HARVARD UNIVERSITY

LIBRARY
OF THE

Museum

of Comparative Zoology

W^ ^i^Tlve

zoology

^''""HaTva/univerBity

BULLETINS
OF

AMERICAN
PALEONTOLOGY

VOL. LVll

1969- 1970

Paleontological Research Institution

Ithaca,

New

York 14850
U.S.A.

MU3.

COivV.-'.

2-0011

LIBRARV

APR 27

i970

HARVARD
UNlVERSlTYi

INDEX
No separate index is included in the volume. Kach number is
indexed separately. Contents of the volume are listed in the beginning of the xolume.

CONTENTS OF VOLUME
Bulletin No.
255.

LVII

Plates

The Ammonite Fauna

Pages

of the Kialagvik

Formation at Wide Bay, Alaska PeninPart II. Sonninia Sowerbyi Zone

sula.

(Bajocian).

By Gerd

256.

E. G.

Westermann

1-226

1-47

227-321

48-62

New Middle Jurassic Ammonitina from

New Guinea.
By G. E. G. Westermann and
T.A.Getty

'z3'-B
i'-'JUS.

BULLETINS
OF

COMP.

200L'.

dec 3

'

HARVARD

AMERICAN *'""'"
PALEONTOLOGY

Vol.

57

No. 255

THE AMMONITE FAUNA OF THE KIALAGVIK

FORMATION AT WIDE BAY, ALASKA PENINSULA.
PART II. SONNINIA SOWERBYI ZONE
(BAJOGIAN)

By
Gerd

E. G.

Westermann

1969

Paleontological Research Institution

Ithaca,

U.

New York
S.

A.

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1969
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BULLETINS
OF

AMERICAN
PALEONTOLOGY
(Founded 1895)

Vol.

57

No. 255

THE AMMONITE FAUNA OF THE KIALAGVIK

FORMATION AT WIDE BAY, ALASKA PENINSULA.
PART II. SONNINIA SOWERBYI ZONE
(BAJOCIAN)

By

Gerd

E. G.

Westermann

November

20,

1969

Paleontological Research Institution

Ithaca,

New York

14850, U.S.A.

Library of Congress Catalog Card Number:

Printed

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in the United States of America

Arnold Printing Company

6If-135

CONTENTS
Page
Abstract

Preface

Stratigraphy

Upper boundary and correlation

of the Kialagvik Formation

Biostratigraphy of the Kialagvik Formation

17

Erycitoidcs hoivclli Zone

17

Sonninla soivcrbyi Zone

18

Otoites saiizei

Zone

Faunal relations and ages of the

22
iS".

soiverbyi Zone assemblages

Conclusions
Diversity
Fossil

30

and paleolatitude

31

localities

Systematic

32

description

36

Repository of types

Measurements

22

of

Family Phylloceratidae

36

ammonite

shells

36

Zittel

36

Genus Phylloceras Suess

36

Genus Partschiccras Fucini

38

Genus Holcophylloccras Spath

40

Family Lytoceratidae Neumayr

42

Genus Lytoccras Suess

42

Family Strigoceratidae Buckman

Genus Hcbctoxyites Buckman

44
44

Faiiiil\-

Oppeliidae

Genus
Fainil\-

Bunarelli

liraJfordiit

Hildoceratidae

47

Buckman

48

Hyatt

52

Genus Psi-udoliocnas Buckman

52

Genus Asthcnocrras Buckman

61

Famil\

Ilammatoceratidae Buckman

Genus Eudmctoccras Buckman

Appendix

/f//4///7o<vvv;j

nuclcospinosum Westermann, 1964

Genus Planammatoccras Buckman

Family Sonniniidae Buckman

Genus

Sonitinia Ba\le

63

72
82

90
92
92

Genus If itchrllia Buckman

108

Genus Prlrkodiifs Buckman

126

Family Otoitidae Mascke

128

Genus Docidonras Buckman

133

Genus Psrudotoilcs

157

Spath

References

166

Russian summary

172

Plates

173

THE AMMONITE FAUNA OF THE KIALAGVIK

FORMATION AT WIDE BAY, ALASKA PENINSULA
PART II. SONNINIA SOWERBYI ZONE (BAJOCIAN)
Gerd E. G. Westermann
McMaster University, Hamilton, Ontario

ABSTRACT
contact between Kialagvik and Shelikof Formations is now drawn at
the base of the Callovian regional unconformity; this accords with the original
definition and eliminates strong heterochroneity.
The Sori/iitiia soivcrbyi (Standard) Zone is represented by, below, at least
35
subgreywacke, greywacke, and mudstone containing the Eudmctoceras
dark shales
amplcctcns assemblage [. amplcctcns zonule] and, above, 50-100
with concretions containing the Pseudocidoccras assemblage [Pscudocidoceras
zonule.] Part of the incompetent shales are often missing due to strike-slip
faulting above the E. amplcctcns zonule. The underlying E. hoivcHi Zone is
separated by 80-100
poorly exposed, unfossiliferous, and little investigated
unfossiliferous
elastics; the overlying 0. sauzei Zone is separated by 110-130

The

sandstone and shale.

The E. amplcctcns zonule has yielded Eudmctoceras (Euaptctoccras) amplcctcns (Buckman), Docidoccras (?), Bradfordia? (Pracoppclia) and Hcbcsoivcrbyi Zone, L. discites
toxyites and is accordingly placed in the lower
Subzone, of the lowermost Bajocian. The Pscudocidoceras zonule includes, below, Sonninia (Euhoploccrasj, Eudmctoceras klimakomphalum (Vacek), and
Docidoccras s.s. also attesting the L. discites Subzone, and, above, abundant
evolute IVitchellia indicating middle to upper S. soivcrhyi Zone.
The Ammonoidea are placed in 15 genera, none new, 13 subgenera with
three new [Sonninia ( Alaskoccras)
Docidoccras (Pscudocidoceras) , Bradfordia} (Pracoppclia)^, 30 species with 13 to 20 new of which nine are named
[Partsc/iiccras ellipticum, Pseudolioccras costistriatum, Sonninia (Euhoploceras)
bifurcata, S.
(Alaskoceras) alaskcnsis, IVitchellia sutneroides, Docidoccras
(Pscudocidoceras) luidebaycjisc, D. (P.) camac/ioi, D.? (P.?) paucinodosum,
Bradfordia} (Pracoppclia) oppeliiformisli, and tivo subspecies [Pseudolioccras
maclintocki fastigatum, Eudmctoceras (Euaptctoccras) klimakomphalum dis,

-S".

coidalc^.

About one-half of the Ammonitina genera and subgenera range higher

than in Europe. Affinities are closest to Europe and secondly to South America;
affinities to the Western Pacific (Western Australia and Indonesia) are weak
and could be accounted for indirectly by migration via Europe and South
America. All genera are also known from other continents and only Pseudotoitcs is restricted to the Pacific. High faunistic diversity and species distribution suggest a somewhat lower latitude or a lower temperature gradient than
at present, or a combination of both factors.

PREFACE
This is the second part of a study of the extraordinarily
rich and well-preserved ammonoid fauna from Wide Bay (formerly Kialagvik Bay) comprising the almost universal assemblage of
the Bajocian Sonninia sowerbyi Zone of the Upper Kialagvik Formation. The first part (Westermann, Bull. Amer. Paleont., vol. 47,
No. 216, 1964) dealt with the Erycitoides howelli Zone which was
correlated with the Lndwigia concava Zone of the Aalenian (formerly called Lower Bajocian)
The great bulk of the E. hoivelli fauna was believed to be endemic to Alaska and the western Canadian Arctic until recently

Bulletin 255

when Sey and Kaladieva

soutliern coast of the

reported the same fauna from the

(1967)

Okhotsk Sea

in

eastern

far

Siberia:

late

Toarcian sandstone with Pseudoliceras beyrichi (Schloenbach) are

overlain by approximately 480 m siltstones which bear at the base
P. inaclititocki (Haughton) s.s. of early Aalenian age, in the middle
part Erycitoides hozvelli (White)

E. (Kialagvikites) spinatus West-

eYn\:inn, Pseudolioceras muclintocki whiteavesi

(White)

P.

aff.

(?)

and Inoccranius sp., and above luocerarmis sp.

and P. (?) aff. P.m. xchitcavesi. These siltstones pass into 370 m
sandstones and siltstones with Inoceramus sp., followed by Upper

P.m.

XL'hiteavesi,

Jurassic beds with erosional luiconformity.

has for the

The

E. hoxvelli fauna

time permitted the accurate dating of the thick

first

Middle Jurassic terrigenous sequences distributed throughout

ern Siberia. Furthermore, Tmetoceros

mann

has

now

also

cf.

east-

T. flexicostatian Wester-

been reported from the southern Andes (West-

ermann, 1967)
Wliile Part

ed

Wide Bay

of this

monograph was

(1 ext-figs. 1,2)

in

in press, the author visit-

August of 1964

for

two weeks dur-

HubOregon State University, assisted

in the field. McMaster University and the National Research Council of Canada financed this short expedition. All transportation
was by air, with commercial airlines to King Salmon at the eastern
end of Bristol Bay and small chartered planes expertly piloted by
bush pilots, via Pilot Point at Ugashik Bay to Wide Bay making
use of the intertidal slope at Preston Creek (Camp A) and a small
ing which 10 days were suitable for field work. Mr. Phillip

bard, then a geology student at

air strip of

(Camp

B)

former

along the sea

Pilot Point

oil

exploration at the

After detailed

the

mouth

collecting in the

S.

of Short Creek

sowerbyi Zone

Camp A, a chartered float plane from

Camp to location B on the northwestern side

cliffs east

moved

fossil

of

where the E. Iimvelli Zone of the coastal bluffs from

Pass Creek to the head of the bay and the important sections along
Short and Anderson Creeks were re-examined. Important technical
advice and aid were given by (Charles Rowett, then at the University of Alaska, R. L. Dettcrman and T. B. Ball of the U.S. Geological Survey, Alaska Branch, and by M. V. Kirk of Shell Oil
of the bay

Company.

monograph was

de-

layed because of the necessary re-investigation and taxonomic

re-

The completion

of the second part of this

Alaskan ammonites,

Pt. II:

Westermann

Scale

Text-fig.

1.

Index

map

in

Km

of south Alaska.

European and South American repreS. sowerbyi Zone ammonite

assemblages. Particularly the Hammatoceratidae and Sonniniidae
have been either in a state of utter taxonomic confusion as reflectvisions of the northwestern

sentatives

of

the

almost universal

NW

Eued in the classification of Buckman (1887-1907; 1909-30:

and
rope) or are poorly known with regard to range variation
affinity (South America)
Type specimens were studied in the
British Museum of Natural History and in the Geological Survey
,

Museum, London, with the help of M. K. Howarth and F. M.

Anderson; the subgenus Sonninia (Euhoploceras) Buckman was
thoroughly revised (Westermann, 1966) the most important classical sections in the southern Andes of Chile and Argentina were reexamined (Westermann, 1967) plastotypes of specimens described
in Tornquist's (1898) monograph were kindly sent by H. K.
;

Schmidt, University of Gottingen, and those of Jaworski's (1926)

monograph were sent by H. K. Erben, University of Bonn, Ger-

many. Of particular interest were the circum-Pacific genus Pseiidotoites Spath and the alleged Australian Zejnistephanus of which

Bl'lleti.x 255

plaster casts ol topotypes were liiriiished by P.

versity ol

Survey.

Western Australia, ami by R.

The author

locations of

(1933)

Lupher

also

re-examined

Bajocian section

at

in

(Coleman, Uni-

5. souwrbyi Zone
Oregon and Crickniay's

1963 the

in east-central

(1911)

}.

\V. Imlay, U.S. Geological

Moiuit [ura. California. Plastotypes of

the sonniniids from the inaccessible old collecting point at

Minewanka
furnished

i)y

in

D.

the Alberta

foothills

Frebold,

(cf.

1957a)

Lake
were

McLaren and H. Frebold, Geological Survey

of

Canada.

The author owes

sincere gratitude to R. \\. Imlay, U.S. Geo-

logical Survey, \Vashington, D.C., for

making

available to

him

all

Zone of Wide
Bay and also for fmnishing the plastotypes of all ammonites described and figined h\ him. The survey collections (USGS) were
made by Walter R. Smith in 1924, S. N. Daviess in 1944, L. B.
Kellinn in 1943 and 1915, and R. W. Imlay and Don Miller in
1948. A small collection was lent by L. G. Hertlein, of California
Academy of Sciences (C.A.S.) in San Francisco, and a few specimens were made a\ailai:)le by V. S. Mallory of the Thomas Burke
Memorial \Vashington State Museum, University of Washington
(U.W.) in Seattle. M. V. Kirk, Shell Oil Company, who loaned
the entile Aalenian and early Bajocian collections of Wide Bay
for the first part of this monograph, furnished stratigraphic information (cf. Part I) and polaroid photographs of a number of
ammonites from the S. soxvcvbyi Zone. However, most imfortiniately, the entire fossil collection had to he retmned to Shell Oil
fossil collections

Company

of the Survey from the

S.

soiverbyi

in 1964 after only a preliminary survey of the

assemblages was completed.

The taxonomy

of the

5.

soxoerbyi

Hammatocera-

tidae was fruitfidly (Hscussed with E. Elmi, Universitd' de Lyon,

and B. Cieczy, Museinn Koiiit, Budapest.
Mrs. Gay Walker redrew the text-figines and Miss V. Elkington
made the prints from films taken by the author.

STRATIGRAPHY
UPPER BOUNDARY AND CORRELATION OF THE
KIALAGVIK FORMATION

The

difficulties of coirelation

and definition

of formational

contacts caused by strong lateral facies changes are enhanced by

Alaskan ammonites,

the development ol
fossils

In
fig.

tlie first

II:

W^estermann

hiati

jxiiiuoiilonnities witli

part of this
,

as

indicated

i)y

which are difficult to detect.

monograph (Westermann 1964a, text-

and by bedding plane

4 opp. p. 338)

Pt.

taults

adopted the lithostratigraphic correlation

and formational boundaries of Shell Oil Company as communicated

by M. V. Kirk, although the alternate position of the formational
contact Kialagvik-Shelikof as placed by the U.S. Geological Sur-

vey had also been indicated. Without direct knowledge of the

exposures

tensive field

chose to reproduce the results based on the more ex-

work which

Geological Sinvey

was

as

admitted by R.

W. Imlay

of the U.S.

Company.

that carried out by Shell Oil

This correlation resulted in a remarkable alleged heterochroneity

of the formational contact transgressing along the

NW side of Wide

Bay through a lateral distance of only 10 km from tlie top of the

E. Iiozoelli Zone (Moose Creek Mt. Kathleen section) to the top
of the O. saiizei Zone (Mt. Mamie section) and back to the E.
hoxoclli Zone (Short Creek section) involving a stratigraphic interval of 200-300 m. Along the southeastern side of the bay, the
contact was placed within the intermediate

my

5.

sowerbyi Zone.

Wide Bay in
and correlation
of the formational boiuidary as originally implied by Capps
(1922) and as amended and defined by Kellum, Daviess, and
Swinney in 1944 (1945) previous to the more recent mapping
Based on

simimer,

1964,

subsequent brief

now adhere

to

field

work

at

definition

the

by Shell Oil Company (luipublished)

The base of the Shelikof Formation was originally
1922)

placed at the regional iniconformity

formity)

(locally

(Capps,

paracon-

developed throughout the "Cold Bay" [Puale Bay] area

including "Kialagvik Bay"

[Wide Bay],

of the

terrigenous and

tuffaceous beds bearing the Callovian guide fossil Cadoceras.

lies directly on the Lower
Middle Jurassic sequence (Aalenian and lower
Bajocian) is present at Wide Bay, which Capps (1922) named the
Kialagvik Formation. However, Capps had only visited the northwest side of tlie bay where the Bajocian (s.s.) is reduced, and he
probably did not see the actual contact which is usually concealed,
nor was he able to compile a complete stratigraphic section from
the isolated exposures {op. cit., p. 95) The upper expanse of the

While

Jurassic,

at

Puale Bay the Shelikof

thick

10

Bulletin 255

Kialagvik Formation becomes evident from the described exposures of "sandstones, sandy shale and conglomerate." Besides
the E. hoiL'clU Zone of the bluffs, Capps' KialagN ik Formation also
included the O. sauzel Zone of location No. 1-113 (10809), about
km upstream Caribou Creek at the west end of the bay. Previously T. \V. Stanton (in Capps, 1922, p. 96) noted that the
1

Souniuia and Inoceramus from this locality are "identical with

forms in No. 33 of Martin's Tuxedni Bay section" (Martin and
Katz,

1912,

p.

61:

lot

No. 33 with "StepJiauoceras, Harpoceras,

[O. sauzei Zone] and are younger than the E. hoicelli

etc.")

semblage. This location

of Kellum, et

al.

(1945,

is

fig.

placed in the "Inoceramus

as-

probably identical with location F 20

"Section near Camp 3" which was
2)
:

sp. C.

et al. (loc. cit.) the sandstone

Subzone." According to Kellum,

bearing this assemblage [O. sauzei

Zone] is separated from the "Hatnrnatoceras Zone" [E. howelli

Zone] beneath by an unconformity and about 125 m of poorly
exposed beds including, above, grey-green silty sandstones with

abundant plant fragments. The "Moose Creek

Company

tion" of Shell Oil

No. 1-113
Formation, 100-120

Mt. Kathleen

includes Capp's original

sec-

Kialagvik

which,

location

however, is placed in the Shelikof

above the formational contact at the top of

the E. howelli Zone.

et al. (1945), who carried out a summer's field work
and produced the geological map of Wide Bay, logically
continued Capps' preliminary work and successfully attempted
to compile and correlate the three sections of Caribou, Anderson,
and Short Creeks. Their placing and correlation of the formational
contact appears essentially correct and in agreement with the original definition. In the Anderson Creek section ("Creek near Camp

Kellum,

in 1944

No. 2
j)any)

'

of Kellum, et ah, "Mt.

Mamie

section" of Shell Oil

Oil (>ompany,
Shell Oil

i.e.

al.

above the sandstones of

at top of "Seymourites subzone"?)

Company

Com-

and Shell
the O. sauzei Zone (by

the contact was placed similarly by Kelhmi, et

The

al-

leged correlation of the E. JioivelU Zone sandstones of Caribou

Creek with the O. sauzei Zone sandstones of Anderson Creek as

proposed by Shell Oil (Company, appears to be due to the stratigraphic approximation of the respective arenite series in the latter
section; the intermittent argillaceous sequence has become thinner,

Alaskan ammonites,

Pt. II:

Westermann

more arenaceous and carbonaceous (Kellum,

Shell location No.

now dated

as

and

is

601

et

lower in

1945,

al.

fig.

2).

(WA

has been re-examined

(F43)

14)

O. sauzci Zone, based on the presence of

Parabigotites crassicostatus Imlay. Shell location No.

10-12

II

section,

tlie

600 (F 42)

poorly fossiliferous, but a single

is

}Docidoceras (Pseudocidoceras), based on preliminary identification

in 1964,

would suggest up

to 50-60

possible

5.

Zone

soioerbyi

in arenaceous-argillaceous and, below, carbonaceous fades.

km northeast of Anderson Creek,

predominantly arenaceous and conglomoverlaid by 30-40 m barren grey shales,

In the Short Creek section, 5

only the E. hoivelli Zone

eratic.

This sequence

is

is

well exposed at the steep right

bank near the mouth of

a tributary.

of silty shale with several thick beds

of conglomeratic sandstone

and lenticular limestone, the lower

Above follow about 150

50

of

Zone [L
cett)

which have now

definitely

1062: Stemjnatoceras

cf.

S.

been placed in the O.

triptolemus

1038: 'Witchellia' adnata Imlay,

Inoceramus

liicifer

WA16:

Eichwald ranges almost

sequence (F4, F24, F25,

WMla)

saiizei

(Morris and Ly-

Parabigotites sp.];

to

the

top of

this

which, therefore, probably belongs

Zone (cf. Imlay, 1955, p. 86) This sequence is conformably overlaid by dark grey shales with diabase
sills which have yielded Callovian ammonites at 140-150 m above
base [A463, F5] and are probably entirely of Callovian age. The
entirely in the O. sauzei

Kialagvik-Shelikof contact
shale

and sandstone

Kellum,

The

et al.

30-40

is,

series

therefore,

at

the

top of the

silty

of the O. sauzei Zone, as placed by

(1945)

barren shales of the Short Creek section separat-

ing the arenite sequences of E. hoxoeUi and O. sauzei Zones

may

be-

These shales can tentatively be correlated with the poorly exposed argillaceous sequences near the
bases of the Caribou Creek and Anderson Creek sections and, perhaps, with the Pseudocidoceras zonule (? and all or part of the
overlying unfossiliferous interbedded shales and sandstones) of the
long in the

S.

soiverbyi Zone.

southeastern side of the bay.

Along the southeastern shore

Creek, the bluffs and

of

Wide

Bay, east of Preston

expose from the base

(1)

10-20

mudstone with some greywacke beds, (2) about

interbedded silty shale and greywacke grading upwards

imfossiliferous

22

cliffs

Bulletin 255

12

subgreywackc

in massive fossiliferous

sowerbyi Zone]:

50-()()

(3)

(?

[E. (iniplcc tens

100 m)

to

zonule of

mostly highly

S.

fossili-

ferous shales with concretions [Pseiidocidoccras zonule of S.

sowerbyi Zone] grading upwards into {A) 80-100
unfossiliferons
interbedded sandy slialcs and sandstones which are topped by (5)

subgreywackes with intermittent

fossiliferous

The

silty

shale [Parabigo-

Zone emerges immediately west of Preston Creek but crops out below sea level
eastward. Tlie small cape just west of the mouth of Preston Creek
which forms tlie base of a high bluff (Shell loc. L 154) belongs
titcs

zonule of O.

to tlie T. ten
tiie

lie

E.

Jioxi'clli

zonide, formerly T. tenue-jlexicostatnm zonule of

highest E. hoxvelU Zone, lower horizons of which are exposed

in the shore

cliff at

L 556) However,
my locations ^V^A
.

5.

Zone].

saiizei

the western

Shell location

end of the Bay

(Shell Iocs.

555,

54 in the E. howelli Zone and

WA

3 and
4 which yielded the lowest probable
sowerbyi Zone faunules are stratigraphically separated by about

80-100

unfossiliferons terrigenous sediments, so that the position

of the zonal boiuidary

At

unknown.

is

most of the bluff west of Preston Creek is roughly

ecjuivalent to the bhdfs along the northwestern shore of Wide
Bay, including the base of the Caribou Creek section, while the
bluff of the E. amplecten.s zonide east of Preston Creek seems unleast

represented at the other side of the Ijay.The Pseiidocidoceras zonule

may

be c(|ui\alent

to

the

30-70

argillaceous

beds overlying

the E. Iioxvelli sandstones at the northwestern side of the bay.

The

Parabigotites zonule and prol)aljly also at least part of the underlying interljedded sandstone-shale sequence belong to the "Dnctylio-

ceras-Inocerarnus sp. C. Zone" [O. saiizei Zone] of Kellum, et

al.

which they believed to be the most persistent zone in the

Wide Bay area. This forms tlie upper thick arenaceous complex of
(1945)

the Kialagvik Formation.

map

However, Kellum, et nl. (1915, p. 7; diagrammatic section on

B) wrongly correlated the southeastern shore exposures with

known sequence

the better

of the other side of

miles from the

Wide

Bay.

The

end of the peninsula

continuing ... in the

from
all the way to the end of the bay and
mountain front beyond to the head of the
"sea cliffs

3i/^

valley",

believed to

represent the "DactyUcoeras-Inocerarniis sp. C. Zone", belong in

Alaskan ammonitks,

Pt.

II:

Westermam

13

S. soxverbyi Zone and in the southwest

upper E. JiowcUi Zone. In their diagrammatic section, the
Kialagvik-Shelikof contact was drawn at the top of the Pseudocidoceros zonule but on their geologic map at the top of the E.

the northeast to the lower

to the

aniplectens zonule and, soutliwest of Preston Creek, probably near

the top of the E. howclU Zone.

LOCALITY SYMBOLS

14

Bulletin 255

Alaskan ammonites,

NW

side

Westermann

Pt. II:

SE

WIDE BAY

of

side

(composite)

(Short Creek)

(Anderson Creek)

15

:)
300 M
TizJ

//

/
Pseudocidoceras

(i}7 /

zle/

^'Z-T .

Vl.1038

/ SOWERBYI

^'V
'

/--T^^

/
VLI062

amplectens zleS

ZONE
/

UPPER
HOWELLl ZONE

fe^^*'^}^

Text-fig. 4.
Tentative correlations of the upper Kialagvik Formation
between the Anderson Creek and Short Creek sections at the northwest side
(part. Keilum, et al., 1945) and the SE. side (composite) of Wide Bay. The
important fossil localities, formational contacts, ammonite zones and zonules

are indicated.

BIOSTRATIGRAPHY OF THE KIALAGVIK FORMATION

ERYCITOIDES HOVVELLI ZONE

The

results of

are consistent with

my
the

brief field

work

in

1964

(Text-figs.

2-9)

faunal sequences as reported from

Kialagvik Formation along the northwestern side of

Wide Bay

the
in

monograph (Westermann, 1964a, p. 339 ff.)

The thicknesses of the faunal zonules and their intervals are confirmed for the Short Creek and Caribou Creek sections except for
the E. howelli zonule which now appears somewhat thicker. Howthe

first

part of this

Bllletin 255

16

CO
-I

O
i

E
t,

>

to

en

Ui.

o-

>.

-~

>0"!'ivD

CO

Nvioorva

U.S.

Phylloceras (Zetoceras)

sp.

IMLAY

Holcophylloceras costisparsum
Lytoceras

1+

eudcsianum (ORB.)

aii. L.

Hcbctoxyiics

(ORB.)

P. zetcs

cf.

Partschiceras ellipiicum

Geology Survey

BUCK

H. hebcs

aff.

Bradfordia? (Praeoppclia) oppeliiformis subgen.

et sp.

Pscudolioccras madintocki fastlgatum subsp. n

Pseudolioccras cost'utriatum

Asthcnoceras

sp.

A. nannodes (BUCK.)

aff.

cuJmctum BUCK
E. (Euaptetoceras) klimakomphalum discoidale

Eudmctoccras

aff. E.

(s.s.)

E. (Euaptetoceras) aff. E. nuctcospinosum

E. (Euaptetoceras) amplectens

(BUCK)

Eudmetoccras s.l.indct. [(J]

Planammatoceras (Pscudammatoccrasf)

subsp. n

WEST

[? Var./subsp. aguilonia

(1)

IMLAY]

Cf.

Sonninia (Euhoploceras) bijurcata

Sonninia (Euhoploceras

sp.

benneri

aff. P.

(HOFFM.)

sp. indet

?)

Sonninia (Alaskoceras) alaskensis subgen.

et sp.

IVitchellia sutncroides sp. n

Pclckodites

Docidoceras

D,
D.
D.
D,
D,
D.
D.
D.

cf.

(s.s.)
(s.s.)

P. pclckus

aff.

BUCK

D. longalvum

(VACEK)

(Pseudocidoceras) <widebayense subgen.

et sp.

5+

(1)

D. viidebaycnsc
(Pseudocidoceras'? ) aii. D. 'widebayensc
(Pseudocidoceras) camachoi subgen. et sp. n
(Pseudocidoceras)

cf.

(Pseudocidoceras) sp.n.

?l7Pscudocidoceras/suhgei\. nov.] sp.n.

?

(Pseudocidoceras?) paucinodosum subgen.

(Trilobiticcras?)

Pseudotoites

cf.

Pseudotoites

cf. P.

sp.n.

et sp.

indet

P. argentinus

ARKELL

transatlanticus

(TORNQ.)

Text-fig. 6.
Occurrences of ammonite species from the 5. soiverbyi Zone
of Wide Bay; numbers of identified specimens indicated. Localities of the
U.S. Geological Survey (U.S.G.S.), California Academy of Sciences (C.A.S.),
Shell Oil Company (Shell), University of Washington (U.W.), and of the
author.

Alaskan ammonites,

Pt.

Westermann

II:

17

FORM

KIALAGVIK
5
"

!l

ill

\
\

w
o

>

>

J>

cf

phyllocerasiZeioceras)

P'es (ORB)

Partschiceras elhpticum sp n
--.

Hoicophylioceros costisparsum

---

Lytoceros

off-

IMLAY

L eudesionum (ORB)

Pseudolioceras mchntocki fosiigotum subs

^.
-

costistriotum sp n

Asthenoceras

A nannodes (BUCK

aff.

Eudmetoceras

Euopletoceras

klimokomphalum

dis

jidale subsp n

omp/ecfens {BUCK )[var/7 subsp osui/omo IML.]

()

Sonninia (Euhoploceras) bilurcato sp

(-? subsp)

E eudmefum BUCK.

aff.

sp ndel,

(E?)

S.

lAlaskoceros) alostfensis sp

n.

Witcheliia sutneroides sp n

--

Pelekodilesct Ppe/ekus BUCK.

(SpoluMes

Docidoceras

?) sp nov

off.

spolions BUCK.

D longolvum (VACEK)

off

(Pseudocidoceras)

wideboyense sp

t*(defcoyense

(P)

cf

(P)

camachoi sp

(P)(') sp

novB

(R)-'

Pseudoloites cf

P orjen/mus ARKELL

/ronsol/ondcus

(TORN

Bradfordio 7 iProeoppelia) oppeliiformis

.-

cf.

Hebetotyites off

K hebei

BUCK.

Composite section of the S. soiverbyi Zone and

Text-fig. 7.
distribution at the SE. side of Wide Bay. Stratigraphic range of
(sections) and relative abundances of species indicated.

ammonite
localities

Bulletin 255

18

Comparison of ihe \ertical ranges of Aalenian and early

Text-fig. 8.
Bajocian ammonite genera and subgenera between southern Alaska and northwestern Europe. Note the higher ranges of many taxa in southern Alaska.
ever, there

is

no evidence

for the previously inferred thickness in-

crease towards the southwest.

Formation

of the Kialagvik

The

concealed.
ule)

E.

teres

Creek

at Short

zonule

fossil

evidence

is

now

named

here

zonule

guide

teres

WB

5,

confirmed in

et al.,

Westermann, wliich

fossil for this

assemblage.
at

The

the

45 Akni,
is,

there-

T. tenue

top of the

WB

confirmed in the same section (my location

48A - 109) by abundant Tmetoceras tenue Wester-

Zone

uses

avail-

100

almost totally

is

Kellum,

(formerly tejiue-jlexicostatum zonule)

E. howelli

as

is

c.

(formerly E. teves-projinuhis zon-

(my location

by abundant Erycitoides

F12)

12

adcHtional

separating lower and upper E. howclli Zone,

the Short Creek Section

fore,

No

the tentative T. scissum zonule since the basal

aijle for

is

mann, with T. jlexicostatum Westermann and T. cf. kirki Westermann; T. tenue is the most distinct species and named guide fossil.
Both zonules also bear the zone index E. hozvelli.
SONNINIA SOWERBYI ZONE

The

zone contains the E. arn plectens zonule below, and

tiie

exceptionally fossiliferous Pseudocidoceras zonide aliove; botli have

been clearly identified only along

km

strip at the southeastern

shore of the bay.

E. cnnplectens zonule.

The

zonule consists of 10-12

mostly

Alaskan ammonites,

Pt. II:

Westkrmann

19

Bulletin 255

20

massive subgreywacke and yields abundantly Endineloceras (Euaptetoceras) arnplecteu.s

(Imlay)

],

(Buckman)

[incl. var.

moderately conmion Docidoceras

pauci nodosum,

n.

sp.,

and

scarce

cf.

D.

Bradjordia? (Praeoppelia) oppeliiforrnis,

(P.) xvidebayense,
n.

sp.,

H. hebes Buckman, Sonninia (Eulioploceras})

tids.

This

or subsp. aguilonia
(Psexidocidoceras

?)

n.

sp.,

Hebetoxyites

aff.

sp.,

and phyllocera-

the "acme-zone" of E. amplcctens, but the species

is

ranges into the E. howelli Zone below, and into the O. sauzei Zone
above.

The

common

only

to this zonule

species

which appears

to

be restricted

D.} paucinodosum which could, therefore, be con-

is

sidered as guide

fossil.

The

and the

Otoitidae, Oppeliidae,

strigo-

ceratid Hebetoxyites attest to a Lower Bajocian [post-L. concava

Zone] age; the common E. amplectens and the stratigraphic posi-

below the Pscudocidoceras zonule, yielding early Soyininia and

Docidoceras s.s., restrict the age correlation to the lower or basal
the (European) Ludxoigia
S. soxverbyi Zone, approximately to
tion

discites

Subzone.

Although the massive character of the subgreywackes (? and

greywackes) and the apparent random orientation and poor preservation of most ammonite shells suggest submarine slumping or
similar processes of mass-wasting, there is no paleontological evidence for significant stratigraphic mixing of the faunas. The common Eudmetoccras amplectens are usually strongly corroded, fractured, and often fragmented wiiich coidd be due to reworking;
alloditliony would be consistent with the known occmrence of this
species in the luitlerlying E. hoivelli Zone. The other ammonites,
all imknown fiom the Aalenian, are mucli better preserved and,
therefore, almost certainly autochthonous. However, even under
the assumption that the E. amplectens are reworked, the E. amplectens zonule

is

to

be placed in the lower part of the

S.

sowerbyi

Zone because of the superimposed Pscudocidoceras zonule.

Pseudocidoceras zonule.
(?

to 80 m)

The

zonule consists of at least 40

shales with calcareous concretions

are separated from the E. amplectens zonule by 10

which probably
-

20

unfossili-

ferous shales. This incompetent interval has been structurally re-

duced or

totally suppressed

wherever the competent E. amplectens

lies below sea level where the

exposed and the contact

sequence appears complete. Thus,
zonule

is

all

or part of the structurally

Alaskan ammonites,

20-30

isolated

Pt.

Westermann

II:

shaley section ol locality

longs at the base of

WA

21

15 apparently be-

Pscudocidoceras zonule. This zonule bears

tlie

most diverse and best preserved known

ammonite assemblages of the S. sowerbyi Zone in North America,
and possibly of anywhere but a few localities of Europe. However,
the fossiliferoiis exposmes known in 1964 were restricted to only

without doubt the

350

400

Bay (from

richest,

length of the shore

loc.

WA

10 to loc.

along the south side of the

cliff

WA

oW

15;

Iocs.

21251 and 12405

were concealed or destroyed in 1964)

The

zonule includes several ammonite assemblages in over-

lapping succession. However, horizontal variation being unknown,

the apparent famristic changes could be due to differences in biofacies,

post-morte?n

and preservation. Nevertheless, the obmore than local significance

drift,

served vertical ranges are probably of

because

(1)

lithofacies varies

little

throughout the zonule,

(2)

ranges vary greatly from species to species with frequent overlaps,

and

there are certain resemblances in the

(3)

ammonite

succes-

sion with other areas.

A
parent.

two-fold faunistic subdivision of the zonule

The

hijurcata, n. sp.,

S.

ividebayense, n.

n. subsp.,

and the

formis, n. subgen.

klimakomphalum
ed

n. sp.,

Pseudolioceras

sp.,

less

common

and

n.

sp.,

n.

subgen. and

n. sp.,

sp.,

D.

mackUntocki fastigatuni

Brodfordia} (Praeoppelia) oppelii-

and Eudtnetoceras (Euaptetoceras)

discoidole, n. subsp.; all are essentially restrict-

to this part except for

the zonule.

strongly ap-

(Alaskoceras) aJaskensis, n. subgen. and n.

Docidoceras (Pseudocidoceras) catnachoi,

(P.)

is

lower part contains abundant Souuinia (Euhoploceras)

The upper

D. xvidebayense which ranges throughout

part bears

abundant Witchellia

D. (Pseudocidoceras) xvidebayense,

the top, moderately

common

Other ammonoids such

n. sp.,

Pseudolioceras costistriatum,

as Phylloceratidae, Lytoceras,

Pelekodites, Docidoceras

s.s.,

siitneroides,

and, especially at

and Pseudotoites are

n.

sp.

Eudmetoceras,

scarce throughout

and Asthenowas found only in a "nest," i.e. a single concretion, in the

middle of this interval. A third assemblage may be present at the
the zonule so that their ranges cannot be established

ceras

poorly
19801,

known apparent
?

USGS

CAS
and the common

19863,

(Pseudocidoceras)

base of the zonule

29011) which
S.

(Iocs.

WA

15, ?

USGS

yields only a few

(Euhoploceras),

S.

D.

(Alasko-

Bulletin 255

and

ceras), Pscudolioccxis,

mon

wood and

fragments of

The
choi

/>.?

The

(PracuppcUa).

only other com-

larger fossils are clusters of Inoceramiis lucifer

is

Eichwakl and

leaflets.

lower assemblage, for whicli D. (Pseiidocidoceras) cama-

chosen as guide

single Docidoceras

fossil,

1).

aff.

has also furnished near the top a

lonirfthniin

(Vacek)

supporting the cor-

Subzone of the lower

sowerbyi Zone. The ujjper assemblage which is probably best

relation of this interval with the L. disci tes

S.

characterized by the e\()Iute WitcJicUia sulneroides, sp.

clearly in

ilie

both or either of the northwest

trigonolis

and WitcJicUia

morphologically

belongs

Emopean Subzones

of Sonninin

indeed
is
between "Zugophorites" Buckman
trigoualis Subzone and typical
of the
hwxiinsciiJa:

IT.

sutneroides

intermediate

subgen.?]

[]\'it( lu'llia
]]'it( licllia

n.,

liigher part of the S. soxvcrbyi Zone, equivalent to

of the

.S'.

laeviusculn Subzone.

IT',

OTOITES SAUZEI ZONE

The ammonoid

assemblage of the Parahigotites zonide has

re-

monographed by Imlay (1964) who has given ample

evidence for O. saiizei Zone. A number of Parahigotites crassicostatus Imlay and some Bradfordia sp., Stemmatoceras cf. 5. triptolemus (Morris and Lycett) "Witchellia" adnata Imlay and, sig[var. or subsp.
nificantly, Eudmetoceras amplectens (Buckman)
cently been

agiiilonia Imlay]

WA
P.

12,

WB

1,

crassicostatus

zonule by 80-100

The

have been identified from

WB

14,

and

zonule

is

Shell localities

separated

from

my

localities

1038 and
the

WA

1062.

1,

The

Pseiidocidoceras

m unfossiliferous arenaceous shales and sandstones.

beds superposed on the Parahigotites zonule are almost barren

but yield abundant luoceramus lucifer Eichwakl

which appears to Ijecome extinct at ilie top of the O. saiizei Zone
(Imlay, 1955, p. 86) The upper O. saiizei Zone assemblage with
of ammonites,

Skirroceras kirsclmeri Imlay from southeast Alaska (Imlay, 1964, p.

B7) appears to be absent at Wide Bay, probably because beds of
this

age are missing under the paraconformity of the Callovian

Shelikof Formation.

Alaskan ammonites,

Pt. II:

Westermann

FAUNAL RELATIONS AND AGES OF THE

ZONE ASSEMBLAGES
The ammonoid fauna
is

in

its

global

of

tlie

relationship

23

SOWERBYI

S.

sowerbyi Zone at

5.

admixture of Pacific elements and a strong regional

character at the species level. Tiiis

is

Wide Bay

European,

essentially

an

with

(endemic)

evident from the following

discussion of the principle areas of development of this assemblage,

American Cordilleras, the southern Andes, Western

and Emope. The Phylloceratina, represented by Phylloceras, PartscJiiceras, HolcophyUoceras, and the Lytoi.e.

the North

Australia

Indonesia,

known

hy Lytoceras, are
cosmopolitan and, therefore, omitted from
ceratina, represented only

pointed out that their relative al^undance

this
is

be entirely

to

discussion;

it

than commonly attributed to eugeosynclinal environments,

ticularly at oceanic margins.
era,

one appears

to

par-

Ammonitina genNorth America while a number

the

be endemic to

of the others have

known from

Of

is

significantly smaller

1 1

described

here longer vertical ranges

other continents, which

is

than

essentially

previously

the

European

epeiric Jurassic (Text-fig. 8)

NORTH AMERICA
Southeast

J /fli/ffl.

the presence of the

S.

Surprisingly, there

is

no good evidence

sowerbyi Zone in Alaska other than at

Bay, although the Bajocian

is

well

known from

for

Wide

the fossiliferous

Tuxedni Group of Cook Inlet (Imlay, 1962, 1964) where the

O. saiizei Zone seems to follow directly on the E. howelli Zone.
However, unfossiliferous beds, which could represent the apparently missing zone, are sometimes intercalated in the thick sedi-

mentary sequences.
Imlay (1964) described from the Tuxedni Group of the Talkeetna Mountains several ammonites which are identical with or
closely related to species occurring at Wide Bay in the 5. sowerbyi
Zone. Sonninia (Euhoploceras) bifnrcata, n.
B33,

pi. 4, figs. 5, 6, 12;

"Sonninia

could possibly belong in the

5.

sp.

(Imlay,

n. sp. undet."; p.

B14)

1964, p.
,

which

sowerbyi Zone, and Witchellia sp.

were described from the "upper half of the lower sandstone." The
former species, however, is said to be usually associated with
Emileia constricta Imlav, such as in the lowest fossiliferous beds

Bulletin 255

24

of the

BI4,

Red

Glacier Formation of the Iniskin Peninsula (op.

pi. 4, fig.

cit., p.

11, table 13). E. constricta

is a close ally of E.
polyshides (^Vaagen) and a good indication of the O. sauzei Zone.

S.

10,

(Alaskoceras) sp.

fig.

1,

"Sonniuio

2;

and

siltstone"

cf. S.

said to

is

alaskeusis, n. sp. (op.

cit., p. B33, pi. 2,

nodata Buckman") occurs in the "upper
be associated with Stephanoceras (Skirro-

aff. S.

ceras) spp., again clear indicators of the O. sauzei Zone. Eudyneto-

ccras (Euaptetoccras) amplectcus

lonia (Imlay)

(op.

B35,

cit., p.

(Buckman)

[var. or subsp. agiii-

pi. 4, figs. 1-4; 9; pi. 5, figs. 4, 7-9;

was described from a 60 m

Zone assemblage, as
well as from the Parabigoiites zonule of Wide Bay. Planammatoceras (Pseudainmatoceras?) aff. P. benneri (Hoffman)
(op. cit., p.
"Witchellia} aguilonia Imlay,

n. sp.")

interval again yielding the typical O. sauzei

B33,

pi.

figs.

3,

known fiom

2-4:

"Souuiuia

cf.

S.

patella

Zone and L.

Waagen")

formerly

Subzone of Europe
and lierc described from_ the 5. soxverbyi Zone of Wide Bay, was
described from the O. sauzei Zone assemblage of the Talkeetna
Mountains. However, the alleged Witchellia cf. W. laeviuscula
de C. Sowerby) (Imlay, p. B35, pi. 7) is not a true Witchellia
(J.
of the

W.

the L. coticava

discites

laeviuscula gioup.

Western Canada.

The

S.

soxcerbyi

Zone

is

usually absent un-

der paraconform O. sauzei Zone or, more frequently,

aninn Zone.

The

only

known

humphriesi-

S.

exceptions are probably part of the

"middle sedimentary division" of the Hazelton Group of Hudson

west-central British Columbia, and a poorly known
horizon in the Fernia Group of "Devils Point" at Lake Minne-

Bay Mountain,

wanka, Alberta.

The

small collection of sorniiniids from scree of

Mountain was described by McLearn

to

.S'.

(1926)

Hudson Bay

and dated

hutnphriesianutn Zone. However, four of the five

as O. sauzei

new

specific

names under Sonniuia and Guhsania McLearn were based on single,

incomjilete, and distorted phragmocones only and are here regarded
as noniina dubia. Guhsania bella McLearn, the type species, has
a discoidal, almost oxycone shell, with strong rectiradiate simple
costae probably bearing lateral tubercles, and appears to be intermediate between Sonninia (Papilliceras) and Fissilobiceras. "Sonninia Hansoni" resembles the Andean S. espinazitensis Tornquist;
".S". skawahi"
and ".V. silveria" may belong to a single species of

Alaskan ammonites,

l:^ W.

Pt. II:

Westermann

25

AUSTRALIA

subgenera
species

Text-fig. 10.
Cosmopolitan affinities of the principle ammonite assemblages of the S. soiverbyi Zone of Wide Bay, possibly indicating migration
routes; based on numbers of common subgenera and species. Note that the affinities between southern Alaska and Western Australia
Indonesia could be
accounted for by migration via Europe and South America, and that the assumption of continental drift would alter the relative position of the AustraloIndonesian area significantly.

Witchellia ("Zugophorites") or possibly partly to Sonninia (Eiiho-

from Lake Minnewanka and evofrom the middle part of the northwest European
S. soiverbyi Zone. Guhsania bella and "Sonninia Hansoni" could
come from a higher level, i.e. the O. sauzei Zone. "Guhsania ramata" is known from a single small fragment only, resembling G.
ploceras), resembling the species

lute Witchellia

bella.

The long known poorly preserved small "Devils Point" assemblage (Whiteaves, 1889) representing the only good evidence

Bulletin 255

26

of the

5. soii'crbyi

(1957a, pp. 48-49,

Zone from

(Euhoplocevas/ or Alaskoceras})
ploceros)

cf. 5.

ceras) sp. (loc.

(op.

cit., pi.

cit., figs.

2,

3)

liere identified

is

bifurcata, n. sp. (op.

cit., pi.

20, fig. \)

S.

(EuJioplo'

and Witchellia ("Zugophorites")

sp.

19, figs. 2 a, b)

Westerji U.S.A.
is

was redescribed by Frebold

with Sonninia
gracilis (Whiteaves)
5. (Euho-

(Jan.ula,

and

19-20)

pis.

\\\

northern California, the

tentatively indicated in the

Shasta County, by

(?)

Sonnina

Mormon Formation

S.

sozverbyi

Zone

Mount

Jura,

of

(Eiilioplorcra.s) "schiiclierti"

(Crick-

may, 1933, }wm. dub.)

The
America

next best representation of the

after

The

Wide Bay

is

.S.

sowerbyi Zone in North

Supplee area of east-central Ore-

in the

was described by Lupher (1941) and was re-inand the author (unpublished),
but tlie fossils ha\e not been described. According to Lupher and
Imlay, the lower part of the A\'^eberg Formation has yielded Tmetoccras, ^vhich in its upper range is associated with Praestrigites and
Docidoceras; this association, not verified by this author, would
extend the range of Tinetoceras into the S. soivcrbyi Zone. Above
follows a rich assemblage of large Sonninia (EiiJioploccras) sp. and
gon.

section

vestigated by Imlay (1964, p. B19)

strongly reminiscent of the

IVilclicUia ("ZugopJwrite.s") sp., both

Lake Minne^vanka forms but preserved too poorly

for

specific

identification; this assemblage probably also includes sparse Strigoceras, Praestrigites},

Docidoceras

and Eudmetoceras?, thus

s.l.,

re-

sembling the assemblages of the northwest European lower and

middle S. sozverbyi Zone. Slightly higher is an assemblage of abundant Asthenoceras sp., known in two specimens from the L. murchisonae Zone of England and from the

S.

sozverbyi

Zone of Wide Bay,

with sparse Witchellia and Sonninia (Euhoploceras). This

Warm

is

fol-

lowed by the O. sattzei Zone

mation which also contains abundant Asthenoceras.
In sunuiiary, the other North American assemblages of the
S. sozverbyi Zone have in common with Wide Bay five to seven
genera and at least six subgenera, all of which, however, also occur in Europe; the small number (two to four) of common species
is probably due to the poor knowledge of the non-Wide Bay asassemblage of the

Springs For-

semblages and specific affinity actually appears to be quite high.

Significantly, Bajocian Pseudolioceras

and

particularly the

common

Alaskan ammonites,

respectively

typical

Pacific

Westermann

Pt. II:

ami Wide Bay forms

27

P.sc'udotoites,'

Docidoceras (Pseudocidoccras) and SotDiiiiia (Alaskoceras) appear

to be absent while the strigoceraticls, present in the 5. sowerbyi

Zone of Oregon, make their appearance in Alaska only in the

O. saiizei Zone. These statements are highly tentative awaiting the
monographic study of the Oregon ammonite assemblage. The apparent close affinity of the Oregon assemblage, the only better
known assemblage of the North American S. sowerbyi Zone except
for Wide Bay, to the northwest European epeiric seas assemblages
may be owing to ecological resemblance.

SOUTH AMERICA

Souther?! Andes.

The

S.

soiverhyi

Zone appears

to

be absent

in the northern and central Andes (Guiana to Peru) and, contrary

to the opinion of Arkell (1954, p. 591; 1956, p. 585) and Imlay
(1964, p. B21) tlie e\'idence for its presence in the soutliern Andes
is far from excellent although approximately every second genus,
,

every fourth subgenus, and probably several species of the

Wide

Bay assemblage are found there (Westermann, 1967). In northern

Chile and northwestern Argentina three major assemblages are
present, stratigiaphically above early to middle (?upper) Aalenian
beds and below the widespread O. sauzei Zone; they have respecgerthi
(Jaworski)
tively been named after Eudmetoceras (?)
and Pseudotoites singularis
"Pleydellia" piichensis (Burckardt)
,

(Gottsche) (op.

The
you'or^A/i

ceras)

cf.

mocrickei

E. gerthi asseml^lage has also yielded

Westermann (1964a) ,. (?) kochi

E.
E. klimakomphalum (Vacek)
,

Eudmetoceras

(Prinz)
(E.)

klimakomphalum

Sonuiuin (Euhoploceras) sp., Zurcheria

"Fontannesia" austroamericnna Jaworski, Tmetoceras cf.
sparse

(?)

E. (Euapteto-

[not Bradfordia, with hollow-floored keel]

(Jaworski)

and probably
sp.,

fit.).

(?)

(?)

T.

scissujn (Benecke) and T. cf. T. flexicostatum Westermann. This

assemblage was dated as upper Aalenian by Jaworski (1926) and
as basal Bajocian L. discites Subzone by Arkell (1956, p. 585)
either date or both may be correct, and the possibly somewhat
;

Mm.

carlottensis Whiteaves, 1876, from the S. humpricsianum Zone of southeastern Alaska and coastal British Columbia, is not a Pseudotoites as supposed
by Arkell (1954. p. 587; 1956, p. 538, 542) but a Zemistephanus (Imlay, 1964,
p.

B52).

Bulletin 255

28

condensed assemblage
cian boundary.

The

now

is

placed at about the Aalenian-Bajo-

"P." puchensis assemblage consists of closely related species

(or a "plexus")

hammatoceratids, possibly intermediate

of late

Burckhardt (1903) imder "HarpoH. Jtialarguensc, n. sp., H. hauthali, n. sp.,

to early sonniniids, described by

ceras pucl\cusc, n. sp.,

WitclielUa argcutina,

Harpocerns

Harpoceras striatuliim Sowerby and

sp.,

n.

klirnnkotupluiltun

and

Vacek,"

well

as

as

abundant

of

Sonninia
This assemblage is only known
from west-central Mendo/a and may be approximately equivaWitchellio}, n. sp. indet.

(Fuhoploccras) adicra

a single nucleus resembling

(W'aagen)

lent in age to the E. gertlii assemblage.

The
and

P. singularis

includes

(Tornquist)

the sparse P.

and probably

quist)

tiniis Arkell,

(?)

Pseudotoites
(n.

(Tornquist)

also P. transatlanticus

S.

evolutum

(Latotoites)

subgen.) spJiaeroceroides (Torn-

Sonninia ("Sonninites")

cens Tornquist and

zitteli

(Papilliceras)

(Gottsche)

P. argen-

S.

intiimes-

espinazitensis

S.

cf.

(Torn-

This assemblage belongs, therefore, in the iq^per part of the

soiverhyi Zone or, possibly, at the base of the O. sanzei Zone.
While the affinity of the E. gerthi assemblage is about equally

quist)
S.

assemblage overlies the E. gcrthi assemblage

common

the

divided between southern Alaska and Europe, the P. singularis

as-

semblage shows additional resemblances to the Australo-Indonesian

area. However, the South American Pseudotoites species are closer
to Alaskan than to west Pacific species and have with the former
probably P. transatlanticus and P. argentinus in common; the Andean P. singularis is morphologically intermediate between the

and early Emileia

typical Australian forms

northwestern Euioj^ean middle to upper

('Tmileites")

sowerbyi Zone.

of the

The

as-

with Alaskan and, especially,

sociated sonniniids Jiaxe

affinities

European

unknown from

species but are

.S'.

the west Pacific margin.

WEST PACIFIC MARGIN

The

S. sowerbyi Zone
Newmarracarra Limestone of Western Australia (Arkell
and Playford, 1954) parts of this assemblage have also been identified from the iMoluccas and New Guinea, but no good evidence
is

from

only well-known larger assemblage of the

tiie

exists for the jjresence of the

zone in eastern Asia.

The

Australian

Alaskan ammonites,

assemblage has in

common

Pt.

Westermann

II:

with Ahiska

three

[Sonninia, Witchellia, Pseudotoites, Docidoceros

genera

four

to

(?)

29

but only one

or possibly two subgenera [the sparse Sonninia (Eitlioploceras) and

?

Docidocera.s (Trilobiticeras)], and no

dominated by the

typically

sjjecies.

The assemblage

European Fontannesia

is

clarkei

[F.

which appears to be absent along the eastern

Pacific margin and by several species of the typically Pacific Pseudotoites, including the microconchiate P. (Latotoites), some of which
are strongly reminiscent of evolute Emileia s.s. and E. (Otoites)
and others of Docidoceras, all of the northwestern European
This assemblage also inS. soiverbyi Zone (Westermann, 1964b)
cludes sparse "coronate cadicones" which were originally identified
with the northwestern American stephanoceratid Zemistephanus
(Arkell, in op. cit.) but are in fact closely related to the Andean
(Crick)

witii syns.]

Pseudotites
as a

(?)

(n.

subgen.)

sphaeroceroides (Tornquist)

as well

few involute Witchellia.

It

appears, therefore, that the relationship of the

S.

sowerbyi

Zone Ammonitina assemblages between Australo-Indonesia and

Alaska is not so close as previously supposed and that it can be
explained satisfactorily by indirect migratory connections
the southeastern Pacific

i.e.

on the one hand and Europe on the

via

other.

EUROPE
All genera of the Alaskan

S.

soiverbyi Zone, except for the

known from Europe, although Pseudolioceras and apparently also the poorly known Asthenoceras are there
restricted to older beds. Of the 12 Ammonitina subgenera (incl.
nominate subgen.) known from Alaska, eight were originally desparse Pseudotoites, are

scribed from northwestern

Europe [Eudmetoceras

s.s.,

E.

(Euap-

tetoceras), Planarnmatoceras (Psendammatoceras), Sonninia (Enhoploceras), Pelekodites s.s., P. (Spathulites), Docidoceras s.s., D. (Trilobiticeras)'],

and

of the species at least

two or three are in

with Europe [Eudmetoceras klimakotnphabtm (Vacek)

(Buckman) ]
tens (Buckman)
? Pelekodites pelekus
,

common

E. amplec-

while five

European species [i.e. to Asthenoceras nannodes (Buckman) Eudmetoceras eudmetum Buckman, Planammatoceras benneri (Hoffman) Docidoceras longalvum (Vacek), Hebetoxyites hebes Buckman]. Of the other four
others are very closely affiliated with
,

BlLLETIN 255

30

Alaskan .sul),<;cncia two have Ijcen identiticd ^\ith certainty only

from Wide Bay [Docidoccras (Pseiidocidoceras) and Bradfordia}
(Pracoppclia)]; one [Sonniuia (Alaskoceras)]
eastern Alaska

and

possii^ly Alberta,

occms also in southand only one [Psciidotoites s.s.]

widespread aroinid the Pacific.

The affinity to the northwestern Emopean assemblages is close
indeed; not only is the e(]uivalence of the E. mnplecteyis and Pseiidois

cidoceras zonides with

serious doubt, but

This

is

the

some

S.

sowerbyi

(Standard)

infra-zonal correlation

is

Zone beyond
also suggested.

particularly apjjarent in the acme-zone succession, from be-

(Buckman) Sonninia (Eiihoploand Witchellia suliwroides, n. sp.; the lower

two acme-zones can be correlated with the /.. discitcs Subzone and
the upper with the S. trigoiialis and W. hiexjiitsciihi Sul)zones or
W. laeviit.sc iihi Sub/one. However, it appears that even in northwestern Eiuope tlie upper two subzones are distinct only in a few
low, of Eiidinetoceras amplectcns

ceras) bifurcata, n.

areas.

sji.,

This infra-zonal correlation

cause the Alaskan vertical ranges for

far

particularly

is

sjjecies,

exceed those in Europe (Text-fig.

8)

surprising be-

subgenera, and genera

Fortunately, as generally

recognized, these range differences pertain to disappearance

more than

to first

much

appearance.

CONCLUSIONS

The
in

S.

sowerbyi

(Standard)

North America; where

Zone

is

most commonly missing

present, the zone

is

usually poorly

fos-

two known exceptions of Wide Bay and eastcentral Oregon. The ammonite fauna of the Kialagvik Formation is in all taxonomic categories related most closely to Europe
and secondly only to South America. The affinities to AustraloIndonesia are less close and could be accoiuited for indirectly by
migration via Europe and South America. All genera are also
known from other continents and only Pseiidotoites, rare in South
Alaska, is restricted to tlie Pacific; 25% of the subgenera and
50-80% of the species appear to be restricted to North America or
siliferous witii the

even to southern Alaska.

Approximately

50%

of the

Ammonitina genera and subgenera

range higher in southern Alaska than in Europe, and the same is

even true for several species. Nevertheless, the Bajocian (Standard)

Alaskan ammonites,

zones of

S.

soxocrhyi, O. saiizei,

identified with confidence

Zone somewhat

and

Pt. II:

Westermann

and

hutnphricsiaiiiini

dispersion

of

can be

a subdivision of the S. soiverbyi

siniihir to that in

Trans-Pacific

.V.

31

Einope is
ammonite

suggested.
species

chning

the

Bajocian as assumed by Arkell (1956, pp. 597-601) is not a necessary

conclusion from the study of distriijutions, at least not for the

[Zemistcphanus is known only from Alaska and

Columbia; Pscvdotoites is rare in North America and
there restricted to southern Alaska] and migration from AustraloIndonesia could have occurred along the continental margin or in
narrow marine channels of the Gondwana continent which at this
time was probably beginning to break apart (compare Irwing,
1964, figs. 10, 17; Hamilton, 1964, fig. 8).

North

Pacific

British

DIVERSITY AND PALEOLATITUDE

first ammonite monograph by Imlay

Based on the

on

several faunal listings, Arkell

(1956, p. 617)

(1953)

and

suggested, that be-

cause of high diversity the assemblages of the southern Alaska Jurasoriginated at a lower latitude than their present position

sic

62N) High diversity lias subsequently been confirmed

Aalenian and other Bajocian ammonite assemblages
(Imlay, 1961, 1962, 1964; Westermann 1964a) and is here again
demonstrated. The distribution tables of these monographs show
that single ammonite assemblages contain commonly between 10
and 20 reasonably abundant species of 6 to 12 genera and three
to six families. Such diversity indicates at least a temperate climate
for the southern Alaska Jurassic seas; cold climate as assumed by
Bain (1963, fig. 8) can certainly be refuted. Actual diversity counts
(57

also for the

are not attempted here because they are not available from other

When compared properly with assemblages

equivalent in horizontal and vertical extent, relative abundances,

Jurassic assemblages.

sample size and taxonomic "splitting," the southern Alaskan

Lower Bajocian ammonite assemblages are closely comparable in
diversity to their classical European equivalents. However, other
larger invertebrates are scarce at Wide Bay except for abundant
thin-shelled Inoceramiis

which might have been pseudoplanktonic.

