3 55-56* 52°13’W 32°52’N 4612
13-3-1 120-121 18°14’°W 06°02’N 4585
13A-1-1 115-116 18°14’W 06°02’N 4585
13A-2-1 90-91* 18°14’W 06°02’N 4585
13A-4-1 95-96* 18°14’W 06°02’N 4585
SILICOFLAGELLATES AND EBRIDIANS: LING 139
13A-5-1 40-41* 18°14’W 06°02’N 4585
LSDH 78P 100-101 168°02E’ 4°31’S 3208
516-517 168°02E’ 4231/5 3208
LSDA 101G 38-39 7°12 5 BoA's 2960
MP 5-1 25-30 133°07’"W 14°22’N 4700
MSN 126G 66-67 154°45'W 24°41'S 4542
MSN 149P 294-296 145°15’W 9°23’N 5100
PROA 88P 275-276 167°14’E 2°56’N 4428
RIS 111P 280-281 133°29'W 14°55’N 4770
WAH 7P 259-260 153°02’W 3°58’N 4992
516-517 153°02’W 3°58/N 4992
WR TR 11 Low (Bo. 202)
WR TR 23 (K9391)
WR TR 39F
WR TR 39F (J.S. 1068)
WR ER 39H*
Hermitage Quarry, south Trinidad, B.W.I. (Bolli,
1957, p. 101)
Near the Retrench trigonometrical station, Gol-
conda Estate, Trinidad, B.W.I. (Bolli, 1957, p.
101; Riedel and Sanfilippo, 1970, p. 510)
Lower Oligocene part of the Oceanic Formation
at Bath, Barbados
Oceanic Formation at Bath, Barbados
Uppermost Eocene part of the Oceanic Formation
at Bath, Barbados
*Samples examined but were barren of silicoflagellates and ebridians, there-
fore not listed in Text-figures 2 and 3.
>
MSN 149P
JYNZI6Pe °
. WAH TP
PROA 88P
is
*LSDH 7aP
°MSN I26G
RIS 11IP
MP 5-1
CHUB 15
“AMPH 6P
= NORTH
caste. AMERICA
WR TR 39F, “ap
© 39F (U.S. 1068) & 39H. =I
$=" WR TR | LOW(BO 202) awents
7" 23(K 9391) °
ciears DSOP 13 813A
AMPH 37P
.
Text-figure 1—Index map showing the geographic location of samples in-
vestigated.
140 BuLLETIN 273
CALCARE-
Ous
Pl NANNO- SAMPLES
WEST COAST PLANK -
EUROPEAN | (CALIFORNIA) TON RADIOLARIAN | (depth intervals
STAGES STAGES ZONES ZONES in cm)
[G friocontho.si triacantha
Dictyocha hexacantha
[Corbisemo recto recta
&. antiqua (without spine)
Septamesocena apiculota
Corbisema bimucronata
Corbisema apiculata
C. triecantha vor. minor
Naviculopsis biopiculata
C. archangelskiana
N. constricta
Ebriopis crenulata
L. furcula vor. minor
Corbisema hostata
L. simplex
Lyramula furcula
Vollacerta hortoni
CHATTIAN Dorcadospyris
(BORMIDIAN) ofeuchus
ZEMORIAN
LATTORFIAN
RUPELIAN
Theocyrtis MP 5-1 25-30
tuberoso WR TR 39F
WR TR 39F (JS. 1068)
—-—?--
REFUGIAN
BARTONIAN
DSDP
13-3-1, 120-121
a= 6-8
6-3-1. {ascii
F's 7-10
6-4-2, fio6 128
Podocyrtis
-94
LUTETIAN ULATISIAN chalara 6-4-3, {?3Sit2
Thyrsocyetis 6-5-1, 33-35
friocontha 6-6-2, I-2
PENUTIAN IZA-I-1, 115-126
PRIABONIAN
NARIZIAN
D>
a
te}
eta] ety gee fee | yee |e
EOCENIE
YPRESIAN BULITIAN
THANETIAN
YNEZIAN
VERY RARE
RARE
PALEOCENE
MONTIAN
COMMON
ABUNDANT
pANae “DANIAN”
CAS 1144 | il
MAESTRI-
CHTIAN
Text-figure 2.—Distribution of silicoflagellates and ebridians in Upper
Cretaceous and Paleogene section. The figure is compiled based on the data of
Berggren (1969), Martini (1970), and Riedel and Sanfilippo (1970, 1971).
munication) indicated that the sediment is Zone NN.1 of calcareous
nannoplankton, and thus is in agreement with that of Foraminifera.
The sample is, therefore, placed at the base of Miocene in Text-
figure 3 of this paper. The Oligocene-Miocene boundary is placed
by Berggren (1969, table 2) at the Globigerinoides Datum, 1.e. 22.5
m.y. and is between N.3 and N.4 Zones of planktonic Foraminifera
which falls within NN.1 Zone of calcareous nannoplankton (Martini
and Worsley, 1970, p. 289). Riedel (1971) placed this boundary
within N.4 planktonic Foraminifera and NN.1 calcareous nanno-
planktonic Zones, and between Calocycletta virginis above and
141
LING
SILICOFLAGELLATES AND EBRIDIANS
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142 BULLETIN 273
Lychnocanium bipes below of the radiolarian Zones. For convenience,
two samples, JYN V 16 P, 231-233 cm and MSN 149 P, 294-296 cm,
are also placed at the base of the Miocene in Text-figure 3.
In Text-figure 4, a combined sequence of the experimental
Mohole drilling samples is shown together with the biostratigraphic
subdivision of the sequence determined by means of calcareous nan-
noplankton by Martini and Bramlette (1963), planktonic Foramini-
fera by Parker (1964), Bandy and Ingle (1970), and diatoms by
Kanaya (1971).
In addition, samples from the following locations were examined,
and some specimens are illustrated in the present paper for com-
parison of the taxa:
Oamaru Diatomite: Near Oamaru, South Island, New Zealand.
A. R. Edwards kindly provided the samples from the following
three sections; all the sample numbers are those of the New
Zealand Geological Survey:
Bain’s Farm (North of Fault) section; S 136/963-970
Forrester’s Hill section; S 136/1096, 1195 and 994
Jackson’s Paddock section; S 136/1194, 914 and 878-882.
Kellogg Shale (KEL): West of Byron, Contra Costa County,
California. According to John W. Ruth of the Standard Oil Com-
pany of California, samples were collected by Alfred R. Loeblich,
Jr., of Chevron Oilfield Research Company from the location where
Mandra (1968, p. 272) sampled, and they were approximately at
4- or 5-foot intervals from the bottom (#1) to the top (#10) of
the present exposure. Thus the total thickness of outcrop is slightly
different from that which Mandra measured in 1948,
Sidney Shale (SID): South of Antioch, Contra Costa County,
California. The samples were collected by John W. Ruth from the
old quarry where Mandra (0. cit.) collected his samples. The
seven samples are: #1, at the base of the exposure; +2, 5 feet; #3,
60 feet; #3-A, 61 feet; 4¢4, 63 feet; #5, 65-73 feet (composite);
+6, 74-83 feet (composite); #7, approximately 88 feet from the
base (the top of the exposure).
Austrian samples: Alfred Bachmann kindly sent me the follow-
ing samples which he had examined for silicoflagellate assemblages;
they are:
SILICOFLAGELLATES AND EBRIDIANS: LING 143
in experi-
For samples with asterisk mark, see also Text-
Sadinuaj WhUDIY{010g
OU'4 2
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(OQNA//8 W=) ODNA//a YI DUEI0SAaY/
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and
Bramlette
AUS 1: “sample G, Walbersdorf im Burgenland (Jungeren
Badener Serie).”
AUS 2: “Limberg bei Maissau (Luschitzer Serie).”
AUS 3: “Ziegelei Ernstbrunn, Waschbergzone (Eggenburger
Serie, Burdigal).”
AUS 4: “Steiehang E Wallern a.d. Trattnach (Oberoligozan).”
A detailed description of these samples has already been presented
by Bachmann and Papp (1968).
144 BULLETIN 273
Finally a series of outcrop samples from Italy studied recently
by Sanfilippo (1971) were analyzed to evaluate silicoflagellate oc-
currences in the Mediterranean region (see discussion and Text-
figure 7).
LABORATORY PROCEDURES
A sample of sediment was trated routinely with hydrogen
peroxide (30%) to remove organic matter, The sediment was then
treated with concentrated HCl to dissolve any calcareous micro-
fossils that might mask or dilute the siliceous microfossils. The re-
maining residue was then decanted and sieved through a 74-p sieve.
Since silicoflagellates and ebridians are generally smaller than 74 p,
this provides an easy way to concentrate and separate them from
larger siliceous microfossils, such as Radiolaria and diatoms. Strewn
slides were made from this residue and dried. Canada balsam was
then used to adhere the cover glass of 22 x 40 mm size,
Counts were made on these strewn slides to determine the rela-
tive abundance of silicoflagellate and ebridian species. Measurements
for an individual taxon were generally made on the major axis on the
basis of at least 20 specimens.
Temporary or wet slides were also prepared by removing the
strewn slides from the oven before the mounting medium became
completely solid. By gently applying pressure in one direction on the
cover glass, specimens in the slide will rotate within the mounting
medium and thus permit an observation from various orientations.
The location for the illustrated specimens in the strewn slides
is indicated by the sample designation (as discussed earlier), fol-
lowed by the slide number, the location by the England finder read-
ing as previously described (Ling and Anikouchine, 1967).
All the slides examined during the present study will be de-
posited permanently in the micropaleontology collection, Depart-
ment of Oceanography, University of Washington, Seattle, Wash-
ington.
SYSTEMATIC MICROPALEONTOLOGY
In spite of an apparently simple configuration in comparison
with other microfossils, the systematics of silicoflagellates are not
so simple as one would expect. The enormous number of variety or
SILICOFLAGELLATES AND EBRIDIANS: LING 145
forma proposed by previous workers has created a large number of
names, and this fact has hindered the utilization of silicoflagellates
and ebridians as biostratigraphic indicators.
It is here considered that these microfossils belong to plant
origin (Loeblich and Tappan, 1968) and thus are discussed accord-
ing to the International Code of Botanical Nomenclature adopted in
Utrecht (Lanjouw, et al., 1966). In the following section, the classi-
fication scheme proposed by Frenguelli (1940), Deflandre (1950a),
Papenfuss (1955), Glezer (1966), and more recently by Bachmann
(1970b ), is generally followed. However, genera and species of each
genus are arranged in alphabetical order. Illustrations of Septa-
mesocena apiculata are placed immediately next to Mesocena for
the convenience of comparison,
Inasmuch as many of the previously reported taxa are involved
during the present study, it has become necessary to review them
critically from the biostratigraphic point of view. Consideration has
been given to these taxa which had been recorded by the workers
as variety and forma or forma, and recognized here whenever they
seem to have either stratigraphic importance or possible paleoeco-
logical implication. In addition, to facilitating better understanding
of previous authors’ observations, specimens from the reference
materials of well-known localities are presented for comparison. A
detailed analysis of microfossils from these localities is currently
being undertaken by the author, and the results will be reported
in later papers as the work progresses.
SILICOFLAGELLATES
Perhaps because of the simple configuration, the terminology
of the siliceous skeletal elements of silicoflagellates has not changed
much since it was first introduced by Gemeinhardt (1930). The
short description prepared here and also Text-figure 5 uniformize
and introduce some new terms used in the later discussion.
Terminology — The typical skeleton of a silicoflagellate consists
of a polygonal basal body ring with radial spines at each corner ex-
tending generally horizontally. From the basal side of the polygon,
basal accessory spines project down- and inwardly. In some species
(such as Dictyocha fibula var. messanensis), there are short acces-
sory spines at the exterior side of the basal body ring.
146 BuLLeTIN 273
The structure within or above the basal body ring is referred to
as an apical structure, which may be in the form of a ring, apical
ring; with an opening or openings; apical window or several apical
windows; or a simple straight bar, an apical bar; or a flattened thin
plate, apical plate. Occasionally short spine(s), apical accessory
spine(s), are present on an apical ring or on an apical bar. The apical
structure and the basal body ring are connected by J/ateral rods.
The spaces delineated by the apical structure, the basal body
ring and also by lateral rods are called lateral window(s) (“Basal-
fenster” of Gemeinhardt, 1930, p. 21) or basal window of Glezer
(1966, p. 9).
Orientation — The shape of the basal body ring and the posi-
tion of the apical structure serve to orient the silicoflagellate speci-
mens, Except for circular or triangular forms, generally the silico-
flagellate skeleton is longer in one direction; this is called the major
axis, and the shorter one, the minor axis. When a silicoflagellate is
orientated in such a way that the basal body ring, radial spines and
lateral rods are on a plane and with the apical structure form a
shape of truncated pyramid, it is called lateral view (edge view of
Loeblich, III, et al., 1968; Ling, 1970); the view directly facing the
apical structure is called an apical view and the opposite is termed
as abapical view (Marshall, 1934; Loeblich, III, et al., 1968, p. 7).
Order SIPHONOTESTALES Lemmermann, 1901a
Genus CANNOPILUS Haeckel, 1887, emend. Bachmann, 1967
Remarks.— As discussed earlier Ling (1970), Loeblich, III,
et al. (1968) and more recently Lipps (1970) considered that
Haeckel’s name should be regarded as a junior synonym of Hali-
calyptra by Ehrenberg (1847), on the basis of samples from the
Barbados Island and Richmond, Virginia. Furthermore, Lipps (1970,
pp. 975-976) suggested that another form, Clathropyxidella described
by Deflandre (1938) also from Barbados, is probably synonymous.
Three of the samples studied are from Barbados; special attention
was made during the microscopic examination to find forms similar
to those previously described, but the search was unsuccessful. Un-
doubtedly further investigation is needed to determine the exact
SILICOFLAGELLATES AND EBRIDIANS: LING 147
APICAL VIEW
LATERAL VIEW
— basal accessory spine — _ ;
a pppical bar
_-——basal body ring——W—
~~
ABAPICAL APICAL
vIEW > —— view
=
™~ .
~— —lateral window — ——
=
~— — — lateral rod —- ———
lateral /
window
major axis
apical accessor
7 spine x41
_7lateral window
f
7 i] ;
\iateral rod ~ —apical plate ~
o- P , if
‘apical window
\
Text-figure 5.—Terminology of the silicoflagellate skeleton (Modified after
Deflandre, 1950a)
nature of these forms and to resolve this taxonomic problem, and in
the present paper the long-recognized Haeckel’s name is applied.
Cannopilus hemisphaericus (Ehrenberg) Pl. 23, figs. 1-5
Remarks. — As discussed earlier (Ling, 1970), the synonymy
of this species has been presented by various investigators; how-
ever, despite their painstaking efforts, the present species has not
been satisfactorily defined. This is apparently due to the fact that
some of the specimens assignable to the species demonstrate inter-
mediate characteristics by possessing a basal body ring like Diste-
phanus speculum; and yet the complex nature of the apical struc-
ture, hemispherical outline and varying numbers of apical windows
148 BULLETIN 273
are common in the genus Cannopilus. Thus Glezer (1966) and Bach-
mann (1970a, b) make a special effort to differentiate between
those with the diameter of the apical structure equal to or less than
that of the basal body ring as Distephanus speculum var. canopil-
oides, and those with the larger diameter as Cannopilus hemi-
sphaericus.
As has been noted from the central north Pacific sediments
(Ling, op. cit.), it is not necessary to have a larger diameter of the
apical structure in order to have a hemispheric outline. It is also
apparent that the number of apical windows cannot be the valid
taxonomic criteria as Ehrenberg or Lemmermann once proposed;
thus it is somewhat puzzling that Glezer (1966, p. 280) again con-
sidered an apical structure with six or more windows as one of the
criteria for Cannopilus; and the taxonomic names of Ehrenberg or
Lemmermann, based on the number of apical windows, appeared
recently (McManus, et al/., 1970, particularly for the description of
Hole 32-34) which had already been well discussed as early as 1964
by Bachmann.
Perhaps the above discussion can be elucidated by Bachmann
(1970b, p. 10) at the end of his emended description of Distephanus
speculum var. cannopiloides which states “Abgrenzung zum nahe
verwandten Cannopilus hemisphaericus vorgenommen werden.”
Therefore, following the previous central north Pacific study,
the specimens showing hemispheric outline of the apical structure in
the lateral view and generally possessing two or more apical windows
and larger than ordinary related forms, such as Distephanus specu-
lum, are all included under the present species. The only exceptions
to the present taxon are those specimens with two small windows,
the size of one window divided by an apical bar, and with the apical
structure definitely not hemispherical but parallel to the basal body
ring. These such as Dictyocha btpartita Ehrenberg (1844a; 1854),
are found in Recent surface sediments from various parts of the
world, including the Bering Sea and Antarctic waters.
Dimensions. — Length of basal body ring, 28-40 p; of radial
spine, 14-26 pu.
Known geologic range. — Miocene.
Occurrence. —C. hemisphaericus is found between the interval
from WR TR 11, Low (Bo. 202), Globorotalia fohsi barisanensis
SILICOFLAGELLATES AND EBRIDIANS: LING 149
Zone or Dorcadospyris alata Zone (Riedel and Sanfilippo, 1970) in
Text-figure 3, and from EM 7-2, 6-7 cm, Luisian (Parker, 1964)
or Helvetian ? (Martini and Bramlette, 1963) in Text-figure 4,
to EM 8-11, 69-70 cm, N.12 planktonic foraminiferal Zone or Om-
matartus antepenultimus-Cannartus (?) pettersson Zone (Riedel
and Sanfilippo, of. cit.). It is interesting to note here that Mandra
(1968) recorded the occurrence of the present species from Relizian
and Mohnian stage samples. A sample EM 7-2, 7-10 cm, which 1s
identified as Relizian by Bandy and Ingle (1970, p. 140, fig. 3; p.
167), did not yield the present species.
Cannopilus picassoi Stradner Pl. 23, figs. 6, 7
Cannopilus picassoi Stradner, 1961, Erdél u. Kohle, vol. 14, No. 2, p. 92,
figs. 101-104.
Cannopilus picassoi Stradner, Bachmann, 1963, in Bachmann et al., Geol.
Ges. Wien, Mitt., vol. 56, No. 1, p. 153, pl. 20, figs. 74-81; pl. 22, figs.
17-19.
Cannopilus picassoi Stradner, Bachmann and Papp, 1968, Giorn. di Geol.,
(2) iceviolei3i5saple 3,etie~ 16.
Cannopilus picassoi Stradner, Hajos, 1968, Geol. Hungarica, ser. Paleont.,
fascs 37 ps 70M pls 6, figs 5.
Cannopilus picassoi Stradner, Bachmann, 1970, im Cat. Fossil, Austriae, No.
1b, p. 14.
Remarks. — When Stradner (1961) originally described the
present species from the Miocene (“Unteres Torton’’) of Frattings-
dorf, Austria, it seems that two morphologically different forms
were included: one with well-developed apical accessory and radial
spines (his figs. 101-102), and the other with both spines poorly
developed, particularly the radial spines (his figs. 103, 104). The
Italian specimens are the only forms found during the present study
which show only the latter type; however, the general form, includ-
ing numerous apical windows and the overall size, agrees with the
specimens from Austria,
Dimensions. — Diameter of apical structure, 30-40 p.
Known geologic range. — Miocene.
Occurrence. — This species is found rarely and only from Italian
samples.
Cannopilus sphaericus Gemeinhardt Pl. 23, figs. 8-10
Cannopilus sphaericus Gemeinhardt, 1931a, Deutsch, Bot. Ges., Ber., vol. 49,
No. 2, p. 104, pl. 10, figs. 3, 4.
Cannopilus sphaericus Gemeinhardt, Frenguelli, 1940, Rev. Mus. La Plata,
n.s., vol. 2, Paleont., No. 7, pp. 48-51, fig. 6e. .
150 BULLETIN 273
Cannopilus sphaericus Gemeinhardt, Deflandre, 1950a, Microscopie, vol. 2,
fies. 95.096, 9702).
Cannopilus sphaericus Gemeinhardt, Stradner, 1961, Erdél. u. Kohle, vol.
14, No. 2, p. 92, figs. 99, 100.
Cannopilus sphaericus Gemeinhardt, Bachmann, 1963, in Bachmann, et al.,
Geol. Ges. Wien, Mitt., vol. 56, No. 1, p. 153, pl. 20, figs. 69-73; pl. 22,
fig. 16.
Cannopilus sphaericus Gemeinhardt, Glezer, 1966, in Cryptogamic plants of
thesUcS'SeRe vol-7pa coils plz. ties 5:
Cannopilus sphaericus Gemeinhardt, Bachmann and Papp, 1968, Giorn. di
Geol., ser. 2, vol. 35, fasc. 2, p. 122.
Cannopilus sphaericus Gemeinhardt, Hajos, 1968, Geol. Hungarica, ser.
Paleont., fasc. 37, p. 70, pl. 6, figs. 1-4, 6.
Cannopilus sphaericus Gemeinhardt, Mandra, 1968, California Acad. Sci.,
Proc., ser. 4, vol. 36, No. 9, pp. 247-248, figs. 66(?), 70.
Remarks. — The present species is characterized by the presence
of larger and consequently fewer numbers of apical windows than
C. picassoi or C. tetraceros, and by the presence of five or six down-
and outwardly directed basal radial spines. Lateral windows are
frequently Jarger in size than the apical windows. Apical accessory
spines at the top and the sides of the spherical apical structure are
either present or absent.
Dimensions. — Diameter of apical structure, 28-40 up.
Known geologic range. — Miocene.
Occurrence. — The specimens assigned to the present species
are found only from experimental Mohole samples during the
present study, and from EM 7-2, 6-7 cm to EM 8-14, 388-389 cm
interval. Thus the top seems to coincide with the boundary of
Helvetian? — Tortonian? in middle Miocene of calcareous nanno-
fossils by Martini and Bramlette (1963); and approximately
Luisian-Mohnian of planktonic Foraminifera by Parker (1964) and
Bandy and Ingle (1970). On the basis of studies mainly from
the land sections of California, the present species is considered
as one of the indices for the Mohnian stage by Mandra (1968),
but recently Ruth (1971) recorded that it ranges from middle
Luisian to almost the top of middle Mchnian.
Cannopilus tetraceres Bachmann and Ichikawa Pl. 23, figsyadee
Cannopilus tetraceros Deflandre, 1949, nomen nudum.
Cannopilus tetraceros Deflandre ex Bachmann and Ichikawa, 1962, Kana-
zawa Univ., Sci. Rept., vol. 8, No. 1, p. 172, pl. 5, figs. 73-75; pl. 9,
Higssoulole
Remarks. — When Bachmann and Ichikawa described the pres-
SILICOFLAGELLATES AND EBRIDIANS: LING 151
ent species in their investigation from the Wakura Beds (1962, p.
172), of Miocene (late) (Fuji and Bachmann, 1969), they indicated
that Deflandre found a similar specimen from Moron, Spain, and,
therefore, they credited the authorship to Deflandre and dated it as
1949. Until now no description or figure has been presented by De-
flandre; therefore the date of 1949 should be considered as nomen
nudum and as invalid. The species should be Cannoptlus tetraceros
Deflandre ex Bachmann and Ichikawa or its abbreviated form as
above and dated as 1962 (Lanjouw, et al., 1966, ICBN Art, 46;
Recommendation 46C).
Dimensions. — Diameter of apical structure, 42 p.
Known geologic range. — Miocene.
Occurrence. — Throughout the present study, the species is
found in sample EM 8-15, 246-249 cm only.
Genus CORBISEMA Hanna, 1928, emend. Frenguelli, 1940
Remarks. — The genus was first proposed by Hanna (1928);
and the emended diagnosis given by Frenguelli (1940, p. 69) which
was later accepted by Deflandre (1950a, p. 47/82), is followed
here. Glezer (1966, p. 223; pp. 252-253) restricted the present genus
to those forms in which the basal accessory spines are “on the
lateral rods at the corners of apical widening,” and thus Glezer con-
sidered only C. geometrica Hanna as belonging to the present genus,
indicating that specimens which have basal accessory spines on the
basal body ring are regarded as belonging to the genus Dictyocha.
Corbisema apiculata (Lemmerman) PISZS shigss ea
Dictyocha triacantha var. apiculata Lemmerman, 1901b, Deutsch. Bot. Ges.,
Ber., vol. 19, p. 259, pl. 10, figs. 19-20.
Dictyocha triacantha var. apiculata Lemmermann, Schulz, 1928, Bot. Archiv.,
vol. 21, No. 2, pp. 247-249, fig. 27.
Dictyocha triacantha var. apiculata fa. aspera Schulz, 1928, Bot. Archiv.,
: vol. 21, No. 2, pp. 247-249, fig. 28.
Dictyocha triacantha var. apiculata fa. late-radiata Schulz, 1928, Bot.
_ Archiv., vol. 21, No. 2, p. 281, fig. 73.
Dictyocha triacantha var. apiculata Lemmermann, Gemeinhardt, 1930, in
Kryptogamen-Flora, vol. 10, pt. 2, pp. 41-42, fig. 30.
Dictyocha triacantha var. apiculata fa. aspera Schulz, Gemeinhardt, 1930,
in Kryptogamen-Flora, vol. 10, pt. 2, pp. 41-42.
Dictyocha triacantha var. apiculata fa. late-radiata Schulz, Gemeinhardt,
1930, in Kryptogamen-Flora, vol. 10, pt. 2, p. 43, fig. 32.
Corbisema apiculata (Lemmermann), Hanna, 1931, Mining in California,
VolaZ7a No.2. ps 198 ples ties 2:
152 BULLETIN 273
Dictyocha triacantha var. apiculata Lemmermann, Deflandres, 1932a, Soc.
France, Microsc., Bull., vol. 1, No. 1, fig. 32.
Corbisema apiculata (Lemmermann), Frenguelli, 1940, Rev. Mus., La Plata,
n.s., vol. 2, Paleont., No. 7, fig. 12h.
Corbisema apiculata (Lemmermann), Stradner, 1961, Erdé] u. Kohle, vol.
14, No. 2, p. 89, figs. 26, 27, 33.
Dictyocha triacantha var. apiculata fa. apiculata Glezer, 1966, in Crypto-
gamic plants of the U.S.S.R., vol. 7, pp. 228-229, pl. 6, fig. 5.
Dictyocha triacantha var. apiculata fa. late-radiata Schulz, Glezer, 1966,
in Cryptogamic plants of the U.S.S.R., pp. 229-230, pl. 6, fig. 1.
Corbisema apiculata (Lemmermann), Mandra, 1968, California Acad. Sci.,
Proc., ser. 4, vol. 36, No. 9, p. 248, figs. 25, 30(?), 35.
Remarks. — The specimens referable to the present species show
a triangular basal body ring with short radial spines at three rounded
apices. Basal accessory spines generally present, short or occasionally
strong and distinct.
Dimensions. — Length of basal body ring, 30-60 up.
Known geologic ranges. — Cretaceous (late) to Eocene (late).
Occurrence. — Throughout the present study, the species is
found in samples from DSDP Hole 6, and in reference samples from
Bain’s Farm section of Oamaru Diatomite, New Zealand, as well as
from Kellogg and Sidney Shale of California.
Corbisema archangelskiana (Schulz) Pl. 23; figaids
Dictyocha triacantha var. archangelskiana Schulz, 1928, Bot. Archiv., vol.
Ze Now 2h pp. 250-25 tig 33) a-caups Zo leehiess 7750781 (er)
Dictyocha triacantha var. archangelskiana Schulz, Gemeinhardt, 1930, in
Kryptogamen-Flora, vol. 10, pt. 2, pp. 45-46, fig. 37a, b(?).
Corbisema archangelskiana (Schulz), Frenguelli, 1940, Rev. Mus., La Plata,
n.s., vol. 2, Paleont., No. 7, fig. 12a.
Corbisema archangeskiana (Schulz), Stradner, 1961, Erdél u. Kohle, vol. 14,
No. 2, p. 89, fig. 25 (only).
Dictyocha triachantha var. archangelskiana Schulz, Tsumura, 1963, Yoko-
hama Munic. Univ., Jour., ser. C-45, No. 146, pp. 48-49, pl. 9, fig. 1; pl.
22, fig. 14.
Dictyocha archangelskiana (Schulz), Glezer, 1966, in Cryptogamic plants
of the UlS-.SiRs -vol.07s (pp.'232-233) spl. 8, figs» 6, 7:
Remarks. — Glezer (1966) correctly assigned figure 6 from
Oamaru, New Zealand, and figure 7 from the eastern slope of the
Urals to the present species, but erred by referring to Gemeinhardt’s
figure 34, which is Dictyocha triacantha var. inermis fa. late-
radiata Schulz (= Corbisema geometrica Hanna, in this paper).
The species is characterized by three elongated triangular lateral
windows, sides of which are subparallel and with narrow but not
pointed apices.
SILICOFLAGELLATES AND EBRIDIANS: LING 153
Dimensions. — Length of basal body ring, 50 up.
Known geologic range. — Cretaceous (late) to Eocene (late).
Occurrence. — Found rarely and only from DSDP sample, 6-4-3,
110-112 cm.
Corbisema bimucronata Deflandre Pl. 24, fig. 1
Corbisema bimucronata Deflandre, 1950a, Microscopie, vol. 2, pp. 63/82-
64/82, figs. 174-177.
Dictyocha bimucronata (Deflandre), Tsumura, 1963, Yokohama Munic.
Univ., Jour., ser. C-45, No. 146, p. 50, pl. 9, fig. 6.
Dictyocha bimucronata (Deflandre), Glezer, 1966, in Cryptogamic plants of
the. W-S.SiR. vol. 7. ps. 253,.pi. 9x tise 2:
Remarks. — This small species possesses sharply truncated
apices with short spines at both edges. Basal accessory spines are
short but generally present. Glezer (1966, p. 233) indicated that
the early Oligocene (?) form from the Ukrainian area, the only
occurrence from Russia, did not possess the basal accessory spines
and she is of the opinion that “arrangement of the supporting spines
(= basal accessory spines in this paper) cannot serve as the major
criterion in defining the genus Corbisema as does Deflandre.”
Dimensions. — Length of the basal body ring, 38 pn.
Known geologic range. — Eocene to Oligocene (?).
Occurrence. — Until now the species has been reported, except
in Russia, only from “Radiolarite 4 Diatomées de Springfield et de
Newcastle, ile de la Barbade”, and was regarded as “?Miocene in-
férieur” but is now considered as Eocene by Loeblich, III, et al.
(1968, p. 18). The specimens found during the present study were
also from Barbados samples, WR TR 39 F (J.S. 1068) and 39 F,
which are both from the Theocyrtis tuberosa Zone, and the strati-
graphic position for the former is also identified as slightly above
the [sthmolithus recurvus Zone (Riedel and Sanfilippo, 1970). Ap-
parently the present species has a limited geologic, as well as geo-
graphic distribution, because no specimen was found from other
areas, including Oamaru, New Zealand, and California.
Corbisema geometrica Hanna Pl. 24, figs. 2-4
Dictyocha triacantha var. inermis Lemmerman, 1901b, Deutsch. Bot. Ges.,
Ber vole 195 p. 259 pple, tics 2c
Corbisema geometrica Hanna, 1928, Jour. Paleont., vol. 1, No. 4, p. 261, pl.
_ 41, figs. 1, 2.
Dictyocha triacantha var. inermis Lemmerman, Schulz, 1928, Bot. Archiv..,
vol. 21, No. 2, p. 249, fig. 30a, b; p. 281, fig. 75.
154 BULLETIN 273
Dictyocha triacantha var. inermis fa. late-radiata Schulz, 1928, Bot. Archiv.,
vol. 21, No. 2, p. 281, fig. 74.
Dictyocha triacantha var. inermis Lemmermann, Gemeinhardt, 1930, iz
Kryptogamen-Flora, vol. 10, pt. 2, p. 43, fig. 33.
Dictyocha triacantha var. inermis fa. late-radiata Schulz, Gemeinhardt,
1930, in Kryptogamen- -Flora, vol. 10, pt. 2, p. 43, fig. 34.
Dictyocha triacantha var. inermis Lemmerman, "Deflandre, 1932a, Soc. France
Microsc., Bull., vol. 1, No. 1, fig. 31.
Corbisema geometrica Hanna, Deflandre, 1940a, Acad. Sci. Paris, C) R.,
vol. 211, No. 19, p. 446, figs. 3-4, 6-8.
Corbisema gceometrica Hanna, Deflandre, 1950a, Microscopie, vol. 2, pp.
53-82—54-82, figs. 134, 136-139.
Corbisema archangelskiana Stradner, 1961 (not Schulz or Deflandre),
Erdol u. Kohle, vol. 14, No. 2, p. 89, figs. 19, 24 (only).
Dictyocha geometrica (Hanna), Tsumura, 1963, Yokohama Munic. Univ.,
Jour., ser. C-45, No. 146, pl. 9, figs. 4, 5; pl. 22, figs. 15-16.
Corbisema geometrica Hanna, Glezer, 1966, in Cryptogamic plants of the
WES:S: REV vollv/ps 2 535ipleag whiz. 7.
Dictyocha triacantha var. inermis fa. inermis Glezer, 1966, in Cryptogamic
plants of the W:S:S:R2 vol) 7 p. 230; pl. 8, figs; 1, 2- ply $2. tizaels
Corbisema geometrica Hanna, Mandra, 1968, California Acad. Sci., Proc.,
ser. 4, vol. 36, No. 9, p. 248, figs. 3-5.
Remarks.— The present species was described originally from
Cretaceous materials of California. Careful examination of more
than ten slides, including both larger and smaller than 74 » fractions
made from topotypic material, CAS 1144, yielded only a few speci-
mens and with apical plate only (PI. 24, fig. 2). Deflandre (1940a)
illustrated that the position of basal accessory spines is variable,
ranging from on the lateral rods near the hyaline apical plate to on
the basal body ring. Glezer (1966), on the other hand, was of the
opinion that the present species should be for those with basal ac-
cessory spines on the lateral rods only, and she also reported basal
accessory spines up to 10 » long. Consequently Glezer considered the
present species to be the only form belonging to the present genus,
and she limited the range for the genus as Cretaceous (late), while
for those Dictyocha triacantha var. inermis fa. inermts, early Paleo-
cene is additionally mentioned.
Hanna included a specimen without hyaline apical plate as a
deviated form within the present species. Apparently the inclusion
of such a latter form led Deflandre to combine those described pre-
viously by Lemmermann (1901b) as Dictyocha triacantha var.
inermis from “Fuur in Jutland” under the present species. Unfor-
tunately no such specimen was observed from the topotypic ma-
terial. There is a possibility that future detailed study of Cretaceous
SILICOFLAGELLATES AND EBRIDIANS: LING 155
materials from California might yield such a specimen as Mandra
(1968) illustrated; therefore, Hanna’s concept is followed here.
Although Deflandre (1950a, p. 54/82) indicated that Dictyocha
triacantha var. archangelskiana Schulz (1928, pp. 250-251, fig. 33a-c)
may also belong to the present species, and Stradner (1961) ap-
parently followed such an opinion, this is not accepted in the present
study.
Dimensions. — Length of basal body ring, 70-90 up.
Known geologic range. — Cretaceous (late) to Eocene.
Occurrence. —C. geometrica as here recognized is found in
Cretaceous California and middle Eocene DSDP materials.
Corbisema hastata (Lemmermann) Pl. 24, fig. 5
Dictyocha triacantha var. hastata Lemmermann, 1901b, Deutsch. Bot. Ges.,
Ber., vol. 19, p. 259, pl. 10, figs. 16, 17.
Dictyocha triacantha var. hastata Lemmermann, Schulz, 1928, Bot. Archiv.,
vol. 21, No. 2, pp. 249-250, figs. 31a-c; 29b (only).
Dictyocha triacantha var. hastata Lemmermann, Gemeinhardt, 1930, in
Kryptogamen-Flora, vol. 10, pt. 2, p. 43, fig. 35a-c.
Dictyocha triacantha var. hastata Lemmermann, Deflandre, 1932a, Soc.
France, Microsc., Bull., vol. 1, No. 1, fig. 28.
Dictyocha triacantha var. hastata Lemmermann, Frenguelli, 1940, Rev. Mus.,
La Plata, n.s., vol. 2, Paleont., No. 7, fig. 12c (only).
Dictyocha triacantha var. hastata Lemmermann, Glezer, 1966, in Crypto-
gamic plants of the U.S.S.R., vol. 7, pp. 231-232, pl. 6, figs. 6-8; pl. 7,
fies, 8 pla sl, fiesyis) 9%
Dimensions. — Length of the basal body ring, 24-30 uy.
Known geologic range. — Paleocene (early) to Eocene (late).
Numerous locations are listed for the occurrence of the present
species from Russia by Glezer (1966).
Occurrence. — Found from the samples of DSDP 6-6-2, 1-2 cm,
Thyrsocyrtis triacantha Zone, to DSDP 13-3-1, 120-121 cm, Podo-
cyrtis chalara Zone (Riedel, 1971).
Corbisema recta (Schulz) Pl. 24, figs. 6, 7
Dictyocha triacantha var. recta Schulz, 1928, Bot. Archiv., vol. 21, No. 3,
pi250.aties052a.0 Db:
Dictyocha triacantha var. recta Schulz, Gemeinhardt, 1930, in Kryptogamen-
Flora, vol. 10, pt. 2, p. 45, fig. 36.
Phyllodictyocha recta (Schulz), Deflandre, 1946, Soc. Bot. France, Bull.,
vol. 93, No. 9, pp. 335-337, fig. 1.
Phyllodictyocha recta (Schulz), Deflandre, 1950a, Microscopie, vol. 2,
p. 45/82, figs. 112, 113.
Dictyocha recta (Schulz), Tsumura, 1963, Yokohama Munic. Univ., Jour.,
ser. C-45, No. 146, p. 49, pl. 9, fig. 2; pl. 22, fig. 8.
156 BULLETIN 273
Description. — Basal body ring, triangular either smooth or
slightly convex; radial spines short at each corner; short basal ac-
cessory spines at the abapical side of the basal body ring directly
below where the lateral rods join with basal body ring; apical struc-
ture simple, consisting of three equal lateral rods meeting at the
center and in some specimens, slightly expanding to form a small
apical plate.
Remarks. — Deflandre’s new genus, Phyllodictyocha, with
Schulz’s variety as the type species, is characterized by having a
flattened and non-tubular basal body (1935, 1946), and Tsumura
(1963, p. 49) stated that no lumen (or hollow structure) can be
found within the basal body. The specimens found from the DSDP
samples failed to show any particular flattened nature of the basal
body ring and the lumen was present in contrast to illustrations by
Deflandre and Tsumura; furthermore the basal accessory spines
seem heavier than those shown by them.
Dimensions. — Length of basal body ring, 30-38 p.
Known stratigraphic range. — Eocene.
Occurrence. — The present taxon is found from samples DSDP
13-3-1, 120-121 cm, Thyrsocyrtis triacantha Zone and Barbados,
WR TR 39 F (J. S. 1068), Theocyrtis tuberosa Zone and is just
above the top of Isthmolithus recurvus Zone (= NP.19 Zone of
Martini, 1970) (Riedel and Sanfilippo, 1970).
Corbisema triacantha (Ehrenberg) Pl. 24, figs. 8-13
Dictyocha triacantha Ehrenberg, 1844a, Verh. K. Preuss. Akad. Wiss. Ber-
lin, Ber., Jahrg. 1844, p. 80.
Not Dictyocha triommata Ehrenberg, 1845, Verh. K. Preuss. Akad. Wiss.
Berlin, Ber., Jahrg. 1845, pp. 56, 76.
Not Dictyocha triommata Ehrenberg, Ehrenberg, 1854, Mikrogeologie, pl.
33, IN, 1S, saree all
Dictyocha trigona Zittel, 1876, Deutsch. Geol. Ges., Zeitschr., vol. 28,
ple 83.) plar2 ties. 6 Gae
Dictyocha trigona Zittel, Riist, 1888, Palaentographica, vol. 34, No. 5/6.
Joy AISI
Dictyocha triacantha Ehrenberg, Lemmermann, 1901b, Deutsch. Bot. Ges.,
Ber, vole 195 pa258plad0s figs S:
Dictyocha triacantha Ehrenberg, Schulz, 1928, Bot. Archiv., vol. 21, No. 2,
p. 247, figs. 24, 25a.
Dictyocha triacantha Ehrenberg, Gemeinhardt, 1930, in Kryptogamen-Flora,
vol. 10, p. 40,, tigss28a (2). bi ((2)eres 29aG)e
Corbisema triacantha (Ehrenberg), Hanna, 1931, Mining in California, vol.
Ai, je ID) aayers ile
Corbisema triacantha (Ehrenberg), Zanon, 1934, Acta Pont. Acad. Sci. Nov.
Lincei, vol. 87, p. 66, pl. n, fig. 8.
SILICOFLAGELLATES AND EBRIDIANS: LING SyvA
Corbisema triacantha (Ehrenberg), Frenguelli, 1940, Rev. Mus., La Plata,
n.s., vol. 2, Paleont., No. 7, fig. 12g.
Corbisema hastata Frenguelli, 1940 (not Lemmermann), Rev. Mus., La Plata,
n.s., vol. 2, Paleont., No. 7, fig. 12b (only).
Corbisema trigona Zittel, Deflandre, 1950a, Microscopie, vol. 2, pp. 52/82-
53/82, fig. 130.
Dictyocha triacantha fa. minor Desikachary and Maheshwari, 1956 (not
Schulz), Ind. Bot. Soc., Jour., vol. 35, No. 3, p. 258, text-fig. 2.
Dictyocha triacantha var. triacantha fa. triacantha Glezer, 1966, in Crypto-
gamic plants of the U.S.S.R., vol. 7, pp. 226-227, pl. 4, figs. 1-6; pl. 5,
figs. 1-4.
Corbisema triacantha (Ehrenberg), Mandra, 1968, Calif. Acad. Sci., Proc.,
ser. 4, vol. 36, No. 9, p. 249, figs. 28, 31.
Remarks. — The basal body ring of this species is triangular in
shape and is on a plane, in comparison with the following taxon; and
the apical plate, if present, is definitely rarely developed. Because
of considerable variation in size and shape, as well as in length of
radial spines, some smaller specimens with less well-developed radial
spines cannot satisfactorily be differentiated from those of Dictyocha
triacantha var. minor.
A good example of such a case is a specimen illustrated by
Schulz (his fig. 25 a) as D. t. fa. minor. It should be noted that
Schulz’s figure 24 for the present species illustrates that the lumen
of the basal body ring is clear at the three corners as well as in the
radial spines but is lost completely at the middle of the basal ring.
Gemeinhardt’s specimen (fig. 28 a) from Mors Island even shows
the septal structure at the apices. Ehrenberg’s D. triommata is
considered synonymous by Gemeinhardt and Glezer but is considered
as D. triacantha var. minor in this paper. Gemeinhardt also in-
cluded another of Ehrenberg’s species, D. trifenestra, with the
present taxon, but judging from the original figure, it may be re-
lated to D. fibula; therefore, D. trifenestra is excluded from the
above synonymy list.
Dimensions. — Length of basal body ring, 18-35 p.
Known geologic range. — Cretaceous to Miocene (late).
Occurrence. — The present species is recovered from middle
Eocene DSDP sediments and Italian samples. From the experimen-
tal Mohole section, the species is recovered from the Luisian inter-
val only (Parker, 1964; Bandy and Ingle, 1970).
Corbisema triacantha var. flexuosa Stradner Pl. 24, figs. 14-17
Corbisema triacantha vor. flexuosa Stradner, 1961, Erdél u. Kohle, vol. 14,
No. 2, p. 89, pl. 1, figs. 1-8.
158 BuLLETIN 273
Dictyocha triacantha var. flexuosa (Stradner), Glezer, 1966, in Cryptogamic
plants of the U.S.S.R., vol. 7, p. 228, pl. 5, figs. 5 (?), 10 (only).
Corbisema triacantha var. flexuosa fa. I1 Bachmann, 1970b, im Cat. Fossil.
Austriae, No. 1b, p. 16.
Remarks.— One of the samples that Alfred Bachmann pro-
vided the writer is from Wallern, Austria, the type locality for the
present variety and typical of such specimens is here presented with
both apical and lateral views (Pl. 24, figs. 14, 15). During the
present study, this taxon was found only from Italian land samples.
Glezer (1966) recorded the occurrences in Russia as late Eocene
to late Oligocene, but only figure 10 from Mangyshlak Peninsula of
late Oligocene age seems to agree with the original concept of
Stradner; while figure 5, an early Oligocene Ukrainian specimen has
a dentate basal body ring, but such is never observed by Stradner
nor so far by the present author from Wallern samples. Figures 6
and 8 of Glezer (op. cit.) are from the Ural-Caspian area and are
of late Eocene age, but the former shows a straight triangular basal
body ring with strongly crenulate (?) surface ornamentation, which
is unusual for these forms. Nevertheless it is better considered as
C’. triacantha var. triacantha, while the latter has short radial spines
as well as a smoothly concaved basal body ring, and may be re-
garded as C. t. var. minor. Therefore, some of the above-discussed
questionable specimens of Glezer have been excluded from the
present taxon.
Dimensions. — Diameter of basal body ring, 20-30 p.
Known geologic range. — Possible Oligocene (late) only.
Occurrence. — Found the present taxon only from Italian out-
crop samples between sample WRE 67-99 and -95 which is identified
as Calocycletta virginis Zone (Sanfilippo, 1971).
Corbisema triacantha var. minor (Schulz) Pl. 24, figs. 18-23
Dictyocha triacantha fa. minor Schulz, 1928, Bot. Archiv., vol. 21, No. 2,
p. 247, fig. 25b (only).
Dictyocha triacantha fa. minor Schulz, Gemeinhardt, 1930, in Kryptogamen-
Flora, vol. 10, p. 41 (not fig. 29a or b).
Not Dictyocha triacantha fa. minor Schulz, Desikachary and Maheshwari,
1956, Ind. Bot. Soc., Jour.. vol. 35, No. 3, p. 258, text-fig. 2.
Corbisema trigona (Zittel), Tynan, 1957, Micropaleont., vol. 3, No. Zapp:
130-131, pl. 1, fig. 1 (only).
Corbisema triacantha fa. minor (Schulz), Bachmann, 1963, in Bachmann,
et al., Geol. Ges. Wien, Mitt., vol. 56, No. 1, p. 151, pl. 17, figs. 28-32
(not fig. 33).
SILICOFLAGELLATES AND EBRIDIANS: LING 159
Dictyocha crux Ehrenberg, Bachmann, 1964 (part), in Ichikawa, ef al.,
Kanazawa Univ., Sci. Rept., vol. 9, No. 1, p. 103, pl. 4, figs. 38-39
(only) (as trigonale Abwandlungen). | '
Dictyocha triacantha var. triacantha fa. minor Schulz, Glezer, 1966, in
Cryptogamic plants of the U.S.S.R., vol. 7, pp. 227-228, pl. 5, figs 7-9.
Dictyocha crux fa. trigona Frenguelli, Bachmann, 1967, im Ichikawa et al,
Kanazawa Univ., Sci. Rept., vol. 7, No. 1, p. 157, pl. 4, figs. 1-6, 8
(only).
Remarks. — The above synonymy list is presented as a first
approach to prepare this taxon for future possible stratigraphic use.
The specimens here considered are smaller than other varieties of
the species, generally with shorter radial spines, and furthermore
they generally have convex sides with rounded apices showing a
roundly triangular basal body ring. Apical structure consists of
lateral rods forming a Y, or occasionally with a small apical plate.
Basal accessory spines are short, if present. Size range and some
larger specimens admittedly approach to its type C. triacantha, and
in some cases the separation is difficult.
Thus Schulz’s figure 25a is considered in the present paper as
C. triacantha, and similar consideration is applied to some of Bach-
mann’s illustrations from Austria and Japan. Schulz’s figure 26b
from Mors Island clearly possesses an apical accessory spine and
so far no such specimen has been observed; the specimen from Mors
Island is excluded from the present taxon.
Dimensions. — Length of basal body ring, 16-25 yp.
Known geological range. — Eocene (early) to Miocene.
Occurrence. — The latest occurrence for the present taxon in
the experimental Mohole section is the same as C. triacantha, and
Cannopilus sphaericus; however in the Mediterranean region, it is
found slightly higher than that of C. triacantha.
Genus DICTYOCHA Ehrenberg, 1839
Dictyocha ausonia Deflandre PL. 25, figs. 1-10
Dictyocha pons Ehrenberg, 1844a, Verh. K. Preuss. Akad. Wiss. Berlin,
Ber., Jahrg. 1844, pp. 64 (in table, name only), 80.
Dictyocha pons Ehrenberg, 1854, Mikrogeologie, pl. 21, fig. 40.
Dictyocha navicula var. pons (Ehrenberg), Lemmermann, 1901b, Deutsch.
Bot. Ges., Ber., vol. 19, p. 258.
Not Dictyocha ausonia Deflandre, Glezer, 1966, in Cryptogamic plants of
the U.S.S.R., vol. 7, p. 252, pl. 9, fig. 1.
Dictyocha ausonia Deflandre, Hajos, 1968, Geol. Hungarica, ser. Paleont.,
fase. 37, pp. 65-66, pl. 3, fig. 11 (only).
For others, see Ling (1970, pp. 88-90) as D. cf. ausonia Deflandre.
160 BULLETIN 273
Remarks. — The present species, Deflandre (1950a) originally
described from Miocene Diatomite of Italy, and at the same time
presented a series of variations. On the basis of central north Pacific
deep-sea sediments, Ling (1970) discussed in detail the present
taxon, as D. cf. awsonia, and expressed some hesitation in accepting
completely Deflandre’s species concept mainly because no complete
range of intraspecific variation was observed at that time, The
present study now reveals that there is a wide range of intra-
specific variation as Deflandre claimed, but radial spines on the
minor axis do not disappear as readily as Deflandre illustrated.
Actually, if radial spines disappear completely, it is perhaps after
the gradual disappearance of two lateral windows on the minor
axis (see Pl. 25, figs. 5-8). A middle Miocene species, Dictyocha
pons, proposed and illustrated by Ehrenberg from Oran, Algeria,
which is similiar to Plate 25, figure 7 here, is thus considered as a
part of variation for the present species. Judging from the illustra-
tion, the size also seems to agree with such a conclusion.
On the other hand, D. regularis Carnevale (1908) is not con-
sidered as conspecific with the present species, as Deflandre (1950a)
indicated and Ling (1970) suggested as a possibility, because of the
definitely much longer radial spines on the major axis; the species
is transferred to genus Naviculopsis and discussed later.
Glezer (1966) recorded the only Russian occurrence and refer-
red to Deflandre’s taxon; but judging from the illustration, as well
as the remarks that Sakhalin specimens possess longer radial spines
on the major axis, and such is not the case for the present species,
the Russian specimens are excluded from the present taxon.
Dimensions. — Length of the basal body ring, 30-50 y; of radial
spines, 6-14 p.
Known geologic range. — Miocene.
Occurrence. — Apparently the present species ranges from WAH
7P, 259-260 cm, Calocycletta virginis Zone and N.5 planktonic
foraminiferal Zone (Riedel and Sanfilippo, 1970) to EM 6-2, 30-32
cm, which is the top of Delmontian and Mohnian or Mohnian
(Parker 1964) and the boundary between Mohnian and Delmontian
stages (Bandy and Ingle, 1970).
Dictyocha fibula Ehrenberg Pl. 25, figs. 11-15
Remarks. — As in the previous investigation of central north
SILICOFLAGELLATES AND EBRIDIANS: LING 161
Pacific sediments, this species encompasses the wide variation and
forma previously reported by various workers from all over the
world (see Mandra 1968; Glezer, 1966).
Dimensions. — Length of basal body ring, 30-45 yw; of radial
spine, 5-16 pu.
Known geologic range. — Eocene to Recent.
Occurrence. — Found from WR TR 11, Low (Bo. 202),
Dorcadospyris alata Zone (Riedel and Sanfilippo, 1970), to modern
sediments in many parts of the world.
Dictyocha fibula var. aculeata Lemmermann Pike259 figse Gn
Remarks. — The synonymy list, discussions and dimensions
were presented previously by the present writer (Ling, 1970).
Known geologic range. — Quaternary.
Occurrence. — This variety is recovered from DSDP 9-5-2, 99-
100 cm, N.22 planktonic foraminiferal Zone (Blow, 1970, pp. 357,
360) to Recent.
Dictyocha fibula var. messanensis (Haeckel) Pl. 25, figs. 18, 19
Remarks.— The detailed taxonomic consideration was pre-
sented by the author (Ling, 1970).
Known geologic range. — Quaternary.
Occurrence. — Like D. f. var. acwleata, this variety is recovered
from DSDP 9-5-2, 99-100 cm to Recent.
Dictyocha fibula var. octagona Tsumura PIN2Z6 figs: 152
Not Dictyocha octonaria Ehrenberg, 1844b, Verh. K. Preuss. Akad. Wiss.
Berlin, Ber., Jahrg. 1844, p. 201.
Not Dictyocha octonaria Ehrenberg, Deflandre, 1950a (part), Microscopie,
vol. 2, fig. 37 (only).
Dictyocha fibula var. Tsumura, 1959, Yokohama Munic. Uniy., Bull., vol.
11, Nat. Sci., No. 1, p. 66, pl. 4, figs. 47-49 (fide Tsumura, 1963).
Dictyocha fibula var. octagona Tsumura, 1963, Yokohama Munic. Univ.,
Jour., ser. C-45, No. 146, p. 55, pl. 10, figs. 11-13; pl. 23, figs. 8-10
(as D. octagona on p. 55).
Remarks. — The illustrated specimens agree with the species
proposed by Tsumura from Tsubaki, Sado Island, Niigata Prefecture
of Japan.
Tsumura questionably correlated his present species with
Ehrenberg’s D. octonaria, but in the original description, Ehrenberg
indicated its close relationship with another form, D. ornamentum,
162 BULLETIN 273
which in turn is related to Distephanus speculum. Tsumura also
correlated his specimen with one (fig. 37) of Deflandre’s Dictyocha
octonaria, but judging from the illustration, Deflandre’s specimen
possesses small apical accessory spines on an apical bar, lateral rods,
and basal accessory spines, and no such specimen has been found
during the present study; therefore, both Ehrenberg’s and De-
flandre’s species are excluded from the present vartety.
Dimensions. — Length of basal body ring, 20-30 pw; of radial
spines, 10-14 p.
Known geologic range.— Listed only as fossil, according to
Tsumura.
Occurrence. — This variety is found from AMPH 6P, 50-51 cm,
Dorcadospyris alata Zone and N.11 planktonic foraminiferal Zone
(Riedel and Sanfilippo, 1970), to EM 7-1, 14-15 cm, N.12 planktonic
foraminiferal Zone, according to Riedel and Sanfilippo (o. cit.).
Martini (1971b) indicated his Dictyocha octagona horizon from the
Deep Sea Drilling Project Leg VII in the equatorial Pacific region,
but did not list it in another article (1971a).
Dictyocha hexacantha Schulz Pl, 263fig73
Dictyocha hexacantha Schulz, 1928, Bot. Archiv., vol. 21, No. 2, p. 255,
fig. 43.
Dictyocha hexacantha Schulz, Gemeinhardt, 1930, in Kryptogamen-Flora,
vol. 10, pt. 2, pp. 55-56, fig. 46.
Dictyocha hexacantha Schulz, Hanna, 1931, Mining in California, vol. 27,
| No. 2; p. 9198; pl. D; fig: 7:
Dictyocha deflandrei fa. hexacantha, Frenguelli, 1940, Rev. Mus. La Plata,
n.s., vol. 2, Paleont., No. 7, p. 65, fig. 14g (only).
Corbisema hexacantha (Schulz), Deflandre, 1950a, Microscopie, vol. 2, pp.
65/82-66/82, figs. 183-187.
Dictyocha hexacantha Schultz, Glezer, 1966, in Cryptogamic plants of the
U.S.S.R., vol. 7, p. 239, pl. 10, fig. 12.
Description. — Basal body ring, triangular with convex side;
three radial spines, short, at the apices; apical structure simple,
consisting of three lateral rods that meet near the center and ex-
tend the distal ends beyond the basal body ring, thus showing six
radial spines in apical or abapical view; basal and apical accessory
spines absent,
Remarks. — Schulz (1928) described and illustrated the present
species from Oamaru, and a specimen here presented (Pl. 26, fig.
3) is from sample S136/964, Forrester’s Hill section of Oamaru
Diatomite, New Zealand.
‘al
SILICOFLAGELLATES AND EBRIDIANS: LING 163
Results of the preliminary observation so far made, including
some Eocene samples from California and New Zealand, reveal that
the shape of the present species is better considered basically as
triangular rather than hexagonal. The so-called six radial spines
from which the name of the present species was apparently derived
actually are three radial spines at the three apices and three exten-
sions of lateral rods of apical structure that intersect approximately
at the middle of the basal body ring. Thus, it may not be surprising
that the apical structure and such distal extension are similar to
“propeller-like structure” that Mandra (1969, lower-right figure
on p. 173) observed from Antarctic deep-sea samples.
Dimensions. — Length of basal body ring, 20-26 u.
Known geologic range. — Eocene to Oligocene (early ?). In
addition to Schulz’s finding from Oamaru Diatomite, Hanna (1931)
reported the present species from Kreyenhagen Shale (Eocene) of
California. Although Mandra (1968) failed to find the present
species during his study, this author found the present species from
Sidney Shale during preliminary examination. Deflandre (1950a)
added the Springfield, Barbados occurrence, which is now regarded
as Eocene age (Loeblich, III, et al., 1968, p. 18), and not Miocene
(early ?) as indicated by Deflandre (op. cit.) or Glezer (op. cit., p.
239). The “early Oligocene (?)” occurrence is reported by Glezer
from Ukrainian, Russian.
Occurrence. — This species is recovered only from DSDP 6-4-1,
6-8 cm, upper part of the Globigerapsis kugleri Zone (Ewing, et al.,
1969) and the Thyrsocyrtis triacantha Zone (Riedel, 1971), The
species is observed in the reference samples from Oamaru Diatomite,
New Zealand, and Sidney Shale, California.