Typically benthonic bivalves, gastropods, and brachiopods, as well

extremely rare, w4iile plant remains, particularly

fragmented wooden trunk, are common at several levels.
as belemnites, are

Bulletin 255

32

Only a few paleomagnetic data are available from North

America for the position of the Jurassic North Pole and all come
from tiie southwestern United States; although the plots are widely

mean

scattered even for data from single formations, a

pole posi-

southward for 8- 12 meridians is

indicated (C^oUinson and Runcorn, 1960)
This would shift the
latitude of the southern Alaskan Jurassic from the present 57 62N.
to aijout 50 N., the present latitudes of northern Oregon and
Vancou\er Island or of southern England and Central Europe.
European latitudes were, however, also appreciably lower than at
tion lelatively displacing Alaska

present. Polar shift, possibly comi)ined with

would

warm

east Pacific cur-

account for temperate Jurassic seas

in the area of soutlicrn Alaska. Nevertheless, the prevalence of

rents as at present,

fully

terrigenous sediments in contrast to the abimdant limestones in the

European Jurassic may suggest

relatively higher latitude for south-

ern Alaska. Furthermore, low^ thermal gradients of the Bajocian

Pacific

Ocean and Atlantic "Ocean"

are indicated by the extraordi-

nary \\ide distributions of several ammonite species which include

Einope, the southern Andes and southern Alaska. Such reduced

gradient could probably be responsible for the recorded diversity

without change

in latitude.

FOSSIL LOCALITIES
L'nitcd States

(12405 collected by
Daviess, 1944;

W.

localities.

R. Smith, 1924; 19028 and 19801 by

S.

N.

19862-19922 by Lewis B. Kellum, 1945; 21251 and

12252 by Ralph
Field Xo.

Geological Survey (USGS) Mesozoic

W. Imlay and Don

J.

Miller, 1948)

Catalogue Xfo.

F-27

12405

SE. side of

Wide

Bay, "1.1/2 mi. from cape

near islands."
44

A Km

F21

19028

[\on F21 of Short Creek section] SE.

Wide Bay, bluff 1.56 km E. of

side of

Preston Creek, about 11.0

of

Hartman

Isl.

km

S.

56

W.

Massive sandstone at base

of section.

44

A Km

72

19801

SE. side of

Wide

Bay, sea

E. of Preston Creek, 9.7

Hartman

Isl.

cliff c.

km

S.

55

1.6

W.

km
of

Base of shaly unit below an

Alaskan ammonites,

Wkstkrmann

Pt. II:

unconformity,

A Km

F55

19862

limy

witli

concre-

Kialagvik Fm.

tions,

45

shale

33

SE. side of

Wide

Bay, sea

and

of Preslon Creek

Harlman

Isl.,

9.0

cliffs 2.5

km

S.

km

E.

W.

of

52.5

elevation 7 m. Shale with

sandstone stringers above massive sand-

below top of Kialagvik Fm.

Bay, sea cliff about 2.1
of Preston Creek and 9.1 km S.

stone, 43

45

A Km

F56

19863

Wide

SE. side of

km

E.

W.

53.5

uses

of

Hartman

(190

Isl.

SW.

of

19862 "where the shale dips into

the sea")

Float from about 45

below

top of Kialagvik Fm., shales with stringers

of sandstone.

45

A Km

F62

19869

SE. side of

Wide

Bay, gully near shore

about 2.3 km E. of Preston Creek.

Interbedded dark grey blocky concretionary shale and thin stringers of grey wealine,

thering limestone,

145

below top of

Kialagvik Fm.

45

A Km

F63

19870

Same

USGS

locality as

talus at base of sea

A Km

F112

19922

from

Probably from

below top of Kialagvik Fm.

West end of Wide Bay, upper Kialagvik Creek at SW. foot of Lone
Hill. About 14 m below top of Kialagvik

about 36
45

19862, but

cliff.

km SW.

of

Fm.
45 Al 103

21251

SE. side of

W.

46.5

of

Wide Bay, sea cliff 8.5 km

Hartman Island. (2.85 km

of Preston Creek).

About 150

S.

E.

below

top of Kialagvik Fm.

48 Al 104

21252

Same
stone

USGS

locality as

stone,

19862.

Gray

silt-

above a massive sand("probably a little lower than lot

about 8

21251").
California

109

Academy

of Science

29011

(CAS)

localities

SE. shore of

Wide

Tatcliff Island.

Bay, 5.5

(1.83

km

km SW.

of

E. of Preston

Bulletin 255

34

Creek)

Black shale "just above contact

of Kialagvik witli Shelikof

C.E.L. No. 68

29014

SE. side of

CAS

2901

Wide

Fm."

Bay, about 0.3

km

(1.83

Dark grey sandstone with

Kialagvik Fm.
Shell Oil

Company

km W.

of

E. of Preston Creek)

basaltic dike,

(Shell) localities

546

SE. shore of

tion."

Fm.

Wide

Bay, about 2.2

km

E.

Creek, in "Mt. Frances Sec-

of Preston

About

15

(for localities

below top of Kialagvik

mentioned in text only,

see Text-fig. 2)

G. E. G. WcsternKniii (\VA) localities

Field No.

W\

SE. side of

Wide

Bay,

.7

km

E. of Preston Creek,

above

end of high bluff, elevation 60 to 70 m. Subgreywacke and mudstone, some silty shale, 50-65 m stratieast

above E. amplectens zonule of

grapliically

Kialagvik Fm.

^VA

Wide

SE. side of

bluff; top of

(Plate 2)

Bay, bluff 1.2

elevation c.5 m. 4

km

E. of

above base of 12

Preston Creek,

interbedded

and greywacke below massive subgreyaynplectens zonule; Kialagvik Fm. (Plate 3,

shale, siltstone

wacke of E.
above, and 4)

\\\

Wide

SE. side of

Bay, subgreywacke bluff 1.6

km

E. of

Preston Creek. Float probably from base of bluff; E.

amplectens zonule, Kialagvik Fm.

WA

SE. side of

W.

Wide

Bay, bluff 1.65

km

[E.

WA

E. of Preston Creek,

Mudstone, greybelow top of l)luff

amplectens zonide] Kialagvik Fm. (Plate 1).

end of subgreywacke
wacke and silty shale about 15
50

of

SE. shore of

more

Wide

Bay, 1.75

bluff.

km

E. of Preston Creek,

westerly of the two large tectonic blocks of massive

subgreywacke east of main bluffs. From talus of massive subgreywacke and some silty shale; E. aynplectens
zonule, Kialagvik

WA

SE. shore of

Fm.

Wide

Bay,

1.8

km

E.

of Preston

Creek,

Alaskan ammonites,

Westermann

Pt. II:

35

more easterly of the two large tectonic blocks of massive

subgreywacke east of the main bluff with basaltic dike.
E. amplectens zonule: about 8 m massive subgreywacke,
highly fossiliferous especially at 2.2 to 0.5
especially

meters of

cideras zonule missing)

WA

10

from top;

mudstone and siltstone, fossiliferous

at top and near base; overlain by several
crumbly shale indicating fault zone (Pseudo-

underlain by 3

(Plate

3,

below)

SE. shore of

Wide
km W.

Bay, 2.45 to 2.55

Creek (4.1
both sides of

of cape at

little

stream. 21

km

E. of Preston

end of Bay),

sea cliff at

dark grey shale with

abundant calcareous black concretions, often in bedding

planes and forming a thick "bed" at 10.5 m above base
of section; highly fossiliferous throughout; Pseiidocido-

Fm.

ceras zonule, Kialagvik

WA

1 1

Wide

SE shore

of

WA

W.

30

of

10)

Bay, 2.5

(Plate 5)

km

E. of Preston

Creek (50

Kialagvik Fm., from base:

dark grey shales with abundant calcareous con-

cretions, especially in lower part,

tionary "bed"

(same

as

WA

and thick concre10.5

10,

m) near

base;

Pseudocidoceras zonule, highly fossiliferous in lower

20 m.

15

and arenaceous shale with few concretions,

silty

poorly fossiliferous.

30

shale with interbedded lenticular sandstone

silty

beds, unfossiliferous.

15

interbedded subgreywacke, sandstone and

silty

shale, unfossiliferous.

W^A

12

SE shore

of

Wide

Bay, 2.15

km

E. of Preston Creek,

elevation about 100 m, top of section beside gully ad-

WA

jacent to

(overlying c.65

13 w.

thick-bedded subgreywacke

unfossiliferous shales

-)-

the W^a

15

section)

WA

13

(only 15

30

and

WA

13w

Wide Bay, 2.48 km E. of Preston Creek

W. of WA
but separated by fault) About

SE. shore of

1 1

some concretions,
from base; Kialagvik Fm.

shale with

16

SE. shore of

Wide

Bay. 2.45

km

fossiliferous at 3

E. of Preston

Creek

36

Bulletin 255

(15

\VA

\V. of

13).

About 20

shale with single

concretions, tossiliferous near base; Kialagvik

^\'A 11

Wide

SE. shore of

Bay, 1.95

low-tide exposure about 30

wacke beds; Kialagvik Em.
\\'.\

15

Wide

SE. shore of

20-30

and

km

Bay, 2.15

from sea

km

Fm.

E. of Preston Creek,
cliff.

Subgrey-

E. of Preston Creek.

some concretions,

shales with

faulted; at 3 to 16

partly slumped
from base moderately fossili-

ferous except for abiuidant Itioccraiiius: Kialagvik

Fm.

SYSTEMATIC DESCRIPTION
REPOSITORY OF TYPES
Specimens collected by the U.S. Geological Survey, which betypes by description or figuring, are deposited in the U.S.
National Museum (USNM) Washington, D.C.; specimens collected
by the author are deposited in the Department of Geology at
'McMaster University (McM.)
Hamilton, Ontario, under the

come

catalogue numbers J
Some fossils are now being returned from
the U.S. Geological Survey to the California Academy of Sciences
.

(CAS) in San Francisco, California.

MEASUREMENTS OF AMMONOIDS
The measurements taken and their abbreviations
as in

the

first

part of this

are the

monograph (Westermann,

same

1964a,

p.

PI. 8, fig. 4; Text-fig.

11

357).

PHYLLOCERATIDAE Zittel, 1884

Subfamily PHYLLOCERATINAE Zittel, 1884
Genus PHYLLOCERAS Suess, 1865
Subgenus ZETOCERAS Kovacs, 1939
Family

Phylloceras (Zetoceras)

Material.

cf. P.

single

1018).

zetes (d'Orbigny), 1850

fragmentary phragmocone with test remains came from the Rscudocidoceras zonule, S. sowerbyi Zone, of

WA

10 at 15

The

(J

specimen

closely
resembles the phragmocone
crushed innbilicus recently described by Imlay (1964, p. B31,
figs.

under P/iyllocera.s rf. P. kunthi Neumayr from the O.

Zone of the Tuxedni Formation in the Talkeetna Mountains,

3,4)

sauzei

with
pi. 2,

Alaskan ammonites,

Pt. II:

Westermann

37

11

Text-fig.
Cross-section
11.
(d'Orb.), phragmocone fragment,

(McM

J 1018)

of
loc.

Phylloccras

WA

IS

in

(Zftoccras)
the

cf.

zctcs

P.

Psrudocidoccras zonule

1-

Alaska. Imlay's specimen also has similar irregularly tetraphyllic

P. zetes has recently been reinvestigated
by Geczy (1967, p. 9) who concluded that in Hungary Zetoceras
cannot be separated from PJiylloceras. Zetoceras is, therefore, placed

saddles (from plastotype)

as a

subgenus in Phylloceras.

The compressed whorl

flanks,

section with weakly convex, converging

narrowly rounded venter, umbilical margin and overhang-

ing umbilical wall, as well as the rectiradiate lirae and the septal
suture match Geczy's specimen from the Pliensbachian of Hinigary.

From

Prince Patric Island in the Canadian Arctic, Frebold

p.

pis.

5,

7,

8,

9,

fig.

2)

described the dubious

which was leased on a single

mold and an undescribed fragment, the only

(Zetoceras) thorsteinssoni

internal

new

ence from P. zetes admittedly being

its

(1961,

species P.

totally septate

certain differ-

younger age; possible

distinc-

and
came from the Arkelloceras beds of the Wilkie Point Formation which probably correspond to the O. saiizei Zone, a date based on the occurrence
of Arkelloceras in the O. sauzei Zone of the Alberta foothills
(Westermann, 1964b) and Wide Bay (Imlay, 1964, p. B53, pi. 28,
tions of the

"new

missing radial

figs. 7-9;

species" include a shallow umlDilical slope

folds.

The

Arctic specimens

generic identity here verified)

38

Jil'I.LETIN

255

fracture

Text-fig. 12.

phragmocone,

Cross-section

loc.

WA

of Partschiceras

8 in the E.

Genus PARTSCHICERAS
Partschiceras ellipticum Westermann,
(?) 1964. Macrophylloccras

sp.

ellipticum, n. sp., incomplete

amplectens zonule

indet.

J 962)

1-

Fucini, 1923

n. sp.

A,

(McM

PI. 8, figs. 1,2;

Imlay

Text

(SE Alaska),

fig.

Pap. 418-B, p. B31, pi. 1, figs. 1-7.

Partschifcras garJanum (Vacek), 1886, subsp. ?, Westermann.
Bay I) Bull. Amer. Paleont., vol. 47, No. 216, p. 453, pi. 76.
(?) 1964. Partsrhirrras cf. P. gardanum (Vacek), 1886, Westermann,

12

Geol.

U.S.

Sur., Prof.

1964.

447,

pi.

74,

figs.

(Wide
id.,

p.

1-3.

Westermann, lOG-^a, p. 453, pi. 76; well-preserved

phragmocone, largely with test, septal suture and inner

//o/o/)'/?6'.

large

whorls exposed. Repository: Cal. Acad.

Sci.

(San Francisco)

12606.

Locus typinis.-Ca\. Acad. Sci. 29017 (field No. 99): "Cliff

exposure on the west shore of the south end of Wide Bay, inside
hook made I)y long sand spit."
Stralutu /}'/>/>;/;?/. Kialagvik Formation, upper E. hozvelli
Zone.
Derivatio notninis.

With

regard to the elliptical whorl

sec-

tion.

Diagnosis.

An

obvolute species of Partschiceras witli com-

pressed elliptical whorl section and simple

weak

costae.

Probable distribution and agc. E. hoivelli Zone to

riesianum Zone of southern Alaska (Wide Bay, Cook

Talkeetna Mountains)

S.

humph-

Inlet,

and

Alaskan ammonites,

Material from the

Pt. II:

sowerbyi Zone.

S.

Westermann

One

39

incomplete phrag-

WA 10 at 2.5 m (J 1013) one phragmocone fragment

of
WA 10 (f 1916) one incomplete phragmocone from
from scree
WA lower part (} 962) All internal molds with test remains,
mocone from

8,

E. amplectens

mation,

Wide

and lower Pseudocidoceras zonules of Kialagvik ForBay.

The

whorls are extremely involute to obvolute

and compressed-elliptical in section; the flanks are evenly convex
sloping to the occluded umbilicus and to the narrowly curved,
Description.

somewhat arched

The
50

mm

venter.

inner whorls bear slightly prosoradiate^

lirae.

At about

diameter, blunt and weak, rectiradiate plications arise on

the outer one-half to two-thirds of the whorl; subsequently, these

simple costae strengthen somewhat, often becoming slightly prosoradiate, and cross more or less straight over the venter where they

reach their greatest relative strength.

The

septal suture has a graded series of slender, essentially

diphyllic saddles.

Comparison. The holotype from the E. howelli Zone and the

specimens from the S. sowerbyi Zone of Wide Bay match the
" Macrophylloceras sp. indet. A" of Imlay (1964, p. B31) from the
O. sauzei and 5. humphriesianmn Zones of southeastern Alaska.
However, possibly because of imperfect preservation, the "vague

Cook Inlet
on the Wide Bay specimens.

flexuous undulations" on the innermost whorls of the

species (loc. cit.) cannot be seen

This species appears

statum (Imlay, 1953,
eastern Alaska which

closely related to Partschiceras grossico-

p. 74, pi. 25)

is

from the Callovian of south-

distinguished in the extremely dense sec-

ondary costation, especially on the venter where the primary costae become obsolete. Significantly, a specimen morphologically exactly intermediate between the Lower Bajocian and Callovian
Alaskan species has been described by Imlay (1962, p. A5) from
the early Upper Bajocian Magasphaeroceras rotimdum assemblage
of the Tuxedni Formation, Cook Inlet; while coiling and whorl
section are the same as in P. ellipticum and P. grossicostation, the
costation is intermediate consisting of continuous primaries anti
Correct spelling for "prorsiradiate" auct., according to Classics
Master University.

Dept.,

Mc-

40

Bulletin 255

relatively strong short secondaries singly intercalated

All other Partsc/iiccras species

(including the

on the venter.
synonym Macro-

phylloccras Spath,

1927) are distinguished by the subrectangular

subsquare whorl section and stronger costation or stronger costation. Some resemblance can be seen to P. gardaninn (Vacek, 1886,
to

from the partly condensed Aalenian to basal

S. Vigilio in the Alps which is distinguished
by the development of an umbilical edge.
The Alaska species agrees in the wiiorl sha}}e and coiling with
Phylloceras hctcrophyllum (J. Sowerby)
type species of Pliyllop. 70, pi. 6, figs. 1-3)

Bajocian beds of

Cap

ceros.

Measurements.
holotype

(phragm.).

Dmm

W%

H%

Alaskan ammonites,

Pt. II:

Westermann

11

Material. One internal mold of a damaged j)hragmocone and

one incomplete body chamber from USGS 21252; one well-preserved
incomplete phragmocone with exposed inner whorls and one fragment, from USGS 19862 (USNM 160921); one incomplete body
chamber with shell remains from scree of
5 (J 1012)
(?) one
crushed fragment from
10 at 22 m (J 1008) one internal mold
of a phragmocone fragment with incomplete body chamber, ? one
juvenile or nucleus from scree of
13 w (} 1011). From tlie
Pseudocidoceras and E. aniplectens zonides of the S. soiverbyi Zone,

WA

WA

WA

Kialagvik Formation,
Description.

Wide

Bay.

The

whorls are strongly involute to obvolute

with moderately compressed elliptical, often somewhat ovate section

end

becoming more or less strongly subrectangular towards the

phragmocone. There are five to seven sigmoid constric-

of the

tions per whorl, distally only slightly projecting,

and usually crossing the venter with

radiate

sometimes

recti-

slight convexity or

The internal mold is almost smooth except for

some barely visible, somewhat falcoid plications on the body chamber which become irregidar and obscure on the middle and inner
very obtuse arch.

flank.

The

septal suture has a

graded sequence of slender diphyllic

saddles.
Disciissio7i.

There

sum Imlay from

is

H.
humphriesiannm, and

perfect resemblance with

the O. saitzei,

S.

costisparS.

suhjur-

Zones of the Cook Inlet region, southeastern Alaska, extending the range of this species throughout most of the Bajocian

catinti

(s.s.).

There can be

little

doubt that the early Upper Bajocian

"Calliphylloceras sp." of Imlay (1962, p. A5,

cal

least in part restricted to the internal

fig.

pi.

1,

fig. 8)

is

identi-

with H. costisparsiini in which the constrictions are also at

15)

mold

(cf.

Imlay, 1964,

pi.

indicating the insignificance of this feature for the

1,

dis-

tinction of Calliphylloceras from Holcophylloceras.

H. ultramontanum (Zittel) from the subWide Bay (Westermann, 1964a, p. 448, pi.
74, figs. 4-7) is distinguished only by the falcoid and strongly projected constrictions which on the externside are strongly convex
Holcophylloceras

jacent E. hoivelli

Zone

cf.

of

or arched comprising almost a right angle.

H. costisparsnrn is morphologically and stratigraphically intermediate between H. ultramontanum (Zittel, 1869, p. 66, pi. 1, figs.

Bulletin 255

42

4-6)

ranging throughout the Aalenian

(-J-? S.

soxverbyi Zone)

H. mediterraneum (Neumayr,

1871, p. 340, pi. 17,

zignodianutn d'Orbigny, 1848,

pi.

per Bajocian to the Tithonian

Tnontanum

is

figs. 2-5)

and

[= H.

182] which ranges from the Up(Wendt, 1963, p. 115). H. ultra-

distinguished by fewer constrictions of falcoid shape,

or bearing lateral "tongues,"

and by more compressed whorls; H.

and stronger costation. In

tncditerraneuni has a larger umbilicus

A. costisparsum the constrictions are essentially restricted to the

internal

diary

is

figs. 6-8)

mokl

resembling CalUphylloceras. Another interme-

tlius

the dubious
,

H. deslongchainpsi

known only

(Brasil,

in a few inner whorls

1895, p. 29, pi.

from the

S.

1,

humphrie-

sianum Zone of Sully in Normandy, which, reversely appears to resemble H. ultrnrnoritanuni in whorl shape and coiling, but H.
mediteiTaneiiin in tlie constrictions. The taxonomic significance
of the shallow linguate depressions projecting mid-laterally from
the constrictions is doubtful since they have been observed seemingly irregularly in a

number

of species (cf. d'Orbigny, 1848, pi.

182, fig. 3).

Measurements.

Dmm

USNM

160921

(phragm.)

(C.70)

W%

H%

U%

Alaskan ammoxNites,

Pt.

II:

Westermann

43

Text-fig. 14.
Cross-section of Lytoceras sp. aff. L. eudesianum (d'Orb.), incomplete phragmocone, loc.
10 in the
Pseudocidoceras zonule (McM J 1005)

WA

The

shell

1.

planulate with advolute circular whorls.

is

The

in-

mold is usually smooth, except for markings of "rhythmic"

growth on a single small phragmocone (J 1010) The test bears
rectiradiate sometimes slightly flexed lirae or riblets of somewhat
ternal

uneven strength, every third or fourth of which is finely crinkled

(festooned) at least on the outer whorls. The septal suture has
relatively simple and regular saddle endings.
This species matches D'Orbigny's figure of the holotype (1846,
pi.

128) except for the missing lamellar flares. Because of the

finer crenulation,

it is

unlikely that the intermittent and

raised crinkled lirae or riblets are the

L. neumayri Pugin (1964, p. 34,

smooth lirae or
Pugin (1964, p.
pi.

1,

fig.

1)

is

remnants of dissolved

pi. 2, figs. 2-3)

is

much

somewhat
flares.

distinguished by

and the absence of flares. L. espinazitum

eudesianum" of Gottsche, 1878, p. 8,
distinguished by regular constrictions. However,

riblets

32; for "L.

Bulletin 255

44

Imlay

(1961. p. B32)

who

recently described but did not figure

probable L. rudcsiamirn from the O. sanzci Zone of southeastern Alaska, mentioned "weak constrictions." Lytoceras eiidesianutn has previously been known to range from the S. humphriesithe

first

amnij Zone to the Lower Callovian of Europe only (Pugin, 1964,

p. 32;

Sturani, 1964b, p. 12; VVendt, 1964, p. 116). AVhile

Lower

Jurassic Lytoceras species persist into the Aalenian where they be-

come increasingly rare, the genus

Lower Bajocian S. sowerbyi and O.
13;

p.

Vacek, 1886,
the

S.

sauzei Zones. Besides the rare

mentioned above, there is L. siibfrancisci Sturani

synonym "L. vaccki" Geczy, 1964 z=: L. francisci

L. espinazittiw

(1964b,

almost luiknown from the

is

pi. 2,

non Oppel) with

range from the Aalenian to

Innnphric.siauiun Zone, which

is

distinguished by the com-

and said to be intermediate between the Toarcian L. francisci and L. eudesiauum. S.

pressed elliptical whorls

(and spiralic

lirae)

siibfrancisci has intermittent crinkled riblets or lirae

whorls and

lirae

witliout flares.
intei

The Alaskan

mediate between

(d'Orb.)

on the inner

with intermittent blimt riblets on the outer whorls,

/..

species

is,

therefore, morphologically

subfraticisci Sturani

and

L.

eudcsianum

Measurements.

Dmm
J1005 (phragm.)

JIOIO (phragm.)

94

W%

H%

U%

Alaskan ammonites,

Westermann

Pt. II:

Text-fig.

15a-b.

45

of

Cross-sections

Hcbetoxyitrs sp. aff. H. hebes Buckman,

microconchs from the E. amplcctens zonule; X 1- a- Complete specimen with lappet, loc.

WA

(McM

plete specimen, loc.

and

tiire,

5.

(McM

a few poorly preserved fragments including

imprint with lappet, from the upper part of

WA

J 1043). b.

WA

WA

All from the E. atnplectens zonule of the

Kialagvik Formation,

Wide

Tlie

S.

Incom-

J 1041).

body chamber
and scree of

sowerbyi Zone,

Bay.

oxycone with involute compressed whorls, converging outer flanks and narrowly
rounded venter. Tiie inner flanks are parallel up to the sharp umbut markedly
bilical margin on the complete specimen (J 1043)
Description.

phragmocone

is

typically

convex beside the raised umbilical edge

1041), causing a

sliglit rise

at the other

specimen

(J

of the flanks at about two-fifths whorl

The phragmocone measures only 20-30 mm in diameter.

The body chamber, only about three-fifths whorls long, becomes
height.

inflated

and assumes

"elliptical" coiling,

egiessing almost radially

and

with the umbilical seam

finally returning to

spiral

coiling;

whorl height is reduced while the venter becomes more broadly

rounded. The aperture has a slightly projected broad ventral lip
and prominent mid-lateral lappets with thickened test. Both preserved lappets (left lappet of J 1043 and right imprint from same)
are incomplete consisting of narrow tongue-like projections. The
complete shell diameter varies between approximately 30 and 45

mm.
The

costation

is

obsolete

on the complete specimen

(J

1043)

except for faint rursiradiate plication visible under oblique illumi-

The somewhat
on the whole ultimate

nation on the outer flanks of the body chamber.

smaller incomplete specimen

(J

1041) bears,

whorl somewhat falcoid rursiradiate blunt costae or plications

Bulletin 255

46

which become obsolete on the mid-lateral spiral rise. The inner

convex part of tlie flank is smootli. Tlie plications die out gradually
beside

tlie

The

venter.

septal suture

ing the small

L/U

si/e.

is

is

only moderately complex, even consider-

well developed

saddles are of similar

graded

series

and

as

deep

as L, the

E/L and

these imibilical elements

si/.e;

form a

with rectiradiate "satldle line".

Discussion. While this is the first description of Hebetoxyites

from the Americas, supposed H. cf. hcbcs and H. cf. clypeus Buckman have i)een recorded by Lupher (1941) from the upper S.
sowerbyi or O. sauzci Zone of east-central Oregon.

The whoil

shape and coiling of the phragmocone and the

good evidence that

septal suture are

affiliated witli Hebetoxyites.

this

is

a strigoceratid closely

literature survey indicates that speci-

mens with complete body chambers

are exceedingly rare in the

The

specimens with incomplete

Strigoceratidae.

small

that

fact

body chambers which may include microconchs as well as immatures, have rarely been figured, may be due to the difficulty of
identifying small strigoceratids since

many

of the diagnostic features

are developed only in the adult macroconchiate

microconchiate body chamber.

phragmocone or

The body chamber

of these micro-

conchs, measuring barely one-half the size of figured macroconchs,

deviates from the described macroconchiate strigoceratids in
egression of the umbilical seam,

well-known and

common

close to the

"elliptical coiling".

This

the
is

microconchiate feature in other families.

Nevertheless, affinity of the

is

i.e.

Wide Bay microconchiate phragmocones

macroconchiate H. hebes Buckman. Consecjuently,

the author does not hesitate to place these specimens in Hebetoxyites

and, furthermore, sees no necessity to create a

sexual" subgenus for these probable male

There

also

is

less discoidal

some resemblance

Bradjordia;

cance suggesting

common

Measurements.

this

is

J1043
J1041

(phragm.)
(phragm.)

to the larger umbilicate

considered of phylogenetic

and

signifi-

ancestors.

Dmm

(apert.)

new "mono-

shells.

43

W%

H%

U%

Alaskan ammonites,

Pt. II:

Westermann

47

Family OPPELIIDAE Bonarelli, 1894

Subfamily

OPPELIINAE

Bradfordia and Oppelia.

[type species

The

Ammonites subradiatus

figured in Arkell, 1951,

et al., 1957, p.

p.

edge".

J.

de C. Sowerby; holotype

re-

51] was defined in the Treatise (Arkell,

275) as "compressed, involute, feebly keeled; with

and

distant falcoid primary ribbing

trolateral

Bonarelli, 1894

genus Oppelia Waagen, 1869

In

contrast,

close fine secondaries

Buckman,

Bradfordia

liomphala Buckman,

on ven-

1910

[type

was defined (loc. cit.) as

comprising the "group of Oppelia praeradiata H. Douville; moderately involute to involute, imkeeled, venter rounded, smooth, umbilical wall steep, umbilical edge sharp or raised; outer half of
whorl sides with fine, somewhat rursiradiate ribbing which is not
species B.

1910]

projected on shoulders. Sutures simple for the family".

The

alleged

occurrence of Bradfordia in Argentina was based on the erroneous

Eudmetoceras (Euaptetoceras) klimakomplialuyn

However, as most recently pointed out
by Geczy (1967, p. 224) Bradfordia is difficult to define and the
Treatise diagnosis as quoted above is unsatisfactory especially with
identification

of

moerickei (Jaworski, 1926)

regard to the costation. In contrast to the Treatise, the original

author (Buckman, 1910, p. 95) excluded O. praeradiata from the

genus because of the slightly projected and truncated costae. Furthermore, only a year prior to the Treatise, Arkell

(1956, p. 166)

placed in Bradfordia three species with strongly projected costae,

i.e.

'Oppelia' gracililobata

Vacek,

'O.'

sub plica tella Vacek,

and

'Harpoceras' bluynius de Gregorio [syn. O. platyomphala Vacek].

The

first

which

two species

edge of these species

cause

differ also in the differentiation of the costae

closely resemble those of

it is

The

is

Oppelia

s.

s.

The

raised umbilical

not a diagnostic feature for Bradfordia, be-

missing in the type species.

septal suture of Bradfordia

is

less

complex, with smaller

and fewer umbilical elements than in Oppelia. However, the dubious "Amblyoxyites" Buckman, 1922, which closely
resembles involute Bradfordia and is usually placed in tentative
synonymy, has the suture of Oppelia.
The Mediterranean 'O.' gracililobata^ 'O.' subplicatella and
even more so the Alaskan B-f oppeliiformis, n. sp., combine feaE, broader L,

48

Bulletin 255

tares ot Bradfordia wiili iliose ol Oppcliii; the sole distinction ot the

Wide Bay

species

from Oppclia

s.

s.

is

in

the narrowly

rounded

venter and probaljly also in the septal siitine which actually appears

intermediate between Bradfordia and Oppelia. According to the

numi)er of characters-in-conimon
ably

I)e

classified

this species

group should prob-

with Oppelia, but the application of the con-

ventional positive weighing of ventral featines woidd place

Bradfordia. This species group

is,

therefore, classified as

in

it

new

sub-

genus Pracoppclia and tentati\cly placed in Bradfordia.

BRADFORDIA Buckman, 1910

BRADFORDIA ? (PRAEOPPELIA) Westermann,
Genus

Subgenus

n.

subgen.

Type species. B.^ (P.) opprliiforniis, n. sp.

Subgenus diagnosis. Ldv^e, strongly compressed, involute,
venter rounded; with distant falcoid primaries and short dense
projected secondaries fading out ventrally or forming blunt chevrons; resembling Oppelia

s.

s.

except for the venter

(and the

less

developed septal suture)

The

subgenus probably includes 'O.' gracililobata

which the inner whorls are poorly known [the inner

Rer/iarks.

V^acek of

whorls figured by Vacek, 1886,

pi.

10, figs. 2, 3.

are not conspecific;

and "O." subplicatella Vacek, a possible

synonym. However, both appear to differ in tlie septal sutme with
less developed E lobes and E/L saddles. All are from the (lower)

fide Ceczy, 1967, p. 225]

S.

soxeerbyi Zone.

Bradfordia? (Praeoppeiia) oppeliiformis Westermann,

PI. 10, figs. 3-6; Pis.

Holotype.

PI. 10, fig. 6; PI. 20;

11,

n. sp.
12; Text-figs. 16a-b

well-preserved complete phrag-

mocone with incomplete crushed body chamber. Repository:

USNM

160922.

Locus
19863

[=

typicus.

45

AKm

on the southeastern

U.S.
F56

Geological

Survey

(float), coll. L. B.

side of

Wide

Mesozoic

locality

Kellum, 1945], sea

cliff

Bay.

Stratum typiciun. ShAes of the upper Kialagvik Formation,

probably in or subjacent to the basal Pseudocidoceras zonule, S.
soiverbyi

Zone;

with Eudmetoceras klimakomphahim

and Docidoceras camachoi, n. sp.

associated

discoidale, n. subsp.,

Alaskan ammonites,

Pt. II:

Westermann

49

Text-fig. 16 a-b
Bradford'ia? (Pracoppdia) opprliiformis, n. sp. a.
Cross-section of phragmocone, loc. USGS 21252 in the Pscudocidoceras zonule
(USNM 160923) X 1- b. External septal suture, loc.
WA-11, lot 37
13180)
X 1.5.

UW

(UW

Derivatio noiniuis.

Reflecting the close affinity to Oppelia

s.s.

Age.

(Lower)

Diagnosis.

.S'.

soioerbyi Zone.

species of D.f (Praeoppelia) with strongly

com-

pressed inner whorls, outer whorls compressed subelliptical with

inner flanks sloping gently towards the somewhat rounded not

markedly raised umbilical margin.
Material. The holotype and three somewhat crushed incomplete internal molds from USGS 19863; one partly crushed internal
mold from USGS 19862; one large phragmocone and one crushed
juvenile or inner whorl from CAS 29011; one well-preserved phragmocone with preserved nucleus and test from USGS 21252 (USNM
160923) one large phragmocone with well-preserved septal suture
from
WA-111, lot 37 (UW 13180); one slightly crushed
phragmocone, internal mold, from
15 at 3 m (J 1039)
one
incomplete phragmocone with preserved nucleus from scree of
5 (J 1040) All from the upper Kialagvik Formation, probably
;

UW

WA

WA

Bulletin 255

50

E.

atnplcctois

to

Pseudocidoceras zonules of the lower

basal

soiverbyi Zone, Kialagvik Formation,

Description.

The

Wide

mm

nucleus at 12-15

S.

Bay.

diameter consists of

smooth, moderately involute, strongly compressed

elliptical

whorls

with shallow umbilicus, the convex sides sloping gently to the umbilical seam. Subsecpiently, a narrow umbilical wall with somewhat

rounded margin develops, the whorls become involute, the inner

flanks flatten but continue to slope toward the umbilicus so that the
greatest

whorl width

costation develops.

is

retained

The immature

mid-laterally,

and the

typical

costation consists of blunt some-

what irregular falcate primaries which are mainly superficial but

become stronger on the outer one-third of the flank, and of short
blunt strongly prosocline secondaries on the rounded shoulders
whicli are singly and in pairs intercalated and cross weakly over
the venter.

The matme phragmocone

retains the strongly compressed in-

However, the umbilicus deepens

somewhat, due to the slightly raised rounded umbilical margin and
the conesponding development of a shallow depression on tlie
lowermost flanks; the whorls remain thickest at or just above the
middle of the flanks. The outer flanks converge gently, often markedly flattened, towards the evenly and narrowly roimded venter
which may widen somewhat towards the end of the phragmocone.
The mature costation consists of somewhat irregular, more or less
strongly concave (distally j^rojected) primaries on the outer half of
the flank, a few of them continuing faintly onto the inner flanks
forming weakly falcate to falcoid costae, and of nimierous prosoradiate short secondaries on the shoidders which become medially
volute wliorls with

convex

flanks.

obsolete or cross the venter convexly

cmved

or arcuate.

The

pri-

maries are sometimes obscurely bundled in pairs at about two-thirds

whorl

lieight,

and the secondaries are born about

as often

by anas-

At the end of
large phragmocones, both primaries and secondaries weaken if the
latter do not become obsolete.
The complete phragmocone measures approximately 120 mm
in diameter. The body chamber is known only from the holotype
where it is incomplete (one-half whorl) Tlie full diameter is estitomosis, bifurcation of primaries or intercalation.

Alaskan ammonites,

mated

at 160

mm.

Westermann

Pt. II:

Tlie umbilicus remains narrow and shallow witli

the umbilical wall rounding into the whorl sides;

broadly rounded.

somewhat

51

The body chamber

irregular

more or

is

externside

tlie

is

almost smooth except for

less rectiradiaie plications

on

the outer

flanks.

The
large

septal suture

and only

is

moderately complex (Text-fig. 166) E is

L is broad and trifid with
.

slightly sliorter than L;

L/U

wide and high as E/L;

radius with L; Uo is
about half as broad as L and the broad Uo/Ug saddle about twothirds as large as L/U2 (2d and 3d morph. lat.)
the other umbilical elements are much smaller with only the outer saddle fully
developed bearing first and second-order frilling. The suture is,
short thick stems; the

saddle

at least as

is

the umbilical lobes terminate at a

common

therefore, intermediate in character

Oppelia with respect to the

size of

between Bradfordia

E and

the

number

of

(s. s.) and

major um-

bilical elements.

Discussion.

This new

species

most

is

closely

related

to

'Oppelia' suhplicatella and 'O.' gracililobata, Vacek spp., of the

Mediterranean lower

sowerbyi Zone, but distinguished by

.S'.

more involute and compressed inner whorls,

the depressed

and

tlie

less

developed circum-umbilical area and, probably, the larger E lobe.

There are

also

some

striking resemblances to the internal

molds

(without the hollow-floored keel) of Eudmetoceras (Euaptetoceras)

amplectens (Buckman) and to a

lesser

degree also to E. (E.) klijna-

komphalum discoidale, n. subsp., both of which are associated.

The phylogenetic origin of the Oppeliidae arising with Bradfordia from Eudmetoceras
et al., 1957, p.

favours this hypothesis

special

interest

lias

repeatedly been suggested

275; Geczy, 1967, p. 225)

is

if

the

evolving lineages giving

PraeoppeUa

regional

is

(Arkell

and the Alaskan evidence

included in Bradfordia. Of

morphological "habitus" of the

rise to the

vexing problem of possible poly-

phyly or ecotypes: the Alaskan E. amplecferis, E. klimakom phalum

and B? oppeliiformis

are all distinguished

from

counterparts in the umbilicus which has a

less

their

Mediterranean

developed umbilical

margin or edge. The same "habitus' 'is also present in northwest

Europe and the southern Andes, so that E. amplectens appears here

Bulletin 255

52

closer to the Ojjpcliidac ancestry than E. kliinakoinpliahnn while

this

is

reversed in the Mediterranean

(Geczy, loc.

cit.)

Significantly, this very earlv oppeliid resembles the type species

of Oppclia

altogether

oppeliiformis

is

more

closely

than typical Bradfordia.

radiata only in the narrowly rounded venter

and the

less

complex

umbilical lobes.
Measuretnetits.

Dmm

holotvpe

(hodvch.)

B?

distinguished from the type species Oppelia sub-

W% H% U%

Alaskan ammonites,

Pt.

II:

Westermann

53

17

Pscudol'ioccras maclinlocki fastigatum, n. subsp.

Text-fig. 17 a-b.
X 1Holotype, cross-section through complete phragmocone. b. Almost complete
phragmocone, loc. CAS 29011 (CAS 13110).
;

a.

mold with some test remains, of complete phragmocone with

beginning of body chamber. Repository: USNM 160924.
ternal

Locks typicus. lJ.S. Geol. Survey Mesozoic locality 21252,

sea cliff on SE side of Wide Bay, Alaska Peninsula.
Stratum typicum. Upper Kialagvik Formation; probably
Pseudocidoceras zonule, 8 m above base.
Derivatio uorniuis. With regard to the whorl section which
tends to develop a fastigate venter.

Age.

S.

soiverbyi Zone, probably lower part.

Diagnosis.

subspecies of P.

ynacklintocki with

umbilical

ridge, strongly converging outer flanks, venter tending to be fas-

becoming obsolete on the inner flank.

incomplete and partially crushed specimens
from USGS 12405; one incomplete body chamber with crushed
phragmocone from USGS 19801; one phragmocone and several
tigate,

and

costae

Material.

Four

Bulletin 255

54

fragments from USGS 19862; holotype from USGS 21252; one almost complete phragmocone, only slightly damaged and one crushed
specimen from CAS 29011; one partially crushed inner whorl with
remains of deformed phragmocone from
10 at 0-1 m; three
moderately preserved inner whorls witli fragments of body chambers
from
15 at 3 m; one small specimen probably with part of
body chamber from
15 at 9 m; one large phragmocone from

WA

WA

WA

U\\'-\VA 111,

lot

37.

mains, fiom the lower

cidoceras zonide)

All are internal molds mostly with test rejxiri

sou'crbyi Zone, Kialagvik. Formation,

No

Rrniarks.

(including lower Pseudo-

of the shales

overlying the E. atnplectens zonule of the

Wide

S.

Bay.

complete description needs

to

be given here

because the suijspecies resembles closely P. macUntocki xchitcavesi

first part of this monogiaph. The moderand the adolescent development of a typical
imibilical margin are the same. However, the whorl section is
usually markedly distinct in the outer flanks whicli converge more
strongly and in the venter whicli is more narrowly roimded beneath

(White) described in the

ately involute coiling

the keel or frequently almost fastigate, at least in the internal

mold. Furthermore, the falcate ribs usually fade on the inner

flank so

the

tliat

umbilical ridge and the shallow lower-lateral

groove tend to become smooth on the

last

while the concave outer part of the ribs

is

phragmocone whorl,
broad but

at least as

blunter than in P. macUntocki xuhiteavesi. These minor differences

are

marked on

average

size

the last two or three whorls of the

phragmocone of

while the end of large phragmocones and the body

chamber may not be significantly different.

There appears to be some morphological overlap between the
two Wide Bay subspecies. However, the sample of this new subspecies is insufficient for statistical analysis and quantification of
the respective features is difficult. Nevertheless, some significant
morphological change has certainly taken place around the
Aalenian-Bajocian i^oundary indicating a new cinono-subspecies.

The

vertical range of P. 7naclintocki

(Haughton)

the three successive subspecies P. macUntocki

(Wliite)

and

P.

rii.

fa.stigatum

the Aalenian and into the

S.

is

now known

soxvcrbyi

Zone

.s.s.,

to

P.

m.

including
xvliiteavesi

extend throughout

of the Bajocian, a time

Alaskan ammonites,

VVestermann

Pt. 11:

luhitcavesi
maclintock't
Text-fig.
18.
Cross-section of PscuJolioceras
(White), lectotype, E. hoivelli Zone of "Wrangel Bay"; shown for comparison;

1.

interval equivalent to the range of the

whole genus

(in the

Toar-

cian) in Europe.

Because of
the

costae

converging outer flanks and the reduction of

this
subspecies resembles typical Toarcian

tiie

stems,

Pseudoliocerns, such as P. compnctile

(Young and Bird)

evolution

of P.

more

closely than

(Simpson)
its

maclintocki fastigaturn

and

P. lythense

ancestral subspecies.

may

therefore

The

be called

"retrogressive".

Measureme7its.

Dmm

holotype

(end phragrn.)

69

W%

H%

U%

P-S

23.5

48

16.5

18

Pseudolioceras costistriatum Westermann,

n. sp.
PI. 13, figs. 3-6; Text-figs. 19,

Holotype.

V\. 6, figs. 3a,b; Text-fig. 19;

Internal

mold

20

of in-

Bulletin 255

56

Text-fig.

Cross-section of Pscudolioceras costistriaium, n.

19.
incomplete body chamber with damaged phragmocone; X !

type,

complete body chamber

witli

holo-

sp.,

damaged phragmocone. Repository:

McM.

J 1056.
Locus typicus.

shore of

Wide

Stratum

Locality

WA

1 1

at

30 m, sea

cliff

at

S.E.

Bay, Alaskan Peninsula.

typicutn.

Upper

Pseudocidoceras

zonule

(19.5

above concretions bed) upper Kialagvik Formation.

Age. S. smverbyi Zone, Bajocian.
,

The holotype, two small immature (? or microspecimens and one fragment of a large phragmocone

Material.
conchiate)

from

WA

30 m; one small immature

11 at

s]X^cimen from

USGS

mains from the

S.

(?

19862. All are internal

or microconchiate)

molds with

test re-

sowerbyi Zone of the Kialagvik Formation,

Wide

Bay.
Diagnosis.

An

involute species of Pscudolioceras with strong

umbilical ridge, tabulale-unicarinate venter, striate costae and weak

intercalatories.

The whorls are involute and strongly comThe innermost whorls, up to approximately 15 mm

Description.
pressed.

diameter, are compressed-oval as in most other species. Subsequent-

Alaskan ammonites.

30

20

Pi. II:

Westermann

57

u
10 -

D (mm)

Scatter for relative umbilical width (U:D) of PseudolioText-fig. 20.

holotype) and P. madintocki ivhitcavesi
ccras costisiriatum, n. sp. (circles,
(White) (dots with visually drawn central line). Note the narrower umbilicus
of the new species.

ly,

H =

the vimbilical slope heightens

and the margin raises to form an

maximal whorl width lies

increasingly strong umbilical edge; the

approximately in the middle of the flattened

sides;

the venter be-

mm

diameter and
comes distinctly tabvilate-unicarinate at about 25
sometimes even somewhat bisulcate on the mature phragmocone;
the hollow-floored keel is not strong but well defined.
The ornament consists of blunt typically falcate principal
costae tending to become obsolete on the inner flank. The flattened
outer costae are striated superficially, often appearing as costellae
compounds, and sometimes weakly fasciculate on the inner whorls.
The falcate striae or costellae bundles are also present on the internal mold of the relatively large holotype, particularly on the
shoulders where they appear as faint intercalated secondaries, and

appear irregularly on the middle of the

also

The

sides.

single con-

sometimes intercalated.
septal suture is of moderate complexity and resembles

cave outer rib

is

that of other Pseudolioceras.

Comparison.

P.

costistriatum

somewhat resembles

P. maclin-

(Haughton) and particularly the subspecies P. m. whiteavesi

(White) from which it is distinguished by the narrower umbilicus,
the tabulate-unicarinate venter, and the falcate striae or costellae.

tocki

Bulletin 255

58

The

only

known

Fscudolioccras witli tai)ulate-iinicarinate venter

is

Buckman which differs strongly

This species is almost homeomorphous with

the early Toarcian P. subcoucnvinn

in coiling
tlie

from
is

and

costation.

peneconteinporary
tlie

Staiiffcriid

(Hypcrlioceras) which

is

absent

Americas; there can be no question, however, that this

a Pseudolioceras

and thus the

latest representative of the

Harpo-

ceratinae.

Measurements.

Dmm

Holotype (body ch.)

W%

H%

U%

Alaskan ammonites,

Pt.

II:

Westermann

59

been described and are apparently unknown from Emope. This is

not surprising if one considers the acute scarcity of G. nannodes
in tlie European L. ynurcfiisonae Zone which abounds in otlier ammonoids.
Dimorphism is well known in Toarcian Grammoceratinae, including Grannnoceras

(cf.

Buckman,

1890, pi. 28, figs. 20, 21; pi. 32,

However, their microconchs are so similar to the inner

whorls of the macroconchs that Buckman did not distinguish them
specifically regarding them as the juveniles. These microconchs with
lateral lappens are of similar size as 'G.' yiatuwdes which is, however, clearly distinct in the much finer or obsolete ornament and the
fig.

3)

laterally sulcate keel.

(Arkell,

1957, p.

Gratnmoceras"

is,

The

diagnosis of Asthenocerns in the Treatise

261; classified as subgenus)

simply as "dwarf

therefore, insufticient; Asthenoceras

is

not the

any described Grammoceras.

The Asthenoceras sample from Wide Bay comprising about 20
specimens comes from a single concretion of the Pseudocidoceras
zonule in the S. soiverbyi Zone. Another much larger sample was
collected by the author from a single lenticular bed in the uppermost Weberg (or basal Warm Springs) Formation near Supplee

microconch

to

in east-central Oregon; the association with evolute Witchellia

cf.

Sonninia (Eiihoploceras)

is

as in

Alaska and indicates

S.

and

soiver-

However, R. Imlay from the U.S. Geological Survey

recently showed the author rich collections of Asthenoceras from
the O. saiizei Zone of the Warm Springs Formation at approximately the same Oregon locality. The Oregon species varies gieatly
in size and suggests the development of dimorphism.
The largest Oregon specimens of 50-70 mm diameter resemble
typical Grammoceras ex gr. G. striatiihim (Sowerby) in the combyi Zone.

pressed oval whorl section and evolute coiling but are distinct in
the laterally sulcate high keel.
fied as

Grammoceras, such

A number

of species usually classi-

as G. fallaciosiim

(Bayle)

have high

hollow-floored keels but these have never been observed to be

laterally sulcate. The Wide Bay species consists of two size groups,
i.e.

mm
last

about 15

mm

large microconchs with small lappets

and 30-35

large macroconchs with somewhat modified ornament on the

whorl. These extremely small adult sizes compare with average

Bulletin 255

60

mm for microconch and 60-120

macroconchs (cf. Buckman, 1890, pis. 27-20). There is
little doubt that both samples represent single dimorphic species
which cannot reasonably be split taxonomically; the Wide Bay
form is here descril^ed as the single species Asthenoceras aff. A.
nonnodes. Because the macroconch of the Dorset species is unknown, a definite identification with that species is not yet posdiameters for Grammoccras of 25-10

mm

for

sible.

The author

has long pondered whetlier to elevate Astheno-

ceras to generic level or whether to retain

Grarmnoceras. Preference

is

now

subgenus of

as

it

given to the generic level for the

following reasons:
1.

2.
3.

The high, laterally sulcate keel is unique.

The ornament is much finer than in Grnrnmoceras.
The great difference in age from Grammoceras

abundance

(and

its

dimorphic species in North America)

Furthermore, the juvenile whorls are probably more widely
umbilicate and more compressed than in other grammoceratids.

The

as

simjjlified suture

generalized
Init

nature,

(here described for the

i.e.

conforming

not restricted to tlicm. There

strongly

even

piojected

less

known

and partly

with

close

is

the Graphoceratidae by Arkell (1957,

is

of a

resemblance in the dense,

fasciculate

Vacekia stephcusi

time)

first

some grammoceratids
growth

Buckman
p. L 262)

lines

(1899),
;

with the
placed in

this species

is

from

the L. rnurcfiisonae Zone of the Inferior Oolite and distinguished

in the different coiling, whorl

shape and keel.

appears that

It

Asthenoceras belongs to a "conservative stem" of Hildoceratidae,

whicli includes Arieticc.ras

and Graynnioceras.

It is

of interest that

in the Pliensbachian Arieticeratinae a similar almost

smooth and

compressed form was produced as an early offshot, i.e. Asaploceras.

Middle Jmassic Hildoceratidae al)ove the /.. opaliwdm Zone
are essentially restricted to the Pacific realm. Of the Harpoceratinae, Pseiidolioceras occurs in

northwestern North America and

probably in Japan (Westermann, 1964,

ceratinae, Asthenoceras, previously

p.

351)

known only

Of
in

the

Grammo-

two specimens

/.. ynurchisonae Zone of Dorset, is locally

Oregon and southern Alaska. It is probably significant

questionable from the

abundant

in

that both are long ranging genera,

i.e.

Pseudolioceras ranging from

Alaskan ammonites,

the upper Toarcian to the

L. miirchisonae

Zone

S.

Westermann

Pt. II:

61

smuerbyi Zone and probably from the

Zone. Tmetoceras reaches at

to the A. sauzei

Aalenian in southern Alaska, Oregon (?)

Andes (Westermann, 1967) and probably Thailand
(Komalarjn and Sato, 1964) Pleydcllia (s.L), which is probably the
latest wide-spread member of the Grammoceratinae in Europe
where it ranges into the lower L. opolinum Zone, has also been described from the lower Aalenian of a single locality in the Canadian
Arctic (Frebold, 1960, p. 23, pi. 12, figs. 1-4). However, the disputed "PleydelUa assemblage" (or 'P' puchensis group) of Mendoza
in the southern Andes (Burckhardt, 1903, pis. 1, 2) is tentatively
placed in advanced Hammatoceratinae or very early Sonniniidae.
least to the top of the

the southern

Asthenoceras

sp. aff.

A. nannodes (Buckman) 1890,

PL

c^

14, figs.

&

1-7,

Text-figs. 21a-c

YouY almost complete macroconchs, three comand two fragmental microconchs with lappets, and about
dozen incomplete, juvenile or fragmentary specimens; all from
Material.

plete

a single concretion of

WA

Wide

tion,

stage,

keeled

(5-6

Following

the

section changes
to fastigate

S.

1037, J

(J

1035)

in the

sowerbyi Zone, Kialagvik Forma-

Bay.

Description.
larval

11m

10 at

Pseiidocidoceras zonule of the

(at

the

first

coiling becomes

globose whorl, the probable

the whorl
from sub-circular to somewhat compressed oval
D) and to strongly compressed oval and
3

mniD)

widely umbilicate;

mm
.

Finally, at 10-12

mm

diameter, the adult size

phragmocone, the sides are somewhat flattened curving sharply to the umbilical seam and into the usually
rounded but sometimes narrowly tabulated externside which carof the microconchiate

ries a

strong blunt keel.

The

microconchiate {i) body chamber

long and terminates at only 15-16

is

about one-half whorl

mm diameter with lateral

lappets.

no sigphragmocone
grows
(2)
to about 22-25 mm diameter, becoming even more compressed
with narrowly convex externside and exceptionally high and narrow hollow-floored keel. The keel attains the typical laterally
Beside the slight egression of the umbilical seam, there

nificant modification.

The macroconchiate

is

Bulletin 255

62

Text-fig.

21

a-c.

Asthrnoceras sp.
man), compiled
V

/'N

Septal

on

section

cross

Silicate

tlie

of

WA

10 in the
of a single concretion, loc.
Psrudocidocrras zonule, magnified, a. At
D
10
D (McM J 1035). b. At 12
D, largest
(McM J 1036). c. At 22
available phragmocone diameter (McM
J 1035a),

mm

mm

chamber was

suture

A. nannodcs (Buckfrom three specimens

aff.

last

mm

phagmocone

wliorl.

The body

originally at least one-lialf whorl long terminating

mm

cHameter. Although not preserved, the aperture was

probably simple in accordance with the reduction in flexure of

at 32-36

the growtli lines at the end of the preserved Ijody chambers.

The ornamentation
ficial,

and

develojjs

sliglitly flexed,

of the inner wliorl

only after 5-7

mm

is

faint, largely super-

diameter.

It

consists

of

strongly projected, extremely fine striae represent-

ing growtli lines and often also of widely spaced similarly flexed

bundles of

striae

or

obscure

plications

which could represent

pooily develoj^ed costae.

On

growth lines and striae bundles condiamber where they become falcoid by increase
of mid-lateral convexity indicating the gradual development of lateral lappets. The few preserved short and narrow lappets, positioned at about two-fifths whorl lieight, are probably all more or
less incomplete. The internal mold of the microconch is either
entirely smooth or bears similar weak irregular striae as the surthe microconch, these

tinue onto the body

face of the shell.

ing

it

The

striae

sometimes continue onto the keel

giv-

a braided appearance.

The macroconch

develops densely spaced strongly projected

on the last whorl. The costae are usually restricted to the

outer flanks and some specimens bear a few blunt plications on
the inner flanks which arise from costae fasciculation. Falcoid
growtli lines are also common. Towards the end of the body
chamber, costae and giowtli lines become less flexed and somecostae

Alaskan ammonites,

Pt.

Westermann

II:

The most complete specimen

times straight.

(PI.

63

11, fig. 5a, h)

has

body chamber wiiich toward the

end bears almost rectiradiate and complete, widely spaced blunt
costae. These macroconchs are fully grown according to the apfive-eighths whorls of preserved

proximation of the

The

last septa.

suture

septal

considering the small

the large
("1st.

12

E/L

and 2d

mm

(Text-fig.

saddle

lateral

21)

mm

is

extremely simple, even

(3.5-4 mm H) only
and the L and U^ lobes
lobes") are frilled, at the first order only. At
size.

At 10

diameter

(externsaddle)

diameter, the suture

is

subammonitic.

still

mm

served at the diameter of 22

suture ob-

only slightly more complex,

is

frilling only of the E/L saddle. E and E/L are

U^ decreases ontogenetically in relative size. There are one
or two more small external umbilical elements of regularly decreasing size. The internal part of the suture is imknown.
Comparison. The microconch of the Wide Bay species resembles the "cotypes" of A. naunodes Buckman. They differ in the

with second-order
large,

smaller size (about 15

the

more

mm

vs.

about 25

regularly bundled costae

if

mm

D) and possibly

also in

developed.

Because the Dorset species is poorly known and represented

by the microconch only, no definite judgment as to the affinity
between the Dorset and Wide Bay forms is possible, and the latter
can only be tentatively referred to A. nannodes.

The Oregon form

retains the dense

is

much

ornament up

conch.

Measurements.

Wide Bay form and

body chamber of the macro-

larger than the

to the

Bulletin 255

64

been published, based on assemblages respectively from France,

Hungary, Bulgaria, and Turkey. The study of Maroccan hammatoceratids by Lelievre (1960) had been overlooked by me. Furthermore, a taxonomic revision of the Hammatoceratidae based
on ontogenetic studies of the septal sutures was carried out by
Schindewolf (1964). The following five new generic and subgeneric names have been given since 1961:
liave

Genus Pseudammatocrras Elmi,

signis Oppel,

1856

13;

1964, p.

type species: Amtnonitfs sub'in-

(Renz, 1925).

Subgenus EuJmrtoccras (RhoJaniccras) Elmi, 1964, p. 60; type species: Hammatocrras rhodarilcum Renz, 1925 (for "Hammatoccras subinsignc" Bayle,
7ion Oppel). Microconchs to EuJrriftoccras s.s.
Genus Sptnarnmatocttas Schindewolf, 1964, p. 340; type species: Hammato-

cnas pugnax

1886. Microconchs.

\'acek,

Subgenus Hammatocrras { PseuJaptctocrras) Geczy, 1966, p. 77; type species:

Harpoccras klirnakomphalum Vacek, 1886; lectotype designated Sapunov,
1964: Vacek's figs.

17,

17a.

Subgenus Hammatocrras (Cscrnyriccras) Geczy, 1966, p. 83; type species:

Hammatocrras vrrpillirrrnsr Roman and Beyer, 1923.

At

least

19

new species names were given and a number of

named by Elmi (1964) and Gcczy (1966) for

infra-specific taxa

keeled Middle Jurassic Hammatoceratinae [exclusive of Erycites,

Abbdsitcs,

and A. (Ambersitcs)]. Both authors considered Abba-

from Erycites, in contrast to the Treatise

267) but in conformity with this monograph (Westermann, 1964a, p. 404) However, Elmi and Lelievre
differed greatly from Geczy in the classification of the keeled hamsites generically distinct

(Arkell, et

al.,

1957, p.

While
Middle Jmassic hammatoceratids in
the single genus Harnmatoceras with two old and two new subgenera, Elmi and Lelievre retained almost all the genera of Buckman. Geczy (p. 31) also doubted the generic distinction of Erycitoides Westermann, 1964. This judgment was based only on my
preliminary abstract (Proc. 1st. Coll. Juras., 1964) and later withdrawn after study of the monographic description (oral communication at 2d Coll. Juras., 1967) More particularly, Geczy regarded
as synonyms of Hauirfuiloceras the "genera" Brcdyia Buckman,
1910, Eudynetoceras Buckman, 1920, Pacliammntoccras Buckman,
1921, Planammatoceras Buckman, 1922, Farannnatoceras Buckman,
1925, Pscudamtnatoceras Elmi, 1963, and the subgenus E. (Rhodaniceras) Elmi, 1963. [For Eryciloides and E. (Kialagvikites) Westermatoceratids, particularly with respect to taxonomic level.

Geczy included

all

keeled

Alaskan ammonites,

niami,

1964, see above].

therefore,

Geczy's

Westermann

II:

"Haniinatoceras

(1964)

65

s.

str.",

named keeled HammatoceraEuaptctoceras Buckman which was retained as sub-

included

tinae except for

Pt.

all

pieviously

genus only because of the supposedly asymmetrical lateral lobe

(L)
Geczy then jjroceeded to distinguish two new subgenera of
Hanimatoceras, again on sutural characters, i.e. H. (Csernyeiceras)
.

with large E and H.

(Pseudaptetoceras) with alleged relatively

simple suture and symmetrical

comparison with E. (Eiiapteto-

in

ceras).
I

have not arrived at any definite opinion regarding the

many of the early and intermediate HammatoceraTaxonomic levels somewhere midway between the ones above

classification of
tinae.

reviewed are strongly suggested, similar to the treatment in the

Treatise (Arkell, et

al.,

1957, p.

267), to the tentative classifica-

monograph and to the recent usage

Sapunov (1964), and Blaison, et al. (1966).

tion in the first part of this

by Bremer (1966),
Hanimatoceras

and defined

(s.s.?)

Hyatt

in the Treatise

(p.

perhaps reasonably diagnosed

is

267)

as

including the synonym

PacJiainmatoceras and as being confined to the upper Toarcian.

Pseudammatoceras Elmi comprising early Aalenian species grouped

around the type species P. subinsigue (Oppel), however, appears
close to Bredyia which was not even mentioned by Elmi. It is possible that Arkell, et al., in the Treatise meant to include these
forms in Bredyia because this is the only genus listed from the L.
opaliuum Zone. Bredyia and perhaps also the related Pseudammatoceras s.s. could be classified as subgenera of Hammatoceras, but
later species of Elmi's Pseiidaminatoceras are

dubious 'Pararnrnatoceras' and even

low)

to

much

closer to the

Planammatoceras

(see be-

Planammatoceras Buckman should be generically distinguished

from Hammatoceras, but it cannot be defined to nontuberculate
forms as in the Treatise (Arkell et al, 1957; p. L 267) This author
(Westermann, 1964, p. 411) and Elmi (1963, p. 89) included in
Planammatoceras the widely umbilicate and tuberculate "H. lorteti
Vacek, non Dumortier" (r= P. romani Elmi) There is a good possibility that P. teniii?isigne (Vacek) and P. planijorine Buckman
which were previously regarded synonymous with P. planinsigne
(Vacek) (loc. cit.) are but the compressed and finely ornate end.

Bulletin 255

66

members

of

sequence,

i.e.

an intergrading penecontemporaneous morphological

a single species, with the other widely umbilicate and
strongly ornate end-member i:)eing named P. romani Elmi. Such a
morphological se([iience would conform with the "Buckman law of
covariation" (Westermann, 1966)
However, the distinction of Plaunmynatoceras, flourishing in
the L. murcJusonae Zone, from typical Eudnictocerns of the L. coucava and lower S. soioerbyi Zones is ill-defined, and I am still hesitant to take a more definite stand regarding their taxonomic separation on the generic level than I did in the first part of this monograph (Westermann, 1964a, jx 412). Tlie genus Plajunnmntocerns
is,

therefore, tentatively retained.

Eudinetoccyas

(s.s.)

is

certainly not sufficiently characterized

by "un enrolement evolute et par leur ombilic plat"

p. 59)

steep umbilical wall

which, in contrast to
tion

and

is

(Elmi, 1963,

usually present in Exidmetoceras

many Plananunatoceras

of similar whorl sec-

coiling, lesults in a stepped umbilicus.

distinctive char-

acter of Eudrnetoceras could be the nimierous long, often fascicu-

and the strong secondaries of the juvenile whorls

However, if tubercles or nodes are present, the
costation seems to resemljle some "Paranimatoceras." I am unable
to find any significant difference in the septal suture between the
type species E. eudnietiini Buckman and similarly shaped other
Aalenian Hammatoceratinae, although some apparently related
species have simpler and less suspensive sutures. The classification
of the Andean 'Hammatoceras' gerthi Jaworski and 'E. eudmetuni'
jaworskii Westermann is now being reinvestigated by the author
based on new material.
A brief study of Buckman's collections from Bradford Abbas
in December
in Dorset (Geological Survey iMuseum, London)
earlier
expressed
opinion
(Westermann,
confiimed
my
has
1964,
1964a, p. 409) that E. euaptctuin' Buckman, the type species of
late primaries
(cf.

l-.H

cm D)

Eudrnetoceras

(Eiuiptetoceras),

cannot specifically

from 'Harpoceras' ajndlUieijorme Vacek. E.

Buckman appears

be

separated

(Euaptetoceras) am-

from E. arnaltheiforrne in
the rounded umbilical margin and smootli inner flank throughout
giowth although the rare occurrence of intermediate forms cannot
plectens

clearly distinct

be ruled out. Tlie flanks of E. nrnplectens are

flat

and subparallel

Alaskan ammonites,

Westermann

Pt. II:

to slightly convergent while the umbilical

broadly rounded
7

cm

(test also

in diameter.

beuge (1955,
Geczy (1966,

The

keeled)

G7

margin and venter are

up to approximately

at least

Mau-

identification of 'E. amplcctcns' of

pi. 6, figs. 4, 5)

with E. kUtnakompliahim (Vacek) by

is, therefore, probably correct. The coiling of

becomes progressively more involute dining
growth with the juvenile whorls resembling E. (Eudmetoceras)
eudmetinn. Furthermore, the simple strong costation is similar as
in juvenile E. eudmetum and E. (Rhodaniceras) Elmi, the supposed microconch. However, juvenile resemblance in whorl section
is closer to E. inferneuse (Roman) because of absent sulci. Thus, the

E.

p. 78)

(Euaptetoccras)

small E. inferense could be the corresponding microconcli to E.

amaltheifortne.

The

secondary costae of

'E.

euapteturn' vary from

and weakly projected to slightly falcoid; these variable

features were used by Elmi (1963, p. 101) to distinguish E. amaltheijorme from 'E. euapteturn.'
Lelievre (1960, p. 34) distinguished the more widely umbilicate Eudmetoeeras (s.s.) from the narrowly umbilicate (at least

rectiradiate

when

fully

grown) E. (Euaptetoceras)

on allegedly

mer supposedly with

and three
metrical

more

L and

four

ments by Geczy (1966,

elements

lobes)

more

The

elements.

on the

significant "genetical" character of

lobes,

may

basis of measure-

regard this multiplication of um-

(by sulDdivision of

U^ or by introduction

and the corresponding ventrad

inter-relation

ventrally located asym-

shift

of

functional dependant of the increase in whorl overlap

lar

based

the for-

positional differences of

as insignificant

p. 77)

i.e.

central position of a symmetrical

elements, the latter with

have already been judged

bilical

at the generic level

significant differences in the septal suture;

as

of
a

and not

taxonomic significance.

new

direct
as a

simi-

influence the suspension of the umbilical

the suspension angle

(between "saddle line" and radius)

(involute) whorls

being usually reduced in greatly overlapping

without significant differences in total suspension. Furthermore,

the less incised suture of the microconchiate E. (Rhodaniceras) can
probably be explained by the small size and is expected to be

matched by juvenile Eudmetoceras. The reference

oilu in Euaptetoceras (Lelievre, 1960,

p. 34,

based solely on Buckmans' figure (1922,

pi.

to the

Geczy, 1966,

asymmetry
p. 29) was

299) of the holotype of

Bulletin 255

68

'. ciiaptctui/i'

and appears

to be

no more than

mere

irregularity.

specimen figured by Maubeuge (1955, pi.

inider the same name appears to have a symmetrical L.

Finally, the

specimens labelled

as

'E.

fig.

6,

No

3)

other

euaptetiuu' were found in Buckman's

collection of the Geological Survey Museimi, nor

am

aware of any

other descriptions of this alleged species from England. Lelievre's

supposition

(loc. cit.)

these differences were also present in

tliat

Maroccan specimens, contradicts

septal suture was not preserved.

The

close affinity

(Euaptctoceras)

may

statement

his

between juvenile Eudmeioceras

be retained

up

s.

and

E.

to the late adolescent stage so

that incomplete shells are difficidt to classify.

uuch'ospinosum \\'estcrmann from the E.

now

s.

the

that

35)

(p.

case in point

hoioclli

is

E.

Zone which

is

transferred from the former to the latter subgenus. Eunpteto-

ccras

subgenus

therefore, included as

is,

part of

first

The

tliis

sid)genus E. {Pseudapteloccras) Gec/y can

E. (Euoptetoceras) only

beside

in Eucbnctoccras, as in the

monograph.

kUmakornpluduin Vacek

is

if

be retained

type species 'Harpoceras'

tlie

sufficiently distinct

from E. (Eiiapteto-

ceras) anialtlicifortne to warrant subgeneric rank.

The

respective

type species, however, were included in the same subgenus by apparently

previous

all

(1966). However,
that

'//.'

authors,
1

Elmi (1964) and Geczy

(Westermann 1964a p. 410)
strigoceratid on the strength of the

including

suggested earlier

klitnokoipIi(diiin

is

"lanceolate" whorl shape and the occurrence of strongly projected

growth

lines

and

spiral

markings on Wide Bay specimens. Other

strigoceratid characters of 'H.' klimakomphaliirn are the small

um-

bilicus with steep umbilicial walls and the reduction of primaries;

possibly a "prerequisite" for the occurrence of striae. Further studies

have now revealed that

(2)

weak and

irregular concentric markings

specimen of the Wide Bay assemblage,
similar markings or weak striae are also known from E. amal-

are present

on only

(1)

a single

theiforme (the holotype, Vacek, 1886, pi.

ense Roman and Boyer (priv. com. Elmi)

9, fig. 1)
,

(3)

and

E. verpillier-

the costation

is

mark-

edly falcate at least on the immature wliorls of strigoceratids while

it is

(4)

straight to slightly falcoid in

the growth

lines

tiie

E. kUnuiknuipluduni group,

are rarely preserved

and therefore poorly

Alaskan ammonites,

known

Pt. II:

in hammatoceiatids, (5) the

ceratids (priv. com. Geczy,

19r)()),

Westermann

lobe

(6)

the

is

69

usually larger in sLrigo-

Wide Bay sample shows

apparent intergradation of the narrowly umhilicate forms with

those having more widely umhilicate and, significantly, tubercidate
inner whorls, and

(7) the umbilicus of the body chamber opens as

hammatoceratids while remaining small in strigoceratids. I,
therefore, conclude, in agreement with Dr. Elmi's tentative opinion
(priv. com., based on photographs) that the suboxycone forms from

in

Zone of Wide Bay and H. kliiiuikonpJialum are

Hammatoceratinae and are to be placed in Euchnctocera.s.
Nevertheless, tlie resemblance of the suboxycone or discoidal
hammatoceratids of the E. klifnakomphalum group from the L. coucava and S. sowerbyi Zones to early strigoceratids such as the conthe

S.

soxvcrbyi

temporary Praestrigitcs Buckman (but probably floinishing slightly

later) is considered to reflect pliylogenetic relationship. Hence, the
Strigoceratidae are probably derived from Eudmctoceras rather than
from Oppeliidae or Graphoceratidae as recently suggested by
Schindewolf (1964, pp. 367, 430). The resemblances of Strigoceratidae to Sonniniidae (Scheurlen, 1928, p. 37; Schindewolf, 1964,
p. 430) can be explained with their common origin in the Hammatoceratinae.

Concerning the alleged subgenus E. (Pseudaptetoceras) Geczy,

contemporary species morphologically intermediate between

several

klimakomphalum (Vacek) and typical E.

(Roman) has the narrowly rounded externside of E. klimakom phalum but stronger primaries and a much wider umbilicus. E. ainplccteus (Buckman) is also
involute but differs in the roiuided imibilical slope and broad
the type species E. (P.)

(Eunptetoceras) are known. E. infernense

externside,

apparently with

much

variation,

as

well

as

in

the

smooth inner flanks. E. tyrrhcnicum (Renz) appears to have the

whorl shape of E. kUmakomphalum but the ornament of E. amplectens. Furthermore, broadening of the externside and rounding
of

the umbilical walls

appears to be highly variable in other,

Hammatoceratinae such as Planaininatoceras plaunisignc

(Vacek) where the whorls can become almost fastigate. Umbilical
width which is at least partly correlated with whorl section,
earlier

appears to vary greatly intra-specifically as well as inter-specifically

Bllletin 255

70

between appareiitl) dose relatives. This is also indicated in the

presiuiied hammatoceratids ['Pleydellia' auct.] figured by
Burckhardt (1900, pi. 20; 1903, pi. 1, 2), all of which differ,
liowever, from typical hammatoceratids in the much simpler septal
suture and tlie low keel. Suture simplification ("degeneration")

Andean

may

be present to

pears to be a
nificance

of

small degree in E. kUmakovi phalurn and ap-

common

feature

among

hammatoceratids; the

late

remains

character

this

doubtful.

Consequently,

sigif

Pseudaptetoceras were retained as a subgenus of Eudmetoceras, the

type

species

would

be

only

its

certain

species.

'Oppelia'

moerickei Jaworski and 'Hamtnatoceras' discus Merla are tenta-

However, E. klimakomphalum
differs by the broad externvs. c. 14%)
and the coarser cos-

tively included in the type species.

renzi

Elmi

(196.3,

side, the larger umljilicus

and appears

tation

10, fig. 2)

p. 79, pi.

(H)'^^'^,

closer affiliated to E. amalthei^orme. 'Deltoto-

ceras' corroyi Cierard, placed in the species-group of E. klitnakojn-

phalnm by Elmi (UX)^, p. 101), has few widely spaced primaries

and could be an involute member of tlie 'H.' sieboldi (Oppel)
group. Consequently, the subgenus H. (Pseudoptetoceras)

synonymous

tively consideretl
is

shared by Blaison,

unsolved

Still

Buckman,
ceras

Jjy

1925,

al.

tenta-

the taxonomic position of Parnminatoceras

which was placed

Arkell, et

is

This opinion

(1966, p. 101)

et al.

is

witli E. (Euaptctoceras).

(1957)

p.

in

synonymy with Plaiiammato-

267) but with E. (Euaptetoceras)

by the present author (1964a, p. 409) on the grounds that tlie holotype of the type species P. obtectum Buckman (not P. rugatum

Buckman
from

the

(Vacek)

].

as erroneously stated

holotype

Bremer

of

'E.

(1966,

p.

in

loc.

cit.)

eunpteturn'
158)

[^

supported

indistinguishable

is

E.

amaltheijorme

this

opinion. This

statement does not necessarily imply that the species or subgenera

are synonymous.
slightly

The whorl

and the externside

in the umbilicus

(= 3.5

is

sides are

flattened converging only

broad. Also the inner whorls as seen

cm D)

of the plastotypes of both type

species are similarly evolute with

flat

inner flanks and vertical

umbilical wall and have the same dense primaries without tubercles.

There may be a slight difference in the costation of the idtimate

phragmocone whorl, the secondaries being born by furcation in
'P.

obtectum' rather than by fasciculating as in

'E.

eunpteturn'.

Alaskan ammonites,

The

following measurements were

H%

Westermann

Pt. II:

made on

W%

71

the plastotypes.

U%

S(l/+ whorl)

'Euaptetoceras euaptetum' holotype (plastotype)

(phragm.)

12.8

cm

45

26

21

12

and
'Parammatoceras obtcctum'
(phragm.)

There

is

holotype

cm D)

(plastotype)

18.5

48

26

14.5

47

11.5

45.5

27

no evidence

17

(at 7.5
C.5

19.5

21

12

18-19

18.5

for "ogivale"

(like a gothic arch)

whorl

section or tuberculate inner whorls of 'Parammatoceras obtectum'

assumed by Elmi (1963, p. 95) who included in this genus a

seemingly diverse group of known and newly named species from
the L. opalinum and L. murchisonae Zones. Some of these species
are moderately evolute, partially nodose forms with oval whorl
section which are grouped about 'H.' sieboldi
(Oppel)
["P.
obtectum Buckman" in Elmi, pi. 6, fig. 1; P. richei Elmi]. In
as

Westermann appears to be morphobetween 'H.' sieboldi and Eudmetoceras so

that I previously assumed a close relationship between 'H.' sieboldi
and Eudmetoceras (Westermann, 1963a, p. 410)
these resemblances are now thought to be probably due to convergence. In
studying Elmi's monograph (1963) one soon becomes aware that
other more evolute and compressed species placed by him in
Parajiimatoceras' are closely affiliated with Planammatoceras [op.
cit.: 'Parammatoceras' auerbachense
(Dorn)
pi. fig. 2 and 'P.'
boyeri Elmi, pi. 7, fig. 1; compare with Planammatoceras planijorme (Buckman) (pi. 11, fig. 1) nr P. planinsigne (Vacek) P.
dubari (Maubeuge) and P. romani Elmi] although the nuclei may
be distinguished by bullae-like strong primaries. The widely umbilicate 'P.' rochei Elmi (pi. 5, fig. 2) resembles closely Eudmetoceras (Eudmetoceras) masticonnensis Elmi, (pi. 9, fig. 5) as well as
several widely umbilicate Pseudammatoceras species of Elmi, i.e. P.
Alaska, E. niicleospijwsum
logically intermediate

'

grandis, P. mouterdi,

and

placed in Parammatoceras.

rugatum Buckman which Buckman

The unusual oblique bullae of 'Param-

P.

matoceras suballeoni' Elmi and of the closely affiliated

'P'

alleoni

Bulletin 255

72

Dumortier (Elmi, 1963, pi. 8) are also present in the closely

grouped early Aalenian group of Pseudammatoceras ornatum Elmi,
P. guUense (Renz) P. durnortieri (Prinz) and probably P. subinsigne (Oppel) type species of Pseudammatoceras Elmi.
The dubious 'Parammatoceras' appears to be synonymous with
E. (Euaptetoceras) unless investigations on new material from the
type locality and upe stratum can demonstrate that the inner
,

whorls of the large type species P. obtectinn are significantly

differ-

ent from those of the E. ajnaltheijorme group and, instead, resemble those of
P.

rugatum

immatme

H. sicboldi group,

tlic

'Parammatoceras' ,

in

as

implied by Elmi. By placing

Buckman implied

strongly nodose

whorls. However, Elmi's transfer of this species to his

new

genus PsciuUnitinatoceras seems to indicate that either Buckman's

classification was a\i ong or that 'Parammatoceras' as understood by
Buckman intergrades with Pseudammatoceras. Consequently, it
appears advisable for the time being (1) to consider 'Parammatoceras' a

uomcfi dubiufu,

(2)

to tentatively transfer the lot of the

(1963) to PseudamPseudammatoceras
as a
matoceras', and (3)
Hammatoceras.
The
cresubgenus of either Planammatoceras or

species included in 'Parammatoceras' by Elmi

to tentatively classify

ation of yet another taxon

Csernyeiceras

Roman and
wliorl section

Geczy,

is

certainly inadvisable.

based on Hammatoceras verpillierense

Boyer, appears to be well separated on the base of

and

septal suture.

EUDMETOCERAS Buckman, 1920

EUDMETOCERAS (EUDMETOCERAS) Buckman,
Genus

Subgenus

Eudmetoceras (Eudmetoceras)

sp. aff. E.

PI.

liuJmrtoirras (s.s.) cf. E. fudmctuin

Bay), Bull. Amer. Paleont., vol. 47, p. 409.

1964.

Material.

One

of another large

from

uses

Wide

Description,

Buckman, Westermann

(Wide

and one small fragphragmocone, well-preserved internal molds

(USNM

21251

15, figs. a,b

large jjhragmocone whorl

ment

soiverbyi Zone,

1920

eudmetum Buckman, 1920

160926). Pseudocidoceras zonule,

S.

Bay, Alaska, Penninsula.

lihe

widely umbili(ale

plnagmocone whorl of 12.7 cm diameter is

(U 38%) and compressed oval in section with

narrow steep umbilical

slope. Tlie externside of the internal

mold

Alaskan ammonites,

Pi. II:

Westermann

73

is narrowly rounded, almost fastigate, and shows remnants of a

narrow sharp hollow-tloored keel. The costation is strong becoming somewhat blunt toward the end ot the preserved phragmocone.
The ultimate half whorl carries nine somewliat bullae-like, rectiradiate primaries which bifurcate or trifmcate at about two-fifths
whorl height. There are 28 partly intercalated, markedly convex
and strongly projected secondaries on the idtimate lialfwhorl which
reach the smooth narrow median band (internal mold) at an angle
of about 45 (corresponding to a ventral costae angle of 90)
Septum and suture are well preserved. The septum is planodis-

culate; the intensely incised sutures

(PI.

15, fig. a)

are terminally

not markedly approximated, indicating that the phragmocone

incomplete.

The

other fragmentary specimen

and blunt costation

sutures

at

about 17-20

cm

L and E

are slender with long thin stems.

has

is

approximated

diameter. All lobes

are equally deep, the

L/Uo

saddle is almost as high but only 2/3 as wide as E/L, U^ is

narrow, asymmetric and somewhat oblique; U;^ (? -|- U4) is strongly

suspensive consisting of two major external indentations

lobes?)

The

saddle boundary

kink over L/Uo.

The

is

(or

therefore strongly bent with the

internal suture has a single

dominant U/I

saddle and adjacent strongly suspensive subdivided umbilical ele-

ments.

Comparison.

The

incomplete

Wide Bay specimen

resembles

the corresponding ultimate halfwhorl of the holotype of E.

tum [plastotype studied] although

shell.

The

the latter

is

eudme-

preserved with the

only differences are in the slightly wider umbilicus, the

more widely spaced primaries and the somewhat stronger

Wide Bay specimen (see measurements
below) Buckman's paratype, a topotype from the L. discites Subzone of Bradford Abbas in Dorset, which is much larger and has

stronger

flexed secondaries of the

.

an incomplete body chamber, has even closer set primaries. However, the inner whorls up to eight cm diameter of the Wide Bay

form are unknown.

Eiidynetoceras masticonnensis Elmi (1963, p. 69, pi. 9, figs. 5a,
from the L. discites Subzone of Maconnais, France, is distinguished from the type species by the nodose inner whorls, which
are not preserved in the Wide Bay specimen, and the stronger
bullae-like primaries of the phragmocone of comparable size (see
b)

38

Alaskan ammonites.

Trav. Geol. Bulg.,

forme (Vacek)].

Ft.

Ser. Paleont., vol.

6,

Westermann

II:

pi.

2,

figs.

2a-c

= ^.

75

cf.

amalthri-

(Pscudaptetoceras) klimakomphalum involutum

1966. Hammatoccras
(Prinz, 1904), Geczy, id., p. 80, pi. 20, figs. 1, 3; pi. 40, fig. 13; pi. 51,
fig. 1; text-fig. 67 [
. cf. amplcctcns (Buckman)].

}non

Lectotype
figs. 17, 17a.

(det.

Sapunov, 1964,

p.

262):

Vacek, 1886,

densed" lenticular limestone beds at Cap

St.

p. 79)

has seen the type specimens, the costation of the lectotype

what more feeble than represented in Vacek's

Measurements of type specimens according
or to original figures (in brackets).

Dmm
(phragm.)

8,

Lago de

Vigilio,

Garda, in the southern Alps. According to Geczy (1966,

lectotype

pi.

probably incomplete phragmocone from the "con-

is

who

some-

figure.
to

Vacek (page 81)

W%

H%

U%

Bulletin 255

76

30 %U

20 %U
VoU

phrogm.

E klimakomphalum

lectotype (L) + topotypes

E.k.discoidale, ho\oV{H)

Enucleospinosum, ho\oV{H)

20

10

off.

nucleospinosum

50

100

200

300

D (mm)

Text-fig. 22.
Scatter with growth lines for relative umbilical width
of Eudrrutocrras (Eiiaptrtocrras) spp., phragmocones. Note the change
from widely iimbiiicate ju\'enile ^vhorls to involute mature whorls at 30-40
D and the close resemblance between E. (E.) klimakomphalum (Vacek) s.s. and
E. (E.) klimakomphalum discoiJalr, n. subsp., while E. (E.) nucleospinosum
Westermann and E. (E.) sp. aff. E. nucleospinosum, respectively from the E.
hoii-elli and S. soaverhyi zones, tend to retain the wide umbilicus.

{U:D)

mm

De.scril)tio>i.
ter,

The

innermost whorls, up to 12-15

are widely uml)ilicate, subcircular, keeled

mm

diame-

and may have simple

costation with or without fine tubercles. Signiticantly, the strongest

ornament

umbilicate

in

is

found in the most widely

fig.

8a). Tliere can be little

form of fine tubercles

luiclci

(paratype, PI. 14,

Alaskan ammonites,

Cross-section

Text-fig. 23.

nucleospinosum
12405

(USNM

Westermann,
160231)

of

Pt. II:

Westermann

Eudmetoceras (Euaptctoccras)
complete phragmocone,

probably

77

sp.
loc.

aff.

E.

USGS

1-

Cross-section of Eudmetoceras (Euaptctoccras) klimakomText-fig. 24.

phalum discoidale, n. subsp., phragmocone (section of ultimate half whorl rotated bv 20), loc. USGS 19863 in the Pseudocidoccras zonule (USNM 160230)
;

1.

doubt that

all

specimens from locality

USGS

12405 belong to a

single variable species.

At 12-15 mm diameter, the whorls become rapidly compressed

and involute. The intermediate phragmocone whorls are tightly
coiled (U-10-16%)

subtrigonal ("lanceolate") in section with nar-

Bulletin 255

78

Text-fig. 25.
Septal suture
(Euaptctoof
Eudmctoceras
crras) klimakomphalum discoid(dr, n. subsp., holotype, phraginocone at 72
D; X 1-5.

mm

row, steep and usually well-separated somewliat curved umbilical

and gradually converging outer

on which
is set a high and narrow, hollow-floored keel. The ornament consists of blunt almost rectiradiate, often weakly rmsiradiate and
somewhat projected costae which in the earlier stages usually reach
over the whole flank and are more or less weakly fascicidate. The
later growtli stages have irregular blunt primaries which split into
two or three secondaries; the inner flanks tend to ijecome smooth
conmiencing from tlie umbilical margin, so tliat beyond 30-50 mm

walls,

subparallel

inner

flanks

flanks forming (internal mold) an acutely fastigate venter

diameter they bear only irregidar radial undulations while the

blunt costae are restricted to the outer one-third of the

The

sides.

one and one-half whorls of the phragmocone,

beyond 60-100 mm diameter, are compressed subtrigonal ("lanceolate") in section with well-roimded imibilical slope and narrowly
rounded to fastigate venter and strong hollow-floored keel. The
broad blunt costation is restricted to the outer third of the whorl
sides while the centre and inner flanks are smooth. The end of the
jjhragmocone, which reaches 130-150 mm diameter, may become
somewhat wider umbilicate.
The body chamber is preserved only in a few small fragments
which show some distant very blunt secondary costation.
The matiue septal suture (PI. 16, fig. If; Text-fig. 25) is highly
complex with long narrow finger-like lobes. L is longer than E and
the U lobes; tlie L/U^ saddle is much smaller than E/L, and the
other lolies and saddles of tlie external suture form a declining
sequence with straight saddle-boundary. Only the inner umbilical
elements are slightly olilitpie. However, tlie suture of the somewhat
last

one

to

Alaskan ammonites,

more evolute immature whorls

Pt. II:

(20-35

Westermann

mm

D)

is

79

clearly suspensive

with two or three oblique umbilical elemeuts.

Discussion. One large specimen and a fragment from

USGS

whorls and by
ings
16,

loc.

21251 of the same zonule differ by extremely involute mature

tlie

preservation of lateral irregidar faint spiral mark-

and strongly projected weak growth lines on the shoidder (PI.

However, typical spiral striae or
fig. 2; PI. 18, figs, la, b)
.

grooves as developed in most Strigoceras

man) described by Imlay

S. humphriesianum Zones

(1964, p.

[cf. S.

languidurn

(Buck-

37) from the O. sauzei and

of southeastern Alaska] are absent.

The

suture is poorly preserved except for some

Although this specimen is probably only an involute variant of E.
klimako7nphalum discoidale the close resemblance to Strigoceras
intensely frilled detail.

is

considered significant.

Another specimen of the same assemblage, which

is

distin-

guished by evolute nodose juvenile and intermediate whorls, is described below as E. (Euoptetoceras) cf. E. nucleospinosinn Wester-

mann.
Comparison. ^\\\f^ new subspecies is distinguished from E.
klimakomphaliim s.s. by the rounded umbilical slope of the outer
whorls and probably also by stronger compressed whorls with more
acutely converging flanks forming a more narrowly rounded, almost fastigate venter, and the somewhat wider spaced costation.
'Hammatoceras discus', Merla and 'Harpoceras (Lioceras) amaltheiforme mut. involutum' Prinz (see Geczy, 1966, p. 80, pi. 20, figs. 1,
3) are both closely affiliated with the restricted species and probably mere varieties. Neither the suture nor the umbilicus of the
as supposed by Merla
and inferred by Geczy (1966, pp. 65, 66, 80) The
distinction of 'H. klimakomphahnn paenamplectens' Geczy (1966,
based solely on the supposedly simpler septal suture of a
p. 81)

former appear to be significantly different

(1934, p. 18)

single possibly corroded specimen, appears dubious. Geczy's state-

(Buckman) has a simpler

klimakomphalum does not agree with my own
perience and other descriptions (see Westermann, 1964a, pi. 68,

ment

that Eiidmetoceras amplectens

ture than E.

su-

exfig.

Elmi (1963, p. 100) is probably

correct in placing 'Oppelia subradiata Sowerby var. tyrrhenica'
la;

Bremer, 1966,

pi. 15, fig.

la).

Bulletin 255

80

Renz

(1925, pi.

2, tigs.

1,

la)

in E. (Euaptetoceras).

from E. klimakoin pJialum

smooth inner half of the whorl

(listingiiished

totally

E.

tyry/ioiinim

closely

resembles

(1926; lectotype here designated:

s.s.

sides

'Oppelia'
pi.

3,

This species

is

apparently only in the

figs.

throughout growth.
mocrickei
12 a-c;

Jaworski

?non

fig.

13)

from the (upper) L. concaxui or lower S. soicerbyi Zone of Mendoza in tiie southern Andes (type specimens and type locality reinvestigated) which has a hollow-floored keel, a slightly rounded
uml)ilical wall and obsolescent j^rimary costae. E. klhuakoin plialum
nioerickei is distinguished from E. kUiunkomphalum s.s., E. k.
discoidale and E. tyrrhrniciirn by the denser, almost rectiradiate and

The new subspecies has broader,

more widely spaced costae than any of the named species and
subspecies. There is no sharp innbilical margin or edge as in E.
hlit/iakoni phdluin s.s. and the flank of the last whorl rounds gently
oiih slightly projected secondaries.

to

the uml)ilical seam, similar as in E. atuplectcns.

agui Ionia' Imlay

(1961, p. B35, pi. 5)

'Witchellia}

from southeastern Alaska,

which is a typical E. (Euaptetoceras), has much more robust whorls

with relatively large immatiue imibilicus and stronger costation.
Measurements.

Dmm

Holotype

(phragm.)

W%

H%

U%

Alaskan ammonites,

Pt.

II:

Westermann

81

phragmocone from USGS 12405 (USNM 160231),

Kialagvik Formation, probably F.seudocidoceras zonule of

large

erbyi Zone,

Wide

upper
sow-

S.

Bay.

The

up to about 40 mm diameter,
and almost planulate with
shallow umbilicus, while the outer one and one-half whorls become
abruptly much more involute (U = 16%) and typically discoidal
Description.

inner whorls,

are widely umbilicate

(U

30%)

with rounded subvertical innbilical wall.

The
rounded
eral

nucleus below 12-15

to

nodes

mm

somewhat depressed
[six

to

seven

per

diameter

in

is

typically "coronate,"

whorl section with strong

half-whorl].

The

next

whorl

latis

compressed oval in section with fastigate venter on the internal

mold; the shell had a sharp hollow-floored keel and carried densely
fasciculate, almost rectiradiate, slightly to moderately projected

which retract from the innermost flanks. The umbilical

seam of these inner whorls lies outside the lateral nodes of the preceding whorls at approximately one-half whorl height.
At 45-50 mm diameter, the rovmded steep umbilical wall develops and the overlap increases rapidly so that the last whorl's
umbilical seam lies on the penultimate whorl's umbilical margin.
The inner flanks are now flattened and subparallel while the outer
costae

flanks converge gently

The

up

to the high,

narrow, hollow-floored keel.

whorl has irregularly branching and somewhat fasciculate, slightly falcoid costae which strengthen on the outer flanks;
there are about 13-14 blunt prosoradiate primaries and 30-32 projecting secondaries on the last one-half whorl of the phragmocone.
Discussion. Eudmetoceras (Euaptetoceras?) nncleospinosum
Westermann (see appendix) is distinguished by the larger "coronate" stage with somewhat stronger nodes which are "impressed" in
the subsequent umbilical wall. E. amaltheijorme (Vacek) which is
last

closely affiliated,

differs

in

the cross-section with well-developed

shoulders and broadly rounded externside. This specimen

phologically intermediate between E.

klimakomphalum

is

mor-

discoidale

and the slightly older E. nncleospinosum from the E.

howelli Zone of Wide Bay, probably reflecting phylogenetic rela-

n.

subsp.

tionship.

Bulletin 255

82

Measurements.

Dmm
USNM

160231

(phiagm.)

94

H%

W%

U%


Alaskan ammonites,

Pt. II:

Westermann

83

30 % U

80
70

60
20 %U

50

40

1-

30
10

%U

20

10

phrogm.

body ch

5 holotype (H) +

topotypes
a

var {subsp?)ogu//on/o, types

50

20

15

100

200

150

300

D(mm)

Text-fig. 26.
Scatter with growth lines for relative umbilical width
of Eudmetoccras (Euaptctoceras) amplcctcns (Buckman), for comparison of the type specimens from England with the sample from the E. amplectens zonule of Wide Bay and 'WitcheU'ial aguiloTiia' Imlay [= var./susp.? of
E. amplcctcns^ from the O. sauzci Zone of southern Alaska. Note the morphogenetic change and the great similarity of the three "samples" with the possible exception of adult phragmocones
however, the holotype of E. amplcctcns
is probably an involute variant.

(U:D)

Euaptctoceras amplcctcns (Buckman), Elmi (S.E. France), Trav.

Labor. Geol. Lyon, p. 72, pi. 10, figs, la, b.
1964. Eudmetoccras amplcctcns Buckman, Sapunov
(Bulgaria), Trav.
Geol. Bulg., ser. Paleont., vol. 6, p. 261, pi. 1, figs, la, b, pi. 3, figs,
1963.

la, b.

1964.

Eudmetoccras amplcctcns (Buckman), 1886 [error for 1889], Wester

Bull. Amer. Paleont., vol. 47, No. 216, p. 417, pi. 67,

mann (Wide Bay),

figs.

1964.

2a-c;

pi.

68,

figs,

la-c;

IFitc/icllia? aguilonia

text-figs.

Imlay

n. sp.,

28,

29.

Imlay (Alaska), U.S. Geol.

Prof. Pap. 418-B, p. B35, pi. 4, fig. 9, pi. 5, figs.

(?)1964. fVitc/icllia? aff. IF.? aguilonia Imlay n.
B36, pi. 4. figs. 1-4.

4,

Sur.,

7-9.

sp.,

Imlay,

id.

ac,

p.

Bulletin 255

84

amplrctnis
(Buckman),
Bremer
(Euaptrtocrras)
Eudmrtoirras
(Turkey), N. ]h. Geol. Palaeont., Abh. 125, p. 159, pi. 15, fig. 1.
klimakomphalum
involutum
(.')1966. Hammatoccras
( Ps(udaptrtocrras)
(Prinz, 1904), Geczv (Hungary), Geol. Hung., ser. Paleont., fasc. 34,
1966.

p. 80, pi. 20, figs. 1,

3.

non 1955. EuJmctocnas amplrctcris (Buckman), Maubeuge

Mem. ^uisse paleont., \ol.

makomphalum (Vacek)].
non
et

71,

p.

34,

pi.

6,

figs.

4,

5,

(Swiss Jura),

[=

E.

cf.

kli-

Eudmctocrras (Euaptetoceras) amplectens Buckman, Blaison

(French Jura), Bull. See. Hist. Nat. Doubs, No. 68, fasc. 4, p.
figs. 1, 2 on p. 101 [=. cf. amalthciforme (Vacek)].

1966.
al.

100,

Nine incomplete and more or less strongly corroded internal molds, partly with body chambers and test remains, and several fragments, from
8 (} 952-968)
? one fragment from
4; two fragments from
5; two internal molds
of phragmocones and body chamber fragments from
12
one almost complete well-preserved phragmocone
(fl2r)2a, b)
from USGS 12105. The collections
5 and
4,
8 are
from the massive greywackc beds [. amplectens zonide] at the base
of the 5. sowerbyi Zone; USGS 12405 is from the superjacent
Pseudocidoceras zonule of the same zone, while
12 is from
the Pseudobigotites zonule of the O. saiizei Zone. All from the
Material.

WA

W\

WA

WA

WA

WA

WA

WA

Kialagvik Formation, southeast shore of

Descriptioji.

species

which

is

This

more

Bay.

highly variable in coiling (umbilical width)

morphogenetically and infraspecifically

the

Wide

appears to belong to a single

material

loosely coiled forms are

more

(Text-fig.

26)

both

As

usual,

strongly ornate than the

very involute forms. Because the involute variants are indistin-

guishable from typical E. amplectens

whole material
evolute forms
lonia'

(Imlay)

is

(U
.

It

S.

soiverbyi

Wide Bay and on

as

the

relatively

'forma agui-

however, improbable that these constitute

Zone and

widely umbilicate forms

at

The abundant

15%) may be distinguished

is,

an inde]>endcnt subspecies or
the basal

(Pis. 20, 21; Text-fig. 26)

placed in this species.

may

species.

Both formas are present

in the O. saiizei Zone, but the

in

more

be relatively more frequent above,

the Iniskan Peninsula. Alternately, the total

intergrading form group could perhaps be included in the subspecies E. aynplectens aguilonia

(Imlay)

Alaskan ammonites,

Pt.

II:

Westermann

The mature shell is large, involute and subdiscoidal with

rounded umbilical slope, broadly rounded venter carrying a hollow-floored keel, obsolescent primaries and densely spaced moderately projected secondaries. The umljilical seam of the last wliorl,
including the body chamber, egresses strongly. The nucleus is
much more widely umbilicate than the mature phragmocone and
may have very blunt, irregular, distantly spaced primaries. The
morphogenesis from this evolute juvenile stage into the typical
involute growth stage occurs at varying sizes; as expected, a longer
retained and more evolute juvenile stage will generally result in
a less involute mature shell, i.e. forma aguilonia (Text-fig. 26)
The nucleus is visible in two specimens (J 965 and USNM
160234;

PI.

21,

fig.

Id)

The innermost whorls

bearing a low sharp keel and moderately evolute

are

(27%

subcircular
at 15

mm

D) becoming slightly compressed oval at 12-15 mm diameter, and

embracing the preceding whorl slightly more than one half. At
20-25 mm diameter the whorls become abruptly more involute and
develop a compressed trapezoidal whorl section with rounded umbilical walls and shoulders. The mature phragmocone of the complete specimen is narrowly imibilicate (11%), discoidal with wellrounded umbilical margin and steep but not vertical umbilical
slope, converging outer two-thirds of the flanks and broadly rounded venter which carries a hollow-floored keel. There are only
some irregular swellings on the inner flanks of the penultimate
halfwhorl while the outer third of the flank carries blunt, moderately projected secondaries, 31 per halfwhorl.

The

other nucleus

(J 965) is from a more widely umbilicate specimen. The whorl

section changes from subcircular to compressed oval at 15

mm

diameter, becomes trapezoidal at 20-25

tively evolute

mm

(27%

at 26.5

mm

D,

c.

mm

D, but remains

26%, at 32

mm

rela-

D, 22%,

at

This specimen has much stronger costae which on the

nucleus are irregularly fasciculate and markedly falcoid with raised,
irregularly arranged bullae-like swellings on the inner flanks. At
about 30-35
D the costae become rectiradiate, moderately projected and restricted to the outer flanks while the inner flanks
have only a few irregular broad swellings. There are about 26-30
secondaries per halfwhorl. This form is probably identical with
46

D)

mm

Bulletin 255

86

'Witclu'llia} alt. IT.? nguilonid' ol

the O. sauzci

Zone

Imlay (1964,

from

pi. 4, figs. 1-4)

of the Iniskin Peninsula.

Other specimens from the fossiliferoiis E. amplectens zonule

(W'A 8) are intermediate in all visible morphological features between the in\olute weakly ornate forms (} 963) and the more
widely imibilicate, more strongly ornate forms

968)

(J

Irregular

broad swellings on the inner flanks which often extend onto the
umbilical slope, can be seen in the uml^ilici of most median to
relatively

widely evolute variants.

One

of

the

evolute

variants

(about five per halfwhorl)

(J 968) has distant blunt primaries
up to a diameter of 110 mm. The secondaries on the mature

phragmocone

of these relatively evolute variants are widely spaced

(about 25 per halfwhorl)

end

of large

phragmocones

and weaken markedly only towards the

at 120-150

tation of the involute variants

mm

diameter, while the cos-

(averaging 30 secondaries per half-

becomes obsolete already at 80-110 mm. Densely spaced,

concave simple growth lines are sometimes visible even
on the internal mold and some low swellings may be developed.
Toward the end of the phragmocone, at about 80-100 mm diameter, the imibilical seam egresses rapidly, the umbilical wall becoming lower and less steep. The full phragmocone diameter is
wliorl)

slightly

mm.
The body chamber

120-160

is

two-tliirds to three-foinths

whorls long;

the two specimens with preserved apertures are approximately 200

mm

in diameter. The umbilical seam egresses strongly until the

apertme embraces only about one-third of the preceding whorl.
While the whorl height is somewhat reduced, the inner height
(dorsum to venter) remains approximately constant. The flanks
flatten and may become slightly concave in the lower third. The
aperture is preceded by a weak constriction of the internal mold

probably indicating thickening of the

moid with broad

21,

fig.

The

peristome

is

and ventral lappet

sig-

(PI.

2).

The

septal

suture

is

highly complex with

which interfinger from one

E;

shell.

mid-lateral projections

line into the next.

slender

is

endings

longer than

the saddles decrease successively in size towards the umbilical

seam following
(U.,

-)-

?)

a straight saddle line; the inner umbilical elements

are usually oblicjue

and somewhat suspensive.

Alaskan ammonitks,

Pt.

II:

Westermann

87

Since

the first volume of this monograph (Westermann, 1961a) went into press, Eudiuetoceras arnplcctens has been
described (1) from the Maconnais in eastern France (Elmi, 1963),
from beds ijetween the L. concavn and S. sowerbyi 'beds'
(2)

Discussion.

[PL. discitcs Suljzone]

of Bulgaria

(Sapunov, 1964),

(3)

possibly

fiom the L. concavn Zone of Hungary (Geczy, 1966; as Hanunnloccras klirnakoniphalu?n involutiitn (Prinz), (4) from the L. discites
Subzone of Turkey (Bremer, 1966), and (5) from tlie O. sauzci
Zone of southern Alaska (Imlay, 1964; as Witchellia} agiii Ionia,
n. sp.)

While the European and Turkish finds, as well as the two

my first volume from the E. howelli Zone

specimens described in

of southern Alaska lie within the generally accepted vertical range

of E. amplectens,

i.e.

Zone

for

L. concava to lower

(upper)

Zones, the range of the species

now extended

is

Wide Bay and probably

also for the

soiaerbyi

S.

into the O. sauzei

Talkeetna Moimtains
southern

of the Iniskin Peninsula (type locality of TF.? agiiilonia')

Alaska. Although the exact level of the Talkeetna Mountains oc-

cmrence
tive,

the

mens.
lay

(as

unknown and

is

sauzei

5.

the zonal assignment, therefore,

well established for the

is

tentaspeci-

and recovered one each of the inmore widely umbilicate forms from the Pseudobigolocality WA 12, more than 100 m above the Pseudo-

Witchellia} agiiilonia)

zonule of

cidoceras zonule. 'Witchellia}

B36)

Wide Bay

have seen the specimens from Wide Bay recorded by Im-

volute and the

tites

Zone

aff.

TF.? aguilonia'

appears indeed to be an evolute variant of

with which

it

is

Imlay (1964,

Imlay

(1964

p.

'TF.? aguilonia'

associated also in the Talkneeta Mountains.

p.

noted the "similar appearance" of the

B36)

'Witchellia} aguilonia, n.

sp.'

and Eudmetoceras amplectens but was

obviously averted by the slight difference in age.

that the former "appears to have

more flexuous

He
less

also stated

regular rib-

bing that tends to be more fasciculate." However, these supposed

differences do not apply for the majority of specimens, including
both figured specimens of 'IF.? aguilonia' [holotype and paratype],
according to

On

my knowledge

of E. aynplectens topotypes.

septal suture as figured by Imlay

from a paratype, as partly seen on the photograph of the holotype, and observed by me on other specimens

the

(1964, pi.

strength of

5, fig. 4)

the

Bulletin 255

88

including

'IF.? aff.

IV.} dguilotiia' of

Imlay

(1964, pi.

4,

fig.

4),

no doubt that 'Witchellia} aguilonia' belongs to the

Hammatoceratinae and not to the Sonniniidae. The suture is extremely complex, and the innbilical elements are suspensive.
there can be

The

majority of the southern Alaskan specimens are distinfrom typical European E. amplectefis in the somewhat
wider umbilicus of the (mature) phragmocone (U^ 15-20% vs.
10-12.5''p) and by the often present widely spaced, strongly irregular swellings on the inner flanks of the inner whorls. However,
these swellings are apparently absent on the holotype and the
large figured paratype of 'IF.? oguilouia'. Furthermore, the morphological variation of the typical European E. atnplectens is jx)orly
known. It is possible that E. atnplcctens of Dorset (type locality)
intergrades with the more widely umbilicate E. amaltheiforme

gtiished

[syn. E. euaptetuin

Buckman]

w4iich has densely spaced primaries.

Although the European E. arnplectens, including the holotype,

reacli the involute growth stage at a small size, some specimens,
such as Buckman's paratype (pi. 180, B, fig. 2) retain the more
widely imibilicate stage until about one-half full size (U^16.5% at
69 mm D, see Text-fig. 26) The fact that no European E. arnplectens lias been described with a mature umbilical width of more
tiian 13%, as is common in the Alaskan forms of both S. sowerbyi
and O. saiizei zones, may be due to "splitting" although this ap,

pears improbable.

In conclusion,

it is

advisable to place the

and the involute weakly

costate

single species E. mnplectens and,

former as forma aguilonia

The

ratlier

more evolute ornate

specimens from Alaska in the

if

necessary,

to distinguish

the

than as subspecies aguilonia.

involute forms of the Alaskan E. arnplectens resemble

E. kliniak<))np}ialiim discoidale n. subsp. from the slightly younger

Pseudocidoceras zonule of

Wide Bay from which

they are distin-

guished particularly in the Inroad shoidders. Significantly, these

species have never been foimd togetlier which is probably owing
to different liabitats,

water.

i.e.

E. arnplectens possibly living in shallower

Alaskan ammonites,

Measurements.

160234 (phragm.)

II:

Westermann

Dmm
USNM

Pt.

W%

H%

U%

89

Bulletin 255

90

cf.

Eudmetoceras
Material.

part

(McM

s.

J 958)

1.

indet. [?

PI. 21, fig. 3

single riglu external

.

WA

mold from

E. amplectens zonule of the

5.

8,

lower

sozverbyi Zone,

upper Kialagvik Formation, \Vide Bay.

Description. At .Hfi mm diameter, the whorls are compressed
oval in section and Avidely imibilicate (33%)
Significantly, the
umbilical seam egresses markedly at the end of the preserved conch
.

Avhich coincides with modification of the costation, suggesting that

tlie

shell

The

was adult. However, the septation

costation

consists

where they

and of
one-quarter whorl, the costae become
i)ullae

is

somewliat

of

invisible.

falcoid

costae

intercalatories.

fasciculate,

partly reduced

On
and

forming
the last
irregu-

larly sj^aced.

T^his

and

specimen
D) of the widely
umbilicate variant of the macroconchiate Endmetoceras amplectens
and coidd, therefore, be the corresponding microconch (male) The
microconchiate subgenus E. (Rhodaniceras) Elmi is distinguished
by ventral sulci and strong, not projected costae.
Discussion.

adidt

probaijly

closely resembles the nucleus

(up

to

c.

25

complete

mm

Genus

PLANAMMATOCERAS

?Subgenus

P.

Buckman, 1922

(PSEUDAMMATOCERAS)

["Ptiramrruitocrras" Buckman, 1925,

Planammatoceras (Pseudammatoceras?)

cf.

P.

Elmi, 1963

rinrn

<//;.]

benneri (Hoffman) 1913

PL
(

22, figs. 2,

Hammatocrras brncri sp. nov., G. Hoffmann (Germany), Strat.

u. Amm. Fauna Unt. Doggers in Sehnde, p. 191, pi. 18, fig. 15.
1964. Sonninia cf. S. patella (Waagen), Imlav (Cook Inlet), U.S. Geol.

.')

1913.

Sur., Prof.

Paper 418-B,

p.

B33,

pi. 4,

figs. 2-4.

Material. One large almost complete well-preserved specimen from uses 21251 (USNM 160235). Psendocidoceras zonule
of tlie
sozverbyi Zone, Kialagvik Formation, Wide Bay.
Description. The inner whorls up to 25-30 mm diameter are
.v.

widely umbilicate, subcircidar in section, and have a thin low keel.

Suljsequently, the whorls

become more involute and increasingly

compressed with suljtriangular section and narrowly rounded venter

with hollow-floored, low keel. The narrow steep umbilical wall be-

comes increasingly more separated from the

converging flanks by the umbilical margin.

slightly

convex and

Alaskan ammonites,

Pt.

II:

Westermann

91

(20 mm D) consists of slightly

and weakly projected simple ribs wliich partly fasciculate. Subsequently, widely and irregularly spaced (4-6/halfwhorl)
strong bullae-like primaries develop on the inner third of the
flanks while the intermediate primaries become obsolete. The secondaries, about four times as numerous as the primaries (20/halfwhorl) are bliuit slightly rursiradiate and somewhat projected on
the shoulders. They die out gradually beside the keel. The middle
of the flank, at the base of the secondaries and beneath the subsequent umbilical seam, is almost smooth so that the secondaries appear disconnected. Towards the end of the phragmocone (120 mm
D) the costae become obsolete with some wide undulations remaining on the inner flank.
The body chamber, three-fourths whorls long, terminates in
the partly preserved simple apertiue (195 mm D)
The inner
flank becomes more compressed and slightly concave beside the

The

costation of the nucleus

rursiradiate

well-developed

umbilical

The

margin.

outer

three-fifths

of

the

rounded shoulders. The

venter is obtusely fastigate. The last halfwhorl is smooth except
for faint somewhat prosoradiate and slightly projected growth lines.
The septal suture is highly complex with typically oblique and
flanks are flat

and converge

to the broadly

suspensive umbilical lobes.

The specimen closely resembles Sonuiuia ex. gr.

(Waagen) for which it was at first mistaken before
the typically hammatoceratid septal suture was observed.
This specimen belongs to the species-group of Hammatoceras'
sieboldi (Oppel) which flourished in the L. murchisonae Zone and
was placed in the dubious 'Paraminatoceras' Buckman by Elmi
The Wide Bay form is distin(1963, p. 65; holotype refigured)
guished from 'H.' sieboldi by the much wider spaced secondaries
and closely resembles 'H.' benueri Hoffmann, originally described
from the upper L. concava Zone (-f- L. discites Subzone) of Hannover, Germany. Elmi (1963, p. 94) placed 'H.' benneri in Pseudamatoceras but separated Hoffmann's similar paratype (fig. 14)
not only specifically (using the species name achillei which had
been tentatively suggested by Hoffmann) but generically by placing
it in Eudmetoceras s.s.
Discussion.

S.

patella

'

Bulletin 255

92

Of special signilicaiue is tlie occurrence of this form in the

Talkeetna Mountains on the Iniskin Peninsula of southeastern
Alaska. The single large specimen described as 'Sonninia ci. S.
patella (Waagen)
by Imlay (1961. p. B33, pi. 4, figs. 2-4) came
from a large collection indicating the O. sauzei Zone (op. cit., table
'

12, p.

B27)

although

it

might

liave originated in a

within the 60-65 thick sequence of

table 11).

My

USGS

locality

rein\estigation of this specimen

lower horizon

324113 (op.

(May

1967)

cit.,

showed

is complex and strongly suspensive in condevelopment in all Sonniniidae. If the origin of this
form in the O. sauzei Zone can be established, the upper range
of the Hammatoceratinae would be extended from the (lower) 5.
sowerbyi Zone to the O. sauzei Zone.

that the septal suiiue

trast to the

Measurcjnents.

Dmm

USNM

160235

(body ch.)

Y{%

W7c

U%

Alaskan ammonites,

tlie

Pt.

highly variable species

single

Westkrmann

II:

S.

adicra

93

can clearly be

tinguished within the European realm of the subgenus

dis-

(Eufioplo-

5.

ceras)

which ranges

but

vertically essentially restricted to the L. discites Subzone, al-

is

from southern England

laterally

though appearing rarely

trigotuilis

.S'.

Marocco

coucava Zone of southern Eng-

in the L.

land and ranging into the

to

Subzone of the middle

S.

soiuerbyi Zone.

The
is

distinction of Sonninia (Eiihoploceras)

certainly not sharp.

Even

pi uqiiaus Bayle, resembles

many

from Souninia

the type species of the latter,

s.s.

pro-

S.

adicra, type species of the former, in

S.

forms have been named such

Buckman and 5. sowerbyi adicroides Hiltermann,
which are now placed in S. adicra. Nevertheless, the sub-

respects; several intermediate

as 5. spinifera

both of

genus has been retained in the Treatise and most recently by Westermann (1966) especially because of stratigraphic usefulness. 5.
(Eiihoploceras) includes Sonniniidae tending to have evolute round-

ed whorls with rursiradiate strong costae, which are retained beyond the spinous stage onto the body chamber, and a thin low
hollow-floored keel. However, some variants become almost or entirely smooth with compressed whorls and no single character is
diagnostic for this subgenus.

In the circum-Pacific realm,

northern California, Oregon,

and Western

described)
less

(Eiihoploceras)

5.

(?)

Australia.

is

known from

Alberta, southern Alaska

The

5.

sowerbyi Zone

is

(here

more or

strongly indicated at all occurrences.

At Mount Jura in Shasta County, California, 'Stiphromorphites

Crickmay (1933, pi. 28, 7wm. dub.), a probable but
poorly known S. (Eiihoploceras), occurs in the Mormon Formation
below beds with S. (Papilliceras) spp. of the O. saiizei Zone and

schiicherti'

well below Norynannites (Itinsaites).

The
central

best published sequence

Oregon (Lupher,

in the

contains a fauna of large

Supplee area of

east-

The
The Weberg Formation

not yet been described.

ciple section in 1962.

is

but the abinidant new forms have

author visited and recollected the prin-

1941)

S.

of the Colpitts

Group

(Eiihoploceras) with typically straight

strong costae on the outer whorls, evolute Witchellia ('Zugophorites'),

early

cf.

Strigoceras [or? Praestrigites],

cf.

Docidoceras

s.l.,

Bulletin 255

94

Eudrnctoceras}, and abundant pelecypods.

Tmetoceras occurs beneath and steplianoceratids several hundred meters above.

From an isolated exposure of the Fernie Group at Lake Min-

newanka near

Banff, Alberta, se\cral poorly preserved

S.

(?)

(Euho-

which are probably conspecific with the Wide Bay form,

were described and tentatively compared with Soiniinia gracilis
Whiteaves by Frebold (1957a, p. 48, 19, pi. 20) They are probably

ploceras),

also associated with JVitchellia ('Zugophorites') (op.

cit.,

pi.

19, fig.

2).

Another poorly known

(1954, pi. 27;

nam

5.

(Eiihoploceras)

is 'S.

playfordi' Arkell

dub.) from the Newmarracarra Limestone of

Western Australia Avhich was figured in the side view of a single

fragment. However, the material did not justify the naming of a
new species and the sjjecific characters remain dubious. The Australian

.S'.

(Eulioploccras)

is

associated with Pseudotoites, as at

Wide

Bay, an involute JVitchellia and abundant Eontauncsia, and therefor belongs in the

S.

sowevbyi Zone.

Souuiuia (Eiihoplorera.s) closely resembles and

is

directly an-

which has similar almost world-wide

distribution and indicates the O. sauzei Zone, although in Europe
it may appear already in the upper S. sowerbyi Zone. S. (Papilliceras) is characterized by the reduction of costae, the retention or
mature re-appearance of medio-lateral spines or nodes, and the
strongly compressed whorl section. However, the delimitation from
S. (Eithoploceras) has to be based on several characters such as whorl
section, umbilical width and ornamentation; where both subgenera overlap in whorl section, 5. (Papilliceras) is distinguished by
more evolute whorls and somewhat stronger ornamentation (Westermann, 1966, text-fig. 6.)
cestral

to

.v.

(Papilliceras)

Sonninia (Euhoploceras) bifurcate Westermann,

n. sp.

Pis. 23-26;
Text-figs. 27-31

Sonninia sp. indet., Frebold (Alberta), Geol. Sur. Canada,

287, p. 48-49, pi. 20, figs. 1-3 [plastercasts studied].
1965. Sonninia? n. sp. indet., Imlay (Cook Inlet), U.S. Geol. Sur.,
(?)19S7.

Paper 418-B,

Holotype.

p.

PI.

B33,

pi. 4,

figs.

5,

6,

Mem.
Prof.

10-12 [specimens studied].

23, figs, la, b; Text-fig. 30a. Internal

mold

of

phragmocone with one-third whorl of incomplete and deformed

body chamber. Repository: USNM 160236.

Alaskan ammonites,

E
E

Pt. 11:

Westermann

95

30

60

80

20

100

W(mm)

D (mm)

Text-fig. 27.

and whorl

Scatter

section

and growth

{H:W)

of Sonniriia

lines for relative umbilical

(Euhoploccras) bifurcata,

width (U:D)
n. sp., from

the (lower) S. soivrrbyi Zone of Wide Bay; dots and solid lines for phragmocones, circles and dashed lines for body chambers. Note the slightly larger
umbilical width of the mature whorls (60-80
D).

mm

Locus

typicus.

south sliore of

U.S.

Wide

Geol.

Survey Mesozoic locality

12405,

Bay, Alaska Peninsula.

Shnles of upper Kialagvik Formation.

Pseudocidoceras zonule of the 5. sowerbyi Zone, Bajo-

Stratum typicutn.
Age.
cian.