Dictyocha sp. Pl. 26, figs. 4-8
Dictyocha fibula var. fibula fa. rhombica Glezer, 1966 (not Schulz, Lemmer-
mann nor Ling), im Cryptogamic plants of the U.S.S.R., vol. 7, p. 249,
DieelSs figss 5,07) (in) =) 9s onlys
Description. — Basal body ring, rhombic with almost straight
sides; radial spines present, two on the major axis always longer,
sometimes longer than those on the minor axis; apical bar along
the minor axis like D. ausonia; four lateral windows of approxi-
mately similar size or two on the minor axis slightly smaller but
so far never disappearing as D. ausonia. Basal accessory spines,
164 BuLLETIN 273
either present on abapical side of junction of lateral rods and basal
body ring, or slightly to the side of the junction, or sometimes
absent.
Remarks. — Because of the nature of radial spines, particularly
those on the major axis, the shape of skeleton is extremely variable,
but the basal body ring is definitely rhomboid.
A similar silicoflagellate was discussed as D. fibula var. rhom-
bica by Schulz (1928, fig. 37) and Ling (1970), and D. fibula var.
rhombus by Lemmermann (1901b), but none of them are related to
the forms illustrated here. Glezer (1966) recorded a few Russian
specimens that seem to be conspecific, but it is believed that Glezer
erred by referring to Schulz’s or Lemmermann’s species. Further-
more Glezer’s figure 4, which is from Mondaino (?), Italy, seems
to belong to D. ausonia; therefore, comparison with Russian ma-
terial will be definitely necessary before the new name can be
established for the present species.
Another Russian Eocene form, D. fibula var. fibula fa. eocaenica
Krotov, according to Glezer (0. cit., p. 249, pl. 15, fig. 6; pl. 32,
fig. 5), is synonymous with D. fibula fa. rhombica Schulz sens
Gemeinhardt, fig. 40a (only) but has dentate surface and is larger
than the present taxon.
Dimensions. — Length of basal body ring, 26-40 y; of radial
spine 6-20 np.
Occurrence. — This species is found in samples ranging from
WR TR 11, Low (Bo. 202), in many Mohole samples to LSDH 78
P, 516-517 cm, Spongaster pentas Zone (Riedel and Sanfilippo,
1970).
Genus DISTEPHANUS Stohr, 1880
Distephanus crux (Ehrenberg) Pl. 26, figs. 9-16
Remarks. — As in previous studies (Ling, 1970, 1971), wide
intra- and infraspecific variation is recognized under the present
species, except those varieties recognized below,
Dimensions. — Length of basal body ring, 24-36»; of radial
spines, 8-24 p.
Known geologic range. — Eocene to Miocene (late). Pliocene
(?), Quaternary (?). Late Miocene to Pliocene occurrence recorded
by Glezer (1966, p. 262) from Noto Peninsula and Kanzawa area,
SILICOFLAGELLATES AND EBRIDIANS: LING 165
Honshu Island, Japan, is more likely to be Miocene (late) only.
Quaternary occurrences reported by Glezer (op. cit.) from inter-
moraine layer, Ladoga area, Raukhiala, Russia, and interglacial de-
posits from Finland need to be reexamined.
Occurrence. — This species is found during the present study
from sample WR TR 23, (K9391), Calocycletta costata Zone
(Riedel and Sanfilippo, 1970), to EM 6-2, 10-11 cm which is con-
sidered as Delmontian by Bandy and Ingle (1970) or upper Mio-
cene by Martini and Bramlette (1963). Mandra (1968) reported
the Californian occurrence for the present species from Relizian,
Mohnian and Delmontian stages.
Distephanus crux var. longispina Schulz Pl. 26, figs. 17-19
Distephanus crux fa. longispina Schulz, 1928, Bot. Archiv., vol. 21, No. 2,
p. 256, fig. 44.
Distephanus crux var. longispina Schulz, Stradner, 1961, Erdél u. Kohle,
vol. 14, No. 2, p. 92, fig. 56 (only).
Dictyocha crux fa. longispina (Schulz), Bachmann and Ichikawa, 1962,
Kanazawa Univ., Sci. Rept., vol. 8, No. 1, p. 169, pl. 2, figs. 16, 17 (?),
18, 19 (only).
Dictyocha crux fa. longispina (Schulz), Bachmann, 1963, in Bachmann et al.,
Geol. Ges. Wien, Mitt., vol. 56, No. 1, p. 148, pl. 16, fig. 16.
Dictyocha crux Ehrenberg, Hajos, 1968 (part), Geol. Hungarica, ser.
Paleont., fasc. 37, p. 65, pl. 3, figs. 8, 9 (only).
Dictyocha crux var. longispina (Schulz), Bachmann, 1970a, Verh. Geol.
B.-A., Jahrg. 1970, No. 2, p. 287, pl. 4, figs. 15-17; pl. 5, figs. 1-9.
Distephanus crux var. longispina Schulz, Bachmann, 1970b, in Cat. Fossil.
Austriae, No. Ib, p. 9.
Remarks. — The present variety is characterized, as Bachmann
described (1970a, p. 287), by the relatively long radial spines along
the major axis, relatively large apical ring, and above all, by a highly
arched apical structure which is in turn supported by high-angled
lateral rods (see Pl. 26, fig, 17).
The sudden and common appearance of this variety in the ex-
perimental Mohole section is significant. With the consideration that
Schulz’s original figure for his forma came from Santa Monica,
California, which is considered as Helvetian stage by Loeblich, IIT,
et al. (1968), the specimens from Mohole sediments are assumed
to be identical with that of Schulz.
Dimensions. — Length of basal body ring, 28-36 y; of radial
spines, 16-28 up.
Known geologic range. — Miocene,
Occurrence. — Throughout the present study, the specimens re-
166 BULLETIN 273
ferred to the present taxon are found only from experimental Mohole
samples, from EM 8-15, 505-506 cm, to EM 8-11, 69-70 cm, the latter
of which is recorded as Ommatartus antepenultimus - Cannartus (?)
petterssoni Zone and N.15 planktonic foraminiferal Zone (Riedel
and Sanfilippo, 1970).
Distephanus crux var. octacanthus Desikachary and Maheshwari
Pl. 26, figs. 20-22
Distephanus crux var. octacanthus Desikachary and Maheshwari, 1956, Ind.
Bot. Soc., Jour., vol. 35, No. 3, pp. 260-261, text-figs. 10, 12, 13; pl. 13,
figs. 8, 9.
Remarks. — As discussed by Desikachary and Maheshwari from
upper Miocene samples of Nicobar Island, the present taxon is char-
acterized by an octagonal basal body ring; and two radial spines
on the major axis are much longer than the other six. The specimens
found during the present study agree with the above original descrip-
tion and illustrations. Apparently the size and shape demonstrated
during the present study show limited infraspecific variation.
Glezer (1966, p. 263) included the present variety within a part
of Distephanus stauracanthus (Ehrenberg), but they are different
morphologically and should be regarded as separate taxa (compare
with Ling, 1970, pl. 19, figs. 7, 8).
Dimensions. — Length of basal body ring, 24-30 »; of radial
spines, 8-16 uw
Known geologic range. — Miocene.
Occurrence. — Martini (197la, b) noticed the limited occur-
rence of the present taxon in the DSDP Leg VII samples and
placed his Dictyocha octacantha horizon near the top of his Cor-
bisema triacantha Zone. This horizon, according to him, is time
equivalent with the middle of NN.6 calcareous nannoplankton, ap-
proximately at the bottom of N.12 planktonic foraminiferal and
middle part of Dorcadospyris alata radiolarian Zones. Throughout
the present study this variety has occurred in the experimental
Mohole samples, interval between EM 7-2, 6-7 cm and EM 7-1, 14-
15 cm.
Distephanus speculum (Ehrenberg) PI. 26, figs. 23, 24; Pl. 27, figs. 1, 2
Remarks. — As pointed out in previous studies from the central
north Pacific deep-sea sediments and Shinzan diatomaceous member,
SILICOFLAGELLATES AND EBRIDIANS: LING 167
Akita Prefecture, Japan (Ling, 1970, 1971), the synonymy for the
present species has been presented by numerous workers on this
subject.
Dimensions. — Length of basal body ring, 18-40; of radial
spines, 6-30 yp.
Known geologic range. — Eocene to Recent.
Occurrence. — D. speculum is found from sample WAH 7P, 516-
517 cm, N.4 foraminiferal and Calocycletta virgins - Lychnocamum
bipes Zones (Riedel and Sanfilippo, 1970), throughout many of the
Mohole and Mediterranean samples to LSDA 101G, 38-39 cm sam-
ple, which is N.17 or 18 foraminiferal and Spongaster pentas radio-
larian zones (Riedel and Sanfilippo, 1970). Occurrence of the present
species is recognized in various parts of the world by many investi-
gators.
Distephanus speculum var. pentagonus Lemmermann | 5) DY a 00S
Distephanus asteroides Haeckel, 1887, Rept. Voy. Challenger, Zool., vol. 18,
p. 1564.
Distephanus speculum var. pentagonus Lemmermann, 1901b, Deutsch. Bot.
Ges., Ber., vol. 19, p. 264, pl. 11, fig. 19.
Distephanus speculum var. pentagonus Lemmermann, Schulz, 1928, Bot.
Archiv., vol. 21, No. 2, pp. 263-264, fig. 57.
Distephanus speculum var. pentagonus Lemmermann, Gemeinhardt, 1930,
in Kryptogamen-Flora, vol. 10, pt. 2, pp. 65-67.
Dictyocha speculum Ehrenberg, Stradner, 1961 (part), Erdél u. Kohle, vol.
14, No. 2, p. 92, figs. 72, 73 (only).
Dictyocha speculum fa. pentagona (Lemmermann), Bachman, 1963, iz Bach-
mann et al., Geol. Ges. Wien, Mitt., vol. 56, No. 1, p. 150, pl. 21, fig. 3.
Distephanus speculum var. pentagonus Lemmermann, Glezer, 1966, in Cryp-
togamic plants of the U.S.S.R., vol. 7, pp. 267-268, pl. 21, figs. 1-5.
Distephanus speculum var. pentagonus Lemmermann, Bachmann, 1970b, in
Cat. Fossil. Austriae, No. Ib, p. 10.
Remarks. —In contrast to the previous work from the central
north Pacific deep-sea sediments (Ling, 1970), an attempt was
made during the present study to examine the biostratigraphic
distribution of the present variety by recognizing it as an inde-
pendent taxon. At least in the experimental Mohole samples, the
present variety has a limited stratigraphic occurrence (see below).
Haeckel (1887) proposed the new species from Challenger sta-
tion 272 but did not give any illustration. As the sample contains
numerous Tertiary Radiolaria (Riedel, 1957), and judging from the
size, it seems that the species named by Haeckel may be considered
synonymous and, therefore, it is included here. Apparently Glezer
168 BULLETIN 273
regarded the present variety in a broader sense by including Lem-
mermann’s Distephanus speculum var. pentagonus fa. armata
(1901b, pp. 264-265, pl. 11, fig. 20) which possesses apical accessory
spines on the apical ring. So far no such specimen has been recovered
in the present study; hence the forma of Lemmermann’s is not
included, Glezer (1966) also synonymized a part of Hanna’s
Distephanus variabilis (1931, pl. E, fig. 8 only), but the size for
the latter is much bigger and the basal body is much thicker than
for the present species. Eocene samples from California need to be
studied in the future, and therefore, Hanna’s species name is ex-
cluded from the present taxon at this time.
Dimensions. — Length of basal body ring, 20-26 y; of radial
spine, 8-16 pu. .
Known geologic range. — Eocene to Neogene, Quaternary (?).
Occurrence for the variety reported by Glezer (1966) from the inter-
moraine layer of Ladoga area needs to be reexamined.
Occurrence. — During the present study, specimens identified as
the present taxon are found only in samples from experimental
Mohole drilling material from EM 8-12, 94-95 cm to EM 8-11, 322-
323 cm. Thus from the analysis of the Mohole samples, they are
recovered from the middle (Parker, 1964) or lower (Bandy and
Ingle, 1970) Mohnian stage. Whether such limited stratigraphic
occurrence is a local phenomenon needs to be examined in the future
in other areas. Mandra (1968) observed the California land occur-
rence only from Delmontian samples.
Distephanus speculum var. pseudocrux Schulz Pay tics 5-0
Distephanus speculum fa. pseudocrux Schulz, 1928, Bot. Archiv., vol. 21,
No. 2, p. 263, fig. 52a, b.
Distephanus speculum fa. pseudocrux Schulz, Gemeinhardt, 1930, in Krypto-
gamen-Flora, vol. 10, pt. 2, p. 64, fig. 55.
Distephanus speculum fa. pseudocrux Schulz, Tsumura, 1963, Yokohama
Munic. Univ., Jour., ser. C-45, No. 146, p. 63, pl. 11, figs. 2, 3.
Distephanus japonicus fa. pseudofibula Glezer, 1966 (part), in Cryptogamic
plants of the U.S.S.R., vol. 7, p. 276, pl. 25, figs. 1, 3, 5 (?) (only).
Distephanus speculum (Ehrenberg), Mandra, 1968 (part), Calif. Acad. Sci.,
Proc., ser. 4, vol. 36, No. 9, p. 262, fig. 76 (only).
Remarks. — The figure (see Pl. 27, fig, 5) illustrated here
agrees with the form originally described by Schulz. Similar apical
structure can be found also in specimens with a pentagonal basal
body ring but rarely, while another figure (see Pl. 27, fig. 6) is
SILICOFLAGELLATES AND EBRIDIANS: LING 169
considered as an example of morphological variation for the present
variety.
Glezer (1966) combined the present and the following taxon
as Distephanus japonicus (Deflandre), which possesses the charac-
teristic apical accessory spine(s) according to original species diag-
nosis but no such specimen was encountered during the present
study. Furthermore as can be seen from the distribution chart
(Text-figure 4), the two varieties are not always found together,
such as in the eastern Bering Sea margin samples (see below);
therefore, they are considered as separate taxa in the present study.
Dimensions. — Length of basal body ring, 24-30 yp; of radial
spine, 8-12 un.
Known geologic range. — Neogene, most likely only Miocene.
Occurrence. — The present taxon is found only from the Mohole
samples interval between EM 8-10, 78-79 cm, and EM 8-9, 148-149
cm. The age of this part is identified as upper Mohnian by Bandy
and Ingle (1970), Delmontian and Mohnian or Mohnian by Parker
(1964), and upper Miocene by Martini and Bramlette (1963).
Distephanus speculum var. pseudofibula Schulz Pl. 27, figs. 8-13
Distephanus speculum fa. pseudofibula Schulz, 1928, Bot. Archiv., vol. 21,
No. 2, pp. 262-263, fig. 51a, b.
Distephanus speculum fa. pseudofibula Schulz, Gemeinhardt, 1930, in
Kryptogamen-Flora, vol. 10, pt. 2, p. 64, fig. 54.
Distephanus speculum fa. pseudofibula Schulz, Deflandre, 1932a, Soc.
France, Microsc., Bull., vol. 1, No. 1, fig. 46.
Dictyocha pseudofibula (Schulz), Tsumura, 1963, Yokohama Munic. Univ.,
Jour., ser. C-45, No. 146, pp. 55-56, pl. 11, figs. 1-3; pl. 24, fig. 2.
Dictyocha pseudofibula var. complexa Tsumura, 1963, Yokohama Munic.
Univ., Jour., ser. C-45, No. 146, pp. 56-57, pl. 11, fig. 4; pl. 24, fig. 1.
Distephanus japonicus fa. pseudofibula Schulz, Glezer, 1966 (part), in
Cryptogamic plants of the U.S.S.R., vol. 7, p. 276, pl. 24, figs. 4, 6-8, 10.
Remarks. — As indicated under the preceding taxon, the com-
bination by Glezer (op. cit.) seems undesirable as well as mislead-
ing; therefore, Glezer’s usage is not followed in this study.
This variety is described originally by Schulz from San Pedro
and Redondo, California, Abashiri of Japan, and Ngemegy of Hun-
gary, and all are fossil forms, Tsumura found his var. complexa
from a Modern sea sample near Japan, and it is the only Quaternary
record.
Recently Hopkins (et al., 1968) reported the variety (as Dicty-
170 BULLETIN 273
ocha pseudofibula) from the eastern margin of the continental
slope of the Bering Sea. By courtesy of David W. Scholl of the U.S.
Geological Survey at Menlo Park, the TT-2 and -4 samples (TT18-
02 and TT18-04 respectively) were examined, and they are illus-
trated here (PI. 27, figs. 11-13) for comparison.
Dimensions. — Length of basal body ring, 26-32 yp; of radial
spine, 7-14 un.
Known geologic range. — Miocene to Quaternary (?).
Occurrence. — Throughout the present study, the present var-
iety 1s found only from experimental Mohole samples, and the
interval between samples of EM 8-10, 78-79 cm, and EM 6-2, 10-11
cm, upper Mohnian to Delmontian. Martini (1971la, b) recognized
his Dictyocha pseudofibula horizon from the DSDP Leg VII, which
is correlated with NN.10 calcareous nannoplankton, N.16 fora-
miniferal and lower part of Ommatartus antepenultimus radio-
larian Zones. Judging from the stratigraphic position, it is possible
that occurrence within the Mohole section is time-equivalent with
that of the above western equatorial Pacific. The Bering Sea dredge
haul samples are, therefore, regarded as of late Miocene age rather
than late middle Miocene as previously assumed (op. ctt., p. 1475).
Genus LYRAMULA Hanna, 1928
Remarks. — Since the genus and two species were first described
by Hanna (1928), the exact nature of the forms has remained
uncertain. Thus, Gemeinhardt (1930) and Frenguelli (1940) ex-
cluded the genus, and Lipps (1970) regarded them as questionable
silicoflagellates and suggested their possible affinities as “the setae
of several species of Chaetoceros-like diatoms, from which the frus-
tule has been dissolved.” This may be possible because of selective
dissolution, the delicate frustules of Chaetoceros are rarely found in
bottom sediments despite the fact that they are abundant in plank-
ton both in numbers of species and individuals (Kanaya and
Koizumi, 1966). However Chaetoceros frustule with setae, but not
the setae alone, have been reported from deep-sea sediments and
Tertiary deposits by several diatom researchers; such as Koizumi
(1968), Kozlova (1964), Jousé (1962), Sheshukova-Poretzkaya
(1967), and the author has also found such specimens from surface
sediments of various parts of the world oceans. Therefore, the forms
SILICOFLAGELLATES AND EBRIDIANS: LING 171
illustrated here from Cretaceous materials under the present genus
Lyramula are too distinctive to be considered as a part of diatoms
at present,
At the same time, microfossils similar to the Lyramula species
have been described by Tsumura (1963) as L. tenwpertica Kokubo
et Tsumura (pp. 30-31, pl. 3, fig. 3; pl. 16, fig. 5) from Japan and
L. tenwipertica var. laevis Tsumura (p. 31, pl. 3, fig. 5; pl. 16, fig.
3) from California. Recently Hopkins (et al., 1969, p. 1477) re-
corded the occurrence of L. tenuipertica from the dredge haul of
TT18-3 (as TT-3) from the continental margin of the Bering Sea.
Numerous specimens were observed similar to Tsumura’s
species during the present study, and an example is shown here.
One specimen is from a sample, DSDP 9-5-2, 99-100 cm, identified
as belonging to the N.22 planktonic foraminiferal Zone (Blow,
1970), and possessing the surface structure of “roughly an alter-
nate scalariform” (Tsumura, 1963, p. 30). Under careful examina-
tion, the specimen does not show the hollow structure of the rods
like the Lyramula species from Upper Cretaceous, and such differ-
ence is apparent under phase contrast microscopy (compare PI.
27, figs. 14, 15 vs. 16, 17). The affinity of Lyramula tenuipertica
remains uncertain, and therefore, is excluded from the present paper.
Lyramula furcula Hanna Pl. 27, figs. 16-18
Lyramula furcula Hanna, 1928, Jour. Paleont., vol. 1, No. 4, p. 262, pl. 41,
figs. 4, 5.
Lyramula furcula Hanna, Deflandre, 1940b, Acad. Sci. Paris, C. R., vol.
211, No. 21, p. 509, figs. 1-4.
Lyramula furcula Hanna, Deflandre, 1950a, Microscopie, vol. 2, p. 61/82,
figs. 163, 165, 167, 169.
Lyramula furcula Hanna, Tsumura, 1959, Yokohama Munic. Univ., Bull.,
vol. 11, Nat. Sci., vol. 1, p. 43, pl. 1, figs. 1-2 (fide Tsumura, 1963).
Lyramula furcula Hanna, Tsumura, 1963, Yokohama Munic. Univ., Jour.,
ser. C-44, No. 146, pp. 29-30, pl. 1, figs. 1; pl. 3, figs. 1-2; pl. 16, figs. 1-2.
Lyramula furcula Hanna, Mandra, 1968, California Acad. Sci., Proc., ser. 4,
vol. 36, No. 9, p. 256, figs. 7, 8, 13.
Lyramula furcula Hanna, Glezer, 1966 (part), in Cryptogamic plants of the
WES SAR, Vile snp caleople 2 figs: 3.05—7-
Remarks. — Deflandre (1950a, figs. 167-169) illustrated speci-
mens with one or more lateral rods as a premutant form of the
present species, but the topotypic sample, CAS 1144, did not yield
such specimens.
172 BuLueTIN 273
Dimensions. — Length of lateral rods (from the base of Y),
45-55 uw.
Known geologic range. — Cretaceous (late) to Paleocene
(early), Glezer reported the early Paleocene occurrence from west-
ern Siberia, Russia.
Occurrence. — The species is found to be abundant but only
in the topotypic material from California, Therefore, Mandra (1968,
p. 242) concluded that the present species seems to be restricted
stratigraphically to the late Cretaceous, at least from California.
Lyramula furcula var. minor Deflandre Pl. 27, figs. 19-21
Lyramula furcula var. minor Deflandre, 1940b, Acad. Sci. Paris, C. R., vol.
211, No. 21, p. 509, figs. 7-10.
Lyramula furcula Deflandre, 1950a, Microscopie, vol. 2, pp. 62/82-63/82,
figs. 170-173.
Lyramula furcula Hanna, Glezer, 1966 (part), in Cryptogamic plants of the
WS:S:Reavoly 74p.22t
Remarks. — Similar to the preceding taxon except that the
present variety is smaller in size (approximately one-half), and
frequently has a third lateral rod; thus the three rods are approxi-
mately 120° apart (see Pl. 27, fig. 21). Glezer (1966) considered
that the present small size is within the range of variation of the
type.
Dimensions. — Length of lateral rods (from the base of Y),
28-35 up.
Known geologic range. — Cretaceous (late).
Occurrence. —L. furcula var, minor is found only from the
CAS locality 1144 and rarely. Mandra studied a sample which came
from nearby this locality but failed to find the variety.
Lyramula simplex Hanna Pl. 28, fig. 1
Lyramula simplex Hanna, 1928, Jour. Paleont., vol. 1, No. 4, p. 262, pl.
41, fig. 6.
Lyramula simplex Hanna, Deflandre, 1940b, Acad. Sci. Paris, C. R., vol.
211, No. 21, p. 506, figs. 5, 6.
Lyramula simplex Hanna, Deflandre, 1950a, Microscopie, vol. 2, pp. 62/82-
62/82, figs. 164, 165.
Lyramula simplex Hanna, Tsumura, 1959, Yokohama Munic. Univ., Bull.,
vol. 11, Nat. Sci., vol. 1, p. 46, pl. 1, fig. 4 (fide Tsumura, 1963).
Lyramula simplex Hanna, Tsumura, 1963, Yokohama Munic. Univ., Jour.,
ser. C-45, No. 146, p. 31, pl. 1, fig. 2; pl. 3, fig. 4; pl. 16, fig. 4.
Lyramula simplex Hanna, Glezer, 1966, in Cryptogamic plants of the
WES SERS voll 75 ps 22d ipler2. figs 2.4.
Lyramula simplex Hanna, Mandra, 1968, Calif. Acad. Sci., Proc., ser. 4, vol.
36, No. 9, pp. 256-257, figs. 9, 10.
SILICOFLAGELLATES AND EBRIDIANS: LING IW
Remarks. — Specimen referred to here has thicker rods than
the L. furcula and clearly possesses fine surface ornamentation, as
Hanna (1928) originally described.
Dimensions. — Length of rods (from base of U), 82-90 ux.
Known geologic range. — Cretaceous (late).
Occurrence. —So far the present species is recovered rarely
from CAS 1144, the type locality for the present species.
Genus MESOCENA Ehrenberg, 1839, emend. Deflandre, 1950a
Remarks. — In a previous article, Ling (1970) discussed in de-
tail the taxonomic problems involved for a species, Mesocena cf.
elliptica. At that time, the observation was based only on deep-sea
sediments from the central north Pacific and also on the illustra-
tions of the reference, Since then the study has been expanded by
including many reference samples, and it is now possible to present
additional data of the detailed investigation.
The occurrence of a silicoflagellate form referable to Mesocena
was first described by Ehrenberg (1839, p. 129), and the figure
was illustrated by Ehrenberg (1854, pl. 22, fig. 41) as Dictyocha
triangula from Caltanisetta, Sicily.
It was Stradner who noticed in 1956 and 1961 for the first
time that some of the Mesocena specimens possess septa (“Scheide-
wand” of Stradner) in the basal body ring. The structure can be
detected if the specimen is carefully examined under the micro-
scope but it is more easily detected and clearly illustrated by utiliz-
ing a phase contrast attachment. Stradner further stated that
Mesocena with septa (Mesocena septatae) were derived from the
genus Corbisema, while those without septa (Mesocena aseptatae)
came from Dictyocha. Bachmann (1970b), then proposed the new
genus Septamesocena for the former.
Mesocena apiculata (Schulz) Pl. 28, figs. 2-4
Mesocena oamaruensis var. apiculata Schulz, 1928, Bot. Archiv., vol. 21,
No. 2, p. 240, fig. 11.
Mesocena polymorpha var. triangula Lemmermann, Gemeinhardt, 1930,
in Kryptogamen-Flora, vol. 10, pt. 2, p. 28, fig. 12a (only).
Mesocena apiculata Schulz, Hanna, 1931, Mining in California, vol. 27, No.
Zaple Distious:
Not Mesocena apiculata (Schulz), Deflandre, 1932c, Soc. Bot. France,
Bull., vol. 79, Nos. 5, 6, p. 499, figs. 34, 35.
Not Mesocena apiculata (Schulz), Tynan, 1957, Micropaleont., vol. 3, No.
2 ps 154. plata 10:
174 BULLETIN 273
Not Mesocena apiculata (Schulz), Stradner, 1961, Erdél u. Kohle, vol. 14,
No. 2, p. 89, figs. 28, 29, 32.
Not Mesocena apiculata (Schulz), Bachmann, 1962, Geol. B.-A., Verh.,
Jiahre? 19625 Nos 2, p: 379; fisss Osa:
Not Mesocena apiculata (Schulz), Bachmann, 1963, in Bachmann, et al.,
Geol. Ges. Wien, Mitt., p. 152, pl. 19, figs. 60-62.
Not Mesocena apiculata (Schulz), Bachmann, 1964, im Ichikawa, et al.,
Kanazawa Univ., Sci. Rept., vol. 9, No. 1, p. 107, pl. 4, fig. 36.
Mesocena aff. apiculata (Schulz), Glezer, 1966 (part), im Cryptogamic
plants of the US.S.R., vol. 7, p. 282, pl. 28, fig. 5 (?); pl. 33, fig. 7,
(not pl. 29, fig. 9).
Not Mesocena apiculata (Schulz), Hajos, 1968, Geol. Hungarica, ser.
Paleont., fasc. 37, p. 67, pl. 4, fig. 3.
Not Mesocena apiculata (Schulz), Bachmann and Papp, 1968, Giorn. di
Geol., ser. 2, vol. 35, fasc. 2, pl. 3, figs. 1, 2.
Not Mesocena apiculata (Schulz), Bachmann, 1970a, Geol. B.-A. Verh.,
Jahrg. 1970, pp. 280, 282, 284.
Remarks. — The above synonymy list is prepared to record
that the various forms were referred to this species by numerous
workers from many parts of the world. VW. apiculata is differentiated
from another species, Septamesocena apiculata Bachmann, which
will be discussed later, by the absence of septal structure at the
apices.
Dimensions. — Diameter of basal body ring, 36-46 »; of radial
spines, 2-6 yp.
Known geologic range. — Paleocene (?), Eocene, Miocene (?).
The type locality for the present species is Oamaru Diatomite, New
Zealand, of Eocene. The oldest occurrence, Paleocene, is reported by
Glezer (1966) from the eastern slope of the Ural region, and it
possesses a dentate basal body ring (pl. 28, fig. 5); such a specimen
has not yet been observed from samples of Oamaru Diatomite. An-
other Paleocene specimen (pl. 33, fig. 7) from the same Russian
location has a smooth basal body ring, but the structure at the
three apices is obscured. It is conceivable that previous students of
silicoflagellates may have overlooked the septal structure, and oc-
currence of the present species may be limited only to the Eocene
age.
Occurrence. — M. apiculata is not found in the deep-sea sedi-
ments examined during the present study. The specimens here
illustrated are from Bain’s Farm (PI. 28, figs. 2, 3) and Jackson’s
Paddock (PI. 28, fig. 4) sections of Oamaru Diatomite, New Zea-
land, for comparison with similar forms, particularly with Septa-
mesocena apiculata.
SILICOFLAGELLATES AND EBRIDIANS: LING 175
Mesocena circulus Ehrenberg Pl. 28, figs. 5, 6
Dictyocha (Mesocena) circulus Ehrenberg, 1840, Verh. K. Preuss. Akad.
Wiss. Berlin, Ber., p. 208.
Mesocena circulus Ehrenberg, Ehrenberg, 1844a, Verh. K. Preuss. Akad.
Wiss. Berlin, Ber., p. 65.
Dictyocha circulus Ehrenberg, Ehrenberg, 1854, Mikrogeologie, pl. 19, fig. 44.
Not Mesocena circulus Ehrenberg, Lemmermann, 1901b, Deutsch. Bot. Ges.
Bers volet9s ps 2517.
Not Mesocena circulus Ehrenberg, Schulz, 1928, Bot. Archiv., vol. 21, No.
2; p. 242, fig. 14.
Not Mesocena circulus Ehrenberg, Gemeinhardt, 1930, in Kryptogamen-
Flora, vol. 10, pt. 2, p. 33, fig. 18.
Mesocena circulus Ehrenberg, Tsumura, 1963, Yokohama Munic. Univ.,
Jour., ser. C-45, No. 146, p. 41 (not pl. 7, fig. 1).
Not Mesocena circulus Ehrenberg, Glezer, 1966, in Cryptogamic plants of
the U.S.S.R., vol. 7, pp. 285-286, pl. 29, fig. 8.
Remarks. — This species is characterized by a circular to oval-
shaped basal body ring with one series of radial spines generally in
a plane.
Confusion about the present species has resulted from the be-
ginning when Ehrenberg gave the original description of the present
species as “D. cellula unica circulari margine dentata Diam. - 1/48.
Fossils in marga Graeciae.” The first illustration given by Ehren-
berg (1854) came from upper Miocene, Aegina, Greece, and the
margin can be interpreted as either coarsely dentate or as possess-
ing about 17 short radial spines, The diameter of the specimen is
over 50 » according to Loeblich, III (et al., 1968, p. 217, fig. 20).
Lemmermann (1901b) added the Barbados and Caltanisetta
(Sicily) as additional localities, and although referring to Ehren-
berg’s description and figure, he indicated the species as “Gehiuse
kreisrund, 20-30 » gross, glatt oder am dussern Rande schwach
gezahnt.” It is thus apparent that Lemmermann’s specimens are
about 1% the size of Ehrenberg’s. Schulz (1928) followed Lemmer-
mann’s description, and the figure given from Mors Island is oval-
shaped. Schulz at the same time observed that a specimen from
Kusnetzk, Russia, has only a 14 p» diameter. Gemeinhardt (1930)
followed Lemmermann’s and Schulz’s concept. Tsumura (1963) also
recorded the present species, but his specimen (pl. 7, fig. 1) seems
to possess two series of radial spines and should be transferred to
M. circulus var. apiculata. Glezer (1966) also referred a specimen
from the Ural (pl. 29, fig. 8) to the present species, but it shows
a finely dentate margin and not radial spines. A similar form is
176 BULLETIN 273
reported by Hajos (1968) and classified as M. elliptica var. circulus
which is later followed by Bachmann (1970b).
Dimensions. — Diameter of basal body ring, 68-72 y; of radial
spines, 3-6 p.
Known geologic range. — Eocene to Miocene.
Occurrence. — The present species is recovered from two
samples, EM 8-12, 26-28 cm, and EM 8-11, 69-70 cm only which
are within Mohnian age by Parker (1964), Bandy and Ingle (1970),
and Tortonian (?) of middle Miocene by Martini and Bramlette
(1963).
Mesocena circulus var. apiculata Lemmermann Pl.-28; figs: 57,0
Mesocena circulus var. apiculata Lemmermann, 1901b, Deutsch Bot. Ges.,
Ber., vol. 19, p. 257, pl. 10, figs. 9, 10.
Mesocena circulus var. apiculata Lemmermann, Schulz, 1928, Bot. Archiv.,
vol. 21, No. 2, pp. 242-243, fig. 15a, b, c.
Mesocena circulus var. apiculata Lemmermann, Gemeinhardt, 1930, in
Kryptogamen-Flora, vol. 10, pt. 2, pp. 33-34, fig. 19.
Distephanus polyactis fa. mesocenoidea Deflandre, 1932c, Soc. Bot. France,
Bull., vol. 79, Nos. 5, 6, fig. 41 (only).
Mesocena circulus var. apiculata Lemmermann, Zanon, 1934, Acta Pont.
Acad. Sci. Nov. Lincei, vol. 87, p. 65, fig. 5.
Paradictyocha polyactis fa. mesocenoidea (Deflandre), Frenguelli, 1940, Rev.
Mus. La Plata, n.s., vol. 2, Paleont., No. 7, p. 53, fig. 8b, d, e (?)
(only).
Paradictyocha apliculata Lemmermann, Frenguelli, 1940, Rev. Mus., La
Plata, n.s., vol. 2, Paleont., No. 7, p. 53, fig. 7h.
Mesocena circulus var. apiculata Lemmermann, Tsumura, 1959, Yokohama
Munic. Univ., Bull., vol. 11, Nat. Sci., No. 1, p. 55, pl. 2, figs. 19-21
(fide Tsumura, 1963).
Mesocena circulus var. apiculata Lemmermann, Tsumura, 1963, Yokohama
Munic. Univ., Jour., ser. C-45, No. 146, p. 41, pl. 6, figs. 7, 8; pl. 20,
figs. 1-2.
Mesocena circulus Ehrenberg, Tsumura, 1963, Yokohama Munic. Univ.,
Jour., ser. C-45, No. 146, p. 41, pl. 7, fig. 1.
Paradictyocha polyactis fa. mesocenoidea Frenguelli, Glezer, 1966, in Cryp-
togamic plants of the U.S.S.R., vol. 7, p. 278, pl. 28, fig. 1.
Remarks. — Ling (1971) discussed the present taxon in detail
on the basis of Japanese Miocene specimens. Frenguelli’s Para-
dictyocha apiculata from Sendai, Japan, seems likely to belong to
the present taxon judging from the illustration.
Dimensions. — Diameter of basal body ring, 64-74 yp; of radial
spines, 3-5 p.
Known geologic range. — Eocene to Miocene. Schulz (op. cit.,
fig. 15c) described the only Eocene form for the present variety
SILICOFLAGELLATES AND EBRIDIANS: LING 177
from Oamaru, New Zealand. Examination of Oamaru Diatomite by
the author so far has failed to recover such a specimen.
Occurrence. — Specimens identified as belonging to the pres-
ent variety were found only from experimental Mohole samples
from EM 7-1, 35-36 cm, to EM 6-2, 10-11 cm, with their occur-
rence apparently limited to only middle and late Miocene.
Mesocena cf. elliptica Ehrenberg Pl. 28, figs. 9-15; Pl. 29, figs. 1-3
Remarks. — As discussed by Deflandre (1932c), Glezer (1966),
and Ling (1970), the taxonomy of the present species is one of
the most uncertain among silicoflagellates. On the one hand, differ-
ent taxonomic names were proposed, based on the number of radial
spines (Ehrenberg, Lemmermann); and on the other hand, all these
forms are grouped in only one species (Deflandre, Glezer). Be-
cause of this and because of the limited number of samples ob-
served at that time, Ling (1970, 1971) referred to these forms as
M. cf. elliptica. It is extremely unfortunate that Hajos (1968)
and later Bachmann (1970b) proposed a new combination and
grouped them as M. elliptica, apparently following Deflandre’s prac-
tice (1932c), but then proposed WM. e. var. quadrangula Bachmann
and Ichikawa (1962) in place of M. e. var, elliptica (see Lanjouw,
et al., 1966, ICBN, Art. 25). Furthermore inclusion of M. circulus
as one of the variety within this species also seems undesirable.
During microscopic examination, an effort was made to seek
the biostratigraphic occurrences of the taxa even by recognizing
separately those distinguished on the basis of the number of radial
spines (see Text-figs. 3, 4, 7).
The following is apparent from the present study:
1. No specimen with three radial spines that may be refer-
able to Dictyocha triangula and not M. apiculata or Septamesocena
apiculata has been found.
2. The four-spine form, “MV. elliptica”, stratigraphically appears
earlier and ends slightly later than the two-spine “MM. diodon”,
3. Only one specimen assignable to M. hexagona Haeckel (see
Ling, 1971) is found from sample EM 8-13, 49-51 cm (“h” in Text-
fig. 4).
4. Only a single specimen similar to MV. polymorpha var. quad-
178 BULLETIN 273
rangula Lemmermann (1901b) is found from sample WR TR 11,
Low (Bo. 202) (“q” in Text-fig. 3).
As in previous articles, a wide range is considered for the
present taxon.
Dimensions. — Length of basal body ring, 43-70 pn.
Known geologic range. — Eocene to Pleistocene
Occurrence. — In the Mediterranean section of Italy (see Text-
figure 7), the initial appearance of so-called “M. elliptica” starts
with the beginning of the Calocycletta costata radiolarian Zone
(Sanfilippo, 1971), and apparently this seems the earliest for this
form so far recognized during the present investigation. From
other samples, the first occurrence is noticed from Trinidad sample,
WR TR 11, Low (Bo. 202), which is assigned to the Dorcadospyris
alata Zone by Riedel and Sanfilippo (1970). The highest strati-
graphic occurrence of the four-spine form is in LSDA 101G, 38-39
cm which is N.17 or 18 foraminiferal Zone or Spongaster pentas
Zone (Riedel and Sanfilippo, 1970).
On the other hand, the two-spine form (“M. diodon’’) is found
only from the Mohole samples, the interval between EM 8-15,
505-506 cm, to EM 6-2, 30-32 cm, ranging fom Luisian to Mohnian.
Mandra (1968) indicated the range of his M. crenulata var. diodon
from Valmonte Diatomite (Mohnian) to Buttle Diatomite (Del-
montian),
The specimen of the “M. hexagona’ is found in a sample of the
lower Mohnian stage, but Mandra found his specimens from the
Buttle Diatomite (Delmontian) of California,
Mesocena cf. elliptica var. minoriformis Bachmann and Papp
Pl. 29, figs. 4-7
Mesocena elliptica minoriformis Bachmann and Papp, 1968, Giorn. di Geol.,
sera 2-vole 55 plo sips 121. tig:
Mesocena elliptica var. minoriformis Bachmann and Papp, Bachmann,
1970b, zz Cat. Fossil. Austriae, No. 1b, p. 13.
Remarks. — Bachmann kindly sent the author possible topo-
typic material from Ernstbrunn, Austria (Aus. 3 in this paper).
The author agrees that the size of the present variety is definitely
smaller than that of M. cf. elliptica, and also accepts that the present
variety has “relativ starken Basalringteile wesentlich von der geo-
logisch jiingeren Art .. .” M. cf. elliptica was also found in the
SILICOFLAGELLATES AND EBRIDIANS: LING 179
sample Bachmann sent, although he did not indicate such occur-
rence in his table.
Dimensions. — Diameter of basal body ring, 28-50 up.
Known geologic range.— Miocene (Burdigalian).
Occurrence. — Similar to M. cf. elliptica, this Bachmann’s var-
iety is found earlier in Italian samples of the Calocycletta virginis
Zone than in another part of the area of the present study, WR TR
23 (K9391), which is the Calocycletta costata Zone (Riedel and
Sanfilippo, 1970) from Trinidad. The highest occurrence is from
sample EM 8-13, 125-126 cm which is lower Mohnian according to
Bandy and Ingle (1970) and Parker (1964) or lower part of Tor-
tonian (?) by Martini and Bramlette (1963).
Mesocena oamaruensis Schulz Pl. 29, figs. 8-10
Mesocena oamaruensis Schulz, 1928, Bot. Archiv., vol. 21, No. 2, p. 240,
fig. 10a, b.
Mesocena oamaruensis Schulz, Gemeinhardt, 1930, in Kryptogamen-Flora,
vol. 10, pt. 2, p. 34, fig. 20.
Mesocena oamaruensis Schulz, Deflandre, 1932a, Soc. France, Microsc.,
Bull., vol. 1, No. 1, fig. 11.
Corbisema oamaruensis (Schulz), Frenguelli, 1940, Rev. Mus., La Plata,
n.s., vol. 2, Paleont., No. 7, p. 64, fig. 13c.
Mesocena oamaruensis Schulz, Tsumura, 1959, Yokohama Munic. Univ.,
Bull., vol. 11, Nat. Sci., No. 1, p. 56, pl. 2, fig. 22 (fide Tsumura, 1963).
Not Mesocena oamaruensis Schulz, Stradner, 1961, Erdél u. Kohle, vol. 14,
No. 2, p. 89, figs. 30, 31.
Not Mesocena oamaruensis Schulz, Tsumura, 1963, Yokohama Munic. Univ.,
Jour., ser. C-45, No. 146, p. 42, pl. 7, fig. 2; pl. 21, fig. 12.
Not Mesocena oamaruensis Schulz, Bachmann, 1964, in Ichikawa ef al.,
Kanazawa Univ., Sci. Rept., vol. 9, No. 1, pl. 4, fig. 37.
Mesocena oamaruensis Schulz, Glezer, 1966, in Cryptogamic plants of the
WES SHR EViOlNa/. eps 2ooe Dla ZSnetion se
Remarks. — One of the forms illustrated here (Pl. 29, fig. 8)
is from sample $136/1096 of the Forrester’s Hill section of Oamaru
Diatomite, New Zealand. It is apparent that the basal body ring
is smooth, without either radial spines or accessory spines. Lumen
is apparent in the specimen but no septa. The specimens from
Kellogg and Sidney Shales of California (see Pl. 29, figs. 9, 10)
beyond any doubt are conspecific with that of Oamaru Diatomite,
except that the Californian specimens are slightly smaller in size.
Stradner’s form is illustrated as having septa as also does
Bachmann’s specimen from the Hojuji diatomaceous mudstone of
Japan; therefore, they are excluded from the present species (see
180 BULLETIN 273
below). Tsumura also described specimens from Jackson’s Paddock,
Oamaru, New Zealand, but his illustration (pl. 7, fig. 2) can be
interpreted as having septa, and another specimen (pl. 21, fig. 12)
as having spines at the radial corners; therefore they are excluded
from the present species.
Dimensions. — Length of basal body ring, 60-90 up.
Known geologic range. — Eocene (late), Oligocene (early ?).
As discussed by Loeblich, III (et al., 1968, p. 18) and is accepted
here, several localities or sections from the so-called Oamaru Dia-
tomite are now considered as late Eocene age. The only Oligocene
but questionable record for the present species is reported by Glezer
(1966) from Dnepropetrovsk of European Russia. Thus it is con-
ceivable that the present species has limited stratigraphic range
of only late Eocene.
Occurrence. — During the present study, no specimen referable
to the present species was recovered from the deep-sea sediments.
The present species is discussed here and illustrated for comparison
purposes to differentiate it clearly from the very similar form,
Septamesocena apiculata, that previous workers assigned erron-
eously.
Genus NAVICULOPSIS Frenguelli, 1940
Remarks. — The silicoflagellate with naviculoid basal body
ring was named by Ehrenberg (1839) as Dictyocha navicula from
Miocene sediments of Zante, Greece. The detailed structure of these
specimens was discussed by Frenguelli (1940, p. 60), who first
noticed that in some specimens the middle part of the basal body
ring where the apical structure joins is sometimes transformed into
a thin flattened form thus losing the general tubular nature; there-
fore, in either apical or abapical view, the tubular skeleton is
suddenly narrowed to capillary form in the middle of the basal body
ring. Deflandre (1950a) later concurred in this opinion, although
both Frenguelli and Deflandre failed to mention that the middle
of the basal body ring frequently shows some constriction. Schulz
observed such a phenomenon and described it as Dicityocha navicula
var. constricta. As can be seen in the illustrations of different taxa
within the present genus, such constriction is found rather com-
monly.
SILICOFLAGELLATES AND EBRIDIANS: LING 181
Another important point that deserves attention for its taxo-
nomic significance has been neglected, In 1950, Deflandre (p. 36/82)
indicated that in rare cases there are some specimens that show
flattening of the two apices of the major axis, and he indicated that
he found such forms in samples from Zante, Greece and Saint-
Laurent-La Vernede, France, but only those from Zante were illus-
trated under the name of Naviculopsis navicula (Ehrenberg) (his
figs. 241-243). Later Stradner (1956, 1961) also noticed this struc-
ture, called “Endflichen”, which exhibits small projections or spines
from both sides of the plates (see 1961, abb. 1 A; figs. 46, 47).
Bachmann (1970a) also illustrated such specimens from his Ernst-
brunn, Austria sample. Note that in these specimens, no radial
spines were observed. The taxonomic significance of such flattened
nature of the apices is apparent and should not be confused with
the small flattened hyaline area found between the tubular basal
body ring of the apices as discussed by Glezer (1966, p. 254) for
N. foliacea.
Finally it should be pointed out here that in some naviculoid
forms these two transformations just described never take place
and thus the tubular nature is preserved completely throughout
the basal body ring.
These three different types within the genus of Naviculopsis
may suggest different phylogenic lineages. However, the author pre-
fers to await further study of these forms, particularly examination
of Eocene samples from New Zealand and California.
Naviculopsis biapiculata (Lemmermann) Pl. 30, figs. 1-4
Dictyocha navicula var. biapiculata Lemmermann, 1901b, Deutsch. Bot. Ges.,
Ber., vol. 19, p. 258, pl. 10, figs. 14 (?), 15.
Dictyocha navicula var. biapiculata Lemmermann, Schulz, 1928, Bot. Archiv.,
vol. 21, No. 2, pp. 244-245, figs. 18, 19.
Dictyocha navicula var. biapiculata fa. aspera Schulz, 1928, Bot. Archiv.,
vol. 21, No. 2, p. 246, fig. 20a, b
Dictyocha navicula var. biapiculata Lemmermann, Gemeinhardt, 1930,
_ in Kryptogamen Flora, vol. 10, pt. 2, pp. 36-37, fig. 24a (only).
Dictyocha (?) biapiculata Lemmermann, Hanna, 1931, Mining in California,
vol. 27, No. 2; pl. D; figs. 5 (?). 6.
Dictyocha navicula var. biapiculata Lemmermann, Deflandre, 1932a, Soc.
France, Microsc., Bull., vol. 1, No. 1, figs. 15-17 (not fig. 18).
Not Dictyocha navicula var. biapiculata Lemmermann, Zanon, 1934, Acta
Pont. Acad. Sci. Nov. Lincei, vol. 87, p. 65, fig. 6.
Naviculopsis biapiculata (Lemmermann), Frenguelli, 1940, Rev. Mus., La
(aa n.s., vol. 2, Paleont., No. 7, p. 60, fig. 11c, d (as fa. normal)
only).
182 BULLETIN 273
Dictyocha navicula var. apiculata Lemmermann, Tsumura, 1959, Yokohama
Munic. Univ., Bull., vol. 11, Nat. Sci, No. 1, p. 58, pl. 3, figs 24-26
(fide Tsumura, 1963).
Dictyocha navicula var. biapiculata Lemmermann, Tsumura, 1963, Yoko-
hama Munic. Univ., Jour., ser. C-45, No. 146, p. 45, pl. 8, figs. 1-3; pl.
2ieetigs: 7-9:
Naviculopsis biapiculata var. biapiculata Glezer, 1966, im Cryptogamic
plants of the U.S.S.R., vol. 7, pp. 255-256, pl. 16, figs. 2, 4-5.
Naviculopsis biapiculata (Lemmermann), Mandra, 1968 (part), California
Acad. Sci., Proc., ser. 4, vol. 36, No. 9, p. 264, figs. 20, 21 (only).
Remarks. — As described and illustrated in his original figure
15, when Lemmermann (1901b) proposed this species, he indi-
cated that the apical structure was in the form of an apical bar
(“Kieselbalken”). Although no reference samples are available from
Dolje, Yugoslavia, Fuur Island or Kusnetzk, Russia, the localities
from which Lemmermann’s new taxon was proposed, samples from
Jackson’s Paddock section of the Oamaru Diatomite are here illus-
trated (PI. 30, figs. 1, 2) in lateral view to show the nature of
the transition of the basal body ring to the apical bar, Also as will
be demonstrated later, the basal body ring of the specimens be-
longing to the present genus always remains on the same plane of
the radial spines in lateral view, and is never warped or curved
upward as Lipps (1970, fig. 3, bottom figure) illustrated.
The above synonymy is prepared by adhering to the original
definition. Thus, Gemeinhardt, although referring his specimen to
that of Lemmermann’s, described the apical structure of his as
“kleineren mitunter besonders breit, bandartig, nach der mitte zu
schmalerwerdend.” Therefore, his figure 24b, which clearly shows
such a broad apical structure is excluded from the present species.
One of Mandra’s specimens (1968, fig. 22) is also excluded on the
same basis and is referred to as N. foliacea. In addition, as Deflandre
(1950a) indicated, Schulz’s fa. aspera is included as a morpho-
logic variation of the present species.
Dimensions. — Length of basal body ring, 40-54 »; of radial
spine, 34-50 p. ;
Known. geologic range. — Eocene to Oligocene (early ?). Pos-
sible early Oligocene occurrence is reported by Glezer (1966) from
several locations of western Siberia.
Occurrence. — The present species is found rarely and from
DSDP 6-6-2, 1-2 cm, upper part of Hantkenina aragonensis plank-
tonic foraminiferal Zone (Ewing, et al., 1969), or Thyrsocyrtis tri-
SILICOFLAGELLATES AND EBRIDIANS: LING 183
acantha Zone (Riedel, 1971) and WR TR 39 F (J.S. 1068) from
Barbados, which is the Theocyrtis tuberosa Zone (Riedel and San-
filippo, 1970).
Naviculopsis constricta (Schulz) Pl. 30, figs. 5-8
Dictyocha navicula var. biapiculata fa. constricta Schulz, 1928, Bot. Archiv.,
vol. 21, No. 2, p. 246, fig. 21.
Dictyocha navicula var. biapiculata fa. constricta Schulz, Gemeinhardt,
1930, in Kryptogamen-Flora, vol. 10, pt. 2, p. 38, fig. 25.
Naviculopsis constricta (Schulz), Frenguelli, 1940, Rev. Mus., La Plata,
n.s., vol. 2, Paleont., No. 7, fig. 11a, b.
Not Naviculopsis constricta (Schulz), Stradner, 1961, Erdél u. Kohle, vol.
14, No. 2, p. 89, pl. 2, figs. 49-52.
Not Dictyocha var. biapiculata fa. constricta Schulz, Tsumura, 1963, Yoko-
hama Munic. Univ., Jour., ser. C-45, No. 146, p. 46, pl. 8, fig. 8.
Naviculopsis biapiculata var. constricta (Schulz), Glezer, 1966, in Crypto-
gamic plants of the U.S.S.R., vol. 7, pp. 257-258, pl. 17, fig. 4.
Description. — Basal body ring, oval, frequently constricted
at the middle where the apical structure joins; two long radial spines
at each end of the major axis; apical structure in the form of an
arched band with more or less uniform width throughout except
broadened slightly where connected with the basal body ring; no
apical nor basal accessory spines present.
Remarks. — Schulz (1928) indicated when he proposed his
new fa. contricta that the middle part of the basal body ring is,
in either apical or abapical view, constricted. As already noted by
Frenguelli (1940) when he discussed the present genus, such a
phenomenon is common among the naviculoid silicoflagellates be-
cause of transformation of the basal body ring from tubular to
flattened nature. The present species can be recognized by the
band-form apical structure and generally shows uniform width
throughout; thus it can be differentiated from those either with an
apical bar like N. biapiculata or with a foliated apical plate such as
N. foliacea. Depending on the width of the apical band, the size
of lateral windows is variable.
Dimensions. — Length of basal body ring, 26-34 »; of radial
spines, 18-36 u.
Known geologic range. — Eocene (late). Glezer listed the age
of Kreyenhagen Formation of California and Oamaru Diatomite,
New Zealand as “late Eocene or early Oligocene” but they are
now regarded as Eocene (see Loeblich, III et al., 1968, p. 18).
Therefore, the geologic range for the present species seems limited
to middle and late Eocene.
184 BULLETIN 273
Occurrence. — Like the preceding taxon, this species is found
only from samples of DSDP and Barbados.
Naviculepsis foliacea Deflandre Pl. 30, figs. 9-11
Dictyocha (?) biapiculata Lemmermann, Hanna, 1931 (part), Mining in
California, vol. 27, No. 2, pl. E, fig. 10 (only).
Dictyocha (?) biapiculata Lemmermann, Clark and Campbell, 1945 (part),
Geol. Soc. Amer., Mem. 10, p. 3, pl. 5, fig. 8 (not others).
Naviculopsis foliacea Deflandre, 1950a, Microscopie, vol. 2, pp. 76/82-77/82,
figs. 235-240.
Naviculopsis foliacea Deflandre, Stradner, 1961, Erd6él u. Kohle, vol. 14, No.
2p. 89 ities 3:
Naviculopsis foliacea Deflandre, Glezer, 1966, in Cryptogamic plants of the
WAStS Reaviole 7, ps 2585 wDlea7ette. 5.
Naviculopsis biapiculata (Lemmermann), Mandra, 1968 (part), Calif.
Acad. Sci., Proc., ser. 4, vol. 36, No. 9, p. 264, fig. 22 (only).
Remarks.— The specimens found during the present study
from the Deep Sea Drilling Project (Pl. 31. fig. 11) agree with the
description and illustration originally given by Deflandre from
Barbados. Although no sample is available from the Kreyenhagen
Shale where Hanna apparently described the present species, speci-
mens from the Kellogg Shale and Sidney Shale of California are
illustrated (PI. 31, figs. 10, 11). The nature of the foliated apical
plate and the size variation of the apical plate in contrast to that
of basal body ring can be seen in these figures.
Note here also that in DSDP samples, there is a flattened
hyaline area between the basal body ring at the apices which is not
evident in specimens from California. Therefore, Glezer is of the
opinion that the present species should be limited to those with
the flattened area. As indicated earlier, however, the significance
of such a flattened area should not be confused with that of the
completely flattened end plate found in N. navicula. It is the au-
thor’s opinion that the presence of the foliated apical plate, from
which the name of the present species is apparently derived, is
characteristic; and the value of the flattened area needs to be
confirmed in the future after more specimens are studied, The
California specimens are, therefore, included in the present species.
Dimensions. — Length of basal body ring, 40-56 u; of radial
spine, 30-36 yp.
Known geologic range. — Eocene. The age of “Diatomo-radio-
larite de Springfield, Ile de la Barbade,” which was considered as
“Oligocéne-Miocéne ?” by Deflandre (1950a, p. 76/82) and which
SILICOFLAGELLATES AND EBRIDIANS: LING 185
is followed by Glezer (1966), is now regarded as Eocene by Loeblich,
DD (et al3 1968, p. 18>):
Occurrence. — N. foliacea here discussed is found in several
DSDP samples of middle Eocene age. With consideration of the
Californian specimens, this species has the highest occurrence at
the top of Eocene or bottom of Oligocene from the Barbados
sample WR TR 39 F (J.S. 1068).
Naviculopsis lata (Deflandre) Pl. 30, figs. 12-16
Dictyocha biapiculata var. lata Deflandre, 1932c, Soc. Bot. France, Bull.,
vol. 79, p. 500, figs. 30, 31.
Dictyocha navicula var. biapiculata Lemmermann, Zanon, 1934, Acta Pont.
Acad. Sci. Nov. Lincei, vol. 87, p. 65, pl. n, fig. 6.
Naviculopsis lata (Deflandre), Frenguelli, 1940, Rev. Mus., La Plata, n.s.,
vol. 2, Paleont., No. 7, p. 61, fig. 11h (?).
Dictyocha ausonia var. regularis Carnevale, Deflandre, 1950a, Microscopie,
vol. 2, pp. 68/82-69/82, fig. 197 (only).
Naviculopsis robusta Deflandre, Stradner, 1961, Erdél u. Kohle, vol. 14,
No. 2, p. 89, pl. 2, figs. 39-41, 43-45 (not fig. 42).
Naviculopsis robusta Deflandre, Bachmann and Papp, 1968, Giorn. di Geol.,
ser. 2, vol. 35, fasc. 2, p. 119, pl. 3, figs. 3, 4.
Naviculopsis lata (Deflandre), Bachmann, 1970a, Verh. Geol. B.-A., Jahrg.
1970, No. 2, pp. 278-279, pl. 2, figs. 1-20; pl. 3, figs. 1-15.
Remarks. — As discussed in detail by Bachmann (1970a), the
present species is apparently similar to another species, Naviculopsis
robusta, from the Eocene Diatomite of Singhiliewski, Russia, but
the latter is characterized by possessing a small triangular, foliated
lateral plate where the basal body ring and apical bar meet. So far
in this study, a specimen with this structure as Deflandre (1950a)
and Glezer (1966) illustrated has not been seen; therefore, it is
agreed that N. robusta has a limited biogeographic distribution.
Some of the present species show a slight constriction in the
middle of the basal body ring where an apical bar forked and
merged, and such a phenomenon can be observed easily from the
abapical view, particularly under phase contrast (Pl. 30, fig. 14),
but in no case does the basal body ring show a flattened nature.
Apparently Deflandre (1950a), by considering the wide range
of variation for Dictyocha ausonia, regarded the specimen of the
present species as within its variation. As discussed earlier, the varia-
tion of a gradual narrowing and the disappearance of radial spines
on the minor axis is not so readily as shown by Deflandre for
Dictyocha ausonia (figs. 194-196) or D. mutabilis (figs. 203-208).
186 BULLETIN 273
Dimensions. — Length of basal body ring, 42-52 p; of radial
spine, 10-16 p.
Known geologic range. — Oligocene (late).
Occurrence. — During the present study, NV. lata is found only
from Mediterranean Italian samples ranging from near the bottom
of Calocycletta virginis Zone to the highest available samples of
Calocycletta costata Zone (Sanfilippo, 1971).
Naviculopsis navicula (Ehrenberg) Pl. 30, figs. 17-19; Pl. 31, fig. 1
Dictyocha navicula Ehrenberg, 1839, K. Preuss. Akad. Wiss. Berlin, Abh.,
Jahrg. 1838, p. 129.
Dictyocha ponticulus Ehrenberg, 1844c, Verh. K. Preuss. Akad. Wiss. Ber-
lin; Ber., p. 267-
Dictyocha ponticulus Ehrenberg, Bailey, 1845, Amer. Jour. Sci. Arts., vol.
48, No. 2, p. 328, pl. 4, fig. 21.
Dictyocha navicula Ehrenberg, Ehrenberg, 1854, Mikrogeologie, pl. 20, No.
1, fig. 43.
Dictyocha navicula Ehrenberg, Haeckel, 1887, Rept. Voy. Challenger, Zool.,
vol. 18, p. 1559.
Dictyocha navicula Ehrenberg, Lemmermann, 1901b, Deutsch. Bot. Ges.,
Ber., vol. 19, pp. 257-258, pl. 10, figs. 12-13 (only).
Dictyocha navicula Ehrenberg, Lemmermann, 1903, in Nordisches Plankton,
Bot. No: 21, p: 27, fig. 90:
Dictyocha navicula Ehrenberg, Schulz, 1928, Bot. Archiv., vol. 21, No. 2,
p. 243, fig. 16b (only).
Dictyocha navicula Ehrenberg, Gemeinhardt, 1930, in Kryptogamen-Flora,
vol. 10, pt. 2, pp. 35-36, fig. 22.
Not Dictyocha navicula (Ehrenberg), Deflandre, 1932a, Soc. France, Mi-
crosc., Bull., vol. 1, No. 1, fig. 13.
Not Corbisema apiculata fa. naviculoidea Frenguelli, 1940, Rev. Mus., La
Plata, n.s., vol. 2, Paleont., No. 7, fig. 12i.
Not Corbisema apicuata var inermis Lemmermann, Deflandre, 1950a, Micro-
scopie, vol. 2, fig. 88.
Naviculopsis navicula (Ehrenberg), Deflandre, 1950a, Microscopie, vol. 2,
p. 77, figs. 241-243.
Naviculopsis navicula (Ehrenberg), Stradner, 1961, Erdé] u. Kohle, vol.
14, No. 2, p. 89, fig. 47 (only).
Dictyocha navicula var. naviculopsis (Deflandre), Tsumura, 1963, Yoko-
hama Munic. Univ., Jour., ser. C-45, No. 146, p. 44, pl. 7, fig. 5.
Not Dictyocha navicula Ehrenberg, Glezer, 1966, in Cryptogamic plants of
the U.S.S.R., vol. 7, pp. 233-235, pl. 9, figs. 4, 5.
Naviculopsis navicula (Ehrenberg), Bachmann, 1970a, Verh., Geol. B.-A.,
Jahrg. 1970, No. 2, fig. 2b (only).
Remarks. — The specimens recovered from Northern Italy agree
in general with the first figure for the present species illustrated
by Ehrenberg from Zante, Greece, except that they are slightly
smaller in size.
In the illustration of Deflandre (1950a), the flattened struc-
ture at the apices of the major axis and the rudimentary spines
SILICOFLAGELLATES AND EBRIDIANS: LING 187
are seen for the first time. As shown here the degree of development
of such a structure differs among the specimens but can be found
only after careful examination (see Pl. 30, fig. 19). Therefore, it is
the author’s opinion that such a structure perhaps escaped notice
by previous workers particularly prior to 1950. The above synonymy
is prepared with this in mind.
In addition to listing the fossil locations, Haeckel reported
(1887, p. 1559) the present species “living in the Atlantic, Stations
352, 354 and off Bermuda, surface”; and later workers such as
Lemmermann and Gemeinhardt, apparently followed these records.
Judging from geological occurrence of the present species (see be-
low), it seems likely that the above record of living forms for the
present species has been either reworked from nearby land deposits
or may be outcrops of Tertiary deposits at the ocean bottom.
Glezer referred Russian specimens all to Eocene, but her illus-
trations clearly show they are different from the present taxon by
possessing the basal accessory spines, as well as a much wider basal
body ring along the minor axis.
Dimensions. — Length of basal body ring, 18-24 up.
Known geologic range. — Miocene.
Occurrence. — The specimen referable to the present N. navi-
cula is found, in the present study, only from the Italian samples
and only from WRE 67-77 and -76 samples of the Calocycletta
costata Zone. Apparently this species has a limited biogeographic
as well as stratigraphic range,
Naviculopsis quadrata (Ehrenberg) Pest. figs 2
Dictyocha quadrata Ehrenberg, 1844c, Verh. K. Preuss. Akad. Wiss. Berlin,
_ Ber., Jahrg., 1844, pp. 258, 267.
Dictyocha quadrata Ehrenberg, Bailey, 1845, Amer. Jour. Sci. Arts, vol. 48,
_ No. 2, p. 328, pl. 4, fig. 22.
Dictyocha quadrata Ehrenberg, Haeckel, 1887, Rept. Voy. Challenger, Zool.,
_ vol.18; (p. 1559.
Dictyocha quadrata Ehrenberg, Lemmermann, 1901b, Deutsch. Bot. Ges.,
Ber., vol. 19, p. 259.
Dictyocha quadrata Ehrenberg, Lemmermann, 1903, in Nordisches Plankton,
‘ Bot., No. 1, p. 27.
Dictyocha navicula var. rectangulare Schulz, 1928 (part), Bot. Archiv., vol.
_ 21, No. 2, pp. 243-244, fig. 17a (only).
Dictyocha navicula var. rectangulare Schulz, Gemeinhardt, 1930 (part),
in Kryptogamen-Flora, vol. 10, pt. 2, p. 36 (not fig. 23).
Naviculopsis rectangularis (Schulz), Frenguelli, 1940, Rev. Mus., La Plata,
_ ns., vol. 2, Paeont. No. 7, p. 60, fig. 11), k.
Dictyocha biapiculata (Lemmermann) var. iberica Deflandre, Colom, 1940,
Las Ciencias, vol. 5, No. 2, p. 351, fig. 9 (fide Deflandre, 1950a).
188 BuLLeETIN 273
Naviculopsis iberica Deflandre, 1950a, Microscopie, vol. 2, pp. 74/82-76/82,
figs. 231-234.
Naviculopsis rectangularis (Schulz), Stradner, 1961, Erdél u. Kohle, vol.
14, No. 2, p. 89, pl. 2, fig. 48.
Dictyocha navicula var. rectangularis Schulz, Tsumura, 1963, Yokohama
Munic, Univ., Jour., ser. C-45, No. 146, pp. 45-46, pl. 7, figs. 8-10; pl.
2s figs =3r
Naviculopsis rectangularis (Schulz), Bachmann, 1970a, Verh, Geol. B.-A.,
Jahrg. 1970, No. 2, p. 284, abb. 3, figs. a-f.
Naviculopsis rectangularis (Schulz), Bachmann, 1970b, im Cat. Fossil. Aus-
triae, No. 1b, p. 17.
Remarks.— The present species is described by Ehrenberg
(1844c) from “Bermuda-Inseln” and is illustrated by Bailey (1845)
from “Bermuda.” These locations are now regarded as actually
from Nottingham, Maryland, and the age as Helvetian stage ac-
cording to Loeblich, III (et al., 1968, pp. 16-17). These, however,
have escaped the attention of previous investigators. Thus Schulz
in 1928 when he proposed his Dictyocha navicula var. rectangulare
from the same locality, failed to mention Ehrenberg’s description
or Bailey’s illustration, and Schulz’s name has been adopted by
various students since that time. It should be mentioned here that
Schulz included two morphologically different forms under his
name. The form considered here is only that with a rectangular-
shaped basal body ring (fig. 17a only). It is possible that NV. tberica
described by Deflandre (1950a) from Spain and Majorca is a
somewhat irregular shape of the present species but still within
the range of intraspecific variation and, therefore, is considered
as synonymous.
Another specimen illustrated by Schulz at the same time (fig.
17b) has a elongated oval basal body ring which was erroneously
referred to recently as N. iberica by Bachmann (1970a, Abb. 1, fig.
Gh dink.)
Dimensions. — Length of basal body ring, 36-52 ,»; width,
24-30 p; radial spine, 6-8 yp.
Known geologic range. — Miocene.
Occurrence. —Only from Mediterranean Italian outcrop
samples, WRE 67-78 to -76, the Calocycletta costata radiolarian
Zone (Sanfilippo, 1971) are the present species recovered.
Naviculopsis regularis (Carnevale) Pl..31, figs; 35
Dictyocha regularis Carnevale, 1908, R. Inst. Veneto, Sci. Lett. Arti, Mem.,
vol. 28, No. 3, p. 35, pl. 4, fig. 28.
SILICOFLAGELLATES AND EBRIDIANS: LING 189
Dictyocha ausonia var. regularis Carnevale, Deflandre, 1950a, Microscopie,
vol. 2, pp. 68/82-69/82, fig. 198 (only).
Remarks. — This lenticular-shaped species possesses relatively
longer radial spines, and the size seems to agree with the middle
Miocene form originally described by Carnevale from Bergonzano,
Reggio Emilia, Italy.
Deflandre considered the present species as within the range
of variation for his Dictyocha ausonia, but as discussed earlier, it
is hard to conceive that two radial spines on the major axis can
become longer, while the other two on the minor axis diminish
(compare PI. 31, figs. 3-5 with Pl. 25, figs. 1-8).
The present species also resembles the preceding species, NV.
lata, by showing some constriction at the middle of the basal body
ring, but is generally smaller in size, and has a slender basal body
ring and no small lateral windows as found in the JN. Jata.
Dimensions. — Length of basal body ring, 33-38 uw; width,
19-24 yw; radial spines, 14-22 yp.
Known geologic range. — Miocene.
Occurrence. — N. regularis is found from the Italian samples
whose age has been identified as Calocycletta virginis and C. cos-
tata Zones (Sanfilippo, 1971).
Naviculopsis sp. Pls fig 6
Naviculopsis navicula Stradner, 1961 (not Ehrenberg), Erdél u. Kohle,
vol. 14, No. 2, p. 89, fig. 46 (only).
Naviculopsis navicula Bachmann and Papp, 1968 (not Ehrenberg), Giorn.
_ di Geol., ser. 2, vol..35, fasc. 2, pl. 3, fig. 5S.
Naviculopsis navicula Bachmann, 1970a (not Ehrenberg), Verh. Geol. B.-A.,
Jahrg. 1970, No. 2, abb. 2, figs. a, c-f (not b).
Remarks. — Although the present species is similar structurally
to NV. navicula, because it possesses the “Endflaichen” (Stradner,
1961), it differs in that the apices on the major axis are sharply
truncated, and it has two short spines at both corners of the plate.
The specimen illustrated here is from Ernstbrunn, Austria, the pos-
sible type locality of Stradner’s specimen for comparison.
Dimensions. —No complete specimen was observed.
Known geologic range. — Miocene (early).
Occurrence. — The specimen assignable to the present taxon
is found only from Ernstbrunn, Austria, and has not been found
in the deep-sea sediments,
190 BULLETIN 273
Naviculopsis trispinosa (Schulz) Pl. 31, figs. 7, 8
Dictyocha navicula var. trispinosa Schulz, 1928, Bot. Archiv., vol. 21, No.
2, pp. 246-247, fig. 23a, b.
Dictyocha navicula var. trispinosa Schulz, Gemeinhardt, 1930, im Krypto-
gamen-Flora, vol. 10, pt. 2, p. 40, fig. 27.
Dictyocha navicula var. trispinosa Schulz, Deflandre, 1932a, Soc. France,
Microsc., Bull., vol. 1, No. 1, fig. 24.
Naviculopsis biapiculata fa. trispinosa Schulz, Frenguelli, 1940, Rev. Mus.,
La Plata, nis:, vol. 2, Paleont., No. 7, fig: lle.
Dictyocha navicula var. trispinosa Schulz, Tsumura, 1963, Yokohama Mu-
nic. Univ., Jour., ser. C-45, No. 146, p. 46, pl. 8, figs. 4-6.
Naviculopsis trispinosa (Schulz), Glezer, 1966, in Cryptogamic plants of
the U.S.S.R., vol. 7, pp. 258-259, pl. 17, fig. 7.
Remarks. — Apparently the present species occurs rarely, and
during the present study, no specimen was found. The specimen
here illustrated is for reference purpose and particularly to present
the lateral view.
Dimensions. — Length of basal body ring, 24 yu; of radial spines,
25 pe
Known geologic range. — Eocene.
Occurrence. — No specimen belonging to N. trispinosa has been
found in the present study. The representative specimen presented
here is from a reference sample, $136/966, Bain’s Farm section,
Oamaru Diatomite, New Zealand.
Genus PARADICTYOCHA Frenguelli, 1940
Paradictyocha polyactis (Ehrenberg) Pl. 31, figs. 9-11
Dictyocha polyactis Ehrenberg, 1839, K. Akad. Wiss. Berlin, Abh., Jahrg.
1838, p. 129.
Dictyocha polyactis Ehrenberg, Ehrenberg, 1844a, Verh. K. Preuss. Akad.
Wiss. Berlin, Ber., Jahrg. 1844, p. 80.
Dictyocha polyactis Ehrenberg, Ehrenberg, 1854, Mikrogeologie, pl. 22,
fig. 50.
Distephanus speculum var. octonaria fa. polyactis (Ehrenberg), Jorgensen,
1899, Bergens Mus. Aarbog 1899, No. 6, pt. 2, p. 50.
Distephanus speculum var. polyactis (Ehrenberg), Lemmermann, 1901b
(part), Deutsch. Bot. Ges., Ber., vol. 19, pp. 265-266, pl. 11, fig. 17.
Distephanus speculum var. polyactis (Ehrenberg), Lemmermann, 1903, in
Nordisches Plankton, Bot., pt. 21, p. 31, fig. 106.
Not Distephanus speculum var. polyactis (Ehrenberg), Gemeinhardt, 1931b,
Deutsche Stidpolar-Exped., vol. 20 (Zool. vol. 12), p. 241, pl. 42, fig. 7.
Distephanus polyactis Deflandre, 1932c, Soc. Bot. France, Bull., vol. 79,
Nos. 5-6, pp. 501-502, fig. 40 (only).
Distephanus speculum var. polyactis fa. decagona Zanon, 1934, Acta Pont.
Acad. Sci. Nov. Lincei, vol. 87, p. 71, pl. n, fig. 47.
Distephanus speculum var. polyactis fa. dodecagona Zanon, 1934, Acta
Pont. Acad. Sci. Nov. Lincei, vol. 87, p. 71, pl. n, fig. 48.
Paradictyocha polyactis (Ehrenberg), Frenguelli, 1940, Rev. Mus., La Plata,
n:s., vol. 2) Paleont, No: 7, p: 52) fig: 7£ 2 1(?) sp. 54. fis) Sa (only)e
SILICOFLAGELLATES AND EBRIDIANS: LING 191
Paradictyocha polyactis fa. polyactis Glezer, 1966, in Cryptogamic plants of
the U.S.S.R., vol. 7, p. 277, pl. 24, figs. 7-11. ,
Paradictyocha polyactis (Ehrenberg), Mandra, 1968 (part), California
Acad; Sct., Proc.,, ser. 4, vol. 36, No. 9, p. 264.
Remarks. — The examination of related forms such as Meso-
cena circulus var. apiculata from various localities convinced the
author that: (a) the present species actually occurs only rarely;
(b) so far no intermediate forms have been found during the
present study demonstrating the morphologic variation of the
gradual disappearance of the apical structure as indicated by Fren-
guelli (1940) or the gradual enlargement of the apical ring to fuse
with the basal body ring indicated by Deflandre (1950a); and (c)
the diameter of the specimen is relatively larger than those of re-
lated forms frequently referred to as Distephanus speculum.
As shown previously (Lemmermann, 1901b; Ling, 1971), in
lateral view, Mesocena circulus var. apiculata has two series of di-
vergent radial spines on the exterior side of the basal body ring.
Thus, should the disappearance of the apical ring as claimed by
Frenguelli or fusion of the apical ring according to Deflandre ac-
tually be the case, at least one series of the radial spines should point
either inwardly or perpendicular to the basal body ring on apical
view, and so far this is definitely not the case, Thus perhaps it is
advisable to limit the present species to the form that is close to
Ehrenberg’s original figure or only to the complete specimen of
Frenguelli (1940, fig. 8a). It is possible that Mandra found the
present species during his California study, but his figures are not
like the present species and, therefore, they are excluded.
Lemmermann (1901b, 1903) illustrated a form, but without
mentioning whether it is a fossil or modern form. Jorgensen (1899)
recorded the present species from plankton samples of the west
coast of Norway but without illustration. Gemeinhardt (1931b)
referred to a single specimen from the Atlantic Ocean (0°18’N,
16°1YW), but as the basal diameter is approximately only 25 un,
(less than one-half that of the specimen found during the present
study — about 60 » here), and as Gemeinhardt himself illustrated
in the same paper that there are wide variations in the number
of spines of Distephanus speculum in the Atlantic, it is concluded
that Gemeinhardt’s specimen is related closely to Distephanus specu-
lum; therefore it is excluded from the present species.
192 BuLLETIN 273
Dimensions. — Diameter of basal body ring, 60-66 yp; of radial
spine, 4-8 pz.
Known geologic range.— Miocene. Glezer (1966, p. 277)
described the Russian occurrence as Neogene.
Occurrence. — P. polyactis thus recognized is found, during the
present study, only in one sample, EM 8-15, 36-37 cm, which is
Luisian, according to Parker (1964), Bandy and Ingle (1970) or
Helvetian according to Martini and Bramlette (1963).
Genus ROCELLA Hanna, 1930
Remarks. — The present genus was proposed by Hanna (1930)
with Rocella gemma as the type by monotypy, and until recently,
this unusual siliceous microfossil has not been recovered.
The exact nature or affinity of these remains is still uncertain.
Hanna (1930) indicated that he was in favor of placing them within
or closely related to silicoflagellates over the wheel-like plate of
holothuridae, radiolarians, sponges, and diatoms, or the possi-
bility of calcite replaced by amorphous silica. Recently Lipps
(1970, p. 981) suggested that they were small valves of a diatom
from which the silica had been dissolved from the “pores.” The
present author has studied deep-sea sediments from various parts
of the world including the equatorial Pacific, but until now diatoms
indicating such a possibility have never been encountered.
Therefore, despite the nature of the genus still remaining un-
known, there is a good possibility of its being considered a bio-
stratigraphic index for deep-sea sediments, and thus it is included
in the present study.
Rocella gemma Hanna Pl. 31, figs. 12-15
Rocella gemma Hanna, 1930, Jour. Paleont., vol. 4, No. 4, pp. 415-416, pl.
40, figs. 8-18.
Remarks. — The specimens found during the present study
agree with the original description and illustration given by Hanna
from the San Gregorio Formation (possible Helvetian) near Purris-
sima, California, which Loeblich, III (et al., 1968) regarded as Baja
California, Mexico.
The present species was recovered from Core 17 of the DSDP
Hole 34 at Lat, 39°28.21’N.; Long. 127°16.54’W., and its age 1s con-
SILICOFLAGELLATES AND EBRIDIANS: LING 193
sidered as “Coccolithus bisectus — Triquetrorhabdulus carinatus
Subzone;” but whether the sample is autochthonous is questioned
(McManus, et al., 1970, pp. 114-115). Core 16 sample from the same
hole, which is regarded as the same age as that of Core 17, was
identified by Bukry and Bramlette (1970, p. 490) as upper Oligo-
cene to lower Miocene from its nannofossil assemblage.
Dimensions. — Diameter, 50-70 p.
Known geologic range. — Miocene.
Occurrence. — The present species is found abundantly in sam-
ples of MSN 149P, 294-296 cm, and rarely in JYN V 16P, 231-233
cm, and these samples are considered here as the base of the Mio-
cene agreeing with the above observation. Therefore, it is considered
that the above DSDP specimens are likely not reworked.
Genus SEPTAMESOCENA Bachmann, 1970b
Septamesocena apiculata (Deflandre) Pl. 29, figs. 11-15
Not Mesocena oamaruensis var. apiculata Schulz, 1928, Bot. Archiv., vol.
21, No. 2, p. 240, fig. 11.
Mesocena polymorpha var. triangula Lemmermann, Gemeinhardt, 1930
(part), iz Kryptogamen-Flora, vol. 10, pt. 2, p. 28.
Mesocena polymorpha var. triangula fa. glabra Gemeinhardt, 1930 (not
Schulz), in Kryptogamen-Flora, vol. 10, pt. 2, p. 28, fig. 12b (only).
Not Mesocena apiculata (Schulz), Hanna, 1931, Mining in California, vol.
Zi Nowe2 sp 198; ple Dy tics 3 ink
Mesocena apiculata (Schulz), Deflandre, 1932c, Soc. Bot. France, Bull., vol.
79, Nos. 5-6, p. 499, figs. 34 (?), 35.
Mesocena apiculata (Schulz), Tynan, 1957, Micropaleont., vol. 3, No. 2, p.
134, pl. 1, fig. 10.
Mesocena apiculata (Schulz), Stradner, 1961, Erdél u. Kohle, vol. 14, No.
7 vs OSE eeh AE AS ey
Mesocena apiculata (Schulz), Bachmann, 1962, Verh. Geol. B.-A., Jahrg.
1962, No. 2, p. 379, figs. 10, 11.
Mesocena apiculat (Schulz), Bachmann, 1963, in Bachmann, ef al., Geol.,
Ges. Wien, Mitt., vol. 56, No. 1, p. 152, pl. 19, figs. 60-62.
Mesocena oamaruensis Schulz, Tsumura, 1963, Yokohama Munic. Univ.,
Jour., ser. C-45, No. 146, p. 42, pl. 7, fig. 2; pl. 21, fig. 12.
Mesocena apiculata (Schulz), Bachmann, in Ichikawa, ef al., 1964, Kan-
azawa Univ., Sci. Rept., vol. 9, No. 1, p. 107, pl. 4, fig. 36.
Mesocena oamaruensis Schulz, Bachmann, in Ichikawa, ef al., 1964, Kan-
azawa Univ., Sci. Rept., vol. 9, No. 1, pp. 107-108, pl. 4, fig. 37.
Mesocena aff. apiculata (Schulz), Glezer, 1966, in Cryptogamic plants of
the U.S:SIR-: vols 7;ips282, pl28, fig. 54(2))3 pl> 29, fig. 9:
Mesocena apiculata (Schulz), Bachmann and Papp, 1968, Giorn. di Geol.,
ser: 2, vol. -35, ‘pl. 3, figs: 1, 2:
Mesocena apiculata (Schulz), Hajos, 1968, Geol. Hungarica, ser. Paleont.,
fase: 3/7, sp. (67,1 pla 4a tis. 3?
Mesocena apiculata (Schulz), Bachmann, 1970a, Verh. Geol. B.-A., Jahrg.
1970, No. 2, pp. 280, 282, 284, pl. 5, figs. 10-16; pl. 6, figs. 1-5.
Septamesocena apiculata (Schulz), Bachmann, 1970b, in Cat. Fossil. Aus-
triae, No. 1b, pp. 13-14.
194 BULLETIN 273
Remarks. —In 1928, Schulz, recognized a silicoflagellate form
with a smooth basal body ring but without septa from Hungarian
sediments and established a M. polymorpha var. triangula fa. glabra.
Gemeinhardt (1930) found the Schulz’s forma unnecessary, but in
his figure 12 b, he illustrated the presence of septa. The above syn-
onymy is prepared after close examination of the figures illustrated
by previous students of silicoflagellates. The new genus, Septameso-
cena, was proposed by Bachmann for the Mesocena with a septal
structure, Stradner (1961) referred the present species to the form
described originally by Schulz as Mesocena oamaruensts var. apicu-
lata from Oamaru, New Zealand, and such practice was followed by
Deflandre, Hanna, and others and until as recently as the early part
of 1970 by Bachmann, The samples from the Oamaru Diatomite
are currently under detailed examination; however, as already
shown here (PI. 28, figs. 2-4) for comparison, so far specimens from
Oamaru which correspond with Schulz’s original figure, have no
septa.
Glezer recorded specimens ranging from early Paleocene to late
Eocene in Russia. The early Paleocene specimen from the eastern
slope of the Urals has no septal structure (pl. 28, fig. 5); the late
Eocene form (pl. 29, fig. 9) from the Ural-Caspian area possess
discontinuous lumen and can be interpreted as being due to septal
structure, but it has a dentate basal body ring.
Dimensions. — Length of basal body ring, 42-58 p; of radial
spines, 2-3 p.
Known geologic range. — Eocene (late) to Miocene.
Occurrence. — The present species is found only in DSDP
13-3-1, 120-121 cm and Italian outcrop samples. Examination of
more Tertiary California samples is required before worldwide
distribution of the present species can be discussed in more detail.
Genus VALLACERTA Hanna, 1928
Vallacerta hortoni Hanna Pl..32) figssslees
Vallacerta hortoni Hanna, 1928, Jour. Paleont., vol. 1, No. 4, pp. 262-263,
pl. 41, figs. 7-11.
Dictyocha siderea Schulz, 1928, Bot. Archiv., vol. 21, No. 2, p. 284, fig.
Sillaeb:
Dictyocha siderea Schulz, Gemeinhardt, 1930, in Krytogamen-Flora, vol. 10,
pt2, pe 56, fey 47.
Vallacerta hortoni Hanna, Deflandre, 1940a, Acad. Sci. Paris, C. R., vol.
211, No. 19, p. 446, figs. 1, 5.
SILICOFLAGELLATES AND EBRIDIANS: LING 195
Vallacerta hortoni Hanna, Deflandre, 1940c, Acad. Sci. Paris, C. R., vol.
Diie Now 235 ps 5985) tgs Le
Vallacerta hortoni Hanna, Deflandre, 1950a, Microscopie, vol. 2, pp. 57/82-
58/82, figs. 144-149.
Dictyocha siderea Schulz, Tsumura, 1959, Yokohama Munic. Univ., Bull.,
vol. 11, Nat. Sci., vol. 1, p. 66, pl. 4, fig. 52 (fide Tsumura, 1963).
Dictyocha siderea Schulz, Tsumura, 1963, Yokohama Munic. Univ., Jour.,
ser. C-45, No. 146, p. 59, pl. 11, fig. 12.
Vallacerta hortoni Hanna, Glezer, 1966, in Crytogamic plants of the U.S.S.R.,
voll 70p.\ 219, pli, fighacs plisiy fig. si:
Vallacerta hortoni Hanna, Mandra, 1968, California Acad. Sci., Proc., ser.
4, vol. 36, No. 9;-p.'267, tgs. 11, 12; 19:
Remarks. — A literature survey reveals that Hanna’s article
was published in January, while Schulz’s article was dated in June
of the same year; Hanna’s name, therefore, has priority,
Although Schulz (1928) also proposed a quadrangular form as
Dictyocha siderea var. quadrata from Prussia, no such variety is
found in the topotypic sample, CAS 1144, from California.
Dimensions. — Diameter of pentagonal plate, 35-45 yw; length
of spines, 5-8 yp.
Known geologic range.—Cretaceous (late) to Paleocene
(early). The Paleocene occurrence is reported from Russia by
Glezer (1966).
Occurrence. — The present species is recovered only from the
topotypic material from California. Mandra (1968, p. 242) indicated
that the present species (together with Lyramula furcala) is found
only from Upper Cretaceous.
Ebridians
Terminology. — Morphologic terms used herein are those com-
monly employed and summarized by Deflandre (1951) and are
shown in Text-figure 6.
Order STEREOTESTALES Lemmermann, 1901a
Genus AMMODOCHIUM Hovasse, 1932
Ammodochium rectangulare (Schulz) Pl. 32, figs. 3-5
Remarks. — For the synonymy, refer to Ling (1971).
Known geologic range. — Paleocene to Miocene.
Dimensions. — Length, 18-24 up.
Occurrence. — From the samples studied, this species is found
from WR TR 11, Low (Bo. 202), Dorcadosypris alata radiolarian
196 BULLETIN 273
Zone (Riedel and Sanfilippo, 1970) to EM 6-2, 10-11 cm, late Mio-
cene.
Genus EBRIOPSIS Hovasse, 1932a
Ebriopsis antiqua (Schulz) Pl. 32, figs. 6-12
Remarks. — For the synonymy of the present species, also refer
to Ling (1971). Since the present species was first reported by
Schulz, wide range of variation has been recognized by earlier in-
vestigators: surface ornamentation of skeletal elements is smooth
or crenulate; with or without anterior and posterior axial spines,
“corne axiale anterieure” and “corne axiale posterieure” of Deflandre
(1951) (see Text-figure 6B).
A B
Text-figure 6.— Terminology of ebridian skeleton. A. Ammodochium
rectangulare, B. Ebriopsis antigua; a = actine, ca = anterior axial spine,
cp = posterior axial spine, o — opisthoclade, p = proclade, r = rhabde,
sa = anterior synclade, sp = posterior synclade, t = triaene, T = triode
(Modified after Deflandre, 1951).
Deflandre (1950b) noticed that in Paleocene Ebriopsis valida,
both the rhabde and one of the opisthoclades are on the same plane
of symmetry, but in Eocene form, E. mesnilt (=E. crenuata in
this paper, see below), there is an angle between the two skeletal
elements, beginning from 12° to between 35° to 50° in the late
Eocene (not Oligocene) Oamaru samples from New Zealand. At
the time of Miocene form, FE. antiqua, the opisthoclade 02 is ap-
proximately on the same plane with one of the actines, Deflandre,
SILICOFLAGELLATES AND EBRIDIANS: LING 197
therefore, proposed the phylogenic lineage of E. valida - E. mes-
nit - E. antiqua.
Although no samples nor specimens from Paleocene sediments
are available for the present study to confirm Deflandre’s observa-
tion, lineage between FE. crenulata - E. antiqua seems agreeable.
In addition to his observation, the following are obvious from the
present investigation:
1. Surface ornamentation is strongly coarse and crenulate in
the middle Eocene specimens of FE. crenulata, and also the skeletal
elements are generally broader than those from the higher horizon.
2. A gradual size increase is noticed from specimens of middle
Eocene to Miocene form.
3. Although forms without anterior and posterior axial spines
are found from middle Eocene samples, the specimens with these
spines occur mostly throughout Miocene and also possibly late
Eocene, but the specimens with such well-developed spines are
rather commonly found in Miocene samples and are also reported
from various parts of the world.
Dimensions. — Diameter, 28-40 p.
Known geologic range. — Paleocene to Miocene.
Occurrence. — The earliest occurrence of this species in the
present study is from DSDP 6-6-2, 1-2 cm, and the latest is found
in EM 6-2, 10-11 cm which is considered upper Miocene by Mar-
tint and Bramlette (1963) or Delmontian by Bandy and Ingle
(1970).
Ebriopsis crenulata Hovasse Pl. 32, figs. 13-18
Podamphora mesnili Deflandre, 1932b, Acad. Sci. Paris, C. R., vol. 194,
No. 24, p. 2172, figs. 6, 7. nom. nud.
Ebriopsis crenulata Hovasse, 1932b, Soc. Zool. France, Bull., vol. 57, No. 4,
p. 281, fig. 4 I, Il.
Podamphora mesnili Deflandre, Deflandre, 1933, Soc. Zool. France, Bull.,
ViOls 517, NOmGsIpp A 516s O19 e522, tiess 11266 113)
Ebriopsis mesnili (Deflandre), Deflandre, 1950b, Acad. Sci. Paris, C. R.,
vol. 230, No. 19, p. 1684, figs. 3-7.
Ebriopsis mesnili (Deflandre), Deflandre, 1951, Biol. France, Belgique.,
Bull., vol. 85, fasc. 1, p. 64, figs. 23-29, 50-54.
Ebriopsis mesnili (Deflandre), Hajos, 1968, Geol. Hungarica, ser. Paleont.,
fascsa7, pawl eples 60 tigw £2:
Remarks. — As clearly discussed by Loeblich, III (et al., 1968,
p. 51), the name Podamphora mesnili appeared in 1932b, but was
a nomen nudum and only validated by a later article dated Decem-
198 BULLETIN 273
ber 13 of the same year, but actual date of publication for the
particular issue was February 15, 1933. However, Hovasse in the
meantime published Ebriopsis crenulata on June 14, 1932, and,
therefore, has priority over Deflandre’s name.
The specimen of this species is characterized by a smaller size
than the preceding species, coarsely crenulated surface ornamenta-
tion and broader skeletal parts. No anterior or posterior axial
spines are found.
Dimensions. — Diameter, 24-32 »
Known geologic range. — Eocene.
Occurrence. —Hovasse described the present species from
Oamaru, New Zealand, and one specimen with a lorica. A specimen
from Jackson’s Paddock section, $136/914, of the Oamaru Dia-
tomite, New Zealand, is here illustrated as a reference (PI. 32,
figs. 13, 14).
Genus PARATHRANIUM Hovasse, 1932c
Remarks.— The present genus was proposed by Hovasse
(1932c, p. 465) with P. tenuipes as the type species. Loeblich, III
(1965, p. 189) discussed Dicladia (Ehrenberg, 1844a, p, 73), and
designated D. ? clathrata (Ehrenberg, 1944a, pp. 70, 79; 1854, pl.
18, fig. 100) as the type, and by so doing indicated that Dicladia
has priority over Parathranium. This is later followed by Loeblich,
III (et al., 1968, pp. 32, 62). Recently, Deflandre (1971) rejected
these opinions.
It should be pointed out here, however, that Ehrenberg’s speci-
men from Richmond, Virginia, illustrated by him shows only two
opisthoclades instead of three, and furthermore the exact nature
for the portion of the proclades is not clear. Further study, particu-
larly from the sample in the vicinity of Richmond, Virginia, is neces-
Sary in order to recognize the characters of Dicladia ? clathrata;
thus at the present time Parathranium Hovasse is used.
Parathranium tenuipes (Hovasse) Pli32, figs 219-25
Thranium tenuipes Hovasse, 1932a, Soc. Zool. France, Bull., vol. 57, No. 2,
p. 123, fig. 5.
Thranium tenuipes Hovasse, Deflandre, 1932d, Soc. Zool. France, Bull., vol.
57, No. 4, pp. 309-311, figs. 31-33.
Parathranium tenuipes (Hovasse), Hovasse, 1932c, Soc. Zool. France, Bull.,
vol. 57, No. 2, pp. 465-466.
SILICOFLAGELLATES AND EBRIDIANS: LING 199
Parathranium clathratum Deflandre, 1951 (not Ehrenberg), Biol. France,
Belgique., Bull., vol. 85, fasc. 1, p. 77, figs. 5, 172.
Parathranium clathratum Deflandre (not Ehrenberg), Hajos, 1968, Geol.
Hungarica, ser. Paleont., fasc. 37, p. 72, pl. 6, figs. 16, 17.
Remarks. — The Californian specimens agree well with the
previous records.
Dimensions. — Length of opisthoclade, 18-22 uy.
Known geologic range. — Miocene.
Occurrence. — The present species was recovered only from
Mohole samples during the present study and ranges from EM 7-1,
14-15 cm, N.12 planktonic foraminiferal Zone (Riedel and San-
filippo, 1970) to EM 8-9, 148-149 cm; thus it encompasses most of
the Luisian and all the Mohnian stages by Parker (1964), Bandy
and Ingle (1970), and Kanaya (1971).
DISCUSSION
(a) General
Having examined in detail the numerous silicoflagellates and
ebridian taxa, it is now possible to investigate from the samples
studied, their occurrences since Maestrichtian time.
The stratigraphic occurrences of the taxa are shown in Text-
figures 2, 3, and 4 with the inclusion of the 28 experimental Mohole
samples. The sample coverage throughout the Neogene section is
considered adequate, strongly contrasting to the Paleogene interval,
where available samples examined during the present study were
clustered in a few stratigraphic age intervals and hence are admit-
tedly inadequate for establishing a definite zonation. However, the
following general statement can be made in ascending order:
Lyramula furcula - Vallacerta hortont assemblage: At least a
part of the Maestrichtian or uppermost Cretaceous section repre-
sented by sample CAS 1144 is characterized by these unique silico-
flagellates. DSDP samples 4-4-1, 13A-2-1, 13A-4-1 and 13A-5-1 are
of Cretaceous age (Ewing, et al., 1969; Maxwell, et al., 1970) but
were unfortunately barren of these siliceous microfossils. Neither
the lower nor upper limit of this assemblage, therefore, can be de-
termined from the present study.
Naviculopsis foliacea assemblage: The middle Eocene interval,
particularly its lower part, is characterized by the presence of
various forms of Navicwopsis and Corbisema species, Occurrence of
200 BULLETIN 273
Ebriopsis crenulata is also limited in this assemblage in the present
study. Although the upper and lower limit of these taxa found in
this interval cannot be decided at this time, preliminary examina-
tion of Oamaru Diatomite (Runangan Stage, NP.17 - NP.18 of
calcareous nannoplankton Zones) and Kellogg Shale and Sidney
Shale samples indicate that they would extend their occurrences to
the upper Eocene and, therefore, sample WR TR 39F (1068), which
is considered slightly below the Oligo-Eocene boundary, may indi-
cate the highest occurrence for these forms. There is a good possi-
bility that the interval could be subdivided in the future.
Rocella horizon: Occurrence of Rocella gemma at the base of
the Miocene, in this paper, or possibly the top of the Oligocene,
according to others, may be a good time marker.
As discussed earlier, it was only last year that R. gemma was
found for the second time since the species was proposed, and for
the first time it was found from deep-sea sediments, Core 17 of the
Deep Sea Drilling Project Hole 34 near California. The sample
was interpreted as upper Oligocene to lower Miocene. However,
whether the sample was of autochthonous origin was questioned at
that time. The present finding from samples JYN V, 16 P, 231-233
cm, and MSN 149 P, 294-296 cm, agrees with the above strati-
graphic occurrence, and, therefore, this species may be a reliable
time indicator.
From this horizon to the next assemblage, several known but
long-range silicoflagellates make their first appearance, such as
Distephanus crux, D. speculum, and Dictyocha ausonia, but further
investigation is needed to establish the assemblage.
Dictyocha fibula var. octagona assemblage: The interval is de-
fined by the range of D. f. var. octagona. The initial appearance of
so-called Mesocena elliptica, Cannopilus hemisphaericus, and Am-
modochium rectangulare is slightly below the base of assemblage,
while the top of assemblage is marked by the latest occurrence of
Distephanus crux var. octacanthus. Within this interval Dictyocha
sp., Ebriopsis antiqua (with spine form), and Mesocena circulus
var. apiculata appear.
From the upper part of this assemblage to the uppermost Mio-
cene, numerous experimental Mohole samples were studied as shown
in Text-figure 4. Dictyocha fibula and Parathranium tenuipes show
SILICOFLAGELLATES AND EBRIDIANS: LING 201
initial appearance at the top of this assemblage. As indicated earlier,
judging from the relationship with other microfossils, particularly
with the radiolarians, this is the interval that Martini (1971a, b)
recognized as his Dictyocha octacantha (Distephanus crux var.
octacanthus in this paper) horizon from Deep Sea Drilling Project
Leg VII of the western equatorial Pacific.
Cannopilus sphaericus assemblage: The top of the underlying
Dictyocha fibula var. octagona assemblage to the latest occurrence
of Cannopilus sphaericus defines the assemblage. The base of the
assemblage is also marked by the earliest occurrence of Distephanus
crux var. longispina, so-called “Mesocena diodon’”, and also coincides
with the initial occurrence of the radiolarian species, Cannartus
(2) petterssoni, according to Sanfilippo and Riedel (1970). The
top of the assemblage is also the highest occurrence of Corbisema
triacantha, C. t. var. minor; therefore, it is conceivable that the
upper limit of Corbisema triacantha Zone of Martini is here. Oc-
currence of Paradictyocha polyactis also falls within this assemblage.
The boundary between this and overlying assemblages seems
to agree well with the Helvetian? — Tortonian? boundary of Mar-
tini and Bramlette (1963), and the Luisian — Mohnian boundary
of Parker (1964) and Bandy and Ingle (1970).
Distephanus crux var, longispina assemblage: The interval from
above the underlying Cannopilus sphaericus assemblage to the last
appearance of Distephanus crux var. longispina is considered as the
present assemblage. The top thus defined coincides also with the
latest occurrence of Cannopilus hemisphaericus and likely also with
that of Mesocena circulus. The limited appearance of Distephanus
speculum var. pentagonus, and the highest stratigraphic occurrence
of “Mesocena elliptica” in Mohole samples are also found within
this assemblage. The top of this assemblage coincides with the top
of middle Miocene of Martini and Bramlette (1963) and is ap-
proximately in agreement with the top of the Mohnian stage of
Parker (1964) and the lower and upper Mohnian boundary of
Bandy and Ingle (1970).
Distephanus speculum var. pseudofibula assemblage: The as-
semblage is defined by the range of Distephanus speculum var.
pseudofibula in the Mohole section which also coincides with the
interval from the top of the underlying previous assemblage to the
202 BuLLeTIn 273
highest occurrence of the silicoflagellates and ebridians in the ex-
perimental Mohole samples. Within this interval, limited occur-
rence of Distephanus speculum var. pseudocrux, last occurrence of
Dictyocha ausonia, Ammodochium rectangulare, “Mesocena diodon”,
and Parathranium tenuipes are observed, and they may mark the
Mohnian-Delmontian boundary according to the study made by
Bandy and Ingle (1970), and Kanaya (1971).
The highest sample bearing silicoflagellates and ebridians from
Mohole, EM 6-2, 10-11 cm, is recognized as Delmontian by Bandy
and Ingle (op. cit.), and Kanaya (op. cit.) and is apparently in
agreement with late Miocene age of Martini and Bramlette (1963).
It is likely that the present assemblage is correlative with the
Dictyocha pseudofibula horizon of Martini (197la, b) from the
western equatorial Pacific.
Above this assemblage, throughout the Pliocene to part of the
Pleistocene interval, silicoflagellates are found sporadically during
the present study, and no distinct assemblage can be recognized
until the next and the last assemblage,
Dictyocha fibula var. aculeata assemblage: The beginning of
modern forms, such as Dictyocha fibula var. aculeata, D. fibula var.
messanensis provides the last and the latest assemblage recognized
during the present study.
Within this assemblage, one horizon can be recognized by the
recurrence of Mesocena cf. elliptica, which brackets the Jaramillo
Event of Matuyama Reversed Epoch, about 0.79 million years in the
eastern equatorial Pacific (Hays, et al., 1969); a similar occurrence
is recognized by Jousé (1969) and Muhina (1969). Ling (1970)
also observed such an occurrence from the central north Pacific
sediments and also added that the latest occurrence of Dictyocha
subarctios and D. cf. ausonia is found at the same horizon.
(b) Mediterranean (Italy) region
The investigation thus far carried out on silicoflagellates and
ebridians from deep-sea sediments of the Pacific and Atlantic oceans
and some land outcrops demonstrates that there are much diversi-
fied taxa showing limited geological occurrence since late Cretaceous
time in the low-latitude area. Reports from the midlatitude of the
western Pacific (Ling, 1971) and the central north Pacific bottom
sediments (Ling, 1970) also agree with such findings.
SILICOFLAGELLATES AND EBRIDIANS: LING 203
Previously, workers such as Ehrenberg, Zanon, and Deflandre
recorded the occurrence of these siliceous microfossils from the Medi-
terranean region, but stratigraphic information about these samples
was often vague, and the exact range of the taxa cannot be deduced
from the existing data.
Samples from Italian localities were here examined in order
to seek the stratigraphic occurrence of taxa in Italy, to compare the
results with those from the present deep-sea sediments of low lati-
tude, and at the same time to cross-correlate with the radiolarian
zones from these samples recognized by Riedel and Sanfilippo
(1971) and Sanfilippo (1971).
The results of the present stratigraphic occurrence are sum-
marized in Text-figure 7, and the following microfloral boundaries
are noted in ascending order:
1. The earliest occurrence of Naviculopsis lata, together with
that of Distephanus speculum and Corbisema triacantha, marks the
boundary between samples WRE 67- (in the following discussion
the prefix WRE 67- will not be repeated) 107 and 112, The boun-
dary of radiolarian zones, Lychnocanium bipes below and Calo-
cycletta virginis above, is drawn between 106 and 107 and thus is
slightly below the present silicoflagellate boundary.
2. The first appearance of Mesocena cf. elliptica and Navt-
culopsis quadrata occurs between samples 95 and 78. The boundary
also marks the highest occurrence of Naviculopsis regularis and
Corbisema triacantha and C. triacantha var. flexuosa. This boundary
thus seems to agree with that of radiolarian zones, Calocycletta
virginis below and C. costata above.
Although minor microfloral changes are recognized between the
interval of samples 112 and 95, and of 78 and 76, the Italian sam-
ples thus can be grouped into three assemblages and thus boundaries
are generally in agreement with the radiolarian results.
Perhaps the most interesting and significant phenomenon found
in the Italian samples in comparison with the low- and midlatitudes
of Pacific and Atlantic deep-sea sediments is the abundance of
Naviculopsis specimens throughout the Italian samples here exam-
204 BULLETIN 273
ined. Apparently some taxa are limited in their biogeographic distri-
bution to the Mediterranean region and are within a part of the
European continent only. However their value as a stratigraphic
index is promising for the European Tertiary section.
M. elliptica)
SILICOFLAGELLATES
FROM ITALY
Mesocena cf elliptica (
Naviculopsis lata
Dictyocha ausonia
Noviculopsis quadrata
Cannopilus picasso/
Distephanus crux
| evicutopsis reguiaris
=a
8
RADIOLARIAN 3
ZONES S
(Sanfilippo, é
1971)
Ez Septamesocena apiculatu
WRE 67-76 EA
Calocycletta “77 Li
costara
gall -78 i
-95 i
eal
Calocycletra
virginis
VERY RARE
RARE
COMMON
a
bipes
Text-figure 7.— Distribution of silicoflagellates from Neogene Mediter-
ranean (Italy) region. For detail study of the samples, see Sanfilippo (1971).
SILICOFLAGELLATES AND EBRIDIANS: LING 205
REFERENCES
ann, A. 4
pie co Eine neue Mesocena-Art (Silicoflagellidae) aus dem Kalifornischen
Eozan. Verh. Geol. [Bundesanstalt] B.-A. Wien, Jahrb. 1962, No.
2, pp. 378-383, pl. 1.
1963. Silicoflagellidae. In Bachmann, A. et al., Mikropaldontologische
Studien im “Badener Tegel” von Frattingsdorf N. O. Geol. Ges.,
Wien, Mitt., vol. 56, No. 1, pp. 145-155, pls. 16-22, figs. 1, 2.
1964. Part. II, Silicoflagellidae und Archaeomonadaceae. In Ichikawa,
W. et al., Fossil diatoms, pollen grains and spores, silicoflagellates
and archaeomonads in the Miocene Hojuji diatomaceous mudstone,
Noto Peninsula, Central Japan. Kanazawa Univ., Sci. Rept., vol.
9, No. 1, pp. 87-118, pls. 1-7, figs. 1-20.
1967. Part II. Silicoflagellidae im lida Diatomit. In Ichikawa, W. et al.,
Fossil silicoflagellates and their associated uncertain forms in lida
Diatomite, Noto Peninsula, Kanazawa Univ., Sci. Rept., vol. 12,
No. 1, pp. 149-172, pls. 2-9.
1970a. Silicoflagellaten aus dem oberésterreichischen Egerien (Oberoli-
gozan). Verh. Geol. B.-A., Jahrg. 1970, No. 2, pp. 275-305, pls. 1-7.
1970b. Flagellata (Silicoflagellata). In Catalogus Fossilium Austriae, Ed.
by H. Zapfe, No. 1b, pp. 1-28.
Bachmann, A., and Ichikawa, W.
1962. The silicoflagellides in the Wakura Beds, Nanao City, Prefecture
Ishikawa, Japan. Kanazawa Univ., Sci. Rept., vol. 8, No. 1, pp.
161-175, pls. 1-10, figs. 1-4.
Bachmann, A., and Papp, A.
1968. Vorkommen und Verbreitung der Silicoflagellaten im Neogen
Osterreichs. Comm, Mediterranean Neogene Stratigraphy, Proc.,
4th Sess., Bologna 1967, Giornale di Geologie, ser. 2, vol. 35,
fasc. 2, pp. 117-126, pl. 3.
Bailey, J. W.
1845. Notice of some new localities of Infusoria, fossil and Recent. Amer.
Jour, Sci. Arts, vol. 48, No. 2, pp. 321-343, pl. 4.
Bandy, O. L., and Ingle, J. C., Jr.
1970. Neogene planktonic events and radiometric scale, California. In
Radiometric dating and paleontologic zonation, Ed. by O. L. Bandy,
Geol. Soc. Amer., Spec. Paper 124, pp. 131-172.
Berggren, W. A.
1969. Cenozoic chronostratigraphy, planktonic foraminiferal zonation
and the radiometric time scale. Nature, vol. 224, No. 5224, pp.
1072-1075.
Blow, W. H.
1970. Deep Sea Drilling Project, Leg. 2, Foraminifera from selected
samples. In Peterson, M. N. A., et al., Initial Reports of the
Deep Sea Drilling Project, U.S. Government Printing Office,
Washington, vol. 2, pp. 357-365.
Bolli, H. M.
1957. Planktonic foraminifera from the Oligocene-Miocene Cipero and
Lengua formations of Trinidad, B.W.1I. U.S. Nat. Mus., Bull.,
No. 215, pp 97-123, pls. 22-29.
Bukry, D., and Bramlette, M. N.
1970. Deep Sea Drilling Project, Leg 5, Coccoliths age determinations.
In McManus, D. A., et al., Initial Reports of the Deep Sea Drilling
Project, U.S. Government Printing Office, Washington, vol. 5,
pp. 487-494.
206 BULLETIN 273
Carnevale, P.
1908. Radiolarie e silicoflagellati de Bergonzano (Reggio Emilia). R.
Ist. Veneto Sci. Lett. Arti. Mem., vol. 28, No. 3, pp. 1-46, pls. 1-4.
Clark, B. L., and Campbell, A. S.
1945. Radiolaria from the Kreyenhagen Formation near Los Banos,
California. Geol. Soc. Amer., Mem. 10, pp. 1-66, pls. 1-7.
Colom, G.
1940. Arqueomonadineas, Silicoflagelados, Discoasteridos fosiles de
Espana. Las Ciencias, vol. 5, pp. 343-356, figs. 1-11.
Deflandre, G.
1932a. Les Silicoflagellés des terres fossiles a Diatomées. Soc. France
Microsc., Bull., vol. 1, No. 1, pp. 10-20.
1932b. Sur le genre Podamphora et ses relations avec les Ebriacées-
Acad. Sci. Paris, C. R., vol. 194, No. 24, pp. 2171-2173, figs. 1-8.
1932c. Sur la systématique des Silicoflagellés. Soc. Bot. France, Bull., vol.
79, Nos. 5-6, pp. 494-506, figs. 1-42.
1932d. Remarques sur quelques ebriacées. Soc. Zool. France, Bull., vol.
57, No. 4, pp. 302-315, figs. 1-40.
1933. Enkystement et stade loriqué chez les Ebriacées. Soc. Zool. France,
Bull., vol. 57, No. 6, pp. 514-523, figs. 1-13.
1935. Sur un Silicoflagelle aberrant, Dictyocha recta (Schulz), a
squelette non tubulaire. Soc. France, Microsc., Bull., vol. 3, No.
4, pp. 115-118, figs. 1-5.
1938. Sur deux microfossiles siliceux énigmatiques (Silicoflagellidées?).
Soc. Bot. France, Bull., vol. 7, No. 3, pp. 90-96, figs. 1-25.
1940a. Sur les affinités et la phylogenese du genre Vallacerta, Silico-
flagellidée du Crétacé supérieur. Acad. Sci. Paris, C. R., vol. 211,
No. 19, pp. 445-448, figs. 1-8.
1940b. L’origine phylogénétique des Lyramula et l’évolution des Silico-
flagellidées. Acad. Sci. Paris, C. R., vol. 211, No. 21, pp. 508-510,
figs. 1-12. :
1940c. Sur une structure reticulée méconnue du squelette des Silico-
flagellidées. Acad. Sci. Paris, C. R., vol. 211, No. 23, pp. 597-599.
1946. Phyllodictyocha nov. gen. Silicoflagellidées et formes affines du
Miocéne de Hongrie. Soc. Bot. France, Bull., vol. 93, No. 9, pp.
335-337, figs. 1-5.
1950a. Contribution a Pétude des Silicoflagellidés actuels et fossiles.
Microscopie, vol. 2, 82 pp., figs. 1-240.
1950b. Sur l’évolution des Ebriediens. Interprétation du genre Ebriopsis.
Acad. Sci. Paris, C. R., vol. 230, No. 19, pp. 1683-1685, figs. 1-8.
1951. Recherches sur les Ebriediens. Paléobiologie, Evolution, Systéma-
tiqgue. Biol. France, Belgique., Bull., vol. 85, fasc. 1, pp. 1-84, figs.
1-238.
1971. Rejet du genre Dicladia Ehrenberg (Diatomée) introduit fallacie-
usement dans les Ebriédiens. In II Planktonic Conf., Proc., Roma,
1970, Ed. by A. Farinacci, vol. 1, pp. 349-353, pl. 1.
Desikachary, T. V., and Maheshwari, C. L.
1956. Fossil silicoflagellates from Colebrook and Nancoori Islands.
Indian Bot. Soc., Jour., vol. 35, pp. 257-264, figs. 1-19, pl. 13.
Ehrenberg, C. G.
1839. Uber die Bildung der Kreidefelsen und des Kreidemorgels durch
unsichtbare Organismen. K. Akad. Wiss. Berlin, Abh., Jahrg. 1838,
pp. 59-147, pls. 1-4, 1 table.
1840. . . . 274 Blatter von ihm selbst ausgefiihrter Zeichnungen von
ebenso vielen Arten. . .. Verh. K. Preuss. Akad. Wiss. Berlin,
Ber., Jahrg. 1840, pp. 197-219.
1844a. Uber 2 neue Lager von Gebirgsmassen aus Infusorien. Verh. K.
Preuss. Akad. Wiss. Berlin, Ber., Jahrg. 1844, pp. 57-97.
SILICOFLAGELLATES AND EBRIDIANS: LING 207
1844b. Resultate seiner Untersuchungen der ihm von der Siidpolreise des
Captain Ross, so wie von den Herren Schayer und Darwin
zugekommenen Materialien tiber das Verhalten des kleinsten
‘Lebens in den Oceanen und den gréssten bisher zugdnglichen
Tiefen des Weltmeers. Verh. K. Preuss. Akad. Wiss. Berlin, Ber.,
Jahrg. 1844, pp. 182-207.
1844c. Untersuchungen iiber die kleinsten Lebensformen im Quellenlande
des Euphrats und Araxes, so wie tiber eine neuen Formen sehr
reiche marine Tripelbildung von den Bermuda-Inseln vor. Verh.
K. Preuss. Akad. Wiss. Berlin, Ber., Jahrg. 1844, pp. 253-275.
1845. Neue Untersuchungen iiber das kleinste Leben als geologisches
Moment. Verh. K. Preuss. Akad. Wiss. Berlin, Ber., Jahrg. 1845,
pp. 53-87.
1847. Ueber die mikroskopischen kieselschaligen Polycystinen als mdach-
tige Gebirgsmasse von Barbados und iiber das Verhdltniss der
aus mehr als 300 neuen Arten besthenden ganz eigenthiimlichen
Formengruppe jener Felsmasse zu den jetzt lebenden Thieren und
zur Kreidebildung. Verh. K. Preuss. Akad. Wiss. Berlin, Ber.,
Jahrg. 1847, pp. 40-60.
1854. Mikrogeologie. Leopold Voss, Leipzig; pp. 1-374, pls. 1-40.
Ewing, M., Worzel, J. L., Beall, A. O., Berggren, W. A., Bukry, D., Burk,
C.A., Fischer, A. G., and Pessagno, E. A. Jr.
1969. Initial Reports of the Deep Sea Drilling Project. U.S. Government
Printing Office, Washington, vol. 1, i-xx + 1-672 pp.
Frenguelli, J.
1940. Consideraciones sobre los silicoflagelados fésiles. Rev. Mus. La
Plata, n.s., vol. 2, Paleont. No. 7, pp. 37-112, pls. 1-4.
Fuji, N., and Bachmann, A.
1969. Silicoflagellaten in Tijiritkawa-Schichten. Kanazawa Univ., Fac.
Educ., Natural Sci., Bull., No. 18, pp. 79-94.
Gemeinhardt, K.
1930. Silicoflagellatae. In MRabenhorst, L., Kryptogamen-Flora von
Deutschland, Osterreich und der Schweiz. Akademische ie aes
gesellschaft, Leipzig, vol. 10, pt. 2, pp. 1-87, figs. 1-69, pl.
1931a. Organismenformen auf der Grenze zwischen eee und
Flagellaten. Deutsch. Bot. Ges., Ber., vol. 49, No. 2, pp. 103-110, pl.
10.