Other occurrences. O. sauzei Zone of the Cook Inlet reS. soiverbyi Zone of Devils Point at Lake Minnewanka
(?)
in Alberta; (?) 5. sowerbyi Zone near Supplee in Oregon.
Derivatio 7iominis. The costae bifurcate commonly on the

gion;

inner whorls.
Diagnosis.

Large,

whorls more or

less

compressed oval

to

subrectangular; phragmocone moderately to strongly evolute with

lateral bullae

and

irregularly bifurcating ribs or with irregularly

on shoulders; body chamber with few strong

somewhat curved simple and bifurcating ribs; simple
straight ribs on large body chambers.
Material. Holotype, two phragmocones with incomplete body
chamber, foia- incomplete phragmocones and one fragment from
fasciculate ribs, robust

rectiradiate or

96

Bulletin 255

1.6

1.5

1.4

'3

1.2

'W
I.I

50

100

D (mm)

200

30

35

40

45

U%

Text-fig. 28.
Scatters and growth lines for whorl sections of Sonninia
(Euhoplocrras) blfurcata, n. sp. (for explanation of symbols see Text-fig. 27).
The whorls tend to grow continuously more compressed, although this is not
necessarily so in every specimen. Right: 'compression' (H:W) of mature
phragmocone whorls {53-92 mm D) is inversely correlated with relative umbilical
width; this is due to "partial logical correlation" since increasing
whorl overlap narrows the umbilicus hut increases whorl height.

1.5

'W

Alaskan ammonites,

Pt. II:

VVkstermann

97

USGS 12405 (USNM ir)02-58, 160239, l()()211); two phragmocones

with incomplete body chambers and two fragments from USGS
19869 (USNM 160240, 160242); several fragments from USGS
19862 and 19870; three moderately preserved specimens from

1-6

one small nuclens from about 16 m (McM. J 990)

and one almost complete internal mold from scree (McM. J 975)
of
10; 12 mostly incomplete or fragmental specimens and
several body cliamljer fragments from
13w; four partly deformed fragments from 3 m, 10 m and 17 m of
15.
Description. The nucleus is incompletely known and only
visible laterally in the umbilicus of a spinose specimen (McM.
diaJ 990) Lateral costae are already present at about three
meter consisting of irregularly strong fasciculate primaries. Strong
bidlae-like lateral spines, seven to eight per halfwhorl, are developed
at about 16
diameter.
The few preserved juvenile stages (10-20 mm) have somewhat
depressed oval whorls with narrow and low sharp keel which, at
least in the internal mold, is accompanied by narrow sulci. The ornament consists of strong blunt bifurcating costae which extend in
full strength up to the sulci, and usually of heavy somewhat bullae-

(McM.

J 981)

WA

WA

WA

mm

mm

like lateral spines.

The

adolescent whorls range from subcircular to moderately

compressed oval

(H/W

1.1

widely evolute (U := 30-40%)

1.5)

The

and from moderately

cept for the compressed weakly ornate variants.

ants possess few strong lateral spines

to

sulci are usually retained ex-

The

spinose vari-

on high primaries and usually

have some weak blunt intercalated primaries without spines. The

major primaries bifurcate or tri furcate at about three-fifths whorl
height and some secondaries are intercalated. All secondaries project

somewhat and retain

their full strength or swell to

form weak

bullae on the shoulders; they are abruptly truncated at the sulci.

The

spinose stage,

of the

if

present,

may be

phragmocone. More commonly,

retained to almost the end

at 25-50

mm

diameter, the

become bidlae. With further growth, spines or bullae finally

weaken and may become obsolete. The nonspinose variants bear
blimt bullae often arising from simple fasciculation, and irregular
much weaker intercalated ribs; other specimens have only weak
spines

ribs

with a few stronger ones irregularly intercalated.

Bulletin 255

98

The last whorl of the phragmocone (80-150 mm D) and the

body chamber (120-250 mm D) are moderately compressed ellip36-41%). The
tical to ovate (H/^V 1.4
1.5) and evolute (U
body chamber egresses markedly only in the more involute forms
resulting in similar body chambers with about 40% umbilical
width. The externside of large phragmocones is rounded, often
flattened, and carries a moderately high narrow keel. Rarely de-

veloped shallow

probably restricted to the internal mold.

sulci are

Tlie stronger primaries and lateral bullae of the body cham-

ber or the lateral bullae become more regular and continue into
one of the secondaries, usually the posterior one, resulting in
single long somewhat ciuved ribs. The remaining weaker secondaries are retained for some time. On most body chambers and also
on some large pliragmocones, the single long ribs strengthen and
become more or less rectiradiate, rarely typically rinsiradiate, and
somewhat projected on the shoulders, while tlie secondaries fade.

The

thin keel

sembles

.S'.

is

As usual

The

proximated

last

adult stage almost perfectly

(Waagen)

re-

larger Sonniniidae

in

served; adulthood

The

retained.

(EtiJiopIoccras) adlrra

is

two

(9), apertiues are not preindicated only by several specimens Avith ap-

to three septal sutures at 85-140

mm

diameter.

moderate complexity and nonsuspensive. The saddle boundary may even be slightly reclined ventrally.

sutiue

septal

is

of

Tlie saddles decrease gradually in

size;

the lobes are slender

trifid.

Discussion.

Wide Bay form

Tliere
a

is

single

no doubt

that all

morphological

S.

(Euhoploccras) from

series,

tiieir

duration

was limited, and that they, therefore, belonged to a single palaeospecies. This is evident in the few larger samples from localities
USGS 12405 and
13w. The latter outcrop includes only a
few meters of the total vertical range of 30-50 m. No evolutionary
trend is apparent throughout the range; however, the compilation

WA

of the section

Of

is

admittedly inaccurate.

this supposed single palaeo-species is the

morphologic inter-relation (co-variation) of evolute roiuid whorls
with strong spinose costae, and of more involute compressed whorls
with fine weak ornament (Text-fig. 29) This is in fidl accordance with the " (1st) Buckman Law of Co-variation" (Westermann,

interest within

Alaskan ammonites,

which was

1966)

Pt.

II:

Westermann

described from the closely related

first

Waagen and which applies to the

known species of ornate ammonites.

ploceras) adicra
if

not to

all,

Two

Coruparison.

incomplete specimens of

(Euho-

this species

were

(1964, p.

from tlie Tuxedni FormaMountains and from the Red Glacier Forma-

10-12), respectively

pi. 4, figs. 5, 6,

tion in the Talkeetna

on the Iniskin Peninsula, both

tion

S.

great majority,

recently described as 'Sonninia? n. sp. indet.' by Imlay

B33,

99

in the

Cook

Inlet region of

southeastern Alaska. Significantly, the former occurrence belongs

and the

possibly

latter

probably in the O. sauzei Zone.

Probably identical with

this

species or closely affiliated are

the three poorly preserved specimens from Devils Point at

Minnewanka, Alberta, which were

Frebold (1957a,
only 80-90

mm

pi.

in

20, figs.

1-3)

Lake
and figured by
indet. Although

briefly described
as Sojiniuia sp.

diameter and probably incomplete or immatme,

they show a similar variety of ornament and apparently also of

coiling

and whorl

section.

Also from Devils Point are the poorly preserved type speci-

mens

of 'Schloenbachia' gracilis A\'hiteaves

under Sonninia

gracilis

and

which was redescribed

5. adicra by Frebold

said to be related to

fig. 1)
The two plastotypes, kindly furnished
Survey of Canada, are distinguished by the
much more evolute whorls and the regular simple straight ribs with
high-set nodes or spines on at least the last three whorls. 5. gracilis

(1957a, p. 48, pi. 19,

by

belongs either to

genus

Geological

tlie

S.

5.

(Euhoploceras) or to the

new Alaskan

sub-

(Alaskoceras) described below.

related form

is

the poorly

known

'Stiphroinorphitcs schii-

Mount Jura in
(1933,
northern California which was based on a single poorly preserved
specimen not available for resttidy. This specimen appears to dif-

cherti'

fer

Crickmay

from

S.

bifurcata

pi.

in

28; noyn dub.) from

the shorter secondaries

which die out

gradually leaving a broad smooth band beside the keel.

The large S. (Euhoploceras) from Supplee in east-central Oregon are too poorly preserved to allow specific identification but
they resemble the Alaskan species more closely than any other
known species.
The most
probably

S.

closely related

playfordi Arkell

named but
(1954)

poorly

known

species

is

from the Perth area of West-

Bulletin 255

100

a-b.
Cross-sections of Sonninia (Eu/ioplocrras) bifurcata,
Holotype, body chamber somewhat distorted, b. Complete phragmocone of compressed and weakly ornate variant, loc. USGS 12405 (USNM
160239).

Text-fig.

n. sp.,

1-

30

a-

it is said to differ from S. adirra chiefly

which are more robust on the shoidder. Howthe costae of this form are much more densely spaced tlian in

era Australia, because

in the secondaries
ever,
S.

bifurcata consisting chiefly of simple primaries.

S.

bifurcata

is

distinguished from

many synonyms; VVestermann,

especially

more

1966)

.S'.

in

adicra

Waagen (and

on the shoulder of spinose variants and

distant primaries witli

tained bifurcation.

its

the stronger secondaries

stronger developed

in the generally

and longer

re-

Alaskan ammonitks,

Pt. II:

Westirmann

Text-fig.

31.

Cross-section

101

of

Sonn-

inia (Eu/ioplocrras)

bif areata, n. sp., fragment of large body chamber, left side

reconstructed, loc.
13\v
J

WA

1048)

Measurements.

(bod}' ch.)

(phragm.)

USNM
USNM
USNM

85

(phragm.)

92

160241

(body ch.)

100

(phragm.)

70

(phragm.)

W%

106

160239

160238

38

61

39

USNM

160240 (body ch.)

110

(phragm.)

80
55

USNM

160242 (body ch.)

McM

J 975

McM

J 981

(McM

1-

Dmm

holotype

c.l30 c.34

(phragm.)

53

(body ch.)

C.120

(phragm.)

79

(phragm.)

150
105

H%

U%

H/W

Dsp*mm

Bulletin 255

102

Sonninia (Euhoploceras?)

sp. indet.

PI. 27, fig. 1

Material. One internal mold with test remains of one-third

whorl phragniocone with remnants of inner whorl and one crushed
internal mold of two-fifths whorl body chamber, probably with
(MciM. J 959a, b) Upper E.
aperture, from
8, upper part
arnplectens zonule of the basal S. soxccrbyi Zone, Kialagvik Forma-

WA

Wide

tion,

Bay.

Description.

The

phragniocone of approximately 70

mm

dia-

meter is moderately evolute with compressed subrectangular section and a thin low hollow-floored keel. The costation is extremely
Avcak. blunt, irregular, and restricted to the sides consistins; of broad
straight or

somewhat curved swellings which fade on the rounded

umbilical margin and

shoulder.

tlie

The

septal suture

is

highly

complex and nonsuspensi\e with straight umbilical elements.

Tlie body chamber of apjjroximately 95 mm diameter is evolute, smooth, and probably compressed elliptical to subrectangular in

section witii

Measurements.

faintly

keeled venter.

W%

Dmm
J 959a

(phragm.)

H%

c.28.5

c.70

species.

Diagnosis. An

S.

alaskcnsis

Westermann,

n.

n.

subgen.

sp.

advolute mediimi-sized subgenus of Sonninia

inverse-trapezoidal

witli

(c.26)

c.39

Subgenus SONNINIA (ALASKOCERAS) Westermann,

Type

U%

whorl-section

(diverging

flanks)

pri-

and rursiradiate with regular prominent ventrolateral spines retained on body cliamljer; secondaries obsolete.
Age. S. soiverbyi and O. sauzei Zones, Bajocian.
maries

bliuit

Southern Alaska (+?? Alberta).

S. (Alaskoceras) shows close affinities

i)f.y/)77;;///o/?.

Affinities.

to the spin-

and widely imibilicate variants of the Eiuopean S. (Euhoploceras) adicra (Waagen)

196f)), such as to the
(cf. Westermann,

ose

specimen figured by Oechsle [195S,

pi.

15, fig. 3; as S.

polyacantha

(Buckman) ] from the Swabian Jura. Both subgenera

have evolute whorls and a small hollow-floored keel set on a weakly

crassijorjnis

roimded, sometimes slightly bisulcate venter, simple long, often

rursiradiate primaries with terminal spines which may be molded

Alaskan ammonites,

in

the subsequent

(Alaskoccras)

is

Pt. II:

umbilical slope,

Westermann

103

and reduced secondaries.

S.

distinguished by the inverse-trapezoidal rather than

subcpiadrate whorl section, with diverging gently convex flanks,

and the stronger more regular and more ventrad spines which are
on blunt, often almost obsolete, strongly rursiradiate costae.
There is little resemblance between S. (Alaskoccras) alaskensis, n. sp.
and the often associated S. (Eulioploceras) bifiircata, n. sp. dining
any morphogenetic stage.
Some resemblance exists to ZiircJicria Douville which may
have nodes or small thorns developed on the shoulders, such as in
Z. paiuispinata Buckman and Z. inconstaus Buckman from the L.
discitcs Subzone of Dorset, and (?) Z. pertinax (Vacek) from the
Alps. However, in Zurcheria the spines or tubercles are weak, a
keel is absent or obsolete, and the whorls are less evolute and more
set

compressed.

Another sonniniid with ventro-lateral nodes is Haplopleiiroceras Buckman which also agrees in the advolute coiling. It is
clearly distinct from S. (Alaskoceras) in the subquadrate whorls,
the sharp strong prosoradiate costae, the presence of a second line
of lateral nodes (with the apparent exception of

and

of single secondaries,

and

H.

tobleri

Renz)

in the blunt probably solid keel.

Sonninia (Alaskoceras) alaskensis Westermann,

n. sp.

PI. 27, figs. 2-7,

Text-figs. 32-33
?1964. Sonninia cf. S. nodata Buckman, Imlay (Cook Inlet), U.S. Geol. Sur.,
? n. subsp.]
Prof. Pap. 418-B, p. B33, pi. 2, figs. 1, 2 [Specimen seen;

Holotype. PI.

27, fig. 3a-c; Text-figs. 32a, 33a. Internal

mold

remains of complete phragmocone and distorted fragments of body chamber. Repository: McM. J 1021.
Locus ty picus. \^oc^\\iy
10 (scree), south shore of Wide
Bay, Alaska Peninsula.
^vith

test

WA

Stratum typicum.

Pseuclocodoceras

zonule of upper Kialag-

vik Formation.

Age.
sp.)

(Lower)

5.

sowerbyi Zone

(-J-?

O. sauzei Zone;

n. sub-

Bajocian.

Distribution. S. soiverbyi Zone at Wide Bay; ?

Zone of Talkeetna Mountains, Cook Inlet region.

Derivatio nominis. Self-evident.

Diagnosis.

As

for subgenus.

also O. sauzei

Bulletin 255

104

Cross-sections

of Souninia (Alaskocfras) alaskensis,

Composite: phragmocone of hoiotype, body chamber
from loc.
13\v
McM J 1023), both Pseudocidocnas zonule, b. Part of
phragmocone at 46 mm I), loc.
15 (McM J 1027; cf. pi. 27, fig. 7). Note
the hollow-floored ventro-iateral spines up to more than 11 mm in length
which were developed from the matrix.
32

Text-fiji.

n.

subgen.,

n.

sp.

WA

a-b.

1-

a-

WA

Mdlcrial. ^'\\e liolotype, one almost complete specimen with

body chamber, and two fragmentary specimens from scree of
10 (McM. J 1021 a) one large plnagmocone from the basal 10 m
of
10; one phragmocone fragment from scree of
13; two
13w (McM. J
poorly preserved incomplete specimens from
two almost complete specimens and one phragmocone from
1023)
(McM.
3 m
15; two almost comJ 1027), 23 m and scree of
specimens
and
one
damaged
phragmocone
from USGS 12405.
plete
Pseudocidoceras zonule and subjacent (?) shales, Kialagvik For-

WA

WA

WA

WA

WA

mation, southeast shore of

Description.
1021 a)

(U

The

Wide

juvenile

Bay.

whorls

30%). During

the next one to

(G-IO

mm

D;

McM.

and moderately evolute

two whorls (15-20 mm D)

are depressed elliptical in section

Alaskan ammonites,

Pt.

II:

Westermann

105

Text-fig. 33 a-b.
Septal sutures of Sonninia (Alaskoceras) alaskcnsis, n.
subgen., n. sp., enlarged, a. Holotype at 17
D, approximated at
H, 50
one-sixth whorl before phragmocone end and probably 'senile', b. Specimen at
18
H, loc.
13
J 1029).

mm

mm

mm

(McM

the whorl section becomes inverse-trapezoidal by

flattening

and

diverging of the flanks and by flattening of the broad venter, and

much more evoliite (U ^ 40-42%) The exmay be slightly bisulcate and carries a small hollow-floored
The ornament commences at two to three mm diameter with

the coiling becomes

ternside
keel.

heavy somewhat fasciculate and probably partly nodose primaries.

At about five mm diameter, lieavy foldlike rectiradiate costae (five
to seven per halfwhorl) develop which terminate on the shoulders
of the internal mold in thick knobs marking the base of the prominent hollow-floored spines of the shell. Beyond 10 mm diameter,
blunt strongly projected secondaries, about two for each primary,
are present beside the venter and the primaries become blunter
and rursiradiate. Weaker primaries without spines, which are so

common

in

S.

(Eiihoploceras), are missing or rare.

Throughout

the

adolescent and mature

whorls are advolute, "riding" with only about

growth

10%

stages,

the

overlap on the

shoulders of the preceding whorl, so that the spines

may be molded

The

diverging flank

into the subsequent shallow umbilical slope.

is

only weakly convex from the umbilical seam to the shoulder and

may become

almost smooth on the inner half. Blunt rursiradiate

primaries arise mid-laterally and strengthen only slightly toward

Bulletin 255

106

mm

in diaThe internal molds have large (up to 6

knobs on the shoulders which in test preservation carry

the shoulders.

meter)

ventrad inclined, extremely strong,

to nine per halfwhorl.

served length which

is

hollow-floored

spines,

eight

mm in preThe longest spines measure

as much as the corresponding whorl height
1 1

and width. The venter of

sulcate and smooth, while

mold

the internal
tlie

shell

is liere

is

usually weakly bi-

flat,

with or without

shallow sulci, or weakly convex carrying a narrow and low hollowfloored keel. The floors of spines and keel are retained up to and
including the first one-fourtli whorl of the body chamber. At the
end of the phragmocone, the whorl-section may become subquadratic, as in the holotype. On the last wliorl of the phragmocone,
weak intercalated primaries without spines and densely spaced
blimt and stiongly projected secondaries are often present.
The body chamber, probably tliree-fifths to three-fourths whorls
in length, is advolute (U = SO^j,) and, therefore, often exfoliated
during fossilization. The flanks remain flattish and divergent but
the venter becomes moderately convex. The thin weak keel and
the prominent spines are retained to the apertine. The primaries
are even more reduced and often become obsolete. Superficially,
fine growth lines or lirae may now be present on the flanks. Blunt
secondaries are usually developed and can be seen weakly also on
the internal mold. The form of the peristome is unknown but it
was probably simple according to the almost straight growth lines
towards the end of the body chamber.
The septum shows an almost cruciform fluting pattern which
is in accordance with the advolute, about as high as broad whorls.
This is simply a modified plano-disculate type, similar to the pattern observed in the Alaskan S. (Enhoploceras) and Witchellia.

The

septal suture

the evolute
boiveri
figs.

(Buckman)

as recently figured

except for

193-197)

elements of

(Text-fig. .S3a-b)

roinid-whorled Sonuinia

tlie

resembles closely those of

piuguis

(Roemer)

and

by Schindewolf (1964,

more reduced

internal

S.

text-

and umbilical

alaskenis; only three full

umbilical lobes are de-

veloped wliile there are normally four or

five in typical Soyiyiinia,

tiiis

.S'.

reduction of imibilical elements being interpreted as the

sult of advolute coiling.

ed Uj

is

asymmetrically

As

re-

in otiier sonniniids, the internally locat-

trifid to

almost bifid; Uj and

U.j,

the ex-

Alaskan ammonitf.s,

Pt.

narrow and deep;

ternal counter-part, are unusually

L may

be asymmetrical tending to be

even considering the relatively small

conchs

Westermann

II:

bilid.

107

also Uv.

Tlie complexity

size ot these

is

and
low

piobable macro-

(females)

Affinities.

The

affinities to other

well-known sonniniids have

already been discussed inider the subgeneric heading; the closest

known

relative of 5. olaskensis

such as the European

-S'.

is

from the Alberta

among

probably

adicra (Waagen)

S.

(Eiihoploceras),

Souniiiia gracilis (Whit-

is distinguished
by the more
more regular costae, and it appears
that at least the inner whorls had tubercles high on the flanks or
shoulders, that the venter of the phragmocone was weakly bisulcate and the whorl section subrectangular to subsquare. S. gracilis
thus seems intermediate between S. (Eiihoploceras) and 5. (Alasko-

eaves)

foothills

evolute whorls with stiffer and

but probably closer to the former, particularly resembling

ceras),

evolute variants of

5. (E.)

adicra as figured by Oechsle (1958,

pi. 15,

fig. 3)

Of

particular

interest

scribed as Sonninia
pi. 2, fig.

1,

2)

cf. 5.

the

is

yiodata

single

specimen recently

Buckman by Imlay

de-

(1964, p. B33,

from the O. sauzei Zone of the Talkeetna MounAlthough certainly a close ally of

tains of southeastern Alaska.

5.

alaskensis,

it

is

distinguished in the regularly present relatively

strong and projected minor costae between the widely spaced spine-

bearing primaries. In the absence of a larger sample, the

tion of this form as either a
lated species of
this

is

5.

new chrono-subspecies

(A.) alaskensis

is left

an extremely ornate variant of

Measurements.

It is

improbable that

alaskensis.

Dm

Holotype (phragm.)

open.

5.

classifica-

or closely re-

W%

H%

U%

P( spines)

Bulletin 255

108

Genus WITCHELLIA Buckman, 1889

Type
1

species.

Aiinnoiuti's

laeviusciilus

].

de

Sowerby,

C.

S24.

The genus was named (Buckman,

monites,

p. 82,

footnote

1)

1889,

Inferior Oolite

Am-

for sonniniids with "slight carina [hol-

low-floored keel] bordered by two

fmrows

on

\vhich are not seen

the test." Also originally included in WitchelUa were 'Am.'

siitiieri

Branco, 'Am.' jugijer Waagen, and 'Am.' deltafalcattis Quenstedt.

few years

later,

Haug

(1893)

placed

IT.

sutueri,

W.

lae-

punctadssima Haug in a single WitchelUa species

group. He, however, also includetl in Witcliellia the younger
species grouj^s of Am. romani Oppel and of Arn. edoitardianiis
viuscula,

and

IT.

d'Orbigny which Buckman had just separated inider Dorsetensia,

as well as his group of II'. sayni Haug which Buckman split into
various other 'genera.' The ensuing confusion has at length been
discussed by P. Dorn (1935, p. 93) Spath (1936, p. 5) Hiltermann
(1939, p. 21), and Oe(hsle (1958, p. 77) and need not be brought
up again. The last two authors concluded that WitchelUa cannot
reasonably be separated from Sotnnriia. The distinction of Dorsetensia from WitchelUa is similarly difficidt and probably arbitrary.
In the Treatise, Arkell, et al. (1957, p. L 270) upheld again
the genus Witcliellia with the diagnosis: "Inclined to be involute
compressed, whorls heightening and smoothing early, long before
,

septation ceases; venter narrow, tabulate and carinate,

commonly

even tricarinate, or becoming fastigate; nucleus as in

Soyiniiiia" ; included in the synonymy were Zugophorites, Sonninbisulcate,

ites,

Gelasinites, Dunclryites,

gella, all of

Buckman, 1922-27

Lectotype of the

Riibrileiites,
(see

type species.

lectotype of IF. laeviuscula

(J.

and

7ai-

author examined

the

Anolkoleites,

below).

The

de C. Sowerby, 1824,

pi.

451,

fig.

1;

Buckman, 1908, pi. 5, fig. 1; Arkell, 1956, pi. 34, fig. 1)

British
iMuseum (Nat. Hist.) (cat. No. 439 50 a) On request
in the
by the author, H. K. Howarth developed the left umbilicus and the

refigured:

specimen

is,

therefore, figured again

(Text-fig. 35)

Sowerby's small 'topotypes' from Dinidry, Somerset, are

all

immature specimens and probably belong to the Hildoceradidae, such as Esericeras, and Hammatoceratidae of the Ujjper
Toarcian and Lower Aalenian. The paratype is described below.
nuclei or

Alaskan ammonites,

The

mm

Pt. II:

Westkrmann

109

is completely septate and

Another one-quarter septate whorl is
preserved near the imibiliciis whicli, according to the egressing umbilical seam, represents the end of the adult phragmocone. At
68 mm D, the whorl height is 33 mm, the whorl width 19.5 mm,
and the umbilical widtli 12 mm. There are seven primaries and
23 secondaries on the ultimate one-half whorl. The inner whorls
(< 45 mm D) are much more evolute. The flanks of the nucleus
(20-25 mm D) slope gently to the umbilical seam but curve more
strongly on the intermediate whorls reaching the seam perpendicularly. At the beginning of the ultimate whorl, the flanks converge gently, curving somewhat more strongly besides the keel without forming marked shoulders. The internal mold is closely similar. Subsequently, however, the coiling becomes more involute; the
umbilical margin, the steep slope, and the marked but weak shoulders and sloping narrow ventral bands of the adult shell develop;
the keel is now strong, high, and hollow-floored. The internal mold

lectotype, 68

in diameter,

largely preserved with test.

is

typically tabulate-unicarinate with moderately sharp shoulders

and

low blunt

The

keel;

however, sidci are absent.

costation of the inner evolute whorls, as seen in the de-

veloped umbilicus,

is

irregular,

bullae of different strength.

bearing small

ultimate phragmocone whorl has

markedly falcoid principal

distant blunt, slightly to

composed

The

partly fasciculate,

ribs

which are

of short prosoradiate primaries curving into single rela-

The

tively strong secondaries.

although modest, strength

greatest,

reached on the middle of the flanks. Other

arise at first

by fascicidation and

tion. All ribs project strongly

later

much weaker

is

secondaries

by furcation and intercalaa short distance from the

and die out

keel.

The

septal suture

is

partly preserved.

The E/L and L/Uo

moderately wide

saddles are of similar size

and weakly

and symmetrically

with rather thin long stems.

bilical lobes are

trifid

frilled,

is

The um-

nonsuspensive.

Other specimens of the type species. The paratype of W.

de C. Sowerby, 1924, pi. 451, fig. 2; Buckman, 1908,
(J.
pi. 14, fig. 2)
came also from Dundry in Somerset. This small
specimen of which the ultimate egressing
D)
evolute
(22.5
one-third whorl appears to belong to the adidt body chamber, may
laevixiscula

mm

Bulletin 255

110

be a microconch. However, there is hardly any way of comparing

this specimen with the much larger lectotype except for the inner

While the

flanks of the inner whorls exposed in the umbilicus.

lectotype

is

deeply unibilicate and tuberculate at comparative

the paratype

is

strongly compressed

and

size,

only weak fasciculate

lias

The paratype becomes tabulate (l)ut not sulcate) only on

body chamber and has a sharp thin keel.
Significantly, Biickman placed in IT. Uwx'iusi ula also more evolute and typically tabulate to bisidcate large sonniniids. However,
these specimens came mostly from Oljorne in Dorset. One of these
specimens (Buckman; Type Ammonites, pi. 745) which was refigcostae.

the

med

in tlie Treatise (Arkell, ct al., 1957, p.

269) closely resembles

the lectotype except that the venter of the internal

edly bisidcate.
collection

My

mold

mark-

is

study of unfigmed specimens in the

Buckman

Museum, London, U.K.)

indicates

(Cieological Survey

an intergrading morphological sequence from

cate to tabulate internal molds and that, consequently, the
that there

type

is

a marginal variant of a single, usually

more or

bicarinate species. This might already have

Buckman

bisid-

is

(1920,

Type Ammonites,

168)

pi.

less

lecto-

strongly

been recognized by

as suggested

by

his in-

valid determination of IF. spinifera as substitute "genolectotype."

IF. spinifera,

here placed in synonymy with

on an almost complete,
specimen which appears

to

based

IF. laexnusciila, is

and markedly

relatively small,

be close to the

mean

bisidcate

of the species

variation.

Buckman's

collection

also

variation from the involute

IF',

suggests

that

there

is

continuous

laeviuscula morphotyjje to the

more

evolute and, as usual, stronger ornate forms. Thus, IF. plalymorpJm

Buckman

(192())

is

considered another variant of

IF.

laeviuscula;

mold is bisulcate-tricarinate on the phragmoconc and strongly tabulate on tlie l)ody (haml)er. At about 70
nun diameter, the uml:)ilicus suddenly narrows fiom 28% to 22-2 1%
tlie

venter of the internal

so that the adidi sjjccimen resembles

Vet the inner wliorls match

specimen

liad originally

Witcliellia sutneri

5,

fig.

2;

lectotype in this respect.

(Branco)

with which the

been identified by Buckman.

(Branco) . The author lias searched for

the holotype which apj^ears to be

pi.

tlie

IF. sutneri

lost.

IF.

sutneri

reproduced here in Text-fig. 34)

(Branco, 1879,

came from

the

5.

Alaskan ammonites,

Pt.

II:

Westermann

11

Text-fig. 34.
Holotype of Witchcllia sutncri (Branco), reproduction
of
original figure (Branco, 1879, pi. 5, figs. 2 a-c,
5a), St. Quentin, Lorrain; apf
prox. XI.
<

Lectotype

of Witchellia laeviuscula
(J. de C. Sowerby)
f
'^^u-i^'^"
eft umbilicus newly developed, Inferior
Oolite, Dundry, Somerset. Note irregular fasciculate costae and lateral tubercles
of nucleus visible in umbilicus;
35.

Bulletin 255

112

O. saitzcl Zone of

soii'crbyi or

France, and was

St. QucMUin near Metz in Lorraine,

amply described. Branco listed the following mea-

mm,

surements: estimated complete diameter 100

meter

mm, whorl height

18.5 mm. The wliorl

61

witlth

the iigmes

and

is

said to

whorl, with steep umbilical wall grading into

sulcate

The

figured \\ith short

U^

umbilicus.

as in

the inner whorls resemble

cornigata

(Waagen)

and

sides

bi-

mm

costae are said to originate simply or in pairs from

small nodes near the

S.

flat

venter beyond 20

(bisculcate-lricarinate in drawing)

diameter.

preserved dia-

mm.

whorl width 17.5 nnn, umbilical

section is less compressed than shown in
be scjuare except for the ultimate half27

(Sowerby)

According

to

The

simple septal suture was

Sonninia soiverbyi. According to Branco,

gingcnsis (Waagen) [? a synonym of
and the penultimate whorl 5. adicra

.S'.

drawing, the ultimate halfwhorl of

tlie

the holotype has seven bullae-like primaries

and 22 strongly

pro-

jected secondaries. At least toward the end, the primaries continue

one of the secondaries which are, therefore, of imeven

strength. The missing last whorl was apparently evolute, accordinto

ing to the slightly egressing umbilical seam.

Tlie study of Buckman's collection of evolute Witchellia from

Oborne in Dorset
are members of a
conspecific with
\V.

by

suggestefl to

me

strongly that

all

alleged species

single intergrading morpfiological sequence

siilneri

\V.

(Branco).

IV.

falcata

and

Buckman and

actinophora Buckman, 1926, had originally l^een so identified

Buckman

(1889),

and

JV.

glauca

Buckman

(1925)

was

distin-

guished because of the presence of irregular nodes only.

Species group, plexus or superspecies
ueri.

Finally

W.

laeviuscula

W.

sul-

even the separation of the involute W. laeviuscula-

type forms from

tfie

largely contemporary,

more evolute W.

type forms could be purely arbitrary. In 1893

Haug

sutneri-

(p. 303)

spoke

of the total morphological intergradation in the Elsassian occur-

and the material from the IV. laeviuscula Subzone (upper S.

Zone) of Frogden Quarry near Oborne in Dorset appears
to support this opinion. However, the evidence is still incomplete
particularly because most specimens consist only of phragmocones
and vary greatly in size, their exact morphogenetic stage and the
ajjertuies being unknown. The similar size range of the involute
and more evolute forms speaks against their interpretation as corrence,

soxi'crbyi

responding sexual dimorphs.

Alaskan ammonites,

this

It

mainly

Pt. II:

northwestern

Witchellia 'species group'

Westermann

European

113

(England,

France)

arranged into a morpliological

is

se-

becomes obvious that coiling, whorl section, and costation

co-vary, i.e. ranging from involute, finely ornate, and tabulate to
evolute, coarsely ornate, and bisulcate forms (Text-fig. 39)
This
cjuence,

it

sequence of alleged

'species'

is:

W. patefactor Buckman, 1923

W. laeviuscula (J. de C. Sowerby) 1824
W. spinifera Buckman, 1920
W. platymorpha Buckman, 1925
W. actinophora Buckman, 1926
W. glauca Buckman, 1925
W. falcata Buckman, 1926
W. sutneri (Branco) 1879
It

appears advisable for the time being to distinguish the two

W.

laeviuscula and W. sutneri and to regard the other

synonyms by arbitrary central division of this morphocline
between 'W. platymorpha' and 'W. actijiophora'.
The W. laeviuscula - W. sutneri species group, plexus or superspecies was abundantly known from southern England and eastern
France where it marks the upper S. sowerbyi Zone, i.e. the W.
laeviuscula Subzone. TF. sutneroides, n. sp. from southern Alaska
species

names

is

as

also a

member

of this 'plexus.'

Older Witchellias.

The

middle
England
and were named Zugophorites zugophorus Buckman (1922) and
These generic names are
Gelasinites gelasinus Buckman (1925)
certainly synonymous with Witchellia and both forms may well be
conspecific. There is a possibility that they are conspecific with
TF. sayrii Haug (1893, p. 309; for Ludwigia corrugata Douville,
non Sowerby) from eastern France. Zugophorites [syn.: Gelasi'nites] is at best a subgenus of the slightly younger Witchellia described above from which it is distinguished by the wider umbilicus and the more regular costae without bullae-like inflated
primaries. W. ('Zugophorites') was probably directly ancestral of
the closely related and only slightly younger IF. laeviuscula - W.

5.

sowerbyi Zone, the

S.

first

witchellias occur in the

trigonalis Subzone, of southern

sutneri 'plexus.'

Related genera.

Similar

taxonomic problems

as

discussed

Bulletin 255

114

above arise on the generic level. Many authors working on this

group have concluded that tliere are no distinct morphological
limits between WitcJiellia and Sornilnia on the one hand and between M'itchcUid and Dorsctcn.sia on the other.

The

WitchclUa ol several moderately involute,

discoidal, and tastigate to weakly tabidate genera

inclusion

more or

less

in

mainly from the O. sanzci Zone, such as in the Treatise (Arkell,

probably does not reflect phyletic relationet al., 1957, p. L 270)
ship. These forms miss the I^isidcate or at least clearly tal)tdateimicarinate venter of Witclicllia and have the sharp umbilical mar,

gin and often the simple costation of Dorsetoisia.

It

appears totally

Buckman
Buckman (1926)

arbitrary to place the tabulate-unicarinate 'Hyaliuitcs'

(1921) in Dorsrtcn.sia but the similar 'Dutidyyites'

in Witclicllia simjjly because

The

discoidal 'Riibrileiites'

tlie

latter

Buckman

is

slightly

more

involute.

can hardly be

(1926)

tinguished from Dorseteri.sia of the D. tecta-subtecta group.

ever, 'Riibrileiites'

dis-

How-

and 'Dundryites' bear the irregidar primaries

Witchellia. 'Anolkoleiites'

Buckman

(1926)

of

an extremely invol-

is

ute and compressed form of this same group.

Buckman

Sonninites

(1923)

is

based on a large phragmocone

of the compressed, moderately involute, almost fastigate

S.

felix

whorls

Buckman

and smooth

wliich has a sharp umbilical margin; the inner

("paratype")

are

keeled without

tabidation

and almost

evenly and simply costate, except for the nucleus of 20 nnii dia-

meter which shows fasciculation

.siniulafis
.S".

celans

as in Witchellia

and Sonninia;

S.

Buckman has irregular nodes on the nucleus; however,

Buckman is a typical oxycone. It appears advisable, there-

fore, to retain

Sonninites as a subgenus (or

genus)

for this inter-

mediary group whicli jjossesses adolescent and adult cliaracters most

similar to Dorsetoisia hui retains the juvenile whorls of Witchellia
(}

and Soiininia or

Souiiinia).

lussilobiceras

Buckman

(1919)

is

distinguished by greater inflation and the complex septal sutine.

The

intricate relationship of WitcJiellia with typical Souniiiia

concerning especially the W. laerniiscula

noted especially by Hiltermann

(1939, p.

W.

siitneri

125)

'plexus'

was

and supported by

Oechsle (1958, p. 77) There is no single diagnostic featiae separating them. 'Typical' WitcJiellia costation occins in Sotuiinia corru.

gata

(Sowerby)

5.

alsatica

Haug, and other

species of Sonninia.

Alaskan ammonites,

Pt.

Wkstermann

II:

115

Wliile many Sotniitiia species become smooth, costation may persist

onto large body chambers in Witchellia such as in the Alaskan
species described herein; strong infraspecific variation in the dia-

meter of the costate stage

may

venter of Witchellia

is

often apparent. Even

the bisidcate

be present at least on the inner whorls of

Sonninia.

Dimorphism.
nevi 'plexus'

The

described ]Vitchcllia lacviuscula

W.

sut-

apparently totally macroconchiate (female), with the

possible exception of 'IF. glauca' Ruckman which is fully grown at
75

mm

is

may be

diameter and

Buckman,

identical with 'Spatulites' spatians

a microconch of 59

mm

diameter bearing prominent

lappets.

The corresponding microconchs (males) to 'typical' Witchellia

were described under the generic names of Pelekodites, Naymoceras
Buckman, 1923, and especially under Maccratites and SpatuBuckman, 1928, from the stratotype and type locality of IF.
laeviuscula. All were combined into the single genus Pelekodites in
the Treatise (Arkell, et al., 1957, p. L 270) which includes most
(?)

lites,

lappet bearing sonniniid microconchs.

The
from the

type species of Pelekodites, P. pelekus

-S'.

trigonalis

dry, Somerset. It

a small,

is

slightly depressed

S.

Buckman,

1923,

is

sowerhyi Zone) of Dun-

moderately evolute microconch with

whorl section, weak only slightly fasciculate rur-

and

siradiate costae,

Subzone (middle

lappets,

whorls of the macroconchiate

and
IF.

it

appears to match the inner

('Zugophorites') from the same

subzone. 'Nannoceras nannomorphiim' Buckman, 1923,

is

probably

somewhat stronger ornate variant of P. pelekus. 'Spatulites' Buckman, 1928, with the only species S. spatians Buckman, 1928, and
the almost identical 'Maceratites' Buckman, 1928, with the only
species M. aurifer Buckman, 1928, came also from the IF. laeviuscula
Subzone of Dinidry. They are more compressed and more strongly
costate than Pelekodites, with irregular fasciculation and some bullae. There is close correspondence to the associated macroconchiate
Witchellia laeviuscula IF. sutneri 'plexus.' 'Spatulites' and 'Maceratites' are certainly not distinct even on the subgeneric level and
a

both type species

may be

conspecific; 'Spatulites' spatians has

page

priority.
If

Zugophorites

of Witchellia,

it

may

[syn.: Gelasinites]

is

distinguished as subgenus

be advisable to classify the possibly correspond-

Bulletin 255

116

ing respective mitroconchs Pelckodites and Spatulites at the same

and to place all three in Witchellia. However, microconchs

and inner whorls ot niacroconchs are in need of reinvestigation
and tiie specific dimorphic correspondences, especially of typical
Pelekoditcs, are still poorly known. If one places Sonninia and
level

WitcliclUa in a single genus, as a

kodites
little

number

of colleagues prefer, Pele-

also to be included in Sonninia as a subgenus.

(s.l.) is

new evidence

availal^le

it

With

appears advisable for the time being

to adiiere to the classification of the Treatise

koditcs as a genus; however, Spatulites

is

and

to retain Pele-

tentatively distinguished

as a subgenus.

Witchellia sutneroides Westermann,

PI.

Holotype.
incomplete)
pository:

sp.

Pis. 28-31;
Text-figs. 36-38, 40-41

Text-fig. 36a; well-preserved

phragmocone, internal mold with part of

USNM

shell.

(?

Re-

160243.

Locus typicutn.

IJ.S.
Wide

soutlieastern shore of

Stratum typicutn.
160

la-c;

28, figs,

n.

Geol. Survey Mesozoic locality 21251,

Bay, Alaska Peninsula.

Pseiidocidoceras

zonule, supposedly about

below top of Kialagvik Formation.

Derivatio /;om/???'5. Resembling TF. sutneri (Branco)
Age. Middle to upper S. soiverbyi Zone, Bajocian.
Diagnosis.

sidcate-tricarinate,

large evolute species of Witchellia, typically bi-

usually

througliout

strongly

including

costate

shoulders and lateral carinae; simple ribs on outer one to two

whorls.

Material.

The

holotype and two phragmocones with body

chamber fragments from USGS 21251

(USNM

160249,

160248);

two complete phragmocones, one with fragment of body chamber

fragmental body
from USGS 12405 (USNM 160244, 160247)
chamber from USGS 19869; one complete juvenile specimen from
USGS 19922; one phragmocone fragment and one body chamber
10 at 14-15.5 m (McM J 949)
one
fragment with test from
11 at 19 m (McM
(?) deformed body chamber fragment from
fragments of single large phragmocone with incomplete
J 945)
one large body
11 at 22 m (McM J 948)
body chamber from
(McM J 950) one (?)
chamber fragment from
13, scree
;

WA

WA

WA

WA

Alaskan ammonites,

Pt.

II:

Westermann

117

Text-fig. 36 a-b.
Cross-sections of JVitchcUia sutneroidcs, n. sp.
X 1Holotype, phragmocone with septal face (lobe axes shaded, saddle axes
indicated by dashed lines). Note tricarinate internal moid and high hollow13 (McM J 950).
floored keel. b. Fragment of large body chamber, loc.
;

a.

WA

immature specimen with incomplete body chamber from \VA 13

at 9 m (McM J 947)
one deformed phragmocone with incomplete
body chamber from
3W, scree (McM J 946) All specimens
are well-preserved internal molds, some with parts of the shell,
from the (upper) Pseudocidoceras zonide of the S. sozverbyi Zone,
;

WA

Kialagvik Formation,
Description.

Wide

The

Bay.

nucleus can only be seen on the holotype;

the second whorl, at a diameter of about 2 mm, bears blunt lateral

diameter, the moderately evolute
nodes or bullae. At 3-4
whorls are depressed oval in section with a blunt low keel bordered
by weak sulci. There are eight primaries on this whorl which ter-

mm

minate on the middle of the flanks in rounded nodes; secondaries

are missing. Subsequently, the strong costation becomes highly irregular, fasciculate in groups of two to four, and single ribs swell
to bullae-like primaries.

increases at about 30

The number

mm

of bullae

diameter from

nine per halfwhorl. However,

blunt ribs per halfwhorl.

fasciculations

four to five to

seven to

bullae are not developed in the

involute and stronger compressed variants which

lar

and

more

have 9-14 irregu-

Bllletin 255

118

^n

Cross-sections and septal sutures of IVitchcllia sutText-fig. 37 a-c.

neroides, n. sp., X 1- a- Compressed variant, part of phragmocone with septal
surface (see Text-fig. 36 a), loc. USGS 12405. b. Part of phragmocone and
body chamber of average specimens, loc. USGS 21251 (USNM 160249). c.
H, 42
D (above) and at 32
H (solid line:
Septal sutures at 18
11 (McM J 948).
same specimen as in 37b, dotted line: loc.

mm

At 20

mm

mm

mm

WA

diameter, the whorl section of the holotype becomes

subsquare and finally moderately compressed subrectangular.

slightly

The

converging whorl sides flatten out, roiniding into the low

steep umbilical wall

bisulcate-tricarinate.

and into the

The

Inroad venter

keel of the shell

is

which

is

usually

high and bladelike

but reduced resemliling the carinae on the internal mold, while

the sulci are somewhat shallower than on the internal mold. Tlie
evolute, less ornate forms become more compressed and almost

Alaskan ammonites,

Pt. II:

Westermann

119

Cross-section of WitchText-fig. 38.

sutneroides, n. sp., largest phragmocone with apparent shell duplication (cf.
PI. 31), loc. USGS 21251 in the Pseudo1160248);
cidoceras zonule
ellia

(USNM

Bulletin 255

120

307oU

60

80

D (mm)

Scatter with growth lines for relative umbilical width

Text-fig. 39.
of the Witchrllia lar-viuscula - sutncri group, IV. laeviuscula Subzone
of England, and of 'Gclasinitcs' gclashius Buckman, iS. trigonalls Subzone of
England. Note the subcontinuous distribution from 21 to 34% U throughout
the mature whorls of the phragmocone (dots and solid lines). A more or less
arbitrary division at about 29% U separates the IV. laeviuscula 'plexus' from
the //'. suttieri 'plexus'.

(U:D)

ovate in wliorl

.section

with narrower venter and weaker developed

sulci.

At 35-50 mm diameter, the costation becomes bifurcate or,

more rarely, triturcate after extension of the lower-lateral bullae
into rectiradiate or

somewliat prosoradiate

maries, 8-13 per halfwhorl, reach their

two-thirds whorl height

primaries.

maximum

and usually continue

The

pri-

elevation at about

straight or slightly

rursiradiate into one of the swollen secondaries.

The

secondaries,

18-25 per halfwhorl, project strongly on the roinided shoulders

and

full strengtli. In the more compressed

and more involute forms, there are mostly somewhat irregular blunt

reach the outer carinae in

much weaker

ribs.

Alaskan ammonites,

Pt.

II:

Westermann

121

D (mm)

Text-fig. 40.
Scatter with growth lines for relative umbilical width
(U:D) of Witchcllia sutncroidfs, n. sp., holotype (H) and catalogue or localifj'
numbers of other specimens indicated, all from the PscudociJoccras zonule of
Wide Bay. Note that the mature phragmocone (dots and solid lines) varies from
26 to 38% U, resembling the IV. sutneri 'plexus' and 'Gelasinites' (Text-fig. 39).

At 80-100 mm diameter, tlie principal ribs become more numerous by continuation of the primaries into single strengthened
secondaries while the others become reduced. The last wliorl of
large shells bears strong, mostly simple, rectiradiate, projected to

somewhat rursiradiate costae, resembling large Euhoploceras. This

strong ornament is usually retained up to the end of the body
chamber, with the exception of the more involute and compressed
variant where it becomes obsolete. All body chambers retain the
striking bisulcate-tricarinate venter.

Bulletin 255

122

13

12

(O)
(path.)

A
10

o
5
Si

O-

()
D mm-

5 I

25-40

60 -end

40-60

27

26

?5

phrogm.

29

28

30

U
+L

32

31

33

34

36

35

Scatter

for 'ornament' (P-primaries, or bullae per half(relative umbilical width, U :D X 100) at different phragmocone diameters of H'ltrhrllia sutrtrroidrs, n. sp., all from the Psrudocidoceras
zonule of Wide Bay. There may be some weak negative correlation, at least
for the smaller size groups, i.e., the more evolute shells may tend to bear fewer
(stronger) primaries or bullae.

Text-fig.

whorl)

vs. 'coiling'

The
is

high infraspecific variation

Biicknian

"(1st.)

Law

of Covariation"

is

another example of the

(VV^estermann, 1966)

There

a complete morphological transition from involute, strongly com-

pressed and weakly ornate forms to evolute, weakly compressed

and strongly ornate forms. In addition, there

The

form.

mm,

is

probably

adult diameter varies from about 100 to

a smaller

more than 200

26% to more than 36% in phragmowhorl section (H/W) of adult phragmocones

and 1.4 (? 1.45) and of body chamber between 1.25

umbilical width from

cones of similar

size,

between

1.15

and

(Text-fig. 40)

.6

The number of primaries is difficult to

may occin* singly or in the form of bullaeThe number of secondaries varies between aI)out 18

determine because they

like l)undles.

Alaskan ammonites,

and 28 per
specimens.

Pt.

II:

lialfwhorl, but clianges little

The

Westermann

on the

last

123

whorls within

plot ol relative umbilical width against

number

of

primaries/bullae suggests weak negative correlation at different

growth stages (Text-fig. 41). This variation is pene-contemporary

since no morphological change was observed throughout the restricted vertical range of this species.

The septum
(Text-figs.

36-37)

is
.

fluted

according to the planodisculate type

The major

lobe-axis

lateral

L-L

concave

is

I.
The
The incomplete re-

fusing medially with the incomplete central lobe-axes of

incomplete lobe-axis of

duced median lobe-axes of E ends

saddle)

reduced.

U^. is strongly

at the

cinved E/L-E/L (external

saddle-axis. In accordance with the different strength of

these septal elements, the suture has a small E, a large

large

(which

is

trifid

with long thin endings)

suspensive) umbilical elements of which

the size of L.

U3.

The

large

The

U^

internal suture has a deep narrow

I/U

saddle.

is

adjacent major indentation

The

general complexity

E/L

saddle, a

and small (non-

usually only one-half

is

either all or part of

and

a single

(frilling)

is

(paired)

moderately

high.

This species closely resembles Witchellia sutneri

from the S. sowerbyi Zone of Europe, from which it is
distinguished by the stronger costation especially on shoulders and
carinae of the last whorls and by the well-developed bisulcatetricarinate venter of shell and internal mold of phragmocone and
body chamber. Also, W. sutneri appears to have more tightly coiled
Comparison.

(Branco)

outer whorls.

Two

small specimens resemble

W.

sutneri (original figures re-

produced, Text-fig. 34) so closely that they were originally identified with that species; they are especially close to 'W. platymorpha'

Buckman (Type Ammonites, pi. 580) from the W. laeviuscula

Subzone of Dorset. However, these Wide Bay specimens appear to
intergrade with the bulk of the sample with which they are, therefore,

regarded conspecific.

whorl (including body

chamber) of W. siitneroides strongly resembles W. ('Zugophorites')
Buckman from the S. trigonalis Subzone (middle S. soiverbyi Zone)
of Dorset. However, the English forms are bisulcate on the internal
mold only and their costation is much weaker becoming obsolete
on the last whorl.
Particularly, the widely umbilicate last

Bulletin 255

124

ir.

sutueroides

ate between the

is,

therefore, morphologically almost intermedi-

European forms

of the

middle and upper

S.

sower-

byi Zone, resj>ectively, but distinct in the stronger costation especially

on the shoidders.

the

Significantly,

same featme

also dis-

from its
European relatives.
Of special interest are two species [uurni)ia dubia'\ described
from British Columbia and imnamed specimens from Alberta. The
tinguishes the associated

S.

(Euhoploceras) bifiircata,

n. sp.

Smvey of Canfrom talus of the Hazelton Group at Hudson Bay Moinitain, British Columbia, was based
on a single incomplete and poorly preserved specimen. The small
conch (52 mm D) is evolute (Ur=38%) compressed elliptical to
slightly subrectangidar in whorl section, and has strong simple
costae bundled in pairs near the umbilicus. The internal mold is
strongly keeled and narrowly bisidcate. Although superficial similarity with W. sutncroides exists, the poorly known Canadian form
is distinct in the absence of primaries or elongated bullae and in
the reduced lateral carinae of the venter. 'Sonninia silveria' McLearn (1926) foinid in the same scree, is based on a single distorted
and incomjilete poorly preserved j^hragmocone, possibly with part
of the Ijody cliamber. Tlie inner whorls up to 45 mm diameter resemble evolute, l)isidcate, and fasciculate variants of W. sutncroides,
but the British (Johmibia form becomes subsequently almost smooth
and the venter simply keeled on narrow tabidation. Both specimens may well lielong to a single (? new) species of Sonninia but
plastercasts

were kindly furnished by

ada. "Sonninia skaicahi"

McLearn

tlie

Geological

(1926)

the specific

names

are to be regarded as

nomina

diibia

(tiiey

were

omitted in the "illustration of Canadian Fossils" by Frebold, 1961)

Sonninia

sp.

indet.

Devils Point at Lake

1957a, p. 49,

pi.

19,

was descriljed from the Fernie Group

Minnewanka near
fig.

2).

The

Banff, Alberta

figured specimen

of a large evolute sonniniid with

is

at

(Frebold,

a fragment

subrectangular whorl section,

moderately bisulcate-tricarinate venter of the internal mold, and

simple rectiradiate projected weak costae which bundle on the ante-

penultimate whorl. The plastercast of this and of another fragment

were kindly fmnished by the Geological Survey of Canada. Both
specimens were apparently associated witli tlie S. (? Enhoploceras)
cf. bifurcaia (op. cit., pi. 20; see
bijnrcata). This form resembles
.S'.

Alaskan ammonites,

Pt. II:

Westermann

and may be
Columbia forms described above.

closely Witchcllin ('Zugophorite.s')

British

125

identical with

tlie

Shell duplication. The largest specimen (USGS 160248) with

an estimated complete diameter of 200-220 cm shows unique
thickening and continuation of apparent test from the sides of the
last phragmocone whorl onto the exposed sides of the penultimate
whorl and over the plugged umbilicus where the inner whorls are

not preserved (Text-fig. 38;

this

apparent secondary

thick,

but on the right side

and one

mm

On the left side of the conch,

only about one-half millimeter
duplicated (locally triplicated ?)

PI. 31)

test
it

is
is

This apparent test generally rounds

off the sharp angles at the umbilical seams and seems to have been
continuous also onto the missing body cliamber. While superficially
closely resembling ammonite test and internally showing similar
to

two

thick.

although often duplicated layering, the material consists of coarsely

crystalline calcite (X-ray fluorescence inspection by B. J. Burley)
which either could have been derived from ammonite test by diagenetic alteration or be

This

little

altered nonbiogenic calcite.

phenomenon

is probably best explained

by late diagenetic nonbiogenic processes in which the surrounding
sediment was separated from the shell (or dilation within the test)
due to differential compaction and the thus developed supplementary natural mold subsequently filled with calcite by slow lateral

shell duplication

secretion. It appears unlikely that this

apparent shell

is

true test

grown animal asymmetrically over both the

entire sides of the conch after destruction of the inner phragmocone
whorls. Such magnitude of secondary shell secretion is unknown
secreted by the fully

in the cephalopods.

While this article was in press, several instances of 'diagenetic

conch hypertrophy' resembling this case were described and analysed by R.

Hollmann

103, pp. 305-334)

(1968, Neues Jahrb. Geol. Palaontol., Abh.

from Upper Jurassic ammonites of East Africa.

Measurements.
Holotype (phragm.)

j^^^
69

^^^

j^^^

U%

p/bullae

126

Bulletin 255

Alaskan ammonites,

whorl section

this

square.

Pt. II:

Westermann

to the end, the slightly larger

The body chamber

127

one becomes subbecomes

of only one-hall whorl length

more evolute and somewhat compressed

elliptical in section,

with

simple narrow tabulations beside the bhnit keel. Both specimens

have

lateral lappets

The

which are probably incompletely preserved.

costation consists of moderately strong irregularly fascicu-

late costae which are almost rectiradiate and slightly flexed on the
phragmocone, but markedly rursiradiate and terminally projected
on the body chamber. The costation is reduced, finally becoming
obsolete on the last one-fourth whorl.

Measurements.-

McM

J 1028 a

(body

ch.)

^^^

^^^

^^^

U%

128

Bli.letin 255

the venter is keeled-tabiilate or bisiilcate. On the larger phragmocone are many irregularly fasciculate ribs, some of which are inflated near the base marking the beginning of WitcheUia-type
bullae.

The

costation of the smaller specimen

much

is

coarser but

similar in style.

mm

The body chambers of the two specimens were 52

and
approximately 32
in respective complete diameters. The whorl
section of the larger specimen is compressed elliptical with strong
keel and rather sliallow sulci on the internal mold. Somewhat con-

mm

vex blunt ribs are developed on the middle of the sides but beobsolete near the aperture. The shell remnants show increas-

come

ingly convex growth striae

An

and

test

thickening near the peristome.

incompletely preserved ventral lappet

(or

rostrum)

is

still

on the internal mold. Tiie sides are somewhat constricted on the internal mold only and extend in small
bisulcate-tricarinate

may be
The body chamber fragment

medio-lateral lappets whicli

incomplete.
of

the

smaller

specimen has

strong rursiradiate ribs whicli are partly somewhat fasciculate.

externside

The

is

septal suture

exactly as in juvenile Witchellia sutner-

is

narrow and moderately deep, the E/L saddle

large and deep, the umbilical lobes small and non-sus-

oides, n. sp.
large,

The

bisulcate throughout.

is

pensive.

This

form strongly resembles the English P.

(Buckman) which has a diameter of 59 mm,
and the smaller 'Maceratites' aurifei- Buckman (34 mm D) it is
Comparison.

(Spatulites) spatians

distinguished by the better developed bisulcate-I)icarinate venter

retained on the body chamber, the stronger ornate shoulders and

probably the simplified costation of the body chamber.

Significantly,

these

same features distinguish the apparently

W. siitneroides, n. sp. from tiie

corres|X)nding macroconchiate

northwestern Eixropean W. laevhiscida

Measurements.-

McM

939

(aperture)

,),^^

^^^

W.

sutneri group.

H^^

U%

Alaskan ammonites,

Pt.

II:

Westermann

129

Family OTOITIDAE Mascke, 1907

[Incl.

Sphaeroceratinae Buckman, 1920]

Otoitids and sphaeroceratids are regarded as two subfamilies

of a single family by this author

(Westermann, 1956, 1961,

in contrast to their treatment of the 'Treatise'

p.

55)

(Arkell, et al. 1957)

Schindewolf (1965, pp. 161-177) recently upheld their separation

at the family level because of minute differences in the umbilical

However, he retained unquestioned

lobes.

in

the single probably dimorphic pair Emileia

transfer of Docidoceras
tidae.

and D.

his

Otoitidae only

E.

(Otoites) after

(Trilobiticeras) to his Sphaerocera-

Schindewolf did not investigate the supposed hammatoceratid

Abbastites, placed in Erycites in the Treatise but generically dis-

tinguished by the present author, which according to the evidence

from Wide Bay is directly ancestral to Docidoceras (Pseiidocidoceras), n. subgenus. Because of the presence of the secondary morphological internal lateral lobe U,,, Abbastites should be transferred
from the Hildocerataceae to the Stephanocerataceae in accordance
with Schindewolf's classification.
D. (TrilobitiIf the supposed dimorphic pair Docidoceras
ceras) is placed in the sphaeroceratids as supposed by Schindewolf,
the distinction between the otoitids and sphaeroceratids is further
diminished by neglecting differences in dimorphism; in the "old"
-

classification

conchs.

The

only

the

otoitids

possessed

bearing micro-

lappet

[and Pachyceratidae]

Tulitidae

are certainly

more

from the otoitids and sphaeroceratids than asserted

by Schindewolf (1965, p. 235) for example, the septal sutme lacks

clearly distinct

the Uq.

This author

most hesitant to follow Schindewolf (1965, p.

dimorphic pair Cadomites - C. (Polyplectites) in the sphaeroceratids because of their unusual septal
structure (Westermann, 1956, p. 242) which he now confirmed.
There can be little doubt about their direct descendance from
is

170) in placing the probable

typical stephanoceratids.

Pseudotoites Spath

is

characterized by planulate subcircular to

rounded ovate inner whorls with inclined and partly smooth umbilical wall, 'cadicone' intermediate whorls, and typically ovate
outer whorls bearing blunt nodes and bullae on the umbilical
Primaries and
margin with reduced primaries (Text-fig. 42)
.

Bulletin 255

130

secondaries are usually prosoradiate.

The

aperture

is

not collared

and the peristome strongly oblique. The author

grateftdly

ceived plastercasts of several Australian

identified by

Arkell, from the Sedgwick

toites, as

Museum. The

topotypes,

re-

internal suture of Pseiido-

investigated in two specimens of P. ex gr. P. singularis

(Gottsche)

from the Argentine Andes, agrees

in detail

with that

given for Emileia and E. (Otoitcs) by Schindewolf (1965, pp. 153,

162), probably including the two-pronged U,. However, the earliest

growth stage could not be developed. Coiling and whorl

tion of P. singularis resemble closely the

more evolute

sec-

species of

Emileia.

The common Alaskan

formerly

"sub-coronates",

tentatively

regarded as new subgenus of Pseudotoites

(Westermann, 1964a,
have throughout depressed
footnote 19, p. 61, pi. 9, figs, la, b)
whorls with mid-lateral edge, sharp
subelliptical
('lenticular')
long primaries, and an undivided Ui lobe, and are, therefore, now
subgen. D.
classified luider Docidoceras (Pseudocidoceras), n.
(Pseudocidoceras) is distinguished from Docidoceras s.s. mainly in
the regularly strong projected (convex) secondaries and in the
oblicjue smooth aperture without collar.
The Australian 'Zeniistephauits' Arkell (1954) which is not
identical with tlie Canadian stephanoceratid Zcinistcplmniis (Imlay,
,

\Vestermann,

1964;

(Pseudocidoceras).

(Arkell, 1954, pi. 39,

1964b,

pp.

61-64),

somewhat lesembles D.

plastercast of the septal surface of Z. corona

fig. 3)

ern Australia, shows the

kindly sent by the University of West-

l)ullate

However, the more striking

pattern of the Otoitidae

similarities are with the

(s.l.).

South Ameri-

can P. (?) sphaeroceroides (Torncjuist) Both the Australian and

South American "coronates" may best be classified imder a new
subgenus of Pseudotoites (Westermann, 1964b, p. 53) although
.

they

may

be related as closely to Docidoceras.

In spite of
material of P.

my

recent extensive search in the type-area, no

(?)

sphaeroceroides

refigured by Arkell

men

of P.

1964b,

pi.

(?)

9,

(1954, pi. 40,

is

figs.

2a-c)

sphaeroceroides previously figured

fig.

7)

new

The holotype was

The complete speci-

available.

has strong tubercles

(internal

(Westermann,
mold) on the

body chamber which are lacking in D. (Pseudocidoceras), and the

whorl section of the phragmocone is depressed ovate with the

Alaskan ammonites,

Pt.

II:

Westermann

131

Cross-section of the inText-fig. 42.

ner and intermediate whorls of Pseudotoites robiginosus (Crick) [det. P. emiliodcs by Arkell; Sedgevvick Museum F
11923], Mount Hill, Western Australia;
for comparison with the Alaskan species.
Note the planulate and rounded inner
whorls.

lateral

edge at about two-fifths whorl-height, while

Wide Bay

pressed elliptical in the

inner whorls of the Australian Z.

known, except

for the

it

is

usually de-

forms. However, the section of the

corona

is

apparently

end of the phragmocone where

it is

poorly
also de-

pressed ovate.

The

Company, the U.S. Geoand by me from the S. soiverbyi Zone of Wide Bay
in summer 1964, is evidence for two distinct but associate otoitid
groups, i.e. subcoronate Docidoceras (Pseudocidoceras) and planumaterial collected by Shell Oil

logical Slavey,

late Pseudotoites.

Tlius, Pseudotoites

is

present in North America, but neither as

formerly supposed (Arkell, 1954,

p.

586)

by 'Amjnonites' carlotten-

Whiteaves from western British Columbia and southeastern

Alaska which is a Zemistephanus (Imlay, 1964; Westermann, 1964b,
nor by the abundant "sub-coronate" Wide
footnote 19, p. 62)
previously {op. cit., pi. 9, fig. 1), but by
suggested
Bay forms as I
sis

the scarce planulate P.

cus (Tornquist) of the

cf.

S.

and P. cf.
Zone of Wide Bay.

argentirius Arkell

soiverbyi

trajisatlanti-

Bulletin 255

132

80

F 7o

Alaskan ammonites,

The major
(2)

clusters are

133

The European Docidoceras

s.s.,

the Alaskan D. (Pseudocidoceras),

Pseudotoites and

West

(1)

Westermann

Pt. II:

(4)

Irian, formerly

1962, end.

17,

fig.

(3)

South American

tlic

the Australo-Indonesian

Pseudotoitcs

[for

West New Guinea, see Visser and Hermes.

17, and for the Sula Islands, Moluccas, see

Kruizinga, 1926,

pi. 6, figs. 1, 2; pi. 12, fig. 3]; a fifth cluster could

be represented by the northwestern European Emileia ('Emileites').
All clusters certainly overlap in time within the lower to middle S.

soiverbyi Zone. Clusters (I) to (4) appear to be subcontiguous in

morphoclinal relationsliip and geographic distribution, the Aus-

tralo-Indonesian Pseudotoites being most distinct from the European Docidoceras s.s., and (2) and (3) successively occupying intermediate positions. Each cluster is chstinguished by a combination
of characters which also occur singly in the others. Yet in southern
Alaska (2) and (4) are associated breaking the linear relationship.
The South American Pseudotoites singidaris (Gottsche) group

appears to be as closely related to E. ('Emileites') as to (1) Docidoceras S.S., both essentially European. Besides the major clusters,

supplementary clusters with different

lateral

distributions occur;

Pseudotoites argejitinus Arkell from the South American cluster

supplements the Alaskan

cluster, the

cadicone P.

(?)

(n.

subgen.)

sphaeroceroides (Tornquist) group supplements the South Ameri-

can and probably also

forms

Australo-Indonesian

the

are, in turn, closely affiliated

of D. (Pseudocidoceras). It
{^Emileites') cluster

clusters.

possible that even the

is

was spread

These

with the Alaskan main cluster

as far as

European

E.

South America.

Genus DOCIDOCERAS Buckman, 1919

Type species by original designation. D. cylindroides, Buckman, 1919, from the 5. soioerbyi Zone, L. discites Subzone, of southern England and France.
The taxonomy of the genus has recently been discussed by this
author (1964b, pp. 51, 56-57) The nominate subgenus D. (Docidoceras) is macroconchiate and includes the three distinct species D.
cylindroides Buckman, D. planulatum Buckman, and D. longalvum
probably synonymous with, or subspecies of these are
(Vacek)
'Coeloceras' limatum Pompecki, D. perfectum Buckman, D. bijorme
Buckman, and D. liebi Maubeuge. All have simple moderately
.

Bulletin 255

134

oblique and typically collared peristomes. Tlie probably corresponding microconchiate forms are included in the subgenus D. (Trilobiticeras) with the only distinct named species D. (T.) punctiim
(Vacek) which includes as subspecies or synonyms T. trilobitoides

Buckman and

T. platygaster

Buckman

[part.]; the aperture bears

lateral lappets.

The
Otoitidae

pertinent featiue distinguishing Docidoceras from other

[including Sphaeroceratinae]

is

the

evolute,

strongly

depressed 'lenticular' whorl section with mid-lateral edge. Other

Otoitidae have ovate whorl sections with the maximal whorl width
or lateral-edge below mid-flank (less than one-half whorl height)
(Text-fig. 43)
at only a

Tiie 'coronate' whorls of Docidoceras

commence

few millimeters diameter and are developed throughout

the phragmocone, although the adult

phragmocone and particularly

body chamber may become somewhat ovate by inflation of the
externside and steepening of the inner flanks (umbilical slope)
As in most other Otoitidae the costation of typical Docidoceras is
more or less rectiradiate, crossing straight over the externside, but
there are some exceptions such as the holotype of D. liebi Maubeuge
D. planiilaturn Buckman]. Unrefuted D. (Docidoceras) and
[?
D. (Trilobiticeras) are known from the northwestern Emopean and
Mediterranean provinces only.
the

Subgenus

DOCIDOCERAS (DOCIDOCERAS) Buckman,

Docidoceras (Docidoceras?)

sp. aff. D.

1919

longalvum (Vacek) 1886

PI. 32,

fig. 5a-c; PI. 33; Text-fig.

Material. One large phragmocone with

chamber, well-preserved internal mold, from
Lower Pseudocidoccras zonide of the
J 937)
upper Kialagvik Formation, Wide Bay.

incomplete

WA

Description.

The

sliell

is

S.

10 at 8

44

body

m (McM

soiverbyi Zone,

typically "serpenticone planulate";

the whorls are strongly depressed elliptical, j)artly slightly ovate

rounded lateral edge, and become increasingly more

embracing the preceding whorls less than one-third. The
costation is prominent on the phragmocone consisting of straight
rectiradiate primaries which trifurcate in mid-lateral tubercles or
nodes into slightly prosoradiate secondaries. The secondaries become more densely spaced and blunt on the end of tlie phragmocone. The preserved one-fourth wliorl of the body chamber is almost smooth.
in section with

evolute,

Alaskan ammonites,

Pt.

II:

Westermann

Text-fig.

44.

Cross-section

135

of

Docidoceras (Docidoceras) sp. aff. D.

longal'vum (Vacek), complete specimen with strong umbolateral groove
on last phragmocone whorl and body
chamber, loc.
10 in the lower
Pseudocidoceras zonule (McM J 937)

WA

The

1.

ultimate one-quarter whorl of the phragmocone and espe-

body chamber fragment have on right and left innermost

strong and well-defined grooves [here
named umbolateral groove] On the phragmocone these are largely
filled with "porous material" which is believed to represent secon-

cially the

flanks

(umbilical slopes)

dary secretion, possibly originally rich in organic material, for

at-

tachment of the paired retractor muscle (see also Jordan, 1968)

On the body chamber, the giooves correspond to all of the inner
flanks and are bordered by a sharp mid-lateral edge (on the left side
apparently raised by fracture) (Text-fig. 44)
The septum is typically bullate with slightly stronger outer
than inner saddle axes. The suture is of moderate complexity with
.

suspensive umbilical elements.

Bulletin 255

136

Discussion.

The

Alpine holotype of D. (D) longalvum has

recently been refigured

English D. perfectum

Bremer, 1966,

p. 52;

(W'estermann, 196-45,

Buckman was drawn

p. 163)

pi.

into

6,

fig.

synonymy

1).

(op.

The
cit.,

matching the holotype of D. longalvum

except for the slightly stronger costation.

The Wide Bay specimen is distinguished from D. longalvum

by the more distant partly prosoradiate and markedly stronger
costation and by the iimbolateral groove of unknown taxonomic
significance, featiues which suggest affiliation with D. (Pseiidocidoceras).

Measurements.

Dmm

McM

J 937 (body ch.)

(phragm.)

140

W7f

H7o

U%

Alaskan ammonites,

is

Westermann

137

by planulate rounded inner whorls, short

distinguished

maries which

Pt. II:

may become

obsolescent,

and prominent

pri-

nodes

lateral

or btillae of the outer whorl.

A single apparently corresponding microconch was associated

with the several abundant macroconchiate D. (Pseudocidoceras)
macroconchs from Docidoceras

species. Just like the

niocone

is

the phrag-

s.s.,

distinguished from D. (Trilobiticeras) in the strongly

prosoradiate ribs. The naming of a new subgenus would be premature and could later become superfluous, if dimorphism can
be established as more material becomes available.

The

D. (Pseudocidoceras) camachoi,

affinity of

sites platystoyniis

Westermann

jacent E. howelli Zone

(1964a, pi. 66,

E/L

n. sp. to
1-2)

Abba-

of the sub-

so striking that their direct phyletic re-

is

lationship appears certain. A. platystomus

the reduced

figs.

distinguished only in

is

saddle and in the alternating secondaries of the

inner whorls.

D. (Pseudocidoceras) bears much resemblance to the 'Emileites'

group, i.e. widely umbilicate Emileia species from the middle and

upper

5.

crater

Buckman;

Zone of southern England,

soiuerbyi

these in turn,

may have

especially to Emileia

evolved directly from

D. cylindroides Buckman of the lower

5. sowerbyi Zone
(Westergroup needs reinvestigation;
it is distinguished from the Alaskan forms by the ovate roimded
whorl section and the retaining of long, thin primaries on the body
chamber.

mann, 1946b,

p. 60)

The

'Emileites'

Docidoceras (Pseudocidoceras) widebayense Westermann,

n. sp.
Pis. 34-37; Text-figs.

Holotype.

PI.

mold with minor

No. 160249.
Locus typicus.
8.5

km

(5.32

mi.)

34, fig. la-d; a well-preserved

test

remains. Repository:

Sea

cliff

4549

complete internal

U.S.

Nat.

Museum,

on southeastern shore of Wide Bay,

west of west end of Hartman

south, 461/4

Island. U.S. Geological Survey Mesozoic locality 21251

(collector

Imlay and Miller, 1948)

Stratum typicum. Pseudocidoceras zonule, supposedly about
160
below top of Kialagvik Formation.
Derivatio yiominis. Nnvaed after the occurrence at Wide Bay.

Age and

occurrence.

S.

sowerbyi Zone. Bajocian,

Common

Bulletin 255

138

Septal sutures of Docidoccras (Pscudocidoccras) , n.

Text-fig. 45 a-g.
subgen., loc. USGS 19862 in the Pscudocidoccras zonule of Wide Bay, magnia-b. at c.l
W, c. at 11.6
fied, a-f. D. (Pcsudocidoccrasj sp. indet. juv.

mm

mm

and

and
2.5

mm H,
mm H; g.

D. Note

d.

at c.2

mm W,

e.

at

mm

(P.) ividchaycnsc, n. sp. at 6

the secondary lobe Un.
i>.

in the Pseudocidoceras zonule, especially in

in the subjacent E.

eastern side of

and

mm
mm

mm W,

H,

f.

mm H

at

and

mm
c.9

the upper part, rare

amplectens zonule; only known from the south-

Wide

Bay.

medium-sized species of D. (Pseudocidoceras),

strongly costate including body chamber.
Material. Three complete and two incomplete specimens
Diagnosis.

from

USGS

12405; one almost complete specimen

mocone from USGS

and one phrag-

19801; one complete specimen, five phragmo-

Alaskan ammonites,

o)

15

Pt. II:

Westermann

139

Bulletin 255

140

70

50

40

30

20

phroqm

wideboyense

cf/off wideboyense

operture

D comochoi
10

(Tnlobitic ') sp nov

50

100

150

D (mm)

Text-fig. 47.
Scatter with several growth lines for relative umbilical
width (U:D) oi Docidocnas (PsruJocidoccras) n. spp. and D. (Trilohitifrras?)
sp. indet. Psrudocidirrras zonule of Wide Bay; Holotype of the type species
phragmocones solid lines,
D. cyliridroidrs Buckman included for comparison
body chambers: dashed lines). Note that the growth lines remain essentially
constant throughout the phragmocone but increase with the body chamber, that
D. (P.) ividr/uiycnsr, n. sp. tends to be slightly more evolute than D. (P.)
camachoi, n. sp., that the microconchiate D. (Trilohiticeras ?) plots subcentrally within the former macroconchiate forms, and that D. (D.) cyliridroidrs
resembles D. (P.) luidehayrnsr in this parameter.
,

cones partly with incomplete body chamber and several fragments

from the USGS 19862 (USNM 160250) several complete phragmocones and one body chamber from USGS 19869; two complete
specimens, one phragmocone, several fragments of phragmocones
and body chambers from USGS 21251; one body chamber with
one small specimen with
pliragmocone fragment from
5;
(?)
;

WA

Alaskan ammonites,

Pt.

Westermann

II:

141

cylindroides

60
80%W

60%*

/TO'ltW

'/

50

40

// 7

20 -

pnrQgm

cf /off

wideboyensa

{Tr.lobitic ?)

50

comachoi

aperture

widetayense

sp nov

100

150

(nim)

Scatter with growth lines for relative whorl width ('thickof Docidoceras (Pscudocidoceras) , n. spp. and D. (Trilobiticeras ?), n. sp. indet., Pscudocidoceras zonule of Wide Bay (same symbols as
Text-fig. 47). Note that width growth ceases or even decreases (growth rate
becomes negative) at the beginning of the body chambers which, therefore, become relatively 'thinner', that there is close resemblance between D. (P.) widebayense, n. sp. and D. (P.) camachoi, n. sp., and that the microconchiate D.
(Trilobiticcras ?) as well as the holotype of the generic type species D.
cylindroides Buckman plot within the distribution of the macroconchs.

Text-fig. 48.

ness"

=:

W:D)

WA

m; six almost complete internal molds

phragmocones and body chambers from
10 at 1 to 22 m (McM J 903, 909-917)
two incomplete phragmocones with body chambers, several large fragments from
1
at 19 to 27 m (McM J 904-906)
two complete internal molds
with test remains, two phragmocones, five fragments from
13
All except one [WA5
E.
at 9 to 16 m (McM J 895-902, 916)
amplecteths zonule] from the Pseudocidoceras zonule of the S.

aperture from

and

10 at 6

several fragmentary

WA

WA

WA
=

soxverhyi Zone, Kialagvik Formation,

Description. ^he^

Already at 6-8

mm

phragmocone

Wide
is

Bay.

'sub-coronate' throughout.

diameter, the evolute whorls are strongly de-

Bulletin 255

142

phragm

body ch
cylindroi^des

10

20

30

40

50

60

70

80

90

100

110

120

130

D (mm)

Text-fig. 49.
Scatter \vith growth lines for density of secondaries (number per half whorl) vs. diameter (S:D) for Docidoccras (PscudociJoccras),
n. spp. and D. (Trilohiticiras ?), n. sp. indet., PscuJocidocrras zonule of Wide
Bay (same symbols as Text-fig. 47). Note that there is large overlap for the
phragmocones of D. (P.) ividcbayensr, n. sp. and D. (P.) camachoi, n. sp., buti
clear separation for the body chambers, that D. (Trilobiticeras ?) sp. plots
within the distribution of the macroconchiate D. (Pscudocidoceras) , and that
the holotype of the type species D. (D.) cylindroidcs Buckman resembles D.
(P.)

camachoi

in this

parameter.

pressed, lenticular, about double as wide as high, with evenly

rounded broad externside, distinct circa-mechan lateral edge and
rounded inner flanks which reach the umbilical seam almost vertically. Prosoradiate primaries extend hom the umbilical seam up
to the lateral edge where they carry small nodes. The densely spaced
secondaries project strongly and cross convexly, sometimes with
median weakening, over the broad externside.
The morphogeny is well known beyond 15 mm diameter. As
shown in the scatter cHagrams (Text-figs. 47, 48) there is little
change in the growth rate for whorl shape and coiling up to the
end of tlie phragmocone. Whorl width varies from 64 to 82 percent
,

Alaskan ammonites,

and the

of the diameter

The

wliorl

Westermann

Pt. U:

143

height approximates one-half of

maximal whorl width) lies

between three-sevenths, more rarely two-fifths, and one-half whorl
height, i.e. just below the middle of the flank. The umbilical width
ranges from 28 to 38 percent of the diameter. The diameter of
the width.

the adidt
ness,

edge

lateral

phragmocone

(or the

varies greatly with strong positive skew-

ranging from probably

less

than 40

mm, and measuring most commonly

to

(?30)

40-50

mm

more than 60

(Text-fig. 46)

There are 10-14 rectiradiate to moderately prosoradiate primaries per halfwhorl on the intermediate and outer whorls of
the phragmocone.

gradually

up

They

seam and strengthen

where
they
edge
carry nodes. Here they

arise at the umbilical

to the lateral

bifurcate or trifurcate into always

more

or less strongly prosoradi-

which usually curve evenly convex over the externside, but are sometimes slightly arched or become medially obsolete.
The density of secondaries on the last one or two whorls of the
phragmocone varies from 25 to 38 per halfwhorl, being most comate secondaries

monly 25-30

The

(Text-fig. 49)

adult body chamber

is from three-quarters whorls to over

one whorl in length and varies in diameter from probably 45 to
100 mm, measiu'ing most commonly 65-85 mm (Text-fig. 46) From
the beginning of the body chamber, the coiling becomes markedly
more evolute (39-47%) Whorl width ceases to increase and there
may be slight 'contraction'. Consequently the whorl section becomes higher and, by reduction of the lateral edge, rounded elliptical to slightly ovate (Text-figs. 47, 48)
At the aperture, the test
is internally greatly thickened causing a broad oblique constriction
on the internal mold, and a broad lip or ventral lappet produces
.

a strongly oblique peristome.

The
sf>ecies,

costation of the

body chamber

consisting of increasingly strong

most distinctive for this

and distant primaries and

is

especially heavy, strongly prosoradiate secondaries

whorl)

(Text-fig. 49)

The

(18-27 per half-

primaries do not markedly retract from

the umbilical seam; they retain greatest prominence over the maximal whorl width where they often carry blunt nodes.
The septum is typically bullate (abullate) with central E-I
lobe axis and complete L-Un lobe axis separating two sub-equal
saddle axes (E/L-I/U and L/U.-U/U) The septum of the re,

Bulletin 255

144

lated D. camachoi n. sp. has recently been figiued under 'Pseiidotoites, n. sp.'

The

(Westermann, 1964b,

septal

suture

(Text-tig.

pi. 9, fig.

45)

la, b)

superficially

is

similar

to

most Otoitidae (including Sphaeroceratinae) and other Middle

Jurassic sphaerocone and cadicone ammonites, with two subequal
morphological external and internal saddles and more or less high
complexity.

The morphogeny

observed in a D. icidebayense (Text-

45g) agrees in detail with that of a nucleus of D. (Pseudocidoceras) sp. indet. from the same locality, so that only the umbilical
fig.

elements at 10

mm

diameter have been figured of the former in

supplementation of the other series (Text-figs. 45 a-f) Up to 1

whorl width, only E, L, U., Uj and I are present with the
umbilical seam situated on the lower inner slope of the Uo/Uj
.

mm

saddle.

At

1.2

adjacent to the

mm

mm

H) U^ develops externally
seam and, simultaneously, on the dorsal side of the
1.5

(0.8

Uj/I (internal)
about 2.5

crest of the

mm

saddle.

mm

Frilling of the suture

com-

H) when U has developed

(1.5
mences at
to a large straight lobe separating the sub-equal Uj/U,, and
U/I saddles, while U, remains undivided. U4 develops adjacent
D)
H and 6
whorl width (2.5
to the seam at about 4
at a stage when all other lobes including Ui are trifid. The shape
some

ing to
frilling

lobes, especially of Uo,

mm

mm

mm

of

is

often strongly asymmetric tend-

Only a moderate degree of complexity by

appear
the lobes remain usually quite broad, in
and
is achieved
bifid.

contrast to Eniileia.

Comparison. -T\\t quantitative studies of D. widehayense and

D. camachoi establish clearly discontinuous variation of the body
chamber costation which in this species is always much stronger
tlian in

D. camachoi

(Text-fig. 49)

D. camachoi

is

usually

(20-

30% overlap) slightly more involute and finer costate throughout

(Text-figs. 47, 49) D. undebayense and D. camachoi are associated
D. camain the lower part of the Pseudocidoceras zonule to which
.

choi appears to be restricted.

Bulletin 255

116

Dmm

McM

W%

H%

U%

F%

Alaskan ammonites,

Measurements.-

USNM

160254

^^^

Pt.

^^^

II:

^^^

Westermann

^^^

^^^,

147

Bulletin 255

148

- USNM 160256.
typicus. Sea cliff on southeastern

Repository.

Locus

shore of

Wide

Bay,

probably U.S. Geology Survey Mesozoic locality 19862.

Stratum ty pic u rn. SlvAes of the upper Kialagvik Formation,
jirobably lower Pscudocidoceras zonule.

Derivatio

no/tiiuis.

In

honour

of

Horacio Camacho,

Inst.

Geologia, Universidad de Buenos Aires, Argentina.

Age and

occurrence.

Wide

eastern side of

S.

Bay;

sowerbyi Zone, Bajocian, along south-

common

in

and possibly

restricted to the

lower part of the Pscudocidoceras zonide.

Diagnosis.
costae

large sjjecies of D. {Pscudocidoceras) with fine

becoming obsolete on body chamber.

Material.

Two

almost complete internal molds, two phragmo-

USGS
mold from USGS

cones and one complete body chamber from

(USNM
(USNM

21251

one complete internal

one complete sj>ecimen with damaged body chamber,
largely with test, from USGS 19863; (? the holotype and) one large
body chamber from USGS 19862; two well preserved, one poorly
preserved and one incomplete internal mold and ? one almost complete specimen from USGS 12405; two almost complete internal
molds and one plastercast from CAS 29011 (CAS 13111) three almost complete internal molds, two body chambers, three fragmentary phragmocones and ? one complete somewhat crushed internal
10 at 2-8 m (McM J 918-921, 923,
moid with test remains fiom
one phragmocone and one fragment from scree of
925, 926)
11 (McM J 924)
one damaged phragmocone with body chamber
from
13w (McM J 922). All from the Pscudocidoceras zonule
160259)

160257)

19801

WA

WA

WA

of the

S.

sowerbyi Zone, Kialagvik Formation,

Description and comparison

phragmocone

is

ivilfi

Wide

D. xvidebayense,

Bay.
n.

,v^.

The

"sub-coronate" and the whorl section strongly de-

pressed lenticular with quasi-midlateral edge throughout, similar

to D. xvidebayense, n. sp. (Text-figs. 46-48)

However, the umbilicus

somewhat larger, varying from 35 to 48% of the diameter,

somewhat finer and denser, with 12 to 17 primaries and (28) 31 to 48 secondaries (>25 mm D) per halfwhorl.
is

usually

and

the costation

Yet there
characters.

is

considerable overlap with D. xvidebayense in

The mature phragmocone

is

all

usually 55 to 80

these

mm

in

Alaskan ammonites,

Pt.

diameter with an observed range from 40

commonly

H9

Westermann

II:

to 85

mm,

as

compared

phragmocone diameter

of D. ividebayeu.se

The body chamber becomes markedly more

evolute while the

to the

(40-60

mm

smaller

D)

(absolute) whorl width remains approximately constant; the adult

diameter

is

most commonly 100-115

mm

65-80

vs.

bayense) with an observed range from 80 to 140

The
trast to

costation weakens

D. ividebayense

what denser

mm

D. in ivide-

mm.

on the adult body chamber,

(Text-fig. 49)

(11-18 per halfwhorl vs.

The

in

con-

primaries are some-

and usually become

10-13)

very blunt, often developing into oblique bullae which at least

on the internal mold tend to retract from the umbilical seam. Yet
some specimens become almost "coronate." The secondaries of the
body chamber are extremely fine and dense, (32) 38-50 per halfwhorl [vs. 18-27 (35?) in D. xvidebayense], and may become obsolete especially ventrally. As in all D. (Pseudocidoceras), primaries
and secondaries are strongly prosocline and the secondaries cross
the externside either evenly convex or somewhat arched.
The section of more or less the last whorl usually becomes
slightly to markedly ovate, with the rounded lateral edge moving
below mid-flank. The steepness of the inner flank and uml^ilical
slope (or umbilical slope) is increased on the lateral mold, especially
if

the

commonly

to the umbilical

preted as the
inally with

present umbolateral groove

seam

mold

much

(PI. 39; PI.

40

fig.

la)

developed adjacent
This groove is inter-

is
.

of a porous secondary shell layer, possibly orig-

organic substance, facilitating the better attach-

ment of the retractor muscles (see also Jordan, 1968)

Septum and suture are identical with D. loidebayejise.
Discussion.

The

differences of the often associated D. wide-

bay ense were given above; however, phragmocones, especially

when

incomplete, cannot always be identified specifically. D. camachoi

bears

much resemblance

Buckman, type

to

Docidoceras (Docidoceras) cylindroides

species of the genus,

which
and

rectiradiate to weakly prosoradiate ribs

collared peristome.

is

distinguished in the

in the less oblique

and

150

Alaskan ammonites,

Docidoceras (Pseudocidoceras

Pt.

?) cf. D.

II:

Westermann

camachoi,

n. sp.

151

PI. 42, fig.

Material. One
part of

tlie

complete body chamber, internal mold with

phragmocone from CAS 29011 (CAS 13120) Associated
.

with D. camachoi,

S.

Wide

soxverbyi Zone, Kialagvik Formation,

Bay.

The specimen

differs

from D. camachoi

in the

more weakly

prosoradiate costae which are retained at almost full strength on the

body chamber. This could be a 'stunted' variant of D. camachoi

which reached adulthood before the ribbing became modified,
similar as described from D. cf. icidebayense.

Measurements.CAS

13120

p^^

^^^

^^^

^^^

85
50

40
64

26
34

48

18

40

sp. indet.

(apert.)

(phragm.)

Docidoceras (Pseudocidoceras),

n.

c.42

PI. 36, figs. 4a, b,

One

incomplete deformed internal mold with

chamber from USGS 19862 (USNM 160252)
one almost complete internal mold from USGS 12405 (USNM
160251)
Associated with D. widebayense and D. camachoi, S.
Material.

pathological body

Wide

soxverbyi Zone, Kialagvik Formation,

This form
camachoi,

by the smaller diameter, the denser costation,

n.sp.,

(about 20 primaries on the two

especially regarding the primaries

last

Bay.

distinguished from D. xuidebayense, n. sp. and D.

is

halfwhorls of the phragmocone)

and the wider umbilicus

(U 40%)
Docidoceras

(?) [?

Pseudocidoceras or

n.

subgen.], n. sp. indet.

figs,

PI. 43,

la, b; Text.-fig.

50

Material. One incomplete phragmocone with crushed body

chamber, largely with shell, from
3 at the base of the amplectens zonule (McM J930)

WA

Description.

This

specimen

is

typically

'coronate,'

Telo-

two whorls of the phragmocone which are broad and strongly depressed 'lenticular' with
rounded lateral edge; the inner flank ('umbilical slope') and the
broad externside are gently and evenly rounded. The widely

ceras-Vike, at least

throughout the

last

spaced primaries consist of blunt plications barely reaching the

umbilical seam. There are strong mid-lateral spines on the

last

three

Bulletin 255

152

Cross-section of DocidoText-fig. 50.

crras \^}Pseudocidoceras or n. subgen.], n.

incomplete phragmocone,
base of the E. amplccicns zonule (McM J 930)
X 1sp.

loc.

indet.

WA

B,

at the

whorls of the phragmocone

(9 to 10

per halfwhorl) which, on the

inner whorls, are molcletl into the subsequent umbilical seam.

The

secondaries are moderately strong, rather dense (10 per halfwhorl)

and weakly prosoradiate.

The body chamber remains strongly depressed with dominant
quasi-midlateral nodes or spines; the inner flanks are smooth, the

secondaries become blunt, stronger prosoradiate and, on the internal mold, partly obsolete.
ly oblicpie

The

aperture

is

peristome, slight constriction and

simple with moderate-

weak

collar.

^\\h

specimen may be a strongly 'coronate' D.

(Psrndocidoceras): however, it is distinguished by the spines, tlie
Affinities.

blunt primaries and

tlie

chamber shows close

affinities to

whole

the

shell

(Tornquist)

The body
South American Pseiidotoites, and

almost rectiradiate secondaries.

resembles

P.

(?)

(see VV^estermann,

subgen.)

.sphaeroceroides

pi. 9, fig.

7); however, the

(n.

1964b,

whorls are not ovate and the secondaries stronger and almost
straight. This species is probably closely related with the associated
D.

(Pseiidocidoccras?) paucinodosurn, n. sp.

by somewhat smaller,
Measurements.

McM

1930

(apert.)

(phragm.)

less

depressed and

Dmm
c.lOO
54

W7f

H%

67

35

less

which

is

distinguished

ornate whorls.

U%

9-10

41

(c.35obsol.)

40

Alaskan ammonites,

Pt.

Westermann

II:

153

Docidoceras? (Pseudocidoceras?) paucinodosum Westermann,

PI. 43, figs. 2a, b; PI. 44, figs.

PL

Holotype.

mold except

slightly

n. sp.
2; Text-fig.

51

complete internal

43, fig. 2a, b. Text-fig. 51;

nucleus,

for

1,

deformed.

CAS

Repository:

13123.

Southeastern side of Wide Bay, Calif. Acad.

29014 (field no. 108/C.E.L. No. 68)
Stratum typicinu. "Dark grey sandstone with basaltic dike",
E. amplectefis zonule, upper Kialagvik Formation.
Locus

typicus.

Sci. locality

Derivatio uotninis.

With

respect to the widely spaced nodes

or spines.
Diagnosis.

species of Docidoceras

[or Pseudotoites}] with

evolute depressed whorls bearing distant reduced blunt primaries,

with strong lateral nodes or spines, and dense, straight to weakly

prosoradiate blunt secondaries.
Material.

The

mold

holotype; one partly distorted internal

WA

and two fragments from scree of

two almost
5 (McM J934)
complete internal molds, somewhat distorted, from subgreywacke
of uses 19828. E. amplectens zonule of the early 5. sowerbyi Zone;
?

WA

one fragment from the O. sauzei Zone of

12 (McM J 1044)
reworked]. All from the Kialagvik Formation at Wide Bay.

[?

Description.

The

approximately 20

mm

innermost whorls are not preserved. From

diameter, the whorls are depressed subellipti-

with more or
rounded lateral edge. The umbilicus is wide (35-40%) moderately deep and subconical. The phragmocone whorls embrace
about one-half, the umbilical seam barely touching the lateral
nodes. The body chamber occupies approximately one full whorl,
egressing especially with the last one-half whorl so that the aperture embraces only about one-quarter of the phragmocone. As
usual, width growth decreases or ceases with the body chamber.
The aperture is simple and moderately oblique, with slight
constriction and weak collar or lateral flange. The full diameter
is 75-85 mm, a modest size for a macroconchiate Pseudotoites or

cal or lenticular to slightly ovate in section, often

less

Docidoceras.

The

costation of the inner whorls

widely spaced

(7-9

per halfwhorl)

(internal

mold)

consists of

blunt primaries, fading away

near the umbilical seam, but ending laterally in strong nodes or

Bulletin 255

154

Cross-section of DociJoText-fig. 51.

crras f (Psrudocidoceras ?) paucinodosum, n. sp., hoiotype, with complete body

chamber;

Spines. Tliere are five to six times as

1-

many

densely

set,

straight or

somewhat prosoracHate secondaries which arise horn and between

the lateral nodes and run evenly over the broad externside.
On the long body chamber, the primaries of the internal mold
liecome blunt plications and retract from the inner flank (umi)ilical slo|>e)
or they may become obsolete. The lateral nodes
remain strong and usually rounded, more rarely bullae-like; but
may weaken at the end. The secondaries usually remain dense
becoming blunt to almost obsolete; however, they may again
increase markedly in strength and spacing near the end of the
body cliamber. A weak umbolateral groove may be developed.
The septum is bullate with the outer saddle axis being slightly
stronger than the inner one. The suture is unknown.
Comparison. The intermediate and outer whorls resemble
Docidoceras, especially D. (Pseudocidoceras), in the dimensions and
the strength of the lateral nodes/spines. However, tlie development
of l)lunt primaries, dense secondaries, and prominent nodes especially on the body chamber are strongly reminiscent of Soutli American Psexidotoites, such as P. trausatlnniiciis (Tornquist) which it
,

Alaskan ammonites,

Pt.

II:

Westermann

155

and coiling or P. (?) (n. subgen.)

which has similar "coronate" inner
whorls. This species is, therefore, intermediate between Docidoceras and Pseudotoitcs, and possibly closest affiliated with P. (?)
sphaeroceroides which siioiild probably be distinguished as a new
also resembles in

sphaeroceroides

thickness

(Tornquist)

subgenus.

Measurements.

Bulletin 255

156

and spacing and pass evenly convex over the externside.

aperture expands laterally into a weak flange with
thickened test which, on the internal mold, is separated from the
body chamber by a constriction. The collar or flange extends into
lateral lappets of which the left one is preserved with 5 mm
length. Their accurate original shape is uncertain but consistent
with the simj)Ie 'spatulate' type of Trilobiticeras (Buckman, 1919,
strength

The

pi.

110).

The

septal suture

is

incompletely preserved; nevertheless, the

two subecjual E/L and L/Uj saddles of Docidoceras and other

Ototidae witli bidlate septum can be recognized.
Affinities. Tlie phragmocone of this typical microconch cannot be distinguished from inner whorls of the much larger D.
(Pseudocidoceras) which abounds in the same bed (compare with
PI. 41, fig. 3; Text-figs. 47-49)

This specimen

is

distinguished from

all

described species of D.

(Trilobiticeras) by the strongly prosoradiate costae with ventrally

convex secondaries, and jMobably by the weaker apertural collar.

Tliese are the same featines which distinguish tlie probably corresponding macroconclis D. (Pseudocidoceras) and D. (Docidoceras).

Suggested
association,

sexual
tliis

diuiorpJiistn.

small sj^ecimen

is

Considering morphology

proI)al)ly tlie

and
male of D. (Pseudo-

Astoimding is the scarcity of this microconch,

specimen in a sample of approximately 500 Docidoceras
cidoceras).

i.e.

a single

s.s.

A num-

ber of possible reasons for such numerical miss-matches have been

suggested

(Westermann, 1964b,

could have caused

needed

this

p.

36), a combination of

strong disproportion.

which

More material

to place this form in

macroconch (female)

Measurements.Shell

L546

(apert.)

(phragm.)

is

dimorphic relationship and

species previously known only from its

to establish clearly tlie suggested

^^^
C.27

^^^

H^^

^j^^

Alaskan ammonites,

Pt.

II:

Westermann

157

Genus PSEUDOTOITES Spath, 1939

Type species by original designation. Stephanoceras leicharti

Neumayr, 1885, from the early Bajocian (s.s.) Newmarracarra Limestone of Western Australia.
Only the Australian representatives have previously been redescribed and revised (Arkell, 1954; Westermann, 1964b) while the
Soutii American species are still poorly known. The Alaskan forms
are described for the first time. My search in 1965 for new material of Pseudotoites in the Andes of Neuquen and Mendoza,
Argentina, was successful only with respect to P. singularis (Gottsche)

and

P.

(Latotoites)

abundantly just below

evolutum

blage of the O. saiizei Zone.

able of the

Emileia

the

No

somewhat dubious

ceroides (Tornquist)
tiniis Arkell, all

Otoites

significant

species

which occur

(Tornqtiist)
-

P.

new material

is

avail-

subgen.?) sphaero-

(n.

(?)

assem-

Papilliceras

P. transatlanticiis (Tornquist)

and

P. argen-

based on single or a few poorly preserved speci-

mens and inadequately described. Furthermore, the holotypes of

P. sphaeroceroides and P. argentinus had apparently not been returned after being lent to the late Dr. W. J. Arkell and were not
available for re-examination.

Subgenus PSEUDOTOITES (PSEUDOTOITES) Spath, 1939

The

following tentative diagnosis

(Pseudotoites): Large Otoitidae

is

here given for Pseudotoites

(macroconchs)

nucleus with sub-

rounded ovate or slightly depressed elliptical whorls and

reduced primaries which are retracted from the umbilical seam;

circular,

intermediate whorls inflated-cadicone with depressed strongly ovate

section, lateral tubercles or bullae absent or

weak; outer one to two

whorls becoming more evolute and compressed, ornament with

strong lateral nodes and bullae, primaries and often secondaries be-

coming obsolete. Costae prosoracHate throughout. Peristome simple

and strongly oblique.
Pseudotoites (Pseudotoites)

cf. P.

argentinus Arkell, 1954

Pis. 45, 46,

Text-figs. 53-56

Cf. 1954. Pseudotoites argentinus n. sp., Arkell (Australia), Phil. Trans. Roy.
Soc. London, B., vol. 237, p. 592, pi. 40, figs. la-c.

Material.

One

almost complete specimen with damaged body

chamber from USGS 19922

(USNM

160262)

one partly crushed

Bulletin 255

158

Text-fig. 52 a-c.
Internal septal suture of PsfuJotoites sp. indet. from the
P. singularis assemblage (S. soivrrbyi or
hasai O. sauzri Zone) of Los Molles (bed

Neuquen, Argentina; magnified, a.

mm W, b. at 4 mm
and 2.5 mm
H, c. at 4.5 mm
and 3 mm H. Note
the secondary Un lolie and the apparently
divided Ui lobe.
6) in
at \.2

USGS 12405 (USNM 160263) one fragmentary

specimen with parts of phragniocone and body chamber from
USGS 19862; two phragmocones witli parts of body chambers from
13w at C.18 m (McM J935, 936) one phragmocone with beginning of body chamber from VVA 10 at 11 m (McM J 938) All
moderately to well-preserved internal molds with test remains from
phragniocone tioin

WA

the P.seudocidocera.s zonule

Formation

at

Wide

Description.

of

the

S.

soicerhyi

Zone,

Kialagvik

Bay.

The

whorls are evolute

(40-45% U)

almost

'serpenticone' thoughout, with roinided siibcircular to slightly de-

becoming strongly ovate and often compressed on the last whorl and body chamber. The umbilical slope
(or inner flank) is only slightly rounded and slopes moderately
steep to the umbilical seam situated well outside the lateral nodes of
pressed ovate section,

the preceding wliorl.

The maximum whorl width

third whorl lieight or e\'en lower

the umbilical seam on

or

less

tlie

on

tlie

last

lies at about onewhorl and close to

body chamber. The externside

is

more

evenly and liighly arched.

The

costation consists of leduced short prosoradiate primaries,

strong lateial nodes or sh(jrt bullae, and prosoradiate densely spaced

blunt secondaries. There are 9-1 1 blunt primaries per halfwhorl on
the inner whorls, oidy slightly increasing in number on the outer
whorls; they arise gradually near the middle of the umbilical slope
and retract from the seam finally becoming obsolete or merely minor

Alaskan ammonites,

ph

50

20

Pt. II:

Westermann

159

Bulletin 255

160

Text-fig. 54.
Scatter with growth lines for relative umbilical width
of PsruJotoitrs spp. (Same symbols as Text-fig. 53). Note that the
growth rate increases throughout the phragmocone and body chamber, and that
separation of species is poor.

(U:D)

The septum

is

typically bullate.

with narrow finger-like endings of

a

common

radius, the

E/L

saddle

The

suture

is

slightly

is

highly complex

U2 and U3 reach
larger than the L/U^

all lobes; E,

L,

on one specimen; the U2/U3 saddle is rather broad

but only one-half as high as the L/Uo saddle; U^ is strongly oblique.
Discussion. \n naming this species for 'Emilcia aff. singunon
laris Gottsche' of Jaworski
(1926, p. 255, pi. 2, figs. 3a-c)
Gottsche, Arkcll (195-1, p. 592) nevertheless figured a new speci-

saddle, at least

Alaskan ammonites.

Pt. II:

Westermann

161

50

\X

40

*'^

K^.

C>X>^^

0-.

30

40

30

50

60

70

W7

Text-fig. 55.
Scatter with growth lines for relative umbilical width vs.
of Pseudotoites spp. from South America and
relative whorl width
Wide Bay (symbols as in Text-fig. 53). The combined parameters for 'coiling'
and 'thickness' illustrate the specific dimensional differences which may or may
not be significant.

(U%:W%)

men from

Espinazito Pass as the holotype. P. argentiniis

is

said to

be "much more evoliite and larger than Gottsche's original", but

neither description nor measurements were given. However, Jaworski's

measurement and

show

figures

that the

phragmocone of

his

specimen has much broader whorls than the holotype and that
Jaworski's specimen is indeed closely affiliated with P. transatlanticus (Tornquist) (Text-figs. 53,54) The inner whorls of the holo.

unknown because the umbilicus was not developed.

The affinities of the Wide Bay form with P. argeyitiniis, probably known only in the holotype, can only be conjectured. The
type are

former

is

almost certainly more evolute

c.

35%, aperture

42-477o

'^-y-

tralian

species,

tinguished by

P.

less

spaced secondaries.

leirharti

45-50%,

is

most

vs.

(end phragmocone

39%)

closely

Among

affiliated

c.

the Aus-

but

dis-

evolute whorls with denser primaries and wider

Bulletin 255

162

Measurements.

USNM

160263

Dmm

W^C'

H7r

u%

Alaskan ammonites,

Pt.

II:

Westermann

163

approximately constant while width appears to contract slightly

and gradually. The simple peristome is somewhat oblique and the

internal mold has a strong lateral flange following a constriction,
probably corresponding to

The

shell thickening.

primaries and especially the secondaries are prosoradiate

throughout.

The

nucleus (20

mm D)

has strong primaries which are

retracted from the umbilical seam, carry small lateral tubercles,

trifurcate into moderately strong, continuous

and

typically

and
'con-

On the intermediate and last whorls, the primaries

become increasingly shorter and more blunt, leaving a smooth band
besides the umbilical seam, and swell distally until they produce
strong bullae; the secondaries become blunt, much denser, weaker,
and finally obsolete on the body chamber leaving only broad plivex' secondaries.

cations.

Septum and sutiue are as in P. cf. argentinus described above;

the suture is complex with suspensive umbilical elements.
Of special interest is the presence on both specimens of a strong
umbolateral groove (here named) adjacent to the umbilical seam,
reaching halfway up the inner flank (or umbilical slope) and extending from the end of the phragmocone to the middle of the
body chamber where it becomes gradually obscure. A faint trace of
this groove is also visible on the inner whorls of the phragmocone.
The umbolateral groove contains remnants of a dark porotis material, the surface of which is in continuation with the internal
mold of the whorl. Common in the described Docidoceras (Pseudocidoceras) from

Wide

Bay,

it is

probably related to the attachment

which has been known in this posifrom slight impressions and organic remnants in many ammonites since G. C. Crick (1898) and has most recently been discussed by F. W. Jordan (1968)
Discussion. The two specimens differ from P. cf. argentinus
apparently only in the broader, more depressed, and slightly more
of the paired retractor muscle
tion

They closely resemble the poorly preserved holoknown specimen) of the somewhat dubious P.
(Tornquist) The holotype (refigured: Arkell, 1954,

involute whorls.
type

(the only

transatlanticus
p.

592,

whorl,

13), preserved with slightly more than one exposed

entirely more or less strongly crushed and its dimensional

fig.
is

features can therefore only be conjectured

(see table

below)

It is

Bulletin 255

164

Text-figs. 56 a-b.
Cross-section of PsruJotoitcs cf. P. argftitinus Arkell,
sunvrrbyi Zone of Wide Bay; XI. a. Almost complete specimen with damaged
body chamber, loc. USGS 19922 (USNM 160262). b. Phragmocone, loc.
13
in the Pstudociduceras zonule (McM J 935).

S.

WA

apparent that the

moderately

The
3)

prolialily

thick, evohite

last

whorl of

and depressed ovate

tlie

evohite specimen described by Jaworski

as 'Ernileia aff. singularis Gottsche'

argentinus,

is

also close to P.

phragmocone

is

in section.

(1926, pi.

and placed by Arkell

transatlanticus, which,

in

2,

fig.

in P.

turn,

is

intermediate in coiling and wiiorl sections between P. singularis

and

P. argentinus.

Alaskan ammonites,

Pt. II:

Westermann

165

Cross-section of PscudoText-fig. 57.

(Tornquist),
cf. P. transatlanticus

toitcs

complete on one side only, loc. USGS

the
Pscudocidoccras zonule
19862
in
(USGS 160264); note the umbolateral
groove on the body chamber; X 1-

Measurements.

Dmm

USNM

160264

(apert.)

W%

H%

U%

Bulletin 255

166

Appendix

to incdsiircnicuts.

H% U%

Dmm W7r

Holotype of P. transatlantlcus (from plastotype)

(end "phragm. body ch.)
37.5
68
c.50
(phragm.)
60
c.50
c.33.5

10
10
10

c.50
c.50

C.48

'E mill HI aff. singulinis

C.54

C.37

c.3+
36.5
c.35

45-50

[aworski", tion Gottsche (from text and figs.)

(body ch.)
(phragm.)

c.80

c.40

c.35

c.65

c.30
c.33

c.40

c.40
9

c.37

c.41

c.42

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1956. Phylogcnie der Stephanocerataceae und Pcrisphinctaceae des Dogger.
Neues Jahrb. Geol. Palaontol., Abh. 103, pp. 233-279.
1964a. The ammonite fauna of the Kialagvik Formation at Wide Bay,
Alaska Peninsula. Part I. Loiver Bajocian (Aalenian). Bull. Amer.
1963.

lirns. Boll. Soc. Paleontoi.

Paleont., vol. 47, No. 216, pp. 325-503, pis. 44-76.

Sexual-Dimorphismus bci
mmonoideen und seine
die Taxionomie der Otoitidae (cinschl. Sphaeroceratinae ;

\96A-h.

Bedeutung fiir
Ammonitina. M.

Jura). Palaeontographica, Abt. A, vol. 124, pp. 33-73, pi. 6-9.

1964c. Occurrence and significance of the Arctic Arkclloceras in the middle
Bajocian of the Alberta Foothills (Ammonitina, Jurassic) Jour. Paleont.,
.

vol. 38, pp. 405-409.

Coi'ariation and taxonomy of the Jurassic ammonite Sonninia adicra

(JVaagen). Neues Jahrb. Geol. Palaontol., Abh., vol. 24, pp. 389-412.
1967. Sucesion de ammonites del Jurasico Medio en Antofagasta, Atacama,
Mcndoza, y Neuquen. Rev. Asoc. Geol. Argentina, vol. 22, No. 1, pp.

1966.

63-73.

Whiteaves, J. F.
1876. Mcsozoic Fossils, vol. 1. On some invertebrates from the coal-bearing
rocks of Queen Charlotte Islands, part 1. Geol. Sur. Canada, pp. 1-262,
pis.

1-32.

from the Rocky Mountains three miles north of the east end of
Devil's Lake. Geol. Sur. Canada, Contr. Can. Palaeont., vol. 1, Part 2,
pp. 163-172.

1889.

Fossils

Zittel, K. A. von
1869. Bemerkungen iiber Phylloceras tatricum Pusch, sp. und einige
Phylloccras-Arten. Jahrb. K. K. Geol. Reichsanst., vol. 19, pp. 59-68,

andere
1

pi.

Bulletin 255

172

$ayHa a>!Monj!T03 d^opMauxu Kna;iarEHK y sajiHsa IL'npoKoro (yay^

Soiia Sonnia sowerbyi
noJiyocTpoB A-iHcnao 4acTb San.
Tepj^ E. r. BecTepMaHH,

Ciofi)*

KoHcneKTo

TpaHKua MeKflv f^opMauHHMH KHa.iarBHK h lUejiHKOEa b HacTon^ee

BpewK npoBe;;ena y ocKosaHHH Ka.njioBvaHCKoro corjiacHH (napaKOH;
3T0 coTjiacyeTCH
MaeT CHJibHuJi reTepoxpaHHSM.

(JiopMHocTK)

ocHOBHhiM onpe.nejieHiieM a hckjdo-

3oKa S. sowerbyi ( cTaHflapTHan) coctoi!T H3 HUNCHeil nacTH no

Kpa{*Heii Mepe To;in;i!i;oit Cojiee 35 MeTpoB, cocTOflinef! h3 noArpefisaKH, rpef^saKii h a.neBpoJiHTOB , coAsp^.a^nx naGop I'.uJmetoceras anplectens [amplectens zonule] h H3 sepxHefi ^aCTH, 50 - 100 MeipoB, cocTOflmefi H3 rjiHHKCTHX cjiaKueB c KOHKpeuHHMu coAepxamiiMH Ha6op Pscudocidoceras [i'seudocidoccras zonule], qacTb HenpoMHbix cjiaHues ^acTO
HexaaTaeT H3-3a cdpocoB pa3;;ejia Ha^ a^plcctcns zonule. HnjKe.-ierKaD;afl 30Ka E, hoKclli
0T;;eJieHa ot BepxHeK 30Jibi 80 - 100 MexpawH
oSHaY.eHHHX, HeMbix h Ma;io:iccj-feflOBaHHbix KJiacTHMHUx nopoAi nepexpHBa-ocafl 30Ha 0. sauzci oTj;e;ieiia necyaHiiKaMH H rjiHKiicTUM cianueM oSrieJl .v.O!r,HOCTb!o b 110 - 130 MeipoB,

Anplectens zonule ;;aj:a Hudmetoccras (Euaptctoccros') amplectens

(BucVman), Docidoceras (?), Bradfordia? (Praconpclia) H Hobctoxyites h noSTOMy ee npi;ypa"-;;iBa-'jT b Hn:sHrJ-'j S; sowerbyi 30Hy, L.discites nofl30Hy, K caMbn.1 HK3aM 6aH0ca.
SoHyjia Pseudocidoccras BK.iKj^aeT , bhh3y, Sonninia (Uuboploccras) Eudtictoccras klinakomphalum (Vacck) ii Docidoceras s. s., noflTBep-?;!iaH TaK>"e l. discites no^soHy h Bbime- oShjib,

HHH

Witchellia

3bojik)T

Ha

yKa3UBa>D-aii>*

cpeflH-ora

h;ih

BepxH?OK)

30Hy

S.

so'.verbyi

Ammonoidea pa3;;ejieHH Ha:

15 po;!,OB, hk oflHoro HOBoro; 13 uoji,VOJIOB - H3 KOTopbiX 3 HOBHX [Sonninia (Alaskoceras)
Docidoceras (Pseudocidoceras) jRradfordia ? (Praeoppelia]
30 bh;;ob - k3 KOTopux ot 13 ;3,o
20 HOBbix M 9 H3 KOTopHX Ha3HBaKiTCH [Partschiceras ellinticun, Pseudolioceras costi striatium, Sonninia (Uuhoploceras) bifurcata, S, (Alaskoceras)
alaskensis, 1','itchellia sutneroides, Docidoceras (Pseudocidoceras) widebayense,
D. (P.) camachoi, Bradfordia?, (Praeoppelia) oppeliiforinis
h 2 no;iBn;|Ta ]
[Pseudolioceras nclintocki fastigatun,' Eudtnetoceras (Euaptetoceras)
klinaKonphaliim discoidale].
,

poAOB h no;ipoflOB Anmonitina pacnpocxpaHeHH'

Cpo;i;cTBO hx Sojiee 6jiH3Koe c EBponon h noTOM c I-CnchoJi AuepHKot:. CpoflCTso c SanaflHoi^ THxooKeaHcxoii ofijiacTbfo (SanaAHofi ABCTpajiJief! h MHAOHesiie-J:) cjiadoe h kof.ho c^HTaTt,
Bee
MTO OHO npoHSOEJio KOCBeHHo, '-!epe3 EBpony h }>cHy.-3 AMepuK.y.
po;ibi c ;ipyrnx MarepiiKOB HsaecTHLi h TOJibKO Pscudotoitcs npnypo^eHhi
BucoKoe ttiayHiiCTJiMecKoe pa3HoodncK;i>DM;iTe'jibHo K Ti'.koN'.y OKeany.
pa3He yKa3HBaeT Ha SoJiee HH3:<yK) niipoTy h/iijih 6ojiee H;j3Kne re'ineCkojio ^o.^OBKH^r

innpe

^eu

paTvpH,
BH;',0B.

EBpone.

xie.M

HacTOflmee BpevH, mto TaKxe noKa3aHo pacnpe^OjieHHeM

PLATES

Bulletin 255

174

Explanation of Plate

from the mouth

Southeast side of Wide Bav, middle part, tele-photograph
Formation forms
Kialagvik
The
bay.
the
of
side
other
the
on
Creek
of Short
formation the upper slopes and
the coastal cliffs and lower slopes, the Shelikof
cliffs.

The numbers

Text-figs. 2-3).

indicate

my

fossil

localities

(WA-prescnpt

omitted,

cf.

Bull. Amer. Paleont., Vol. 57

Plate

Bull. Amer. Paleont., Vol. 57

Plate

Alaskan ammonites,

Pt.

II:

Westermann

Explanation of Plate

175

Bajocian section of the upper Kialagvik Formation at southeast shore of

1
and 4), viewed from the
Bay, 1.7 km east of Preston Creek (Iocs.
east. The bluff is formed by the massive subgreyvvackes of the E. amplectcns
overlain by several meters of deteriorated
zonule (lower
so'u:crbyi Zone)
shale marking a bedding-plane fault zone (F) which has suppressed the shales
with concretions of the Pscudocidoccras zonule; the middle of the section consists of unfossiliferous
interbedded silty shales, sandstones and mudstones
bearing a sedimentary dyke (s.d.) in a small steeply dipping fault; at the top
are thick-bedded greywacke, subgreywacke siltstones, and mudstones of the
Parahigotitcs zonule (O. sauzci Zone).

WA

Wide

.S".

Bulletin 255

176

Explanation of Plate

Coastal bluff of the E. amplectens zonule (lower S. soiverbyi Zone). Above:

east of Preston Creek (Loc.
massi\e fossiliferous subgreywacke
2)
underlain by poorly fossiliferous interbedded silty shale, mudstone and greywacke (cf. PI. 4). Below: 1.8 km east of Preston Creek (Loc.
massive
8)
fossiliferous subgreywacke truncated by steep fault with basaltic dyke; in the
middleground are Bajocian interbedded shales, sandstones, and mudstones with
at least most of Pseudocuiocrras zonule missing (probably bedding-plane fault
above . amplectrtis zonule)
in the background are shales, siltstones, and
sandstones of the Callovian Shelikof Formation.
1.2

WA

km

WA

Plate

Bull. Amer. Paleont., Vol. 57

-i'-'j

sr

"^i'K
1

^Wtl

v*^'

^ A?^

'

vij^ v^ ^'

'

.V'

Bull. Amer. Paleont., Vol. 57

a\ :!

Plate

Alaskan ammonites,

Pt. II:

Westermann

177

Explanation of Plate 4
Coastal bluff of the E. amplcctens zonule (lower
agvik Formation; 1.2 km east of Preston Creek (ioc.
cf. Pi. 3). Left: Interbedded silty shales,
mudstones,

S.

soiucrbyi Zone)

WA

Kial-

marked by arrow;

and greywackes, below'

and massive subgreywacke with fissility due to bedding-plane faulting'
(cf. Pi'
2), above. Right: Detail of greywacke beds of locality
2, graded with sole

WA

structures.

178

Bulletin 255

Explanation of Plate

Coastal cliff of the PsiudociJocnas zonule (S. so'ujcrbyl Zone), Kialagvik

2.5 km east of Preston Creek (loc.
10; marker bed at 10.5 m
indicated); highly fossiliferous beds \vith calcareous concretions. The overlying unfossiliferous interbedded silty shales, sandstones, and mudstones, and
the greywacke and subgreywacke beds of the O. sauzf't Zone are largely covered by vegetation. Abo\e follow the shaley, arenaceous, and partly carbonaceous and tuffaceous beds of the Callovian Shelikof Formation.

Formation;

WA

Bull. Amer. Paleont., Vol. 57

Plate

Plate

Bill. Amer. Paleont., Vol. 57

*-'::%
;7

It,
,'.-J

Alaskan ammonites,

Pt. II:

Westermann

Explanation of Plate

179

Detail of the coastal bluff of the lower E. hoivcUi Zone, middle Kialagvilc
Formation; northwest shore of Wide Bay, near Pass Creek (Iocs.
9-11).
Somewhat weathered muddy subgreywacke, above, and mudstone, below, with
calcareous concretions containing abundant wood remains and ammonites', particularly Erycitoidcs lioixflli (White) and E. (Kialagvikitcs) kialag'vtkfnsis

WB

(White).