1931b. Die Silicoflagellaten der Deutschen Siidpolar-Expedition 1901-
1903. Deutsche Siidpolar-Exped., vol. 20 (Zool. No. 12), pp. 221-
258, pls. 42-43.
Glezer, Z. |.
1966. Silicoflagellatophyceae. In Cryptogamic plants of the U.S.S.R., Ed.
by M. M. Gollerbakh, Akad. Nauk. SSSR, Botanicheskii Institut
IM V. A. Komarova. (Translated from Russian by Israel Program
for Scientific Translations Ltd., Jerusalem, 1970), vol. 7, pp. i-iv +
1-330, pls. 1-33, figs. 1-28.
Haeckel, E.
1887. Report on the Radiolaria collected by H.M.S. Challenger during
the years 1873-1876. Rept. Voy. Challenger, Zool., vol. 18, pp.
i-clxxxviii + 1-1803, pls. 1-140.
Hajos, Marta
1968. Madtraalji Miocene Uledekeinek Diatomi (Die Diatomeen der
Miozdnen Ablagerungen des Mdtravorlandes). Geologica Hungar-
ica, ser. Paleontologica, fasc. 37, pp. 1-401, pls. 1-63.
Hanna, G. D.
1928. Silicoflagellates from the Cretaceous of California. Jour. Paleont.,
vol. 1, No. 4, pp. 259-264, pl. 41.
208 BULLETIN 273
1930. A new genus of Silicoflagellates from the Miocene of Lower Cali-
fornia, Jour. Paleont., vol. 4, No. 4, pp. 415-416.
1931. Diatoms and silicoflagellates of the Kreyenhagen Shale. Mining
in California. California Div. Mines Rept., State Mineralogist,
vol. 27, pp. 187-201, pls. A-E.
Hays, J. D., Saito, T., Opdyke, N. D., and Burckle, L. H.
1969. Pliocene-Pleistocene sediments of the equatorial Pacific: Their
paleomagnetic, biostratigraphic and climatic record. Geol. Soc.
Amer., Bull., vol. 80, No. 8, pp. 1481-1513, figs. 1-16, pl. 1.
Hopkins, D. M., et al.
1969. Cretaceous, Tertiary, and Early Pleistocene rocks from the con-
tinental margin in the Bering Sea. Geol. Soc. Amer., Bull., vol. 80,
No. 8, pp. 1471-1480, figs. 1-2, pls. 1-2
Hovasse, R.
1932a. Note préliminaire sur les Ebriacées. Soc. Zool. France, Bull., vol.
57, No 2, pp. 118-131, figs. 1-12.
1932b. Second note sur les Ebriacées. Soc. Zool. France, Bull., vol. 57,
No. 4, pp. 278-283, figs. 1-6.
1932c. Troisiéme note sur les Ebriacées. Soc. Zool. France, Bull., vol. 57,
No. 5, pp. 457-476, figs. 1-27.
Jorgensen, E.
1899. Protophyten und Protozoen im Plankton aus der norwegischen
W estkiiste. Bergens Museums Aaarbog, No. 6 (pt. 2), pp. 1-112,
pls. 1-4, tables 1-1xxxii.
Jousé, A. P.
1962. Stratigraphic and paleogeographic researches in the northwestern
part of the Pacific Ocean. Izd. Inst. Okeanologii, Akad. Nauk SSSR,
Moscow, pp. 1-258, pls. 1-80, tables 1-31.
1963. Problems in stratigraphy and paleogeography of the northern part
a the Pacific Ocean. Okeanologia, Akad. Nauk SSSR, vol. 3, No.
6, pp. 1017-1028, figs. 1-2.
1969. Silicoflagellates in bottom sediments of Pleistocene and late Plio-
cene in the Pacific Ocean. In Micropaleontology and organogenous
sedimentation in the Oceans, Ed. by A. P. Jousé, Nauka, Moscow,
pp. 105-114, figs. 1-4.
1971. Diatoms in Pleistocene sediments from the northern Pacific Ocean.
In The Micropaleontology of Oceans, Ed. by B. M. Funnel] and
W. R. Riedel, Univ. Press, Cambridge, pp. 407-421, pls. 31.1- 31.3,
figs. 31.1-31.7
Kanaya. T.
1971. Some aspects of Pre-Quaternary diatoms in the oceans. In The
Micropaleontology of Oceans, Ed. by B. M. Funnell and W. R.
Riedel, Univ. Press, Cambridge, pp. 545-565, pls. 40.1-40.6.
Kanaya, T. and Koizumi. I.
1966. Interpretation of Diatom thanatocoenoses from the North Pacific
applied to a study of core V20-130 (Studies of a deep-sea core
V20-130, pt. 4) Tohoku Univ., Sci. Repts., ser. 2 (Geol.), vol. 37,
No. 2, pp. 89-130, figs. 1-6
Koizumi, I.
1968. Tertiary diatom flora of Oga Peninsula, Akita Prefecture, North-
east Japan. Tohoku Univ., Sci. Repts., ser. 2 (Geol.), vol. 40, No.
3, pp. 171-240, pls. 32-35, figs 1-15.
Kozlova, O. G.
1964. Diatoms of the Indian and Pacific sectors of the Antarctic. 1z-
datel’stvo, Nauka, Inst. Oceanology, Akad. Nauk SSSR. (Trans-
lated by Israel Program for Scientific Translations, Ltd., Jeru-
salem, 1966), pp. 1-191, pls. 1-6.
SILICOFLAGELLATES AND EBRIDIANS: LING 209
Lanjouw, J. (Chairman), et al.
1966. International Code of Botanical Nomenclature adopted by the
tenth International Botanical Congress, Edinburgh, August 1964.
Kemink en Zoom N. V. Utrecht, Netherland, pp. 1-402.
Lemmermann, E.
1901la. Beitrdge zur Kenntniss der Planktonalgen. Deutsch. Bot. Ges.,
Ber., vol. 19, pp. 85-95.
1901b. Silicoflagellatae. Deutsch. Bot. Ges., Ber., vol. 19, pp. 247-271, pls.
10-11.
1903. Klasse Silicoflagellatae. In Nordisches plankton, Ed. by K. Brandt
and C. Apstein. (Botanischer Teil), No. 21, pp. 25-32, figs. 89-108.
Ling, H. Y.
1970. Silicoflagellates from central north Pacific core sediments. Bull.
Amer. Paleont., vol. 58, No. 259, pp. 85-129, pls. 18-20.
1971. Silicoflagellates and Ebridians ‘from the Shinzan diatomaceous
mudstone member of the Onnagawa Formation (Miocene), North-
east Japan. In 11 Planktonic Conf., Proc., Roma, 1970, Ed. by A.
Farinacci, vol. 2, pp. 689-703, pls. 1, 2, figs. 1-3.
Ling, H. Y., and Anikouchine, W.A.
1967. Some spumellarian Radiolaria from the Java, Philippine, and
Mariana Trenches. Jour. Paleont., vol. 41, No. 6, pp. 1481-1491,
pls. 189-192, text-figs. 1-5.
Lipps, J. H.
1970. Ecology and Evolution of Silicoflagellates. North American Pale-
ont. Convention, Proc., pt. G, pp. 965-993.
Loeblich, A. R., Jr., and Tappan, H.
1968. Fossil Phytoflagellates. Geol. Soc. Amer. Spec. Paper No. 101,
paelZ5:
Loeblich, A. R., III.
1965. Type designations for Dicladia (Ebriaceae), Periptera (Baccillario-
pee aey: Gymnocella, Phaeodactylis and Phaeosphaera (Radio-
laria). Proc. Biol. Soc. Washington, vol. 78, pp. 189-192.
Loeblich, A. R., Ill, Loeblich, L. A., Tappan, H., and Loeblich, A. R., Jr.
1968. Annotated index of fossil and Recent, stlicoflagellates and ebridians
with descriptions and illustrations of validly proposed taxa. Geol.
Soc. Amer, Mem. 106, pp. i-xl7-1-319, pls. 1-53.
McManus, D. A., et al.
1970. Initial Reports of the Deep Sea Drilling Project. U.S. Government
Printing Office, Washington, vol. 5, pp. i-xxi -+ 1-827.
Mandra, Y. T.
1968. Silicoflagellates from the Cretaceous, Eocene, and Miocene of
California, U.S.A. California Acad. Sci., Proc., ser. 4, vol. 36, No.
9, pp. 231-277, figs. 1-83, tables 1-7.
1969. Silicoflagellates: A new tool for the study of Antarctic Tertiary
Climates. Antarctic Jour. U.S., vol. 4, No. 5, pp. 172-174.
Marshall, S. M.
1934. The silicoflagellata and Tintinnoinea. Sci. Rept. Great Barrier
Reef Exped., vol. 4, No. 15, pp. 623-662.
Martini, E.
1970. Standard Paleogene calcareous nannoplankton zonation. Nature,
vol. 226, No. 5244, pp. 560, 561.
1971a. Standard Tertiary and Quaternary calcareous nannoplankton. In
II Planktonic Conf., Proc., Roma, 1970, Ed. by A. Farinacci, vol. 2,
pp. 739-785, pls. 1-4, tables 1-6.
1971b. Neogene Acne: from the equatorial Pacific. In Winterer,
E. L., et al., Initial Reports of the Deep Sea Drilling Project,
US: Government Printing Office, Washington, vol. 7, pp. 1695-
1708, figs. 1-3, tables 1-4.
210 BULLETIN 273
Martini, E., and Bramlette, M. N.
1963. Calcareous nannoplankton from the Experimental Mohole Drill-
ing. Jour. Paleont., vol. 37, No. 4, pp. 845-856.
Martini, E., and Worsley, T. R.
1970. Standard Neogene calcareous nannoplankton zonation. Nature,
vol. 225, No. 5229, pp. 289-290.
Maxwell, A. E., et al.
1970. Initial Reports of the Deep Sea Drilling Project, U.S. Govern-
ment Printing Office, Washington, vol. 3, pp. i-xx + 1-806.
Muhina, V. V.
1963. Biostratigraphic correlation of bottom deposits at station 3802 in
the Equatorial zone of the Pacific. Okeanologiia, vol. 3, No. 5,
pp. 861-869, figs. 1-3.
1966. The boundary between sediments of Quaternary and Tertiary age
in the Pacific (on the basis of diatom analysis). Oceanology, vol.
6, No. 1, pp. 99-110, figs. 1-5.
1969. Biostratigraphy of sediments and some questions of paleogeogra-
phy of the tropical region of the Pacific and Indian Oceans. In
Micropaleontology and organogenous sedimentation in the Oceans,
Ed. by A. P. Jousé, Nauka, Moscow, pp. 52-84, figs. 1-10.
Papenfuss, G. F.
1955. Classification of the algae. In Miller, R. C. (chairman), A cen-
tury of progress in the natural science. California Acad. Sci.,
San Francisco, pp. 115-224.
Parker, F. L.
1964. Foraminifera from the experimental Mohole drilling near Guada-
lupe Island, Mexico. Jour. Paleont., vol. 38, No. 4, pp. 617-636,
pls. 97-102, tables 1-3.
Riedel, W. R.
1957. Radiolaria: a preliminary stratigraphy. Swedish Deep-Sea Exped.,
Rept., vol. 6, fasc. 3, pp. 61-96, pls. 1-4.
1971. Radiolarians from Atlantic Deep-Sea Drilling. In I1 Planktonic
Con., Proc., Roma, 1970, Ed. by A. Farinacci, vol. 2, pp. 1057-
1065, figs. 1-8, tables 1, 2.
Riedel, W. R., and Sanfilippo, Annika
1970. Radiolaria, Leg 4, Deep Sea Drilling Project. In Bader, R. G., et
al., Initial Reports of the Deep Sea Drilling Project, U.S. Govern-
ment Printing Office, Washington, vol. 4, pp. 503-575, pls. 1-15.
1971. Cenozoic Radiolaria from the western tropical Pacific, Leg 7. In
Winterer, E. L., et al., Initial Reports of the Deep Sea Drilling
Project, U.S. Government Printing Office, Washington, vol. 7,
pp. 1529-1672, pls. 1-8, figs. 1-3, tables 1-8.
Riedel, W. R., Ladd, H. S., Tracey, J. I., and Bramlette, M. N.
1961. Preliminary drilling phase of Mohole Project: Summary of coring
operations (Guadalupe Site). Amer. Assoc. Petr. Geol., Bull., vol.
45, No. 11, pp. 1793-1798.
Rust, D.
1888. Beitrage zur Kenntnis der fossilen Radiolarien aus Gesteinen der
Kreide. Palaeontographica, vol. 34, pp. 181-213.
Ruth, J. W.
1971. Late Tertiary diatoms and silicoflagellates of the California coastal
area. Geol. Soc. Amer., Cordilleran section (abstract), p. 188.
Sanfilippo, Annika
1971. Neogene radiolarians of the Mediterranean and western Pacific.
In II Planktonic Conf., Proc., Roma, 1970, Ed. by A. Farianacci,
vol. 2, pp. 1121-1127, figs. 1, 2.
SILICOFLAGELLATES AND EBRIDIANS: LING Abt
Sanfilippo, Annika, and Riedel, W. R.
1970. Post-Eocene “closed” theoperid radiolarians. Micropaleont., vol.
16, No. 4, pp. 446-462, pls. 1-2.
Schulz, P.
1928. Beitrdge zur Kenntnis fossiler und rezenter Silicoflagellaten. Bot.
Archiv., vol. 21, No. 2, pp. 225-292, figs. 1-83.
Sheshukova-Poretzkaya, V. S.
1967. The Neogene marine Diatomaceae of Sakhalin and Kamchatka.
Izd. Leningrad Univ., pp. 1-327, text-figs. 1-8, tables 1-19, pls.
1-50.
Stohr, E.
1880. Die Radiolarienfauna der Tripoli von Grotte Provinz Girgenti
in Sicilen. Palaeontographica, vol. 26, pp. 69-124, pls. 1-7.
Stradner, H.
1956. Uber fossile Silicoflagelliden aus dem Tertiar Osterreichs. Univ.
Wien, Dissertation (unpublished).
1961. Uber fossile Silicoflagelliden und die Méglichkeit ihrer Verwen-
dung in der Erdélstratigraphie. Erdél. u. Kohle, vol. 14, No. 2,
pp. 87-92, pls. 1-3, fig. 1.
Tsumura, K.
1959. Komentariita figuraroj de silikoflageluloj. Yokohama Munic. Univ.,
Bull., vol. 11, Nat. Sci., No. 1, pp. 35-90, pls. 2-4.
1963. A systematic study of silicoflagellatae. Yokohama Munic. Univ.,
Jour., ser. C-45, No. 146, pp. 1-84, pls 1-28.
Tynan, E. J.
1957. Silicoflagellates of the Calvert formation (Miocene) of Maryland.
Micropaleont., vol. 3, No. 2, pp. 127-136, pl. 1, figs. 1-3.
Zanon, D. V.
1934. Silicoflagellate fossili Italiane. Acta Pont. Acad. Sci. Nov. Lincei,
vol. 87, pp. 40-82, figs. 1-3, pl. 1
Zittel, K. A.
1876. Uber einige fossile Radiolarien aus der norddeutschen Kreide.
Deutsche Geol. Ges. Zeitschr., vol. 28, pp. 75-86, pl. 2.
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PLATES
214
Figure
1-5.
6,7.
8-10.
11, 12.
13-17.
18.
BuLweTIn 273
EXPLANATION OF PLATE 23
All figures 500 unless otherwise indicated.
Cannopilus hemisphaericus (Ehrenberg) ........................00000.. 148
1. EM 7-1, 14-15 cm, L-2 (V42/0); 2, 3. Different focus levels,
EM 7-1, 14-15 cm, L-2 (E37/3); 4. EM 7-1, 14-15 cm, L-2
(R27/0); 5. EM 8-12, 94-95 cm, L-2 (018/1).
Cannopilus picassoi Stradmer ....................ccccccecteenereeeenes 149
6. WRE 67-77, R-2 (C43/1); WRE 67-77, L-7 (N27/3).
Cannopilus sphaericus Gemeinhardt .................0...cccees 150
8. EM 7-2, 6-7 cm, L-2 (L18/4); 9. EM 8-15, 505-506 cm,
L-2 (E33/0); 10. EM 8-14, 388-389 cm, L-2, (042/0).
Cannopilus tetraceros Bachmann and Ichikawa ........................ 151
11, 12. Different focus levels, EM 8-15, 246-249 cm, L-2
(V24/4).
Corbisema apiculata (Lemmermann) .. .................0......ccccees 152
13. $136/964, L-2 (U30/4); 14. SID 1, R-1 (E7/1); 15. KEL
9, R-1 (M23/3); 16. DSDP 6-6-2, 1-2 cm, R-1 (V34/4) ;
17. DSDP 6-6-2, 1-2 cm, L-2 (H18/3).
Corbisema archangelskiana (Schulz) ate tee 152
18. DSDP 6-4-3, 110-112 cm, L-4 (G29/3)
BuLL. AMER. PALEONT., VOL. 62 PLATE 23
BULL. AMER. PALEONT., VOL. 62
PLATE 24
14-17.
18-23.
SILICOFLAGELLATES AND EBRIDIANS: LING 215
EXPLANATION OF PLATE 24
All figures 500 unless otherwise indicated.
Corbisema bimucronata Deflandre .......
1. WR TR 39F (J.S.1068), L-2 (C41/4).
Corbisema’ .geometrica) Hanna. £2... 22... J.ccc0.5sksc.--deccscecesaveectessovnees 154
2. CAS 1144, R-5 (D30/1); 3. DSDP 6-6-2, 1-2 cm, R-2
(K17/4), X400; 4. DSDP 6-6-2, 1-2 cm, R-2 (J36/3), 400.
Corbisema hastata (Lemmermann)
5. DSDP 12-3-1, 120-121 cm, L-2 (P18/1).
Corbisemayrectan(Schulz)e..2 0 ee re te cess ate 155
6. DSDP 13-3-1, 120-121 cm, L-2 (U18/0); 7. DSDP 13-3-1,
120-121 cm, L-2 (P40/0).
Corbisema triacantha Ehrenberg ....00.00.0...0...ccccccccceecececceee 156
8, 9. Different focus levels, KEL 10, R-1 (J48/0); 10. KEL 7,
L-2 (K37/0); 11. WRE 67-99, L-2 (M37/0); 12. WRE 67-99,
L-3 (D22/0); 13. AMPH 6P, 50-51 cm, L-2 (L22/3).
Corbisema triacantha var. flexuosa Stradner
14. AUS 4, L-2 (W17/0); 15. Lateral view, AUS 4, L-2
(W25/0); 16. AUS 4, L-3 (K26/0); 17. WRE 67-96, L-2
(F26/0).
Corbisema triacantha var. minor (Schulz) .... ..................... 158
18. EM 8-15, 246-249 cm, L-2 (S7/0); 19. WRE 67-96, L-2
(K2273) e205 Keb 9s e-2) (12/2) 21D SDP 6-4-2, 7=10
cm, L-2 (R32/0); 22. DSDP 6-4-3, 110-112 cm, L-2 (Z32/2) ;
23. DSDP 6-6-2, 1-2 cm, R-2 (M39/4).
216
Figure
1-10.
11-15.
16,17.
18, 19.
BuLLeTIN 273
EXPLANATION OF PLATE 25
All figures 500 unless otherwise indicated.
Dictyocha ausonia Deflandre 2.0.02... ccceceee cette 160
1. WRE 67-77, L-7 (R18/2); 2. WRE 67-99, L-2, 019/0; 3.
WAH 7P, 259-260 cm, L-2 (Y13/4); 4. WAH 7P, 259-260
cm, L-2 (Q10/2); 5. EM 8-9, 148-149 cm, L-2 (D19/2); 6-10.
WR TR (K9391), L-2 (U47/4); 7. WR TR 23 (K9391) L-2
(M17/4); 8. WR TR 23 (K9391), L-2 (N23/2); 9. WR TR
23 (K9391), L-2 (H10/4); 10. WR TR 23 (K9391), L-2
(N28/0).
Dictyocha fibula Ehrenberg ............... ete: PE Reson: Gi
11. WR TR 11, Low (Bo. 202), L-2, (W34/1); 12. WR TR
11, Low (Bo. 202), L-2 (K33/4); 13 PROA 88P, 275-276
cm, L-2 (H28/1); 14. EM 6-2, 30-32 cm, L-2, (Y18/0);
15. AMPH 6P, 17-18 cm, L-2 (N17/2).
Dictyocha fibula var. aculeata Lemmermann ...................... 161
16. DSDP 9-5-2, 99-100 em, L-2 (U29/3); 17. DSDP 9-5-1,
8-9 cm, L-2 (L23/0).
Dictyocha fibu'a var. messanensis (Haeckel) | .........0...0........ 161
18. DSDP 9-5-2, L-2, 99-100 cm, L-2 (Y32/4); 19. DSDP
9-5-2, 99-100 cm, L-2 (016/2).
PLATE 25
BULL. AMER. PALEONT., VOL. 62
uw oo aT,
wt
ty
Oey
PLATE 26
Bulut. AMER. PALEONT., VOL. 62
Figure
I, 24,
17-19.
20-22.
23, 24.
SILICOFLAGELLATES AND EBRIDIANS: LING 217
EXPLANATION OF PLATE 26
All figures <500 unless otherwise indicated.
Page
Dictyocha fibuia var. octagona Tsumura ............ treats SGI
1. AMPH 7P, 17-18 cm, L-2 (P7/3); 2. EM 7-1, 35-36 cm,
L-2 (W8/4).
Dictyocha jhexacanthacochulz 4 iv...cnulugete...2ceeelso)...8 162
3. $136/964, L-2 (D10/0).
Diciyocha sp. ... sie «ile ee RE 164
4. AMPH 6P, 17-18 cm, L-2 (W9/0); 5. EM 8-15, 505-506
cm, L-2 (L12/4); 6. EM 8-11, 322-323 cm, L-2 (S8/0) ;
7. EM 8-11, 69-70 cm, L-2 (B33/4); 8 EM 8-11, 69-70
cm, L-2 (S14/2).
Distephanus crux (Ehrenberg) 1b RAE eR OP ces LOD
9. EM 8-11, 421-423 cm, L-2 (G8/4); 10. EM 8-12, 94-95 cm,
L-2 (E17/1); 11. AMPH 6P, 17-18 cm, L-2 (¥9/0); 12.
EM 6-2, 30-32 cm, L-2 (N46/2); 13. $136/880, L-2 (N30/4) ;
14. WRE 67-77, L-4 (N49/2); 15. WRE 67-77, L-7 (Z36/2) ;
16. WRE 67-77, L-7 (G32/0).
Distephanus crux var. longispina Schulz ................... LOD
17. Oblique lateral view, EM 8-14, 388-389 cm, L-2 (P31/4) ;
18, 19. Different focus levels, EM 8-13, 125-126 cm, L-2
(M19/0).
Distephanus crux var. octacanthus
Desikachary and Maheshwari ........ A eee eed | el 66
20. EM 7-2, 6-7 cm, L-2 (G17/2); 21. EM 7-1, 35-36 cm, L-2
(V24/1); 22. EM 7-1, 14-15 cm, L-2 (U25/4).
Distephanus speculum (Ehrenberg) .....000......00000.0cceccceceeeeeeeeee. 167
23. WAH 7P, 516-517 cm, L-2 (N14/0); 24. WRE 67-112,
L-2°(¥17/0).
218
Figure
152:
3, 4.
5-7.
8-13.
14, 15.
16-18.
19-21.
BuLLeTIN 273
EXPLANATION OF PLATE 27
All figures 500 unless otherwise indicated.
Distephanus speculum (Ehrenberg) ....................0::cccccceescceteeeees 167
1. EM 8-12, 26-28 cm, L-2 (J41/1); 2. EM 6-2, 10-11 cm,
W-2 (W31/0).
Distephanus speculum var. pentagonus Lemmermann ........ 167
3. EM 8-11, 322-323 cm, L-2 (F11/0); 4. EM 8-11, 322-323 cm,
L-2 (G35/4).
Distenphanus speculum var. pseudocrux Schulz .................... 168
5. EM 8-9, 148-149 cm, L-2 (D45/4); 6. EM 8-9, 148-149 cm,
L-2 (V29/0); 7. EM 8-9, 148-149 cm, L-2 (L44/0).
Distephanus speculum var. pseudofibula Schulz .................... 169
8, 9. Different focus levels, EM 8-10, 383-384 cm, L-2 (G26/3) ;
10, (EM. ‘6-2, 10-11 “cm, L-2 (J40/0); 11: EE! 18-020 Rs
(G30/0); 12, 13. Different focus levels, TT 18-02, R-1
(N23/0).
Lyramula tenuipertica Kokubo et Tsumura ............................ 171
14. DSDP 9-5-2, 99-100 cm, L-2 (U14/4); 15. the same
specimen of figure 14 under the phase contrast photomicro-
graph.
Lyramula furcula Hanna ......0.:....6.cecc100005. See ee 171
16. CAS 1144, L-2 (C24/1); 17. the same specimen of figure
16 under the phase contrast photomicrograph; 18. CAS
1144, L-3 (G26/4).
Lyramula furcula var. minor Deflandre | .................000...000....... 172
19. CAS 1144, L-3 (J21/0); 20. CAS 1144, L-3 (Q23/3);
21. CAS 1144, L-3 (Q23/3).
BULL. AMER. PALEONT., VOL. 62 PLATE 27
BuLL. AMER. PALEONT., VOL. 62 PLATE 28
5, 6.
a8.
9-15.
SILICOFLAGELLATES AND EBRIDIANS: LING 219
EXPLANATION OF PLATE 28
All figures 500 unless otherwise indicated.
Page
Eyramulae simplex: Hannas forte: eee ee 172
1. CAS 1144, L-2 (Q36/4).
Mesocena apiculata (Schulz) ..............0.....cccccceccceseseceeseeeseneeeees 173
2. $136/966, L-2 (L13/0); 3. the same specimen of figure 2
under the phase contrast photomicrograph; 4 §$136/880,
L-2 (R5/4).
Mesocena circulus Ehrenberg 2...............0.cccccccccccccceeeeeceeeeeeeeeeeees 175
5. EM 8-11, 69-70 cm, R-1 (S43/4) 400; 6. EM 8-11, 69-70
cm, L-2 (F17/2).
Mesocena circulus var. apiculata Lemmermann ................... 176
7. EM 8-12, 94-95 cm, L-2 (L13/2); 8 EM 8-15, 246-249
cm, R-1 (N43/3).
Mesocena cf. elliptica Ehrenberg |......0.00.0.0000000..0000.0cc eee atzte/
9. EM 6-2, 30-32 cm, L-2 (D10/2); 10. EM 8-11, 69-70 cm,
L-2 (L32/3), 400; 11. EM 8-15, 505-506 cm, L-2 (W26/4) ;
12. AMPH 6P, 50-51 cm, L-2 (D13/2); 13. AMPH 6P, 50-51
cm, L-2 (J31/3); 14. WRE 67-77, L-2 (026/1); 15. WRE
67-76, L-2 (M13/1).
220
Figure
1-3.
4-7.
8-10.
11-15.
BuLLeETIN 273
EXPLANATION OF PLATE 29
All figures 500 unless otherwise indicated.
Mesocena cf. elliptica Ehrenberg ........00.0...........0.:cceeeseeeeees 177
1. LSDA 101G, 38-39 cm, L-2 (E44/1); 2. EM 8-13, 49-51 cm,
L-2 (C8/3); 3. WR TR 11, Low (Bo. 202), L-2 (E42/2).
Mesocena cf. eliiptica var minoriformis Bachmann and Papp 178
4. AUS 3, L-2 (08/4); 5. AUS 3, L-2 (G16/4); 6. EM 8-13,
L-2 (R37/1); 7. EM 7-2, L-2 (D18/2).
Mesocena oamarvensis SchulZ ..................cccccceccccccecccceceeveeeeeeeeess 179
8. S$136/1096, L-2 (J11/0), 400; 9. KEL 6, R-2 (S23/2);
10.SID 3, R=1) (J50/2).
Septamesocena apiculata Bachmann |... cee 193
13. WRE 67-99, L-3 (S41/4); 14. the same specimen of
figure 13 under the phase contrast photomicrograph; 15.
DSDP 13-3-1, 120-121 cm, L-2 (S10/2).
BuLL. AMER. PALEONT., VOL. 62 PLATE 29
PLATE 30
BULL. AMER. PALEONT., VOL. 62
Fs ites
aero.
rigs tn
=
ya Fv
oan
Figure
1-4.
9-11.
12-16.
17-19.
SILICOFLAGELLATES AND EBRIDIANS: LING 2a
EXPLANATION OF PLATE 30
All figures 500 unless otherwise indicated.
Naviculopsis biapiculata (Lemmermann) 2.0.0.0... 181
1, 2. Different focus levels, lateral view, S136/914, L-2
(R31/4); 3. DSDP 6-6-2, 1-2 cm, L-3 (Q17/0); 4. WR TR
39F (J.S.1068), L-3 (R40/1).
Naviculopsis constricta (Schulz) ..0..000000....000....... ea el oS
5. S136/878, L-2 (N48/0); 6. Lateral view, S136/1096, L-2
(C23/0); 7. KEL 9, R-1 (E46/3); 8. DSDP 6-6-2, 1-2 cm,
R-2 (W46/0).
Naviculopsis foliacea Deflandre ........0.............50ccceeecceetteee es 184
9. Lateral view, KEL 3, R-1 (V47/3); 10. Oblique abapical
view, KEL 8, R-1 (G13/0); 11. DSDP 6-6-2, 1-2 cm, L-2
(G33/3).
Naviculopsis; latan (Deflandre) ie rrr caer ee 185
12. AUS 4, L-3 (U36/3); 13. the same specimen of figure 12
under phase contrast photomicrograph; 14. Abapical view,
under the phase contrast photomicrograph, WRE _ 67-100,
L-2 (034/2); 15. AUS 4, L-3 (X31/3); 16. WRE 67-76,
L-4 (W16/3).
Naviculopsis navicula (Ehrenberg) ..............000....00...c cece 186
17. WRE 67-77, L-4 (K49/0); 18. WRE 67-77, L-2 (C29/0);
19. a part of the same specimen of figure 18 with higher
magnification, 1250.
222
Figure
12-15.
BULLETIN 273
EXPLANATION OF PLATE 31
All figures <500 unless otherwise indicated.
Page
Naviculopsis nmavicula (Ehrenberg) .....000................000ccccceceeee 186
1. WRE 67-77, L-6 (W39/2).
Naviculopsis quadrata Ehrenberg ......0.00000......0...ccceeccccecece ees 187
2. WRE 67-77, L-6 (W40/0).
Naviculopsis regularis (Carnevale) .....000....0000...cccc eee 188
3. WRE 67-95, L-2 (G12/); 4. WRE 67-112, L-2 (037/0); 5.
WRE 67-99, L-4 (K32/1).
Naviculopsis, Sp* 2.20.6. S80 ee... eee 189
6. AUS 3, L-2 (M22/0).
Naviculopsis trispinosa (Schulz) oo....000.0..ccccccccccccceeeeeceeeeeeees 190
7, 8. Different focus levels, lateral view, $136/966, L-2 (Q26/0).
Paradictyocha polyactis (Ehrenberg) (0 ooo...cccccceccee 190
9. EM 8-15, 36-37 cm, L-2 (R45/1); 10. EM 8-15, 36-37 cm,
L-3 (E16/0); 11. EM 8-15, 36-37 cm, L-4 (R9/0).
Rocella gemma Hanna PRR R kN en Ee 192
12. JYN V 16 P, 231-233 cm, L-2 (L48/0); 13. MSN 149P,
294-296 cm, L-2 (Y16/2); 14. MSN 149P, 294-296 cm, L-2
(P27/4); 15. MSN 149P, 294-296 cm, L-2 (C19/3).
BULL. AMER. PALEONT., VOL. 62 PLATE 31
BULL. AMER. PALEONT., VOL. 62 PLATE 32
Figure
IP
3-5.
6-12.
13-18.
19-21.
SILICOFLAGELLATES AND EBRIDIANS: LING 223
EXPLANATION OF PLATE 32
All figures 800 unless otherwise indicated.
Page
Wallacerfarhortonii blanma se tie oo eee tee te a cate nese eearee nce 194
1. CAS 1144, L-3 (N34/4), 500; 2. CAS 1144, L-3 (N18/3),
< 500.
Ammodochium rectangulare (Schulz) .....00.0..0.0000..c. 195
3, 4. Different focus levels, EM 7-1, 35-36 cm, L-2 (C10/0) ;
5. $136/880, L-2 (M7/4).
Ebriopsis antiqua (SCHULZ). ees cee ee cee . 196
6, 7. Different focus levels, EM 6-2, 10-11 cm, L-2 (C38/0) ;
8. EM 7-1, 36-36 cm, L-2 (S33/3) ; 9. EM 8-9, 148-149 cm,
L-2 (G28/1); 10. EM 8-15, 505-506 cm, L-2 (W32/0) ;
11, 12. Different focus levels, DSDP 6-6-2, 1-2 cm, R-1
(R11/0).
Ebriopsis crenulata HOvasse ooo.........cccccceecccee ee ceeeceeeeees 197
13, 14. Different focus levels, $136/914, L-2 (U23/0); 15-17.
Different focus levels, DSDP 6-6-2, 1-2 cm, L-2 (H11/2);
18. DSDP 6-4-3, 110-112 cm, L-5 (X20/2).
Parathranium tenuipes HovaSsSe_ ..............0....0cc0ccccccceeeeeeeeeeeeeeeeeeee 198
19, 20. Different focus levels, EM 8-9, 148-149 cm, L-2 (J29/1) ;
21. EM 8-15, 505-506 cm, L-2 (V17/1).
INDEX
Note: Light face figures refer to page numbers. Bold face figures refer
to the plate numbers.
A
var. aculeata,
Dictyocha fibula 25 161, 202, 216
alata, Dorcadospyris.. 149, 161, 162,
166, 178, ae
Algerian jfpecereeses 60
Ammodochium .......... 195, 196
Antanrcticiwmc 148
antepenultimus,
Ommatartusee. 149, 166, 170
antiqua, Ebriopsis ..32 196, 197, a,
apiculata,
Corbisema .......... 23 151, 152, 214
Mesocena _............ 28 173, 174, 177,
193, 219
Paradictyocha ........ 176
apiculata,
Septamesocena ..29 145, 174, 177,
180, 193, 220
forma apiculata,
Dictyocha triacan-
tha var. apiculata .. 152
var. apiculata,
Dictyocha navicula.. 182
Dictyocha
triacantha ............ 151, 152
Mesocena
CiGCULUS) 22... 28 175, 176, 191,
200, 219
Mesocena
oamaruensis ........ 173, 193, 194
aragonensis,
Hantkenina ............ 182
archangelskiana,
Corbisema .......... 23 152, 154, 214
Dictyochawe 152
var. archangelskiana,
Dictyocha
tiacantha ee 152, 155
forma armata, Diste-
phanus speculum
var. pentagonus .... 168
aseptatae, Mesocena .. 173
forma aspera,
Dictyocha navicula
var. biapiculata ...... 181, 182
Dictyocha
triacantha var.
apieulata, ....0.008 151
asteroides,
Distephanus ............ 167
PRET ENIC Rs ioe ee 191, 203
ausonia, Dictyocha 25 159, 163, 164,
185, 189, 200,
202, 216
cf. ausonia, Dictyocha 159, 160, 202
Austria (or Austrian).. 142, 149, 158,
159, 178, 181,
189
B
Bain’s Farm section .. 142, 174, 190
Bat DAGOS) Foc sen kaart 139, 146, 153,
163, 175, 183,
184, 185
barisanensis, Globoro-
taliaetonsivee nso: 148
Bering Sa) ..-fe4 ee 148, 169, 170,
171
Bermudaneseest.. ee 187
biapiculata,
Dictyocha (7?) ........ 181, 184
Naviculopsis _...... 30 181, 182, 183,
184, 221
var. biapiculata,
Dictyocha navicula 181, 182, 185
Naviculopsis
biapiculata .......... 182
bimucronata,
Corbisema _.......... 24 153, 215
Dictyochay =. oee 154
bipartita, Dictyocha .. 148
bipes, Lychnocanium.. 142, 167, 203
bisectis, Coccolithus .. 193
bromia, Thyrsocyrtis.. 137
Buttle Diatomite ........ 178
Cc
California (or
Californian) 72. 142, 153, 154,
155, 163, 165,
168, 169, 171,
172, 173, 178,
181, 184, 185,
191, 192, 194,
195, 199, 200
California Academy
of Science (CAS) .... 138, 154, 171,
172, 173, 195,
199
Cannopilusie 146, 148
var. cannopiloides,
Distephanus
SPEGulun | -ce.. cat 148
224
INDEX
carinatus, Trique-
trorhabdulus’ =... 193
Chaetoceros_................ 170
chalara, Podocyrtis .... 155
circulus,
Dictyochal ys 175
Mesocena ......... 28 175, 176, 177,
201, 219
var. circulus, Meso-
cena elliptica .......... 176
clathrata, Dicladia ?.. 198
clathratum,
Parathranium _........ 199
Clathropyxidella ........ 146
var. complexa, Dictyo-
cha pseudofibula .... 169
constricta,
Naviculopsis ........ 30 183, 221
forma constricta, Dic-
tyocha navicula var.
biapieulata™ .20..4.2.. 183
var. constricta,
Dictyocha
Naviculasee ee 180
Naviculopsis
biapiculata —2....- 183
Corbisema’-2=....2.0 32 151, 153, 199
costata,
Calocycletta 20... 165, 178, 179,
186, 187, 188,
189, 203
crenulata,
Ebriopsis °2..))).. 32 196, 197, 198,
200, 223
crux,
Mictyocha. .......0 159, 165, 169
Distephanus ....... 26 164, 200, 217
forma decagona, Dis-
tephanus speculum
Var spolyactis) ..........
Deep Sea Drilling
Project (DSDP)
190
i. 135, 136, 137,
138, 153, 155,
156, 157, 161,
162, 166, 171,
182, 184, 185,
192, 193, 194,
197, 199, 200,
201
Spssaynbet sos 160, 165, 168,
170, 178, 197,
202
Delmontian
Dicladian =. ee 198
Dichvoehawe ee: 151, 159
diodon, Mesocena ...... Wiis LitosreOue
202
var. diodon, Meso-
cena crenulata ....... 178
Distephanus! 2. 164
forma dodecagona, Dis-
tephanus speculum
var. polyactis ....... 190
E
I DTIODSISsa ee cee eee 196
elliptica, Mesocena .... 177, 178, 200,
201
cf. elliptica,
Mesocena ....... 28, 29 173, 177, 178,
179, 202, 203,
219, 220
var. elliptica, Meso-
cena elliptica .......... 177
forma eocaenica, Dic-
tyocha fibula
Vite, IACI, cessonsgnenne 164
experimental Mohole
(EM) (or Mohole) .. 135, 137, 138,
142, 149, 150,
15 573159;
160, 162, 165,
166, 167, 168,
169, 170, 176,
Wiel S819)
192, 196, 197,
199, 200, 201,
202
F
fibula, Dictyocha .25 157, 160, 200,
216
Painilan dagen s =. fee. 165
var. flexuosa,
Corbisema
triacantha ....... 24 157, 158, 203,
215
Dictyocha
ftiacantha 158
foliacea,
Naviculopsis _...... 30 181, 182, 183,
184, 199, 221
Forrester’s Hill
SGCtiOniie 32 3...04eeee! 142, 162, 179
lergcha(ec ae). pe nee 181
furcula, Lyramula 27 171, 172, 173,
195, 199, 218
BuUreSland ses ae 182
225
INDEX
G
gemma, Rocella ...31 192, 200, 222
geometrica,
Corbisema ........... 24 151, 152, 153,
154, 155, 215
forma glabra, Meso-
cena polymorpha
var. triangulaliies:.. 193, 194
Globigerinoides
DAtuIMG ree 140
GreCCe eo ececs teetsee- 3s 175, 181, 186
H
Halicalyptraly -..-----:- 146
hastata, Corbisema..24 155, 157, 215
var. hastata, Dictyo-
cha triacantha 155
TSIGIRVEUTEI, socennspaecceeeoeee 149, 150, 165,
188, 192, 201
hemisphaericus,
Cannopilus _.......... 23 147, 148, 200,
201, 214
hexacantha,
Corbisema _............-.. 162
Dichyochay. .eese.. 26 162, 217
forma hexacantha,
Dictyocha
deflandrel | ............. 162
hexagona, Mesocena .. 143, 177, 178
Hojuji diatomaceous
mudstones |.)......-.---- 179
hortoni,
Vallacerta ........... 32 194, 195, 199,
223
JERTROERIAY ssa pecedeansesece 169, 194
I
iberica,
Naviculopsis ............ 188
var. iberica. Dictyo-
cha biapiculata ...... 187
forma inermis, Dictyo-
cha triacantha
Vat INeGMIS ese. 154
var. inermis,
Corbisema apiculata 186
Dictyocha
triacant has... ee 153, 154
Italy (or Italian) ........ 149, 157, 158,
159, 164, 178,
179, 186, 187,
188, 189, 202,
203, 204
J
Jackson’s Paddock
Sectlonue ee ates 142, 174, 180,
182, 198
Japan 2: | ees 159, 161, 165,
167, 169) 176;
176
japonicus,
Distephanus) 222. 169
Jutland 2...382eoe 154
K
Kellogg Shale ............ 142, 152, 179,
183, 184, 200
Kieselbalken: ...............: 182
Kreyenhagen Shale .... 163, 184
kugleri,
Globigerapsis .......... 163
L
WadOa: tse nce 168
var. laevis, Lyramula
tenuipertica ............ 171
lata, Naviculopsis .30 185, 186, 189,
203, 221
var. lata, Dictyocha
biapieulatal -se-core 185
forma late-radiata,
Dictyocha triacan-
tha var. apiculata 151,152
Dictyocha triacan-
tha var. inermis .. 152, 154
forma longispina,
Dictyocha crux ...... 165
Distephanus crux .. 165
var. longispina,
Distephanus crux 26 165, 201, 217
TAU STAN ee ee 149, 150, 157,
178, 192, 199,
201
loyramiulay ae eee 170, 171
M
Miryam died essences 188
Matuyama Reversed
EypoOchiae ee 202
Mediterranean ............ 159, 167, 202,
Pu 203, 204
mesnili,
IE DRIOpSISHee ee 196, 197
Podamphora ............ 197
Mesocena 2.2... 145, 173, 194
226
INDEX
forma mesocenoidea,
Distephanus
polyactis ....... 176
Paradictyocha
polyactis ........ 45 176
var. messanensis,
Dictyocha fibula 25 145, 161, mie
IMGXIGOMLS Soke ; 192
forma minor,
Corbisema
triacantha 158
Dictyocha
triacantha .... 157,158
Dictyocha
triacantha var.
iracanihale- 159
var. minor,
Corbisema
triacantha ...... 24 158, 201, 215
Dictyocha
friacanthal 22.2. 157
Lyramula
furculave. er, 172, 218
minoriformis, Meso-
cena elliptica .......... 178
var. minoriformis,
Mesocena elliptica .. 178
Mesocena cf.
elliptica ........... 29 178, 220
Mohnian, -2..22.08...) 149, 150, 160,
165, 168, 169.
LOS Ge S:
179, 199, 201,
202
Moron) Spain®s............ 151
Morsi sland ........:....<. 157, 159
mutabilis, Dictyocha.. 185
N
navicula,
Dictyocha 7-4... ‘ 180, 186
Naviculopsis 30, 31 181, 184, 186,
187, 189, 221,
222
Naviculopsis .....:........ 160, 180, 181,
199, 203
forma naviculoidea,
Corbisema apiculata 186
var. naviculopsis,
Dictyocha navicula. 186
Nicobar Island ............ 166
INOGEWAYE o)c:eo 191
re)
Oamaru (or Oamaru
Diatomite) .......
oamaruensis,
Corbisema ...
Mesocena ...........29
octacantha, Dictyocha
var. octacanthus, Dis-
tephanus crux ...26
octagona, Dictyocha ..
var. octagona, Dictyo-
cha shibulas. 26
octonaria, Dictyocha .
ornamentum,
Dictyocha
Pacific
Paradictyocha
Parathraniume 2.
forma pentagona,
Dictyocha speculum
var. pentagonus,
Distephanus
speculum 27
pentas, Spongaster ....
petterssoni,
Cannartus (?)
picassoi,
Cannopilus .......... 23
Phyllodictyocha .........
polyactis,
Dictyochaws
Distephanus ............
Paradictyocha .. 31
forma polyactis,
Distephanus specu-
lum var. octonaria
Paradictyocha
polyactis =...
var. pOlyactis, Diste-
phanus speculum .
227
142, 152, 153,
162, 163, 174,
177, 179, 180,
182, 183, 190,
194, 196, 198,
200
179
179, 193
166, 201
166, 200, 201,
217
161, 162
161, 200, 201,
17
161, 162
161
135, 148, 162,
166, 167, 170,
173, 192, 201,
202, 203
190
198
167
167, 201, 218
164, 167, 178
156
190, 191, 192,
201, 222
190
191
190,
INDEX
pons, Dictyocha 159, 160
var. pons, Dictyo-
cha naviculale. 159
ponticulus, Dictyocha 186
forma pseudocrux,
Distephanus
speculum) =)... 168
var. pseudocrux,
Distephanus
Speculum! 22-72 27 168, 202,218
pseudofibula,
Dictyocha 22... 169, 170, 202
forma pseudofibula,
Distephanus
Japonicusn.....4e.2 168, 169
Distephanus
speculum) se. 169
var. pseudofibula,
Distephanus
speculum ....... 27 169, 201, 218
Q
var. quadrangula,
Mesocena elliptica.. 177
Mesocena
polymorpha ........ 141, 177, 178
quadrata,
Dichyoenay 4... 187
Naviculopsis ........ 31 187, 203, 222
var. quadrata, Dictyo-
cha siderea .............. 195
recta,
Corbisema ............ 24 155, 215
Dictyochae..t- 155
Phyllodictyocha . 155
var. recta, Dictyocha
tracanthase 155
rectangulare,
Ammodochium. ... 32 195, 196, 200,
202, 223
var. rectangulare,
Dictyocha navicula.. 187, 188
rectangularis,
Naviculopsis .......... 187, 188
recurvus,
Isthmolithus ............ 153, 156
regularis,
Dictyocha ........... 160, 188
Naviculopsis
_... 31188, 189, 203,
222
var. regularis, Dictyo-
cha ausonia
Relizian
185, 189
149, 165
forma rhombica,
Dictyocha fibula
var. fibula 163, 164
var. rhombica,
Dictyocha fibula .... 164
var. rhombus,
Dictyocha fibula .... 164
robusta, Naviculopsis 185
ROCA te 35 eee 192, 200
Russia (or Russian) .. 153, 155, 158,
160, 164, 165,
iPr Ges als),
180, 182, 185,
187, 194, 195
S
Septamesocena .......... 173, 193, 194
septatae, Mesocena .... 173
Siberag 2 eee 182
Sicily? bie ee Itsy, les
siderea, Dictyocha ... 194, 195
sidney, ‘Shales... 142, 152, 163,
179, 184, 200
simplex, Lyramula 28 172, 219
Siphonotestales .......... 146
SDAIN fa. eee eee 188
speculum,
Dictyocha, 222... 167
Distephanus . 26, 27 147, 148, 162,
166, 167, 168,
191, 200, 203,
PAL, PA:
sphaericus,
Cannopilus ......... 23 149, 150, 159,
201, 214
stauracanthus,
Distephanus ........... 166
Stereotestales ............ 195
subarctios, Dictyocha.. 202
T
tenuipes,
Parathranium ... 32 198, 200, 202,
223
Thranivm ee 198
tenuipertica,
lbyramulal eee 27 171, 218
tetraceros,
Cannopilus ..........23 150, 151, 214
Mortontanieren cee 149, 150, 176,
179, 201
228
INDEX
triacantha,
Corbisema ............ 24 156, 157, 159,
166, 201, 203
Dictyocha,=— 4... 156
HV ESOCYLLISM see 155, 156, 163,
182, 183
forma triacantha,
Dictyocha triacan-
tha var. triacantha.. 157
var. triacantha, Cor-
bisema triacantha .. 158
triangula, Dictyocha . Ilgey, Lee/z
var. triangula, Meso-
cena polymorpha 173, 193
trifenestra, Dictyocha 157
trigona,
Corbisemas- 158
Dietyochay 156, 157
forma trigona,
Dictyocha crux ..... 159
Atreab aN Gl, Ve cey ee ee cane 139, 178, 179
triommata, Dictyocha 156, 157
trispinosa,
Naviculopsis ....... 31 190, 222
forma trispinosa,
Naviculopsis
bpliapiculata =o... 190
var. trispinosa, Dic-
tyocha navicula z 190
tuberosa, Theocyrtis.. 137, 153, 156,
183
V
valida, Ebriopsis . 196, 197
Vallacertameee- ine 194
Valmonte Diatomite . 178
variabilis,
Distephanus .... 168
WATERED co sccmosncceepseccatcote 146, 198
virginis,
Calocycletta ....... .. 140, 158, 160,
167, 179, 186,
189, 203
WwW
Wakura Beds .............. 151
229
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BUEREE TINS
OF
AMERICAN
PALEONTOLOGY
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MUS. COMP. ZOOL
mea ote LIBRARY
Vol. 62 =
R 9 19/3
No. 274 FE
HARVARD
UNIVERSITY
SILURO-DEVONIAN MICROFAUNAL
BIOSTRATIGRAPHY IN NEVADA
By
W. A. McCLeLLAN
1973
Paleontological Research Institution
Ithaca, New York 14850, U.S.A.
PALEONTOLOGICAL RESEARCH INSTITUTION
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BULLETINS OF AMERICAN PALEONTOLOGY
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BWIREEATINS
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Vol. 62
No. 274
SILURO-DEVONIAN MICROFAUNAL
BIOSTRATIGRAPHY IN NEVADA
By
W. A. McCLeLLan
January 29, 1973
Paleontological Research Institution
Ithaca, New York 14850, U.S.A.
Library of Congress Card Number: 72-86844
Printed in the United States of America
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CONTENTS
Page
JéN| OSL S| cee PRs Tap as Ae ES ng A Ue ee Be PE naa TE 235
PACK OW GG SIL e Its fae eeees were ee eee ee ae ee eee ce Sn eee 235
Tin tO GUC et orig cee a ee oO Rl rE sca MeLct jenna een eee 235
Stratignapliy: | a 08 ie wees Se ee ie Be A ie eae Repeat) 238
Balleontol ogynctetcee ee ee ee ee ee Pies ees) A LRN ee 246
LEE SESE) (Yo ia aE ee Bet tn SR AN EE BRE Mit Poa 252
PESTO SE REN Tc EY pe cn ee RE RR bee ce ea nN ih Seago cde degaieseeee 257
Systematicmp alle on tol o pry ge eee see ee tee ec ee cates a oe 2 2a ons Wetec zan as ee 275
MoLaminite rida cee etn eee eee ee ee SO er WS Ae 275
(QELS ETO IE ee ek Ae eee A A en ee 320
WG Gallntiygpnte OSte tye sees tae ce ese eet eee ear ees, eee ee NR ete ea te heats eA td 345
References @ite digests 5 tke Minis Binary cele eh A Aes tle as Sono Si ca Seer eh aa cPoaseeanaete 351
Plates scans ree rR ue ben ee ipo: | Petey WIR Sewn, pit eee 7 eet desc ie 357
TABLES
Table Page
1. Extended geographic distribution of known Silurian genera .................. 249
2. Extended stratigraphic range of known SpeCies ..............-..--.c--ccssccsecsesesee-s 249
Sa CON OC OMS yg ee ce areata tec ht re NEM ee 5 OE DE 252
4. Family Astrorhizidae, abundance and stratigraphic range ......................- 259
5. Subfamily Psammosphaerinae, abundance and stratigraphic range ...... 259
6. Subfamily Saccammininae, abundance and stratigraphic range ................ 260
7. Subfamily Hemisphaerammininae, abundance and stratigraphic range .. 260
8. Additional families, abundance and stratigraphic range _.........................- 263
oe Order Laleocopidaystraticraphicy range) ese eee ee ere ere eee eee eee 264
1OsOxdersPodocopidawstratignaphic ran ce mses. cece eee eee 264
11. Stratigraphic occurrence of species at March Spring ....................::..:--+-- 265
12. Stratigraphic occurrence of species at Ikes Canyon .......................---.------- 266
13. Stratigraphic occurrence of species at Rabbit Hill ......................:......-00 267
14. Stratigraphic occurrence of species at Copenhagen Canyon ................-...-- 268
15. Stratigraphic occurrence of species at Lone Mountain ......................--.---- 268
16. Stratigraphic occurrence of species at Wood Cone ....................-::--::-00ee0--0-- 269
Wee otratioraphic occurnence Of species) at. Gatecl ithe es esses eececccaceseeoeesceoeeeeee 269
18. Stratigraphic occurrence of species at Pete Hanson Creek .....................-.. 270
19. Stratigraphic occurrence of foraminifers at Willow Creek ..................4 271
20. Stratigraphic occurrence of ostracodes at Willow Creek ....................... 272
21. Stratigraphic occurrence of species at Coal Canyon ..............2..---:::--00------ 273
22. Stratigraphic occurrence of species at Cortez Canyon ........................0.-.---- 274
ILLUSTRATIONS
Figure Page
iy dindex map) of centrall Neva die eee cee cee tes cee cee srens coe = pee c cee sc eeccecrecen cco cceecceeesnen=s 237
2. Stratigraphic relationships of Silurian and Lower Devonian
EORINE ACTON Sealine CE MG all RIN ayced Cl elec ea ete eo ene ae 239
3. Siluro-Devonian Lithosomes in central Nevada ........ Bde cess t cwate tebe aaustaae 240
Roraminiteralimassemb laren Zz OMe si eee es ceree cerca eens cee cee cee arenes eee 258
ah a
AWAITS
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tr@me tenes
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(Lived) (err
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rian geires 7h hangeep jadi
SILURO-DEVONIAN MICROFAUNAL
BIOSTRATIGRAPHY IN NEVADA
W. A. McCLeELLAN
ABSTRACT
From a regional microfossi] study of Silurian and Lower Devonian strata
in central Nevada, a large and diversified fauna of Foraminiferida and
Ostracoda is described in detail for the first time. The foraminifers consist
of 22 genera and 45 species. Two of the genera, Anictosphaera and Atelika-
mara, and the following species are new: Anictosphaecra progressa, Asche-
monella bastillensis, Atelikamara incomposita, Haplophragmoides antiquus,
Hemisphaerammina bipatella, Sorosphaera compacta, Sorosphaera inflata,
Storthosphaera malloryi. Recognition of Haplophragmoides antiquus in the
Lower Devonian of central Nevada is the earliest record of the family Lituo-
lidae. The ostracode fauna contains 29 genera and 20 identifiable species.
Of these, Longiscapha is a new genus, and Aechmina cortezensis, Longiscapha
nevadensis, Rudderina mutaspina, Thlipsura bispinosa, Ulrichia obesita, and
Winchellatia fragilis are new species.
The Foraminiferida recorded here have been combined with previously
known Silurian species from the Midwest to establish a tentative series of
assemblage zones for the Silurian and Lower Devonian of North America,
which are based on dominance of the different subfamilies. The Nevada
fauna represents the Upper Silurian, characterized by the Psammosphaerinae,
and the Lower Devonian, characterized by the Astrorhizinae and Hemi-
sphaerammininae. The Lower and Middle Silurian assemblages, best known in
the Midwest, have been divided into the following three zones: 1) Albion —
Hippocrepininae, Rhizammininae, Ammodiscinae; 2) lower Niagaran — Sac-
cammininae, Ammodiscinae; 3) middle Niagaran — Hippocrepininae, Psam-
mosphaerinae, Hemisphaerammininae. In general, there is not sufficient evi-
dence for widespread correlations on the basis of the ostracodes recognized
here, though several genera and species of the Nevada fauna compare closely
with those known from Appalachian strata of uppermost Silurian and Lower
Devonian age.
The physical environment shows local variances with time which are
recognized regionally as lithosomes. Association of the foraminifers with the
several lithosomes suggests overall similarity to associations in other parts
of North America. Associations between ostracodes and the various lithosomes
remain uncertain due to their sporadic and limited occurrence.
ACKNOWLEDGMENTS
The writer wishes to especially thank Dr. V. S. Mallory for
his critical review and suggestions on several important points in
this study, Dr. John Kepper also assisted in many ways. The
Atlantic Richfield Company generously contributed field expenses
and research facilities. Additional contributions for field expenses
were made by Humble Oil and Refining Company. The University
of Nevada, Las Vegas, provided research funds for scanning electron
microscopy. The Paleontological Museum of the University of Cin-
cinnati in Cincinnati, Ohio, provided type specimens for study.
INTRODUCTION
Paleontological studies of Silurian and Lower Devonian rocks
in the Cordilleran region have been mostly centered around the
236 BULLETIN 274
brachiopod faunas. Detailed microfossil reports have dealt exclu-
sively with conodonts, though recent progress has been made with
chitinozoans. Analysis of the combined brachiopod, conodont, and
graptolite faunas has made possible the faunal subdivision of these
strata, and certain of the elements have proven useful for inter-
regional correlations.
Foraminiferal studies of Middle Paleozoic rocks have never
been made west of Oklahoma. The work of Ireland (1939) from that
area is the only detailed study of the Upper Silurian and Lower
Devonian. The only report of ostracodes from these strata in
Nevada is that of Berdan in Gilluly and Masursky (1965).
Foraminiferida and Ostracoda are reported here in detail for the
first time from Nevada.
In addition to the systematic description of species, the fauna
has been applied as a biostratigraphic tool, and the age relation-
ships of the formations, previously established on macrofossils or
other bases, are re-examined in a regional framework throughout
central Nevada, (Text-fig. 1). At this time the microfauna cannot
be applied biostratigraphically in as refined a manner as the
brachiopods and graptolites.