130

Bulletin 255

Explanation of Plate
Coastal bluff of the upper E. hoivelli Zone,
unfossiliferous; west end of northwest shore of

middle Kialaffvik FormaUon;

Wide Bay Bedded sandstones

mudstone, followed by massive
overlaid by conglomerate and angular-bedded
subgreywacke.

Bull. Amer. Paleont., Vol. 57

Plate

Plate

Bull. Amer. Paleont., Vol. 57

'

^'^m
I

Alaskan ammonites,

Pt. II:

Westermann

Explanation of Plate
Natural size

if

not otherwise indicated.

Page

Figure
1,2.

181

Partschiceras ellipticum Westermann, n. sp

38
Kialagvik Formation, S. sowrrbyi Zone, Wide Bay. la-c.
Damaged internal mold, totally septate, from 8 m below top
of E. amplcctcns zonule. Loc.
8.
McM J 962. 2. Damaged
internal mold with septal suture, totally septate, from the
Pseudocidoccras zonule. Loc.
10 at 2.5m. McM J 1013.

WA
WA

3a-c.

Holcophylloceras costisparsum Imlay

Kialagvik Formation, S. soiverbyi Zone, Wide Bay. Damaged
internal mold with septal suture, totally septate. Loc. USGS

4.

Phylloceras (Zetoceras) cf. P. zetes (d'Orb.)

Kialagvik Formation, S. soiverbyi Zone, Pseudocidoccras zonule.
Fragment of phragmocone with septal suture and test remains.

19862.

Loc.

USNM

WA

40

160921.

10 at 15m.

McM

J 1018.

36

Bulletin 255

182

Explanation of Plate
Natural size

if

not otherwise indicated.

Page

Figure
la-b.

Holcophylloceras costisparsum Imlay

40

so^vrrhyi Zone, Wide Bay. Damaged

Kialag\ik Formation,
phragmocone with incomplete one-half whorl body chamber,
internal mold with test remains. Loc. USGS 212S1, USNM
S'.

160920.
2, 3a-b.

Lytoceras sp. aff. L. eudesianum (Orb.)

Kialagvik Formation, S. soiccrbyi Zone, Pscudocidoccras zonule,
\\'ide

WA

Bay.

Totally

septate,

incomplete

McM

1010,

internal

mold.

Loc.

Large phragmocone,
probably with beginning of body chamber, internal mold with
test

10,

basel 2 m.

remains. Loc.

WA

3a-b.

10 at 14-16m.

McM

J 1005.

42

Plate

Bull. A.mek. Paleoxt., Vol. 57

"^i^

Plate

10

Alaskan ammonites,

Pt. II:

Explanation of Plate
Natural size

if

Westermann

183

10

not other\vise indicated.

Figure
l-2b.

Page
sp. aff. H. hebes Buckman
Kialag\ik Formation, S. soivcrhyi Zone, E. amplcctens zonule,
Wide Bay. la-h. Complete specimen with left lappet. Log.
5. McM J 1043. 2a-b. Almost complete specimen. Loc.
8
upper part. McM J 1041.

Hebetoxyites

44

WA
WA

3-6.

Bradfordia? (Praeoppelia) oppeliiformis Westermann,

n.

sp.

Kialagvik Formation, S. soivcrhyi Zone, Wide Bay. 3a-c. Phragmocone with incomplete crushed body chamber; 3a. inner
whorls; 3b. section of phragmocone. Loc. USGS 21252. USNM
160923. 4a-b. Inner whorls of specimen with incomplete body
chamber from the E. amplcctens zonule. Scree of Loc.
5.
McM J 1040. 5. Venter of penultimate phragmocone whorl;
same specimen as PI. 12, fig. 1. 6. Holotype penultimate
phragmocone whorl; see PI. 11, figs. a-c.

WA

48

Bulletin 255

184

Explanation of Plate
Natural size

if

11

not otherwise indicated.

Figure
a-c.

Page

Bradfordia? (Praeoppelia) oppeliiformis Westermann,

n.

sp.

?) S. soiverbyi Zone, Wide Bay.

Holotype, phragmocone %vith one-half whorl, incomplete and
damaged body chamber, b-c. Ventral views of ultimate and
penultimate half-whorls with section. Loc. USGS 19863,
160922 (see PI. 10, fig. 6).

Kialagvik Formation, (lower

USNM

48

Bull. Amer. Paleont., Vol. 57

Plate

11

Bull. Amer. Paleont., Vol. 57

Plate

12

Alaskan ammonites,

Pt. II:

Explanation of Plate
Natural

size

if

Westermann

12

not otherwise indicated.

Figure
1, 2.

Page
Bradfordia? (Praeoppelia) oppeliiformis Westermann, n. sp.
Kialagvik Formation, S. soiuerbyi Zone, Wide Bay. 1. Complete
phragmocone. Loc. CAS 29011. CAS 13109. (see PI. 10, fig.
5). 2. Probably complete phragmocone. Loc.
WA-111 lot
37.

UW

UW

13180.

48

Bulletin 255

186

Explanation of Plate
Natural size

if

13

not otherwise indicated.

Page

Figure
1, 2.

Pseudolioceras macklintocki fastigatum

Westermann, n. subsp
Kialagvik Formation, S. soivrrbyi Zone, Wide Bay. la-b. Holotype, phragmocone with incomplete body chamber. Loc. USGS
21252. USNM 160294. 2. Phragmocone, internal mold. Loc.

CAS
3-6.

29011.

CAS

13110.

Pseudolioceras costistriatum Westermann,

3a-b.

Hoiot> pe,

almost

complete

internal

n.

56

sp

mold with one-half

whorl body chamber, from the upper Psfudocidoccras zonule.

11 at 30m. McM J 1056. 4a-b. Small
?juv.) specimen, partly with test; +b detail of 4a, X2. Loc. USGS 19862,
USNM 160295. 5. Small ( ?juv.) specimen with some test, from
the Psfudocidocrras zonule. Loc.
11 at 30m. McM J 1056a.
phragmocone. Same loc. as 5. McM
6. Fragment of large
Loc.

WA

WA

J 1056b.

52

Bull. Amer. Paleont., Vol. 57

Plate

13

Bi'LL.

Amer. Paleont., Vol.

57

Plate

14

Alaskan ammonites,

Westermann

Pt. li:

187

Explanation of Plate 14
Natural

size

if

not otherwise indicated.

Figure
1-7.

Page

Asthenoceras sp. aff. A. nannodes (Buckman) $ and 9

From a single calcareous concretion of the Kialagvik Formation,

61

WA

Psiudocidoccras zonule,
soivcrbyi Zone, Loc.
10 at 11m.
la-b. Almost complete microconch
lb X2.
J 1037c.
2a-b. Complete microconch with lappet; 2b X2.
J 1037b.
3a-b. Complete microconch with exfoliated body chamber and
base of lappets; 3b X2,
J 1037a. 4a-b. Incomplete
specimen with some test and preserved septal suture, probably
septate up to end, macroconch ?; 4b X2.
J 1036a. 5 a-b.
Almost complete macroconch with one-half whorl body chamber, internal mold.
J 1035b. 6. Macroconch with about
one-quarter whorl body chamber and test remains.
J 103 5. 7. Macroconch with one-quarter whorl body chamber,
internal mold.
J 1035a.
-S".

McM
McM

McM

McM

McM

McM

McM

8, 9.

Eudmetoceras (Euaptetoceras) klimakomphalum

discoidale Westermann, n. subsp
sowcrbyi Zone, Wide Bay. 8. Paratype,
small
?juv.) phragmocone with crushed body chamber, test
remains and keel. Loc. USGS 12405, USNM 160928. 9. Phragmocone, internal mold. Loc. USGS 12405. USNM 160929.

Kialagvik Formation,
(

S.

75

188

Bulletin 255

Explanation of Plate
Natural size

if

15

not otherwise indicated.

Page

Figure
a-b.

Eudmetoceras (Eudmetoceras)

aff.

E.

eudmetum

Buckman
Kialagvik Formation,
internal mold. Loc.

72
S. soivcrbyi

USGS

Zone,

21252,

Wide

USNM

Bav. Fully septate

160926.

Bull. Amer. Paleont., Vol. 57

Bull. Amer. Paleont., Vol. 57

Plate

16

Alaskan ammonites,

Pt. 11:

Explanation of Plate
Natural

size if not

189

16

otherwise indicated.

Page

Figure
1, 2.

Westermann

Eudmetoceras (Euaptetoceras) klimakomphalum discoidale

Westermann, n. subsp
Kialagvik Formation, S. soiverbyi Zone, Wide Bay. la-f. Holomold; Ic-d, nucleus of same specimen, XI and X2 le, ultimate half-whorl removed; If, septal
suture on ultimate half-whorl, X2. Loc. USGS 12405. USNM
160927. 2. Fragment, probably of the same subspecies, with
test showing strongly projected growth lines. Loc. USGS 21251,
type, fully septate internal
;

USNM

160928.

75

Bulletin 255

190

Explanation of Plate
Natural

size if not

17

otherwise indicated.

Page

Figure
la-c.

Eudmetoceras (Euaptetoceras) klimakomphalum

discoidale Westermann, n. subsp.
Kialag\ik formation, S. soivrrhyi Zone, Wide Bay. Incomplete

75

large phragmocone; lb, penultimate whorl, partly with preserved hollow-floored keel. Loc. USGS 19863.
160230.

USNM

2a-b.

Eudmetoceras (Euaptetoceras)

sp. aff.

E.

nucleospinosum

Westermann

81

Kialagvik Formation, S. soiverbyi Zone, Wide Bay. Large phragmocone, partly crushed; 2b, inner whorls of same specimen.
Loc.

USGS

12405.

USNM

160231.

Bull. Amer. Paleont., Vol. 57

Plate

17

Bull. Amer. Paleont., Vol. 57

Plate

18

Alaskan ammonites,

Pt. 11:

Explanation of Plate
Natural

size if not

Westermann

18

otherwise indicated.

Figure
la-b.

191

Page

Eudmetoceras (Euaptetoceras) klimakomphalum

discoidale Westermann, n. subsp
Kialagvik formation, S. soivrrhyi Zone, Wide Bay. Incomplete

Cf.

75

phragmocone with approximately one-quarter whorl body

chamber and test showing faint concentric markings and
strongly projected growth lines. Loc. 21251. USNM 160232.
2a-b.

Eudmetoceras (Euaptetoceras) klimakomphalum

discoidale Westermann, n. subsp
Kialagvik formation, S. sovjerbyi Zone, Wide Bay. Incomplete
large phragmocone. Loc. USGS 19863, USNM 160233.

75

Bulletin 255

192

Explanation of Plate
Natural

19

size if not otherwise indicated.

Figure

Page

Eudmetoceras (Euaptetoceras) amplectens (Buckman)

83

?subsp. 'aguilonia' Imlay]

Formation, S. soiverbyi Zone, E. amplectens zonule,

[var. or

Kialagvik
Wide Bav.

0.75. Loc.

Largest

WA

8.

specimen,

McM

968.

with

aperture

(D 200mm)

Bull. Amer. Paleont., Vol. 57

Plate

19

Bull. Amer. Paleont., Vol. 57

Plate

20

Alaskan ammonites,

Pt. II:

Westermann

193

Explanation of Plate 20
Natural

size if not

otherwise indicated.

Page

Figure
a-c.

Eudmetoceras (Euaptetoceras) amplectens (Buckman)

83

?subsp. 'aguilonia' Imlay]

Kiaiagvik Formation, S. soiucrbyi Zone, E. amplectens zonule,
Wide Bay. a, damaged specimen with aperture, Loc.
8
upper part; b-c, part of body chamber removed.
[var. or

WA

Bulletin 255

194

Explanation of Plate 21
Natural

size

if

not otherwise indicated.

Figure
1,2.

Page

Eudmetoceras (Euaptetoceras) amplectens (Buckman)

Kialagvik Formation,
soivrrbyl Zone, Wide Bay. la-d.
.S".

83
Fully
outer

internal mold with test remains; Id, part of

whorls removed. Loc. USGS 12405. USNM 160234. 2. Fragment with aperture, from E. amplectens beds. Loc.
8.

septate

McM
3.

Cf.

WA

J 952.

Eudmetoceras

s.l.

indet. [?

c^

90

Kialagvik Formation, S. soicerbyi Zone, E. amplectens zonule,

Wide Bay. Plasticine cast of natural mold. Loc.
8 lower
part.

McM

WA

958.

Bull. Amer. Paleont., Vol. 57

Plate

21

Bull. Amer. Paleont., Vol. 57

Plate

22

Alaskan ammonites,

Pi. 11:

Westermann

195

Explanation of Plate 22
Natural size

if

not otherwise indicated.

Figure
a-b.

Page

Planammatoceras (Pseudammatoceras
(Hoffman)

?) sp. aff. P.

benneri

Kialagvik Formation, S. soivcrhyi Zone, Pseudocidoccras zonule,

Wide Bay. Large specimen with aperture (D 195mm); a,
X 0.7; b, last half-whorl of body chamber and part of phragmocone removed, X 1. Loc. USGS 21251. USNM 160235.

90

Bulletin 255

196

Explanation of Plate 23
Natural size

if

not otherwise indicated.

Page

Figure
1-2.

Sonninia (Euhoploceras) bifurcate Westermann,

Kialag\ik Formation,

phragmocone

A^.

soivrrhyi Zone,

Wide

n.

sp

Bay. la-b. Holo-

one-quarter whorl of somewhat

mold. Loc. USGS 12405.
160236. 2. Body chamber fragment, slightly crushed.
Loc. USGS 19869.
160237. 3a-b. Inner whorls of
spinose variant; 3b, ultimate one-quarter whorl removed.
type,

crushed

body

\vith

chamber,

internal

USNM

USNM

Loc.

USGS

12405.

USNM

160238.

94

Bull. Amer. Paleont., Vol. 57

Plate

23

Bull. Amer. Paleont., Vol. 57

Plate

24

Alaskan ammonites,

Pt. II:

Westermann

197

Explanation of Plate 24
Natural

size

if

not otherwise indicated.

Figure
1-2.

Page
Sonninia (Euhoploceras) bifurcate Westermann,

n.

sp

Kialagvilv Formation, S. soiuerbyi Zone, Wide Bay. la-c. Phragmocone with beginning of body chamber, compressed and
weaiv'ly ornate variant. Loc. USGS 12405.
160240. 2.
Incomplete phragmocone with one-half whorl of body chamber, spinose variant. Loc. USGS 19869.
160240.

USNM

USNM

94

198

Bulletin 255

Explanation of Plate 25
Natural size

if

not otherwise indicated.

Page

Figure
1-2.

Sonninia (Euhoploceras) bifurcate Westermann,

n.

sp

Kialag\ik Formation, S. soiverbyi Zone, Pseudoc'tdoceras zonule,

Wide Bay. la-b. Large body chamber fragment with weak
secondaries, internal mold. Loc.
15 at 10m. McM J 1049.
Incomplete phragmocone with somewhat crushed bodv
2.
chamber fragment. Loc. USGS 19869. USNM 160242.

WA

94

Bull. Amer. Paleont., Vol. 57

Plate

25

Bull. Amer. Paleont., Vol. 57

Plate

26

Alaskan ammonites,

Pt. II:

Westermann

199

Explanation of Plate 26
Natural

size

if

not otherwise indicated.

Figure
1-2.

Page
Sonninia (Euhoploceras) bifurcate Westermann,

n. sp.

Kialagvik Formation, S. soiufrhyi Zone, Wide Bay.

1
Phragmocone with one-half whorl of body chamber, terminally
deformed; average form, from the Pseudocidoccras
zonule
Loc.

WA

10.

McM

975.

2.

Phragmocone with one-half

average form

whorl of somewhat crushed body chamber;

Loc.

USGS

12405.

USNM

160241.

94

Bulletin 255

200

Explanation of Plate 27
Natural

size if not

otherwise indicated.

Figure
1.

Page

Sonninia (Euhoploceras ?) sp. indet

Incomplete phragmocone, from E. amplccte?is zonule,
Zone,

Kialagvik Formation. Loc.

WA

8,

Wide

102
5. sonuerbyi

Bay,

McM

J 959a.

2-7b.

Sonninia (Alaskoceras) alaskensis Westermann,

Kialagvik Formation,

n.

sp

103

soivrrbyi Zone, Pscudocidoceras zonule

subjacent shales). Wide Bay. 2. Internal mold of phrag(
mocone. Loc.
13w,
J 1023a. 3a-c. Holotype, phragmocone with some test and hollow-floored spines developed
from the surrounding matrix, from scree of the Pscudocidoceras zonule; 3c, ultimate half-whorl removed. Loc.
10.
J 1021. 4. Inner whorls of phragmocone, internal mold,
Loc.
15 at 22m.
J 1024. 5. Almost complete internal
mold with test and sockets of hollow-floored spines. Scree
of Loc.
10.
J 1020. 6a-b. Nucleus of phragmocone,
partly with test; 6b
10.
2. Loc.
J 1021a. 7a-b.
Parts of large phragmocone with hollow-floored spines developed from surrounding matrix, ventral views. Loc.
15
at 3m.
J 1027.

S.

WA

McM

WA

McM

WA
WA

McM

McM

McM

WA

McM

WA

Bull. Amer. Paleont., Vol. 57

Plate

27

Bull. Amer. Paleont., Vol. 57

Plate

28

Alaskan ammonites,

Pt. II:

Westermann

201

Explanation of Plate 28
Natural

size if not

otherwise indicated.

Figure
l-2b.

p^g^
Witchellia sutneroides Westermann, n. sp
116
Kialagvik Formation, S. suivrrbyi Zone, Wide Bay. la-c. Holotype, phragmocone with test remains. Loc.
USGS 21251.
USNM 160243. 2a-b. Almost complete internal mold with onehalf whorl of body chamber, compressed, involute and weakly
ornate

USNM

variant;
160255.

2b,

penultimate

whorl.

Loc.

USGS

12405

Bulletin 255

202

Explanation of Plate 29
Natural size

if

not otherwise indicated.

Page

Figure
1-2.

Witchellia sutneroides Westermann, n. sp

116
Kialagvik Formation, S. soivcrbyi Zone, (upper) Pseudocidoceras zonule. Wide Bav. la-b. Incomplete phragmocone. Loc.
10 at 14-16m. McM J 949. 2. Hollow-floored keel, partly
broken, on beginning of body chamber, with conellae-like
10 at 13-14m. McM
substance on keel floor, X 3. Loc.

WA

WA

J 943.

3a-b.

116
Witchellia cf. W. sutneroides Westermann, n. sp
Incomplete internal mold with one-half whorl of body chamber.
S. soicerbyi Zone, Kialagvik formation, Wide Bay. Loc. USGS
19868.

USNM

160247.

Bull. Amer. Paleont., Vol. 57

Plate

29

}
Bull. Amer. Paleont., Vol. 57

Plate

30

Alaskan ammonites,

Pt. II:

Westermann

203

Explanation of Plate 30
Natural

size if not

otherwise indicated.

Figure

Page
l-3b.

Witchellia sutneroides Westermann, n. sp

Kialagvik Formation, S. sowcrbyi Zone,
Wide Bay lab
Large body chamber fragment. Scree of Loc.
13. McM
^50. 2a-b. Immature (or microchonchate
?) specimen with
one-half whorl of body chamber. Loc. USGS
19922 USNM
160246.
3a-b
Immature {}) specimen with one-qua'rter
whorl of body chamber. Loc. 12405. USNM
160347

WA

uq

Bulletin 255

204

Explanation of Plate
Natural size

if

31

not otherwise indicated.

Figure
a-c.

Page
Witchellia sutneroides Westermann, n. sp
116
Kialagvik Formation,, S. soiucrhyi Zone, Wide Bay. Largest
phragmocone, with apparent shell duplication (see Text-fig.
38), X 0.9; b-c, the single photograph has been cut because
of space

limitation.

Loc.

USGS

21251.

USNM

160248.

Bull. Amer. Paleont., Vol. 57

Plate

31

Bi

LL. A.MER.

Paleoxt., Vol. 57

Plate

32

Alaskan ammonites,

Westermann

Pt. II:

205

Explanation of Plate 32
Natural

size

if

not otherwise indicated.

Figure
l-2b.

Page
Pelekodites cf. P. pelekus Buckman
126
Kialagvik Formation, S. soivcrhyi Zone, Pseudocidoccras zonule,
Wide Bay. la-b. Complete specimen with test and lappet; lb

X2

Loc.

WA

vation and
3-4b.

loc.

10 at

11m.

as fig.

McM
2b

J
2.

1028b. 2a-li.
J 1028a.

Same

preser-

McM

Pelekodites (Spatulltes ?) n. sp. aff. P. spatians (Buckman)

127
Kialagvik Formation, S. soiucrbyi Zone, upper Pseudocidoccras
zonule. Wide Bay. 3. Incomplete specimen with apparently
adult body chamber. Loc.
11 at 32m. McM J 1294. 4a-b.
Somewhat crushed phragmocone with part of body chamber
including aperture with lappet.
11 at 22m. McM J 939.

WA

WA

5a-c.

Docidoceras (Docidoceras ?) sp. aff. D. longalvum

(Vacek)
134
Phragmocone with incomplete body chamber; 5a-b, inner whorls
of fig. 5c (same specimen PI. 33, figs. a-b).

Bulletin 255

206

Explanation ok Plate
Natural

size if not

33

otherwise indicated.

Figure
a-b.

Page
Docidoceras (Docidoceras
(Vacek)

?) sp. aff.

D. iongalvum

Kialag\ik Formation, S. soivrrhyi Zone, lower Pscudocidoccras

zonule. Wide Ba\. Internal mold of pliragmocone and part
of body chamlier; b, enlarged (X 3) upper right sector of fig.
a, showing umbolateral groove and porous filling material.
(Inner whorls and \entral views of same specimen on PI. 32,
figs. 5a-c). I.oc.
10 at 8m. McM J 93 7.

WA

134

Plate

33

Bull. Amer. Paleont., Vol. 57

Plate

34

Alaskan ammonites,

Pt. II:

Westermann

207

Explanation of Plate 34
Natural size

if

not otherwise indicated.

Figure
l-3c.

Page
Docidoceras (Pseudocidoceras) widebayense

Westermann,

n.

137

sp.

Kialagvik Formation, S. soivrrbyi Zone, Pseudocidoceras zonule,

Wide Bay. la-d. Holotype, complete internal mold; Ic, body
chamber removed. Loc. USGS 21252. USNM 160239. 2. Septum
near end of phragmocone. Loc.
13 at 13-16m. McM J 899.
3a-c. Complete internal mold except for aperture; 3c-d, last
1 1/2 whorls removed. Loc.
13 at 13-16m. McM J 897.

WA
WA

Bulletin 255

208

Explanation of Plate
Natural

size

if

35

not otherwise indicated.

Page

Figure
l-2d.

Docidoceras (Pseudocidoceras) widebayense

Westermann, n. sp
137
Kialagvik Formation, S. soiucrhyi Zone, Pseudocidoceras zonule,
Wide Bay. la-b. Almost complete internal mold with aperture.
Loc.
10 at 11m. McM J 917. 2a-d. Complete internal
mold with aperture; 2c-d, body chamber removed. Loc.
13 at 9m. McM J 896.

WA

WA

Bill. Amer. Paleont., Vol. 57

Plate

35

Bull. Amer. Paleont., Vol. 57

Plate

36

Alaskan ammonites,

Pt. II:

Westermann

209

Explanation of Plate 36
Natural size

not otherwise indicated.

if

Figure
la-b.

Page

Docidoceras (Pseudocidoceras) widebayense

Westermann,

n.

sp.

Kiaiagvik Formation,
ceras Zonule,
aperture. Loc.
2, 3.

S.

137
soiverhyi

Zone,

Wide Bay. Complete

USGS

19862.

USNM

Docidoceras (Pseudocidoceras ?)
Westermann, n. sp. [6 ?]

(lower)

interna!
16025.

sp. aff. D.

Pseudocido-

mold except for

widebayense
147

Kiaiagvik Formation,
soiverbyi Zone, Pseudocidoceras zonule.
Wide Bay. 2a-b. Damaged small internal mold with some test
remains and aperture. Loc.
10 at 6m. McM J 933. 3.
Damaged small internal mold with aperture, possibly with
.S".

lateral lappet. Scree of loc.

4,5.

WA
WA McM
10.

J 932.

Docidoceras (Pseudocidoceras) n. sp. indet. A

151
Kiaiagvik Formation, 5 soivcrbyi Zone, Wide Bay. 4a-b. Damaged phragmocone and body chamber with aperture. Loc.

USGS 12405. USNM 160251. 5. Somewhat crushed specimen

with pathological body chamber fragment. Loc. USGS 19862.

USNM

160252.

Bulletin 255

210

Explanation of Plate 37
Natural

size

if

not otherwise indicated.

Page

Figure
1-5.

Docidoceras (Pseudocidoceras)

Westermann,

n.

cf.

D. widebayense

146

sp

Kialag\ik Formation, S. sotvcrbyi Zone, Pseudocidoceras zonule,

Wide Bay. la-b. Almost complete internal mold with test
13
at
remains on phragmocone, with aperture. Loc.
13-16m. McM J 928. 2a-b. Almost complete internal mold
13 at 13with one-half whorl of body chamber. Loc.
16m. McM J 927. 3a-b. complete, slightly damaged internal
mold. Loc. USGS 19862. USNM 160253. 4a-b. Phragmocone
with crushed body chamber and aperture. Loc. USGS 19801,
USNM 160255. 5. Phragmocone with crushed body chamber
and aperture. Loc. USGS 21252, USNM 160255.

WA
WA

Bull. Amer. Paleont., Vol. 57

Plate

37

Bull. Amer. Paleont., Vol. 57

Plate

38

Alaskan ammonites,

Pt.

II:

Explanation of Plate
Natural

size if not

VVestermann

211

38

otherwise indicated.

Figure

Page
Docidoceras (Pseudocidoceras) camachoi

Westermann, n. sp
147
Kialagvik Formation, S. soivrrhyi Zone, Wide Bay. Holotype,
complete internal mold with test remains. Probably loc USGS
19862.

USNM

160256.

Bulletin 255

212

Explanation of Plate 39
Natural

size

if

not otherwise indicated.

Page

Figure
a-b.

Docidoceras (Pseudocidoceras) camachoi

147
Westermann, n. sp
Kialagvik Formation, S. sowerhyi Zone, Wide Bay. Complete
internal mold with minor test remains and umbolateral groove
with 'porous material.' Loc. USGS 19801, USNM 160257.

Bull. Amer. Paleont., Vol. 57

Plate

39

Bull. Amer. Paleont., Vol. 57

Plate 40

Alaskan ammonites,

Pt. II:

Westermann

213

Explanation of Plate 40
Natural

size

if

not otherwise indicated.

Page

Figure
l-3b.

Docidoceras (Pseudocidoceras) camachoi

147
Westermann, n. sp
Kialagvik Formation, S. soivcrbyi Zone, Wide Bay. la-b. Almost
complete internal mold with pronounced umbolateral groove
(on both sides), partly filled with 'porous material' and
bridged by the shell. Loc. CAS 29011. CAS 13111. 2a-b. Internal mold of average phragmocone, Loc. USGS 12405,
USNM 160258. 3a-b. Nucleus of phragmocone from the
Pseudocidoceras zonule. Loc.
10 at 6-7. 7m. McM J 921.

WA

Bulletin 255

214

Explanation of Plate
Natural size

if

41

not otherwise indicated.

Page

Figure
1-2.

Docidoceras (Pseudocidoceras) camachoi

147
Westermann, n. sp
Kialagvik Formation, S. soivrrhyi Zone, Pseudocidoceras zonule,
Wide Bay. la-b. Complete specimen except for aperture,
phragmocone with test; lb, cross section of phragmocone.
Loc. USGS 21251. IISNM 160259. 2. Relatively small, incomplete internal mold with test remains and weak umbolateral
groove. Loc.

3a-b.

WA

10 at 1.8m.

Docidoceras (Pseudocidoceras)

Westermann,

n.

McM

cf.

923.

D. camachoi

sp

Kia!ag\ik Formation, S. soivrrhyi Zone, Wide Bay. Nucleus of

phragmocone, internal mold. Loc. USGS 12405, USNM 160260.

148

Bull. Amer. Paleont., Vol. 57

Plate

41

Plate

Bull. Aimer. Paleont., Vol. 57

/
/'

\..
i *4 V

42

Alaskan ammonites,

Pt, II:

Westermann

215

Explanation of Plate 42
Natural

size if not

otherwise indicated.

Figure
la-b.

Page
Docidoceras (Pseudocidoceras) camachoi

Westermann, n. sp
147
Kialagvik Formation, S. soiacrhyi Zone, Wide Bay. Alnnost complete internal mold with aperture and minor test remains;
lb.
2.

penultimate phragmocone whorl. Loc.

Docidoceras (Pseudocidoceras

Westermann,

n.

?) cf. D.

CAS

29011.

CAS

13121.

camachoi

sp

Kialagvik Formation, S. soiuerbyi Zone, Wide Bay. Complete internal body chamber mold, with damaged phragmocone. Loc.

CAS

29011, Cx'\S 13120.

151

Bulletin 255

216

Explanation of Plate 43
Natural

size

if

not otherwise indicated.

Page

Figure

lab.

Docidoceras

(?) [?

n. sp. indet.

Pesudocidoceras or

n. subg.],

151

Kialagvik Formation, 5. soivcrhyi Zone, base of E. amplectcns

zonule, Wide Bay. Phragmocone with crushed beginning of
body chamber, largely with test. Loc.
3.
McM J 930.

WA

2a-b.

Docidoceras

(Pseudocidoceras

Westermann,

n.

?)

paucinodosum

sp

Kialagvik Formation, S. soiverhyi Zone, E. amplcctctis zonule,

Wide Bay. Holotype, complete internal mold with aperture;
2b X 1.1 (see also PI. 44, fig. 1). Loc. CAS 29014. CAS 13123.

153

Bull. Amer. Paleont., Vol. 57

Plate

P^^^^.,

lb

43

Bull. Amer. Paleont., Vol. 57

Plate

44

Alaskan ammonites,

Pt. II:

Westermann

217

Explanation of Plate 44
Natural size

if

not otherwise indicated.

Page

Figure
l-3b.

Docidoceras? (Pseudocidoceras

Westermann,

n.

?)

paucinodosum
153

sp

Kialagvik Formation,
sotvcrby't Zone, E. amplcctcus zonule,
Wide Bay 1. Holotype (see Pi. 43, figs. 2a-b). 2. Complete
but partly crushed internal mold, with shallow umbolateral
groove filled by test. Loc.
5, McM J 934. 3a-b. Almost
complete, slightly deformed internal mold. Loc. USGS 19828.
.S'.

WA

USNM
4a-c.

160261.

Docidoceras? (Trilobiticeras

?) n. sp. indet.

Kialagvik Formation, S. soivrrbyi Zone, E. amplcctens zonule,

Wide Bay. Complete but partly deformed internal mold with
test remains and large lateral lappet. Loc. Shell L 546. Collections of Shell Oil

Co.

155

Bulletin 255

218

Explanation of Plate 45
Natural

size if not

otherwise indicated.

Figure

Page

1,2.

Pseudotoites cf. P. argentinus Arkell

158
Kialag\ik Formation, S. soivrrhyi Zone, Wide Bay. la-b. Almost complete internal mold with damaged body chamber
and minor test remains. Loc. USGS 19922. USNM 160262.
phragmocone. Loc. USGS 12405, USNM
2. Partly crushed
160263.

Bull. Amer. Paleont., Vol. 57

Plate 45

Bull. Amer. Paleont., Vol. 57

Plate

46

"^'^ili

*''

2a

2b

Alaskan ammonites,

Pt. II:

Westermann

219

Explanation of Plate 46
Natural size

if

not otherwise indicated.

Figure
l-2b.

Page
Pseudotoites cf. P. argentinos Arkell
158
Kialagvik Formation, S. soivrrbyi Zone, Pseudocidoccras zonule. Wide Bay. la-b. Phragmocone with crushed
fragment of
body chamber; lb, polished section. Loc.
13w at c.l8m.

McM

WA

935.

2a, b.

Phragmocone

of body chamber. Loc.

WA

with

10 at 11m.

damaged beginning

McM

938.

Bulletin 255

220

Explanation of Plate 47
Natural

size

if

not otherwise indicated.

Page

Figure
a-c.

162
Pseudotoites cf. P. transatlanticus (Tornquist)
Kialag\ik Formation, S. soiccrbyi Zone, io\ver Pscudocidoccras
zonule. Wide Bay. Complete internal mold, other side damaged; a X 0.8, b-c X 1-1; showing umboiateral groove and
section. Loc.

USGS

19862,

USNM

160264.

Bull. Amer. Paleont., Vol. 57

Plate

47

INDEX
Note: Light face type refers to pages. Bold face type refers to plates.

Aalenian, upper
Abbasites
acanthodes, Sonninia
achillei,

Banff, Alberta

17,27
64,129,137
92

Guhsania
cf. benneri, Planammatoceras
22
beyrichi, Pseudobella,

Eudmeto91

ceras
actinophora,
Witchellia

lioceras

112

biforme, Docidoceras

28,92,98

adicra, Sonninia

Sonninia

107, 112

Sonninia

23-26

boweri, Sonninia

93
11,22

boyeri,

27
27

Alberta

6
133

23
24, 103

21,26,99,
102
24

British

21,23,26,
124
106

Param-

matoceras
Bradford Abbas,
Somerset
Bradfordia
Bredyia

alaskensis,

Alaskoceras
Alaskoceras

24,28

bifurcata,

adicroides,

adnata, Witchellia ....

aguilonia, Witchellia
Alaska, S. E

94
24

71

58,66,73,92
22,46,48
64,65

Columbia

24, 131

Bulgaria

64

alleoni,

Parammatoceras

....

Alps
Sonninia
amaltheiforme,
alsatica,

Eudmetoceras

71
103,136
114

Cadomites
California,

Calliphylloceras
Callovian

66, 68, 70,

camachoi,
Docidoceras

81,88
amplectens,

Eudmetoceras

Ammonites

40
10, 11, 16

Argentina

Chile
clypeus,

Cook

28
45,46

Arkelloceras
Asaploceras
Asthenoceras
auerbachense,

Parammatoceras

....

28,131
60
37
60
21,26,126
71

aurifer,
115, 128

131

celans, Sonninites

6,7,73,130,
157
114
27

Anolkoleiites

Fontannesia

21,139-142

S. Vigilio,

carlottensis.
10, 11, 15

Andes

Maceratites
Australia
austroamericana,

38-42

Alps
Caribou Creek section

147

section

argentina,
Witchellia
argentinus,
Pseudotoites
Arieticeras

Cap

19-21 20, 22, 24, 51,

56, 69, 79, 82

Amblyoxyites
Anderson Creek

129
26
41

144
27
46

Hebetoxyites
Alaska
23, 39, 41, 79,
99

Inlet,

...

compactile,
Pseudolioceras
constricta, Emileia
Colpitts Group

....

55
23
93

corona,

Zemistephanus

130

corroyi,

Deltotoceras
costisparsum,
Holcophylloceras

70
..

40

costistriatum,

Pseudoliceras ...8,9

28,130

21

crassicostatus,

27

221

Parabigotites

11,22

Index
crassiformis. Sonninia

eudmetum.
Eudmetoceras

aff.

102
137

polyacantha
crater, Emileia
Csernyeiceras

64,65,72

amus

sp. C.

fallaciosum,

Inocer-

Zone

lum

42

21,51,81

Hammato-

ceras

Docidoceras
Docidoceras camachoi
assemblage
Dorset
Dorsetensia
dubari, Parammatoceras

Dundryites

16,

70,79
28, 93

22
88,103,123
114
71
114

dumortieri, Pseudam-

matoceras
Dundry, Somerset

72
108, 115

Emileia
Emileites
Erycites
Erycitoides howelli

Zone
Erycitoides teres
zonule
Esericeras

Argentina
espinazitum,
Lytoceras

Euaptetoceras
euaptetum,

Eudmetoceras
eudesiamum,
Lytoceras

Euhoploceras

Eudmetoceras
Eudmetoceras ampletens zonule
Eudmetoceras gerthi

assemblage

52
114

13

24,94,124
24

Fissilobiceras
flexicostatum,

Tmetoceras
Fontannesia
Frogden Quarry,
Oborne, Dorset

6,18,27
94
112

G
gardanum,
40

Partschiceras
Gelasinites
gelasinus,
Gelasinites

113
113

gerthi,

23,24,129
28,132,137
64, 129

27,66,73
66
112
112,115

Eudmetoceras
Hammatoceras

38, 39

gingensis, Sonninia
giauca, Witchellia
gracililobata, Oppelia

Sonninia/
Schloenbachia

47, 48, 50

gracilis,

5,10, 11,137

18
108

espinazitensis,

Sonninia
Espinazito Pass.

52

fastigatum
maclintocki
felix, Sonninites
Fernie Group,
Alberta

ellipticum,

Partschiceras

59
112

..

lioceras

24,25,99

Eudmeto14,16,17,18

Witchellia
fastigatum, Pseudo-

falcata,

ceras klimakomphadiscus,

Grammoceras

12

..

deslongchampsii,
Holcophylloceras
Devils Point,
Alberta
discoidale,

28. 157

D
Dactylioceras

29,66
18,29,137

15

Europe, N.
evolutus. Pseudotoites

24,28

25, 94, 99, 107

Grammoceras
grandis, Pseudam-

59,60
71

matoceras
grossicostatum,
Partschiceras

39
24

Guhsania
161

guliense,

Pseudam72

matoceras
43
64,67

70,71

42
21.24, 121

26,67,72.93

H
Hammatoceras
Hammatoceras Zone
Hannover, Germany
hansoni, Sonninia
Haplopleuroceras
hauthali, Harpoceras
Hazelton Group

64
10
91
24
103
28

24,124

12

26

Hebetoxyites
Hebetoxyites

222

10

20,28
20,28

Index
Inoceramus
Ludwigia concava
Zone
Ludwigia discites
Subzone

40
40
73

howelli, Erycitoides

Hudson Bay Mountain,

British

Columbia

Hungary
Hyalinites

Hyperlioceras

24,25
37,64
114
58

infernense,

67,69
60, 111

31

Macrophylloceras
malarguense,
Harpoceras

10

Marocco

sp. C.

Iniskin Peninsula,

Eudmetoceras
intumescens,
Sonninia
involutum, Harpoceras
amaltheiforme mut.

28

Minnewanka Lake,

79

Alberta
moerickei, Eudmetoceras

42
24,94, 124

27,47,70,80
28

Moluccas
27, 66,

mouterdi, Pseudammatoceras
Moose Creek - Mt.

73

71

Kathleen section

Skirroceras

10,16
26,93

Mormon Formation

8
18
22

Tmetoceras

kochi, Eudmetoceras
kunthi, Phylloceras

71,73
28,80

Holophylloceras

Kialagvik Formation

klimakomphalum,
Eudmetoceras

28
64

Eudmetoceras
Mendoza, Argentina
mediterraneum,

kirshneri,

Mount

Hill,

West
131

Australia
Mount Jura,
California
Mt. Mamie section

21,27,28,64,
64, 66, 68, 74
27
36

26,93,99
10

N
Nannoceras

Nannoceras

New

Pseudo161
133
133

liebi,

lorteti,

neumayri, Lytoceras

79

Strigoceras

Docidoceras
limatum, Coeloceras
aff. longalvum,
Docidoceras
32, 33

Guinea
Newmarracarra
Limestone
aff.

22, 29,

ceras

29,

58

24, 107

nucleospinosum,

Eudmetoceras

133

115
43
28, 133

28,94,157

nodata, Sonninia
17

80

Hammato-

Los Molles,
Argentina

14

nannomorphum,

24, 108

languidum,
leicharti,
toites

115

nannodes,
Asthenoceras

aff.

laeviuscula,

Witchellia

73

39,40

82

insignoides,

kirki,

6,52
..

masticonnensis,

24,86,99

jaworskii Eudmetoceras [eudmetum]

115,128

Maceratites
maclintocki,
Pseudolioceras
Maconnais, France

103

Alaska

26,58,60
42

Zurcheria

Inferior Oolite

20,22,27,29,
92, 103

Lytoceras

Eudmetoceras

24,93

Ludwigia murchisonae
Zone

inconstans,

Inoceramus
Inoceramus
Subzone

11,22

lucifcr,

helerophyllum,
Phylloceras
Holcophylloceras

65
178

Oborne, Dorset
obtectum, Parammatoceras

223

110,112

70,71

Index
Okhotsk Sea,
6
47

Siberia

Oppelia

Prinz Patric Island,

Canadian Arctic
propinquans, Sonninia

Pseudammatoceras

opeliiformis, Bradfordia? .10,11,12

37
65
64, 65, 71, 72,

90
21,26,136

20,21,48

Praeoppelia

20,21,48
26, 46, 59

10,11,12
Oregon, east-central
ornatum, Pseudammatoceras

72

Pseudocideras
Pseudocidoceras
zonule
Pseudolioceras
Pseudotoites

11,12,20
22,26,52
27,28,94,
129, 130, 157

Otoites sauzei

Zone

22

10,

11,23,41,

Pseudotoites singularis

79, 84, 92,

assemblage
Puale Bay
pugnax, Spinammatoceras
punctum, Docidoceras

107, 113

P
Pachammatoceras

65

Parabigotites [crassicostatus] zonule

Parammatoceras
65
Partschiceras
parvispinata,

Zurcheria

11,22,84
69
40
103

patefactor,

113
92

Witchellia

Sonninia
paucinodosum,
Docidoceras
Pelekodites

patella,

20
21

pelekus,
Pelekodites
32
perfectum, Docidoceras
Perth, W. Australia
pertinax, Zurcheria
Phylloceras
pinguis, Sonninia
planinsigne,
cf.

..

Planammatoceras
Planammatoceras

115,126
133
99
103
36
106

65,69,71
65,69,90

planulatum,
Docidoceras
planiforme,

133

Planammatoceras

65,71

platymorpha,
Witchellia
platystomus,
Abbasites
playfordi, Sonninia ....
Pleydellia
Pleydellia puchensis

assemblage
Polyplectites
polyshides, Emileia
Praestrigites

Praeoppelia
Preston Creek

110,123
137

94,99
61

28,61,70
..

129
24

26,39
21,48
11

28, 130

64
134

Index
singularis, Pseudotoites

27,133,157,
160
24, 124
28,92

skawahi, Sonninia
Sonninia
Sonninia sowerbyi

18

Zone

Talkeetna Mountains,
Alaska
23,24,36,92,
99, 107

tecta, Dorsetensia

Planammatoceras

Sonninia trigonalis

Subzone
Sonninites

South America
spatians, Spatulites
Spatulites

..

sphaeroceroides,
Pseudotoites
spinifera, Sonninia

....

22,30,93,
113, 115, 123
114
27,130,152,
154
115
115,127

28,130,152,
154, 159
93, 110

Stemmatoceras
11

triptolemus
stephensi, Vacekia

60

Stephanoceras humphriesianum Zone

Stephanoceras

24, 41,

18

thorsteinssoni,

Phylloceras
Tithonian
tobleri, Haplopleuroceras

37
42
103

transatlanticus,

Pseudotoites

28, 131, 154,

159, 162
129, 155

Trilobiticeras
trilobitoides,

Trilobiticeras
triptolemus,

134

Stemmatoceras
Tmetoceras
Tmetoceras scissum

79

24
93
58
58

Stiphromorphites
Stoford, Somerset
Stauffenia
Strenoceras sub-

furcatum Zone

41

striatulum, Grammoceras
Strigoceras
suballeoni, Param-

matoceras
subconcavum,
Pseudolioceras

28,59
79,93

11,22
26,61,94

zonule

18

icostatum zonule

12

Tmetoceras tenue
zonule

Tuxedni Bay
Tuxedni Formation
Tuxedni Group
Turkey

12
lo

23,36,39,99
23, 36, 39, 99
64

tyrrhenicum,

Eudmetoceras

69,79
79

Oppelia
71

58

ultramontanum,

subfrancisci,

Holcophylloceras

43

Lytoceras

41

Ammo64,72
64,72

nites

Hammatoceras
Pseudammatoceras

64, 72

subplicatella,

Oppelia

47,48,50
79
42
26,59,93,99

subradiata, Oppelia
Sully,

65

teres, Erycitoides

Tmetoceras tenue-flex-

(Skirroceras)

subinsigne,

114
18

tenue, Tmetoceras
tenuinsigne,

Normandy

Supplee, Oregon
suterni, Witchellia
sutneroides,
Witchellia

Swabian Jura

...

110, 123

V
vaceke, Lytoceras

43
60

Vacekia
verpillierense,

Hammatoceras

...

64,68

W
Warm

Springs

Formation
21,116

Weberg Formation

102

Western Australia

225

59
93
28, 93, 94, 157
26,

26, 59,

Index

whiteavsi,
Pseiidolioceras

widebavense,
Docidoceras
Wilkie Point

Formation
Witchellia
Witchellia sutneroides

assemblage

Z
6,53

Zemistephanus
Phylloceras
Zetoceras
zignodianum,
Holcophylloceras
Zugophorites

zetes,

20,21,137
37
108

^^

zucnena

226

lou
36
3b

42
22, 25 26 93
^'^'
27,
<ii, lol
-n^.^

^3 -B
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BULLETINS

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AMERICAN
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NEW MIDDLE

JURASSIC AMMONITINA

NEW GUINEA

By
Westermann

G. E. G.

AND
T. A. Getty

I
1970

Paleontological Research Institution

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1 1970

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BULLETINS
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AMERICAN
PALEONTOLOGY
(Founded 1895)

57

Vol.

No. 256

NEW MIDDLE

JURASSIC AMMONITINA

FROM

NEW GUINEA

By
G. E. G.

Westermann and T.

A.

Getty

Department of Geology, McMaster University, Hamilton, Canada,

and Department of Geology, University College, London

February

11,

1970

Paleontological Research Institution

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CONTENTS
Page
Abstract

231

Introduction

231

Acknowledgments

233

Stratigraphy of the

Kemaboe Valley area

Paleontology

235

237

Previous literature on Bajocian-Callovian ammonites

237

Systematic descriptions

238

Faunal

affinities

and ages

288

References

298

Plates

303

NEW MIDDLE JURASSIC AMMONITINA FROM

NEW GUINEA
G. E. G. Wf.stermann and T. A. Getty

ABSTRACT
The Lower Bajocian to Middle Callovian Ammonitina of an unlocalized
collection from the Kemaboe Valley of West Irian (formerly West or Dutch

New

Guinea) is described; several uppermost Jurassic to basal Cretaceous

ammonite species of the same collection are recorded only. Most of the species
and genera are either new to the Indo-Malayan Archipelago or have previously
been wrongly classified. Docidoceras s.s., Stephauoccras s.s., Bullatimorphitcs
(Treptoccras), and Cobbanitcs
?) are new to the entire Southeast Asia-Aus(

tralia

area.

The Lower Bajocian (Sonninia soivrrbyi Zone) is clearly indicated by the

Mediterranean Docidoceras (Docidoceras) longalvum (Vacek) showing close
affinity to the Anatolian subspecies D. limatum (Pompeckj), and by Fontannesia sp. with affinity to the West Australian F. clarkei (Crick) and possibly being identical with 'Grammoceras' kUiani (Kruizinga) from the Sula Islands
in the Moluccas. Middle Bajocian is represented by the almost cosmopolitan
Stephanoceras ex gr. S. humphriesianum (Sowerby) and by the probably endemic species S. (Stcmmatoccras}) etheridgei (Gerth), possibly present in both
dimorphs. The presence of Bullatimorphitcs (Treptoccras) aff. B. nhligi
(Popovici-Hatzeg) and {})Cobbanites aff. C. englcri (Frebold) suggests Upper
Bathonian age or Lower Callovian age, and affinities with Europe-Western
Asia and the northern Cordilleras respectively. Early Middle Callovian is
clearly indicated by the 'Indie' Subkossmatia and Eucycloceras (?). Curiously
enough, the usually ubiquitous Macroccphalites sd. assemblage of the Lower
Callovian is missing.
The unnamed new subgenus of Bullatimorphitcs ?, including B. costidcnsus,
n. sp., and the associated Irianitcs, n. gen., based on the much discussed
'Coeloceras' mocrmanni Kruizinga from the Sula Islands, are probably Callovian and possibly Middle Callovian in age; it is suggested that Bullatimorphitcs ? (n. subgen.) is phylogenetically intermediate between true Bullatimorphitcs (Middle Bathonian to Lower Callovian) and the eucycloceratids, particularly Subkossmatia (Middle Callovian). The family Macrocephalitidae as
defined in the 'Treatise,' by the same token, would then be of polyphyletic
origin, including the macrocephalitids proper which evolved from sphaeroceratids, as well as the eucycloceratids. The Eucycloceratidae Spath are, therefore, provisionally again separated, at least at the subfamily level.
The previous finds of Bajocian to Callovian Ammonitina in New Guinea
and the Indonesian archipelago are re-examined.

INTRODUCTION
Like

all

previous ammonite collections from

New

Guinea, the

under study were obtained ex situ from stream bed and

river bed pebbles of more or less worn and broken concretions.
Consequently, no stratigraphic evidence is available; however, one
important faunal association is known: a single fragment of a concretion contained Irianites cf. /. moermanni (Kruizinga)
and
Bullatimorphites ? (Treptoceras ?) costidensus, n. sp. Unfortunately, the other fossils before reaching us had already been removed
from the concretions and segregated without indicating associafossils

232

Bl LLETIN 256

New Guinea Ammonites: Westermann & Getty

tions. Nevertlieless,

new
known

southeast Asian

be published.

The ammonites

this collection ot largely

scure forms from the important but

archipelago

is

judged worthwhile

to

233

little

or hitherto ob-

described herein represent the majority of a collection

made

the course of an ethnological expedition by Dr. C.C.F.M.

Le Roux

during 1939-1940

to the region of the

Kemaboe

in

Valley in the Cen-

Ranges of central West Irian (formerly West New Guinea or

Netherlands New Guinea) (Text-figs. 1,2). Probably all of the
fossils were obtained along the bed of the Iwaboe River, a left
tributary of the Kemaboe entering about four kilometers below
tral

Zanepa.

The complete collection, permanently stored at the Rijksmuseum van Geologie en Mineralogie (R.G.M.) in Leiden, The
Netherlands, comprises approximately 270 ammonites, many fragmentary, of which more than 200 were sent to us and the remainder
briefly

studied in

the

Leiden

collections.

All

are

preserved in

black calcareous mudstone containing variable amounts of free

calcite

and disseminated

pyrite,

matrix of the same material. This

together with some

mode

silica,

in a

was noted
by Boehm (1913) in ammonites from Windesi, northern Lenggeroe
area

(Text-fig.

1,

loc.

3)

of preservation

but with the difference that the

silica

was a more prominent constituent of the concretions. The state

of preservation of the material is good although in some cases
the inner whorls have been crushed on one side or, more rarely,
destroyed. Some whorls have been partly crushed and a few have
been compressed dorso-ventrally. Exposed whorls have also suffered
from transportation.
In August 1968, G. Westermann studied the main Jurassic
ammonite collections from Indonesia (including West Irian) during a brief

visit to the

following institutions in

The

Netherlands:

Rijksmuseum van Geologie en Mineralogie, Leiden; Mineralogical

and Geological Museum, University, Delft; Geological Institute,
University, Amsterdam; and the Royal Shell Exploration Production Laboratory in Rijswijk near

Den Haag.

ACKNOWLEDGMENTS
For the loan of the collection and of several holotypes we
thank Miss G. E. de Groot and Mr. W. U. Boon van Strein of the
Rijksmuseum van Geologie en Mineralogie in Leiden. For their

Bulletin 256

234

New Guinea

Limestone

slates &
phyllites

sandstones &
sandy shales
Intrusions
Text-fig.

2.

to

Kembelangon
Formation
Faults

Dip

Geotectonic

Herbes, 1962).
stones

Group

map of the Kemaboe Valley area (after Visser and

limestones belong to the New Guinea Limestone; the sandshales and slates and phyllites belong to the Kembelangan

The

sandy

Formation.

help during the

visit of

G. Westermann,

we

also

thank Mr. Schuif

Museum

van Mineralogie en Geologie in Delft, Professors

and
H. J. MacGillavry of the Geologisch Instituut,
Hermes
J. J.
University of Amsterdam, and Dr. R. Lagaaij from the Bataafsche
of the

Internationale Petroleum Mact/scappij N. V. in Rijswijk. Dr.

van den Boogaard of the Geological

Institute

Amsterdam

M.
fur-

nished photographs of type specimens. Comparative fossil material

and plastotypes were made available by Dr. J. Sornay, Museum

New Guinea

Ammonites: Westermann & Getty

235

National d'Histoire Naturclle, Paris, Drs. D. McLaren and H. Frebold of the Geological Survey of Canada, Ottawa, and Dr. R.

Jordan of tlie Niedersachsisclies Landesamt fiir Bodenforschung,

Hannover, Germany. Important discussions were contributed by
Dr. J. Callomon from the University of London and Drs. R. Enay
and C. A. Mangold from the University of Lyon. This project was
supported by a grant of the National Research Council of Canada.

STRATIGRAPHY OF THE KEMABOE VALLEY AREA

Although forming one of the major

east-west valleys in cen-

New

Guinea, the Kemaboe Valley (Text-fig. 2) has remained

poorly known, largely because of the surroimding rugged terrain
tral

and dense
esia

forests; it

is

not shown on the geological

map

of Indon-

prepared by the Direktorat Geologi Indonesia and published

by the United States Geological Survey in 1965 but on the maps

published by the Netherland Nieuw Guinea Petroloeum Maat(referred to hereafter as the N.N.G.P.M.)
(Visser and
Hermes, 1962) The lower course of the Kemaboe is unknown, but
it is presumed to enter one of the tributaries of the Waipoga,
which empties into the eastern side of Geelvink Bay.
According to Visser and Hermes (1962) the thick, coarse and
fine elastics with minor carbonate are placed in the Kembelangan
Formation, comprising Jurassic and Cretaceous, which is most
fully developed in the Lenggeroe area southwest of Geelvink Bay

schappij

(Text-fig.

with

'B',

1)

Here

it

comprises the

'A', 'B',

'C, and 'D'

'C, and 'D' forming a sec[uence in an

upward

members
succession

which 'A' is, at least partly, the lateral equivalent. The 'A' and
'C members are predominantly argillaceous, while the 'B' and 'D'
members are predominantly arenaceous. The 'B', 'C and 'D'
members are restricted to the southwestern Lenggeroe area west of
of

the Jakarti Fault Zone, while the 'A'

member

occurs only east of

it.

shales and mudabout 1200

stones, with intercalated limestone bands and massive limestones
at the base, but becomes progressively metamorphosed eastward

The

'A'

member

passing into schists

type locality

consists of

and

phyllites. Fossils are

(Nanggoebi Valley)

not

common

at the

but along the strike, both to

and south, concretions derived from this member have

yielded numerous ammonites indicating Bajocian, Bathonian ?,
Callovian, and Tithonian (Boehm, 1913; Visser and Hermes,
the north

Bulletin 256

236

1962) North of the Lenggeroe area the 'A' member is overlain with
apparent disconformity or, more probably, with fault contact, by the
.

pelagic limestones of the

Upper Cretaceous Imskim Formation,

ap-

parently suppressing the upper part of the 'A' member. This was
supported by the discovery of Lower Cretaceous ammonites in the
'A' member in the Central Ranges by Gerth (1965)
The 'B', 'C,
and 'D' members were originally described from a well at Etna
Bay where 'B' consists of approximately 600 m sandstone, becoming argillaceous above and overlying the nonmarine Tipoema Formation. The lower part is of Upper Bajocian to Lower Oxfordian
age [based on Grammatodon virgatus (J. de C. Sowerby) ] and has
been correlated with the 'A' member while the upper argillaceous
sandstones, the 'B' member proper, contains Lower Cretaceous
.

Foraminifera.

The

superposed 'C

grey shales and

member

silty shales,

consists of

approximately 700

with thin sandstones.

The

foramini-

fauna indicates Lower Cretaceous, below, and Upper Cre-

feral

taceous, above.

The
150
is

member is another sandstone unit, about

Upper Cretaceous Foraminifera, which
by Cenozoic limestones, but may locally reach upward

overlying 'D'

thick, also

overlain

containing

into the Cenozoic.

The

'A'

member

is,

therefore, the eastern equivalent of at least

most of the 'B' member.

Outside the Lenggeroe area, the Kembelangan Formation has
only been divided into the argillaceous-arenaceous undifferentiated
'BCD' members bearing Jinassic ammonites, and the argillaceous
'A'

member,

also bearing Jurassic fossils.

southward location of the arenaceous

source of the

Both
the

1000

consistently

more

suggests a southern

elastics.

facies of the

Kemaboe

The

facies

Kembelangan Formation

Valley area.

The

'A'

member

intensely folded soft black slates

are developed in

consists of

and

more than

phyllitic slates, with

and orthounknown. The beds show a progressive increase in metamorphism to tlie north and northwest, while to the
south they are overlain, apparently partly with fault contact and
partly with disconformity, by the sandstones and sandy shales of
the undifferentiated 'BCD' members. Southwest of the Kemaboe
intercalations of black marly limestones, silty sandstones

quartzites; the base

is

TABLE

STAGE

We

.
,

New Guinea

Ammonites: Westermann & Getty

(near Wissel Lakes)

Valley

this

237

arenaceous unit contains Upper

Cretaceous Foraminifera. As in the Lenggeroe area, incompetent

upper

'A'

beds appear to have been tectonically suppressed, both

by a fault zone (here the North Paniai Fault

Zone) and the more argillaceous beds to the northeast are tliicker
tiian the arenaceous beds to the southwest. Tiiere would seem to
facies are separated
,

be a possibility that the depositional environment of the Kembel-

angan Formation was

tectonically controlled during Jurassic

and

Cretaceous times.

PALEONTOLOGY
PREVIOUS LITERATURE ON BAJOCIAN-CALLOVIAN AMMONITES
ammonites have been recorded and figured from a
and central New Guinea (Textfig. 1)
The literature was reviewed particularly by Arkell (1956)
and Visser and Hermes (1962) and a revised account of most
specimens figured up to date from the Bajocian to Callovian is
given here in table form (Table 1) The important early descriptions by Etheridge (1890) and especially by Boehm (1913) were
taxonomically revised by Spath (1928) in the course of his work
on the ammonites of Kutch, India, and by Westermann (1956
a,b) The Bajocian to Callovian ammonites figured give good evidence for only the earlier Callovian and Middle Bajocian (s.s.),
because Boehm's figured fragment of a 'Stephanoceras daubenyi"
(pi. 3, fig. 1) cannot be clearly identified with Cadomites sp. However, the subgenus Chondroceras? (Praetulites) Westermann which
was based on "Sphaeroceras godohense Boehm" Kruizinga, non
Boehm, from the Sula Islands and New Guinea, has recently been
described from the Upper Bajocian P. parkinsoni Zone of the
Jurassic

number

of localities in western

Venetian Alps (Sturani, 1964 a,b) The holotype of Stephanoceras

(Stemmatoceras}) etheridgei Gerth (1927) is here refigured (Textfig. 8)
The fragment figured under the same name {op. cit., fig. 2)
however, is a microconchiate Stephanoceras s.l., probably of the
subgenus Itinsaites. Gerth's (p. 226) "Sphaeroceras cf. biillatum
.

d'Orb.", which

is

here figured (Text-fig. 10)

(Treptoceras) uhligi
,

pean Bullatimorphites

The

is

close to the Euro-

(Popovici-Hatzeg)

small collection of ammonites recorded but not figured by

Martin (1911; see

also Visser

and Hermes,

studied in the Rijksmuseum of Leiden.