The physical stratigraphic record is represented by a series
of lithologically distinct bodies (lithcsomes) which persisted, or
were recurrent, over an area for a considerable time. Lithosomes
are recognized and presented here for two reasons. First, they
demonstrate the changing depositional patterns across the region,
an understanding of which is necessary to correctly interpret and
correlate the various formational units. Second, they represent the
environment which must be related to the biota for ecologic in-
terpretations,
All microfossils which are figured or otherwise recognized as
identified material have been deposited in the micropaleontology
collections of the Geology and Paleontclogy Division, Thomas
Burke Memorial Washington State Museum, at the University of
Washington in Seattle. Locality numbers are designated by the
prefix UWA, and a description of each locality is given in the
Locality Register.
SILuRO-DEVONIAN MiIcroFAUNA: McCLELLAN
ree eh
W INNEMUCCA |
hy BATTLE CARLIN
ee ROUNTAINE— 23
BEOWAWE
MITCHELL -
CREEK
TIELEGRAPH
- NORTHUMBERLAND:
: CANYON :
= Limits of Study
= Town
= Major Section
. = Other Sections
Fig.1: INDEX MAP OF CENTRAL NEVADA
237
238 BULLETIN 274
STRATIGRAPHY
Silurian and Lower Devonian rock units in Nevada comprise
the major part of the Tippecanoe Sequence as demonstrated by
Smith (1966) in his study of this sequence throughout western
North America. Within the boundaries of the present study (Text-
fig. 1) these rocks have been preserved in considerable thicknesses
beneath the Wallbridge Discontinuity (Wheeler, 1963). They pre-
sent a complete lateral series of marine rock types, from back-reef
dolomites to reefoid units, fore-reef carbonates and shales, and
finally the eugeosynclinal cherts, black shales, and quartzites. The
various formational units (Text-fig. 2) are briefly outlined here
and then considered, as applicable, in a presentation (Text-fig. 3)
of the lithosomal units. The lithosomes more accurately illustrate
the important lithic variables of the physical environment, which
may be then related to the biological variables in the discussion
of paleoecology. Previous workers have discussed the formations,
and the purpose here is to develop an integrated picture of the
Silurian and Lower Devonian environment. Correlations (Text-
fig. 2) are based primarily on this integrated picture and the
Foraminiferida and Ostracoda.
FORMATIONS AND AGES
The first recognition of rocks of Silurian age was by Hague
(1892) who included all carbonates in the region ranging from
Upper Ordovician through Silurian time as the Lone Mountain
Limestone. He referred to those rocks, believed to be Devonian in
age, as the Nevada Limestone. Subsequent work has resulted in
several restrictions of these units as originally defined, as well as
nomenclatural additions, In recent years relative agreement has
been reached on the nomenclature by most workers. Merriam
(1940) restricted the name Lone Mountain to a massive dolomite
unit representing Late Silurian and possibly earliest Devonian
sedimentation. To the east, outside of this area, the Lone Moun-
tain Dolomite, thus restricted, merges with several other dolomite
formations which have different lithologic characteristics. The Lone
Mountain Dolomite has been demonstrated by Winterer and Mur-
phy (1960) to be a dolomitized reefoid complex cropping out in a
band striking north-northeast through east-central Nevada. The
reef grew in a westerly direction during Silurian time and _ per-
sisted into the early Devonian.
SILuRO-DEVoNIAN MicroFAuNA: McCLELLAN 239
West Fast
FROUAPUPUAUAHAD LRGRDURAPUDRDAPUDGATRAARINORADRRDAADOD
S | | | McColley Canyon
€ D a
fe) 5 .
sy) 8 et
® =
oe ee McMonnigal Rabbit
tS o .
: x Hill
° Lone
| Mountain
i Dolomite
|
| |
| ! z Masket
| £&
fore Ss
H = Shale
| a
c | | Roberts Mountains |
5 |
E | |
v | |
' =
Uv
©
= Bastille |
or Diana |
i
| > | |
> | |
= Gatecliff Hanson Creek
to} {
en te ee eee
Fig. 2: STRATIGRAPHIC RELATIONSHIPS OF SILURIAN
AND LOWER DEVONIAN FORMATIONS IN CENTRAL NEVADA
cia)
maa My
bf AD a.
i074 By
Mo 8
“ad | :
v, j =
A eaten on
: NE rar 8
zt ACEH : :
0 we pe---*--- =O 7) — = == »| Pe te
SS Whatate ai
: Pa os 507 ea ar a na | fj
gS
Pee Le
Soe AD Ne ara4y Hoe cabal
Spr. e> Gop eprasas#®.. Cy
PPLE GP ag
| AY ab! = sans
abel EAC
ACTRESS oo.
ek LH ——a.
SECECEACE RT :
SRT |
tal \ i Ne a :
LW N
LHL
J
\
aed i i aysiu :
a Mi MA : 4 2
; ni REE : ae
] / aT = Siena
| [| E ae
A
l ge
q
nha
ST ee
COPENHAGEN
CAMYON
<
(a)
<
=
Zz WwW
| 3
1
iP
Astrorhizinae (1)
Hemisphaerammininae (2)
DEVONIA
HELDERBERG
GEDINNIAN
ie
=
=)
— t
a
—S Psammosphaerinae (2)
| =
| {
| |
| | |
=i a
| a | =
Sipe phassle Wy aks ie
im | oO > Hemisphaerammininae (2)
f eeevhal, ies
ee ee er Psammosphaerinae (2)
| |
| Hippocrepininae (1)
ree | Saccammininae (2)
Lid
=
eri Ammodiscinae (3)
f=) |
} ae i
—— =
eee | Ammodiscinae (3)
| | ALBION | | Rhizammininae (1)
| iF Hippocrepininae (1)
Fig.4: FORAMINIFERAL ASSEMBLAGE ZONES
TAB Bs 4 f SILURIAN LOWER
t DEVONIAN
Family ASTRORHIZIDAE
Abundance and
Stratigraphic Range
Hyperammina casteri
Hyperammina constricta
Hyperammina curva =a
Marsipella torta
=]
ery bifurcata ee
Rhabdammina cylindrica | ———_—_}—s__1 V7
BBtersc PA
Rhabdammina major =|
Rare; 1-2 (——) Common: 6-10 (gma)
Uncommon: 3-5 (c™-) Abundant: over 10 (cum)
DEVONIAN
Subfamily PSAMMOSPHAERINAE
Cayuga
(Helderberg)
Abundance and
Stratigraphic Range
Ludlow
Anictosphaera progressa TTATTA___}
Ceratammina cf. C. cornucopia
Psammosphaera cava
Psammosphaera laevigata
Raibosammina aspera
Sorosphaera compacta
Sorosphaera confusa
Sorosphaera inflata
Sorosphaera osgoodensis
Stegnammina contorta
Seem cre
Storthosphaera malloryi | W777]
V7T/I 3
Rare: 1-2 ( ) Common: 6-10 (waz)
Uncommon: 2-5 (ct) Abundant: over 10 ( gmp)
TABLE: 6 LOWER
DEVONIAN
SILURIAN
(Helderberg)
Subfamily SACCAMMININAE
Abundance and
Stratigraphic Range
Sorostomasphaera waldronensis
Stomasphaera brassfieldensis
:
a
eae
Rare: 1-2 (——) Common: 6-10 (gza)
Uncommon: 3-5 (t=) Abundant: over 10 (dag)
TABLE: SILURIAN LOWER
DEVCNIAN
Subfam. HEMISPHAERAMMININAE
{Helderberg)
Abundance and
Stratigraphic Range
Atelikamara incomposita
Hemisphaerammina coronata
Hemisphaerammina discoidea?
Metamorphina gibbosa
Metamorphina tholus
Webbinelloidea hattini
Webbinelloidea hemispherica
Webbinelloidea atf W. nodosa
Webbinelloidea ventriquetra
Rare: 1-2 (——) Common: 6-10 (Zz)
Uncommon: 3-5 (t=) Abundant: over 10 ( gm)
Stturo-DeEvontan MicroFaAuNnA: McCLeLLan 261
nammina Moreman (1930) species and Anictosphaera progressa
characterize the very latest Ludlovian strata and Helderbergian
strata. The Hemisphaerammininae appear to be much more im-
portant in the Lower Devonian, especially Webbinelloidea Stewart
and Lampe (1947). An exception to this is Hemisphaerammina
bradyi which is most common in the Upper Silurian. Though not
as clearly defined as the Hemisphaerammininae, the family
Astrorhizidae is also better represented in the Lower Devonian.
However, nearly all of the astrorhizid species are known elsewhere
from all levels within the Silurian.
Pursuant to the above discussion, an informal zonation of the
Silurian and Lower Devonian in central Nevada is shown on Text-
fig. 4. The assemblage zones defined here are combined with those
characteristic of Llandovery strata to lower Wenlock strata in
southern Indiana. Further studies on the Upper Silurian in other
areas may provide a refinement of the assemblages evident in
Nevada.
Little work has been done on Devonian foraminifers, making
it difficult to establish any consistent relationship patterns. Ireland
(1939) reported several Silurian species from the Lower Devonian
Haragan Shale of Oklahoma, In addition he considered Ceratam-
mina cornucopia, Metamorphina bipartita, and Psammonyx max-
wells as restricted to the Lower Devonian. Other species, similarly
restricted, have since been found in the Silurian. Ceratammina cf.
C. cornucopia was found in Nevada in the latest Silurian; the other
species were not encountered, The Haragan foraminifers, and those
from Helderbergian beds here, have many characteristic Silurian
species. It appears that a gradual transition occurs, with introduction
of more and more Devonian species. The subfamily Hemisphaeram-
mininae is the dominant group.
Middle Devonian foraminifers are well known only in Ohio
(Stewart and Lampe, 1947; Summerson, 1958). Two widely ranging
Silurian species, Psammosphaera cava Moreman and Sorosphaera
osgoodensis Stewart and Priddy occur commonly in these beds.
Hemtsphaerammina Loeblich and Tappan (1964) and Webbinel-
loidea Stewart and Lampe (1947) are the dominant genera how-
ever. The Devonian species Hemtsphaerammina discoidea (Sum-
merson) has been reported in the Silurian Waldron Shale by Mc-
262 BULLETIN 274
Clellan (1966, p. 487) and questionably recognized here in the Rab-
bit Hill (Table 13). Webbinelloidea hemispherica Stewart and
Lampe, also known in the Silurian is more common in the Devonian,
Table 7 shows the increasing abundance and number of species of
these two genera in the younger strata of this study. There is still
a more marked difference between the Lower Devonian species of
Nevada and the Middle Devonian of Ohio than between any rocks
in the Silurian. Perhaps the most characteristic feature of the
foraminiferal fauna is the stratigraphic limitation of Stegnammina
Moreman (1930) and Fairliella Summerson (1958), As presently
known, the several species of Stegnammina are restricted to strata
no younger than Lower Devonian, and those of Fairliella occur only
in the Middle Devonian.
Because the earliest Silurian sediments studied here are Wen-
lock in age and the Waldron Shale in Indiana is lower Wenlockian,
there is only a small overlap between the two areas. There are
several species which characterize the overlap. Sorosphaera con-
fusa Brady which occurs sporadically throughout the Silurian, is
present in similar quantities in both Indiana and Nevada. Psam-
mosphaera laevigata White is common in the Lower Silurian and
only present in the lowest strata in Nevada. Similarly, Glomospira
Rzehak (1885), characteristic of the Lower Silurian, is represented
in Nevada by G. stluriana which occurs rarely in the same units as
Psammosphaera laevigata White. The only Silurian records of
Aschemonella Brady (1879) are an unnamed species found in the
Waldron Shale (Hattin, 1960, p. 2016) and A. bastillensis McClel-
lan found here in the Bastille Limestone of early Wenlockian age.
The total stratigraphic range of the ostracodes is shown on
Table 9 and Table 10, each representing one of the two orders to
which the species belong. Most of the species were found only in
strata of Helderbergian age. One of the most commonly occurring
species Phlyctiscapha keslingi Copeland has a greater stratigraphic
range than previously known. It is likely that many other species
would also prove to be more long lived if additional material were
studied in detail. The restricted ranges in Table 9 and Table 10
should be used with caution for that reason. Winchellatia fragilis
McClellan is the youngest known species of the Sigmoopsidae, an
Ordovician family, and its uppermost Silurian occurrence here may
StturRo-DEVONIAN MicroFAuNA: MCCLELLAN 263
SILURIAN LOWER
TABLE; 8
DEVONIAN
Additional Families
(Helderberg
Abundance and
Stratigraphic Range
Wenlock
Ammodiscidae
Glomospira siluriana
Toloypammina tortuosa
Hormosinidae
Aschemonella bastillensis
Lituolidae
Haplophragmoides antiquus
Endothyridae
Nanicella dainae?
Rare: 1-2 (—)
Uncommon: 3-5 (C—)
Common: 6-10 (mz)
Abundant: over 10 ( umm)
be stratigraphically useful. Limbinaria Swartz in Swartz and Whit-
more, 1956 is restricted to upper Ludlow and lower Helderberg
strata in eastern North America and has a similar occurrence here.
Local stratigraphic distribution of the foraminifers and ostra-
codes is shown on Tables 11-22, including columnar sections and
sample localities, From these distributions the previously discussed
composite range charts were compiled. It is evident that the oc-
currences are locally discontinuous for the most part. This may be
due in part to transportation of dead specimens, with the resultant
faunule thus representing a thanatocoenose for most of the species.
This is possible for some of the sample localities in the Willow
Creek Section (Table 19). The ostracodes are even more locally
restricted which clearly limits their biostratigraphic use without
detailed study,
TABLE: 9
SILURIAN
Order PALEOCOPIDA DEVONIAN
Stratigraphic Range Lidlow Helderberg
BEYRICHICOPINA
Aechmina cortezensis
Aechmina equilateralis
Aechmina longior
Aechmina aff, A. phantastica
| Bolbiprimitia teressaccula
t
Hollinella sp.
| Kloedenia aparchoides
L
imbinaria sp.
Myomphalus? sp.
Phlyctiscapha keslingi
a ea
ee a
fcewentes necturie | | =]
Ulrichia obesita Cia Ea
frees
abeynei oe
[Welleropsiejersevensie |
mene een Poa
KLOEDENELLOCOPINA
Eukloedenella aff. E. dalhousiensis areal ene rel a
Parahealdia aff, P, pecorella eS ass
TABLE: 10
Order PODOCOPIDA SILURIAN DEVONIAN
Stratigraphic Range Wenlock Ludlow Helderberg
PODOCOPINA | ae,
Acanthoscapha aff,A. navicula iz —- |
Berounella sp.
Bythocypris aff. B. alcocki
Bythocypris? sp.
Longiscapha nevadensis
METACOPINA
Birdsallella sp.
Thlipsura bispinosa
Tubulibairdia cf,T. chaleurensis
1
T. aff. T. chaleurensis |
Tubulibairdia sp. [Rios |
=.
265
SILURO-DEVONIAN MicroFAUNA: McCLELLAN
jbulisay ees aed x
ePplsajluilweuo4
\ Cee
/OOL O
Buluds youew
i
|
|
ny
im
HHH
"|
INI
i
|
it
|
|
|
=
O
=
E
O
O
=
=
U
STRATIGRAPHIC OCCURRENCE OF
SPECIES AT MARCH SPRING
Table 11:
BULLETIN 274
266
"ds Ze1yasAaqijan
Byisaqo e1ydidin
SISUadNAa|eYd “| “ye elpalequjnqny
Puldsejnw eulsappny
"ds sayiiisend
(Burjsay eydeasiyoKiyd
SISUaISNOUl|ep “FZ “Jye ej jauapaod|ynq
‘ds gsiadAo0uyAg
yoo e “g “sye Siid
2pode4}sQ
Pdsaydsiway eaplojjauiqqam
eyenose PUlWwWeiny{
~TAsO; [ew esaeydsouys0jys
jueWIOW PUlWWweUba,s
éeysaqay eulwweubays
edipuljAD eulwWeuUbays
®}40}U09 eulwweUba}S
‘ds exsaeudsoios
PJE;JUL eLaeYdsSouOS
esnjyuod esaeydsoios
ediipuljAd eulwwepgeyy
edadse eulwwesoqiey
eaed esaeyudsowwesd
2}914}SU09 PUlWWwesadAH
peig eulwwesaeydsiway
Ppyiajlulweso4
{OOL O
uoKkueg say|
iE
R| RI CIR
yaxSeW
mo|pny
yoo, uaM
common
Cc
uncommon,
U
rare,
R
STRATIGRAPHIC OCCURRENCE OF SPECIES AT IKES
Table 12:
CANYON
267
SILURO-DEVONIAN MicroFAuNA: McCLELLAN
Erno Ye nO OG ee
PolJaydsiway eaplojjaulqqam
fUI}}eY4 Caplo|jaulqqom |>.
iAsO]JewW esaeydsoyjyso0}S
ej0ju09 eulwweubays ies ae
epliajiulweso4
| aye
oIeeraeRrS | i
i H il il HH
IItH ¥1qqey
abundant
A
Cc
rare U=
R
,
O
E
=
O
Ut
uncommon,
Table 13: STRATIGRAPHIC OCCURRENCE OF
SPECIES AT RABBIT HILL
‘ds eydeosiyIAlUd
"ds e|/auapao|,0aNn
TED eulmwweubays
common
epliajlulweso4
uncommon, C
0001 fo)
uleyUNOW au0y
rare, U
R
BuLLETIN 274
iKpeag Pienaar
Pplsajylulwmeio4
ommon,
= 5979
uy
HT tit TL
ivaatt i neath
6 , HH ii i ne HHH
stat ttdiataiet vit Tr tReath
uoKkuey) uabeyuadog
R=rare, othe
mojpny
268
OCCURRENCE OF SPECIES AT
Table 15: STRATIGRAPHIC
LONE MOUNTAIN
Table 14: STRATIGRAPHIC
OCCURRENCE OF SPECIES
AT COPENHAGEN CANYON
269
SItuRo-DeEvontan MicroFAuna: McCLettan
SISUaPjaljsseig eraeydsewo}s | «
eiadse PulmWwesoqgiey
PAPD PLAPUCGSOWWeS,
epliaji ulweso4
———
‘ OL
JJl]Ia}eO
&
o
BARD PlaeUdsOWWeSY
epliajiulweso4
aUu0) poom
ule}UNOW au0]
Table 17; STRATIGRAPHIC
OCCURRENCE OF SPECIES
AT GATECLIFF
Mo|pny
i
nce
Table 16: STRATIGRAPHIC
OCCURRENCE OF SPECIES
AT WOOD CONE
BULLETIN 274
270
Eeyeinqnjyispenb eulwweisnu,
Ssluejnbueipenb eulwweubays
Zeysaqay eulWweuba}s
S1SUapoobso esaeyudsosos
edlspuijAd PUlMWepqgeuYy
EIR ae UEC SOUS Ee a evo | me conn
UO UE Be nm
\Apeiq eulWwWwesaeydsiway
common
Pplsajlulweso4
6002
"6003
6004
O
i
oO
ls
is
[@)
0)
(e
5
H] HHH HHI
: Hal i
ays q q 7
Ot ;
HHH HH iH
IE}UNOW S}saqoy
(oS SS
100G O
U
yaad UosSUeH ajad
=rare,
R
STRATIGRAPHIC OCCURRENCE
OF SPECIES AT PETE HANSON CREEK
Table 18:
271
McCLELLAn
S1LtuRO-DEVONIAN MICROFAUNA
erjanbisjuan
eSopou “M ‘sje
edijaydsiway
1U1}}eU
49 Ze UE euES ay
epliajluimeso4
ABO MRO oeaOno HAA
ods 0 wnt! Sv DSHenTHGOgn HANA
Hebets st
49a49 MOIIIM
~
c
uo
U
ie
>}
O
a
S
common, A
R=rare, U=uncommon,
STRATIGRAPHIC OCCURRENCE OF FORAMINIFERS AT
Table 19:
WILLOW CREEK
BuLLeETIN 274
chaleurensis
. B. alcocki
le
Willow Creek
Ostracoda
erditia? sp.
airdia sp.
erounella sp.
Saccarchites saccularis
Winchellatia fragilis
Tubulibairdia cf.
Roberts Mountains
= |/5]/52i|=
X|X|—1X|X]X]| XTX] XI] xX
Table 20: STRATIGRAPHIC OCCURRENCE OF
OSTRACODES AT WILLOW CREEK
DAS:
McCLeLLAn
Stturo-DEVONIAN MICROFAUNA
Sisuakassal sisdoisajjam
Sijesayejinba eulwyodey
B}911}SU0D BUulwwedad
Ppliajlulweso4
SSS
,00S )
uokuey jeoy
ae
s
cn Wnt in
mo|pny
yO; UaM
~
=
oO
5e)
[=
2
Oo
ia)
A
:
C=common
uncommon,
U
rare,
R
STRATIGRAPHIC OCCURRENCE OF SPECIES AT COAL
Table 21
ANYON
r
~
BULLETIN 274
©5980
°598I
uncommon
274
SiSuadnajeyd “| “Jye elpsleqijnaqny
x
x
x
Xx
=i aes
x| xX] Xx] -
DANG ae
S| eles
PjNDIAeU “YW “Jye EYdedsouUedY
Ppliojlulwedso4
|
Table 22: STRATIGRAPHIC OCCURRENCE OF
HHH EIU Ae
ea |
,OOL O
ul
a
THEE
uoAuey Za}109
rare,
R
SPECIES AT CORTEZ CANYON
S1LtuRO-DEVONIAN MicroFAUNA: McCLELLAN 275
SYSTEMATIC PALEONTOLOGY
Order FORAMINIFERIDA Eichwald, 1830
Suborder TEXTULARIINA Delage and Herouard, 1896
Superfamily AMMODISCACEA Reuss, 1862
Family ASTRORHIZIDAE Brady, 1881
Subfamily ASTRORHIZINAE Brady, 1881
Genus RHABDAMMINA MM. Sars in Carpenter, 1868
1869. Rhabdammina M. Sars in Carpenter, Royal Soc. London, Proc., vol. 18
(1868), No. 114, p. 61.
Type species. — Rhabdammina abyssorum M. Sars (1869, zbid.,
poly),
Description. — Test free, tubular, straight or branching with
elongate arms, wall agglutinated, usually well cemented, apertures
at open ends of tubular arms.
This primitive genus, whose species are straight or branching
tubes, is similar to several other genera. The most closely related
is Astrorhiza Sandahl (1858) which also branches but differs in
possessing a central chamber. Linear species of Rhabdammina,
such as R. linearis Brady or R. cylindrica Glaessner, have often
been misidentified, usually as Bathysiphon Sars (1872) species. This
is especially true in studies of Paleozoic foraminifers because of poor
test preservation. Bathysiphon should be partly defined, however,
on the basis of a wall structure consisting of an inner layer of
cemented sponge spicules covered by an outer layer of sand grains,
or other mineral matter, with calcareous or siliceous cement. This
wall structure was well illustrated by Cushman (1910, p. 32) in his
discussion of Recent species of Bathysiphon. The wall structure of
Rhabdammina is clearly shown by the scanning electron micro-
graphs on Plate 33. A related third genus, Marsipella, uses sponge
spicules, sand grains, or other foraminifer tests indiscriminantly in
forming its tests. In contrast, Rhabdammina exclusively employs
mineral grains, such as quartz, in the construction of its test. It is,
therefore, probable that many identifications of Bathysiphon, based
on completely arenaceous structured material, should be referred to
Rhabdammina.
276 BULLETIN 274
Rhabdammina bifurcata Browne and Schott Pl. 33, figs. 8, 9; Pl. 39, fig. 4
1963. Rhabdammina bifurcata Browne and Schott, Bull. Amer. Paleont., vol.
46, No. 209, p. 198, pl. 48, figs. 4, 5.
Description. — Test free, consisting of three nearly straight,
branching, tubular arms united in a single plane to form a Y-shaped
test, lower arm of the Y normally being longer than the other two;
wall finely arenaceous and well cemented; apertures at open ends
of tubes.
Measurements. — Dimensions of figured hypotypes as follows:
Test Length Diameter Short Arm Long Arm
34682 0.22 mm 0.07 mm 0.11 mm 0.18 mm
34683 indet. 0.11 mm 0.21 mm indet.
Occurrence. — This species was found only at locality UWA
5991, where it is a rare element of the fauna. Its only other known
occurrence is in the Osgood Formation of Indiana of Llandovery,
or lowest Niagaran age, and its presence in Nevada extends the
range of the species to the lowest Devonian.
Rhabdammina cylindrica Glaessner Pl. 33, figs. 10, 11; Pl. 40, fig. 4
1937. Rhabdammina cylindrica Glaessner, Moscow Univ. Lab. Paleont., Prob.
Paleont., vol. 2-3, p. 354.
1941. Bathysiphon exiguus Moreman, Stewart and Priddy, Jour. Paleont.,
vol. 15, No. 4, p. 370, pl. 54, fig. 7.
1942. Bathysiphon exiguus Moreman, Dunn, Jour. Paleont., vol. 16, No. 3,
p. 322, pl. 42, fig. 27.
1961. Bathysiphon exiguus Moreman (in part), Mound, Indiana Dept. Con-
serv., Geol. Sur., Bull. No. 23, p. 36, pl. 3, figs. 17-20.
1963. Bathysiphon exiguus Moreman (in part), Browne and Schott, Bull.
Amer. Paleont., vol. 46, No. 209, p. 232, pl. 52, fig. 7, 8.
1966. Bathysiphon exiguus Moreman, McClellan, Bull. Amer. Paleont., vol.
50, No. 230, p. 462, pl. 36, figs. 1-5; pl. 40, figs. 1-5.
Description. — Test apparently free, straight cylindrical tube;
wall finely arenaceous, poorly to well cemented; aperture at open
ends of tubular test.
Measurements. — Hypotype 34684 diameter 0.09 mm, length
0.84 mm; 34685 diameter 0.11 mm, length 0.67 mm.
Occurrence. — Widespread throughout the area; present in
varying degrees of abundance at all stratigraphic horizons within the
several limestone formations. Figured specimens from UWA 5973.
Discussion.—The species Bathysiphon exiguus Moreman
SILURO-DEVONIAN MicroFAuNA: McCLELLAN Bt
(1930) has frequently been applied to specimens having a straight,
cylindrical or tubular test consisting of apparently arenaceous
material. Moreman’s description (p. 46) of this species mentioned
the character of a slightly tapering test. Possibly Moreman’s speci-
mens should also be referred to the genus Rhabdammina on the
basis of wall material. If so, they would be distinct from the species
R. cylindrica Glaessner (1937) by virtue of the slight taper of the
test. The species named by Glaessner is limited to cylindrical, non-
tapering forms. Subsequent authors have identified material, which
is cylindrical and nontapering, as Bathystphon exiguus. Mound
(1961) and Browne and Schott (1963) included a variety of forms
under the latter species, some of which can be referred to the
former R. cylindrica. McClellan’s (1966) material contained one
specimen (U.C. 37783) which is properly a Bathysiphon but should
be assigned to another species due to the constrictions present on
the test.
Rhabdammina major de Folin IAL OB y. 30 0h al|
1887. Rhabdammina major de Folin, Naturaliste, Paris, vol. 9, No. 2, p. 127
(fetae).
Description. — Test free, large, cylindrical, twisted, branching
tube; wall composed of single layer of fine to medium sand grains
with moderate cement; apertures at open ends of branching tubes.
Measurements. — Hypotype 34686 length 1.63 mm, diameter
0.33 mm, branch length 0.37 mm,
Occurrence. — This species is present only at locality UWA 6022
in shaly limestones. It has not previously been reported from the
Silurian.
Discussion. — The Y pattern of branching in Rhabdammina
bifurcata is distinctive from the type of branching in R. major, and
the twisted nature of most specimens of the latter is also a distinc-
tive characteristic. Previously, R. major has been recorded as a
Recent species, but it is found here in rocks of Middle Ludlow age.
Subfamily RHIZAMMININAE Rhumbler, 1895
Genus MARSIPELLA Norman, 1878
1878. Marsipella Norman, Ann. Mag. Nat. Hist., ser. 5, vol. 1, p. 281.
Type species. — Marsipella elongata Norman (1878, ibid., pl.
16).
278 BULLETIN 274
Loeblich and Tappan (1964, p. 186) redescribed the genus as
follows:
Test free, consisting of undivided tubular, cylindrical, or elongate-
fusiform chamber, which may be slightly twisted; wall of agglu-
tinated sand, sponge spicules, or tests of other foraminifers; aper-
tures at open ends of tube.
The above description is more in keeping with modern generic
concepts than the original of Norman which appears to be re-
stricted to the type species. An important feature is the variety of
wall material allowable in this primitive foraminifer. While Mar-
stpella elongata used sand grains and sponge spicules, several species,
including those in the Paleozoic, used only the sand grains, This
suggests a close relationship to Rhizammina Brady (1879) which
uses the same materials in two layers, but whose test is not com-
monly twisted and fusiform. Bathysiphon Sars (1872) shows more
specialization, consistently having an inner layer with spicules and
an outer one of sand grains.
Marsipella torta Stewart and Priddy Pl33ietien7
1941. Marsipella? torta Stewart and Priddy, Jour. Paleont., vol. 15, No. 4,
p. 370, pl. 54, fig. 4.
Description. — Test free, cylindrical, twisted tube, tapering
toward apertural end; wall thin, medium to finely arenaceous, mod-
erately cemented; aperture terminal at end of tube, additional
apertures not presently known from incomplete specimens.
Measurements. — Length of the figured hypotype 34673 is 1.16
mm; larger diameter 0.30 mm, smaller, apertural diameter 0.18
mm.
Occurrence. — This species occurs rarely at two localities repre-
senting a time span from middle Ludlow to Helderberg age. It has
previously been reported from the Osgood Formation of Indiana,
which is earliest Niagaran, or middle Llandovery. Hypotype from
locality UWA 6022.
Discussion. — Stewart and Priddy, in discussion of the generic
position of this species, believed that it was more twisted than nor-
mal for the genus, and that it lacked sponge spicules in the test.
Because the wall structure is known to be variable, their generic
classification seems correct. Specimens recovered from the Nevada
material are somewhat damaged so that no example adequately
demonstrates the complete variability of the species. The tapering
SILURO-DEVONIAN MicroFAUNA: MCCLELLAN 279
test is characteristic and serves to separate specimens from un-
branched examples of Rhabdammina major.
Marsipella sp. Ply soy tiga
Description. — Test free, subcylindrical, moderately contorted,
tubular; wall fine to medium sand grains poorly cemented; aper-
tures at ends of tube though poorly defined due to damaged tests.
One end with narrow lip at aperture.
Measurements. — Length of figured specimen 34674 is 1.09 mm,
diameter of undamaged portion 0.12 mm.
Occurrence. — Rarely present at locality UWA 5973 in strata of
Helderbergian age. Similar individuals referred to this genus have
been reported from the Middle Silurian Waldron Shale in Indiana
by Hattin (abstract, 1960, p. 2016).
Discussion. — The two specimens recovered here both have one
end of the test flattened, apparently by external forces, so that the
original form is indeterminate. The test is less contorted than that
of M. torta, the wall composition less consistent, and there is little
cement. What appears to be minute openings of the test surface
may be due to the poor cementation.
Subfamily HIPPOCREPININAE Rhumbler, 1895
Genus HYPERAMMINA Brady, 1878, emend. Conkin, 1961
1878. Hyperammina Brady, Ann. Mag. Nat. Hist., ser. 5, vol. 1, pp. 433-434.
Type species. —Hyperammina elongata Brady (1878, ibid., pl.
20s tig. Zab).
The generic concept of Hyperammina that Conkin first dis-
cussed (1954, pp. 167-168) and later (1961, p. 254) proposed as an
emended description is that which is followed herein.
Description. — Test free, arenaceous; with an elongate tubular,
singular or branching second chamber, which may be nontapering,
may taper toward the proloculus, or in a few species taper toward
both the aperture and proloculus; aperture open or constricted
slightly to strongly; interior smooth; exterior rough, may be marked
by transverse constrictions of varying strength.
Hyperammina casteri Conkin Pl. 33, figs. 2, 6
1961. Hyperammina casteri Conkin, Bull. Amer. Paleont., vol. 43, No. 196,
p. 260, figs. 6, 7; pl. 20, figs 1-18; pl. 26, figs. 7, 8.
280 BULLETIN 274
Description. — ‘Test free, proloculus shape variable, usually
oblate to spherical, followed by straight second chamber which
expands gradually from constriction near proloculus to diameter
greater than the proloculus; wall thin to thick, finely arenaceous,
well cemented; aperture terminal on second chamber. Microspheric
forms have a small, pointed proloculus and rapidly expanding
second chamber giving the test the appearance of an elongated
cone.
Measurements. — Microspheric specimen 34670 has a_ test
length of 0.58 mm, proloculus diameter of 0.06 mm, apertural end
diameter of 0.16 mm. Specimen 34669 has broken second chamber
and indeterminate length, but proloculus diameter is 0.18 mm.
Occurrence. — This species is poorly represented here in Mid-
dle Silurian strata at only two localities in the northern Roberts
Mountains (UWA 5989, UWA 6024).
Discussion. — Megalospheric forms of H. casteri are similar to
those of H. glabra Cushman and Waters (1927), but the proloculus
diameter is larger in relation to the second chamber than in the
latter species. Conkin (1961, pp. 261-263) presented a thorough
discussion of the affinities of H. casteri to other species with which
the writer is in agreement. This species was first reported from
earliest Mississippian strata where it is abundant in dark shales,
silts, and calcareous shales. McClellan (1966, p. 460) found it to
be dominant in lower energy environments of the Waldron Shale.
In the present material it occurs in argillaceous and cherty lime-
stone beds.
Hyperammina constricta Gutschick and Treckman Pl. 33. fig. 4
1959. Hyperammina constricta Gutschick and Treckman, Jour. Paleont., vol.
33, Now2:5ps 237, pls 34, figs. 117-19) text fig. lamin:
Description. — Test free, proloculus spherical; second chamber
long, straight or slightly curved, bearing regularly spaced, slight
constrictions; wall thin, finely arenaceous, well cemented; aperture
terminal opening on second chamber. Microspheric stage not present
in this material.
Measurements. — Figured hypotype 34671 has length of 0.76
mm, maximum diameter 0.13 mm, minimum diameter 0.10 mm at
constrictions.
S1LURO-DEVONIAN MicroFAuNA: McCLELLAN 281
Occurrence. — Present only in rocks of lowest Devonian age
at localities UWA 5974 and UWA 5985.
Discussion. — Previously this species has been known only
locally from Silurian and Mississippian rocks in Indiana. Its presence
in Nevada indicates a wider geographic occurrence. In every case
it is found in relatively clean limestones indicative of a preference
for higher energy conditions than those of some hyperamminids,
such as H. casteri which preferred a quiet, muddy environment, and
H. rockfordensis (Conkin, 1961, pp. 270-272).
Hyperammina curva (Moreman) Pim3o. figs 5
1930. Bathysiphon curvus Moreman, Jour. Paleont., vol. 4, No. 1, p. 45, pl.
Seatipsee9 pelOs
1961. Hyperammina curva (Moreman), Mound, Indiana Dept. Conserv.,
Geol: Sur., Bull. No, 23, p: 35, pl. 3, figs. 13-16.
Description. — Test free, with megalospheric proloculus small
and bluntly rounded, microspheric pointed; second chamber long,
gently tapering, and slightly curved with no constrictions; wall
thin, finely arenaceous, well cemented; terminal opening of second
chamber serves for an aperture.
Measurements. — Specimen 34672 has a broken second cham-
ber; remaining length is 0.44 mm, diameter 0.15 mm.
Occurrence. — This is the most common Hyperammina species
known from these rocks in Nevada. It was recovered from several
horizons, indicating a presence throughout the entire sequence of
strata studied here. The stratigraphic range of H. curva is here
extended at least into the lowest Devonian by its occurrence in the
Rabbit Hill Limestone. Figured specimen from locality UWA 6024.
Discussion. — The original type specimens have the proloculus
missing, hence Moreman’s mistaken identification as a species of
Bathysiphon. However, they are well tapered as in Hyperammina.
Mound (1961, p. 35) recovered a large number of specimens from
the Brassfield Limestone of Indiana which compared closely with
Moreman’s types and also possessed proloculi.
Both of these collections are from rocks of Llandovery age in
which the species is more abundant than in the younger rocks of
central Nevada. Ecologically the Brassfield is probably similar to
the silty limestone and argillaceous limestone of the Roberts Moun-
tains Formation and Rabbit Hill Formation. Other occurrences of
282 BULLETIN 274
H. curva (Ireland 1939; Dunn, 1942), in the Illinois Basin, are in
more pure carbonates, but the abundance of the foraminifer is not
stated in these instances.
Family SACCAMMINIDAE Brady, 1884
Subfamily PSAMMOSPHAERINAE Haeckel, 1894
Genus PSAMMOSPHAERA Schulze, 1875
1875. Psammosphaera Schulze, Komm. Untersuch. deutsch. Meere in Kiel,
Jahresber., vol. 1872-73, p. 113 (fetae).
Type species. —Psammosphaera fusca Schulze (1875, zbid.).
Cushman’s (1918, p. 34) redescription is given below:
Test free or attached, single chambered, usually spherical, no definite
aperture, the pseudopodia making their way out through the interstitial
openings between the elements of the test; wall of sand grains, mica
flakes, sponge spicules, or other foraminiferal tests firmly cemented.
This genus, with its simple, nonspecialized form, is one of the
most primitive foraminifers known and is a common constituent of
arenaceous foraminiferal faunas from the Middle Ordovician to
Recent. It is found in almost every environment which agglutinated
foraminifers have inhabited. In the Silurian and Lower Devonian
of Nevada Psammosphaera has been found in rocks with lithologies
varying from calcareous, dark, organic shales to reefoid dolomites.
In several instances it is the only foraminiferal genus represented.
This situation may be incurred by specimen transport into a
thanatocoenose, but tubular and polythalmous, spherical genera
would also be subject to the same transportation if such were the
case. It appears more likely that Psammosphaera was more primitive
and able to tolerate a greater variety of ecologic conditions. At the
same time, the genus is often the most abundant one in a well-
developed, diversified faunule. A possible explanation for this
situation may be a lack of intensive competition among the foram-
inifers due to the small number of individuals present.
Psammosphaera cava Moreman Pl. 34, fig. 21
1930. Psammosphaera cava Moreman, Jour. Paleont., vol. 4, No. 1, p. 48, pl.
(Gh, tier, TA
Description. — Test free, spherical; wall thick to thin, outer
layer of medium to coarse sand grains and inner pseudochitinous
layer, well cemented but not an excess on the surface; no apparent
aperture.
SILURO-DEVONIAN MicroFAuNA: MCCLELLAN 283
Measurements. — Hypotype 34679 has a diameter of 0.25 mm.
Occurrence. — Widespread, generally abundant relative to
other foraminifers in these rocks. Specimen 34679 at locality UWA
6010.
Discussion. — Spherical test shapes with a single arenaceous
layer over an inner pseudochitinous one, such as P. cava, are prone
to deformation after death of the animal, Therefore, a wide range
of variation in shape may result, all of which represent the same
species. Several new species have been erected by authors for these
shapes. McClellan (1966, pp. 468-470) reviewed the synonymies
which are referable to Psammosphaera cava.
Psammosphaera laevigata White Pl. 34, fig. 26; Pl. 39, fig. 1
1928. Psammosphacra laevigata White, Jour. Paleont., vol. 2, No. 3, p. 183,
pl. 27, figs. 1a,b.
Description. — Test free, spherical, wall finely arenaceous with
abundant cement, surface appearing smooth and silky; no aperture
apparent, other than small, interstitial pseudopodia openings.
Measurements. — The diameter of the figured specimen is 0.24
mm.
Occurrence. — This species has been recorded as a minor faunal
element from the Lower Silurian of Indiana and is similarly present
here in the lower part of the Roberts Mountains Formation at
locality UWA 6031.
Discussion. — White’s type material appears to have been flat-
tened. This flattening is not uncommon with Paleozoic species of
Psammosphaera, but the examples here do not show similar distor-
tion. The fine wall texture and abundant cement makes them readily
distinguishable from specimens of P. cava,
Genus ANICTOSPHAERA McClellan, n. gen.
Type species. — Anictosphaera progressa McClellan, n. sp.
Description. — Test free, multilocular, globular to spherical
chambers closely appressed to loosely joined, broad openings be-
tween chambers; wall consists of single layer of agglutinated sand
grains, well cemented; no definite aperture.
The most important feature of this genus is the presence of
broad internal openings between the chambers, hence the origin
of the name (Greek, anictos = open). The test has several cham-
284 BULLETIN 274
bers and no aperture, features which characterize many Psam-
mosphaerinae. Other members of the subfamily which are poly-
thalmous have no visible openings between the chambers. There is
some indication of multilocular development in Storthosphaera
Schulze (1875) indicated by the outward projections from the cen-
tral chamber, which give the appearance of incipient chambers
formed by an infolding of the test wall. Such “pseudochambers” are
not evident on Antctosphaera; instead, well-developed chambers
bearing recognizable sutures are readily observable.
Arenosphaera Shchedrina (1939) appears similar to the type
species Anictosphaera progressa, but close inspection reveals the
multiple apertures between the agglutinated grains of the wall.
Thus, Shchedrina’s genus belongs in the subfamily Saccammininae.
All other characters of the two genera point to a close phylogenetic
relationship.
Anictosphaera progressa McClellan, n. sp. Pl. 34, figs. 1-5
Description. — Test free, multilocular, subspherical chambers
tightly appressed with rectilinear boundary bearing a generally
deep suture; presently known only as bilocular forms which have
one chamber larger than the other; internal opening between cham-
bers wide and circular; wall thin, medium to finely arenaceous,
well cemented; no aperture known.
Measurements. — Diameters of the smaller chamber varies
from 0.08 mm to 0.19 mm. Larger chamber diameter ranges from
0.22 mm to 0.27 mm. The ratio of diameters between the cham-
bers is most often approximately 2:1. Wall thickness is fairly
constant at 0.02 mm.
Small Larger
Pl. 34 Spec. Number Chamber Chamber
Rig, 1 34651 paratype 0.19 mm 0.27 mm
Fig. 2 34652 paratype 0.08 mm 0.17 mm
Fig. 3 34653 paratype 0.14 mm 0.27 mm
Fig. 4 34654 holotype 0.10 mm 0.24 mm
Fig. 5 34655 paratype 0.11 mm 0.22 mm
Occurrence. — This species is present in rocks from the middle
Roberts Mountains Formation to Rabbit Hill Formation. This
SILURO-DEVONIAN MiIcrRoFAUNA: McCLELLAN 285
represents a range of middle Upper Silurian (Ludlow) to lowest
Devonian age. Specimen 34651 from locality UWA 6020; 34652 and
34653 from UWA 6010; 34654, and 34655 from UWA 6009.
Discussion. — The 2:1 ratio between the two chambers occurs
commonly but might represent microspheric individuals, The para-
type 34651, has a ratio of 3:2 that may indicate a megalospheric
form. It is believed that this represents a particularly robust in-
dividual, and there is insufficient material available to seriously
consider the possibilty of dimorphism. No. 34653 does not have a
strong suture between the chambers, but this is most probably only
an individual variant in this respect, though it was first believed
to be a dimorphic form.
The most common occurrence of Anictosphaera progressa is in
thin-bedded argillaceous limestones, but it is also present in the
more nearly pure carbonates as well, From the known occurrences
and associations it does not appear that any significant ecologic
interpretations can be made.
Genus CERATAMMINA Ireland, 1939
1939. Ceratammina Ireland, Jour. Paleont., vol. 13, No. 2, p. 194.
Type species. —Ceratammina cornucopia Ireland (1939, ibid.,
p. 196, figs. A-31, 32).
The description given by Ireland (p. 194) is as follows:
Test free, horn-shaped; wall composed of fine sand grains; surface
smooth; aperture not apparent.
Ceratammina cf. C. cornucopia Ireland Pl. 34, fig. 25
Description. — Test free, near horn-shaped but only slightly
curved, apparent constriction near larger end of horn; wall thin,
finely arenaceous, well cemented; no apparent aperture.
Measurements. — Length of figured specimen is 0.26 mm,
larger diameter 0.17 mm, smaller diameter 0.11 mm.
Occurrence. — This species is rarely present at locality UWA
6017 of uncertain stratigraphic position in the Upper Silurian. Ire-
land (1939, p, 196) reported it as a distinctive Lower Devonian
form.
Discussion. — The referred specimen is not well preserved,
showing a fracture along one side of the test. The constriction near
the larger end may be externally induced, and also be partly re-
286 BULLETIN 274
sponsible for the indistinctness of the horn shape. Because of this
damage the specimen cannot be definitely determined as a Cera-
tammina cornucopia.
Genus Raibosammina Moreman, 1930
1930. Raibosammina Moreman, Jour. Paleont., vol. 4, No. 1, p. 50.
Type species. — Raibosammina aspera Moreman (1930, 1bid.,
pl. 6, figs. 13-15), designated lectotype.
The genus, as originally designated, included forms which have
subsequently been considered as species of Stegnammina Moreman
(1930), including the original type R. mica Moreman. Loeblich
and Tappan (1964, p. 196) included all Razbosammuna species
within Stegnammina. McClellan (1966, p. 474) discussed the rela-
tionships of these genera and suggested that Raibosammina aspera
might well be better considered as generically separate from other
Stegnammina species. It is, therefore, necessary to emend the
original description of Moreman which did not restrict the genus
to only branching forms. The following emendation is considered
necessary:
Test free, subcylindrical, chamber may be crooked, irregularly branch-
ing; wall of variable thickness, arenaceous, poorly to well cemented, no
aperture apparent.
As thus described the genus is monotypic and Ratbosammina aspera
is, therefore, designated as typical.
Raibosammina is closely related to Stegnammina which has
several similar subcylindrical species, but it differs from the latter
in having a branching test. The only other species previously identi-
fied as a Raibosammina is Stegnammina mica (Moreman), a sub-
cylindrical, nonbranching, monothalmous form,
Raibosammina aspera Moreman PI-33, fis, 125°Pi-- oo) tee
1930. Raithosammina aspera Moreman, Jour. Paleont., vol. 4, No. 1, p. 50, figs.
13-15.
1964. Stegnammina aspera (Moreman), Loeblich and Tappan, Treat. Invert.
Paleont., Pt. C, Protista 2, vol. 1, p. 196
Description. — Test free, subcylindrical and branching, ends
bluntly terminated; wall of poorly sorted sand grains, moderately
cemented; no apparent aperture, numerous pseudopodia openings
on surface.
S1LtuRO-DEvVoNIAN MicroFAuNA: McCLeELLAn 287
Measurements. — Figured hypotype 34681 has diameter of 0.29
mm and length of 0.71 mm.
Occurrence. — Uncommon at localities UWA 5985 and UWA
5996.
Discussion. — This species has been recognized in rocks from
Upper Ordovician to Lower Devonian age. It does not appear to
have obvious habitat preferences nor to occur in abundance at any
time during its known geologic range. It has a fairly wide geographic
distribution, being present in faunas from Indiana, Oklahoma, and
Nevada.
Genus SOROSPHAERA Brady, 1879
1879. Sorosphaera Brady, Quart, Jour. Micr. Sci., vol. 19, new ser., pp. 28-29.
Type species. — Sorosphaera confusa Brady (1879, ibid., pl. 4,
fras. 18% 19 )t
Brady’s original description follows:
Test free, irregular; consisting of a number of convex or spheroidal
chambers, either discrete or more or less embracing, irregularly crowd-
ed together. Walls thin, loosely arenaceous in texture. General aper-
ture, none. Long diameter of large specimens, 1/6 inch (4.5 mm.)
Several species of this genus have been erected on the basis of
differing number of chambers. It can readily be demonstrated that
these species are all within the expected range of variation of a
single species. The difference in number of chambers may be due
to breakage and separation, or ontogeny. Grubbs (1939, p. 544)
suggested the former and Hattin (personal communication) found
a complete ontogenetic sequence of the genus which suggests the
latter, in the Waldron Shale. The writer concurs with these opinions
and has here considered two and three chambered forms as frag-
ments or neanic individuals of species having a greater number
of chambers, In the case of an isolated chamber it is impossible
to recognize Sorosphaera from Psammosphaera cava unless a frag-
ment of a second chamber is present.
Sorosphaera confusa Brady Pl. 34, fig. 24
1879. Sorosphacra confusa Brady, Quart. Jour. Micr. Sci., vol. 19, new ser.,
oe Ay Rk Ze sitery ain ai)
Description. — Test free, multilocular, consisting of a variable
number of subspherical chambers connected closely in a random
manner with adjoining walls somewhat appressed, entire test having
288 BULLETIN 274
a massive appearance; wall thin, usually finely arenaceous, well
cemented; no apparent aperture.
Measurements. — Chamber diameters of the hypotype 34687
are 0.29 mm, 0.43 mm.
Occurrence. — Infrequently present at several localities and
stratigraphic levels. Hypotype from locality UWA 6030.
Discussion. — Sorosphaera confusa has been previously recog-
nized throughout strata of Silurian age, and here is found in rocks
of lowest Devonian age. None of these occurrences are in abundance.
Though typically occurring in argillaceous or silty limestones, it has
also been found in dolomitic limestones.
Sorosphaera osgoodensis Stewart and Priddy
Pl. 34, figs. 15, 16; Pl. 39, fig. 6
1930. Sorosphaera tricella Moreman, Jour. Paleont., vol. 4, No. 1, p. 49, pl. 5,
figs. 12, 14.
1932. Sorosphaera tricella Moreman, Croneis, et al., Science, vol. 75, No. 1935,
pl. 138.
1941. Sorosphaera osgoodensis Stewart and Priddy, Jour. Paleont., vol. 15,
No. 4, p. 371, pl. 54, fig. 11.
1942. Sorosphaera bicella Dunn, Jour. Paleont., vol. 16, No. 3, p. 325, pl. 42,
figs. 17, 18.
1942. Sorosphaera multicella Dunn, ibid., p. 325, pl. 42, figs. 19a, b.
1942. Sorosphaera tricella Moreman, Dunn, ibid., p. 324, pl. 42, fig. 15.
1947. Sorosphaera bicelloidea Stewart and Lampe, Jour. Paleont., vol. 21,
No. 6, p. 534, pl. 78, fig. 6.
1958. Sorosphaera bicella? Dunn, Summerson, Jour. Paleont., vol. 32, No. 3,
Pao sieepla Si, figs 13.
1958. Sorosphaera bicelloidea Stewart and Lampe, Summerson, ibid., p. 81,
fig. 14.
1961. Sorosphaera bicella Dunn, Mound, Indiana Dept. Consery., Geol. Sur.,
Bull. No. 23, p. 33, pl. 3, figs. 4-6.
1961. Sorosphaera tricella Moreman, Mound, ibid, p. 34, pl. 3, figs. 7-10.
1963. Sorosphaera bicella Dunn, Browne and Schott, Bull. Amer. Paleont.,
vol. 46, No. 209, p. 211, pl. 49, fig. 16.
1963. Sorosphaera tricella Moreman, Browne and Schott, ibid., p. 212, pl. 49,
fia oe
1966. Sorosphaera bicella Dunn, McClellan, Bull. Amer. Paleont., vol. 50, No.
230, p. 472, pl. 37, 41, figs. 7-9.
1966. Sorosphaera tricella Moreman, McClellan, ibid., pl. 37, 41, fig. 10.
Description. — Test free, multilocular, consisting of any num-
ber of spherical to subspherical chambers, usually joined and always
in a single plane which may be somewhat distorted; wall thin,
fine to coarsely arenaceous, moderately cemented; no apertures ap-
parent, infrequent minute and poorly defined pseudopodia openings
present.
StLtuRO-DEVONIAN MicroFAuNA: McCLeELLAN 289
Measurements. — Four-chambered figured specimen 34688 has
chamber diameters of 0.15 mm, 0.15 mm, 0.16 mm, 0.13 mm. Speci-
men 34790 has diameters of 0.15 mm, and 0.13 mm.
Occurrence. — This species, though not widespread or abundant
in Nevada, is present at several horizons. In Silurian rocks and
Devonian rocks of other areas from which it has been reported, it
also occurs in modest numbers. Hypotypes from locality UWA
6018.
Discussion. — The several synonymous species listed above con-
sist of individuals described as having two or three chambers and
are here considered invalid. The figured specimens show that
breakage readily occurs by separation between chambers. Speci-
men 34688 has four complete chambers and a large part of a fifth
preserved. Specimen 34790 has only a small part of a third chamber
remaining, The writer concludes that such accidental breakage is
responsible for the several species names applied by previous
authors.
The inclusion of two-chambered and three-chambered forms in
the species S. osgoodensis is sometimes difficult to accomplish be-
cause the chambers are attached in a single plane. This is especially
true with two-chambered specimens when there are few individuals
in a sample. S. osgoodensis specimens having only two chambers
may be confused with S. irregularis Grubbs (1939) or S. subcon-
fusa Dunn (1942). There presently is some doubt as to the validity
of the latter two species, and they may well be S. osgoodensis. The
chart below illustrates their relationships.
S. osgoodensis S. irregularis S. subconfusa
1) Loosely joined, Chambers closely Closely joined,
spherical chambers joined, spherical subspherical chambers
2) Growth-planar Growth-irregular Massive, irregular
appearance
3) Well cemented Very well cemented Well cemented
(this feature has questionable specific validity)
290 BULLETIN 274
Sorosphaera inflata McClellan, n. sp. Pl. 34, figs. 11, 12; Pl. 40, fig. 1
Description. — Test free, multilocular, chambers subspherical,
presently known only as bilocular with one chamber two or three
times larger than the other, chamber wall appressed where closely
joined, with external suture moderate to deep, no openings between
chambers; wall thin, medium to finely arenaceous, poor to well
cemented; no aperture known.
Measurements. — Paratype 34689 has smaller chamber diameter
0.15 mm, larger chamber 0.30 mm, and wall thickness 0.05 mm.
Holotype 34690 with smaller chamber diameter 0.08 mm, larger
chamber 0.27 mm, wall thickness 0.02 mm.
Occurrence. — This species is rare, occurring at locality UWA
5976 and locality UWA 5987. Both localities are probably of latest
Ludlovian age.
Discussion. — The distinctive feature of Sorosphaera inflata is
the pronounced inequality of size between the two chambers with
subequal diameters. Loeblich and Tappan (1964, p. 196) included
Arenosphaera perforata Shchedrina (1939) as a species of Soro-
sphaera. This, then, would appear to be a closely related species
because it has similar inequality in chamber size. However, A.
perforata has an opening between chambers and multiple openings
in the test wall, and thus actually should be considered a genus in
the Saccammininae. If this is true, S. inflata, which has no recog-
nizable openings between chambers or in the exterior wall, would
have no known close relatives.
Only two-chambered forms are currently known, but it is likely
that specimens with more chambers may be found in the future.
Although ecologic considerations of this new species must of
necessity be limited by the quantity of material and its limited
geographic extent, the two occurrences are in medium-bedded, fine-
grained limestones. One of these bears common crinoidal fragments.
The lack of argillaceous material might suggest somewhat higher
energy conditions.
Sorosphaera compacta McClellan, n. sp. PI. 34, figs. 13, 14; Pl. 40, fig. 2
Description. — Test free, massive, multilocular, subrounded to
subangular, chambers closely packed in an irregular mass and
nearly indistinct externally, adjoining chambers tightly appressed
Situro-DEVONIAN MicroFAuNA: McCLeELLAaNn 291
with linear sutures faint to nonexistent; wall thin, medium to finely
arenaceous, moderately cemented; no apparent apertures.
Measurements. — Holotype 34691 has chamber diameters of
0.18 mm, 0.19 mm, 0.14 mm. Paratype 34692 has diameters of
0.11 mm, 0.13 mm, 0.17 mm.
Occurrence. — This species is present in the Lone Mountain
Formation and Roberts Mountains Formation, with a stratigraphic
range through the Upper Ludlow. Holotype 34691 from locality
UWA 5974 and 34692 from locality UWA 5992.
Discussion. — Sorosphaera confusa is the most closely related
species to S. compacta, but the latter is more massive in appearance
and has more closely joined chambers. In the holotype, chambers
can only be recognized because of the broken test. The figured para-
type has a slightly depressed suture separating one of the three
chambers. The chambers can also be externally recognized on an
unfigured paratype, 346791 which has two partially damaged cham-
bers.
Sorosphaera compacta was recovered from samples without
silt or mud. This would seem to indicate a preference for relatively
higher energy conditions. Associated invertebrates include brachio-
pods, Bryozoa, corals and crinoids in abundance. The microfauna
is usually quite diversified and includes ostracodes, conodonts and,
in one instance, tentaculitids. This fauna has been preserved in
limestones and limey dolomites suggestive of well aerated waters
having bottom currents.
Sorosphaera sp. Pl. 34, fig. 20
Description. — Test apparently free, multilocular, chambers
subspherical where not badly damaged, overall test configuration
indeterminate; wall thin, composed of fine sand grains well
cemented; aperture unknown.
Measurements. — Diameter of measurable chamber 0.11 mm,
other chamber diameter is indeterminate.
Occurrence. — Present only at locality UWA 5985.
Discussion. — Though the specimen is partially broken and one
chamber much distorted, this species is not referable to S. confusa
which also occurs at the same locality, for the well-developed cham-
ber is more nearly spherical and discrete than those of that species.
292 BULLETIN 274
S. confusa and this species are the only sorosphaerids present in the
McMonnigal Limestone.
Genus STEGNAMMINA Moreman, 1930, emend. McClellan, 1966
1930. Stegnammina Moreman, Jour. Paleont., vol. 4, No. 1, p. 49.
Type species. — Stegnammina cylindrica Moreman (1930, tbid.,
pHAgs pli tie; a2):
The emended description of McClellan (1966, p. 475) is given
below.
Test free, monothalmous, cylindrical to angular, straight to curved;
wall thin to thick, agglutinated, poorly to well cemented; aperture not
apparent.
Stegnammina contorta McClellan Pl. 34, fig. 22
1966. Stegnammina contorta McClellan, Bull. Amer. Paleont., vol. 50, No. 230,
p. 476, pls. 36, 40, figs. 17, 18a, b.
Description. — Test free, monothalmous, cylindrical and twist-
ed, ends rounded, chamber tubular and narrow; wall thick, fine to
medium arenaceous grains, poorly cemented; no aperture apparent.