1962, p. 54)

The

was

re-

"Quenstedtocerasf"

Bulletin 256

238

(p.

97)

from the Digoel River

cephalitid

(st.

12117)

is

probably a macro-

and the "Macrocephalites?" from B-River

Bullatirnorphites} macroconch.

(p.

95)

from
the Sepik River (loc. 7) contained the early Callovian "Macrocephalites keeuxi'cusis y" Boehm besides the more abundant Upper
Jurassic ammonites. Arkell and Donovan, respectively identified
(/// Visser and Hermes, 1962)
a smaller collection from Roemberpon Island (Text-fig. 1, loc. 2) and a larger collection from claystone concretions in riverbeds between Geelvink and Etna Bays
in the southeastern Lenggeroe Area (loc. \)
added to the previously described forms were the circumpacific genus Pseudotoit.es and
the poorly known 'Normannites' moennanui (Kruizinga) which
was originally placed in Coeloceras and is here redescribed and
placed in the new genus Irianites. The figured specimens were restudied at the Shell Company in Rijsvijk and partly given on
loan. The taxonomic revision concerns particularly the fragments
(Text-fig. 3) of supposed Upper Bajocian "Baculatoceras" sp.
[^^Garantiaiial (Visser and Hermes, end. 17, figs. 23 a,b) which
is here identified with the Tithonian to Berriasian Blanfordiceras
novaguiy-iense Gerth (1965), a close ally (Psubsp.) of the Himalayan B. xvallichi (Gray)
and the merely recorded Bajocian
"}I.Mbyrintlioceras sp." (op. cit., p. 55) which is a typical Callovian
Subkossmatia. The most recent description of ammonites from the
Schliiter's

collection

(1929)

by Gerth (1965) includes besides Upper

Jurassic to Berriasian forms only the record of "Macrocephalites
keeuwensis" Boehm.
Central Ranges

(loc.

6)

SYSTEMATIC DESCRIPTIONS
HILDOCERATACEAE Hyatt,

Superfamily

1867

Family (?)HILDOCERATIDAE Hyatt, 1867

Subfamily

(?)GRAMMOCERATINAE Buckman,
FONTANNESIA Buckman, 1902

1905

Genus
Fontannesia

?1926.

clarkei (Crick), 1894 [?F. clarkei ssp. kiliani

Pis. 48,49, Text-figs. 4-5
(Kruizinga), 1926]

aff. F.

Grammorrras Kiliani
54,

p.

33, pi.

1,

n.

sp.,

Kruizinga (Sula

Is.),

Jb.

Mijnwezen,

vol.

fig. 2.

Material. Four large and two small, almost complete specimens with body cliambers; one good phragmocone; one large
phragmocone fragment, and one small body chamber with crushed
phragmocone from Kemaboe Valley.

New Guinea

Ammonites: Westermann & Getty

Occurrence.

West

Irian,

Misol and

Description.

The

inner

whorls

(?)

Sula Islands.

subquadrate

are

239

some-

to

rounded flanks and a flattened

There are dense, often somewhat

Avhat subrectangiilar, with gently

venter bearing a blunt keel.

fasciculate

and prosoradiate

ribs,

sometimes irregular in strength

but without tubercles.

The

intermediate whorls become compressed rounded sub-

rectangular, usually developing a narrow, moderately steep

bilical slope,

poorly separated from the flattened flanks, and

umdis-

on which the strongly projected rib endings may

form ventrolateral 'carinae'. The venter is more or less broadly
tabulate with strong blunt keel and, at least on the internal mold,
frequently bisulcate. The ribs remain usually dense, rarely fasciculate, and straight to prosoradiate, with projection on the shoulders, leaving narrow smooth bands on umbilical slope and venter.
The outer one or two whorls are usually subrectangular with
rounded umbilical slope, flat flanks and tabulate-unicarinate venter
which at the end of the body chamber may become weakly bisulcate
or slightly fastigate. The keel is prominent but blunt and 'solid'
(unfloored) throughout. The umbilicus is shallow and only moderately wide (Ur=32-38%)
The costae may remain dense and
more or less straight (PI. 49, figs. 4 a,b) but usually become coarse
and widely spaced, often slightly falcoid fasciculate or more rarely
bifurcating and strongly projected on the outer flank and shoultinct shoulders

der; dying out

on the lowermost

flank, the ribs reach their greatest

strength on the uppermost flanks

may form

a discontinuous

and on the shoulders where they

'carina.'

'simple' with strongly projected

pressed shells
the

septal suture

E/L and L/U

smaller raised

as sug-

(PI. 48. figs. 3 a,b)

(Westermann, 1966), the more involute and comtend to be more densely and weakly costate than

more evolute and

The

aperture was probably

rostrum or ventral lappet

gested by a medium-sized specimen

As usual

The

less

compressed ones.

(Text-fig. 4 a-c)

is

simple with subequal

bipartite saddles, moderately slender trifid L,

U2 followed

at

much

maturity by two almost straight

much

smaller umbilical elements (U3, U4 part.) along a slightly

sinking saddle line but ending at approximately the same radius.

The

internal part consists of a prominent

I/U saddle and

smaller saddle separated by a small oblique weakly bifid

much

Ux

(?)

Bulletin 256

240

Classification of Fontannesia.lt is evident from the discussion

above that the genus resembles the Grammoceratinae of the Hildoceratidae family more closely than the Sonniniidae in which
it has been placed invariably. The principal reason for this classification of Fontannesia has undoubtedly been its stratigraphic oc-

currence in the Sonfiinia soxverbyi Zone where

the

first

it is

accompanied by

"true" sonniniids, and the almost entire absence of the

Hildoceratidae, except for the distinct Tmetoceras and the scarce

Astheuoceras in

tiie

Aalenian of Europe

(cf.

Treatise, p.

254)

It

3.
BlanforJicfias icallic/ii noiuiguinctisc Gerth (1917), newly developed specimen from South Geelvink Bay (loc. 4), previously figured under
'Baculatocnas sp.' (Donovan, in Visser & Hermes, 1962, end. 17, fig. 23).
(Shell Research Lab., Utrecht, s.s. 215a) xl.

Text-fig.

has been shown recently that the Grammoceratinae Pseudolioceras

and Astheuoceras range abundantly

northeastern Pacific realm
that

into the Bajocian

(Westermann, 1969b)

Grammoceras and Pleyclellia continue well

same area (Frcbold, 1960, pi. 12; Frebold,

in the

It

(s.s.)
is

in the

probable

into the Aalenian

et al., 1969, p. 31,

above first Tmetoceras). The stratigraphic bias

against placing Fontannesia in the Grammoceratinae should be
eliminated. Certainly the affiliation through descent of Fontarwesia
pi.

1,

figs.

16,17:

with Grammoceras
lution of Sonninia

s-l.

can not seriously be doubted while the evo-

from Hammatoceratidae is generally acceptThus, the Sonniniidae if retained as understood in the Treatise
would be polyphyletic at the family level.
Comparison. The species shows a similar wide range of aped.

s.l.

New Guinea

Ammonites: Westermann & Getty

241

parently continuous variation as

is apparent in tlie European type

grammoceroides (Haug) and in the Australian F. clarkei
(Crick) A large number of probable morphotypes of F. grammoceroides were described as several 'species' by Buckman (1892)
from the L. discites Subzone of the Inferior Oolite of Bradford
Abbas, Dorset. A similar series of forms of F. clarkei was figured by
Arkell (in Arkell and Playford, 1954), probably including the
morphotypes 'F. etheridgei (Whitehouse)
'F. fairbridgei Arkell'

species F.
.

',

and

'F.

whitehousei Arkell', which range from compressed weakly

ornate to more inflated strongly ornate forms. F. clarkei, from the

S.

sowerbyi Zone of the Newmarracarra Limestone,

is

distinguished

Text-fig. 4.
Septal suture morphogeny of Fontannesia aff. F. clarkei (Crick)
[? ssp. kiliani (Kruizinga)] from Kemaboe Valley, at whorl heights of (a)
5mm, (b) 12mm, and (c) 20mm. (R.G.M. Leiden, st. 12685).

from F. grammoceroides s.l. by the more prominent keel, which is

on a clearly tabulate or even slightly bisulcate venter and retained on the body chamber, by the more irregular often fasciculate
and sometimes bifurcating ribs, and by the somewhat wider umbilicus with gentler slope; the relatively involute compressed vari-

set

ants of F. clarkei [T. whitehonsei'~\ agree in coiling with the rela-

Bulletin 256

242

5.
Holotypes of (left) "Gi ammo c eras Kiliani" Kiuizinga (xl) and
(right) "Grammoccras Baumbcrgcri" Kruizinga (xl.25)
both from the Siila
Islands. (Geol. Inst., Univ. Amsterdam, F. 9882 and F. 9883).

Text-fig.

lively

evolute variants of

and

'fairbridgei'

and 12-14%,

grommoceroides. Significantly,

the ventral featmes

species closely resembles F. clarkei

and the irregular

costae;

however,

and
more prominent on

in the slightly smaller umbilicus

projected costae which are

more

approacli the keel

the

s.l.,

more

in

differs

it

strongly

the shoulders

and

closely.

As already pointed out by Arkell

1954, p. 567)

the

morphotypes comprise only 4-5%

respectively, of the Australian sample.

The New Guinea

somewhat

/'".

'xvhiteliousei'

(in

Arkell and Playford,

the Sula Islands specimens described by Kruizinga

under the new names of 'Grammoccras' baumhergeri,

refigured Text-figs. 4,5) and 'Harpoccras' arietitiforme, resemble almost the Western Australia assemblage, except
(1926, pi.

'G.' kiliani

1)

(iiere

that 'G.' kiliani

is

'H.' arietitijorme

somewhat more involute than

is

bisulcate.

According

to

F. clarkei

s.s.

Kruizinga (1926,

and

p. 39)

Gramsaddle;
smaller
external
and
the
moccras in its higher complexity
the suture is as in Fontannesia. All three names were based on
the septal suture of 'G.' kiliani differs strongly from that of

from ex situ concretions found

alleged Toarcian age is, therefore,

single or a few incomplete specimens

as stream

pebbles and their

New Guinea

Ammonites: Westermann & Getty

mere conjecture. The prominently tabulate-carinate

sulcate venter of these forms

is

holotype of Foutanuesia clarkei

fig.

16)

The

and

in the inner

also present in
(refiguretl

to

weakly

bi-

the fragmentary

by Spath, 1939,

whorls of a topotype

243

pi,

2,

(loc. cit., fig, 2)

'G.' kiliani and 'G.' baurnbcrgcri were reand are here refigured. The holotype of '//.' arietitiforme however, appears to be lost. The holotype of 'G.' baumbergeri is a fully grown specimen (55 mm diameter) with a threequarters whorl long probably complete body chamber and approximated last few septa. The internal mold of the venter at 30
mm D is narrowly tabulate-carinate and was probably tricarinate
on the shell. The simple septal suture is poorly preserved. The weak
costation of the nucleus is fascicidate and somewhat irregidarly
tuberculate; on the intermediate whorls, the costae withdraw from
the umbilical seam and become obsolescent on the relatively small
body chamber. The septal sutme drawn by Krvuzinga (p. 40) is
probably 'simplified' due to corrosion. The holotype of 'G.' kiliani,
also an internal mold, is still incomplete at a diameter of approxi-

holotypes of

investigated

"Harpoceras sp."
Text-fig. 6.
from the "Dogger", west coast of
Rijksmuseum Delft, 14903) xl.

[=

? Fontannesia aff. F. clarkei (Crick)],

Jefbie, Misol Archipelago. (Single specimen;

Bulletin 256

244

mately 66

mm

tation

similar but

is

whicli includes part of the

much

stronger

body chamber. The

than on

'G.'

cos-

baumbergeri,

somewhat

fasciculate except for the ultimate halfwhorl where the

innermost flanks become smooth. Part of the Moluccas collection
of Brouwer (which includes the type specimen of Kruizinga)
Avhich is in tlie Rijksnuiseum \i\\\ Mineralogie en Geologie in Delft,

contained eight body cliamber fragments and one almost complete

specimen labeled 'Harpoccras sp., Dogger,' from west coast of

The body chamber fragments, preserved

molds of grey limestone and varying from 18 to 28

Jefbie, Misol Archipelago.

as internal

mm

in wliorl height, resemble the holotype of 'G.' kilinni, although the

costae are generally

more widely

spaced.

The

single large speci-

men, strongly pyritized, with rather well-preserved phragmocone

(70 mm D) and crushed one-half whorl incomplete body chamber,
resembles 'G.' bamnbergeri in tlie strong compression and early
loss of costation,

which, in turn,

is

closely allied

with Fontanjiesia

whitehousei Arkell (1954), probably a variety of F. clarkci (Crick,

The septal suture is well preserved in this specimen and
1894)
.

closely agrees with those of the Australian

fig.

1;

shown

form (Spath, 1939, Textmore complex than

Arkell, 1954, pi. 29), being markedly

in Kruizinga's

(1926,

p.

40)

figure of 'G.'

Affinity with early Bajocian Fontannesia

baumbergeri.

certainly closer than

is

any other known genus, and the Sula Island forms, 'G.'
and G. baumbergeri, are tentatively placed in this genus.
to

kiliani

Of interest is also the occurrence of Fonia^inesia ci. F. clarkei

Turkey (Bremer, 1966, pi. 16, figs. 2 a,b) i.e. geographically
much closer to the European occurrences of Fontannesia.
Most of the New Guinea forms resemble most closely 'Grammoceras' kiliani, which is tentatively regarded as a subspecies of
in

F.

clarkei.

The

close affinity

is

evident by the presence in the

among

Australian assemblage of relatively involute forms

ally

evolute and costate morphotype 'fairbridgei'

the usu-

(Arkell,

1954,

pi. 27, figs. 4,6)

Superfamily

STEPHANOCERATACEAE

Neumayr, 1875

Family OTOITIDAE Mascke, 1907

Genus DOCIDOCERAS Buckman, 1919

Docidoceras (Docidoceras) longalvum (Vacek) 1886,
limatum (Pompeckj) 1897
1886.

Corlorcras loJiqalvum Vacek

12, p. 99,

pi.

17',

figs.

1,

2.

(S.

of.

subsp.
PI. 50, figs. 1 a-d

Alps), Abh. K.K. geol. Reichsanst., vol.

New

1897.

Guinea Ammonites: Westermann & Getty

Cocloccras limatum Pompeckj

49, p. 745, pi. 31, fig.

1922.
1925.

(Anatolia),

deiitsche

Z.

geol.

245

Ges.,

vol.

5.

Docidoceras prrfectum Biickman (England), 'Type Ammonites,' pi. 314.

Cocloccras lougalvutn Vacek \ar. trapan'icum Renz (Sicily), 'Monte San
Giuiiano (Monte Erice)', p. 30, pi. 1, fig. 6.

Material.

and some

Kemaboe

test

complete internal mold, with aperture

single

remains; inner Avhorls crushed on one side, from

Valley.

Description.

The

outer

phragmocone whorls

are

strongly

depressed sublenticular, with lateral edge at mid-flank, and loosely

coiled, almost serpen ticone.

radiate,

commencing

The

primaries are strong and

recti-

seam and strengthweak bullae on the lateral

directly at the umbilical

ening to small tubercles

or,

finally,

edge. The secondaries, usually in threes with one or two attached

weakly to a primary, are dense, blunt and weakly prosoradiate
crossing somewhat convexly over the broad venter.
The body chamber, li^ whorl in full length, ceases width

growth

after

about one-half whorl while height growth continues,

becoming less depressed; the umbilical seam

the cross section thus

egresses gradually so that the aperture

ceding venter.

The

primaries, at

first

merely
still

'rides'

somewhat

on the

pre-

bullae-like,

weaken rapidly and retract from the umbilical seam. The secondaries become wider spaced and blunt, nearly dying out mid-ventrally at mid-length of the body chamber. Throughout the last
one and one-half whorls, a prominent umbolateral groove is developed on the internal mold along the lower umbilical slope,
filled with a porous, probably secondary shell material, and covered
This groove has recently been observed in a
number of south Alaskan Docidoceras and Pseiidotoites species
where the 'porous material' was interpreted to serve the better attachment of the principle retractor muscle (Westermann, 1969b).
by the outer

At

shell.

the aperture

is

a simple oblique prominent flange or collar

of strongly thickened test

the flanks

on one

(4

mm)

on the internal mold

side

preceded by a constriction of

only.

The

flange

but better preserved on the mold.

is

partly visible

The septum

is

typically bullate with

two complete (paired) saddle axes (E/LI/Un and L/U-U/Ui closely resembling the topotype of Vacek (reThe suture is poorly
figured: Westermann, 1964, pi. 6, fig. 2a)
.

preserved.

Comparison.

The

specimen

closely

resembles

'Coeloceras

Bulletin 256

246

limatinn'

Pompeckj

recently been

from Anatolia (Turkey)

from the L. discites Subzone,

which has

(1897)

reclescril:)ecl

soiverbyi

5.

Zone, of the same area

(Bremer, 1966) in association with Eudmetoceras (Euaptetoceras) cf. E. amplectcns (Buckman)

E. cf.
,

dorsatum

(Merla)

'crassispinata'

ceras

(?)

(Crick)
is

15,

2a, b)

iigs.

Buckman [=
(Bremer)

transiens

F. clarkei

litudtinn

(pi.

(E.)

5.
,

and

Sonninin (Eiihoploceras)

adicra

var. ivhitelioiisei Arkell.

distingiiisiied

primaries more or

less

(Waagen)

significantly,

],

Docido-

Eontnnnesia

cf.

Docidoceras longalvum

from D. lotignhum

s.s.

only in the longer

reaching up to the subsequent umbilical

seam of the 'serpenticone' shell. The primaries of the New Guinea

specimen do not, however, reach the length of D. I. limatiim on
the body chamber, while the secondaries are usually less convex on
the European forms.
Docidoceras perjcctum Buckman from England and 'Coeloceras' trapanicum Renz from Sicily were included in D. longalvum,
representing subspecies at the most, after restudy of the type material from San Vigilio (Westermann, 1964).

The

other

known

flated and, usually,

species of Docidoceras

more

involute.

Of

s.s.

special

are

much more

interest

is

in-

the re-

semblance in the projected ribs and presence of an umbolateral

groo\e Avith the south Alaskan Docidoceras spp. which are presently
being described imder a new subgenus (Westermann, 1969b)
Measurements.
end phragm.
body ch.
aperture

W%

U%
49

31

47
46

24

34

Dmm

\\%

53

27

65
93

50
54

(?)Docidoceras (Docidoceras) sp. indet.

P
17
19
c.22

S
55

45

PI. 50, fig. 2

Material. A single incomplete 1/2 whorl body chamber with

poorly preserved remains of phragmocone, internal mold.
Discussion.

On

the last whorl of

tlie

phragmocone, strong

rectiradiate primaries bearing extended tubercles are visible.

The

body ciiamber fragment is medium evolute, tlie section being depressed and markedly ovate with the loiuided lateral edge at approximately 2/5 whorl height. The {primaries are prominent and
somewhat bullae-like, dividing somewhat irregularly into three and
sometimes four blunt moderately prosoradiate secondaries which
are markedly convex on the venter.
This specimen resembles the afore-described Docidoceras long-

New Guinea

Ammonites: Westermann & Getty

247

alvum (Vacek) from whicli it is distinguished only in the more

inflated, somewhat ovate and more involute whorls. However, without preserved septum or suture, the slight possibility exists that
this

a stephanoceratid.

is

Measiirements.

H%

W%

U%

55

31

51

(c.40)

70

31

45

Dmm
body

ch.

"

Family

15

50

42

STEPHANOCERATIDAE Neumayr, 1875

STEPHANOCERAS Waagen, 1869

Genus

Stephanoceras (Stephanoceras) aff. S. humphriesianum

(J. de C. Sowerby) 1825 9
PI. 51, figs.
Material.
three

last

One

a-b; Text-fig. 7

fragmentary internal mold with parts of the

whorls of phragmocone and

the

beginning of body

chamber, one side damaged.

Description. The ultimate and penultimate whorls of the
phragmocone are weakly depressed subelliptical, slightly ovate in
section with the maximal whorl width at about 2/5 whorl height,
with gently sloping inner flanks (umbilical slope) and somewhat

The idtimate whorl is evolute, overlapping only

about one-quarter of the preceding whorl while the penultimate
whorl almost reaches the nodes of the antepenultimate whorl; thus,
flattened venter.

the inner whorls appear to have been less evolute.

The

egression of

the umbilical seam seems to have continued with the body chamber, giving the shell a 'planulate' appearance.

The ornament

of the antepenultimate

whorl

consists of sharp,

ending in tubercles. The primaries of the

last two whorls are also sharp but somewhat rursiradiate and curved
anteriorly. From the distinct round lateral tubercles arise more or
less rectiradiate dense and fine secondaries in groups of three or,
more rarely four, which cross straight over the venter. At the beginning of the body chamber, both primaries and secondaries be-

rectiradiate primaries

come

blunt.

The septal suture (Text-fig. 7) is complex; externally with

large E/L saddle, slender and deep L of the same length as E, broad
bifid L/Uo saddle and strongly oblique Uo; internally with large
I/U saddle, deep oblique Un and much smaller oblique VjU^
saddle.

Discussion.

The

genus Stephanoceras

s.l.

is

in a state of utter

Bulletin 256

248

Adult

septal suture of Strp/iaiioccras (Stcphanoccras) aif. S.

de C. Sovverby), at approximately 10 cm diameter, from
Valley (R.G.M. Leiden, st. 126191).

Text-fig.

7.

humphricsianum

Kemaboe

(J.

confusion and grossly

(Westeimann, 1964,

'split'

p.

66

at the
ff)

suggests that a small fraction

suffice.

The

'planulate' S.

stedt spp.,

this

[^

brodiaei

5.

group

it

is

(J.

and

and involute and also

F.A. Quenall of which are usually

Sowerby)

S. rniitabile,

characterized by moderately fast increasing,

.S'.

JiinnpJiriesianwn Zone of

generally restricted to the

and derived from

.S'.

Queen Charlotte
Zone

huinpJiriesinnnin

Mascke

(Skirroceras) of the O. saiizei Zone. Stemmatoceras

and distinguished only by

the

pressed sublenticular whorls with well-defined

more prominent primaries or

of

To

cnamanoi

S.

the 'serpenticone' but otherwise similar subgenus

difficult to separate

tion

would

inflated

belongs also the northeastern Pacific

(McLearn) from the

S.

is

of the taxa

subelliptical whorls with fine costae bearing tubercles.

'plexus'

Islands;

more

'plexus' of S. iimbilicum

associated; this

rounded

(10-20 per cent)

JiumpJniesianiim , type species, almost

certainly intergrades with the

more common

genus-group and species levels

Experience on several continents

is

edge and
intergrada-

intraspecifically

features,

with Slephanoceras

s-s.

is

strongly de-

lateral

bullae; however, there

probably interrelated

as morphogenetically,

more

as

well

on the one hand and

Teloceras Mascke on the other, the latter becoming distinct usually

only

if

large

and

fully

grown. Consecjuently, Stemmatoceras and

Teloceras are best distinguished from Slephanoceras at the subgeneric level only. In the northeastern Pacific area, the

Teloceras complex of a single bed (Rock Creek

S.

humphriesiaymm Zone of Alberta,

ed into 12

(macroconchiate)

for

'species'

Stemmatoceras-

Member)

in the

example, has been divid-

comprising an apparently

New

Guinea Ammonites: Westermann & Getty

249

continuous sequence of shells with more or less depressed sublenticular whorls bearing coarse primaries of different prominence;
the first named form is S. charlottcnsis (Whiteaves) which reS. triptolemiis (Morris and Lycett) with the inner whorls
(Stemmatoceras) and the outer whorls of Stephanoceras s.s.
Both S. (Stemmatoceras) and S. (Teloceras) seem to occur in the

sembles

of

O.

S.

saiizei

and

S.

humphriesianum Zones, although the latter is

Zone (if not there misidentified) In Europe

rare in the O. sauzei

(Teloceras) seems to range into the

subfurcatum Zone, as
does S. (Steynmatoceras) in the Western Interior of the United
States and southeastern Alaska (Imlay, 1962, 1967)
however, some
of the Western Interior forms could be Cadomites. The microconchs of StephaJioceras s.l. which have usually been classified under Normanyiites s.l., are still difficult to match on the specific and
even on the subgeneric levels and their discussion is not again attempted here (Westermann, 1964)
However, there can be no
serious doubt about their 'generic' correspondence.
The New Guinea specimen resembles closely S. humphriesianum (J. de C. Sowerby) and especially '5. caamanoi' McLearn
which is here regarded as a junior synonym; the occurrence of
somewhat curved primaries as in our single specimen may be somewhat more frequent in North American Stephanoceras than in the
S.

S.

European

representatives, but this

ably not diagnostic feature.

is

a variable

Our specimen

and therefore probfrom 'typical' S.

differs

humphriesianum, in the broader and slightly less evolute whorls,

thus being intermediate to S. mutabile (F.A. Quenstedt)
Measurements.

Bulletin 256

250

inner wliorls with shell remains crushed on one side, ultimate

whorl worn.
Description.

The

inner whorls

(<23

mm

D)

are

medium

evolute and almost planulate, with moderately depressed, some-

what

subelliptical section.

The

penultimate and ultimate whorls,

the latter probably belonging to the body chamber,

more depressed and sublenticular

in section

become rapidly

with the lateral edge

The inner flanks slope gently to the umbilical seam

and the ventral area is evenly rounded.
The ornament is strong throughout consisting of prominent
straight and only slightly prosoradiate primaries which arise rapidly from the umbilical seam but withdraw from it on the ultimate
whorl; they bear roimd tubercles at least on the better preserved
inner whorls where they are partly overgrown by the ultimate
whorl. The secondaries of the inner whorls are concealed; on the
ultimate whorl, they arise in twos or, more rarely, threes from the
primaries, cinving forward but bending backward before reaching
the venter which they cross straight at full strength, thus suggesting weak tabulation. Septum and suture are not preserved.
at mid-flank.

Comparison.

The

etheridgei (Gerth)

here refigured
type,

specimen resembles the holotype of

from the Vogelkopf Peninsula (?) which

( ?
(Text-figs. 8 a-c)

The

S.
is

inner whorls of the holo-

although poorly preserved, can be seen to bear similarly

prominent primaries with

than tlie end of the conch

Text-fig.

2.

Holotype

of

tubercles; they are also

as

is

evident from

tlie

more planulate

relatively

narrow

Strphanocnas (Strmmatorrras) etheridgei (Gerth)

(loc. 1 ?). (R.G.M. Leiden, st. 1272) xL

$, from the Vogelkop Peninsula

New Guinea Ammonites: Westermann & Getty

beginning of the ultimate

251

However, the holotype becomes

the secondaries become j;)rosoradiate only on the ultimate one-half whorl and ventral straightening is only faintly suggested. However, this may be partly due
to the somewhat smaller size.
The ventral costal features of our specimen are imknown from
any other Stephanoceras s.l. and may be characteristic of fully
grown S. etheridgei, together with the remarkable change of whorl
section. The secondaries are also stronger than in most species of
5. (Stemmatoceras) under which name it is kept with modest confidence. Nevertheless, there is some resemblance to the new genus
Irianites in the change of whorl section and the unusual secondaries,
so that this specimen was originally classified with the much more
abundant form.
The West Irian specimens identified by Donovan (iyi Visser
and Hermes, 1962) as Stephanoceras (Teloceras) aff. S. indicum
(Kruizinga) and 5. brodiaei (J. Sowerby) which were reinvestigated and partly further developed from the matrix, are probably
"Coeloceras" indicum
this
species.
Kruizinga
identical with
less

inflated

wiiorl.

than our specimen;

[?

nomen dubium]

(1926)

the poorly preserved holotype of

which

9.
Holotype of "Coeloceras Indicum" Kruizinga [= Steplia?ioceras
(Teloceras) indicum Kruizinga 9], from the Sula Islands. (Geol. Inst., Univ.
Amsterdam, F. 9884) x6.5.

Text-fig.

Bulletin 256

252

is

here refigured

(Text-fig. 9)

Measurements.

Dmm

belongs also to Stephanoceras

s.l.

Dmm

Bulletin 256

254

Comparison

"Sphanoccras cf. hullatum d'Orb."

(above)
Bullatimorphitcs (Treptoccras) aff. B. uhligi
(Popovici-Hatzeg) from supposed Callo\ian of the Wairori River (loc. 1),
with (below) a topotype of B. (T.) "sucvicum (J. Roemer)" [ B. uhligi'\
from the O. aspidoidis Zone, Upper Bathonian, of Hildesheim, Germany (Niedersachs. Landsamt Bcdenforschung) xl. Original Coll. B.F. B./N.L. f B han-

Text-figs. 10-11.
(Gerth, 1927, p.

226)

[=

nover 6537.

?Genus BULLATIMORPHITES Buckman, 1921

Bullatimorphites

?, n.

sp.

PI. 52, figs. 1-3

Two phragmocones with incomplete body chamone pliragmocone; two fragments of large body chambers of
which one has part of the penultimate whorl; all rather well-preserved internal molds with minor test remains.
Descriptio7i. The inner whorls (< 30-35 mm D) are moderately involute with slightly depressed rounded whorls; the intermediate whorls are involute with compressed cadicone section in
Alatcridl.

bers;

New Guinea

The

Ammonites: Westermann & Getty

255

and sharp primaries which

below mid-flank into two, more rarely three very
dense, prosoradiate and continuous secondaries. The outer whorls
(>45-60
D) become gradually much more evolute, depressed
and typically cadicone with vertical umbilical wall bounded by a
weakly rounded margin marking the broadest whorl width. The
primaries retract from the umbilical wall and become rapidly more
costation consists of curved, dense,

divide just

mm

widely spaced swelling into lower-lateral bullae while

the

sec-

ondaries remain dense, being about five times as abundant as the

primaries.

The

full

diameter

is

mm.

estimated at 100-120

ap-

It

pears probable that the large body chamber fragments belonged to

the

same

species as the smaller incomplete specimens.

The

juvenile septal sutme at a few millimeters diameter

(R.G.M. Leiden, st. 126194) has an oblique 'internal lateral lobe'
separating the two saddles and a shallow (U^) lobe at the umbilical
seam, in all closely resembling the early suture of B.} (Treptoceras

?)

n.

sp.

therefore, probably

Ui.

The mature

(Text-fig.

12).

The

external part of the suture

erately complex, consisting of subequal

ed by a narrow
resembles

'internal

homologous with the primary

trifid

L which

is

(st.

lateral

1st

126195)

E/L and L/U

approximately

except for the somewhat smaller

as

lobe'

is,

umbilical lobe,

deep

size;

is

mod-

saddles dividas E;

the other

U^
um-

elements are small and straight, the saddle line rising somewhat toward the umbilical seam.
Discussion. The inner phragmocone whorls resemble certain
Bullatimorphites Buckman, especially B. (?) sojamim (Boehm,
1912, pi. 35, figs. 2a-b) from the Bathonian-Callovian of the Sula
Islands^) which also has dense, long, and curved primaries on the
bilical

Text-fig. 12.
Juvenile incomplete (internal) septal suture
phites ? (Treptoceras) n.sp.
diameter, from
$ at 2.5
(R.G.M. Leiden, st. 126204).

mm

of

Bullatimor-

Kemaboe Valley

1 This
species is probably also present in the M. macrocephalus Zone of
Chos-Malal in the Argentina Andes (Westermann, 1967, fig. 2).

Bulletin 256

256

phragniocone but

is

more
Uo lobe. The intertwo specimens, are more compressed

distinguished by coarser secondaries,

evolutc and depressed whorls, and the multifid

mediate whorls, at least of

tlian in any known (macroconchiate) tulitid. The body whorl Avith
its gradual luicoiling, with steep uml^ilical slope and with bullaelike primaries is, however, more like that of Tulites (Rngijerites)

Buckman,

(Boehm, 1912,

slightly

resembling

T.

(R.)

(?)

godoheusis

from the Sula Islands assemblages; T. (?) godoheusis differs in the more evolute and much
more coarsely ornate phragniocone and the broad bifid Uo lobe.
The late Middle Bajocian Cliondroceras (Defonticcras) McLearn also shows some stiperficial resemblance but is distinguished
by more involute inner whorls, rounded outer whorls, smaller size,
and particularly by the abullate septum with Up. C. (?) (Praetulites) AVestermann (1956) which is probably of late Bajocian (-f-?
Bathonian) age (Sturani, 1964a), has a prominent lateral edge
and short primaries.
The compressed inner whorls, the variocostate (strengthening)

pi.

primaries,

slender

Uo

35,

figs.

la,b)

also

and the somewhat

rising

umbilical lobes with

are all reminiscent of the Eticycloceratinae;

however

the umbilical elements of the suture are by far not so strongly

raised as in that

group

(at least

not in the mature shell)

and the

outer whorls differ in section.

?

Subgenus

TREPTOCERAS

Enay, 1959

Treptoccras was proposed by Enay (1959) for small Tulititidae

combining the features of the inner wliorls of Bullatimorphites
with those of tiie apertme of Schwaudorjia but bearing a more
oblicjue apertmal constriction.
Such a combination was first shown in the figure of A. microstoma d'Orbigny (1846, pi. 143, figs. 3,4) which Arkell (1954, p.
110) considered a synthetogram since none of the material in
D'Orbigny's collection possessed lappets. Lappets do occur, however, in 'A. microstoma' Quenstedt (1886, pi. 78, fig. 4) which
Arkell

(loc.

genus.

New

1958, p. 66)

cit.),

therefore,

probably, a new
Germany (Westermann,

considered to be,

material from northwestern

and eastern France (Enay, 1959) confirmed the

ence of lappets.

pres-

New Guinea

Because of the

much

Ammonites: Westermann & Getty

dimorphic relationship with

probable

larger Bullathnorphites, but in the absence of

correspondences, Treptoceras

257

is

known

the

specific

preliminarily classified as a sub-

genus of tliat genus in accordance with the convention regarding

probable sexual dimorphs (Callomon, 1963; Westermann, 196-1).
More than one-half of the sphaerocones from the Kembelangan Formation cannot be definitely identified as to sex, i.e. macroconch
or microconch but can be classified as Bullatimorphites
cludes the microconchiate B.

if this

in-

(Treptoceras).

Most specimens here described from West Irian differ from

species of B. (Treptoceras) in the more regularly coiled
and not markedly 'contracted' body chamber, in the denser, more
strongly prosoradiate ribs, and probably, in the slender Uo ("second lateral") lobe.
B. (Treptoceras) is known from the Middle Bathonian to the
Lower Callovian. Important occurrences are in the Middle Bathonian of Crussol (Enay, 1959) the Upper Bathonian of the Paris
Basin (de Grossouvre, 1888) and of the Weser Mountains near
Hildesheim in Lower Saxony (J. Roemer, 1911; Westermann,

known

1956b)

the lower Callovian of the Paris Basin

(Corroy, 1932), of

Kutch in India (Spath, 1931), and, probably, of the southern

Andes (unpublished)
While this paper was in press, the senior author was informed
about new and unpublished findings of later occurrences of microconchiate Bidlatimorphites. Dr. A. Zeiss of the University of Er-

W. Halin of the Geological Survey BadenWiirttemberg recovered single specimens from the Middle Callovian
langen-Niirnberg and Dr.

and Upper Callovian

P. athleta

Zone of the Swabian Jura. Unfor-

tunately, the specimens or photographs

Bullatimorphites

(?)

(Treptoceras

?) sp. aff.

were not available.

B.

(d'Orbigny) 1846 i

Material.

One

microstoma
PI. 53, figs. 2 a-c

complete internal mold with partly preserved

aperture and exposed penultimate whorl.

The small shell (48 mm D) is relatively evo(U of phragmocone

23% of D) and weakly inflated for the
genus; the body chamber is only weakly elliptically coiled and not
markedly 'contracted.' The last phragmocone whorl is moderately
Description.

lute

depressed ovate, the vertical umbilical slope gently rounding into

.
,

Bulletin 256

258

maximal whorl width

and with evenly rounded ventral

the flanks with the

at

height,

area.

about one-third whorl

The

dense strongly

prosoradiate primaries divide just below mid-flank into two or

three dense, sharp,

somewhat back-curved secondaries which

cross

almost straight over the venter.

The body chamber,

length,

just over

three-quarters of a whorl

imwinds somewhat more strongly

curves at the end, residting in weakly

the beginning

at

'elliptical'

coiling.

in

and

Whorl

height ceases to grow and whorl width growth decreases so that

more depressed (H/W 0.71>0.62>0.59)

remain dense and become prosoradiate, high and
sharp on the last one-half of the body chamber, now reaching well
beyond mid-flank and bifurcating regidarly into the almost
straight secondaries. Immediately before the aperture, the whorl
contracts slightly and a terminal constriction truncates the ribs
obliquely. From the expanding flange extend ventro-lateral lappets
the whorl section becomes

The

j)rimarics

(one

preserved)

Althougli

incompletely

preserved,

the

lappet

was probably small and simple.

The mature external septal suture has subequal E/L and L/U
("1st and 2nd lateral") saddles divided by a narrow L lobe, and
a slender and trifid U2. The smaller umbilical elements are not
markedly suspensive and are straight to slightly oblique.
Comparison. The specimen differs from all previously described species of B. (Trepioceras) in the body chamber which
is more regularly coiled, not 'contracted' (except for the aperture)
and becomes more depressed rather than rounded, as well as in
the slender Uo lobe. It is closest to the middle to late Bathonian
B. (T.) microstoma (d'Orl)igny) (Enay, private comm.) a species
,

(De Grossouvre, 1888). B. (T.)

laiirculi Enay and B. (T.) crimarietisis Enay are more compressed
with prominent ventral flares at the aperture. The other known

exliibiting appreciable variability

species

of

the

(Boehm, 1912,

subgenus are more involute. B. (B.}) sofanum

from the Sula Islands
p. 150, pi. 35, figs. 2 a,b)

appears to iiave similar inner whorls but

variocostate, the

is

much

body chamber bearing widely spaced

Measurements.

larger

ribs.

and

New Guinea

Ammonites: Westermann & Getty

Bullatimorphites (Treptoceras) sp.

(Popovici-Hatzeg) 1905

One

Material.

damaged

259

B. uhligi

aff.

PI. 53, figs.

internal

a,b, 3 a,b

mold with incomplete

aper-

one incomplete specimen with end of phragmocone, one-half

\vhorl of body chamber and mold of penultimate whorl; both from
ture;

Kemaboe Valley. One damaged but otherwise complete inmold from the 'Callovian' of the Wairori River (original

the

ternal

of Gerth, 1927, p. 226: "Sphacroceras

Leiden,

cf.

bullatiim d'Orb.";

R.G.M.

12118).

St.

The phragmocone is globular with strongly deand involute whorls (U

15% of D) and dense
prosoradiate sharp primaries which bifurcate or trifurcate at
about mid-flank into dense straight secondaries. The complete body
Description.

pressed ovate

chamber, a

whorl in length, has typically 'elliptical' coiling

at the beginning and at half-length, and
contracted resulting in a much less depressed whorl secfull

with marked geniculation

is

laterally

The umbilical slope becomes shallow and the ventral area

weakly convex. Primaries and secondaries, now in pairs, become
somewhat more widely spaced but blunter. The aberrant ventral
costae feature on the body chamber of the smaller specimen is
tion.

obviously pathological, probably a result of pallial injury.

aperture

is

marked by an oblique

("1st

last septal

and 2nd

The
lat-

probably bearing the base of ventro-lateral lappets.

eral flange very

The complex

constriction followed by a

lateral")

suture has broad subequal

saddles

and slender

E/L and L/U

L and Uo

lobes; the

following smaller umbilical elements are straight and somewhat

raised.

The specimen from

fig. 1, loc. 1)

fig.

10)

it

the Callovian of Wairori River

now
two Kemaboe

described by Gerth (loc.

closely resembles the

cit.) is

(Text-

illustrated (Text-

Valley specimens

described above.

Comparison.
Bathonian and

The
(?)

specimen

closely

lower Callovian

B.

resembles
(T.)

uhligi

the

Upper

(Popovici-

from Romania was refigured by Arkell

This species includes 'Sphaeroceras
Suevicum' J. Roemer (1911) (text-fig. 12), which is common in
the O. aspidoides Zone (? and Lower Callovian) of the Hildesheim
area in northwestern Germany (Westermann, 1958, pp. 66,67;
probably also 'B. microstoma microstoma' with lappet) and is
Hatzeg)
(1954,

the holotype

text-fig.

36,

right)

Bulletin 256

260

Lower Callovian forms described from Franconia

(Kuhn, 1939, pi. 3, fig. 31; pi. 7, figs. 2,9) B. iihligi is distinguished
from our specimen by the smaller size and the blunter more widely
spaced ribs at least on the body chamber. B. (B-}) sofanum
(Boehm) is larger, more evolute, and more coarsely ribbed, esB.
pecially on the venter. According to Enay (priv. comm.)
jnicrostoma (d'Orbigny) is distinguished from B. uhligi mainly in
closely allied to

the somewhat wider imibilicus; however, according to Westermann's own brief study of D'Orbigny's original specimen in the
Museum National d'Histoire Naturelle, Paris, this supposed differential feature needs closer study.

Bullatimorphites

(?)

(Treptoceras

Westermann and

(?)

costidensus

Getty, n. sp. (5?)

PI. 54, figs. 1-4

Holotype.-P]. 51, fig. 3 a-e (R.G.M. Leiden, st. 126203); almost complete body cliamber (5/4 whorls ) and part of phragmocone. From pebbles of the Kemaboe Valley, Kembelangan Formation,

West

Irian.

Almost

medium-sized (? microconchiate) Bullatibody

chamber without strong elliptical coilmorphites, involute,
ing or contraction; costation extremely dense, sharp and prosoZ)/V7g7?o5/.s.

radiate throughout, with long primaries.

Fifteen

body chambers, more or less complete but

with some remnants or molds of
phragmocones, several apertural constrictions, but no complete
apertiue; also several smaller fragments. All from pebbles of the
Material.

usually

somewhat

Kemaboe

distorted,

Valley.

Description.

\i

least

the intermediate

and outer phragmo-

cone whorls are tightly coiled (U

18% of D) and depressed
ovate in section, with the vertical umbilical wall rounding into
the flanks.

The phragmocone

is,

therefore,

typically sphaerocone

85%. The body chamber, about

unwinds
gradually,
tlie coiling becoming
whorl
length,
in
a full
compressed by cessaand
becomes
more
only weakly 'elliptical,'
only
weak
or no 'contraction.'
tion of width growth; but there is
The umbilical wall becomes more shallow as the seam egresses. The
final diameter is between 50 mm and 60 mm, except for a single
specimen whicli shows markings of segmental growth and is 70

with a 'thickness' (W/D) of 65

to

New Guinea

mm

The

large.

densely ribbed.

Sc

Getty

261

outer whorls, including the body chamber, are

thin sharp primaries arise more or less recti-

The

on the umbilical wall but soon swing strongly forward.

radiate

They

Ammonites: Westermann

divide at about mid-flank into two or three dense secondaries

which on the flanks are also strongly adoraly inclined. The secondaries tend to bend backward besides the venter which they
cross more or less weakly convex. However, the connection between
primaries and secondaries is weak and the secondaries are occasionally intercalated.

The

is marked by a strongly oblique constriction of

mold truncating the costae. A ventral flare is missing

aperture

the internal

or weak. Although no peristome

is

complete, small remnants seem

were present.
(R.G.M. Leiden, st. 126232)
with subequal E/L and L/U ("1st and 2d lateral")
to indicate that ventro-lateral lappets

The

septal suture

slender, deep-rooted trifid to multifid

last septal

L and Uo

lobes.

is

complex,

saddles

and

The exposed

sinface of the holotype shows also the raised smaller

umbilical elements near the seam,

the

subec[ual

two "internal

and the bullate (eubullate?) septal structine.

Although it is not certain that the aperture had lappets, this
form is probably a large microconch (or male)
Comparison. The phragmocone resembles the Upper Bathonian and (?) Lower Callovian B. (T.) iihligi (Popovici-Hatzeg)
discussed above, except for the denser more strongly inclined ribs
and the more slender Uo. B. (?) sofaniijn (Boehm) differs in being
more loosely coiled, in the stronger secondaries and the larger,
modified body cliamber; but it has a similar Uo lobe. The body
chamber of this New Guinea species is more regularly coiled without marked contraction, and more densely ribbed with strongly inclined primaries and secondaries than in any described Bullatimorphites. The mutual resemblance of the different New Guinea
forms here described and their similarity to the Eucycloceratinae
probably reflect some phylogenetic relationship.
It is possible that this form is the microconch (male) complement to the clearly macroconchiate (female) Bullatimorphites? sp.
saddles"

n.

described above although the inner whorls of the latter are

usually

more compressed

(cf. Pis.

52,54)

262

New Guinea

Ammonites: Westermann

Getty

Sc

263

ber, the primaries extend beyond mid-flank to about 3/5 2/3

whorl height and the furcation may become somewhat irregular,
with some intercalatories. One specimen (PI. 55, fig. 3) differs

somewhat

in

the

numbers

larger

which vary

of secondaries

in

length on the body chamber.

The

marked by an oblique constriction truncating

is no conspicuous ventral flare. This is followed adorally by a narrow collar and broad, probably simple ventro-lateral lappets. Their shape is, however, unknown due to inaperture

is

the costae; but there

complete preservation.

The septum

is eubullate with two complete

(paired) saddle
and shallow smaller umbilical elements around the seam.
The mature septal suture has accordingly, subequal external and
internal "1st" and "2nd lateral saddles," and straight (non-sus-

axes,

pensive)

sutural elements.

(Text-fig.

12;

2.5

mm

Uo

D)

as

is

slender.

The

early juvenile suture

developed from the most complete

U^ lobe which is only one-third as deep

somewhat oblique, asymmetrically bifid (modified tri-

specimen, has a shallow

and

as I

This is significantly like early sutures

(Westermann, 1956a, text-figs. 4,5;
Schindewolf, 1965, text-fig. 263) and good evidence for the normal
("orthochronic") development of the umbilical lobes; the morpho'internal lateral lobe.'

fid)

figured from Bullatimorpliites

logical internal lateral lobe

sphaeroceratids,

is,

therefore,

macrocephalitids,

U^

(yiot

Un

kosmoceratids,

as in otoitids,

and cardiocera-

tids)

Co7?iparison.

This

new unnamed

closely B.} (T.?) costidensus, n. sp.

species

from which

it

resembles
is

most

distinguished

and the thinner, more evolute whorls. Of preNew Guinea form

is best compared with the Swabian B. microstoma
(d'Orbigny) of
Quenstedt (1886, pi. 78, figs. 3,4) (Enay, private comm.) which
is, however, distinguished by the more involute phragmocone, the
rectiradiate costae, the presence of a ventral flare, and the elliptical coiling of the body chamber. Specimens similar to those of
Quenstedt were figured from the Lower Callovian of Franconia
(Kuhn, 1939, pi. 6, fig. 3), and the Upper Bathonian of eastern
by the smaller

size

viously described B. (Treptoceras) species, this

France

As

(Enay, 1959,
in at least

pi.

7b,

fig.

7).

most of the other questionable Tulitidae here

264

Bulletin 256

described, there

is

also

an

affinity to the inner whorls of Eucyclo-

ceratinae: Subkossinatin, Eucycloccras

and

Idiocycloceras

[? also

the Oxfordian or Titlionian Grayiccras Spath], which they also resemble in the slender \].y, however, the umbilical elements of the

suture are not raised but follow a radial saddle line. In contrast,
resemblances to the Middle Bajocian Labyrinthoceras or Chondro-

Cadomites (Polyplertites) (Callomon, private comm.)

mere homeomorphs, particularly because of the
significant differences in the septum and its suture.
Of the previously published Indonesian material there is some
resemblance to the widely umbilicate and densely ribbed 'Macrocephalites keeuwensis y' Boehm (1913, p. 14, pi. 4, figs. 2a, b only)
from West Irian, which could be an Idiocycloceras.
ceras or even

are considered as

^ge.

The

stratigiaphic position of this species

part the same as that of Irianites

with which
Ic)

/.

is

at least in

moermanni (Kruizinga)

was found associated (PI. 55, fig. 4) Because of the

moerman7ii with Bositra biichi (Roemer) (PI. 58,
the age of both forms is post-Pliensbachian and pre-Kimit

association of
fig.

cf.

/.

meridgian.

Measurements,

New Guinea Ammonites: Westermann & Getty

suture.

They were

(?)

all

included in the Macrocephalitidae in the

(Arkell, ct al, 1957, p.

'Treatise'

265

294). At least in the studied

Eiicycloceras, the early juvenile septal suture has strongly raised

umbilical elements, indicating that the septum

the "internal lateral lobe"

Uj

is

as in

eubullate,

is

the Tulitidae

x,

and not Un

Middle Jurassic sphaerocones including the macro(Westermann, 1956a, p. 258; Schindewolf, 1965, Abb.
179) The suture differs from Tulitidae (and Bajocian

as in all other

cephalitids
248, 262, p.

sphaerocones) in
?

blage.'

tlie

slender

U^ but resembles the Bullatimorphites

Kemaboe Valley 'pebble assem-

here described from the

n. spp.

similar sutural pattern

may

possibly also be present in the

Upper Jurassic mayaitids although their juvenile stages are still

unknown and there is exceptionally strong variation in the adults
according to the figures of Spath (1928) It is tentatively suggested
elsewhere (Westermann, 1970) to classify both the mayaitids and
.

the eucycloceratids either as subfamilies of the Macrocephalitidae

or, alternatively, to

place

Genus
(?)

them

in a single separate family.

EUCYCLOCERAS

Spath, 1924

Eucycloceras intermedium Spath, 1928 ($

?)

Boehm

PL

1912.

Macrocephalites krruivcnsis,

1928.

graphica, Suppl. 4, pi. 38, fig. 3 a,b only.

Eucycloceras intermedium, nom. nov., Spath (Cutch),
N. S., vol. 9, mem. 2, p. 210, [for preceding].

sp.

nov.

/3,

(Sula

56, figs. 2 a,b

Is.),

Palaeonto-

Paleont.

Indica,

Material. Single fragment of one-half whorl of incomplete

body chamber and remnant of penultimate whorl, internal mold.
Description. l^he body chamber fragment (50 mm D) is
moderately involute and strongly compressed with flat sides, vertical umbilical wall and narrow, slightly tabulate venter. The ribs
are sharp, moderately dense and slightly falcoid. The curved prosoradiate primaries bifurcate or trifurcate at or just below midflank; the secondaries are rectiradiate but on the shoulder markedly projected and prominently arched over the venter. The septal
suture is moderately complex with raised umbilical elements on
ultimate and penultimate whorls (suggesting that the septum is
eubullate)
Discussion. This specimen closely resembles the holotype
from the Sula Islands, but the specimens figured from northwestern
New Guinea by Boehm (1913, pi. 3, fig. 3, ? 4; pi. 4, fig. 4, ? 5) under

Bulletin 256

266

the

same original name of "Macrocephalites keeuwensis ^" are

probably more involute and more densely ribbed; these forms seem
to be intermediate to Subkossmaiia which they resemble in the
coiling

and ribbing, while they appear

to

have the venter of

Eiicycloceras.
?

Eucycloceras

sp. indet., $

PL

56, figs. 1 a,b

Material. A single small, almost complete specimen, with

approximately one-half whorl of body chamber.
Description. The specimen, Avhich may be either a juvenile
stage or a microconch, resembles closely the apparently complete

microconch figured by Boehm (1913, pi. 4, figs. 3 a,b) from northwestern New Guinea luider "Macrocephalites keeinveyisis y var.
D) has medium evolute whorls
bifurcata." The nucleus (5-10
with depressed oval section and ventrally arched secondaries. The
D) becomes less depressed, only slightly
body chamber (20-33

mm

mm

broader than high; somewhat prosoradiate primaries of medium

strength and spacing bifurcate at almost mid-flank into rectiradiate secondaries
ceptible arch.

The

which

cross over the venter

with hardly per-

septal suture has strongly raised umbilical ele-

ments as early as at
septum is eubullate.

mm

diameter, strongly indicating that the

Genus SUBKOSSMAIIA Spath, 1924

The

distinction of Subkosstnatia from Eucycloceras appears to

be poorly defined, due

to the presence of intermediate

that separation at the subgeneric level

is

forms so

suggested here, although

separation at the generic level was retained in the 'Treatise.'

Subkosstnatia obscura Spath, 1928, boehmi

Westermann and
Holotype.
16, text-fig.

Getty, n. subsp.

Macrocephalites

9 and

PI. 56, figs. 3,4

keeincensis

pi. 5, fig. 2 [yion

Boehm,

/?

-y

Boehm,

1913, p.

19I2J, from Mamapiri,

N.W. New Guinea.

The use of the

Boehm (1912, 1913)

suffices a

y 8

and

by
comprehensive

their combinations

to distinguish varieties of his

species Macrocephalites keeuwensis does not constitute the estab-

lishment of a new name, because they directly contravene Article

New Guinea

Ammonites: Westermann & Getty

267

llg of the International Code of Zoological Nomenclature (1964)

They

are,

therefore,

When

without nomenclatural status and are not

(1928, p. 212) emended Boehm's name to

'Macrocephalites keeinvensis beta-gamma,' it is clear tiiat he did

available.

Spatli

not intend to propose a

new name, because he

still

attributed

its

authorship to Boehm. This action of Spath's could be regarded as

an unjustified emendation. It certainly cannot be regarded as the
establishment of
Arkell

the

(1956, p. 448)

new name

'Siibkossmatia

beta-gamma/

as

erroneously stated.

Because the species figured by

Boehm

(1913, pi.

5,

fig.

2)

is

from the true Macrocephalites ('Dolikephalites') keenwensis (Boehm) it is renamed here Subkossmatia obscura Spath
boehmi, n. subsp.
Diagnosis. A subspecies of S. obscura with broad almost quadrate whorls, body chamber with short primaries and dense secdifferent

ondaries.

Material.

One

body chamber complete except

for

ventral

part of aperture, with incomplete penultimate whorl; one incom-

body chamber with fragment of penultimate whorl; both inmolds with test remains, from the Kemaboe Valley.
Description. Both specimens resemble closely the holotype
from Mamapiri. The moderately involute last phragmocone whorl
plete

ternal

has extremely dense, sharp, strongly prosoradiate primaries which

become much more

chamber.

and shorter on the body

body chamber and penulti-

coarse, widely spaced,

The whorl

section of the

rounded trapezoidal, almost subquadrate, with

and flat slightly converging flanks which
round into the broad only moderately convex and very slightly
tabulate venter. On the body chamber, the primaries divide unevenly, sometimes dichotomously between 2/5 and 1/2 whorl
height into mostly three weakly prosoradiate secondaries which
cross prominently over the venter. The last suture of the more
complete specimen shows a slender Uo and highly raised umbilical
elements; the septum is typically bullate with two subequal external and internal 'lateral' saddles. Similarly raised umbilical elements with narrow Uo are preserved on the penultimate whorl of
mate whorl

is

vertical umbilical slope

the other specimen.

The

aperture

is

marked on

the internal

mold by

a strongly

Bulletin 256

268

oblique curved constriction at least on the flanks; shell and ventral part of the aperture are not preserved.

Comparison.

This

new

subspecies

is

distinguished from Sub-

Waagen, 1875,
non "Ammonites Opis" J. de C. Sowerby]
by the somewhat broader whorls (end phragmocone: H/W 1.07
vs. '' 1.27)
and the shorter primaries and denser secondaries at
least on the body chamber. Contrary to Spath's opinion (1928, p.
212) both have the same coiling. The similar 5. opis (J. deC. Sowkossniatia obscura Spath [for "Stephanoceras Opis"
p. 1-40, pi. 36, figs.

a,b;

erby)

[holotype refigiued by Spath, 1928,

pears to be distinguished from

S.

pi.

38,

figs.

2 a,b]

ap-

obscura mainly in the more com-

pressed whorls with arched venter and ribs, although no view of

the
ber,

phragmocone venter of S. opis was given and its body chamwhere Spath's (p. 211) reported measurements were taken,

seems to be slightly crushed; even to Spath

distinction appeared doubtful.

are

more

strongly prosocline

The

(loc.

cit.)

the specific

primaries of the outer whorls

on the holotype

of 5. opis than

on

the holotype of 5. obscura, although the inner whorls appear to be

similarly ribbed. These differences may be less significant than

the broader whorl section, shorter primaries

of the

New Guinea

tinguish this form

and denser secondaries

form; however, the aiuhors hesitate to

more than

dis-

from the obviously very similar Indian forms which cannot here be revised.
A closely affiliated if not identical form from New Guinea
was figured by Boehm (1913) under "Macrocephalites keeuwensis /?". The larger specimen (op. cit., pi. 4, fig. 4) still has most^^of
the oblicpie aperture with strong constriction and peristomal collar.
The fragmentary smaller and more evolute specimen figured under
the same name (op. cit., pi. 3, fig. 4) may also belong to this subspecies. The supposed "? Labyrinthoceras sp." of Donovan (in
Visser and Hermes, 1962, j). 55) which was restudied in the Shell
collection, Rijswijk, has also the typical costation and septal suture of Subkossmatia and a similar whorl section to the new subspecies.

at the subspecific level

New Guinea

Measurements.

Ammonites: Westermann & Getty

269

W%

Dmm
R.G.M. Leiden,

88
60

84
figs,

aperture
phrag.

of

Boehm

37.5

36.5

49

46

45
;

43

c.36
47.5

PERISPHINCTACEAE

29.5
19.5
c.l9

enlarged?)

text-fig. 9

c.llO
C.60

Superfamily

U%

126210

st.

aperture
end phrag.
St. 126211

body ch.
Holotype (from

H%

c.43

c.26

49

17

Steinmann, 1890

Family PERISPHINCTIDAE Steinmann, 1890

(?)

Subfamily 'PSEUDOPERISPHINCTINAE' Schindewolf, 1925

[= Siemiradzkiinae Westermann,

1958;

Grossouvriinae Spath, 1930]

Genus COBBANITES Imlay, 1962

The genus Cobbanites, based on the type species C. talkeelnamis Imlay (1962) was compared by its author with the ChoffatiaSiemiradzkia

[including Pseudoperisphinctes^

assemblages of Po-

Neumayr, 1871) and northwestern Germany (see Westermann, 1958), respectively of Upper Bathonian-Lower Callovian
and of Upper Bathonian age. Cobbanites was said to be distinguished by the weaker and denser primaries on the body chamber,
the more strongly projected secondaries, and to be generally
characterized by the strong, projected constrictions which also distinguish it from the otherwise similar Procerites. Cobbanites is
land

(see

homeomorph of the Upper Bajocian LeptosphincBecause of the absence of a 'coronate' stage, Cobbanites is
probably best placed in the 'Pseudoperisphinctinae' for which was

actually a close
tinae.

substituted the

name Siemiradzkiinae

Pseudoperisphinctes

since

the nominate genus

subgenus of Sieiniradzkia (Westhowever, the name Grossouvriina Spath,

at best a

is

ermann, 1958, p. 83)

1930, would have priority if Siemiradzkia and Grossoinnia are retained in the same subfamily.
All hitherto described species are from the Upper Bathonian
to Lower Callovian of western North America.
;

Cobbanites

engleri (Frebold) 1957

PI. 51, figs. 3 a,b

well-preserved internal

mold with incomplete

(?) sp. aff. C.

Material.

One

7/8 whorl body chamber, one side slightly crushed.

Bllletin 256

270

Description.
rate

becoming

The

slightly

maries appear at 2

whorls are evolute and rounded siibquad-

compressed on the body chamber.

mm

diameter.