Measurements. — Length of figured hypotype 34696 is 0.57
mm and diameter 0.12 mm.
Occurrence. — Present in small numbers at several localities
representing a general Ludlow age. Specimens at locality UWA 6010,
which is very near the probable Silurian-Devonian boundary, are
the latest known occurrence for the species. Hypotype 34696 is
from locality UWA 6024.
Discussion. — Stegnammina contorta evidently has no strong
preference for a particular energy level of its habitat. It has most
frequently been recovered from sediment consisting of mixed argil-
laceous and calcareous material, The species is also present in lime-
stones with little argillaceous material, though there is some indica-
tion that this is not the preferred environment.
Stegnammina cylindrica Moreman Pl. 34, fig. 18
1930. Stegnammina cylindrica Moreman, Jour. Paleont., vol. 4, No. 1, p. 49,
yoNk: i, sated. 1.
1942. Stegnammina cylindrica brevis Dunn, Jour. Paleont., vol. 16, No. 3,
1. 325, pl. 42, fig. 25.
Description. — Test free, monothalmous, cylindrical, ends flat-
tened; wall thin, finely arenaceous, well cemented; no definite aper-
ture, but smal] pseudopodia openings present on test surface.
StLtuRo-DEvoNIAN MIcRoFAUNA: MCCLELLAN 293
Measurements. — Figured specimen 34697 measures 0.22 mm
in length and 0.13 mm in diameter.
Occurrence. — This species was found at several localities in
the middle Roberts Mountains Formation, or its equivalent, repre-
senting middle Ludlow time. Though well known elsewhere from
older Silurian strata it has not been recognized in the older units
within this area. Presence of this species at locality UWA 5978
lends paleontological support to the assignment of these beds to the
Roberts Mountains Formation rather than to the Hanson Creek, as
suggested in the stratigraphic discussion previously. The figured
hypotype is from locality UWA 6017.
Discussion. — The length of Stegnammina cylindrica has been
shown by Mound (1961, p. 25) to range from 0.17 mm to 0.50 mm.
The subspecies, S. cylindrica brevis Dunn supposedly varies from
Moreman’s type in being only half as long. However, it is still well
within the range determined by Mound for S. cylindrica specimens.
The occurrence noted above is the youngest yet known for the
species sensu stricto. Mound (1961, p. 25) and Browne and Schott
(1963, p. 210) considered Stegnammina elongata Ireland (1939) as
synonymous with S. cylindrica. The writer believes that S. elongata
presently is not well enough known to be so placed taxonomically.
The main reason concerns the use of length-width ratios in the
two species. McClellan (1966, p. 475) suggested the likelihood of
a fairly constant 2:1 ratio between these dimensions for S. cylindrica.
Measurements made on specimens recovered here agree with that
ratio. The ratio in S. elongata, if similarly constant, would be much
greater, but no measurements have been made for a comparison.
The only reported occurrence of Stegnammina cylindrica in the
Devonian is that by Mound (1961) and Browne and Schott (1963).
Following the above discussion of that species and S. elongata, as
considered herein the specimens at locality UWA 6010 represent the
youngest occurrence of S. cylindrica.
Stegnammina hebesta? Moreman Pl. 34, fig. 19
1930. Stegnammina hebesta Moreman, Jour. Palecnt., vol. 4, No. 1, p. 50, pl.
7, fig. 13.
Description. — Test free, subcylindrical, robust, ends rounded,
diameter about two-thirds the length; wall thin, finely arenaceous
and well cemented; no apparent aperture.
294 BULLETIN 274
Measurements. — Length of the figured specimen is 0.21 mm
and diameter 0.14 mm.
Occurrence. — Hypotype 34698 is from the McMonnigal Lime-
stone at locality UWA 5985. Forms referred to this species are also
present at locality UWA 6000. Both localities are of Helderbergian
age,
Discussion. — The type specimen of Moreman is more robust
than any of those in the present material, having a diameter about
three-fourths the length. This feature is one of the main reasons
for separating S. hebesta from the less robust species S. cylindrica.
The material at hand is poorly preserved for the most part, and the
thin test walls are fragile, making measurement of the length-
diameter ratio difficult. The figured specimen, though slightly de-
formed, seems typical, however, and has a ratio of 3:2 which is
between S. cylindrica and S. hebesta. The consistency found within
examples of the former species precludes the identification of this
material with them. Therefore, it has been referred to S. hebesta
with some question.
Ecologically this species had a preference for a higher energy
environment as indicated by its occurrence in nearly pure carbonate
rocks. In contrast, Stegnammina cylindrica was found in rocks
indicative of quieter conditions as well as consistently occurring at
lower stratigraphic horizons.
Stegnammina moremani (Dunn) Pl 3s4 tig V7 Pi soe thica?
1942. Thekammina moremani Dunn, Jour. Paleont., vol. 16, No. 3, p. 326, pl.
42, fig. 22.
1961. Thekammina moremani Dunn, Mound, Indiana Dept. Conservy., Geol.
Sum, Bulli Nos 23, ps 26) ple i, fiz: 29:
1964. Stegnammina moremani (Dunn), Loeblich and Tappan, Treat. Invert.
Raleont., Pt. ©) Protistay2) vol. 1) ps 196.
Description. — Test free, triangular in outline, slightly de-
pressed; wall thin, finely arenaceous, well cemented; no aperture
apparent, except minute openings on test surface.
Measurements. — Length of hypotype 34699, measured from
triangle apex to opposite side is 0.21 mm for each side.
Occurrence. — This is a rare species, present only at locality
UWA 5985 in the McMonnigal Limestone. The only previous re-
port of S. moremani is from the Lower Silurian Brassfield Limestone
in Missouri and Indiana.
StturRO-DEVvoNIAN MicroFrAuna: McCLeLian 295
Discussion. —Loeblich and Tappan (1964 p. 196) included
species of Ratbosammina Moreman (1930) and Thekammina Dunn
(1942) within the generic limits of Stegnammina stating that
“modifications of chamber shape are not regarded as generic in
importance”. Because Thekammina had been erected on the basis
of its angular test shape it falls within the species limits set by
them.
Stegnammina quadrangularis (Dunn) P34 tes load
1942. Thekammina quadrangularis Dunn, Jour. Paleont., vol. 16, No. 3, p.
326, pl. 42, fig. 23.
1961. Thekammina quadrangularis Dunn, Mound, Indiana Dept. Conserv.,
Geol. Sur., Bull. No. 23, p. 25, pl. 1, fig. 28.
1963. Thekammina quadrangularis Dunn, Browne and Schott, Bull. Amer.
Paleont., vol. 46, No. 209, p. 210, pl. 49, figs. 13, 14.
1964. Stegnammina quadrangularis (Dunn), Loeblich and Tappan, Treat.
Invert. Paleont., Pt. C, Protista 2, vol. 1, p. 196.
Description. — Test free, boxlike with planar sides; wall thin,
fine to coarsely arenaceous, poor to moderately cemented; no aper-
ture apparent.
Measurements. — Hypotype 34700 has length 0.44 mm and
width 0.42 mm. Hypotype 34701 has length 0.64 mm and width
0.31 mm.
Occurrence. — Specimen 34700 is from locality UWA 5992
and 34701 from locality UWA 6019, both in the Lone Mountain
Dolomite. The species is also uncommonly to rarely present at
several other localities from Ludlow to Lower Helderberg age.
Discussion. — For the same reasons as those presented in the
discussion of S. moremant, this species belongs within the limits of
Stegnammina and the generic name Thekammina should be sup-
pressed as a junior synonym.
The test shape of S. quadrangularts varies substantially from
nearly square to elongate, always with the boxlike form. Most
specimens here were nearly square. The height of the test is usually
indeterminate because the thin, fragile wall is so easily damaged
and distorted.
This species occurs most frequently in silty and shaly lime-
stones indicating a probable preference for quieter water. How-
ever, the two hypotypes are from a distinctly different environment,
the reefoid conditions in the Lone Mountain Dolomite. It is one
of the few foraminifers found in this environment, and its presence
296 BULLETIN 274
here is the only known occurrence in such conditions as well as its
youngest stratigraphic position,
Genus STORTHOSPHAERA Schulze, 1875
1875. Storthosphaera Schulze, Komm. Untersuch. deutsch. Meere iz Kiel,
Jahresber, vol. 1872-73, p. 113.
Type species. —Storthosphaera albida Schulze (1875, ibid.,
pl. 2).
Description. — Test free, monothalmous, irregular chamber,
wall finely arenaceous, poorly to well cemented, aperture indefinite.
The surface of the test wall may be irregular with protuber-
ances of various configurations but not possessing any aperture. The
interior, despite the irregularities of the test which suggest incipient
chambers, lacks a definite indication of subdivision that might sug-
gest Sorosphaera.
Storthosphaera malloryi McClellan, n. sp.
Pl. 34, figs. 6-10; Pl. 40, fig. 3
Description. — Test free, irregular, globular chamber appears
lumpy on exterior; wall thin, very finely arenaceous, well cemented;
no aperture apparent.
Measurements. — Maximum test size for the figured speci-
mens averages 0.26 mm.
Pl. 34 Spec. Number Test Size
Fig. 6 34703 holotype 0.29 mm
Big. /, 34704 paratype 0.19 mm
Fig, 8 34705 paratype 0.22 mm
Fig. 9 34706 paratype 0.26 mm
Fig. 10 34707 paratype 0.33 mm
Occurrence. — Present throughout the stratigraphic interval at
several localities. Holotype 34703 from locality UWA 5985; para-
types 34704, 34705 at UWA 5985; 34706 at UWA 5976; and 34707
at UWA 6010.
Discussion. — This species varies considerably from the type,
S. albida, which has an irregular wall thickness and heterogeneous
development of ridges and protuberances on the surface. In con-
trast, S. malloryi has a thin wall and a lumpy, or globular, surface
appearance. The lumpy nature actually resembles chambers but
SILURO-DEVONIAN MiIcROFAUNA: MCCLELLAN 297
smaller lumps make up most of the surface of these pseudochambers.
Interiorly there is no indication of subdivision into a multilocular
growth, hence this is regarded as a species of Storthosphaera.
Previously Storthosphaera has been known only from the
Recent and generally from cold waters. In the present material
it ranges from the Middle Silurian to Lower Devonian. There seems
not to be any preference for a particular environment other than
not being found in the high energy condition of the reefoid dolomites.
Subfamily SACCAMMININAE Brady, 1884
Genus SOROSTOMASPHAERA McClellan, 1966
1966. Sorostomasphaera McClellan, Bull. Amer. Paleont., vol. 50, No. 230, pp.
478, 479.
Type species. — Sorostomasphaera waldronensis McClellan,
(1966, zbid., pls, 37, 41, figs. 1-5).
The original description of McClellan (1966, p. 478) is as
follows:
Test free, multilocular, consisting of a variable number of globular
to spherical chambers irregularly attached to each other; chambers may
be loosely connected or appressed where adjoined; wall thin, finely
arenaceous, well cemented; single, rounded aperture on each chamber,
no openings between chambers.
Sorostomasphaera differs from Sorosphaera Brady (1879) in
possessing apertures, and from Saccamminoides Geroch (1955) in
having consistent chamber diameters each with an aperature.
Sorostomasphaera waldronensis McClellan Pl. 35, figs. 7,8
1966. Sorostomasphaera waldronensis McClellan, Bull. Amer. Paleont., vol.
50, No. 230, pp. 479-80, pl. 37, 41, figs. 1-5.
Description. — Test free, multilocular, variable number of
chambers which are subspherical, chambers loosely or firmly joined,
wall thin, very finely arenaceous, well cemented; single circular
aperture on each chamber.
Measurements. — Diameter of single chamber is 0.30 mm on
hypotype 34694. Diameter of 34695 is 0.29 mm. These measurements
compare closely with the original types (McClellan, 1966, p, 480).
Occurrence. — Rarely present at two localities, UWA 6028
(hypotypes) and UWA 6025 in the Willow Creek section. The
stratigraphic interval is Wenlockian to lower Ludovian, in the lower
and middle Roberts Mountains Formation.
298 BULLETIN 274
Discussiton.— The presence of this species here extends its
stratigraphic range, but it is significantly absent from localities in
younger strata which indicate quite similar physical environment
conditions. McClellan (1966, p, 379) stated that the genus was
not particularly specialized ecologically, and its occurrence in
Nevada is similar to that discussed by him. Thus it is possible that
Sorostomasphaera waldronensis is biostratigraphically a_ useful
species restricted to rocks of middle and upper Niagaran age,
Genus STOMASPHAERA Mound, 1961
1961. Stomasphaera Mound, Indiana Dept. Conserv., Geol. Sur., Bull. No.
S35 Oy Cade
Type species. —Stomasphaera brassfieldensis Mound (1961,
thid., pl. 2, figs. 9-13, text fig. 3).
Mound’s (p. 28) generic diagnosis is as follows:
Wall rough, thick to thin, medium to coarsely arenaceous, grains
poorly to well cemented; test free, subangular to spherical, most com-
monly subspherical or spherical; aperture single, round or oval.
This is a common Lower Silurian genus in Indiana but rare
from younger rocks, It occurs rarely in central Nevada no higher
than lower Ludlovian, or uppermost Niagaran.
Stomasphaera brassfieldensis Mound Pl. 36; fig. 7
1961. Stomasphaera brassfieldensis Mound, Indiana Dept. Conserv., Geol. Sur.,
Bull. No. 23, p. 28, pl. 2, figs. 9-13, text fig 3.
Description. — Test free, monothalmous, subspherical, slightly
produced around aperture; wall thin, medium to finely arenaceous,
well cemented; single, round aperture.
Measurements. — Hypotype 34702 diameter is 0.62 mm and
aperture 0.14 mm.
Occurrence. — The figured specimen is from locality UWA 6022
in the Roberts Mountains. The species is also present at locality
UWA 5983 in the Bastille Limestone. Rare at both localities.
Discussion. — The similarity between S. brassfieldensis and
Saccammina Sars (1869) species is close except for the aperture.
Saccammina species have the aperture at the end of a short neck
which is lacking on the well-preserved Stomasphaera brassfieldensis
in the Bastille strata and Roberts Mountains strata. Sorostoma-
sphaera waldronensis also has affinities with Stomasphaera brass-
fieldensis, but is consistently more nearly spherical, has no modi-
SILURO-DEVONIAN MicroFAUNA: MCCLELLAN 299
fication of the test in the apertural area, and has a thinner and
more finely arenaceous wall. These features allow identification even
of isolated chambers.
Genus THURAMMINA Brady, 1879
1879. Thurammina Brady, Quart. Jour. Micro. Sci. new ser., vol. 19, pp. 45-47.
Type species. —Thurammina papillata Brady (1879, sbid., p.
45) subsequently designated, Cushman, 1910, p. 57.
Brady (1879) failed to provide a generic diagnosis and Cush-
man (1910, p. 57) subsequently described the genus and designated
the type species.
Test typically free, usually nearly spherical, but in some species com-
pressed, chamber single and undivided in typical species; wall thin,
composed of fine sand with more or less chitin; apertures several to
many at the end of nipple-like protuberances of the surface, occasional-
ly wanting.
There are a number of lower Paleozoic species of Thurammina
but seldom is any one of them abundant. The material studied
herein has each of the species sparsely present at only one or two
localities. McClellan (1966, p. 481) did not find the genus to be
generally useful as an environmental indicator. Certain species may
nonetheless be ecologically restricted in the Siluro-Devonian sedi-
ments of central Nevada.
Thurammina arcuata Moreman Pl. 35, fig. 18
1930. Thurammina arcuata Moreman, Jour. Paleont., vol. +, No. 1, p. 54, pl.
Geties. 2. 3.
Description. — Test free, monothalmous, arcuate, inflated poly-
gonal outline; wall thin, finely arenaceous, well cemented; aper-
tures several, at ends of short, broad protuberances.
Measurements. — Width of hypotype 34708 is 0.37 mm. Aper-
ture neck length 0.04 mm,
Occurrence. — Uncommon, figured specimen from _ locality
UWA 6018, in the McColley Canyon Formation. Also present in
the McMonnigal Limestone at locality UWA 5985. Both forma-
tions are of Lower Devonian age.
Discussion. — The specimens recovered here are more robust
than Moreman’s figured types, and the short nodes bearing the
apertures are better developed. The polygonal and arcuate outline
is consistent with both Moreman’s (1930) specimens and those of
Browne and Schett (1963).
300 BULLETIN 274
Thurammina arcuata has previously been recorded from Lower
and Middle Silurian argillaceous limestone strata. McClellan (1966,
p. 460) listed this species as one limited to lower energy environ-
ments in the Waldron Shale of Indiana. The specimens from cen-
tral Nevada were found in less argillaceous carbonates, suggestive
of higher energy conditions.
Thurammina quadritubulata? Dunn Pl 3b tigao
1942. Thurammina quadritubulata Dunn, Jour. Paleont., vol. 16, No. 3, p.
334, pl. 43, fig. 22.
Description. — Test free, monothalmous, spherical; wall thin,
composed of medium-sized sand grains, moderately cemented; four
apertures at ends of short, broad tubules arranged in pairs on dif-
ferent planes.
Measurements. — Diameter of hypotype 34709 is 0.51 mm;
aperture necks 0.035 mm.
Occurrence. — This species is rarely present at locality UWA
6004 in the Roberts Mountains Formation.
Discussion. — The apertures of the specimens studied herein
show a variation in arrangement from that described by Dunn as
normal for the species. He described the apertures (p. 334) as:
four blunt tubular necks arranged in opposite pairs, the horizontal
plane of one pair cutting the vertical pair of the other at right angles.
Viewed from above, however, the four tubes do not lie in vertical planes
through the center of the sphere but are offset on opposite sides of
perpendicular vertical central planes.
These specimens have one pair of apertures arranged at opposite
poles. The other pair does not form a plane normal to the first
pair, because one aperture is at an angle of less than 90 degrees. It
is believed that the above arrangement only represents a variation,
though positive identification of the specimens as T. quadritubulata
was not possible because comparison with the holotype could not
be made.
Thurammina trituba Dunn P1.\35, fig: 17
1942. Thurammina tributa [sic] Dunn, Jour. Paleont., vol. 16, No. 3, p. 334,
pl. 43, fig. 26.
Description.— Test free, monothalmous, triangular outline
slightly inflated; wall thin, finely arenaceous, well cemented; aper-
tures number three, at ends of short nodes forming corners of tri-
angular test outline.
StLtuRO-DEVONIAN MicroFauna: McCLeLLan 301
Measurements. — Lengths of the three sides from aperture to
aperture are 0.30 mm, 0.30 mm, 0.35 mm on figured hypotype 34710.
Occurrence. — Rarely present in the Gatecliff Dolomite at
locality UWA 5998.
Discussion. — The specimen described here is slightly more ro-
bust than characteristic for the species. Its occurrence in the Gate-
cliff Dolomite compares well stratigraphically with the occurrence
in the Osgood Limestone reported by Dunn (1942, p. 334). Kay
and Crawford (1964, p. 438) suggested a Llandovery age for the
Gatecliff on the basis of the graptolite fauna and the Osgood is in
nearly the same stratigraphic interval. Because the species is only
rarely present and has been reported only from Missouri and central
Nevada, too much importance should not be placed on its bio-
stratigraphic occurrence.
Thurammina tubulata Moreman IDL, Bh), sake, ay)
1930. Thurammina tubulata Moreman, Jour. Paleont., vol. 4, No. 1, p. 52,
jl Op sakes fh
Description. — Test free, monothalmous, spherical, wall thin,
finely arenaceous, abundant cement; apertures at ends of variously
positioned tubes.
Measurements. — Diameter of figured specimen 34711 is 0.22
mm and aperture neck is 0.04 mm long.
Occurrence. — This species occurs rarely at locality UWA
6015 only.
Discussion. — The single specimen recovered here has several
of the tubules mostly broken off leaving only the bases. Those which
are intact are long and definitely indicate the specimen to be a
T. tubulata rather than T. papillata which has shorter, nodular pro-
jections.
Subfamily HEMISPHAERAMMININAE Loeblich and Tappan, 1961
Genus HEMISPHAERAMMINA Loeblich and Tappan, 1957
1904. Webbinella Rhumbler, Archiv. Protistentkunden, vol. 3, pt. 1, p. 228.
1957. Hemisphaerammina Loeblich and Tappan, U.S. Nat. Mus., Bull. 215,
pp. 223, 224.
1958. Fairliella (in part) Summerson, Jour. Paleont., vol. 32, No. 3, pp.
5519, 556;
Type species.— Hemisphaerammina batalleri Loeblich and
Tappan (1957, ibid. p. 224, pl. 72, fig. 3).
302 BuLLeETIN 274
The redescription given by Loeblich and Tappan (1964, p. 202)
is as follows:
Test attached, consisting of single hemispherical chamber; may have
bordering flange; wall agglutinated, with considerable cement; no
apparent aperture.
This genus was established by Loeblich and Tappan (1957)
for those agglutinated species of Webbinella Rhumbler (1901) which
were excluded when the type W. hemispherica was recognized as a
polymorphinid. Certain species of Fairliella Summerson (1958) can
also be allotted to Hemisphaerammina. F. dicantha Summerson and
F. clitellata Summerson are retained in that genus as distinctly
different from Hemtsphaerammina, following the decision of McClel-
lan (1966, pp. 484, 485).
Hemisphaerammina bradyi Loeblich and Tappan
Pl. 35,. fig. 1; Pl) 36; hex
1957. Hemisphaerammina bradyi Loeblich and Tappan, U.S. Nat. Mus., Bull.
215, ps 224, pl? 72) figs 3.
Description. — Test attached, monothalmous, hemispherical,
highly convex with side wall forming a sharp angle at junction
with base; dorsal wall thickness greater than the ventral, finely
arenaceous, well cemented; no apparent aperture.
Measurements. — Figured hypotype 34663 has diameter of
0.31 mm and height of 0.28 mm.
Occurrence. — The hypotype is from locality UWA 6018. The
species is common at all stratigraphic intervals in the area.
Discussion.— The most favorable environment for H. bradyi
appears to have been that in which well-bedded limestones were
deposited from waters bearing little argillaceous material, It is
commonly found in samples with a diversified foraminiferal and
invertebrate faunule, that implies relatively higher energy condi-
tions. The only previous Paleozoic record of this species is in the
Waldron Shale of Indiana (McClellan, 1966, p. 485) where it is in
calcareous shales, but again, is associated with a diversified faunule
of a type similar to that of central Nevada.
Hemisphaerammina? cf. H. bradyi Loeblich and Tappan PI. 35, fig. 2
Description. — Test attached, monothalmous, hemispherical,
moderately convex; wall thin, finely arenaceous, base thinner than
convex dorsal wall, well cemented; no apparent aperture.
StturRo-DEVONIAN MIcROFAUNA: MCCLELLAN 303
Measurements. — Specimen 34664 has diameter of 0.28 mm
and height of 0.13 mm.
Occurrence. — Present at locality UWA 6028 associated with
typical specimens of Hemisphaerammina brady.
Discussion. — Hemisphaerammina bradyi is typically hemi-
spherical and highly convex, the test being nearly as high as it is
broad. Variants with partially collapsed tests can usually be readily
recognized. This form is questionably referred to the genus Hemi-
sphaerammina because of the abnormally low convexity which
seems not to be caused by collapse. A closely related genus,
Webbinelloidea Stewart and Lampe (1947), has several species
which also have low convexity but differ from Hemisphaerammina
in the manner of junction between the dorsal and ventral walls (see
generic discussion of Webbinelloidea).
Hemisphaerammina coronata (Ireland) Pl. 35, fig. 4; Pl. 38, fig. 4
1939. panes lle coronata Ireland, Jour. Paleont., vol. 13, No. 2, p. 198, fig.
-11.
Description. — Test attached, monothalmous, elongate hemi-
spherical, broad flange with projecting spines; wall thin, finely
arenaceous, well cemented; no aperture apparent.
Measurements. — Test length of figured specimen 34665 is
0.15 mm, width 0.13 mm, height 0.11 mm, wall thickness 0.01 mm,
width of flange 0.02 mm.
Occurrence. — Only a single specimen was recovered from
locality UWA 6004 in the midlde Roberts Mountains Formation.
Discussion. — This specimen is much smaller than Ireland’s
holotype, whose length is 0.87 mm, and probably represents a neanic
individual. A scanning electron micrograph shows the dorsal wall
having the grains arranged in concentric layers, a pattern not pre-
viously recognized in other specimens. Only a small part of the
diagnostic basal flange is preserved but Hemisphaerammuina casteri
McClellan (1966), the only other species with a broad flange, has
a very thick flange with no spines, and, therefore, cannot be con-
fused with H. coronata.
Hemisphaerammina discoidea? (Summerson) PIF35) figs"3
1958. Fairliella discoidea Summerson, Jour. Paleont., vol. 32, No. 3, p. 557,
pl. 82, fig. 20, text figs. 6a, b.
304 BULLETIN 274
1966. Hemisphaerammina discoidea (Summerson), McClellan, Bull. Amer.
Paleont., vol. 50, No. 230, p. 486, pls. 38, 42, figs. 2a, b, 3.
Description. — Test attached, monothalmous, slightly ovoid,
strongly biconvex, dorsal and ventral surface joined along highly
sinuous line which is sharply defined with thickened test wall; flange
questionably present as wedge edge of sinuous junction line; wall
thin to thick, medium to coarsely arenaceous, poorly cemented; no
apparent aperture.
Measurements. — Width of figured specimen is 0.26 mm and
height is 0.22 mm.
Occurrence. — Rarely present only at locality UWA 6010, of
earliest Devonian age.
Discussion. — This species has previously been reported from
the Columbus Limestone (M. Devonian) of Ohio, and Waldron
Shale (M. Silurian) of Indiana. The specimen here differs from
those of earlier reports in the configuration of the junction line
between the dorsal and ventral surfaces. The Devonian forms
(Summerson, 1958, p. 557) possess a straight to slightly sinuous
junction line, while those from the Silurian (McClellan, 1966, p.
486.) show more variation, Specimens from both of these occur-
rences have a definite wedge-shaped flange. Specimen 34666 has
a much more highly sinuous junction line and may be lacking the
wedge-shaped flange. Unfortunately the flange is difficult to rec-
ognize and for this reason identification of the species was ques-
tioned.
The actual configuration of the junction line probably is not
constant for the species. The varying curvature of the line could
readily be a function of the attaching surface. Those specimens
with a highly sinuous line have correspondingly highly convex ven-
tral and dorsal walls.
Hemisphaerammina bipatella McClellan, n. sp.
Ply 35) figs) 56> Ply 38 etissoaD
Description. — Test attached, monothalmous, dorsal side mod-
erately convex and elliptical, ventral side slightly convex and hemi-
spherical; heavy flange at contact of upper and lower surfaces is
wide adjacent to long axis of the dorsal ellipse of the chamber and
narrow to absent along short axis giving overall test a circular
outline, flange merges gently into chamber surface without suture
SILURO-DEVONIAN MIcRoFAUNA: MCCLELLAN 305
and is indistinct from ventral portion of chamber; wall thick,
medium to finely arenaceous, grains may be arranged in concentric
layers; poorly to moderately cemented; no aperture apparent.
Measurements. — Holotype 34668 has chamber length 0.30
mm, width 0.18 mm, total diameter 0.31 mm, height 0.15 mm;
flange width 0.05 mm, thickness 0.04 mm, Paratype 34667 cham-
ber length is 0.20 mm, width 0.12 mm, total diameter 0.20 mm,
height 0.11 mm; flange width 0.04 mm, thickness 0.03 mm.
Occurrence. — Present only at locality UWA 5990 in the basal
beds of the Roberts Mountains Formation.
Discussion. — Hemisphaerammina bipatella is most closely re-
lated to H. casteri McClellan (1966), both species characterized
by a broad, thick marginal flange. It is also closely related to H.
coronata (Ireland, 1939) in showing a tendency to develop the
test with concentric layers of sand grains, but the latter species
has a much thinner flange. H. discoidea (Summerson, 1958) has a
biconvex test much like H. bipatella, but a narrower flange and
more variable shape. All the specimens recovered here closely re-
semble the holotype though some are less well preserved.
Isolated chambers of the multilocular Metamorphina imbricata
McClellan (1966) look somewhat similar to H. bipatella in their
thick, broad flange and moderate convexity, The chamber is hemi-
spherical, the bordering flange shows a constant width around it, and
the ventral surface is planar rather than convex. H. bipatella can-
not be a Metamorphina species because that genus is only known
to have a planar surface of attachment, is multilocular, and is not
known to have wall material arranged in an orderly manner. None
of the specimens here show any evidence of multilocularity.
Interpretations of ecology must remain inconclusive until the
species has been recognized more widely. It was found as a common
constituent of a limited foraminiferal faunule, lacking associated in-
vertebrates. It occurs in an argillaceous, cherty limestone, suggest-
ing a rather quiet bottom habitat with low circulation.
Genus ATELIKAMARA McClellan, n. gen.
Type species. — Atehkamara incomposita McClellan, n. sp.
Description. — Test attached, inflated dorsally, ventral sur-
face planar to angular; interior incompletely divided into incipient
chambers by walls which may be slightly developed to semicom-
306 BULLETIN 274
plete; wall thin to thick, composed of poorly sorted sand grains,
well cemented; aperture indefinite.
The distinguishing feature of this genus is the incipiently de-
veloped chambers, hence the origin of the name: Greek atelis =
incomplete, kamara = chamber. Storthosphaera Schulze (1875)
resembles Atelikamara externally in the irregular shape, but the
latter differs in being attached, and, therefore, belongs in the Hemi-
sphaerammininae. The same difference separates this new genus
from Anictosphaera n. gen. which has incomplete walls dividing the
chambers of a free test,
Atelikamara does not have definite chambers like multilocular
genera in the same subfamily. In comparison with unilocular gen-
era, none of them has any suggestion of division into chambers.
Thus, Atelikamara is distinct in its character of the partial test
subdivision. The degree to which subdivision takes place is appar-
ently quite variable, as partially developed walls may be present
as well as only very slightly developed partitions on the same
specimen. Often the partial chambers are separated by a depression
in the exterior test wall, such that they present the appearance
of nodes on the surface of the test.
Atelikamara incomposita McClellan, n. sp.
Pl. 36, figs. 1-3, 6; Pl. 40, figs. 5a,b
Description. — Test attached, irregular outline, conical to ir-
regularly inflated dorsally, ventral surface usually planar, may be
angular; interior subdivisions incomplete to nearly lacking; ex-
terior surface may haye nodular appearance with slight depressions
corresponding to interior subdivisions; wall thin to thick, basal
wall unknown, composed of poorly sorted sand grains, abundant
cement; no apparent aperture, scattered pseudopodial openings be-
tween grains on dorsal surface.
Measurements. — Dimensions of the type specimens suggest
two distinct sizes, but the differences are caused by grain sizes in
the wall construction.
Pi 36 Specimen Length Height Wall
Fig. 1 34658 holotype 0.63mm 0.38mm 0.04mm
Fig. 2 34793 paratype 0.67mm 0.45mm 0.04mm
Fig. 3 34657 paratype 0.34mm 0.15mm 0.015mm
Fig. 6 34659 paratype 0.37mm 0.14mm 0.015mm
SILURO-DEVONIAN MicroFAuNA: McCLELLAN 307
Occurrence. — Holotype 34658 and paratype 34793 are from
locality UWA 6022 where the species is common, Paratype 34657
is from locality UWA 6004, and paratype 34659 from locality
6007. All localities are in the middle and upper parts of the Rob-
erts Mountains Formation.
Discussion. — The external form of this species is distinctive
by its irregularity. In the holotype the lumpy appearance only
moderately suggests the internal form, The paratypes have a more
globulose surface suggesting corresponding internal partial sub-
division but actually have little development of the partial chamber
walls. Several unfigured specimens from the type locality lack sur-
face expression of the incipient chamber development inside. The
basal surface is completely lacking in every specimen and likely con-
tained no agglutinated material, and, therefore, it has not been
preserved. It is apparent from specimen 34793 that the surface
of attachment was variable and partially responsible for the form
of the entire test. This individual was attached in a narrow crevice,
evidenced by the sharp angularity of the base, and was, therefore,
restricted in growth to a less spreading habit than specimens at-
tached to a flat surface.
There are two distinct sizes suggested in the measurements
of the figured specimens. Other foraminifers from locality UWA
6022 have utilized coarser grains than normal in constructing their
tests. The large size and thick wall of Atelikamara incomposita
specimens at this locality has resulted from use of coarse sand
grains in the test wall. Examples from other localities all used finer
grains resulting in a thin wall and smaller test size. Since all the
foraminifers recovered from locality UWA 6022 have used larger
grains than normal, it is clear that the grain size was determined
by the coarseness of available material rather than a preference
of the organism. For this reason the size differences of A. incom-
posita do not represent two species, because grain size alone, in a
primitive foraminifer, is not of specific value.
Atelikamara incomposita occurs commonly only in the upper-
most beds representing the argillaceous and cherty limestone litho-
some. It occurs rarely in thick-bedded limestones slightly lower
stratigraphically. All these occurrences are in the Roberts Moun-
tains Formation where it is approximately uppermost Niagaran
308 BULLETIN 274
age, The argillaceous and cherty limestones suggest a preference
by the organism for low energy, quiet water conditions.
Genus METAMORPHINA Browne, 1963
1963. Mctamorphina Browne, in Browne and Schott, Bull. Amer. Paleont.,
vol. 46, No. 209, p. 223.
Type species. — Webbinella tholus Moreman (1933, Jour. Pal-
eont., vol. 7, No. 4, p. 395, pl. 47, figs. 8, 10).
Following is the generic diagnosis given by Browne im Browne
and scnott (1965, p: 223):
Test attached, plano-convex, varying in outline from circular to oval
to linear; often surrounded by a marginal flange; single chambered
to multichambered; wall of fine sand grains, smooth to roughly finished
with the basal wall thin and frequently missing; aperture not apparent.
This genus is characterized by its thin basal wall and marginal
flange. Hemisphaerammina differs from Metamorphina in being
single chambered often having an orderly arranged wall material,
and having higher dorsal convexity. The type species M. tholus
(Moreman) was formerly considered a Webbinella until Loeblich
and Tappan (1957, p. 224) reevaluated the latter genus (see gen-
eric discussion of Hemisphaerammina). Metamorphina was erected
for those arenaceous, multilocular forms of Webbinella, subse-
quently,
Metamorphina tholus (Moreman) Pl. 35; figs 133) P1939), Hees
1933. Webbinella tholus Moreman, Jour. Paleont., vol. 7, No. 4, p. 395, pl.
47, figs. 8, 10.
1963. Metamorphina tholsus (sic) (Moreman), Brown and Schott, Bull. Amer.
Paleont., vol. 46, No. 209, p. 225, pl. 51, figs. 1-9.
Description. — Test attached, plano-convex with low convexity
widening to the basal plane, may have marginal flange; number of
chambers variable, closely appressed with rectilinear contact and
well-developed suture; wall of fine or poorly sorted sand grains,
well-cemented, basal wall thin and often missing; aperture not
apparent.
Measurements. — Hypotype 34676 has test length of 0.35 mm
and width of 0.22 mm. Chambers are each 0.18 mm long and
small, third chamber length is 0.07 mm.
Occurrence. — Rare, only at locality UWA 6010.
Discussion. —M, tholus is a common species of the Lower
Silurian but has not been reported from younger rocks except in
SILURO-DEVONIAN MiIcrRoFAUNA: MCCLELLAN 309
the Mississippian Lodgepole Limestone of Montana (Gutschick,
Weiner and Young, 1961, p. 1205). In Nevada it presently is known
only at one locality, but further study, especially in the argillaceous
limestones in which it most often occurs in other areas, should
reveal a more widespread occurrence in this region. The specimen
figured here has less well sorted wall material than is customary
for the species, In addition, the wall dividing the two chambers
does not extend to the base as is normal, (best seen on PI. 39, fig.
5). An interesting feature of the specimen is the small third cham-
ber which apparently was only partially developed.
Metamorphina gibbosa (Ireland) Plead, figs 16
1939. Webbinella gibbosa Ireland, Jour. Paleont., vol. 13, No. 2, p. 198, figs.
B-23, 24.
1963. Metamorphina gibbosa (Ireland), Browne and Schott, Bull. Amer.
Paleont., vol. 46, No. 209, p. 224, unfigured.
1966. Metamorphina gibbosa (Ireland), McClellan, Bull. Amer. Paleont., vol.
50, No. 230, p. 490, pl. 38, figs. 4a, b, 5; pl. 42, figs. 4a, b, 5.
Description. — Test attached, polythalmous, plano-convex, mar-
ginal flange narrow, may be incomplete, chambers closely joined
with faint suture defining contact plane; wall thin, basal wall very
thin and commonly absent, finely arenaceous, well cemented; no
aperture apparent.
Measurements. — Hypotype 34675 has test length of 0.44 mm,
chambers 0.24 mm, 0.18 mm, and 0.13 mm; test height 0.18 mm.
Occurrence. — Present in rare numbers only at locality UWA
6030 in the lower Roberts Mountains Formation of Wenlockian
age.
Discussion.— The diagnostic features of this species are the
very narrow marginal flange and faint, slightly depressed, dorsal
sutures between chambers. M. tholus (Moreman) may also have
only a narrow flange, but its sutures are more prominent and de-
pressed. An additional feature, which appears with relative consis-
tency, is the high convexity of M. gibbosa compared to other
species, such as M. tholus and M. imbricata.
McClellan (1966, p. 492) considered these three species to
represent a phylogenetic lineage with M. gibbosa the most primitive
and M. imbricata the most advanced. The former species has been
recorded from the Lower and Middle Silurian of several areas,
while the latter is known only from the Middle Silurian of Indiana.
310 BULLETIN 274
If A. gibbosa is more primitive and characteristic of the Lower
Silurian, then its stratigraphic position in central Nevada may be
significant. The one locality at which it occurs has a foraminiferal
faunule quite similar to that found in the Waldron Shale of south-
eastern Indiana (McClellan, 1966). These comparative faunal ele-
ments from the Waldron are species commonly found in Lower
Silurian rocks elsewhere. This evidence suggests correlation between
the Waldron Shale and basal units of the Roberts Mountains For-
mation based on the foraminifers, M. gibbosa and Glomospira
sluriana (see discussion of that species) are probably the most
diagnostic species.
Genus WEBBINELLOIDEA Stewart and Lampe, 1947
1947. Sorosphaeroidea Stewart and Lampe, Jour. Paleont., vol. 21, No. 6,
p. 534.
1947. Webbinelloidea Stewart and Lampe, ibid., pp. 534, 535.
Type species. — Webbinelloidea similis Stewart and Lampe,
(1947, tbids.p:535, pl. 78, tig:-8)-
Description. — Test attached, multilocular, usually plano-con-
vex, chambers hemispherical, globular, to subangular and usually
highly convex; junction of base and sides forms an obtuse angle;
wall thick to thin, well cemented, finely to coarsely arenaceous;
no apparent aperture.
This genus resembles Metamorphina Browne (1963) in general
form and habit but lacks the marginal flange and thin basal wall
of that genus. Single chambered specimens differ from Hemz-
sphaerammina Loeblich and Tappan (1957) on three counts: firstly,
many examples of the latter species have a marginal flange like
that of Metamorphina; secondly, Webbinelloidea joins the basal, or
ventral, wall to the dorsal with a contact that forms a rounded,
bluntly obtuse angle often greater than 90 degrees. A comparable
feature on Hemisphaerammina forms a sharp, V-shaped angle of
not more than 90 degrees. Thirdly, several species of Hemisphaer-
ammuina have test wall material oriented in a regular, layered man-
ner, while species of Webbinelloidea have yet been found to only
randomly place grains in the test wall.
Sorosphaeroidea Stewart and Lampe (1947) differs only in
the polygonal outline of the chambers, This genus, therefore, differs
in traits which seem to be only of specific character. A more de-
Stturo-DeEvonian MicroFauna: MCCLELLAN Si
tailed discussion has been presented by Conkin and Conkin (1970,
p. 4).
The material here recovered and referred to Webbinelloidea
is so placed in accordance with the generic description of Stewart
and Lampe (1947) and Summerson (1958). Recently Conkin and
Conkin (1970) have restudied the type material of the former au-
thors plus new material from the Devonian of Ohio. As a result
they have revised the genus and emended the description, provid-
ing four morphological groups all within the type species W. similis.
They also have demonstrated that Webbinelloidea contains an aper-
ture on the dorsal surface. Hence the genus would properly be in
the subfamily Saccammininae. Of the Nevada species, only W.
hemispherica has been revised by Conkin and Conkin, the others
not being included in their study. The material recovered here does
not show any evidence of an aperture even under scanning electron
microscopy (Pl. 38, figs. 1, 2). Therefore the original generic con-
cept of Webbinelloidea is followed here, though it is recognized
that more, well-preserved material may necessitate future revision.
In view of Conkin and Conkin’s discoveries (1970) the entire re-
lationship of Webbinelloidea, Metamorphina, and Hemisphaeram-
mina should be closely studied for probable phyletic revisions.
Webbinelloidea hattini McClellan Ple3sbs tie. 12
1966. Webbinelloidea hattini McClellan, Bull. Amer. Paleont., vol. 50, No. 230,
p. 495, pls. 38, 42, figs. 10, 11.
Description. — Test attached, one- or two-chambered forms
known, plano-convex with high conical convexity of upper surface,
junction of basal surface forms rounded angle, suture between
chambers deep, rectilinear; wall thin to thick, fine to coarsely
arenaceous, moderately cemented; no apparent aperture.
Measurements. — Hypotype 34713 has diameter of 0.31 mm and
height of 0.26 mm.
Occurrence. — The figured specimen is from locality UWA
6025. Also present at locality UWA 6009, which gives a range of
upper Wenlock to lower Helderbergian age in Nevada.
Discussion. — The high conical convexity of the dorsal sur-
face is characteristic for the species. Multilocular specimens are
further characterized by the deep, rectilinear suture between cham-
312 BULLETIN 274
bers. Webbinelloidea similis Stewart and Lampe (1947) is similarly
multilocular but has a less conical form and the chambers are
more loosely connected.
Webbinelloidea hemispherica Stewart and Lampe
Pl. 35, figs. 9,10; Plessiaigad
1947. Webbinelloidea hemispherica Stewart and Lampe, Jour. Paleont., vol.
Zi eNos6. ps 555siplaeoyetiosyelansp:
Description. — Test attached, apparently monothalmous, cham-
ber hemispherical and strongly convex, basal edge slightly rounded;
wall thick, moderately to coarsely arenaceous, well cemented; no
apparent aperture.
Measurements. — Hypotype 34714 has a diameter of 0.21 mm
and height of 0.11 mm. Hypotype 34715 diameter is 0.25 mm and
height 0.16 mm.
Occurrence. — Hypotype 34714 is from locality UWA 5990 and
34715 from locality UWA 6010, also present at several other locali-
ties and stratigraphic horizons.
Discussion. — This species differs from W. hattini on its lesser
convexity. Stewart and Lampe (1947, p. 535) separated this species
from W. similis on the basis of its unilocularity, and in the posses-
sion of a thicker basal wall and higher convexity. The first charac-
ter was probably due to an insufficient number of specimens. None
of the material studied here shows any definite evidence of being
other than single chambered, and, therefore, is all referable to W.
hemispherica. W. similis presently is only known from the Middle
Devonian of Ohio, whereas W. hemispherica is known at horizons
ranging from Middle Silurian to Middle Devonian. W. hemispherica
specimens recognized here do not compare adequately with those
of Conkin and Conkin (1970) to be similarly considered a W.
similis as revised by them.
Webbinelloidea aff. W. nodosa Summerson
Pl 35, figs: 117 15; Pl) 36, tice2
1958. Webbinelloidea nodosa Summerson, Jour. Paleont., vol. 32, No. 3, p.
555, pl. 82, fig. 12.
Description. — Test attached, apparently monothalmous, square
outline, ventral surface planar, dorsal surface highly convex with
vertical sides; upper surface divided into subequal quadrants by
rectilinear sutures and each quadrant is convex forming a node;
wn
StturRo-DEVONIAN MiIcroFAUNA: MCCLELLAN 3]
junction between dorsal and ventral surfaces forms rounded right
angle; wall thin to medium, composed of sand grains, well cemented;
no apparent aperture.
Measurements. — Figured specimen 34716 has length of 0.26
mm, nodes 0.13-0.14 mm, height 0.19 mm, wall thickness 0.015 mm.
Specimen 34717 has length 0.30 mm, nodes of 0.15 mm, height of
0.22 mm, and wall thickness 0.01 mm.
Occurrence. — Specimen 34716 from locality UWA 6018; 34717
from locality UWA 6010, both rare in occurrence,
Discussion. — Summerson (1958 p, 555) characterized Webb-
inelloidea nodosa by its two rows of nodes on the upper surface
and its rectangular outline. He further described the species to be
roughly hemispherical in section, and to have a slight peripheral
lip where the basal surface joins the upper. The Nevada speci-
mens are more nearly rectangular in section, the sides being verti-
cal, however, they possess the nodes and similar outline. They also
differ in the junction of the upper and lower surfaces, appearing
as a rounded angle instead of a peripheral lip. Because of the sev-
eral similar features, as well as the obvious differences also pres-
ent, the material herein is deemed to have affinities for W. nodosa.
Criteria for erecting a new species are considered insufficient,
though perhaps these specimens, in view of the generic revision
of Webbinelloidea by Conkin and Conkin (1970), should be con-
sidered a species of Ateltkamara, n. gen. There is a similarity in
development of incipient chambers and no apparent apertures.
Webbinelloidea ventriquetra McClellan PIS35e fice 14:
1966. Webbinelloidea ventriquetra McClellan, Bull. Amer. Paleont., vol. 50,
No. 230, p. 496, pls. 38, 42, figs. 6, 7a, b.
Description. — Test attached, bilocular, high dorsal convexity,
ventral surface formed by two planar surfaces separated by an
acute angle which bisects the longitudinal axis of the test; cham-
bers closely joined with a rectilinear boundary and deep dorsal
suture; wall of medium thickness, finely arenaceous, moderate
amount of cement; no aperture apparent.
Measurements. — Specimen 34718 has an overall length of 0.56
mm, individual chambers are 0.30 mm and 0.26 mm, and height
is 0.31 mm.
314 BULLETIN 274
Occurrence. — Rare, present only at locality UWA 6024 in the
Roberts Mountains Formation.
Discussion. — This species bears a much sharper angle at the
junction of the dorsal and ventral walls than is characteristic for
the genus. In this respect it is more like species of Hemisphaeram-
mina, but that genus is restricted to unilocular forms, The only
other species of Webbinelloidea which does not have a planar basal
surface of attachment is W. globulosa McClellan (1966) which has
a concave ventral wall. Such a configuration suggests a more ex-
posed living position for the animal in contrast to W. ventriquetra,
whose test is probably indicative of habitation in protected places,
such as in a crack or small crevice.
Family AMMODISCIDAE Reuss, 1862
Subfamily AMMODISCINAE Reuss, 1862
Genus GLOMOSPIRA Rzehak, 1885
1860. Trochammina [in part] Jones and Parker, Quart. Jour., Geol. Soc.
London, vol. 16, p. 304.
1885. Glomospira Rzehak, Naturforsch. Vereins Briinn, Verhandl., vol. 23
(1884), p. 126.
Type species.—Trochammina gordialis Jones and Parker,
(1860, ibid., p. 304).
Following is Cushman’s (1959, p. 96) redescription of the
genus.
Test free, with a proloculum and long, tubular, undivided, second
chamber winding about its earlier coils in various coils in various
planes; wall arenaceous with much cement; aperture at end of the
tube,
This genus frequently occurs in Lower Silurian rocks, but is
poorly known from Middle Silurian through Devonian time.
Glomospira siluriana Ireland Pl. 36, fig. 5
1939. Glomospira siluriana Ireland, Jour. Paleont., vol. 13, No. 2, p. 201,
figs. B-27, 28.
Description. — Test free, proloculus followed by tubular second
chamber coiled in varying planes around it with four or five revolu-
tions, tube inflated; wall thin, finely arenaceous, well cemented;
aperture at end of tube which is slightly uncoiled.
Measurements. — Specimen 34661 has minimum diameter of
0.37 mm and maximum diameter 0.53 mm,
Wt
StLtuRo-DEvontIAN MicroFauna: McCLeLLANn 3
Occurrence. — This species is rare; present only at locality
UWA 6030 in the basal beds of the Roberts Mountains Formation.
Discussion. —Glomospira siluriana is a common. species in
Lower Silurian rocks from other regions. Its occurrence in the basal
Roberts Mountains Formation of Nevada is the youngest yet
recorded. In combination with Metamorphina gibbosa, it may be
useful in correlating the lowest Roberts Mountains beds with Mid-
dle Silurian rocks of Indiana. Further discussion of its stratigraphic
implications is presented in the section on biostratigraphy.
Subfamily TOLYPAMMININAE Cushman, 1928
Genus TOLYPAMMINA Rhumbler, 1895
1879. Hyperammina [in part] Brady, Quart. Jour. Micr. Sci., new ser., vol.
19. pe teeps 33:
1895. Tolypammina Rhumbler, Kong]. Gese!l. Wiss. Gottingen, Nachr, p. 83.
Type species. —Hyperammina vagan Brady (1879, ibid., p. 33,
pl. 24, figs. 1-5).
The generic redescription of Cushman (1959, p. 98) is as
follows:
Test attached, with proloculus and long, tubular, undivided, second
chamber, earliest portion sometimes coiled, later irregular; wall
arenaceous with much cement; aperture at the end of the tube.
Iituotuba Rhumbler (1895) has a similar habit of irregular
growth in the second chamber. In comparing these two genera
Loeblich and Tappan (1964, p. 213) considered that Tolypammina
“differs from Litwotuba in lacking the early streptospiral coil.”
Tolypammina tortuosa Dunn Pl. 36, fig. 4
1942. Tolypammina tortuosa Dunn, Jour. Paleont., vol. 16, No. 3, p. 341,
p. 44, figs. 19-21, 32.
Description. — Test attached, proloculus spherical, followed by
long tubular second chamber which commonly coils around pro-
loculus in early stages, later portion irregularly winding; wall thin,
finely arenaceous, well cemented; aperture at end of tubular second
chamber.
Measurements. — Hypotype 34712 tube diameter is 0.18 mm.
Occurrence. — This species is rare and present only at locality
UWA 6018 in rocks of lower Helderbergian age.
Discussion. — This is a well-known species in Lower Silurian
rocks and has also been widely reported from the Lower Mississip-
316 BULLETIN 274
pian. While its occurrence in Nevada is Devonian and fills a gap
in the record, it is rare and only present locally.
Conkin (1961, p. 305) considered all species of Tolypammuna,
including T. tortwosa, which occur in Lower Mississippian rocks,
to be characteristic in shallow, warm, near shore, calcium rich sedi- —
ments. He found that often the genus is common in environments
which lack other organisms. T. tortuosa occurs here in the fine-
grained basal beds of the McColley Canyon Limestone as part of a
large microfauna. These limestone beds immediately overlie massive
reef units of the Lone Mountain Dolomite. They indicate shallow,
warm, alkaline conditions during deposition, which should have been
quite favorable to species of Tolypammuna.
Superfamily LITUOLACEA de Blainville, 1825
Family HORMOSINIDAE Haeckel, 1894
Subfamily ASCHEMONELLINAE Eimer and Fickert, 1899
Genus ASCHEMONELLA Brady, 1879
1879. Aschemonella Brady, Quart. Jour. Micr. Sci., new ser., vol. 19, pt. 1,
p. 44.
Type species. — Aschemonella scabra Brady (1879, ibid., p. 44,
pl. 27, figs. 1, 2, 4-11).
Loeblich and Tappan (1964, p. 214) redescribed the genus as
given below:
Test free, consisting of tubular or inflated chambers in single or com-
monly branching series; wall thin, finely to coarsely agglutinated,
firmly cemented; apertures several, rounded, at ends of tubular necks.
Described species of this genus are known from the Cretaceous
to Recent. Cushman (1959, p. 90) mentioned that somewhat similar,
partially divided, tubular forms are known as far back as the
Silurian, but he did not document his statement. The only Paleozoic
occurrence, of which the writer is aware, is that listed by Hattin
(1960, p. 2016) from the Middle Silurian Waldron Shale in south-
eastern Indiana.
Aschemonella bastillensis McClellan, n. sp. Pl 36; fis. 8
Description. — Test free, with tubular chambers inflated on
apertural side, constricting ring divides chambers from tubes which
connect them, tubes slightly expanding in direction of chamber
addition; wall thin, finely arenaceous, well cemented; apertures
S1LuRO-DEVONIAN MicroFAUNA: MCCLELLAN SL7
round and single for each chamber, at end of short neck on inflated
side of chamber.
Measurements. — Holotype 34656 has the following dimensions:
maximum chamber diameter 0.11 mm, connecting tube minimum
diameter 0.04 mm and maximum indeterminate (distorted), length
of apertural neck 0.04 mm, and aperture diameter 0.055 mm.
Occurrence. — Rare, present only at locality UWA 5982 in the
Bastille Limestone of Wenlockian age.
Discussion. — The presence of this genus in the Silurian is the
first occurrence of the superfamily Lituolacea, indicating consider-
able evolutionary development of Foraminifera early in their his-
tory. The reported occurrence of the genus in the Waldron Shale
(Hattin, 1960, p. 2016) appears correlative with that in the Bastille
Limestone, and substantiates other biostratigraphic evidence of the
probable age equivalence of these two units.
The lower beds of the Bastille Limestone, from which Aschemo-
nella bastillensis was recovered, are a thick-bedded, coarse-grained,
biosparite. Residues obtained from samples treated with dilute
hydrochloric acid contained abundant crinoidal fragments and com-
mon well sorted and rounded, frosted, medium-sized quartz grains.
These components suggest the sediment was deposited under con-
ditions of relatively high energy.
Family LITUOLIDAE de Blainville, 1825
Subfamily HAPLOPHRAGMOIDINAE Maync, 1952
Genus HAPLOPHRAGMOIDES Cushman, 1910
1910. Haplophragmoides Cushman, U.S. Nat. Mus., Bull. 71, pt. 1, p. 99.
Type species. — Nomionina canariensis WOrbigny (1839, in
Barker-Webb and Berthelot, Hist. Nat. Iles Canaries, vol. 2, pt. 2,
ps AZSixpl. 2, figsn33.n34):
Following is the redescription of Cushman (1959, p. 102):
Test free, planispiral, of several coils, usually not completely involute;
chambers simple; wall single, arenaceous or with sponge spicules,
firmly cemented, amount of cement varying greatly in different species;
aperture simple, at the base of apertural face of the chamber.
This well-known and common genus is here reported for the
first time from strata older than Carboniferous, in beds of lowest
Devonian age.
318 BULLETIN 274
Haplophragmoides antiquus McClellan, n. sp. Rit ie tigsadle?,
Description. — Test free, planispiral and involutely coiled,
seven chambers in last whorl; chambers slightly inflated, narrowly
arcuate in section; sutures radiate and partially indistinct; prom-
inent, broad, umbilical plug present; wall finely arenaceous, grains
aligned along trace of sutures, moderately cemented; aperture a
thin interiomarginal slit, small accessory apertures near basal-lateral
surface of chambers in last whorl.
Measurements. — Maximum diameter of holotype 34792 is 0.31
mm and axial diameter is 0.11 mm.
Occurrence. — Rare, only found at locality UWA 6010 in the
Rabbit Hill Formation of lowest Helderbergian age.
Discussion. — The two most distinctive features of Haplophrag-
moides antiquus are the broad umbilical plug and pinpoint accessory
apertures. The latter are only recognizable on electron micrographs.
The presence of these apertures has not previously been noted on
any known species of Haplophragmotides. There is the possibility
that H. antiquus is generically distinct, but until the type species
of Haplophragmoides is studied by electron microscopy for presence
of similar apertures, such distinction cannot be made, Other char-
acters of the species appear to be rather conservative when com-
pared to the various complexities which have been added in many
of the other species of the genus (1.e. the strongly lobate chambers
of H. canariensis). This new species compares most nearly to H.
pusilum Hoglund (1947) from the Recent. The latter species has
similar sutures and manner of growth but lacks the umbilical plug
and is more robust.
Suborder FUSULININA Wedekind, 1937
Superfamily ENDOTHYRACEA Brady, 1884
Family ENDOTHYRIDAE Brady, 1884
Subfamily LOEBLICHIINAE Cummings, 1955
Genus NANICELLA Henbest, 1935
1935. Nanicella Henbest, Washington Acad. Sci., Jour., vol. 25, p. 34.
Type species. — Endothyra gallowayi Thomas (1931, Jour.
Paleont., vol. 5, No. 1, p. 40, pl. 7).
Description. — Test free, planispirally coiled, mostly evolute,
numerous low chambers; wall finely granular calcareous, two layers
SILURO-DEVONIAN MicroFAuNA: MCCLELLAN 319
which may be similar or the outer one darker and thinner; aperture
interiomarginal equatorial slit,
Thomas (1931, p. 40) originally defined the type species as be-
longing to the genus Endothyra Phillips (1846). Henbest (1935, p.
34) considered the species to be generically different from other
Endothyra species. His considerations were based on the less inflated
chambers, more regular coiling, and the distinct boundary between
the spiral and septal walls possessed by the /. galloway1, which he
then placed as the type species for Nanicella.
This genus has been previously recorded from rocks no older
than Devonian in Iowa and the USSR. Its presence in Nevada ex-
tends its range to very earliest Devonian time (lower Helderberg).
Nanicella dainae? Chernysheva Pl. 36, figs. 9, 10
1940. Nanicella dainae Chernysheva, Soc. Nat. Moscow Bull., new ser., tome
48 (Sect. geol. tome 18), No. 5-6, p. 127.
Description. — Test free, discoid, planispirally coiled, evolute,
laterally compressed; chambers slightly inflated, almost of equal
height in each whorl, number 12-14 in last whorl; sutures irregularly
radial, shallow; wall finely granular, two layers unrecognizable;
aperture interiomarginal slit.
Measurements. — Diameter of 34677 is 0.70 mm and thickness
0.18 mm. Specimen 34678 has a diameter of 0.52 mm and thickness
0.15 mm.
Occurrence. — The figured specimens were found at locality
UWA 6013 in the upper Roberts Mountains Formation of lowest
Helderbergian age. Also present at locality UWA 5973 in the Rabbit
Hill Formation, slightly younger in age.
Discussion. — The specimens studied here possess all the fea-
tures of Nanicella dainae, with the possible exception of the narrow,
irregularly radial sutures. Preservation is inadequate to positively
recognize the sutures, hence the questioned identification. Com-
parisons with other Nanicella species shows N. dainae to be most
closely related to the type NV. gallowayi. Chernysheva (1940, p. 127)
considered N. dainae to have a somewhat more robust test with a
rounded, rather than acute, periphery, and less distinct sutures. In
addition, the chambers of N. gallowayt are more rapidly expanding
in each whorl and have a much greater height.
320 BULLETIN 274
Namcella dainae has previously been recorded only from Lower
Carboniferous (lower Visean) rocks in the southern Ural Moun-
tains of the USSR.
Subclass OSTRACODA Latreille, 1806
Order PALAEOCOPIDA, Henningsmoen, 1953
Suborder BEYRICHICOPINA Scott, 1961
Superfamily BEYRICHIACEA Matthew, 1886
Family BEYRICHIIDAE Matthew, 1886
Genus BOLBIPRIMITIA Kay, 1940
Type species. —Halliella fissurella Ulrich and Bassler, 1923.
Bolbiprimitia teresaccula Swartz and Whitmore Pl. 41, figs. 1-3
1956. Bolbiprimitia teresaccula Swartz and Whitmore, Jour. Paleont., vol. 30,
No. 5, p. 1082, pl. 104, fig. 18.
Description. — Carapace subovate; straight hinge extending for
nine-tenths of entire length; anterior cardinal angle obtuse, pos-
terior a right, or slightly acute angle; valve strongly convex, anterior
margin broadly rounded; flattened border extends along margins,
with velate ridge in male valves; Ss forms a small rounded depres-
sion slightly anterior of midpoint.
Measurements. — Figured specimens, all male valves, have the
following dimensions:
PLOEX Spec. Number Length Height
Fig. 1 34731 1.04mm 0.67mm
Fig. 2 34732 0.64mm 0.44mm
Fig. 3 34735 0.72mm 0.52mm
Occurrence. — The hypotypes were all found at locality UWA
5974; also present at locality UWA 5975. Both occurrences are in
the upper Roberts Mountains Formation.
Discussion. — The original type for this species is a female
valve, All the specimens found here are male valves, which com-
pare elosely in all characters of the shell, and in addition have a
velate ridge. The elongate dimorphic pouch occupies the position
of the velgate ridge in female valves. The small rounded Ss on
these specimens is distinctive from the elongate cleft of Bolbiprimitia
limbata. Otherwise the species is similar.
Stturo-DeEvonian MicroFAuNA: McCLeLLan 3
bo
fomnk
B. teresaccula has previously been known from the Manlius
Limestone of uppermost Silurian age in New York. Its presence in
the upper beds of the Roberts Mountains Formation in central
Nevada, appears to represent approximately the same stratigraphic
position.
Genus KLOEDENIA Jones and Holl, 1886
Type species. — Beyrichia wilckensiana Jones, 1855.
Kloedenia aparchoides Swartz and Whitmore IAL Gaby aay y/
1956. Kloedenia aparchoides Swartz and Whitmore, Jour. Paleont., vol. 30,
No. 5, p. 1067, pl. 103, figs. 11-14.
Description. — Carapace subovate, dorsally truncated, anterior
cardinal angle obtuse, posterior nearly a right angle; hinge line long,
straight; convex anterior margin; valve moderately convex, slightly
trilobate; S» slightly posterior to middle, S; halfway from S. to
margin; flattened border along margin.
Measurements. — Specimen 34744, not mature, has a length of
0.70 mm and height of 0.42 mm.
Occurrence. — Present only at locality UWA 6024 in the Rob-
erts Mountains Formation.
Discussion. — This rarely occurring species is represented here
by an immature individual which does not have the border as well
developed as in the types of Swartz and Whitmore (1956). In
addition, the lobation and sulci are barely recognizable, which is
also probably due to the immaturity of the specimen.
Genus MYOMPHALUS Swartz and Whitmore, 1956
Type species.— Myomphalus dorsinodus Swartz and Whit-
more, 1956.
Myomphalus ? sp. Pl. 41, fig. 13
Description. —Carapace nearly subrectangular, only slightly
ovoid and dorsally truncated; hinge straight, three-quarters of
length of shell, cardinal angles obtuse; shell moderately convex,
S, and Ss shallow, elongate, poorly developed; exterior and pos-
terior margins convex, marginal border distinct and narrow.
Measurements. — The figured specimen has a length of 0.74
mm and height of 0.44 mm.
Occurrence. — Specimen 34750 is from locality UWA 5981
BULLETIN 274
o>)
bo
bo
in the lower Wenban Limestone. Also present in the McMonnigal
Limestone at locality UWA 5994. Both formations are of Helder-
bergian age.
Discussion. — The genus Myomphalus is described as being
subovoid, with a weak Sz and bordering rounded knobs. On the speci-
mens found here, the median sulcus is bordered by two raised areas,
but there is also an indication of an elongate S,. The shell outline
varies from the characteristic subovoid form, being nearly rectangu-
lar with convex anterior and posterior margins, The shell outline
may appear subrectangular because of an elongate dimorphic pouch
which is ill-defined. The material recovered here is insufficient and
not adequately preserved to be specifically identified, or even defin-
itely recognized as belonging to the genus Myomphalus.
Genus PHLYCTISCAPHA Kesling, 1953
Type species. —Phlyctiscapha rockportensis Kesling, 1953.
Phlyctiscapha keslingi Copeland Pl. 41, figs. 5, 6
1962. Phylctiscapha keslingi Copeland, Geol. Sur. Canada, Bull. 91, p. 36, pl.
IX, figs. 20-24.
Description. —Carapace subovaie, dorsally truncate; hinge
straight, about two-thirds total length, dorsal corners rounded,
cardinal angles obtuse; shell strongly convex, greatest width in
female is median and in male is posteromedian; anterior and pos-
terior margins convex, entire margin and narrow border separated
from lateral shell surface by a furrow; left valve overlaps right valve,
especially along ventral margin.
Measurements. — The length of 34756 is 0.68 mm and height
0.38 mm. Specimen 34757 has length 0.56 mm and height 0.44 mm.
Occurrence. — The figured hypotypes are from locality UWA
5985 in the Lower Devonian McMonnigal Limestone, where the
species is abundant. It has been found widely distributed through-
out the area in rocks no older than upper Ludlow in age.
Discussion. — One of the most abundant ostracode species in
central Nevada, Phlyctiscapha keslingi is restricted to rocks of
latest Silurian and early Devonian age, In every case it occurs in
fore-reef carbonates or calcareous shales. The figured specimens ap-
pear to be representative adults in size but are less than one-half
the size of Copeland’s types. Perhaps the Nevada specimens are
dwarfed, though their abundance would suggest that they were
a)
S1turo-DEVONIAN MicroFauNnA: McCiLe__an Side
quite satisfied in the environment represented here. If they are not
dwarfs, then Copeland’s specimens may be abnormally large in-
dividuals.
Phlyctiscapha sp. Pl: 41, fig. 4
Description. — Carapace elongate subovoid, dorsally truncate;
hinge straight, about two-thirds total length, dorsal corners rounded,
cardinal angles obtuse; shell strongly convex, greatest width 1s
median; anterior and posterior margins strongly convex, entire
margin with narrow border separated from lateral surface of valve
by shallow furrow.
Measurements. — Figured specimen 34758, a female valve,
has a length of 0.41 mm and height 0.27 mm.
Occurrence. — Specimen 34758 is from locality UWA 5991 in
the lower McColley Canyon Formation which is Helderbergian in
age. Also present in the Roberts Mountains Formation at locality
UWA 6024.
Discussion. — This form is quite similar to P. keslingi in most
of its characteristics. It differs in being more elongate in outline
and has a more broadly convex posterior margin. A further con-
sideration is the occurrence, Phlyctiscapha sp. was found in dolo-
mite and argillaceous, cherty limestone samples, which represent
much different environments than the fore-reef carbonates and cal-
careous shales in which P. keslingt was found.
Genus SACCARCHITES Swartz and Whitmore, 1956
Type species. —Saccarchites saccularis Swartz and Whitmore,
1956.
Saccarchites saccularis Swartz and Whitmore Pl. 41, fig. 12
1956. Saccarchites saccularis Swartz and Whitmore, Jour. Paleont., vol. 30,
No. 5, p. 1080, pl. 103, figs. 18, 19.
Description. — Carapace subovate in outline, truncate dorsally;
hinge straight, about four-fifths of length, cardinal angles obtuse and
subequal; shell moderately convex, greatest width is median; an-
terior margin more convex than posterior, narrow submarginal bor-
der at both ends but Jacking on ventral margin.
Measurements. — Length is 0.37 mm and height 0.26 mm on
specimen 34772.
Occurrence. — Rare; at locality UWA 6026 in the Roberts
Mountains Formation.
324 BULLETIN 274
Discussion. — The specimen figured here appears to be an
immature instar in which the lobation and sulcus are not recogniz-
able. Adult specimens from the uppermost Silurian Manlius Lime-
stone of New York are about five times larger (Swartz and Whit-
more, 1956, p. 1080). The Manlius examples also are somewhat
younger than those of central Nevada, which are still within the
Upper Silurian.
Genus VELIBEYRICHIA Henningsmoen, 1954
Type species. — Beyrichia moodeyi Ulrich and Bassler, 1908.
Velibeyrichia? sp. Pl. 41, fig, 16
Description. —Carapace subovoid, elongate; hinge straight,
nearly equal to shell length, cardinal angles both near right angles;
margins of valve with well-developed, striated frill, which is at a
sharp angle to the plane of contact between valves; greatest width
is apparently slightly postero-median, median sulcus is a shallow
groove extending to midpoint of height, poorly developed lobes
border this sulcus.
Measurements. — Figured specimen 34785 has a length of 0.93
mm and indeterminate height about % of length.
Occurrence. — Present only at locality UWA 5985 in the Mc-
Monnigal Limestone.
Discussion. — All the specimens recovered here are poorly pre-
served and none adequately exhibit all the features of the genus.
The lobation is less distinct than normal for the genus, but the frill
is usually mostly preserved and the general shape is that of Veli-
beyrichia. The figured specimen appears to be a male left valve with
only the anterior portion of the frill preserved,
Genus WELLERIOPSIS Swartz and Whitmore, 1956
Type species. — Welleriopsis diplocystulis Swartz and Whit-
more, 1956.
Welleriopsis jerseyensis (Weller) Pl. 41, figs. 8, 9
1903. Beyrichia jerseyensis Weller, Geol. Sur. New Jersey, vol. 3, p. 255, pl.
ey anes, 5.
1956. Welleriopsis jerseyensis (Weller), Swartz and Whitmore, Jour. Paleont.,
vol. 30, No. 5, p. 1075, pl. 105, figs. 14-16.
Description.—Carapace subovate, dorsally truncated by
straight hinge line for three-fourths of length; anterior cardinal
angle bluntly obtuse, posterior angle more sharply obtuse; anterior
SILURO-DEVONIAN MicroFAUNA: McCLELLAN 325
margin convex with greatest point anteroventral, posterior margin
straight in upper part, curving gently into ventral margin, low
marginal ridge is most prominently developed anteriorly; lateral
surface moderately convex, median sulcus as narrow moderately
deep furrow extending from hinge to two-thirds the distance to
margin, L» well developed, broad, L slightly produced dorsally.
Measurements. — Specimen 34786 length is 0.74 mm and height
0.48 mm; 34787 measures 0.67 mm and 0.41 mm.
Occurrence. — Uncommon, found only in locality UWA 5973
in the Rabbit Hill Formation.
Discussion. — The two specimens figured here appear to be
immature and do not have the marginal ridge well developed ven-
trally. The occurrence of this species in Nevada is in strata of
Helderbergian age, younger than the uppermost Silurian occur-
rence recorded to date in eastern North America.
Beyrichiidae ? genus Pl. 41, fig. 16
Description. — Carapace broadly oval in outline, dorsally trun-
cated; hinge gently concave dorsally, two-thirds of length; cardinal
angles obtuse, anterior one more bluntly so; anterior margin mod-
erately convex, posterior slightly convex, maximum height from
mid-ventral point, narrow marginal border bounded by small fur-
row; surface strongly convex, abundant small spines and few large
spines, Ss a long, deep furrow slightly anterior of midpoint and
reaching three-fourths of height toward ventral margin, S; shallow,
curving around posterior of prominent median Ly», L; is broad, less
prominent; dimorphic pouch enlarged anteroventrally, merging dor-
sally with L,; extends ventrally to pceint directly below Ly».
Measurements. — Length of 34789 is 0.30 mm and height 0.75
mm.
Occurrence. — Rarely present at locality UWA 6024 in the
middle Roberts Mountains Formation.
Discussion. — The lobation and dimorphic pouch suggest that
this form belongs in the Beyrichiidae. The figured specimen, to
which the description refers, is believed to be a female right valve.
Another specimen, which has been broken, apparently lacks the an-
teroventral dimorphic pouch and is probably a male. The two speci-
mens are not sufficiently well preserved to compare closely with
any particular beyrichiid genera, and may represent a new genus.
oS)
bo
ON
BULLETIN 274
Superfamily DREPANELLACEA Ulrich and Bassler, 1923
Family AECHMINIDAE Boucek, 1936
Genus AECHMINA Jones and Holl, 1869
Type species. — Aechmina cuspidata Jones and Holl, 1869.
Aechmina equilateralis Bassler Pl. 41, figs. 18, 19
1941. Aechmina equilateralis Bassler, Washington Acad. Sci., Jour., vol. 31,
No. 1, p. 24, fig. 14.
Description. —Carapace nearly equilateral, greatest height
one-third of length from anterior; hinge straight, long; cardinal
angles equally obtuse; anterior and posterior margins convex, an-
terior slightly more so, ventral margin has maximum convexity in
anteroventral area, entire margin denticulate; lateral surface great-
est convexity anteroventral, stout mid-dorsal spine with broad base
is directed slightly posteriorly.
Measurements. — Specimen 34721 has a length of 0.69 mm and
height 0.37 mm. Specimen 34722 length is 0.74 mm and height 0.44
mm.
Occurrence. — This is a commonly occurring species at locality
UWA 5973 in the Rabbit Hill Formation. Fragments found at
locality UWA 6024 have questionably been referred to this species.
Discussion. — No mention was made by Bassler (1941, p. 24)
of dimorphism in this species. The figured hypotype 34772 appears
slightly more enlarged in the anteroventral region of the valve,
thereby distorting the equilateral outline. This enlargement may
possibly be the representation of a dimorphic pouch.
Aechmina longior Bassler Pl. 41, fig. 14
1941. Aechmina longior Bassler, Washington Acad. Sci., Jour., vol. 31, No. 1,
p. 24, fig. 13.
Description. — Carapace subequilateral, greatest height at mid-
point; hinge straight, long, cardinal angles equally bluntly obtuse;
anterior and posterior margins convex, entire margin denticulate;
lateral surface has greatest convexity slightly anterior of midpoint,
mid-dorsal spine is very long, slender, directed posteriorly.
Measurements. — Dimensions of hypotype 34723 are length
0.59 mm and height 0.35 mm,
Occurrence. — Present in the Rabbit Hill Formation only at
locality UWA 5973.
SttuRo-DEvoNnIAN MicroFAuNA: McCLeLLAN 327
Discussion. — This species is similar to Aechmina equilateralts
except for the dorsal spine. A. Jongior is distinguished by the very
long and slender dorsal spine, which is quite distinct from that of
A, equilateralis. Both species are present at the same locality; the
latter occurring in greater numbers.
Aechmina aff. A. phantastica Kesling Pi ate fie a
1953. Aechmina phantastica Kesling, Mich. Univ., Mus. Paleont. Contr., vol.
Wi joy So. fol, al ames, Joh,
Description. —Carapace subrectangular, dorsally truncate;
hinge straight, about three-fourths of length; cardinal angles obtuse,
subequal, posterior margins moderately convex, ventral margin
slightly convex; dorsally directed, slender curved spines at point of
cardinal angles; lateral surface moderately convex, broad based,
stout, mid-dorsal spine directed perpendicular to commissure plane
and slightly posteriorly, and spine is bluntly terminated.
Measurements. — Length of specimen 34724 is 0.71 mm and
height is 0.50 mm.
Occurrence. — Rare, present only at locality UWA 5973 in
the Rabbit Hill Formation.
Discussion. — The form and position of the prominent mid-
dorsal spine on this specimen is identical to that of Aechmina
phantastica. Overall dimensions are also quite similar. This form dif-
fers in possessing long, slender, curved spines arising from the ends
of the hinge line. It also has a less semicircular outline, greater
width, and the major spine is not granuloreticulate as in A.
phantastica. The extra dorsal spines (posterior one is broken at
base) are the most significant characteristics of this form. Coupled
with the other differences from A. phantastica, it probably repre-
sents a separate, new species, but the single ?LV found here is
believed to be insufficient for establishment of a species.
Aechmina cortezensis McClellan, n. sp. Pl. 41> figss 1i15
Description. — Carapace ovoid to subellipsoidal, partially trun-
cated dorsally; hinge straight, about two-thirds length, gradually
merging with margin at ends; anterior and posterior cardinal angles
subequal, broadly obtuse; anterior margin and posterior are equally
stongly convex, ventral margin moderately convex; lateral surface
moderately convex with greatest width median to slightly antero-
median; dorsal median spine broad based, long, directed outward at
328 BULLETIN 274
angle of 40-60 degrees from commissure plane, distal half may
curve slightly posteriorly.
Measurements. — Holotype 34725 has length of 0.57 mm,
height 0.30 mm, and length of spine 0.41 mm, Paratype 34726, a
RV, has length 0.55 mm and height 0.29 mm.
Occurrence. — The type specimens are present at locality UWA
5980 in the lower Wenban Limestone.
Discussion. — Aechmina cortezensis is most similar to A. longior
Bassler (1941) which is present in probable correlative beds of the
Rabbit Hill Formation. The dorsal median spine and shell size of
A. cortezensts are similar to A. longior. This species differs, however,
in having more strongly convex anterior and posterior margins,
broader cardinal angles, shorter hinge line, and lacks the denticles on
the margin,
There is some variability in the shape of the valves in this new
species, the most typical being subellipsoidal, but they may also be
nearly ovate. In addition there is an indication of Jateral swelling in
the anteroventral area, as on figured specimen 34726, that may be
distortion or possibly indicate a dimorphic pouch.
Family BOLLIIDAE Boucek, 1936
Genus ULRICHIA Jones, 1890
Type species. — Ulrichia conradi Jones, 1890.
Ulrichia obesita McClellan, n. sp. Pipa ehicshai-s
Description. —Carapace subrectangular to ovoid; hinge long,
straight; cardinal angles obtuse, subequal; anterior and _ posterior
margins equally convex, marginal rim distinct, forming sharply
defined, narrow shelf; lateral surface strongly convex, finely reticu-
late in concentric pattern in adult, marginal rim more coarsely reticu-
late; anterior dorsomedian node round, posterior node forms narrow,
strongly developed ridge with dorsal end more posterior than ventral
end, Ss forms a narrow short, deep groove between nodes.
Measurements. — Dimensions of the figured specimens are as
follows:
Pl. 42 Spec. Number Length Height
Fig. 1 34780 holotype 0.67mm 0.43mm
Fig. 2 34782 paratype 0.44mm 0.30mm
Fig. 3 34783 paratype 0.37mm 0.26mm
SILURO-DEVONIAN MiIcrRoFAUNA: MCCLELLAN 329
Occurrence. — The type specimens were present at locality
UWA 5985 in the McMonnigal Limestone. Also present at locality
UWA 5986 in the same area and formation.
Discussion. — This species differs from others within the genus
in the shape of the dorsomedian nodes, the posterior one being a
ridge rather than the characteristic subcircular form. It is also
much more highly convex laterally, than is customary for species of
Ulrichia. The reticulate surface pattern is recognizable only on adult
forms, such as the holotype. The reticulation continues, more coarse-
ly, on the narrow, marginal rim, which is not the usual pattern for
the genus.
Ulrichia sp. Pl. 42, figs. 4, 7
Description. —Carapace subrectangular, truncate dorsally;
hinge long and straight; cardinal angles subequal; margin convex,
narrow, well-defined rim; lateral surface moderately elongated,
sharply defined, posterior node smaller, subcircular, Ss shallow,
broad, extending down to midpoint.
Measurements. — Specimen 34783 length is 0.92 mm and height
0.61 mm. Dimensions of 34784 are 0.78 mm and 0.48 mm,
Occurrence. — Present only at locality UWA 5973 in the Rab-
bit Hill Formation.
Discussion. — This form differs from Ulrichia spinosa in the
shape of the nodes but otherwise has many similarities. It also
resembles U. acricula in the position and form of the nodes. How-
ever, the marginal rim is narrower and the shell is more robust than
the lowly convex shell of U. acricula.
Bolliidae ? genus PIS 43 hiess 16h 7
Description.—Carapace subquadrate, dorsally truncated;
hinge long, straight, nine-tenths of length; cardinal angles equal,
obtuse, nearly right angles; anterior margin gently convex, pos-
terior slightly convex, marginal rim very narrow and sharply de-
fined; lateral surface strongly convex, greatest width at midpoint;
anterior node slightly elongate, sharply elevated, positioned just
anterior of midpoint; posterior node forms linear, prominent, knife-
like ridge beginning at dorsal margin and merging posteroventral of
midpoint into a lower, well-defined ridge that curves ventrally
around anterior node and continues up and around until terminat-
330 BULLETIN 274
ing at dorsal margin directly above anterior node; S; forms shallow,
arcuate depression between anterior median node and curving
ridge, S2 is deep, linear groove slightly posterior of midpoint,
Measurements. — The length of 34734 is 0.38 mm and height
0.29 mm. Specimen 34735 has been broken but dimensions are ap-
proximately 0.45 mm and 0.35 mm.
Occurrence. — The figured specimens are from locality UWA
5985. Also present at UWA 5994 and UWA 5975 in strata of latest
Silurian to early Devonian age.
Discussion. — This form has affinities for both Ulrichia Jones
(1890) and Bollia Jones and Holl (1886). The separate anterior
dorsomedian node is similar to that of Ulrichia. The posterior node
and ventral ridge extending from it are suggestive of Bollia. The
ridge in Bollia connects with an anterior lobe, whereas the ventral
ridge on this form extends instead in an arcuate line around the
anterior node to the dorsal margin. This last feature is distinctive
from other bolliids, thus a generic placement cannot be made. There
is neither sufficient, nor adequately preserved material to be de-
scribed.
Superfamily HOLLINACEA Swartz, 1936
Family HOLLINIDAE Swartz, 1936
Genus HOLLINELLA Coryell, 1928
Type species. — Hollinella dentata Coryell, 1928.
Hollinella sp. Pl. 42, fig. 6
Description. — Carapace ovoid, truncated dorsally; hinge line
long, straight; cardinal angles obtuse, anterior more broadly so; an-
terior margin strongly convex, posterior tapers smoothly into ven-
tral margin, well developed, broad striate frill extends along entire
margin; lateral surface strongly convex, Ls well developed, round-
ed node, Ls low, broad, and indistinct; median sulcus between lobes
is shallow, short, extending only in curving pattern along posterior
side of Le.
Measurements. — Dimensions of 34743 are length 0.76 mm,
height 0.51 mm and width of frill 0.15 mm.
Occurrence. — Present only at locality UWA 6024 in the Rob-
erts Mountains Formation.
S1turRo-DEVoONIAN MicroFAuNA: McCLeELLAN 331
Discussion. — Well-preserved specimens of this form are rare,
and, therefore, specific identification could not be made. The median
sulcus is not as broad as normal for most species of the genus, and
the L3 is poorly developed.
Family SIGMOOPSIDAE Henningsmoen, 1953
Genus WINCHELLATIA Kay, 1940
Type species. — Winchellatia longispina Kay, 1940.
Winchellatia fragilis McClellan, n. sp. Pl) 42: fig.5
Description. —Carapace ovate, dorsally truncate; hinge line
long, straight; anterior cardinal angle barely obtuse, posterior is a
right angle; anterior margin gently convex, ventral nearly straight,
curving evenly into posterior margin which is straight; lateral sur-
face has greatest width posteroventral of midpoint; S; very shallow,
narrow groove, S» deep, beginning at midpoint and widening from
a narrow groove upward to near dorsal margin where it bends
slightly forward; Le forms rounded low node outlined by S; and Sz,
L3 an indistinctly raised area with greatest height posteroventral;
histial ridge fine and sharply defined extends from near anterior
cardinal angle around ventral part of valve, ending posteroventrally
where posterior margin begins; ventromedian based spine extends
laterally from broad base, with thickened posterior margin, other-
wise very thin and flat, appears winglike with backward directed
tip.
Measurements. — The holotype male, 34788, has length of 0.42
mm and height 0.34 mm,
Occurrence. — Present only at locality UWA 6024, where it is
rare in Roberts Mountains beds of Upper Silurian age.
Discussion. — The fragile winglike spine on each valve is dis-
tinctive from similarly placed nodes on other species in the genus.
In other characters this new species resembles Winchellatia minne-
sotensis Kay (1940) which has an acute posterior cardinal angle,
while that of the former is a right angle.
Previously the genus has been reported only from Middle
Ordovician strata.
>)
Los)
bo
BULLETIN 274
Superfamily PRIMITIOPSACEA Swartz, 1936
Family PRIMITIOPSIDAE Swartz, 1936
Subfamily PRIMITIOPSINAE Swartz, 1936
Genus LIMBINARIA Swartz, 1956
Type species. — Limbinaria multipunctata Swartz, 1956.
Limbinaria sp. Pl. 42, figs. 8, 9
Description. —Carapace subrectangular, dorsally truncate;
hinge long and straight; cardinal angles obtuse, subequal; anterior
and posterior margins lowly convex, ventral margin nearly straight;
lateral surface has large, slightly elevated medial area with sub-
rounded median pit; continuous narrow ridge parallels margins,
separating them from medial area, is discontinuous dorsally; sur-
face of valve coarsely punctate.
Measurements. — Specimen 34746 length is 0.92 mm and height
indeterminate.
mm. Specimen 34746 length is 0.92 mm and height indeterminate.
Occurrence. — Rare, only present at locality UWA 6024 in the
Roberts Mountains Formation.
Discussion. — The presently known specimens of this form are
not well enough preserved to be specifically identified. They are
similar to Limbinaria multipuncta Swartz (1956), except in having
a subrectangular outline. The dorsal marginal area appears to vary
somewhat also with a different configuration of the ridge that bor-
ders the media] area.
Suborder KLOEDENELLOCOPINA Scott, 1961
Superfamily KLOEDENELLACEA Ulrich and Bassler, 1908
Family KLOEDENELLIDAE Ulrich and Bassler, 1908
Genus EUKLOEDENELLA Ulrich and Bassler, 1923
Type species. — Eukloedenella umbilicata Ulrich and Bassler,
1923:
Eukloedenella aff. E. dalhousiensis Copeland Pl. 42, figs. 10, 11
1962. Eukloecdenella dalhousiensis Copeland, Geol. Sur. Canada, Bull. 91, p.
42, p. X, figs. 1-6.
Description. — Carapace elongate, ovate; hinge straight pos-
teriorly, right valve hinge slightly angulated over left valve at mid-
point; posterior margin convex, ventral nearly straight, anterior
SiLturRo-DEVONIAN MicroFAUNA: MCCLELLAN 333
margin has greatest convexity anteroventrally; greatest width pos-
teromedial, valves lanceolate in dorsal view; median sulcus faint to
indistinguishable.
Measurement. — Specimen 34740 measures 0.45 mm in length
and 0.22 mm in height. Specimen 34741 is 0.37 mm in length and
0.19 mm in height.
Occurrence. — Present only at locality UWA 5985 in the Lower
Devonian McMonnigal Limestone.
Discussion. — The specimens described here compare well to
Eukloedenella dalhousiensis Copeland (1962) in most characters. A
notable exception is the lack of the distinct median sulcus of that
species. Since E. dalhousiensis adults are typically about twice the
size of the specimens here, it is possible that this material either
represents early instars in which the sulcus is not recognizable,
or if such instars should exhibit the sulcus, the use of open nomen-
clature is justified until more detailed information is available.
E.. dalhousiensis has been recorded in the upper Dalhousie beds
of New Brunswick, Canada, which have been called Helderbergian
in age (Copeland, 1962, p. 23). The age of the McMonnigal Lime-
stone, from which these specimens were taken, has also been con-
sidered Helderbergian on the basis of its megafauna (Kay and Craw-
ford, 1964, p. 440).
Genus NEOKLOEDENELLA Croneis and Funkhouser, 1939
Type species.— Neokloedenella prima Croneis and Funk-
houser, 1939.
Neokloedenella ? sp. Pl. 42, fig. 12
Description. —Carapace subrectangular, lanceolate in dorsal
view; hinge straight, overlap indeterminate; anterior cardinal angle
very bluntly obtuse, posterior angle also obtuse; anterior margin
strongly convex, posterior moderately so, ventral margin nearly
straight; lateral surface highly convex, median sulcus unrecogniz-
able, greatest width in posterior third of valve.
Measurements. — Specimen 34751 has length of 0.56 mm and
height 0.29 mm.
Occurrence. — Present at locality UWA 5991 in the lower
McColley Canyon Formation.
Discussion. — The poorly preserved male RV figured here re-
334 BULLETIN 274
sembles the Neokloedenella sp. reported by Stover (1956, p. 1111)
from the Middle Devonian Windom Shale of western New York.
Family GEISINIDAE Sohn, 1961
Genus PSEUDOLEPERDITIA Schneyder, 1956
Type species. — Pseudoleperditia tuberculifera Schneyder, 1956.
Pseudoleperditia ? sp. Pl. 42, ges
Description. — Carapace subquadrate, dorsally truncated; hinge
straight, long; anterior cardinal angle obtuse, posterior forms a right
angle; anterior margin nearly straight, ventral margin curves into
slightly convex posterior; lateral surface strongly convex, greatest
width ventromedian, L» forms a small, rounded node with median
sulcus directly adjacent, forming shallow, elongate depression;
prominent, broad, short spines directed upward from lateral sur-
face of valve at points of the cardinal angles; margin lacks border
but has short denticles anteriorly and ventrally.
Measurements. — Length of 34759 is 0.45 mm and height 0.30
mm.
Occurrence. — Rarely present only at locality UWA 6024 in
the middle Roberts Mountains Formation,
Discussion. — This specimen has been questionably referred to
Pseudoleperditia Schneyder (1956), which is monotypic, on the basis
of the similarity in the lobation and cardinal spines. It seems to
differ in lacking the marginal border of P. twberculifera, having den-
ticles instead, in a straighter hinge line, less anterior convexity,
and in having the greatest width ventromedianly. The last character
may be caused by an elongate brood pouch in this position, The
type of hingement is indeterminate which further hinders positive
identification, as it is an important character of the genus.
Superfamily LEPERDITELLACEA Ulrich and Bassler, 1906
Family LEPERDITELLIDAE Ulrich and Bassler, 1906
Genus PARAHEALDIA Coryell and Cuskley, 1934
Type species. — Parahealdia pecorella Coryell and Cuskley,
1934.
Parahealdia aff. P. pecorella Coryell and Cuskley _ PI. 42, figs. 14, 16, 17
1934. Parahealdia pecorella Coryell and Cuskley, Amer. Mus. Nat. Hist., Nov.
748, text fig. 17.
StturRo-DEvontAn MicroFAuNAa: McCLELLAN 335
Description. — Carapace elliptical, female shorter and more ro-
bust, more sharply convex in posterior margin than in anterior;
hinge long, straight; margins smooth, dorsally slightly convex; lat-
eral surface strongly convex, greatest width in posterior third where
two short spines are directed posteriorly, upper spine at a low angle
posterodorsally, bases of spines separated by narrow, flattened area;
median sulcus very shallow and broad, extending down from dorsal
margin for about one-third of height.
Measurements. — Specimen 34777, female RV, measures 0.46
mm by 0.33 mm.
Occurrence. — Abundant at locality UWA 5980. Also, ques-
tionable specimens present at locality UWA 5981; both in the
lower Wenban Limestone.
Discussion. — These specimens differ from Parahealdia pecorella
in being slightly convex dorsally, the spines are somewhat closer
together, and the sulcus is broader and less distinct. In addition,
female specimens are shorter and more rounded in outline.
Superfamily PARAPARCHITACEA Scott, 1959
Family PARAPARCHITIDAE Scott, 1959
Genus PARAPARCHITES Ulrich and Bassler, 1906
Type species. — Paraparchites humerosus Ulrich and Bassler,
1906.
Paraparchites sp. P42 fis. 15
Description. — Carapace broadly ovate, strongly convex; hinge
about two-thirds of length, cardinal teeth indistinct; small, narrow,
marginal ridge extends posteroventrally to anteroventrally, dorsal
margin nearly straight; surface smooth, nonlobate, nonsulcate.
Measurements. — Length of 34754 is 0.64 mm and height 0.48
mm.
Occurrence. — Figured specimen from locality UWA 6024. Al-
so occurs at locality UWA 5974. Both localities are in the upper
two-thirds of the Roberts Mountains Formation.
Order PODOCOPIDA, Miiller, 1894
Suborder PODOCOPINA Sars, 1866
Superfamily BAIRDIACEA Sars, 1888
Family BAIRDIIDAE Sars, 1888
Genus BAIRDIA McCoy, 1844
Type species. — Batrdia curta McCoy, 1844.
336 BULLETIN 274
Bairdia sp. Pl 43y ficss sles
Description. — Carapace lanceolate in outline, acuminate an-
teriorly, maximum height one-fourth of length from posterior; LV
with short, small spine at midpoint of posterior margin, RV with
short, small spine forming anterior point, LV overlaps RV dorsally
on margin between spines; lateral surfaces smooth, maximum width
near midpoint, tapering gradually to acuminate anterior.
Measurements. — Specimen 34727 length is 0.55 mm plus spines
(0.04 mm each) and height 0.29 mm.
Occurrence. — Present only at locality UWA 6024 in the mid-
dle Roberts Mountains Formation,
Discusston.— The genus Bairdia seems indicated by the man-
ner in which the valves are joined as well as the form of the shell
and presence of small spines. Positive identification and compari-
sons to other, possibly related, species could not be made because
of a lack of material.
Genus BYTHOCYPRIS Brady, 1880
Type species. — Bairdia bosquetiana Brady, 1866.
Bythocypris aff. B. alcocki Copeland Pl. 43, figs. 3-5
1962. Bythocypris alcocki Copeland, Geol. Sur. Canada, Bull. 91, p. 46, pl. X,
figs. 16-20.
Description. — Carapace elongate reniform, broadly ovate pos-
teriorly; hinge straight, in anterior half; LV overlaps RV on entire
margin; greatest height median to posterior, greatest width postero-
median; margin nearly straight ventrally, anterior margin with
maximum point of convexity median to ventral; lateral surface
moderately convex, smooth.
Measurements. — Specimen 34737 is 0.56 mm by 0.29 mm;
34736 measures 0.46 mm by 0.22 mm.
Occurrence. — Specimen 34737 from locality UWA 5974, and
34736 from locality UWA 6026. This form is widely distributed
throughout the area in strata from Ludlovian to Helderbergian in
age. It occurs most frequently in rocks of the latter age.
Discussion.— One of the more common constituents of the
ostracode fauna in central Nevada, specimens referred to this
species show a wide variation in shape. The shel] outline varies from
S1LturRo-DEVONIAN MicroFAUNA: MCCLELLAN 337
reniform to elliptical and the point of greatest width from posterior
to anteromedian in position. Bythocypris alcocki Copeland (1962 )
is reniform and has a maximum width posteriorly. The left valve
overlaps the right along the entire margin, in the same manner as
shown by whole specimens from Nevada. B. phaseolina Ulrich and
Bassler (1923) is similar to B. alcocki but more ellipsoidal and
elongate. The specimens studied herern show some resemblance to
B. phaseolina as well as apparent gradation from that species to
B. alcockhi.
Bythocypris ? sp. Pl. 43, figs. 6
Description. — Carapace subreniform, arcuate dorsally, later-
ally tumid; hinge straight, questionably anterior half; ventral mar-
gin straight to slightly concave, forming acute angles where join-
ing anterior and posterior margins, dorsal margin strongly convex;
lateral surface highly convex, greatest width median to ventro-
median, maximum height at midpoint; valves dorsally depressed
on median marginal area, may be depressed to form flattened area
along ventral margin.
Measurements. — Dimensions of specimen 34739 are length
0.48 mm and height 0.29 mm.
Occurrence. — Figured specimen from locality UWA 5974.
This form is present at several localities in beds ranging from Lud-
low to Helderberg in age.
Discussion. — Species of Bythocypris are commonly reniform
to ovate and not ventrally flattened as is this form, The dorsal de-
pression of the valves along the hinge is suggestive of a similar
structure present on Newsomites Morris and Hill (1952). How-
ever, it is believed that these specimens are generally more compar-
able in form to that of Bythocypris.
Family BEECHERELLIDAE Ulrich, 1894
Genus ACANTHOSCAPHA Ulrich and Bassler, 1923
Type species. — Beecherella navicula Ulrich, 1891.
Acanthoscapha aff. A. navicuia (Ulrich) Pl. 43, figs. 7, 8
1891. Beecherella navicula Ulrich, Amer. Geol., vol. 8, No. 4, p. 203, pl. 2,
figs. 8, 9.
Description. — Carapace elongate, spindle-shaped; hinge long,
>)
Loe)
(oe)
BULLETIN 274
adont; flattened flange of margins greatly reduced; LV with very
short dorsoposterior spine, anterior spine in contact with margin,
RV with anterior end produced into short broad spine,
Measurements. — Specimen 34719, a RV, length is 0.74 mm and
height 0.30 mm; 34720, a LV, length 0.57 mm and height 0.21 mm.
Occurrence. — Present only at locality UWA 5981 in the lower
Wenban Limestone of Lower Devonian age.
Discussion. — These specimens differ from Acanthoscapha navi-
cula, which occurs in Lower Devonian rocks of New York, in several
respects. They are considerably smaller, the anterior and posterior
spines are much shorter, the sheil is less acuminate anteriorly, and
the anterior spines contact the margin. The original types of A.
navicula have a narrow marginal surface between the spine and
margin.
Genus LONGISCAPHA McClellan, n. gen.
Type species. — Longiscapha nevadensis McClellan, n. sp.
Descriptwon. — Carapace elongate, scaphoid, middle third ven-
traliy constricted, anterior spindle-shaped, posterior convex; hinge
straight, long; lateral valve surfaces smooth, strongly convex with
broad dorsum, compressed at ends, LV slightly overreaching RV;
spines lacking, or reduced, on LV only.
The general scaphoid form and adont hinge place this genus
definitely in the family Beecherellidae. The constriction of the
median portion of the valves is similar to that of Ulrichella Boucek
(1936), but is only represented as a height reduction, as the great-
est width of the shell is in the same area. This genus is less acum-
inate than other genera within the family, and the spine is poorly
developed.
Longiscapha nevadensis McClellan, n. sp. Pl. 43, figs. 9-11
Description. — Carapace scaphoid, anterior spindle-shaped; dor-
sal margin straight, anterior margin bluntly acuminate, ventral
margin gently concave through medial third of shell, posterior
strongly convex, anterior and posterior low marginal ridge defined
by very shallow furrow; lateral surface of valve smooth, strongly
convex, slightly convex dorsum, greatest height at end points of
hinge; short, broad based, dorsal spine directed forward from antero-
dorsal position on LY.
SILURO-DEVONIAN MicroFAuNA: MCCLELLAN 339
Measurements. — Holotype 34747 length is 0.59 mm and height
0.22 mm. Paratype 34748 length is 0.53 mm and height 0.19 mm and
paratype 34749 length is 0.64 mm and height 0.22 mm.
Occurrence. — This species was found only at localities UWA
6024 (type material) and UWA 6026 in the upper Roberts Moun-
tains Formation at Willow Creek, which is of probable Ludlow age.
Discussion. — This species is characterized by the depressed
anterior and posterior with the accompanying marginal border, and
the greatest height being at the ends of the hinge line.
At both localities, from which Longiscapha nevadensis was re-
covered, the strata are cherty, argillaceous limestones. The bedding
is very thin, with the shale and chert in laminae and stringers, sug-
gestive of quiet bottom conditions and slow deposition. The micro-
fauna is large and diversified here, but other invertebrate fossils are
rare,
Superfamily CYTHERACEA Baird, 1850
Family BEROUNELLIDAE Sohn and Berdan, 1960
Genus BEROUNELLA Boucek, 1936
Type species. — Berounella rostrata Boucek, 1936.
Berounella sp. Pl. 43, fig. 13
Description. —Carapace subquadrate, with a long spine ex-
tending from posterior dorsal margin; anterior cardinal angle pro-
duced into long, thin spine directed dorsally and curving forward;
lateral surface strongly convex, L; is anteromedian to ventral; deep,
straight sulcus begins at dorsal margin above L; and extends to mid-
point; ventral lobe leading from L, backward until merging with
L;, has long, delicate lateral spine curving posteriorly from postero-
ventral point; L; broad, rising from posterodorsal position to merge
with ventral lobe; marginal flange developed only posteroventrally.
Measurements.— Length of specimen, exclusive of spine is
0.58 mm and height 0.29 mm.
Occurrence. — Only one specimen is known, from locality UWA
6024 in the Roberts Mountains Formation.
Suborder METACOPINA Sylvester-Bradley, 1961
Superfamily HEALDIACEA Harlton, 1933
Family HEALDIIDAE Harlton, 1933
Genus HEALDIA Roundy, 1926
340 BuLLETIN 274
Type species. — Healdia stmplex Roundy, 1926.
Healdia ? sp. PI. 43; fig.-12
Description. — Carapace ovate; dorsal and ventral margins
moderately convex, anterior and posterior both rounded; lateral
surface strongly convex, greatest width posterior, surface smooth
with posterior shoulder bearing two very short and blunt spines.
Measurements. — Specimen 34742 length is 0.63 mm and height
0.39 mm.
Occurrence. — Found rarely, only at locality UWA 5981 in the
lower Wenban Limestone.
Discussion. — This form is poorly represented in the area and
specimens cannot be positively identified as having healdiid hinge-
ment and overlap of the valves. On the basis of shape and surface
features it has been referred to the genus Healdia. The degree and
direction of overlap could not be determined, so it is possible that
the generic assignment is incorrect.
Family CAVELLINIDAE Egorov, 1950
Genus BIRDSALLELLA Coryell and Booth, 1933
Type species. — Birdsallella simplex Coryell and Booth, 1933.
Birdsallella sp. Pl. 43, fig. 14
Description. — Carapace elongate ellipsoidal, RV overreach of
LV noticeable only on dorsal and ventral borders, dorsal view wedge-
shaped; anterior margin convex, posterior nearly truncate, dorsal
and ventral margins straight; lateral surface smooth, greatest width
posterior; posterior swelling developed into strong ridge projecting
backward to margin.
Measurements. — The figured specimen has a length of 0.74
mm and height of 0.45 mm.
Occurrence. — This genus occurs only at locality UWA 5980 in
the lower Wenban Limestone.
Family PACHYDOMELLIDAE Berdan and Sohn, 1961
Genus PHANASSYMETRIA Roth, 1929
Type species. — Phanassymetria triserrata Roth, 1929.
Phanassymetria sp. Pl. 43, fig. 20
Description. —Carapace subovate, asymmetrical, angular in
S1LtuRO-DEVONIAN MicroFAuNA: McCLeELLAN 341
cross-section; dorsum convex, forming longitudinal shoulders with
straight hinge in groove between, ventral margin straight; LV over-
laps RV strongly on venter and is the larger valve; maximum
width slightly posteroventral of midpoint; lateral surface coarsely
punctate.
Measurements. — Length of figured specimen is 1.03 mm and
height is 0.65 mm,
Occurrence. — Present at locality UWA 5980 in the lower
Wenban Limestone, from which only one specimen was recovered.
Discussion. — This specimen differs from the known species of
Phanassymetria, but it is not possible to establish a new species
until additional material has been found.
Genus TUBULIBAIRDIA Swartz, 1936
Type species. — Tubulibairdia tubulifera Swartz, 1936.
Tubulibairdia cf. T. chaleurensis Copeland Pie43. figs, 15,19
1962. Tubulibairdia chaleurensis Copeland, Geol. Sur. Canada, Bull. 91, p. 47,
pl. X, figs. 32-36.
Description. — Carapace ovate, elongated; hinge straight, about
half of total length; dorsal margin slightly convex, anterior and
posterior strongly so, ventral margin straight; LV overlaps RV,
most pronounced ventrally; lateral surface smooth, highly convex,
greatest width posterior giving ovoid shape in dorsal view, maximum
height slightly posterior of median.
Measurements. — Specimen 34776 length is 0.55 mm and height
0.29 mm.
Occurrence. — Specimen 34776 is from locality UWA 6024. This
species is widely distributed in the area in strata ranging from Lud-
lovian to Helderbergian age.
Discussion. — Tubulibairdia chaleurensis has been previously
recorded in the type area of New Brunswick, Canada, where it
occurs in the Lower Devonian Dalhousie beds. The species described
here occurs in strata of similar age. It differs from Copeland’s types
in being more elongate with a straight venter, and the maximum
width is closer to median.
Tubulibairdia aff. T. chaleurensis Copeland Pl. 43, fig. 18
1962. Tubulibairdia chaleurensis Copeland, Geo}. Sur. Canada, Bull. 91, p. 47
pl. X, figs. 32-36.
342 BULLETIN 274
Description. — Carapace ovate; hinge straight, about half of
total length; dorsal and ventral margins gently convex, anterior and
posterior strongly convex; LV overlaps RV except dorsally; lateral
surface smooth, robust with high convexity, greatest width at mid-
point and tapering evenly to both ends, dorsum rises above hinge
line.
Measurements. — Specimen 34778 has a length of 0.63 mm and
height of 0.43 mm.
Occurrence. — This form was found only at locality UWA 5985
in the McMonnigal Limestone.
Discussion. — In general outline, viewed laterally, this form re-
sembles Tubulibairdia chaleurensis more so than the preceding
species. Differences are seen, however, in the median maximum
width, and the valve overlap, which is continuous on the anterior
border in contrast to Copeland’s types. With these significant dif-
ferences the present material has affinities for, but is distinct from,
T. chaleurensis.
Tubulibairdia sp. Pl... 43, figs2t
Description. —Carapace ovate; entire margin convex; overlap
of valves indeterminate; lateral surface smooth, strongly convex,
maximum width posterior.
Measurements. — Specimen 34779 length is 0.66 mm and height
is 0.44 mm.
Occurrence. — Several specimens of single valves were found
at locality UWA 5973 in the Rabbit Hill Formation of Lower
Devonian age.
Discussion. — The single valves occurring at the above locality
appear to belong to Tubulibairdia, although unrecognizable specifi-
cally. Their characters are more generalized than the previously
described species which compare to T. chaleurensts.
Superfamily QUASILLITACEA Coryell and Malkin, 1936
Family QUASILLITIDAE Coryell and Malkin, 1936
Genus QUASILLITES Coryell and Malkin, 1936
Type species. — Quasillites obliquus Coryell and Malkin, 1936.
Quasillites sp. Pl. 44, figs. 1-3
Description. — Carapace subquadrate, ends rounded; LV over-
SttuRo-DEVvoNIAN MicroFAUNA: McCLELLAN 343
laps RV except on dorsal margin; lateral surface moderately convex,
greatest width median, ornamented with fine ridges and grooves;
both valves have flattened dorsum with shoulders forming narrow
ridges for entire length, LV may also have ventral groove setting
off part of valve which overlaps RV; LV has anterior and posterior,
short, blunt spines placed near ventral margin; RV may have similar
matching pair, only one, or no spines.
Measurements. — The figured specimens have the following
dimensions: specimen 34760 has length 0.48 mm and height 0.25
mm and specimen 34761 length is 0.44 mm, height 0.22 mm.
Occurrence. — Present only at locality UWA 5985 in the Mc-
Monnigal Limestone.
Discussion. — None of the known species of Quasillites ade-
quately fits the description of these specimens, which are more
equilateral than described species as well as having a distinctive
development of the lateral spines. Stover (1956, p. 1124) discussed
the problem of variability encountered in Quasillites species. In
the present material considerable variation is suggested, such that
it is deemed advisable not to erect a new species until more speci-
mens are available for study.
Family ROPOLONELLIDAE Coryell and Malkin, 1936
Genus RUDDERINA Coryell and Malkin, 1936
Type species. — Rudderina extensa Coryell and Malkin, 1936.
Rudderina mutaspina McClellan, n. sp. Pl. 44, figs. 48
Description.—Carapace elongated subtriangular, triangle
hypotenuse formed by central margin; hinge straight, extending
for over 0.8 of greatest length; dorsal margin straight, anterior
slightly convex, venter straight to convex rearward, posterior
margin strongly convex; well developed, flattened marginal ridge
anteriorly separated from lateral surface by narrow furrow, similar
posterior ridge mostly indistinct; LV with convex medial portion of
dorsal margin, lateral surface smooth, greatest height anterior, great-
test width posteroventral; single, long, tapering, finely pointed
spine arises from point of greatest width; the spine is directed
anterolaterally in early instars, progressively changing to a postero-
lateral direction in adults.
344 BULLETIN 274
Measurements. — Dimensions of the several instars figured are
listed below:
Pl. 44 Spec. Number Length Height
Fig. 4 34763 paratype 0.47 mm 0.23 mm
Pig. e 34764 paratype 0.47 mm 0.23 mm
Fig. 6 34765 paratype 0.65 mm 0.30 mm
Fig. 7 34766 paratype 0.65 mm 0.30 mm
Fig. 8 34767 holotype 0.76 mm 0.38 mm
Occurrence. — The type specimens are from locality UWA 5985
in the McMonnigal Limestone. Also present at Jocality UWA 5984
and UWA 5975. These occurrences range in age from upper Lud-
lovian to Helderbergian.
Discussion. — The single spine is distinctive for this species.
Previously known species have two spines, one directed anteriorly
and one posteriorly near the valve margins. The spine of Rudderina
mutaspina is more nearly median though still near the ventral
border. This single spine appears to change direction with growth of
the individual (see Pl. 44, figs. 4-8), suggesting considerable varia-
tion within the species,
Rudderina spp. Pl. 44, figs. 9-12
Description. — Carapace subtriangular, greatest height anterior;
long, slender, curving spine directed posterolaterally from median
area at point of greatest width; some specimens also with small,
curving, dorsal spines at ends of straight hinge line.
Occurrence. — Specimens figured here are from locality UWA
6024; also at UWA 6026. Both occurrences in the Roberts Moun-
tains Formation.
Discussion. — Several different forms have been included here
which appear to belong under the genus Ruddeérina, because of the
general outline and spine development.
Superfamily THLIPSURACEA Ulrich, 1894
Family THLIPSURIDAE Ulrich, 1894
Genus THLIPSURA Jones and Holl, 1869
Type species. — Thlipsura corpulenta Jones and Holl, 1869.
Stturo-DEvoNIAN MicroFAuNA: McCLeLLANn 345
Thlipsura bispinosa McClellan, n. sp. Pl. 44, figs. 13-15
Description. — Carapace subovate; dorsal margin convex, ven-
tral straight, anterior and posterior strongly convex with maximum
convexity of posterior below midline; lateral surface strongly con-
vex, greatest width posterior, flat depressed area rises from posterior
margin to form shoulder which leads from posteroventral margin to
dorsal position at about one-third of distance to anterior of valve;
two anteriorly directed furrows lead from shoulder, with small
node representing maximum convexity point of valve and separating
the furrows; marginal to the furrows are two short spines projecting
from the shoulder, the ventral spine directed posteriorly and the
dorsal one posterodorsally directed,
Measurements. — Following are the dimensions of the figured
type specimens.
Pl. 44 Spec. Number Length Height
Fig. 13 34773 holotype 0.90 mm 0.52 mm
Fig, 14 34774 paratype 0.69 mm 0.41 mm
Fig. 15 34775 paratype 0.92 mm 0.56 mm
Occurrence. — Specimens 34773 and 34774 are from locality
UWA 5980 in the lower Wenban Limestone. Specimen 34775 is
from locality UWA 5973 in the Rabbit Hill Formation,
Discussion. — The most closely related species to Thlipsura
bispinosa is T. furca Roth (1929), which has a similar outline and
two furrows. The latter differs in lacking the spines and having a
better defined central node separating the furrows.
This species is not widely distributed in the area and is known
only from the Lower Devonian. Where it does occur, it is one of
the more common constituents of the fauna.
LOCALITY REGISTER
Numbers in (_) refer to location numbers given for the sections
on the index map, Fig. 1.
Coal Canyon (15) — Long. 116°28’, Lat. 40°2’30”, Horse Creek
Valley Quadrangle
UWA 5973 Located at top of hill on east side of Coal Canyon at
elev. 6905’, NE 1/4, SE 1/4, SE 1/4 Sec. 17, T25N,
UWA 5974
UWA 5975
UWA 5976
BULLETIN 274
R49E, north end of Simpson Park Range. Rabbit Hill
Fm.
Located west and 830’ down section from UWA 5973
at elev. 6440’. Roberts Mountains Fm.
Located south of UWA 5974, at elev. 6310’, on south
line 1300’ west of E corner Sec. 17. Roberts Mountains
Fm.
Located on slope on west side of Coal Canyon, at elev.
6440’, southeast of UWA 5975. Roberts Mountains
Fm.
Copenhagen Canyon (7) — Long. 116°24’, Lat. 39°12’30”, Horse
UWA 5977
UWA 5978
UWA 5979
Heaven Mountain Quadrangle
Located at top of cliff on west side of road in Copen-
hagen Canyon at elev. 7100’; NW 1/4, SW 1/4, SW
1/4 Sec. 36, TI6N, R49E. Roberts Mountains Fm.
Located near base of cliff, east and down section 430’
from UWA 5977, at elev. 6970’. Roberts Mountains
Fm.
Located on west side of road, 350’ south and 180’
down section from UWA 5978, at elev. 6865’. Roberts
Mountains Fm.
Cortez Canyon (16) — Long. 116°38’ Lat. 40°11’, Cortez Quad-
UWA 5980
UWA 5981
rangle
Located on north side near top of hill which forms east
side of Cortez Canyon, at elev. 6000’, by claim monu-
ment located 13,025’ NI9W of SW corner Sec. 6,
T26N, R48E. Wenban Limestone
Located north and 55’ down section from UWA 5980,
at elev. 5900’. Wenban Limestone
Gatechff (2) — Long. 116°47’30”, Lat. 39°00’, Wildcat Peak
UWA 5982
UWA 5983
Quadrangle
Located 1/2 mile west of Caesar Canyon, on north side
of Mill Canyon at elev. 7840’, 30’ above base of cliff
where canyon is crossed by stock fence. Bastille Lime-
stone
Located on slope above and 220’ up section from UWA
5982, at elev. 7920’. Bastille Limestone
StLtuRO-DEVONIAN MicrRoFAUNA: McCLELLAN 347
Ikes Canyon (5) — Long. 116°44’, Lat. 39°03’, Dianas Punch Bowl
UWA 5984
UWA 5985
UWA 5986
UWA 5987
UWA 5988
Quadrangle
Located on west side of Copper Mt., overlooking Ikes
Cabin, on north side of canyon, at elev. 8360’ and
6700’ S87W from NW corner Sec. 17, T14N, R46E, in
the Toquima Mountains. Masket Shale
Located north and 10’ up section from UWA 5984, in
small saddle at elev, 8320’. Base of McMonnigal Lime-
stone
Located northwest and 45’ up section from UWA
5985, on north side of saddle at elev. 8290’. McMon-
nigal Limestone
Located southwest of UWA 5986 on nose of hill 250’
above bottom of canyon, at elev. 8010’, 1/4 mile east
of Ikes Cabin. Masket Shale
Located south and 390’ down section from UWA 5987,
at elev. 7790’. Masket Shale
Kelley Creek (13) — Long, 116°20’30”, Lat. 39°56’, Roberts Creek
UWA 5989
UWA 5990
Mountain Quadrangle
Located 6600’ due south of BM 6109 which is in NW
1/4 Sec. 21, T24N, R50F, on slope of ridge on east
side of Kelley Creek at elev. 6880’, near mouth of
canyon on north flank of Roberts Meuntains. Roberts
Mountains Fm.
Located northwest and down section 135’ from UWA
5989, at elev. 6850’, Roberts Mountains Fm.
Lone Mountain (10) — Long. 116°16’, Lat. 39°35’, Bartine Ranch
UWA 5991
UWA 5992
Quadrangle
Located 12,000’ north and 6200’ east of the NE cor-
ner of Sec. 1, T19N, R50E, on southwest side and
near top of Lone Mountain, at elev. 7200’. Lone
Mountain Dolomite
Located due south and 830’ down section from UWA
5991, at elev. 7040’. Lone Mountain Dolomite
March Spring (4) — Long. 116°44’, Lat. 39°02’45”, Dianas Punch
UWA 5993
Bowl Quadrangle
Located 7550’ west of NW corner, Sec. 20, T14N,
R47E on top of hill west of small spring and 4% mile
UWA 5994
UWA 5995
UWA 5996
UWA 5997
BULLETIN 274
south of Ikes Canyon, at elev. 8200’. McMonnigal
Limestone
Located east and 70’ down section from UWA 5993,
on hillside at elev. 8160’. McMonnigal Limestone
Located east and 130’ down section from UWA 5994,
at elev. 8090’. Masket Shale
Located east and 100’ down section from UWA 5995,
at elev. 8050’. Masket Shale
Located east and 130’ down section from UWA 5996,
at elev. 7980’. Masket Shale
Mill Canyon (3) — Long. 116°48’, Lat. 39°01’, east side of To-
UWA 5998
quima Mts., Wildcat Peak Quadrangle
Located at base of ridge on the west side of the north
fork of Mill Canyon, 3600’ north of junction with the
main canyon, at elev. 8280’. Gatecliff Dolomite
Northumberland Canyon (1) — Long. 116°50’, Lat. 38°54’, 7 miles
UWA 5999
south of Mill Canyon in Toquima Mts., unsurveyed
Located at base of hill on north side of Northumber-
and Canyon, 100 yards west of the mouth of Perkins
Canyon and approximately one mile west of national
forest boundary, at elev. 7800’. Perkins Canyon Fm.