They soon become

The

strong,

prial-

most bladelike and prosoradiate, arising rapidly on the umbilical

margin and reaching maximal height at or slightly above midflank; here they bear small tubercles and divide into strongly projected secondaries which at least on the ultimate whorl, are medially
more or less completely interrupted by a smooth band. On the
long body chamber, the primaries weaken and lengthen gradually
while tubercles are missing. There are three strong oblique constrictions on the ultimate whorl and possibly also on the inner
whorls.

The

the end of the

somewhat approximated. The complexity is only moderate but this may be due to
'senility.' The E/L saddle is large and bipartite, L somewhat asymmetrically trifid, L/U2 bifid and much smaller than E/L, while
the umbilical elements are small and moderately suspensive.
Comparison. There is close resemblance to C. engleri (Frebold) especially to the paratype from the basal Callovian Gryphaea
l)ed of Alberta (Frebold, 1957, pi. 40, fig. 1) of which a plaster
cast was kindly supplied by Dr. Frebold. However, C. engleri is
more compressed and has a more complex septal suture. C. talkeetJianus Imlay, from the Upper Bathonian and Lower Callovian of
southern Alaska and Montana, differs also by more compressed
whorls, larger size and less inclined ribs.
There is also close resemblance to Leptospliinctes, especially
the subgenus VeryyiispJiinctes Buckman which locally abounds in
the Upper Bajocian of Europe. While L. (Vermisphinctes) agrees
in the moderately complex septal suture, its innermost whorls are
usually distinguished by being 'coronate' and the ribs less strongly
internal

septal

suture

phragmocone where the sutures

is

preserved

at

are probably

prosocline. Nevertheless,
iety of forms,

still

tlic

Leptosphinctinae include a large var-

difficult to classify

and

largely

homeomorphic

with the 'Pseudoperisphinctinae' being separated partly on their

age difference.

New Guinea

Measurements.

Ammonites: Westermann & Getty

271

Bulletin 256

272

%
(?)Un
,-i.

-.^"1

Text.fig. 13.
Septal suture morphogeny of Irianitcs mocrinatnii (Kruizinga)
6 ,comi)iled from four specimens from Kemaboe Valley; at whorl height of (a)
0.9 mm. (R.G.M. Leiden, st. 126230), (b) 1.0 mm, (c) 2.0 mm, (d) 2.5 mm.
(st.
(st.
126213), (e) 3.5
(st.
126231), (f) 5.5 mm, and (g) 12

mm

mm

126220).

and MidcUe

Bajociaii XortntDinitcs

(Donovan

in Visser

mes, 1962) and jjrobable Slephauoceras (Boehm, 1908)

conch of

/.

(1.

rnoerwanni

lias

either not been

and Herthe macro-

foimd previously

New Guinea

Ammonites: Westermann & Getty

more probably, has

or,

ceras

273

usually not been separated from Stephano-

Previous age estimates for Irianites have thus ranged from

s-l.

the early

Lower

Middle

Jurassic to the middle

Middle Jurassic

(incl.

Callovian)

is

now

Jurassic,

while late

indicated by faunal asso-

ciation.

The taxonomic
Stephauocerns
is

now

s.l.

position of 'Coeloceras' indicum Kruizinga in

(Jaworski, 1933; Arkell, 1956;

Donovan,

op.

cit.)

confirmed; before examining the holotype, the senior author

considered

it

to

be the macroconch of

Coeloderocevas Spath

/.

moermanni.

distinguished by ventro-lateral instead

is

of mid-lateral tubercles, the loss of umbo-lateral tubercles or bullae,

the

somewhat arched

The

denser

and the umbo-lateral

ribs,

prominent

The
is

and probably the simple aperture.

venter,

Dactylioceratidae are similarly distinguished besides having

if

occurring

much

tubercles or bidlae are

less

it all.

stephanoceratid microconch Normannitcs Munier-Chalmas

not tabulate and never bears umbo-lateral bullae or tubercles.

The complete septal suture (Text-fig. 13) with its reduction of

Uo and

!!, and its overall

(Upper Bajocian to

the adjacent saddles, the deep probable

Parkinsoniidae

simplicity, resembles certain

Lower Bathonian)

The U modus

Ui")

of Parkinsonia Bayle

Abb.

10)

(originally as "heterochronic

was observed by Westerinann (1956a

(1965, pp. 206-212) who, therefore,

and by Schindewolf

placed the Parkinsoniidae in the superfamily Stephanocerataceae.

However, the
Mascke,

Uj,

is

fact that in the

usually absent

apparently closely related Garantiana

(but present in a few

Schindewolf,

taxonomic usefulness of
this character, at least regarding higher taxonomic levels. Nevertheless, there is resemblance to the Parkinsoniidae also in the tabulation of the venter, in the tendency to evolute coiling, and finally,
in the ribbing with the exception of the lunbo-lateral bullae and
1965, p. 213-218), suggested the restricted

the ventral continuity.

also

present

in

the

somewhat

similar suture development

Morphoceratidae

which

probably

Buckman (Schindewolf, 1965, pp. 228-231)

the umbilical elements of morphoceratids are straight

Asphinctites

features such as constrictions are absent in Irianites.

superficial resemblance

to

certain

Buckman which

however,

and other
There is also

Callovian Kosmoceratidae,

particular to the microconchiate Torricellites

miceras

is

include

Buckman and

in

Giiliel-

also bear bituberculate primaries; however.

274

Bulletin 256

the whole family without

known exception

fluting type of the septum, with

and

saddles',

mann,
trast,

externally, the large

the abullidisculate

lias

two subequal 'internal

L/Uo

saddle and

lateral

U2 (Wester-

1956, pp. 242,265; Schindewolf, 1965, pp. 187-191). In conseptum of Irianites is of the modified planulate type such

the

as in the stephanoceratids and in most perisphinctids. There is also

some resemblance to certain Reineckeiidae (Callovian)
However, as indicated above, identification of U^ in Irianites is
somewhat tentative. The deep indentation of the primary 'internal
lateral saddle' might have originated at its base, which is the flank
of (?) Ui, and could then be considered a part of a bifid Ui; this
modus is said to be common among the Lower and early Middle
Jmassic ammonites as well as among the late Middle and Upper
Jurassic Perisphintaceae
(Schindewolf, 1962-66)
However, this
splitting of Uj is usually more or less symmetrical and the early
.

sutural elements are mucli higher in early Jurassic forms; furthermore, Kruizin.ga's interpretation of the Irianites suture bearing a
single indented broad 'second lateral saddle'

composed

is false;

this structure

two small saddles enclosing a small Uo.

There is stronger resemblance in the suture to the Perisphinctidae,
particularly to their Middle Jurassic representatives which have an
inclined inner branch of Ui according to Schindewolf (1966, p.
is

actually

332,

of

(in the 'Pseudoperisphinctinae' originally interpreted as

ff.)

'heterochronous

The

U/)

possibility

berriasellid

that Iriayiites

tion set hy Bositra biichi

Irianites

is

Tithonian or Neocomian

excluded through the pre-Kimmeridgian age limita-

is

moermanni

(Roemer)

(Kruizinga, 1926)

(PI. 58, fig.

Ic)

$
Pis. 57,58; Text-figs. 13-16, 20-24

aff. Braiki'nridgii J. Sovverby sp., Boehm (Barbar Is.),

Neues Jb. Min. Geol. Pal., B.B. 25, p. 330, text-fig. 4, pi. 12, fig. 3
[same specimen as Jaworski, 1933, pi. 11, fig. 8]
Corlocrras ynorrmanni n. sp., Kruizinga (Sula Isl.), Jb. Mijnwezen, vol.

?l90i. Stcp/ianoccras

1926.

1933.

54, p. 44, pi. 13, fig. 2, text-fig. on p. 44.

Corlocrras morrman/ii Kruizinga 1926, Jaworski (Niederl. Indies), Neues
Jb. Min. Geol. Pa!., B.B. 70, B, p. 321, text-fig. 6, pi. 11, fig. 1 [holotype

refigured].
?1933. Corlocrras aff.
1956.
1962.

morrmanni Kruizinga, Jaworski,

id.,

p.

323,

pi.

11,

fig.

[same specimen as Boehm, 1908, pi. 12, fig. 3].

Corlodrroceras Imorrmanni (Kruizinga), Arkell (World), p. 441.
Normannitrs cf. morrmanni (Kruizinga), Donovan in Visser and Hermes
(West New Guinea), Verb. Kon. Ned. Geol. Mijnbouwk, Genoot, Vol.
8,

20,

text-fig. 7

p.

55,

end.

17,

figs.

24 a-c.

New Guinea

Ammonites: Westermann

Sc

Getty

275

Fifteen

more or less complete specimens, several

incomplete or juvenile specimens and several fragments (approximately 50 more specimens were retained and only briefly inspected
in the Rijksmuseum, Leiden)
from the Kemaboe Valley, West
Material.

Also

Irian.

two largely crushed specimens from Taliabu, Sula

Islands.

The data for the principal morphological feawere plotted in bivariate scatter diagrams (Text-figs. 21-24),
together with the corresponding data of the macroconchiate /. cf,
I. fnoermanni $
(see below)
Description.

tures

The

small microconchiate shell, mostly between 50 and 65

(range 45-80

mm)

in

maximum

diameter,

is

out with evolute, slowly expanding whorls.

the nucleus

is

mm

'serpenticone' through-

The whorl

section of

depressed oval becoming subcircular at about 8

more or

mm

markedly tabulate; and the outer whorls are subquadrate with narrow steep
umbilical wall, flattened sides and venter (Text-fig. 20) The body
chamber is 5/8 to 7/8 whorls in length, becomes slightly more
evolute and terminates in a lappet-bearing aperture. The exact
shape of the lappets is iniknown but simple spatulate form is suggested. Besides this more common form of about 50-65 mm diameter,
there are a few specimens which are distinguished by somewhat larger size of up to 80 mm and slightly more inflated whorls (PI. 58, figs.
diameter; the intermediate whorls are

less

3,4)

The

largest lappet-bearing specimens

measured 80 mm. This

less

rather than a separate taxon

common form

(retained in Leiden)
is

probably a variety

cannot
be solved with the available specimens without stratigraphic information. However, this question is of importance since the holo(subspecies?)

a question that

type (see below) appears to belong to this form.

The

costation consists of characteristic bulli-tuberculate pri-

maries with peri-umbilical bullae and mid-lateral tubercles, and

high sharp secondaries in groups of twos and threes with bullae-

on the shoulders and some weakening over the

mode of costation is more or less strongly
developed at least on the intermediate and outer whorls while the
nucleus up to 8-12 mm diameter may be densely and simply costate
with bladelike strongly prosoradiate primaries (20-27 /whorl) and
similar sharp secondaries. Towards the end of the body chamber.
like

swellings

venter.

The

described

Bulletin 256

276

tlie

primaries usually lose bullae and tubercles, again becoming

bladelike.

Finally,

simple

bladelike

ribs

may

exceptionally

be

on the internal mold, throughout most of the shell

(PI. 57, fig. 4)
Some reduction of bullae and tubercles is also
present in tlie larger slightly more inflated specimens.
The mature septal suture (Text-fig. 13 g) is weakly incised
with less than average complexity. The morphogeny has been observed commencing at about 3 mm diameter; however, the series of
observations is composed from several specimens. At 0.9 mm whorl
height (H)
the internal sutine appears to have a broad single
saddle which is weakly indented by a lobe on the outer part of its
crest. Slightly later, this lobe is in almost straight to somewhat
oblique position dividing a larger inner from a slightly to markedly
present, at least

smaller outer saddle; this element

is

almost certainly !)

(earlier

called "lieterochronic Uj"), rather tlian a i^ranch of a bifid Uj.

and broad;
L is broad and short-stemmed; the adjacent L/Uo saddle and U^ are
small, more or less straiglit, and tlie following U2/U3 saddle is of
is

broad and deep; the adjacent external saddle

similar

size,

The

suture then drops sharply beside the seam to

moderately low, simple

adjacent to the seam

U.-j

is

(?) U-j.

belong to Ui)
Hololype and 'qua.si-lopotypes'.
in the Sida Islands,
;

The

small

probably the reduced Uj

and U4 would be developed and

(1933)

large

the tlnee elements almost appearing as a single broad

bipartite saddle.

the

is

all

The

element dorsally

no
would

(alternately,

sutural elements

holotype, found ex situ

was newly described and figured by Jaworski

Jawoiski's photograph, his drawing of the body

chamber

whorl section and measurements, as well as Kruizinga's (1926)

original drawing of the septal suture which was verified by Jaworski, are heie reproduced
(Text-fig. 14)
However, the search for
the type specimens in the Mineralogical and Geological Museum
in Delft where the collection was originally deposited, as well as in
the Geological Institute in Amsterdam, wheie the other type specimens described by Kruizinga are now dejjosited, was unsuccessful.
The holotype was apjjarently somewhat developed from the matrix
by Jaworski enabling him to make more accurate measurements
of the last whorl; the body chamber (at about 1/3 whorl length)
is somewhat compressed and not depressed as originally described.
.

New Guinea

Ammonites: Westermann & Getty

277

14.
Holotype of Iriariitrs moermanrii (Kruizinga)
$. Lateral
view and whorl section reproduced from Jaworski (1933, pi. 7, fig. 7, and textfig 6); septal suture reproduced from Kruizinga (1926, Text-fig. on p. 44).
Approximately xl.

Text-fig.

The end

of the

septal sutures

phragmocone

(see

is

marked by

approximated
50 mm D;
well preserved while about
several

Kruizinga's original figure)

at

c.

only 1/3 whorl of the body chamber is

an additional 1/4 whorl is damaged so that the original

diameter was close to 70

maximum

mm,

approaching the largest specimen

described herein. However, none of the body chambers of the New
Guinea specimens is as strongly compressed as the holotype
Fortunately, two partly
(W/H
0.8; Jaworski's measurement)
crushed specimens from Tangi, Wai Miha, Taliabu, in the Sula

(Geological Inst.,
Islands, were found in Brouwer's collection
Amsterdam, catalogue No. Z 8639) from which the missing holotype
was taken. These specimens bear the original labels "Coeloceras
moermanni" and probably are the ones mentioned in the text by
Kruizinga (1926, p. 45, 3 rd. line) The larger one of the 'quasitopotypes' is here figured (Text-fig. 15) There can be little doubt
that these specimens are conspecific with the holotype and with the
microconchs described herein from West Irian. Significantly,
.

Bulletin 256

278

Iiiaiiitcs
motrmatuii (Kriiizinga)
$. 15, 'quasi-topotype'
from Tangi, Wai Miha, Taliabu in the Suia Islands. (Geol. Inst., Univ.
Amsterdam, Z 8639) xl. 16, specimen from South Geelviniv Bay, West Irian
(also figured in Visser and Hermes, 1962, end. 17, figs. 24 a-c) (Shell Res.

Text-figs. 15-16.

Lab., Utrecht,

s.

s.

190 d)

xl.

Brouwer's collection from

macroconch

(Text-fig. 18)

chiate Irianites

cf.

/.

Wai Miha,
which

also includes a single large

closely resembles the

macrocon-

moermanni here described from West

Irian.

Because the search for the holotype is not yet concluded and since
the 'quasi-topotypes' are poorly preserved, no lectotype or neotype
is

now

designated.

Comparison and dinwrphic correspondence. The single fragmentary specimen consisting of incomplete body chamber and part
of the phragmocone, which was described by Boehm (1908, p. 330,
pi. 12, fig. 3, text-fig. 3) from Babar Island, approximately 600 km
southwest of New Guinea, was also studied by Jaworski (1933,
p. 323, pi. 11, fig. 8, text-figs. 7a, b) and probably correctly placed
in close affinity to

and drawings of
(Text-fig.

17)

/.

nioermayuii.

The photogiaphic

reproductions

important specimen are here reproduced

According to Jaworski who had the specimens at
this

New Guinea

Ammonites: Westermann & Getty

279

Text-fig. 17.
Irianitrs (?) aff. /. mocrmanni (Kruizinga)
$, from the
island of Babar, N.E. of Timor. Side view from Boehm (1908, pi. 12, fig. 3);
ventral view and section of ultimate and penultimate whorls from Jaworski
(1933, pi. 11, fig. 8 and text-fig. 7a-b). Approximately xl.

hand, the Babar specimen

is

distinguished from

/.

moermanni

merely by the "absence of the peri-umbiHcal tubercle-line" (translated from German)

However, Jaworski's drawings of the whorl.

Babar specimen has bladelike primaries

and more depressed, nontabulate whorls. Generic identity is highly
probable, although the Babar specimen appears to belong to a new
section indicate that the

species.

The

striking similarity of the

phragmocone with the inner and

intermediate whorls of the macroconchiate Irianites

manni

(Text-figs. 20-24)

latter form.

One cannot

that their relationship

Age.

Two

cf.

/.

moer-

discussed under the description of the

help but draw the tentative conclusion

that of

complementary sexual dimorphs.

incomplete valves and one almost complete valve

of Bositra buchi (F. A.

ing as "Buchii")

is

is

Roomer)

(1936, pi, 4,

=: Posidonia alpina

and

fig. 8;

original spell-

P. ornati auct.]

were

embedded in the matrix at the aperture of a complete although

damaged specimen of Irianites moermanni $ (R. G. M. Leiden, st.
126217) from Kemaboe Valley. The side view of the ammonite
(PI. 58, fig.

a)

shows one of the incomplete pyritized valves (the

280

rest

Bulletin 256

is

closely

resembles

the

figured by Jefferies and

to these authors,

(op.

and could tlierefore not be prepared)

is shown enlarged (PI. 58, fig. 1 c)
and

inside the peristome

the ahnost complete valve

(it.,

p.

Bathonian-Callovian

Minton

who have

specimens

(1965, pi. 19.

reviewed

much

figs. 4, 7)

recently

According

of the previous literature

156-157), B. buchi ranges from the Toarcian to the

Oxfordian and

its

upper limit coincides with that of the genus and

of the entire posidoniid family. This most unamijiguous bivalve

occurrence therefore significantly

indirectly

that

associated with

of
it

the

in a single

probably, post-Pliensbachian.

Measurcincnts.

restricts the

age of Irianites

block

to

and

were
pre-Kimmeridgian and,

Bulla tiinorphites-UkG

forms

wliich

New Guinea

Text-figs. 18a-b.
section of

cf. I. mocrmanni (Kruizinga)

?, from the Derr-ta
Taliabu, Sula Islands. (Rijksmuseum Delft, 14885) xl.

(18b: apertural view, see

I.

281

Irianites

Wai Miha,

Irianites cf.

Ammonites: Westermann & Getty

p.

282.)

moermanni (Kruizinga)

Diagnosis.

Shell

large; inner

Pis. 59-62; Text-figs. 19-24

whorls planulate, tabulate, with

bituberculate primaries; outer whorl inflated, coronate, Sternmatoceras-to Teloceras-like.

Material.

One

and at least 20 moderfragmented specimens; several im-

well-preserved specimen

ately to poorly preserved partly

mature specimens probably identical with this macroconchiate form.

All are internal molds with minor test remains without apertures,
from the Kemaboe Valley.

Bulletin 256

282

Text-fig. 18b (see lext-fig. 18a,

Description.

The

inner whorls

(<

p.

281)

40-55

mm

diameter)

are

evolute with slightly depressed to subcircular section, soon becoming

tabulate and

somewhat subquadrate. The primaries are bullate at

and luberculate at mid-flank where they divide

the umbilical edge

into two secondaries with occasional intercalatories.

are raised

which

The

secondaries

on the shoulder and somewhat weakened over the venter

tliey

go straight

across.

The outer whorls (> 40-55 mm diameter) become rapidly

much more inflated, through positive allometric width growth, and
depressed subelliptical in section (Text-figs. 20,21,23) the ventral
tabulation weakens or becomes obsolete, finally being lost in the
;

New Guinea

Text-fig. 19.

Adult

Ammonites: Westermann & Getty

283

septal suture of Irianitcs cf. /. moermanni (Kruizinga)

whorl height. (R.G.M. Leiden, st. 126222).

2, at approximately 25

mm

largest whorls. Tlie umbo-lateral bullae persist for

some time mov-

ing somewhat higher on the inner flank, and finally fuse with the

high on the lateral edge, to form short

lateral ttibercles, situated

bladelike ribs.
intercalaries,

The
cross

the broad venter.

'coronate'.

largest

strong secondaries, in twos with frequent single

more

The

or less straight

outer

11/^-2

The body chamber

more or

less

occupies nearly one whorl in the

complete

diameter, but the aperture

is

and undiminished over

whorls are therefore typically

mm

specimens measuring 80-90

not preserved in any. Taking into

consideration several large fragments, the terminal diameter was

approximately 90-120

The

mm.

adult septal suture (Text-fig. 19)

is

weakly incised only.

broad and short-rooted; the adjacent simple ('second') lateral

saddle L/Uo is about as high as the external saddle E/L and only
slightly higher but much broader than the oblique 'third lateral'
saddle U0/U3, which is separated by the small simple lobe Uo; the
suture drops sharply adjacent to the umbilical seam with one or
is

two small oblique elements probably belonging

to U3; the internal

suture consists of a dissimilar saddle pair, the inner one being

about twice as large as the outer one, separated by a large almost
straight

'internal

lateral'

lobe

(U?)

the

lobe adjacent to

the

seam (? Uj) is approximately as deep as the 'internal lateral' lobe.

Comparison and dimorphic corresponderice. AXlhough. superficially

resembling Stemniatoceras

or,

if

fully

grown,

Teloceras

(both probably best classified as subgenera of Stephanoceras) this

form is clearly distinguished by the planulate and tabulate inner
,

Bulletin 256

284

the presence of umbo-lateral bullae,

^vhoils,

septal suture.

On

and by the simpler

up to about

the other hand, the inner whorls

mm

50
diameter match perfectly the microconchiate Irianites
nioermanui (Kruizinga) described above, in all studied features,
i.e. whorl section, coiling, costation and septal suture. Although the
,

apertuie
certain

not

is

preserved,

indicators

scatter

diagrams

relative

width (W:D)

the modified large outer whorls

macrochonchiate,

of

i.e.

female,

shells.

are

The

whorl section (W:H),

width (U:D) and density
of primaries (P) of all specimens at hand from Kemaboe Valley,
in( hiding several immature specimens of intermediate size wliich
could not be assigned to eitlier micro- or macroconch, illustrate this
resemblance. Whorl width is negatively allometric for the innermost
whorls, becoming ajiproximately isometric for microconchs {$) and
matroconchs (9) at 20-25
D (W~30-40% of D) .Thereafter
(Text-figs.
,

21-24)

for

relative umbilical

mm

[>

10-55

(35)

differ strongly

now

width

is

^\hilc

isometry

mm

D], however, the macroconchs here described

from the microchonchiate

strongly positive allometric

is

/.

rnoervmnni

up

to the

shells: whorl
end of the conch,

approximately retained in the microconchs

(ex-

cept for insignificant negative allometry at the end of the body

chamber)

Since whorl height grows

similar in both forms

(not shown)

sembles that for whorl width;

i.e.

more or

less

isometric

and

is

the plot for whorl section re-

there

is

strong positive allometry

mature macroconchs only, W/H changing from 1.0-1.3 to

to 1.4-1.8. Growth of umbilical width is isometric and equal in both
forms from the innermost whorls to about 45 mm diameter
(38-49% of D) but thereafter becomes slightly positive for the
body chamber of the microconchs and possibly very weakly negative for the mature macroconchs. The density of primaries (numin

the

ber/whorl)

is,

again, identical for both forms; the juvenile reduc-

tion of density uj) to 15-25 (30)

mm

diameter

is

followed by an in-

crease from about 20-25 primaries per whorl to 27-35

whorls, altiiough there

is

on the mature

possibly a slower increase in the macro-

The strong resemblance between both forms clearly indimorphic correspondence. Significantly, a closely related
form is also present (Text-fig. 18, 140 nun D) in the Brouwer collection from Wai Miha, Sula Islands (Miner, and Geol. Museum
Delft, No. 14885), togetiier with two microconchiate / moermanni

concirs.

dicates

New Guinea

Text-figs. 20a-d.
a-b,

/.

cf.

/.

126229); c-d,

Cross

Ammonites: Westermann & Getty

section

of Irianites, gray indicating body

285

chamber;

moermanni (Kruizinga) $ (R.G.M. Leiden, st. 126221 and

/. moermanni S
(R.G.M. Leiden, st. 126212 and 126220). xl.

Bulletin 256

286

PMRAOM

New Guinea

50

Ammonites: Westermann & Getty

287

U/W
PHRASM

Bulletin 256

288

Aleasurctnents

New Guinea

Ammonites: Westermann & Getty

289

mm

Scatter

of whorl section, i.r. whorl width against whorl height

(W:H), for Irianitcs, whole collection at hand from Kemaboe Valley (as in
Text-fig. 21). Only a single individual growth line is indicated (body chamber
dotted).

Text-fig. 23.

Docidoceras (Docidoceras) loiigalviun

(Pompeckj)

limatum
(1)

D. (D.) sp. indet.

Stephanoceras (Stephanoceras)
(J.

cf.

de C. Sowerby)

(1)
aff. S.

humphriesianum
(1)

290

Bulletin 256

30

(&)&

/$.^*

5"---

25

PHRASM

20

c3"

'

New Guinea

B. (T.
(?)

}) n. sp.

Ammonites: Westermann

&:

Getty

291

A. $

Cobbnnites

(8)

alF. C.

engleri (Freehold)

[??

Leptosphinctes]

(1)

Irinuites mocrjiunnii
/. cf. /.

(Kruizinga)

(65-75)

moermanni (Kruizinga)

(25-30)

Callovian:
(?)
?

Eucycloccras intermedium

E. sp. indet., $

($})

S\)dii\\

(1)

(1)

Subkossmatia obsciira boehmi,

n. subsp.

(2)

Upper Tithonian-Valanginian:
Blanjordiceras wallichi novaguineyise Gerth

(2)

B. sp. indet.

(4)

Hitnolayites

aff.

H. nederburghi Boehm

(1)

Olcostephaiius ('Rogersites') sp. indet.

(1)

Most of the Lower Bajocian (s.s.) to Middle Callovian ammonite species, as well as most genera here described, are either
new to Indonesia (including West Irian) or have previously been
wrongly

classified.

Stephanoceras

s.s.

Bullatimorphites (Treptoceras) ,
,
Cobbanites are new to the entire south-

Docidoceras

and

(?)

eastern Asia-Australia area; only Irianites appears to be endemic to

eastern Indonesia.

Loiuer Bajocian (Text-fig. 25

(Crick)

may

^)

Fontannesia

(Kruizinga) from the Sula Islands, which,

may
F.

if

clarkei

indeed a Fontannesia,

best be classified as a subspecies of F. clarkei

cf.

aff. F.

be identical with the dubious 'Grammoceras kiliani'

F. clarkei has recently

(Bremer, 1966)

s.l.

Significantly,

been described from

the lower 5. sowerbyi Zone of Turkey in the easternmost Mediterranean occurrence of Fontannesia Anatolia appears also to be the
.

only hitherto

(Pompeckj)
extended to
the

S.

known

occurrence of Docidoceras lojigalvum limatiim

(op.cit.) the distribution of

New

Guinea.

The two

soxcerbyi Zone, probably

zone (of Europe) which

This close

is

its

which

species are

lower

now

tentatively

part, the L. discites

also the age of the

affinity suggests direct

is

good indicators

for

Sub-

Anatolian occurrence.

migration from the eastern Medi-

New Guinea (wherever it may have been located

during Jurassic time)
Surprisingly, unquestionable Docidoceras
has not been reported from the fossiliferous Western Australian
terranean to

Bulletin 256

292

O Docidoceras
A

ncl.

Text-figs.

25a-b.

Known

in

the

Kemaboe

North polar projection of the Recent globe. (25b, see

Normanni

s.

I.

tes

s.

I.

geographic distribution of the Bajocian-Callovian

Ammonitina genera and subgenera occurring

on

s.s.

tephanoc eras

assemblage of the same age

(Akell,

1954)

p.

Valley; plotted
293.)

with the possible ex-

ception of the single microconchiate Docidoceras? (Trilobiticeras

depressum (Whitehouse)

and the Iragments

podus' Arkell (op.

32, figs.

cit., \A.

1957)

descrijjtion

are

aff.

7)

anli-

a,b)

Tlie supposed American occmrences

1956,

of 'Otoites

ol

Fo)itannesin

(Arkell)

based respectively on a single record without

or figure

from

Ijiiiish

(Columbia

(Crickmay,

1930)

New

Guinea Ammonites: W'estermann & Getty

BuUatimorphites
(

incl.

Eucy cloce
Cobbanites

p.

s.

Treptoceros

Subkossma

Text-fig. 25b (see 25a,

293

and

ia
ra

I.

292]

and probably on the 'Fontannesia austroamericana' Jaworski

from Mendoza, Argentina, a single prol:)ably microconchiate

men

(1926)
speci-

of inicertain affinities. Reliable occurrences of Fontannesia

are therefore restricted to Europe, Anatolia, Australia,

and

New

Guinea.

The
closely

1969,

'reticulate'

related
Bull.

distribution
i.

e.

Amer.
of

geographic distribution of Docidoceras and

Otoitidae

was

Paleont.,

Docidoceras

Europe, N.W.

Africa,

recently

vol.
s.s

57.

resembles

Anatolia,

(Westermann,

discussed

No. 255)

and

that

New

The confirmed
of

Fontannesia,

Guinea,

while

Bulletin 256

294

North America

from

records

either

refer

to

southwestern

the

Ahiskan D. (Pseudocidoceras) or to records from Oregon (Lupher,

1941) However, some early Bajocian stephanoceratids recently described by Imlay (1964) from southeastern Alaska under Stephano.

ceras (Skirroceras) could possibly be Docidoceras

yielchianus Imlay,

Unfortiniately,

pi.

15,

fig.

6,

and

S.

s.s.

S.

[cf.

(K.) juhlei, pi. 16,

the complete septal sutures of the

best

(K.)

fig.

6].

material

cannot be investigated because this would mean breaking up

type specimens. However, one undescribed specimen of 5. (K.)
(Imlay's identification) from U.S. Geological
cf. S. nelcliianum
Survey locality 24120 has a suture intermediate between that

and Steplianoceras with a dissimilar pair of 'internal

by an almost straight !!; significantly,
the listed assemblage from this locality (Imlay, 1964, Table 12)
suggests early Bajocian age ['Souninia cf. S. nodata Buckman'
r= S. (Alaskoceras) aff. S. alnskensis Westermann; '5. cf. 5. patella
(Hoffmann)
Waagen' =i Plauaiinnatoceras cf. P. bemieri
Although
this
Sowerby)'].
'Witchellia cf. lacviuscula (J. de C.
of Docidoceras

saddles divided

lateral'

species

is

(subgen.

probably correctly placed in Steplianoceras

therefore

Skirroceras

or

Kumatostephaniis)

this

early

Alaskan

steplianoceratid suggests the evolution of the family from evolute

Docidoceras, D. longalvum group, of the Otoitidae.

Middle Bajocian.
toceras}) etheridgei

(Text-fig. 25 a)

(Gerth)

Steplianoceras (Stemma-

based on an incomplete macroconch

(holotype here refigured)

is apparently known only from New
Guinea and probably distinct from the large but poorly known S.
indicum (Kruizinga) originally described from the Sula Islands.
Probably two of the 'Stemmatoceras' from the Langgeroe area
figured in Visser and Hermes (1962, end. 17, figs. 16 a,b and 19 a-c)
,

belong

to E. etheridgei

itinsae

(figs.

20 a,b)

while others resemble closely

S.

albertense

S.

(Teloceras)

and 'Itinsaites'
hum phriesianum Zone of the
(figs.

22 a,b)

itinsae
McLearn spp., from tlie .S.
North American Cordilleras. Closely affiliated with S. (Teloceras)
itinsae (McLearn) and
palliseri (McLearn) also described from
the northern CorcUlleras, is the Mamapiri
(Langgeroe area)
,

.S'.

specimen figured by Boehm (1913, text-fig. 3, pi. 3, fig. 2). Significantly a similar specimen has recently also been found in the
Malargiie area of Mendoza, Argentina (unpublished)

New Guinea

Of

Ammonites: Westermann & Getty

particular interest

ceras of the

5.

295

the discovery of a typical Stephano-

is

now appears

hinnphriesianurn gioup which

almost

caamanoi

(McLearn) is one of
and a specimen
from north-central Chile has been recorded (Westermann, 1967, p.
Several subgenera of Stcphanoceras which are in part inter68)
cosmopolitan in distribution;

.S".

possibly several northeastern Pacific representatives

mediate

to

Cadomiles and of early Upper Bajocian

the Western Interior United States

(5.

undulatum Burckhardt)
Westermann, 1967).

(Imlay,

1961,

age, occur in

1967), Mexico

Chile and Argentina

(Steinmann,

1881;

(Upper) Bathoni an -(Middle)

the

Kemaboe

The

Callovia7i.

larger part of

Valley ammonites appears to be endemic to the eastern

Indonesia area and cannot be dated accurately, but

some confidence in this interval. Surprisingly, the

is

placed with

collection did

not contain a single Macrocephalites

s.l., probably the most ubiand common group in the whole Indo-West Pacific Callovian. However, the presence of the Middle Callovian eucycloceratids
leaves no doubt that this stage is represented.

cjuitous

Of

particular interest are several species, all tentatively classi-

under Bullatimorphitcs, with both macro-and microconch;

the genus was previously almost unknown from the circumpacific
area, with the single exception of B. sofaniini (Boehm) from the
Sula Islands. All species, however, differ from the typically Europeansouthwestern Asian Bidlatimorphites in the dense and usually
curved ribbing of at least the inner whorls, the more regularly
coiled body chamber, and the relatively narrow Uo lobe. In the
costation they resemble the Middle Callovian Eucycloceratinae
Eucycloceras , Siibkossmatia, and Idiocycloceras which, west of Geelvink Bay and on the Sula Islands appear to be profusely represented but were largely misidentified with Macrocephalites spp.
(Boehm, 1912,1913). Significantly, Bullatimorphites sofaniim
(Boehm) and ? B. (Treptoceras) sp. have recently also been found
in the M. ^nacrocephalns Zone of Neuquen in the southern Andes
fied

(unpublished)

The most important

aff.

species for correlation

B. iihligi (Popovici-Hatzeg)

River

(Gerth, 1927, p. 226:

refigured here for the

first

which

is

B. (Treptoceras)

Wairori
hidlatum d'Orb.";

also occurs at the

"Sphaeroceras

cf.

time, Text-fig. 10)

B. iihligi [" B. sue-

Bi'LLETiN 256

296

(Roemer)

viciis

" aiict.]

and

closely attiliated B. crimaciense

["A. microstoma" , Qiienstedt 1886, non d'Orbigny]

Enay

occur in the

lacies of the Upper Bathonian and Lower Callovian of

Eiuope (Germany, Romania) and "B. sucviciis" was recorded
from the Upper Bathonian of the Pamirs (fide Arkell, 1956, p. 403)

shaly

The

microstoma

single B. (T.) aff. B.

(d'Orbigny)

Upper Bathonian European species.

The more abundant new sjiecies which

Middle

resembles the

to

are only

tentatively

placed in BuUatimorphites, however, appear intermediate to the

Middle Callovian Eucycloceratinae which they resemble except for

the much more depressed whorl section. Only the most common
form, B.

costidensus,

is

named although

at least

three different

forms arc present.

Morphological affinity with tlie Eucycloceratinae suggests that

most of the tentative BuUatimorphites may be phylogenetic intermediates between Tulitidae and Eucycloceratinae, i.e. that the
latter subfamily evolved from Tiditidae rather than from Sphaeroceratinae as assumed previously, a derivation retained only for the
Macroccj^halitinae (Macrocephalites s.L). This supposition would
also imj^Iy that this Indonesian BuUatimorphites ? costidensus group
is

among

tidae,

the last

not representing the

if

and possibly

of

Middle Callovian

latest part of the Tuli-

age.

Such age would also

explain the absence in the collection of Macrocephalites

woidd, of course, a Bathonian age)

At least in part associated with the B.

s.L

(as

costidensus group

dimorphic Irianites
moermatnii (Kruizinga) a species and genus probably endemic to
eastern Indonesia-New Guinea. /. moermanni is also associated with
Bositra buchi (Roemer) whicli has a known vertical range from the
(same concretion)

the

is

almost

certainly

Toarcian
occurs

the Oxfordian. Another unnamed species probably

undated beds of Babar Island, northeast of Timor

to

in

(Boelim, 1908; Jaworski, 1933). Irianites

is

somewhat

tentatively

placed in the Perispliinctaceae and possibly closest affiliated with

'Psc'udopcrisjjliinc tinae'.

Callovian.
fjy'

Boehm

Avitli

The. New

(1913, pi.

Subkossuialia

species Suhkossmatia

Guinean 'Macrocephalites keeuwensis

2), identified by Spath (1924, p. 212)

Kutch
(losely resembles the
'heta-gavrma'
5, fig.

opis

(Waagcn)

and the

closely

related

S.

New Guinea Ammonites: Westermann

obscura Spath and

The

n.subsp.
is

is

here given the proper

name

Getty

Sc

5.

297

obscura boehmi,

Sula Islands species Eycycloceras intermedium Spath

probably also present. Both forms appear intimately related and

the genera are probably best iniited.

Eucycloceras and Subhossmatia are typical of the greater Indian

Ocean

area,

although Eucycloceras

Africa (Arkell, 1957,

p.

also

is

reported from North

291). In the famous Kutch section, both

to the upper Rcineckeia rehmanni Zone

which is now placed in the lower but not
basal part of the Middle Callovian, equivalent to the higher parts of
the northwest European A', jason Zone, where Macrocephaliles s.lhas become extinct (Callomon, 1955)

genera are restricted

(Spath, 1933, p. 676)

The

perisphinctid

Cobbanites

(?)

aff.

C. engleri

(Frebold) re-

from Alberta and to a lesser

degree, C. talkeetuanus Imlay from the Upper Bathonian and
Lower Callovian of southern Alaska and Montana.
In summary, the studied ammonites from the Kemaboe Valley
clearly indicate the Lower and the Middle Bajocian (exclusive of
Aalenian) and the lower Middle Callovian, while Middle Bathon-

sembles this basal Callovian species

(probably Upper Bathonian)

ian-Lower Callovian
suggested; however,

it

Callovian are present.

is

possible that only Bajocian

The

in the

Pacific in the

(Middle Bathonian

tentatively

and Middle

faunal affinities are strongly Mediter-

ranean in the Lower Bajocian

somewhat eastern

is

to)

(S.

sowerbyi Zone), cosmopolitan,

Middle Bajocian and mainly 'Indie'

Middle Callovian, with weak en-

and strong endemism in the later faunas.

Bajocian
to Callovian ammonite genera now
all
Considering
Lower Bajocian is repre(Table
known from New Guinea
1)
Pseuclotoites; the Midand
sented by Docicloceras s.s., Fontannesia
dle Bajocian by diverse Stepliauoceras s.l. and Chondroceras;
doubtful Upper Bajocian by Chondroceras }(Praetulites) and pos-

demism

in the Bajocian

by Cadomites ? [recorded only; but not 'Baculatoceras' in

and Hermes, 1962, which is Blanfordiceras]; possible Bathonian by Tulites} [recorded only] and the Cadomites} mentioned
above and Bullatimorphites of the B. uhligi and (?) B. microstoyna groups; Lower Callovian by diverse Macrocephalites s.l.;
(lower) Middle Callovian by Subhossmatia, Idiocycloceras, and

sibly

Visser

Eucycloceras

(?)

Bulletin 256

298

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J.

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Enay, R.
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Etheridge, R.
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Ammonites: Westermann & Getty

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Frehold, H.
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The Jurassic Fernie (ruuj) in the C.aiitidian Rocky Motiiilains

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Gerth, H.
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^Eiji 71CUCS J'orkommcn dcr hatliyalni Ccphalopodcn-Facirs dcs

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Ammonitrn

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Jurassic (Bathonian or early Callovian) ammonites from Alaska and

Montana. U.S. Geol. Sur., Prof. Paper 374-C, pp. 1-32, pis. 1-8.
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Jaworski, E.
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und Dactylioceraten des Lias von

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251-334, pis. 7-11.

and Minton, R. P.
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Jefferies, R. P. S.,
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Palaeontology, vol.

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Kuhn, O.
1939.

Die Ammonilen des frankischen Calloviums. Nova Acta Leopoldina,

n.f., Bd. 6, pp. 451-532, pis. 48-57.

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voL 54, le6ed., pp. 13-85, pis. 1-14.
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Lupher, R.
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Jurassic stratigraphy of central Oregon. Geol. Soc.
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Bull. 52,

Martin, K.
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Mascke, E.
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Paldozoischc, mesozoische
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und kdnozoischc sedimente aus dcm

Samml. Geol. Reichsmus. Leiden, Ser. 1,

Die Stephanoceras Verwandten in den Coronatenscliichten von

Norddeutschland. Gottingen, 38 pp.

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300

Neumayr, M.
Die

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C('j>li(il()j)()(lt >i-l-(iu)ia

dcr OoUtlic

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lialiii

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Die Ammonites

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1875.

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19

Ammoniliden.

Orbigny, A.

d'
1842-51. Palcontologie

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fratigaise;

Ce[>lial()l)odei

I.

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Pompeckj,

J. F.

1897.

Paldonlologische und straligrapJtisciie Notizen

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Quenstedt, F. A.
1883-8. Die Ammonitcn dcs Schivdbischcn Jura. Three

Analolien.

aiis

29-31.

pis.

vols.,

text

and

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Renz, C.
lieitrdge

1925.

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scliAveiz.

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F.
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(Monte Erice)

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Die Fauna der Aspiduides-Srhiclilen

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de C. Sowerby

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Mineial Condiology of Cual lUitain,

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See Bull, .\incr.

I'alconi., \ol. 48, 1965, pp. 429, 430.

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Mem.

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1927-1933. Revision of the Jurassic Ceplinlopods of Kaclili (Cutch). Metn.

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//;

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I. a
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Mayaitinae

XXII

as

Int. Geol.

PLATES

304

Bi'LLETiN 256

Expi.-WATioN OF Plate 48
Xatiiral size

if

not otherwise indicated

Page

Figure

Fontannesia

aff.

F. clarkei

(Crick) [? subsp.kiliani

(Kruizinga)]

238

complete specimen, with 2/3 whorl body chamber, last two

septa not clearly approximated (R.G.M. Leiden, st. 126181).
Incomplete specimen with 1/2 whorl body chamber (R.G.M. Leiden,
St. 126182).

la-c. -Almost
2.

3a-b. Slightly

damaged specimen with

chamber of 2/3
(R.G.M. Leiden,

whorl, last
st. 126183).

partly preserved aperture, body

septa not markedly approximated.

probably immature specimen, body

phragmocone (R.G.M. Leiden, st. 126184).

4a-b. Small,

chamber

and

crushed

Bull. Amer. Paleont., Vol. 57

Plate 48

Plate 49

Bull. Amer. Paleont., Vol. 57

lb
la

m
2a

/*^^^r

' 3b

New Guinea

Ammonites: Westermann & Getty

305

Explanation of Plate 49
Natural

size if not

otherwise indicated

Figure

Page

Fontannesia

aff. F.

clarkei (Crick) [V subsp. kiliani

(Kruizinga)]
la-c.

Wholly

238

specimen, last
preserved nucleus (Text-fig. 4)
septate

septa

approximated,

(R.G.M. Leiden,

st.

with
126185).

\vell-

grown specimen, compressed, and weakly ornate,

1/2 whorl incomplete body chamber (R.G.M. Leiden, st.

2a-b. Probably not fully

with
126186).

3a-b. Incomplete

specimen

with

1/2

whorl

body chamber,

partly crushed, with approximated last sutures;

(R.G.M. Leiden, st. 126187).
4a-b.

b,

phragmocone

nucleus of same

Immature (?), probably almost complete specimen with 2/3 whorl

body chamber (R.G.M. Leiden, st. 126188).

Bulletin 256

306

Explanation of Plate
Natural size

if

50

not otherwise indicated

Page

Figure

lad. Docidoceras (Docidoceras) longalvum (Vacek)

limatum (Pompeckj) 9
Complete

cf.

specimen with aperture; note umbolateral

porous filling (R.G.M. Leiden, st. 126189).

2.

Docidoceras (Docidoceras) sp. indet. 9

bodj- chamber
and remnants
(R.G.M. Leiden, st. 126190).

244

subsp.

groove

with

246

(?)

Incomplete

of

penultimate

whorl

Bull. Amer. Paleont., Vol. 57

Plate 50

/>-

^r

^-f^

la

lb

id

Bull. Amer. Paleont., Vol. 57

Plate

New Guinea

Ammonites: Westermann & Getty

Explanation of Plate
Natural size

if

51

not otherwise indicated

Page

Figure
la,b.

307

Stephanoceras (Stephanoceras)
(J. de C. Sowerby) $

aff.

S.

humphriesianum
247

Fragment of phragmocone with damaged beginning of body chamber.

(R.G.M. Leiden, st. 126191).
2a,b.

3a,b.

Stephanoceras (Stemmatoceras?) etheridgei (Gerth) ?, $ ?

252
Probably almost complete specimen with 2/3 whorl body chamber
(R.G.M. Leiden, st. 126192).
Cobbanites (?) aff. C. engleri (Frebold)
Almost complete specimen with 7/8 whorl body chamber.
Leiden,

st.

126193).

269
(R.G.M.

Bulletin 256

308

Explanation of Plate
Natural size

if

52

not otherwise indicated

Page

Figure

Bullatimorphites
la,b.

n. sp.

St.

254

Somewhat crushed phragmocone with beginning

(R.G.M. Leiden,

of

body chamber

126194).

phragmocone with 1/4 whor! incomplete body chamber

(R.G.M. Leiden, st. 126195).
Fragment of large bodv chamber \vith remnant of penultimate
whorl (R.G.M. Leiden, st. 126196).

2a-c. Undistorted
3.

?,

Plate 52

Bull. Amer. Paleont., Vol. 57

>feMLCf.

la

lb

^i^^^s^"

h^is^p'

Bull. Amer. Paleont., Vol. 57

r^.

Plate 53

New Guinea

Ammonites: Westermann & Getty

Explanation of Plate
Natural size

if

309

53

not otherwise indicated

Page

Figure

la,b. Bullatimorphites (Treptoceras) aff. B. uhliqi (Popovici-Hatzeg) $ 259

Specimen with aperture, beginning of boiiy chamber damaged, with
ventral costae anomaly (R.G.M. Leiden, st. 126197).
2a-c.

Bullatimorphites
(d'Orbigny) 6

(?)

Treptoceras?)

aff.

B.

microstoma
257

Complete, slightly damaged specimen with exposed part of penultimate whorr( R.G.M. Leiden, st. 126199).
3a-b. Bullatimorphites (Treptoceras) aff. B. uhligi (Popovici-Hatzeg)
Incomplete whorl of contracting body chamber with end of phragmocone (R.G.M. Leiden, st. 126198)."

259

Bulletin 256

310

Explanation of Plate 54
Natural size

if

not otherwise indicated

Page

Figure

(Treptoceras?) costidensus Westermann

and Getty, n. sp. {S ?)
260
End of bodv chamber with probably incomplete aperture (R.G.M.

Bullatimorphites

1.

Leiden,
2a, b.

3a-e.

4a-c.

st.

'l26200).

Complete but somewhat deformed body chamber with

aperture (R.G.M. Leiden, st. 126201).

incomplete

Holotype, almost complete body chamber of over one whorl and part
of phragmocone; b,c, lateral and apertural views of last septum; e,
part of body chamber removed (R.G.M. Leiden, st. 126203).

Almost complete body chamber of one whorl

126202).

(R.G.M. Leiden,

st.

Bull. Amer. Paleont., Vol. 57

Plate 54

Bull. Amer. Paleont., Vol. 57

Plate 55

^^tfi^fi

Ic

lb

^>^I!S5S5^

r
Y

'

rj
'vv

2b

New Guinea

Ammonites: Westermann

Explanation of Plate
Natural size

if

8c

Getty

311

55

not otherwise indicated

Page

Figure

Bullatimorphites

(Treptoceras?),

n. sp.

la-d.

Complete body chamber with aperture (lappets) and phragmocone,

right side damaged and distorted (R.G.M. Leiden, st. 126204).

2a-b.

Almost complete specimen (R.G.M. Leiden, st. 126205).

Almost complete body chamber with parts of crushed phragmocone,
secondaries exceptionally dense (R.G.M. Leiden, st. 126206).

3.

4a-b.

impression
Ir'tanitcs
fragment with
of
cf.
moermanni
(Kruizinga) and J5.? (T.?), n.sp. A.; b, latter species after removal
of former, composed of body chamber fragment and plasticine mold
of inner whorls (R.G.M. Leiden, st. 126207).

Rock

262

Bulletin 256

112

Explanation of Plate
Natural size

if

56

not otherwise indicated

Page

Figure

lab. ?Eucycloceras
Soniewiiat

sp. indet.,

distorted

c5

specimen

chamber (R.G.M. Leiden,

st.

266

with approximately
126208).

1/2

whorl

body

Eucycloceras intermedium Spath, $ ?

265
Incomplete 1/2 wluirl hod\ chamber and part of penultimate whorl
(K.CJ.M. Leiden, st. 126209).

2a-b. (?)

3,4.

Subkossmatia obscura Spath boehmi Westermann and Getty,

n.subsp
almost complete body chamber and part of penultimate whorl
(R.G.M. Leiden, st. 126210). - 4a, b, incomplete body chamber \vith
remnants of inner whorls (R.G.M. Leiden, st. 126211).

3a, b,

266

Plate 56

Bull. Amer. Paleont., Vol. 57

*-/S

la

<-^

3a

Bull. Amer. Paleont., Vol. 57

Plate

57

New Guinea

Ammonites: Westermann & Getty

Explanation of Plate
Natural size

if

313

57

not otherwise indicated

r'age

Figure

moermanni (Kruizinga)

Irianites

la-d. Nearly perfect specimen with incomplete aperture (base of lappets),

nucleus damaged; b,c, part of aperture removed to show whorl
section
2.

(R.G.M. Leiden,

Almost

damaged
Leiden,
3a, b.
4.

5a-c.

st. 126212).
(approximated sutures)
with
adult specimen
almost 3/+ whorl body chamber (R.G.M.
incomplete,
126213).

complete
st.

Probably complete specimen except for aperture, with 3/4 whorl

body chamber (R.G.M. Leiden, st. 126214).
Complete specimen with base of lappets, approximated sutures and
2/3 whorl body chamber (R.G.M. Leiden, st. 126215).
Probably complete specimen except for aperture, with 3/4 whorl
bodv chamber and most of the test preserved (R.G.M. Leiden, st.
126216).

274

Bllletin 256

314

Explanation of Plate
Natural size

if

not otherwise indicated

Page

Figure
Irianites
la-c.

58

moermanni (Kruizinga) $

Complete body chamber of 7/8 whorl (? base of lappets) with

damaged phragmocone; c, Bositra buchi (F.A. Roemer) I'Posidonia'
auct.] at aperture of same specimen, X 2. (R.G.M. Leiden, st.
126217).

2a, b. Irianitrs sp. ju\-. with 2/3

Leiden,
3a, b.

whorl incomplete bodv chamber (R.G.M.

126218).

Bodv chamber fragment with

Leiden,

4a, b.

st.

st.

lappets of large inflated form (R.G.M.

126219).

Almost complete specimen with 3/4 whorl body chamber and approximated sutures; large inflated form (R.G.M. Leiden, st. 126220).

274

Bull. Amer. Paleont., Vol. 57

Plate 58

Bull. Amer. Paleont.. Vol. 57

Plate 59

Id

New

Guinea Ammonites: Westermann & Getty

Explanation of Plate
Natural size

if

59

not otherwise indicated

Page

Figure
Irianites cf.
la-d. Slightly

I.

moermanni (Kruizinga)

damaged

preserved
2.

315

Leiden, st.
Cross-section of fragment of
(R.G.M. Leiden, st. 126222).

largest

281

almost complete specimen

bodv chamber; d, nucleus of same,
126221).
but

with

phragmocone

1-5

in

whorl
(R.G.M.

3/4

collection

Bulletin 256

316

Explanation of Plate 60
Natural size

if

not otherwise indicated

Page

Figure
Irianites cf.
la-c.
2.

I.

moermanni (Kruizinga)

Incomplete specimen with 1/2 whorl preserved body chamber and

parts of phragmocone (R.G.M. I.eiden, st. 126223).
Ahnost full whorl (7/8) of incomplete body chamber, partly crushed,
and parts of phragmocone (R.G.M. Leiden, st. 126224).

281

Plate 60

Bull. Amer. Paleont., Vol. 57

la

^Uv-

Bull. Amer. Paleont.. Vol. 57

Plate 61

New Guinea

Ammonites: Westermann & Getty

Explanation of Plate
Natural

size

if

317

61

not otherwise indicated

Page

Figure
I.
moermanni (Kruizinga) 9
Damaged phragmocone with 1/4 whorl fragment

281

Irianites cf.
la-c.

2.

3a-b.

(R.G.M. Leiden, St. 126225).

Large body chamber fragment
?

(R.G.M. Leiden,

of body
st.

chamber

126226).

Juvenile 5, with full one whorl body chamber which is probably

complete on left side; b, X 0.9 (R.G.M. Leiden, st. 126227).

Bulletin 256

318

Explanation of Plate 62
Natural

size

if

not otherwise indicated

Page

Figure
Irianites cf.
la-c.

2a-f.

I.

moermanni (Kruizinga)

Immature specimen -with one

(R.G.M. Leiden, st. 126228).

Immature specimen with one

d,

f,

full

juv

whorl incomplete body chamber;

(R.G.M. Leiden, st. 126229).

full

inner whorls of same, x2

whorl incomplete body chamber

281

Bull. Amer. Paleont., Vol. 57

Plate 62

INDEX
Note. Light face type refers to page numbers. Bold face type
fers to plate

re-

number.
269
Chondroceras
237, 256, 264
Chos-Malal, Argentina
255
clarkei, Fontannesia
aff
48,49
238
Choffatia

adicra,

246

Sonninia

246, 249, 270

Alaska
alaskensis, Sonninia
Alaskoceras
Alberta

294
294
248, 252, 270

albertense,

294
256, 296

Stephanoceras

Ammonites
amplectens,

Otoites

....

Argentina

246
246, 291
292
293-295

arietitiforme,

48,49

238
269
237, 244, 246
271, 273

Asphinctites

Asthenoceras
Australia
austroamericana,

Fontannesia

293

B
238, 240

Baculatoceras
Barbar Island
baumbergeri,

Grammoceras
benneri,

Planammatoceras
Blanfordiceras

boehmi,
Subkossmatia

56

Bositra
braikenridgii,

Stephanoceras
brodiaei,

Stephanoceras
British Columbia
buchi, Bositra

Bullatimorphites
bullatum,
Bullatimorphites
bullatum,
Sphaeroceras

caamanoi,
Stephanoceras

Cadomites
Catacoeloceras

....

248,295
237, 249, 264,

295
271

charlottensis,

Stephanoceras

Bullatimorphites

54

249
295

260, 263

crassispinata,

Sonninia
crickmayi,

Kanastephanus
242
273
239
241, 291

Harpoceras

Chile

Fontannesia
Cobbanites
Coeloceras
Coeloderoceras
cosstidensus,

Eudmetoceras
Anatolia
antipodus,

clarkei kiliani,

246

252

crimaciensis,

Bullatimorphites
Crussol

daubenyi, Cadomites
Stephanoceras
Defonticeras

258, 296

257

237
237
256

IXDKX

Grammoceras

240

grammoceroides,
Fontannesia

longalvum,
Docidoceras

50

241
264
273

Grayiceras
Gulielmiceras

244

M
mackenzii,

Kanastephanus

H
243

Harpoceras
Hildesheim

254, 259

291

Himalayites

humphriesianum,
Stephanoceras

aff.

51

247, 295

Idiocycloceras

Imskim Formation

indicum,
Stephanoceras

Mamapiri
Mendoza
Mexico
microstoma.

Ammonites
53 256, 263, 296
Bullatimorphites 53
254, 263
Misol
243
moermanni,

intermedium, ?
Eucvcloceras

56

Irianites
itinsae,
c^

Itinsaites

241

Irianites

265
238,251,271

Irianites

294
237,290,294

271,274
274, 277

Coeloceras
Coeloderoceras

251,286

Inferior Oolite

Stephanoceras

264
236

252
252
238, 295
294
266, 294
293, 294
295

Stephanoceras
Macrocephalites
Malargue

cf.

Normannites
Moluccas
Montana

57,58 238, 271, 274,

277
59-62
280
237, 274
243
270

mutabile,

Stephanoceras

248

N
Jakarti Fault Zone
juhlei,

Stephanoceras

235
294

n.

sp.

Bulla-

timorphites
n.

K
keeuwensis,
Macrocephalites
Kemaboe Valley

238,265,266
234

Kembelangan
Formation

235
kiliani,
Grammoceras
238, 242
294
Kumatostephanus
Kutch
237,257,296.
297

L
Labyrinthoceras

238,264,268

sp.

55

262

52

254

Bulla-

timorphites ?
nederburghi,
Himalayites
nelchianum,
Stephanoceras

291

294
295

Neuquen
Newmarracarra
Limestone
Normannites
North Paniai Fault
Zone

241

271,273
237
264

Nothocephalites
novaguinense,
Blanfordiceras

238,240

laeviuscula,

Sonninia
Witchellia

294
294

laurenti,

Bullatimorphites

Lenggeroe
Leptosphintes
limatum, Coeloceras
Docidoceras

O
obscura,

258
235,237,294
270
245
245

Subkossmatia
Olcostephanus
opis, Stephanoceras
Subkossmatia

Oregon

320

266,268
.

291
268
268
294

Index
P

suevicus,

palliseri,

294
296
257
273
294

Stephanoceras
Pamirs
Paris Basin
Parkinsonia
patella, Sonninia
perfectum,
Docidoceras

245, 246

Planammatoceras

...

Pleydellia

Poland
Polyplectites
Praetulites
Procerites

Pseudolioceras
Pseudoperisphinctes
Pseudotoites

294
240
269
264
237, 256
269
239
269
237, 245

Bullatimorphites
Sula Islands

T
Taliabu
talkeetnanus,
Stephanoceras

248
237

Islands

Quenstedtoceras

Teloceras

248, 283, 286,

Tipoema Formation
Torricellites
transiens,

Docidoceras
trapanicum,
Docidoceras
Treptoceras

Zone
Rock Creek Member
Roemberpon Island
Rogersites

Romania
Rugiferites

246
256
238
269

Siemiradzkia
skidegatense,
Stephanoceras
Skirroceras
sofanum,
Bullatimorphites

252
248, 294

255 258,261,
295

indet,

50
56

294
236
273
246
246
237,255
292

Stephanoceras

249
256
244,291

Tulites

U
297
248
237
291
259, 296
256

San Vigilio
Schwandorfia
Sepik River

triptoleumus,

Reineckeia rehmanni

Docidoceras
? Eurycloceras

269,270
277

Tangi

Turkey

sp.

275,277

Trilobiticeras

Queen Charlotte

254,259
237,240,255,
265, 276, 277, 284

246
266

Stemmatoceras

248, 249, 283,

Subkossmatia
suevicum,
Sphaeroceras

238, 264, 266

294

259

uhligi, Ballatimorphites aff

umbilicum,
Stephanoceras
undulatum,
Stephanoceras

Vermisphinctes

53 237,254,259,
295

248

295

3 2044 072 271

Date Due

711