(U. Ord? )
Pete Hanson Creek (12) — Long. 116°20’, Lat. 39°53’, Roberts
UWA 6000
UWA 6001
UWA 6002
UWA 6003
UWA 6004
Creek Mountain Quadrangle
Located in high saddle on ridge separating two forks
of Pete Hanson Creek, 2300’ north and 2800’ west
of Roberts Creek Mountain, at elev. 9150’. McColley
Canyon Fm.
Located due west on side of hill and 1850’ down sec-
tion from UWA 6000, at elev. 8680’. Lone Mountain
Dolomite
Located west and 405’ down section from UWA
6001, at elev. 8570’ in low saddle. Roberts Mountains
Fm.
Located west and down section 125’ from UWA 6002,
in low saddle, at elev. 8570’. Roberts Mountains Fm.
Located west and 170’ down section from UWA 6003,
at elev. 8600’ on small nob at end of saddle. Roberts
Mountains Fm.
Stturo-DEVONIAN MicroFAuNA: McCLELLAN 349
UWA 6006
UWA 6007
UWA 6008
Rabbit Hill
UWA 6009
UWA 6010
UWA 6011
UWA 6012
UWA 6013
Located west and 170’ down section from UWA 6004,
at elev. 8560’. Roberts Mountains Fm.
Located west and 310’ down section from UWA 6006,
at elev. 8500’. Roberts Mountains Fm.
Located west and 740’ down section from UWA 6007,
at elev. 8330’. Roberts Mountains Fm.
(6) —Long. 116°25’, Lat. 39°12’, Horse Heaven
Mountain Quadrangle
Located in middle of flat top of Rabbit Hill %4 mile
west of road on north side of Whiterock Canyon, at
elev. 7200’, Rabbit Hill Fm.
Located SSE and 400’ down section from UWA 6009,
on edge of steep slope at elev. 7120’. Rabbit Hill Fm.
Located east and 55’ down section from UWA 6010,
at elev. 7070’. Roberts Mountains Fm.
Located east and 25’ down section from UWA 6011,
at elev. 7050’. Roberts Mountains Fm.
Located east and 25’ down section from UWA 6012,
base of the hill at elev. 7030’. Roberts Mountains Fm.
Telegraph Canyon (21) — Long. 116°04’, Lat. 40°06’, Mineral Hill
UWA 6014
UWA 6015
Quadrangle
Located on low ridge 3900’ west of BM 6912 at top
of pass above Telegraph Canyon; SW 1/4, SW 1/4,
NE 1/4, Sec. 36, T26N, R52E, at elev. 6850’, Rob-
erts Mountains Fm.
Located east and 2080’ up section from UWA 6014,
on nose of hill at elev. 7040’. Roberts Mountains Fm.
Twin Springs Hills (9) — Long. 116°24’45”, Lat. 39°30’, Antelope
UWA 6016
UWA 6017
Tyrone Gap
Peak Quadrangle
Located on low rise on east side of Twin Springs Hills
in NW 1/4, NE 1/4, NW 1/4 Sec. 25, T19N, R49E,
at elev. 6360’. Roberts Mountains Fm.
Located southwest and 260’ up section from UWA
6016, at elev. 6380’. Roberts Mountains Fm.
(19) — Long. 116°04’30”, Lat. 39°49’, Garden Valley
Quadrangle
350
UWA 6034
BULLETIN 274
Located 250’ east of center of WL, Sec. 1, T22N,
R52E at south end of hogback, at elev. 6000’. Lone
Mountain Dolomite
Willow Creek (14) —Long. 116°19’, Lat. 39°57’, Roberts Creek
UWA 6018
UWA 6019
UWA 6020
UWA 6021
UWA 6022
UWA 6023
UWA 6024
UWA 6025
UWA 6026
UWA 6027
UWA 6028
UWA 6029
UWA 6030
Mountain Quadrangle
Located 1150’ east and 3100’ south of SE corner
Sec. 15, T24N, R50E, on point of ridge on east side
of Willow Creek, at elev. 7100’. McColley Canyon
Fm.
Located northwest and 320’ down section from UWA
6018, at elev. 6950’. Lone Mountain Dolomite
Located west and 1505’ down section from UWA
6019, at center EL Sec. 22, T24N, RSOE, at elev.
6590’. Roberts Mountains Fm.
Located NNW and 110’ down section from UWA
6020 at elev. 6550’ Roberts Mountains Fm.
Located west and 435’ down section from UWA 6021,
at elev. 6320’, Roberts Mountains Fm.
Located south and 70’ down section from UWA 6022,
at elev. 6310’. Roberts Mountains Fm.
Located west and 50’ down section from UWA 6023,
beside trail behind ranch house, at elev. 6280’. Rob-
erts Mountains Fm.
Located west and 230’ down section from UWA 6024,
on hill on west side of Willow Creek, at elev. 6500’.
Roberts Mountains Fm.
Located southwest and 310’ down section from UWA
6025, at elev. 6540’. Roberts Mountains Fm.
Located west and 355’ down section from UWA 6026,
at elev. 6500’. Roberts Mountains Fm.
Located southwest and 130’ down section from UWA
6027, on north flank of mountains at elev. 6610’.
Roberts Mountains Fm.
Located west and 330’ down section from UWA 6028,
at elev. 6580’. Roberts Mountains Fm.
Located north and 155’ down section from UWA
6029, on small northward projecting nose of hill,
S1LuRO-DEVONIAN MicroFAUNA: MCCLELLAN 351
midway between Willow and Birch creeks, at elev.
6500’. Roberts Mountains Fm.
UWA 6031 Located NNW and 105’ down section from UWA
6030, at elev, 6455’ approximately 50’ above base of
hill. Roberts Mountains Fm.
Wood Cone (8) — Long. 116°09’, Lat. 39°23’, Bellevue Peak Quad-
rangle
UWA 6032 Located 2700’ west and 1100’ north of BM 7201 by
road north of Wood Cone Peak, on slope of small nob
of hill, at elev. 7160’. Roberts Mountains Fm.
UWA 6033 Located southwest and 750’ down section from UWA
6032, at elev. 7080’. Roberts Mountains Fm.
The following sections were studied, but no microfossils were
recovered from the samples:
Bailey Pass (20) — Long. 116°06’, Lat. 39°55’, Garden Valley
Quadrangle, from top of pass down east side of Sul-
phur Spring Range
Cole Creek (23) — Long. 116°07’, Lat. 40°37’, Carlin Quadrangle,
» 172 See 34, F32N; R52E
Fourmile Canyon (18) — Long. 116°33’, Lat. 40°12’, Cortez Quad-
rangle, at mouth of canyon
Mill Canyon (17) — Long. 116°35’, Lat. 40°12’, Cortez Quadrangle,
at boundary of Lander and Eureka counties
Mitchell Creek (22) — Long. 115°35’, Lat. 40°09’, Sherman Moun-
tain Quadrangle, west side of southern Ruby Moun-
tains
Sheep Corral Canyon (11) —Long. 116°34’, Lat. 39°56’, Walti
Hot Springs Quadrangle, SW corner Sec. 14, T23N,
R48E
REFERENCES CITED
Barker, R. W.
1960. Taxonomic notes on species figured by H. B. Brady in Foramini-
fera of Challenger Expedition. Soc. Econ. Paleont. and Min., Spec.
Publ. 9, 238 pp.. 115 pls.
Barker-Webb, P. and Berthelot, S.
1839. Foraminiferes, in Hist. Nat. Iles Canaries. Vol. 2, pt. 2, Zool., pp.
119-146, pls. 1-3.
gy BULLETIN 274
Bassler, R. S.
1941. Ostracoda from the Devonian (Onondaga) chert of west Ten-
nessee. Washington Acad. Sci. Jour., vol. 31, No. 1, pp. 21-27, figs.
1-37.
Brady, H. B.
1878. On reticularian and radiolarian Rhizopoda (Foraminifera and
Polycystina) of North Polar Expedition of 1875-76. Ann. Mag.
Nat. Hist., ser. 5, vol. 1, pp. 425-440, pls. 20, 21.
1879. Notes on some reticularian Rhizopoda of the Challenger Expedition.
Quart. Jour. Micr. Sci., n. s., vol. 19, pp. 28-60, pl. 5
Browne, R. B., and Schott, V. J.
1963. Arenaceous Foraminifera from the Osgood Formation at Osgood,
Indiana, Bull. Amer. Paleont., vol. 46, No. 209, pp. 191-242, pls.
48-52.
Carlisle, D., Murphy, M. A., Nelson, C. A., and Winterer, E. L.
1957. Devonian stratigraphy of Sulphur Springs and Pinyon ranges,
Nevada. Amer. Assoc. Petr. Geol., Bull., vol. 41, No. 10, pp. 2175-
2191, 3 text-figs.
Carls, P., and Gandl, J.
1969. Stratigraphie und Conodonten des Unter-Devons der ostlichen
Iberischen Ketten. Neues. Jahrb. Geol. Paleont., Bd. 132, No. 2, pp.
155-218, 3 text-figs.
Chernysheva N. E.
1940. K. stratigrafii nizhnego Karbona Makarovskogo rayona yuzhnogo
Urala po faune foraminifer. Soc. Nat. Moscow, Bull., n. s., tome
, 48 (sect. geol. tome 18), No. 5-6, p. 127, pls. 1, 2.
Clark, D. L., and Ethington, R. L.
1966. Gomadants and biostratigraphy of the Lower and Middle Devonian
of Nevada and Utah. Jour. Paleont., vol. 40, No. 3, pp. 659-689,
pls. 82-84, 10 text-figs.
Conkin, J. E.
1954. Hyperammina kentuckyensis, n. sp. from the Mississippian of
Kentucky. Cushman Found. Foram. Res., Contr., vol. 5, pt. 4, No.
119, pp. 165-169, pl. 31.
1961. Mississippian smaller Foraminifera of Kentucky, southern In-
diana, northern Tennessee, and southcentral Ohio. Bull. Amer.
Paleont., vol. 43, No. 196, pp. 131-368, pls. 17-27, figs. 1-43.
, and Conkin, B. M.
1970. Middle Devonian arenaceous Foraminifera of central Ohio. Micro-
paleont., vol. 16, No. 1, pp. 1-14, pls. 1-4.
Copeland, M. L.
1962. Canadian fossil Ostracoda, Conchostraca, Eurypterida, and
Phyllocarida. Geol. Sur. Canada, Bull. 91, pp. 18-51, pls. 5-10.
Coryell, H. N.
1928. ie new Pennsylvanian Ostracoda. Jour. Paleont., vol. 2, No. 4,
. 377-381, pls. 51.
Coryell, H. N. and Cuskley, V. A.
1934. Some new Ostracodes from “White Mound” section of Haragan
Shale, Murray Co., Oklahoma. Amer. Mus. Nat. Hist., Novitates,
No. 748, 12 pp., 18 figs.
Coryell, H. N., and Malkin, D. S.
1936. Some Hamilton Ostracodes from Arkona, Ontario. Amer. Mus.
Nat. Hist., Novitates, No. 891, 20 pp., 38 figs.
Croneis, C., Dunn, P. H., and Hunter, D.
1932. Pre-Carboniferous Foraminifera. Science, vol. 75, No. 1935, p. 138.
Stturo-DeEvoNIAN MicrorAunA: McCLe.LLan 353
Cushman, J. A.
1910. Foraminifera of the North Pacific Ocean. U.S. Nat. Mus., Bull.
71, pt. 1, pp. 1-134.
1918. Foraminifera of the Atlantic Ocean. U.S. Nat. Mus., Bull. 104, pt.
IS pp l-1ils spt) 2, ppel-11i-
1959. Foraminifera. Ed. 4, Cambridge, Mass., 605 pp.
Dapples, E. C.
1959. Behavior of silica in diagenesis. in Silica in Sediments. Soc. Econ.
Paleont. Min., Spec. Publ. No. 7, H. A. Ireland ed., pp. 36-54,
figs. 1-0.
1967. Silica as an agent in diagenesis. in Diagenesis in Sediments. Devel.
in Sed. 8, G. Larsen and G. V. Chilingar ed., Amsterdam, pp.
323-342, 1 text-fig.
de Folin, L.
1887. Les Rhizopodes reticulaires. Naturtiste, Paris, vol. 9, No. 2, p. 127,
text-fig. 12a.
Dunn, P. H.
1942. Silurian Foraminifera of the Mississippian Basin. Jour. Paleont.,
vol. 16, No. 3, pp. 317-342, pls. 42-44.
Ellis, B. F., and Messina, A. R.
1952 + suppl. Catalogue of Ostracoda. Amer. Mus. Nat. Hist., Spec. Publ.
Gilluly, J., and Masursky, H.
1965. Geology of the Cortez Quadrangle, Nevada. U.S. Geol. Sur., Bull.
1175, pp. 1-97, pls. 1, 2, figs. 1-10.
Glaessner, M. F.
1937. Studien iiber Foraminiferen aus der Kreide und dem Tertidr des
Kaukasus; I-Die Foraminiferen der dltesten Tertiarschichten des
Nordwestkaukasus. Moscow Univ., Lab. Paleont., Prob. Paleont.,
vols. 2, 3, p. 354.
Grubbs, D. M.
1939. Fauna of the Niagaran nodules of the Chicago area. Jour. Paleont.,
vol. 13, No. 6, pp. 543-560, pl. 61, 62.
Gutschick, R. C.
1959. Arenaceous Foraminifera from the Rockford Limestone of northern
Indiana, Jour. Paleont., vol. 33, No. 2, pp. 229-250, pls. 33-37, 3
text-figs.
_ Weiner, J. L., and Young, L.
1961. Lower Mississippian arenaceous Foraminifera from Oklahoma,
Texas, and Montana. Jour. Paleont., vol. 35, No. 6, pp. 1193-1221,
pls. 147-150, 5 text-figs.
Hague, A.
1892. Geology of the Eureka district, Nevada. U.S. Geol. Sur., Mono-
graphs, vol. 20, pp. 34-217.
Hattin, D. E.
1960. Waldron Foraminifera in Indiana. Geol. Soc. Amer., Bull., vol.
71, No. 12, pt. 2, abstr., p. 2016.
Henbest, L. G.
1935. Nanicella, a new genus of Devonian Foraminifera. Washington
Acad. Sci., Jour., vol. 25, p. 34.
Ireland, H. A.
1939. Devonian and Silurian Foraminifera from Oklahoma. Jour.
Paleont., vol. 13, No. 2, pp. 190-202, 75 text-figs.
354 BULLETIN 274
Johnson, J. G.
1962. Lower Devonian—Middle Devonian boundary in central Nevada.
Amer. Assoc. Petr. Geol., vol. 46, pp. 542-546, 3 text-figs.
1965. Lower Devonian stratigraphy and correlation, northern Simpson
Park Range, Nevada. Bull. Canadian Petr. Geol., vol. 13, No. 3,
pp. 365-381.
, and Boucot, A. J.
1968. Llandovery to Givetian brachiopod zonal sequence in the Silurian
and Devonian otf central Nevada. Geol. Soc. Amer., Cordilleran
Sect., Program, Ann. Mtg., abstr., p. 69.
Jones, T. R., and Parker, W. K.
1860. On the rhizopodal fauna of the Mediterranean, compared with
that of Italian and some other Tertiary deposits. Quart. Jour.
Geol. Soc. London, vol. 16, pp. 292-307.
Kay, M.
1940. Ordovician Mohawkian Ostracoda; Lower Trenton Decorah
fauna. Jour. Paleont., vol. 14, No. 4, pp. 234-269, pls. 29-34.
1960. Paleozoic continental margin in central Nevada, western United
States. Internat. Geol. Congr. Rept., 21st sess., Copenhagen, Den-
mark, pt. 12, pp. 93-103.
, and Crawford, J. P.
1964. Paleozoic facies from miogcosynclinal belt in thrust slices, central
Nevada. Geol. Soc. Amer., Bull., vol. 75, No. 5, pp. 425-454, pls.
1-6, 9 text-figs.
Kesling, R. V.
1953. Ostracods of the family Aechminidae from the Arkona Shale of
southern Ontario. Michigan Univ., Mus. Paleont. Contr., vol. 11,
No: ey pp: 1-105 plas
Klapper, G.
1969. Conodont succession in central Nevada. Geol. Soc. Amer., Cordil-
leran Sect., Program, Ann. Megt., abstr., p. 72.
Krauskopf, K. B.
1959. Geochemistry of silica in sedimentary environments, in Silica in
Sediments. Soc. Econ. Paleont. Min., Spec. Publ. No. 7, H.
A. Ireland ed., pp. 4-19, figs. 1-6.
Loeblich, A. R., and Tappan, H.
1957. Eleven new genera of Foraminifera. U.S. Nat. Mus., Bull. 215,
pp. 223-232, pl. 72.
200 ———————
1964. Treatise on Invertebrate Paleontology. R. C. Moore ed., Pt. C
Protista 2, vols. 1, 2, 900 pp.
Martinsson, A.
1962. Ostracodes of the family Beyrichiidae from the Silurian of Got-
land. Uppsala Univ., Paleont. Inst., Publ. No. 41, 369 pp., 203
text-figs.
McClellan, W. A.
1966. Arenaceous Foraminifera from the Waldron Shale (Niagaran)
of southeast Indiana. Bull. Amer. Paleont., vol. 50, No. 230, pp.
447-518, pls. 36-42.
Merriam, C. W.
1940. Devonian stratigraphy and paleontology of the Roberts Mountains
region, Nevada. Geol. Soc. Amer., Spec. Paper 25, pp. 1-114.
1963. Paleozoic rocks of Antelope Valley, Eureka and Nye counties,
Nevada. U.S. Geol. Sur., Prof. Paper 423, pp. 1-55, pl. 1, 2, 8 text-
figs.
On
SILURO-DEVONIAN MIcRoFAUNA: McCLELLAN 35
, and Anderson, C. A.
1942. Reconnaissance survey of Roberts Mountains, Nevada. Geol. Soc.
Amer., Bull., vol. 53, No. 12, pt. 1, pp. 1675-1728, pls. 1-4, figs. 1, 2.
Moore, R. C., ed.
1961. Treatise on Invertebrate Paleontology, Pt. Q Arthropoda 3, 442 pp.
1962. Treatise on Invertebrate Paleontology Pt. W Miscellanea, 259 pp.
Moreman, W. L.
1930. Arenaceous Foraminifera from Ordovician and Silurian lime-
stones of Oklahoma. Jour. Paleont., vol. 4, No. 1, pp. 42-59, pls. 5-7.
1933. Arenaceous Foraminifera from the lower Paleozoic rocks of
Oklahoma. Jour. Paleont., vol. 7, No. 4, pp. 393-397, pl. 47.
Mound, M. C.
1961. Arenaceous Foraminifera from the Brassfield Limestone of south-
eastern Indiana. Indiana Dept. Conserv., Geol. Sur. Bull. No. 23,
38 pp., 3 pls.
Murphy, M. A.
1968. Stratigraphy of the Roberts Mountains Formation in the type
region, Geol. Soc. Amer., Cordilleran Sect., Program, Ann. Mtg.,
abstr., p. 86.
Nolan, T. B., Merriam, C. W., and Williams, J. S.
1956. Stratigraphic section in the vicinity of Eureka, Nevada. U.S. Geol.
Sur., Prof. Paper 276, pp. 28-52, pl. 2.
Norman, A. M.
1878. On the genus Haliphysema. Ann. Mag. Nat. Hist., ser. 5, vol. 1,
pa 28ihepls 6s digs
Rhumbler, L.
1895. Entwurf eines natiirlichen systems der Thalamophoren. Gesell.
Wiss. Gottingen, math-physik. Kongl., Nachr., No. 1, pp. 51-98.
1904. Systematische zusammenstellung der recenten Reticulosa. Archiv.
Protistenkunde, vol. 3, pp. 181-294, text-figs. 1-142.
Roberts, R. J., Hotz, P. E., Gilluly, J., and Ferguson, H. G.
1958. Paleozoic rocks of north-central Nevada. Amer. Assoc. Petr.
Geol., Bull., vol. 42, No. 12, pp. 2813-2858, 11 text-figs.
Roth, R.
1929. Some Ostracodes from the Haragan Marl, Devonian of Oklahoma.
Jour. Paleont., vol. 3, No 4, pp. 327-372, pls. 35-38.
Rzehak, A.
1885. Bemerkungen iiber einige Foraminiferen der Oligocaén Formation.
Naturforsch. vereins Briinn, Verhandl., vol. 23 (1884), pp. 123-
129"
Sars, M.
1869. In Carpenter, W. B., On the rhizopodal fauna of the deep sea. Roy.
Soc. London, Proc., vol. 18, No. 114, pp. 59-62.
1872. In Sars, G. O., Underspgelser over Hardangerfjordens fauna.
Vidensk.-Selsk. Christiania, Forhandl., vol. 1871, pp. 246-255.
Schulze, F. E.
1875. Zoologische ergebnisse der Nord-seefahrt, 1872, I. Rhizopoden. II:
Komm. Untersuch. deutsch. Meere in Kiel, Jahresber., vol. 1872-
73, pp. 99-114, pl. 2.
Sharp, R. P.
1942. Stratigraphy and structure of the southern Ruby Mountains,
Nevada. Geol. Soc. Amer., Bull., vol. 53, No. 5, pp. 647-690.
356 BULLETIN 274
Shchedrina, Z. G.
1939. Nowvyg rod peschanistykh Foraminifera iz Arkticheskikh Morey.
Akad. Nauk SSSR, Doklady, n. s., vol. 24, No. 1, p. 94-96.
Smith, D. L.
1966. Tippecanoe sequence in western North America. Unpubl. Ph.D.
thesis, Univ. Washington, Seattle, 83 pp., 6 figs.
Stewart, G. A.
1936. Ostracodes of the Silica Shale, Middle Devonian of Ohio. Jour.
Paleont., vol. 10, No. 8, pp. 739-763, pls. 100-102.
, and Priddy, R. R.
1941. Arenaceous Foraminifera from Niagaran rocks of Ohio and In-
diana. Jour. Paleont., vol. 15, No. 4, pp. 366-375, pl. 54.
, and Lampe, L.
1947. Foraminifera from the Middle Devonian Bone Beds of Ohio. Jour.
Paleont., vol. 21, No. 6, pp. 529-536, pls. 78, 79.
Stover, L. E.
1956. Ostracoda from the Windom Shale (Hamilton) of western New
York. Jour. Paleont., vol. 30, No. 5, pp. 1092-1142, pls. 111-119, 8
text-figs.
Summerson, C. H.
1958. Arenaceous Foraminifera from the Middle Devonian limestones
of Ohio. Jour. Paleont., vol. 32, No. 3, pp. 544-558, pls. 81, 82, 7
text-figs.
Swartz, F. M.
1936. Revision of the Primitiidae and Beyrichiidae, with new Ostracoda
from the Lower Devonian of Pennsylvania. Jour. Paleont., vol. 10,
No. 7, pp. 541-586, pls. 78-89.
, and Oriel, S. S.
1948. Ostracoda from Middle Devonian Windom beds in western New
York. Jour. Paleont., vol. 22, No. 5, pp. 541-566, pls. 79-81, 4 text-
figs.
, and Whitmore, F. C.
1956. Ostracoda of the Silurian Decker and Manlius limestones in New
Jersey and eastern New York. Jour. Paleont., vol. 30, No. 5, pp.
1029-1091, pls. 103-110, 4 text-figs.
Thomas, A. O.
1931. Late Devonian Foraminifera from Iowa. Jour. Paleont., vol. 5,
No. 1, pp. 40, 41, pl. 7.
Ulrich, E. O.
1891. Beecherella, a new genus of lower Helderberg Ostracoda. Amer.
Geol., vol. 8, No. 4, pp. 197-204, pl. 2.
Weller, S.
1903. The Paleozoic faunas. New Jersey Geol. Sur., Paleont., vol. 3, 462
pp., 53 pls.
Wheeler, H. E.
1963. Post-Sauk and Pre-Absaroka Paleozoic stratigraphic patterns in
North America, Amer. Assoc. Petr. Geol., Bull., vol. 47, No. 8,
pp. 1497-1526.
White, M. P.
1928. Some index Foraminifera of Tampico embayment area of Mexico.
Jour. Paleont., vol. 2, No. 3, pp. 177-215, pls. 27-29.
Winterer, E. L., and Murphy, M. A
1960. Silurian reef complex and associated facies, central Nevada. Jour.
Geol., vol. 68, No. 2, pp. 117-139.
PLATES
The University of Nevada provided the funds for the illustrations of this paper.
358
BULLETIN 274
EXPLANATION OF PLATE 33
All figures «60
Page
Rhabdammina major de Folin. ...0...0.......00.00cccccoceccceecceceee 277
#34686 (X30)
Hyperammina: casteri-Conkan 6... 5.:<...0.5<-scec0cesececcste couse eee 279
2. Microspheric form, #34670. 6. Megalospheric form,
# 34669.
Marsipeilan'Sps foe cea seen a ee ee 279
# 34674.
Hyperammina constricta Gutschick and Treckman ......... 280
Megalospheric form, #34671.
Hyperammina curva (Moreman) .....0000.....0oocccccocccccceeeecceeee 281
Megalospheric form, #34672.
Marsipella torta Stewart and Priddy ..................... eS
# 34673.
Rhabdammina bifurcata Browne and Schott .................. 276
8. £34682. 9. #34683.
Rhabdammina cylindrica Glaessner 0... 276
10. #34684. 11. #34685.
Raibosammina aspera Moreman .......... figs ie fla ae, eed ee 286
# 34681.
BuLL. AMER. PALEONT., VOL. 62 PLATE 33
BULL. AMER. PALEONT., VOL. 62 PLATE 34
Figure
1-5.
6-10.
11, 12.
13, 14.
15, 16.
17.
18.
19.
20.
21.
22.
23, 27.
25.
S1tuRo-Devontan MicroFauna: McCLetian 359
EXPLANATION OF PLATE 34
All figures «60
Page
Anictosphaera progressa McClellan, n. sp. . 284
1. Paratype, showing interior, #34651. 2: Paratype, #34652.
3. Paratype, #34653. 4. Holotype, # 34654. 5. Paratype,
# 34655.
Storthosphaera malloryi McClellan, n. sp. 296
6. Holotype, #34703. 7. Paratype, #34704. 8. Paratype,
# 34705. 9. Paratype, #34706. 10. Paratype, #34707.
Sorosphaera inflata McClellan, n. sp. 0... . 290
11. Paratype, #34689. 12. Holotype, #34690. Showing wall
dividing two chambers.
Sorosphaera compacta McClellan, n. sp. ......000000.. 290
13. Holotype, #34691, broken wall shows divisions between
three chambers. 14a, b. Paratype, #34692, shows slight
constriction of test and walls dividing three chambers on
broken side.
Sorosphaera osgoodensis Stewart and Priddy .................. 288
15. T'wo-chambered form with part of third, #34790. 16.
Four-chambered specimen, #34688.
Stegnammina moremani (Dunn) ............c ei ccecceeeeeeeeeeee, 294
# 34699
Stegnammina cylindrica Moreman. .......0.............cccccceeeeeeeeeee. 292
# 34697
Stegnammina hebesta? Moreman |..........000...0ccccccecceeeeeeeee 293
# 34692
Sorosphaeraws ptm se cc ssoe estes caste ete Soe ae 291
# 34693 :
Psammosphaera cava MoOreman ooo.........ccccccccccccccccccccceeceeceeeeees 282
# 34679
Stegnammina contorta McClellan .oooo.2.0o.o oo oococccceeeeeeeeee 292
With damaged test wall, #34696.
Stegnammina quadrangularis (Dunn) ......0000.0...00...... ee. 295
23. Top view, #34700. 27. Top view of broken specimen,
# 34701.
Sorosphaera’ confusa/ brady =. eee. 287
# 34687
Ceratammina cf. C. cornucopia Ireland ................................ 285
With partially broken wall, #34660.
Psammosphaera laevigata White 2000s. 283
# 34680
Figure
12.
13.
14.
16.
fie
18.
19)
20.
BuLLeETIN 274
EXPLANATION OF PLATE 35
All figures 60
Hemisphaerammina brady Loeblich and Tappan .............. 302
Side view, #34663.
Hemisphaerammina cf. H. bradyi Loeblich and Tappan .. 302
Side view, #34664.
Hemisphaerammina discoidea? (Summerson) ................... 303
Side view, #34666.
Hemisphaerammina coronata (Ireland) ...................... oe 303
4a. Top view, 4b. Side view. #34665.
Hemisphaerammina bipatella McClellan, n. sp. ................. 304
5. Paratype, top view, #34667. 6a. Holotype, top view, 6b,
end view, #34668.
. Sorostomasphaera waldronensis McClellan ...............0........ 297
7. £34694. 8. With part of second chamber remaining,
# 34695.
Webbinelloidea hemispherica Stewart and Lampe ............ 312
9. Top view, #34715. 10. Side view, #34714.
Webbinelloidea aff. W. nodosa Summerson ........................ 312
lla. Side view, #34717. 11b. Top view. 15. Top view,
# 34716.
Webbinelloidea hattini McClellan 00.0000... 311
Side view, #34713.
Metamorphina tholus (Moreman) .............. Sees ee eae 308
13a. Top view. 13b. Ventral view; #34676
Webbinelloidea ventriquetra McClellan ..............000000000000.... 313
Ventral view, #34718.
Metamorphina gibbosa (Ireland) .......000000.......cecceccceccccccee 309
16a. Top view. 16b. Ventral view. 16c. Side view; #34675.
Thurammina trituba Dunn 0... .c..ce:6.-..scssusadecssteactesssspsseoes ecco 300
#34710
Thurammina arcuata Moreman. ..........................0..000.cceee 299
# 34708.
Thurammina tubulata Moreman. ....0.00......0.....00000.ccccceeeceeeee 301
# 34711.
Thurammina quadritubulata? Dunn |... 300
# 34709.
BuLL. AMER. PALEONT., VOL. 62 PLATE 35
BULL. AMER. PALEONT., VOL. 62 PLATE 36
Figure
9, 10.
Stturo-Devonian MicroFAuNA: McCLELLAN 361
EXPLANATION OF PLATE 36
All figures 60
Page
Atelikamara incomposita McClellan, n. sp. .....00000000.......... 306
la. Holotype, top view, #34658. 1b. Ventral view. 2. Para-
type, top view, #34793. 3. Paratype, top view, #34657.
Tolypammina tortuosa Dunn .......... eA ee ee ee . 815
#34712
Glomospira siluriama Ireland ......000.0........0cccccccccccececcceeeeeteeeeee 314
+ 34661
Atelikamara incomposita McClellan, n. sp... 306
# 34659. Top view with attached, #34662.
Stomasphaera brassfieldensis Mound ..........0..0.0....00.00000000005. 298
# 34702
Aschemonella bastillensis McClellan, n. sp. ......00..0000.0..00.... 316
Nanicella dainae? Chernysheva. .................0.0...c:cccccceeceeee . 319
9. #34678. 10. #34677.
362 BULLETIN 274
EXPLANATION OF PLATE 37
Figure Page
1. Haplophragmoides antiquus McClellan, n. sp. ............. 23 318
Scanning electron micrograph 300, #34792. Needle-like
fragment on surface of specimen is foreign material.
ion Scanning electron micrograph 200, #34792. Main aper-
ture at point A, aligned grains indicate suture at S.
BULL. AMER. PALEONT., VOL. 62 PLATE 37
PLATE 38
BULL. AMER. PALEONT., VOL. 62
SttuRo-DeEvonian MicroFAuNA: McCLELLAN 363
EXPLANATION OF PLATE 38
Figure Page
1. Webbinelloidea hemispherica Stewart and Lampe .......... 312
# 34715, top view 160.
2. Webbinelloidea aff. W. nodosa Summerson ..... .......... eer 312
# 34717, top view 120.
3. Hemisphaerammina bipatella McClellan, n. sp. .................. 304
3. Holotype, end view 160; #34668. 3b. Top view 160.
4. Hemisphaerammina coronata (Ireland) ..........0.00..0........... O05:
Top view showing ordered arrangement of grains, 320;
#34665.
364 BULLETIN 274
EXPLANATION OF PLATE 39
Figure Page
1. Psammosphaera laevigata White 2.000.000.0000 283
# 34680, «240.
2. Stegnammina moremani (Dunn) ......... ...................c.ccc0eeeeeeees 294
Partially broken specimen 240; #34699.
3. Raibosammina aspera Moreman ..........0.0.........:cccccccccceeeeeeeee 286
# 34681, X80.
4. Rhabdammina bifurcata Browne and Schott .........000000...... 276
Showing interior and wall structure, X120; #34683.
5. Metamorphina tholus (Moreman) .....................00..ccccseeceeeeeeecees 308
Ventral view and interior, X160; #34676.
6. Sorosphaera osgoodensis Stewart and Priddy .................... . 288
# 34688, 120.
PLATE 39
nN
©
4
°
>
a
Z
fe)
a
4
<
a
oe
a
=
<
=
|
5)
Q
N
ve)
=|
S
>
a
a
2)
Q
4
<
Ay
8
fa
5
<
S
=)
a)
Figure
Situro-DeEvoniaAn MicroFauna: McCLeLLAn 365
EXPLANATION OF PLATE 40
Page
Sorosphaera inflata McClellan, n. sp. o.......0..0cccccceceeeeeeeeeeees 290
Holotype shows wall dividing two chambers of partially
broken test, X160; #34690.
Sorosphaera compacta McClellan, n. sp. oo......cccccccccece 290
Holotype, broken wall shows divisions between three cham-
bers and lack of definite sutures, X160; #34691.
Storthosphaera malloryi McClellan, n. sp. .........0...000.00000.... 296
Holotype, 160; #34703.
Rhabdammina cylindrica Glaessner ......0....0.0..00cccccccccccseeeeeeee. 276
X80; #34685.
Atelikamara incomposita McClellan, n. sp. ..................0000008.. 306
5a. Holotype, top view, X80; 334658. 5b. Ventral view,
X80; #34658.
366
eT:
12.
13.
14.
16.
We
LG SAS:
BULLETIN 274
EXPLANATION OF PLATE 41
All figures «45
Page
Bolbiprimitia teresaccula Swartz and Whitmore ................ 320
1. #34731 RV. 2. #34732 RV. 3. #34733 LV.
Phlyctiscaphia:Sp:, 223..A0¢42.. ene eee eee eee 323
# 34758, RV.
Phiyctiscapha keslingi Copeland ..............0.0...0..0ccccececeee 322
5. Male, #34756 LV. 6. Female, #34757 RV.
Kloedenia aparchoides Swartz and Whitmore .................... 321
H# 34744.
Welleriopsis jerseyensis (Weller) ......0.00000...ccccccccceeceeeeeeeeeeees 324
8. #34786 RV. 9. #34787 RV.
Velibeyrichia? “Sp. 25:2 2iw....c.0.c. eee 324
# 34785.
Aechmina cortezensis McClellan, n. sp. .............00.0.0cee. 327
11. Paratype RV, #34726. 15. Holotype, oblique view of
LV showing spine of RV, #34725.
Saccarchites saccularis Swartz and Whitmore .................. , BES
#34772 LV.
Myomphalus?* Sp... ..:....22:c.e%ts.. sc tcieh cd ee ee ee 321
# 34750, RV.
Aechminajlongionsbasslererr ee 326
# 34723.
Beyrichiidae ? genus. ............255 oe eee 325
#34789 (X23).
Aechmina aff. A. phantastica Kesling ..........0.......000000..... BAN
#34724.
Aechmina equilateralis Bassler ...........0.......ccccccceeeeee 326
18. #34721 LV. 19. #34722 LV.
BULL. AMER. PALEONT., VOL. 62 PLATE 41
BULL. AMER. PALEONT., VOL. 62 PLATE 42
Figure
13.
i416, 17.
15.
S1turo-DEvonIAN MicroFauna: McCLeELLAN 367
EXPLANATION OF PLATE 42
All figures «45
Page
Uirichia obesita) McClellan: me Spe is... ccce.-ceiccecedeceecvnceseeosss 328
1. Holotype RV, #34780. 2. Paratype LV, #34781. 3. Para-
type RV, #34782.
UW ichiiams pameette k enc hc in enc a ee Renee coe Seene, Oey
4. #34783. 7. #34784.
Winchellatia fragilis McClellan, n. sp. ...............000000... 331
5a. Holotype LV, #34788. 5b. Ventral view showing pos-
teriorly directed lateral spines.
EON nell ages Deemer 5 se eee eee tec ee care tee eee ats 330
#34743.
Limbinaria sp. .................... goa a a eane Oe, Seana aero a son BYE
8. #34746. 9. #34745.
Eukloedenella aff. E. dalhousiensis Copeland ............ nee 332
10. Dorsal view, #34740. 11. #34741 RV.
Neokloedenella? sp. oo... Peer eh 333
# 34751 RV.
PSeucOlemerciiid: SD... ilsiycc2. ee oer teoet oteh oo eee ea. eee 334
# 34759.
Parahealdia aff. P. pecorella Coryell and Cuskley .... .. 834
14. #34777 RV. 16. #34753 RV. 17. #34752 LV.
ParapanchitesmS} ges. cote cea erences ae Meet eaee te 335
# 34754, RV?.
Figure
15, 19.
16517.
18.
20.
21.
BULLETIN 274
EXPLANATION OF PLATE 43
All figures «45
Page
Bairdia (Sp). <....0...08...0:. dee. ae eee 336
12384727 RVE2. 3534728 Ve
Bythocypris aff. B. alcocki Copeland |.....232...) ee 336
3. 334736 LV. 4. #34737 RV. 5. #34738 LV.
Bythocypris? Sp; ci. natn eee 35i//
# 34739.
Acanthoscapha aff. A. navicula (Ulrich) ...............0............ 337
7. #34719 RV. 8. #34720 LV.
Longiscapha nevadensis McClellan, n. sp. ................0..00000.. 338
9. Holotype RV, #34747. 10. Paratype, dorsal view, #34748.
11. Paratype LV, #34749.
Fealdia (SDs. .c...7..68. eee Ree Err re eee Py ec cubbnccoce:. 340
# 34742.
Berounelia: ‘Spi tei). c0d. neces eee ee eee 339
# 34729, RV.
Birdsallella sp. 2.0.5.5. tin. eee eee 340
# 34730.
Tubulibairdia cf. T. chaleurensis Copeland .......................... 341
15. #34776 RV. 19. Ventral view, #34776. ‘
Bolliidae .? . 317, 318
Carlini os ee 351
casteri,
Hyperammina .... 33 255, 279-281,
303, 305
cava,
Pasmmosphaera 34 253, 256, 261,
282, 283, 287
‘Ceratammina ............ 255, 261, 285,
286
chaleurensis, Tubuli-
bairdia aff. ... 43 341, 342
chaleurensis, Tubuli-
bairdia cf. .. ......43 251, 341
chitellata, Fairliella.. 302
370
INDEX
GoalaCanyonwe-.-ce es 242, 245, 247,
345, 346
GolewCreek tks eee 351
Columbus Limestone .. 304
compacta,
Sorosphaera . 34, 40 253, 254, 290,
291
confusa,
Sorosphaera ....... 34 262, 287, 288,
291, 292
conradi, Ulrichia ...... 328
constricta,
Hyperammina ... 33 254, 280
contorta,
Stegnammina ... 34 250, 256, 292
Copenhagen Canyon... 243, 346
Copper Mountain ...... 347
cornucopia,
Ceratammina ........... 261, 285, 286
cornucopia,
Ceratammina cf. 34 255, 261, 285
coronata,
Hemisphaeram-
mina ............ 35, 38 249, 303, 305
Webbinella .............. 303
corpulenta, Thlipsura 344
Cortez 1S eens 346, 351
Cortez Canyon .......... 250, 346
cortezensis,
Aechmina ............ 41 PAIL, BPAL
CUrta Balrdia <0.) 335
curva,
Hyperammina ... 33 281, 282
curvatus, Icriodus .... 252
curvus, Bathysiphon.. 281
cuspidata, Aechmina.. 326
cylindrica brevis,
Stegnammina .......... 292-294
cylindrica, Rhab-
dammina ....33,40 256, 275-277
cylindrica,
Stegnammina ..... 34 292-294
Cytheraeea t..).h........... 339
D
dainae, Nanicella ...... 319, 320
dainae?,
Nanicellame.. 36 249, 256
Dalhousie beds .......... 251, 333, 341
dalhousiensis, Euklo-
edenella aff. ..42 251, 332, 333
dentata, Hollinella .... 330
Diana Limestone ........ 241, 242
Diana’s Punch Bowl. 347
dicantha,
Rainwellae 2. 302
diplocystulis,
Welleriopsis 324
discoidea,
Fairliella 303
discoidea,
Hemisphaerammina 261, 303-305
discoidea ?, Hemis-
phaerammina ..... 35 256
Distacodus? sp. .......... 252
dorsinodus,
Myomphalus ............ 321
E
elongata,
Hyperammina ........ 279
Marsipella .............. 277, 278
Stegnammina ........... 293
RnGothynay pe eee 318, 319
equilateralis,
Aechmina ............ 41 251, 326, 327
Eukloedenella ............ ADIL, BBVA, ate:
exiguus,
Bathysiphon eee 276
expansus, Icriodus .... 252
extensa, Rudderina .... 343
F
EFairliella’ ~2-2............ abana), 302
fissurella, Halliella .. 320
Fourmile Canyon ...... 351
fragilis,
Winchellatia ...... 42 251, 262, 331
fundamentatus,
Spathognathus ...... 252
furca, Thlipsura ........ 345
fusca, Psammosphaera 282
USUI ae eee 318
G
gallowayi,
Endothyra 318, 319
Nanicellay =. 319
Garden Valley ............. 349, 351
Gateehii, oo focccon. 346
Gatecliff Dolomite .... 244, 247, 301,
348
gibbosa,
Metamorphina ... 35 309, 310, 315
Webbinella i 309
glabra,
Hyperammina 280
globulosa,
Webbinelloidea ...... 314
Glomospira’......... 249, 262, 314,
315
Sf
INDEX
gordialia,
Trochammina ........ 314
H
rane ee ae 320
Hanson Cree
Formation o6..--...5- 241, 243-245,
293
hattini,
Webbinelloidea ..35 250, 311, 312
Haplophragmoides .... 249, 256, HAs
Haragan Shale ............ 250, 261
Healdia.!¢ a 339, 340
Healdtaz sp. >...::..---3 43 340
hebesta,
Stegnammina .......... 293
hebesta?,
Stegnammina ...... 34 250, 254
Hemisphaerammina .. 249, 255, 256,
261, 301-305,
308, 310, 311,
314
hemispherica, Web-
binelloidea ....35, 38 262, 302, ae
Hindeodella sp. .......... 252
Hollinellageee--- 330
Hollinella sp. .......... 42 330
Horse Creek Valley .. 345
Horse Heaven
Mountainee.)= 346, 349
humerosus,
Paraparchites ........ 335
Hyperammina ............ 249, 254, 255,
279, 282, 315
I
leniodus =. 252
Icriodus spp. .............. 252
Tkes Canyon: .....:..:-:-.:- 243, 246, 347,
348
incomposita,
Atelikamara ..36, 40 255, 305, oe
inconstans,
Trichonodella ........ 252
inconstans,
Trichonodella cf. .. 252
inflata,
Sorosphaera .34, 40 290
imbricata,
Metamorphina ........ 305, 309
irregularis,
Sorosphaera ............ 289
J
jerseyensis,
IB CyIC hice eee 324
Welleriopsis ...... 41, 251, 324
K
Kelly Greek ......0::4.... 241, 242, 347
keslingi,
Phlyctiscapha_ ....41 251, 262, 322,
323
Kloedenellocopina 332
Filoedenia: «....6:.0..2:6 321
L
laevigata, Psam-
mosphaera ....34, 39 255, 262, 283
Lagenammina ............ 249
Laketown Dolomite .. 241
latericresens,
Teriodus... 252
limbata,
Bolbiprimitia .......: 320
IMBMaAria —. ois: .ca 263, 332
Limbinaria sp. ...... 42 250, 332
linearis,
Rhabdammina ........ 275
eibuOtubas &..meneeoe 249, 257, 315
Lodgepole Limestone 309
Lone Mountain .......... 245, 347
Lone Mountain
Dolomites 228. oe 238, 241-245,
247, 254, 291,
295, 316, 347,
348, 350
Lone Mountain
Limestone ................ 238
longior, Aechmina 41 326-328
Longiscapha ................ 251, 338, 339
longispina,
Winchellatia ............ 331
M
major,
Rhabdammina _....33 250, 255, 277,
278
malloryi, Stortho-
sphaera .......... 34, 40 254, 296
Manlius Limestone .. 321, 324
Mareh Spring «.....222" 244, 347
Marsipella’ de. .es2ca.: 249, 256, 275,
277-279
Marsipella sp. ........ 33 256, 279
Masket Shale .............. 241-244, 246,
247, 256, 347,
348
maxwelli, Psammonyx 261
McColley Canyon
HOLM Aabionmnes eee 244, 254, 299,
316, 323, 333,
348, 350
372
INDEX
McMonnigal
Mimestone. 22.0... 243, 244, 292,
294, 299, 322,
324, 329, 333,
342-344, 347,
348
media,
Ozarkodina aff. ...... 252
Metacopina .................. 339
Metamorphina ............ 249, 255, 261,
305, 308, 311,
315
mica,
Raibosammina ........ 286
Stegnammina .......... 286
Mille Canyon) -2.-........ 346, 348, 357
Mineral Hill. 349
minnesotensis,
Winchellatia ............ 331
Mitchell Creek .......... 244, 351
moodeyi, Beyrichia .... 324
mormani,
Stegnammina 34, 39 250, 254, ote
95
Thekammina .......... 294
multicella,
Sorosphaera ............ 288
multipunctata,
imbinariae 0. 332
mutaspina,
Rudderina ........ 44 251, 343, 344
Myomphalus ............... ByYA yy)
Myomphalus? sp. .41 251, 321, 322
N
INanICeliaweeet.. ee 249, 256, 318,
319, 320
navicula, Acantho-
scaphayratf. ...0.2 43 337, 338
Beecherella ............ sbi
Neokloedenella .......... 333, 334
Neokloedenella
PSUS sci ne ee 42 333
Newsomites ................ SBM/
Nevada Limestone .... 238, 244
nevadensis,
Longiscapha ....... 43 251, 338, 339
New Brunswick ......... 333, 341
nodosa,
Webbinelloidea ...... 312, 313
nodosa, Webbinel-
loidea aff. ... 35, 38 312
Nonioninaiae.22........... 317
Northumberland
Canyon 246, 348
fe)
obesita, Ulrichia ...42 251, 328
obliquus, Quasillites.. 342
Osgood Formation .... 253, 257, 276,
278, 301
osgoodensis,
Sorosphaera .34, 39 261, 288, 289
Ozarkodina -7..0.-.02-- 252
P
Palaeocopida .............. 320
Banderodus) ~..0.........- 252
papillata,
Thurammina .......... 299, 301
Parahealdiay 250, 334, 335
Paraparchites .............. 335
Paraparchites sp. ..42 335
pecorella, Para-
healdia aff. ....... 42 334, 335
perforata,
Arenosphaera ........ 290
Perkins Canyon ....... : 348
Perkins Canyon
Mormatione 348
Pete Hanson Creek .. 348
Phanassymetria .......... 340, 341
Phanassymetria sp. 43 340
phantastica,
Aechmina aff. ....41 327
phaseolina,
Bythocypris ............ 337
Phlyctiscapha ............ PASSA Va BY -ed
323
Phlyctiscapha sp. 41 323
Podocopinaw.......-..- 335
Prionodina’ sp. =: 252
prima,
Neokloedenella ...... 333
Primitiopsinae ............ 332
progressa,
Anictosphaera 34 254, 261, 283-
285
PSammonyxe eee 261
Psammosphaera ........ 249, 253, 255,
256, 261, 282,
283, 287
Pseudoleperditia ....... 334
Pseudoleperditia?
Dit eee 42 334
pusillum,
Haplophragmoides 318
Q
quadrangularis,
Stegnammina .... 34 250, 253, 295
Thekammina ....
373
INDEX
quadritubulata,
Thurammina _.......... 300
quadritubulata?,
Thurammina ..... 35 300
@uasiliteS” °28\....-..---: 342, 343
Quasillites sp. ........ 44 251, 342
R
TPyslojods JeGUUL 55.5 sce sesooes 349
Rabbit Hill
mormatione 1 242-244, 262,
281, 284. 318,
319, 325-329,
342, 345, 346,
349
Radiolariae. .p.csseee-e-e- 252
Raibosammina ............ 249, 256, rot
Rhabdammina ............ 249, 250, 253,
255, 256, 275-
277, 279
Rhizammina <...........-..: 278
Roberts Mountains .... 245, 280, 347
Roberts Mountains
Kormation 2... 241-248, 281,
283, 284, 291,
293, 297, 298,
300, 303, 305,
307, 309, 310,
314, 315, 319-
SPALL BPBY ayA3).
330-332, 346,
347-350
Roberts Creek.
Mountain =... 3 347, 348, 350
rockfordensis,
Hyperammina ........ 281
rockportensis,
Phlyctiscapha ........ 322
rostrata, Berounella. 339
Ruby Mountains ........ 351
Rud derinaee 1: 251, 343, 344
Rudderina spp. ...... 44 344
S
Saccamminay ss... 249, 298
Saccamminoides ........ 297
Sacearchites! =...) ee. 251, 323
saccularis,
Saccarchites ........ 41 2514320
scabra,
Aschemonella ......... 316
Sheep Corral Canyon 351
Sherman Mountain .... 351
siluriana,
Glomospira ........ 36 262, 314, 315
similis,
Webbinelloidea ...... 310-312
simplex,
Birdsallelia —o5-. 340
JECCRIIGIIE, ~. Jgeeesanansternoe 340
simplex,
Panderodus cf. ...... 252
Simpson Park Range.. 242, 247, 346
Spathognathus 252
Spathognathus spp. .. 252
Sorosphaera’ -.....2.-.- 249, 253, 254,
257, 261, 262:
287-292, 296,
297
Sorosphaera sp. ... 34 291
Sorosphaeroidea ........ 310
Sorostomasphaera .... 249, 250, 255,
297, 298
spinosa, Ulrichia ...... 328, 329
Stegnammina ............ 249, 250, 253,
254, 256, 261,
262, 286, 292-
295
Stomasphaera ............ 249, 250, 298
Storthosphaera .......... 254, 284, 296,
297, 306
subconfusa,
Sorosphaera .......... 289
Sulphur Spring Range 256, 351
symmetricus,
KeriOdUSi ee ee 252
Synprioniodina sp. .... 252
T
Telegraph Canyon .... 242, 256, 349
Tentaculitids .............. 252, 291
teresaccula,
Bolbiprimitia ..... 41 250, 320,321
hentulariinay 3. 275
Thekamminay ee 294, 295
LED SULANe eee 251, 344, 345
tholus, Meta-
morphina ...... 35, 39 255, 308, 309
Webbinella .............. 308
Thurammina 25s... 249, 250, 254,
299-301
Tippecanoe
SEQUENCE 5.0. .catesscccse 238
Tolypammina ............ 2A9, 250, 254,
315, 316
Toquima Range ........ 246, 347, 348
torta,
Marsipella .......... 33 249, 278, 279
tortuosa,
Tolypammina ... 36 250, 254, cae
tuberculifera,
Pseudoleperditia .... 334
374
INDEX
tubulata,
Thurammina ..... 35 254, 301
Tubulibairdia ......... 251, 341
Tubulibairdia sp. . 43 342
tubulifera,
Tubulibairdia 341
Turitellellay 4. 249, 257
triangularis,
Belodella 252
tricella,
Sorosphaera ....... 288
Trichonodella 252
triserrata,
Phanassymetria ..... 340
trituba,
Thurammina 35 300
Trochammina 314
Twin Springs Hills 349
Tyrone Gap 349
U
Wilrichellap eee 338
Wirichiae ee 251, 328-330
Ulrichia sp. 42 329
umbilicata,
Eukloedenella 332
unicostatus,
Panderodus 252
Ural Mountains ... 320
Vv
vagan,
Hyperammina ...... 315
Velibeyrichia 324
Velibeyrichia?
sp. 41 251, 324
ventriquetra,
Webbinelloidea . 35 250, 255, 313,
314
WwW
Wallbridge
Discontinuity .......... 238, 244
Waldron Shale ......... 248, 257, 261,
262, 279, 280,
287, 300, 302,
304, 310, 316.
317
waldronensis, Soro-
stomasphaera ... 35 250, 255, 297
298
Walti Hot SPHNES 351
Webbinella 301, 302, 308,
309
Webbinellodea 249, 250, 255,
261, 262, 303.
310, 314
Welleriopsis 251, 324
Wenban Limestone 248, 244, 247,
250, 322, 328,
335, 338, 340,
341, 345, 346
Whiterock Canyon 349
wilckensiana,
Beyrichia 321
Wildcat Peak ... 346, 348
Willow Creek 242, 245, 263,
297, 339, 350.
351
Winchellatia 251, 262, 331
Windmill Formation 242
Windom Shale 334
Wood Cone 351
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MOAN:
LII.
LIII.
LIV.
LV.
LVI.
LVII.
(ONS 225-230) eno Sin Ps 4-2 DIS py crecrce ce eat earn areeccnccnetecnnceenecreeness
Venezuela and Florida cirripeds, Antarctic forams, Lin-
naean Olives, Camerina, Ordovician conodonts, Niagaran
forams.
(Nos: 23-232). 420 pp.,’-10., plas: =
Antarctic bivalves, Bivalvia catalogue.
(Noa:-233,;230).. 9387. pp. 43 ple.
New Zealand forams, Stromatoporoidea, Indo-Pacific, Mio-
cene-Pliocene California forams.
(Maas 237ec08) 0 S88, DD, 5 Se a aeons
Venezuela Bryozoa, Kinderhookian Brachiopods.
(Nos. 239-245). EHO!) pisyee 5 gp) Sem tecce ccectracectesee ee eeeeeee ev cncteoare rence
Dominican ostracodes, Texan pelecypods, Wisconsin mol-
lusks, Siphocypraea, Lepidocyclina, Devonian gastropods,
Miocene Pectens Guadaloupe.
(Nos. 246-247). GH TPAD oO Ut LS greene tae en eee een ee
Cenozoic corals, Trinidad Neogene mollusks.
(Nos. 248-254). Sy PHN 0 8 epee OS as 0) Pp ena sea cause Ao ete eee ee ee
American Foraminifera, North Carolina fossils, coral types,
Belanski types, Venezuelan Cenozoic Echinoids, Cretaceous
Radiolaria, Cymatiid gastropods.
(Nos. 255-256). S2 TU D DSA OSD Smee ene een) eee
Jurassic ammonites.
LVIII. (Nos. 257-262). S05 pps ws 9 spl ster nee yee oles recone ee ee
Cretaceous Radiolaria, Cretaceous Foraminifera, Pacific
Silicoflagellates, North American Cystoidea, Cincinnatian
Cyclonema, new species Vasum.
LIX. (No. 263). SHAE DD eee a Re ee ee See eee
LX. (
Bibliography of Cenozoic Echinoidea.
Nos. 264-267). SSDP ie OS DISa ae eee eee ee neers
Jurassic-Cretaceous Radiolaria, cirripeds, Bryozoa, paly-
nology.
LXI. (Nos. 268-270). 3 GSA DD rs UD Seece ta eae ee eee ee
Mollusks, Murex catalogue, Cretaceous Radiolaria.
WSEN (CIN 20 EAST A sy a ee sec cieeeeee cere ace
Volumel.
II.
Til.
EV.
VI:
VII.
Trace fossils, ammonoids, Silicoflagellates, microfauna.
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studies, Carboniferous crinoids, Cretaceous jellyfish,
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Tertiary Arcacea, Mississippian pelecypods, Ambonychiidae,
Cretaceous Gulf Coastal forams.
(Nos. 38-41). AAR Tt S Sik DISA erect note cence ee ee
Lycopsids and sphenopsids of Freeport Coal, Venericardia,
Carboniferous crinoids, Trace fossils.
(Nos. 42-45). DANTE S01 8h estab) a) LN me en eR Reesor ai eee aes
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XXXV.
XXXVI.
XXXVII.
XXXVIII.
XXXIX.
XL.
N. Ye, 10017,.US:A2
(Nos. 80-87))5).334. ppi7 27) pls es ee
Mainly Paleozoic faunas and Tertiary Mollusca.
(Nos588-945)23064pps 3 0mplss ee eee
Paleozoic, Mesozoic, and Miocene fossils.
(Nos: 955100)! 420 appt SSipisy\ eo) eee
Florida Recent, Texas and South America Cretaceous,
Cenozoic fossils.
(Nos. <1Q1-708).) 376 hppe S36. opis. Ae ee ee
Tertiary mollusks, Paleozoic Venezuela, Devonian fish.
(Nose l09S114) 412) ppans4 ips.) cst -8 ee ee ee
Paleozoic cephalopods, Cretaceous Eocene, forams.
(Nos: DIS=116).. 73S pp S520 pls: 4. eee
Bowden forams and Ordovician cephalopods.
(No: a) eo S63 pp 65> plaints ne ee
Jackson Eocene mollusks.
(INos sl S3128) 745 Sm ppiea 7. aps. re
Mollusks, crinoids, corals, forams, Cuban localities.
CNos31295133)'-294 “pps. 39) piss. kel
Silurian cephalopods, crinoids, Tertiary forams, Mytilarca.
(INos:2134-139)s “448 pps 516 plss, 2.
Devonian annelids, Tertiary mollusks, Ecuadoran strati-
graphy paleontology.
(Noe; 140-145).:-4000 pp, 19) pls. ..2.3..2 8) eee
Forams, cephalopods, ostracods, conularid bibliography.
(INos:7146-2154).2386