Palynology of the 

Upper Campanian (Cretaceous) 

Judith River Formation, 

North-Central Montana 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 

Palynology of the 

Upper Campanian (Cretaceous) 

Judith River Formation, 

North-Central Montana 

By BERNADINE D. TSCHUDY 

GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 

Identification of fossil pollen and 

spores in rocks from the type area 

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON 1973 
UNITED STATES DEPARTMENT OF THE INTERIOR 

ROGERS C. B. MORTON, Secretary 

GEOLOGICAL SURVEY 

V. E. McKelvey, Director 

Library of Congress catalog-card No. 72-600366 

For sale by the Superintendent of Documents, U.S. Government Printing Office, Washington, D'.C. 20402 

Price: Paper cover-,1.25, domestic postpaid; '1.00, GPO Bookstore. Stock No. 2401-00291 
CONTENTS 

Albstract -----------------------------------------

Introduction -------------------------------------- The Judith River Formation -------------------

Sample localities -----------------------------

Processing of samples and photography ---------

Procedure of study----------------------------

Depository of types and other illustrated 

specimens ---------------------------------- Alcknowledgments ____________________________ _ 

Pollen-spore zone·s _______________________________ _ 

Systematics --------------------------------------

Triletes __ ------------------------------------

Genus Cyathidites Couper, 1953 ------------

Genus Deltoidospora Miner, 1935 emend. 

Potonie, 1956 _____________________ _: ____ _ 

Genus Stereisporites Pflug iri Thomson and 

Pflug, 1953 ----------------------------- Genus Leptolepidites Couper, 1953 emend. 

Potonie, 1966 --------------------------- Genus Baculatisporites Pflug and Thomson 

in Thomson and Pflug, 1953 --------------

.Genus Osmundacidites Couper,l953 -~------­

Genus Kuylisporites Potonie, 1956 -----,-----

Genus !schyosporites Balme, 1957 -----------

Genus· Cicatricosisporites Potonie and 

Gelletich, 1933 emend. Potonie, 1966 _____ _ 

Genus Ghoshispora Srivastava, 1966 sensu 

Leffingwell, 1971 ------------------------ Genus Foraminisporis Krutzsch, 1959a _____ _ 

Genus Polycingulatisporites Simoncsics and 

Kedves, 1961 emend. Playford and 

Dettmann, 1965 -------------------------

Genus Trisolissporites n. gen --------------

Genus Gleicheniidites Ross, 1949 emend. 

Skarby, 1964 ---------------------------

Genus Camarozonosporites Pant ex Potonie, 

1956 emend. Klaus, 1960 ----------------

Monoletes ------------------------------------

Genus Laevigatosporites Ibrahim, 1933 

emend. Schopf, Wilson, and Bentall, 1944 __ 

Genus Umbosporites Newman, 1965 ________ _ 

Genus Microfoveolatosporis Krutzsch, 1959a _ 

Zonates -------------------------------------- Genus Seduc,tisporites Chlonova, 1961 ______ _ 

Genus Zonalapollenites Pflug in Thomson 

and Pflug, 1953 -------------------------

Bisaccates ----------------------------------- Genus Alisporites Daugherty, 1941 ________ _ 

Genus Abietineaepollenites Potonie, 1951 ex 

Delcourt and Sprumont, 1955 -------------

Page 

1 

1 

1 

1 

1 

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3 

3 

6 

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6 

6 

7 

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7 

8 

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14 

14 

Systematics-Continued 

Bisaccates-Continued 

Genus Cedripites Wodehouse, 1933 ---------

Genus Pityosporites Seward, 1914 emend. 

Manum, 1960 ---------------------------

Genus Podocarpidites Cookson ex Couper, 

1953. emend. Potonie, 1958 ---------------

Genus Pristinuspollenites n. gen. -----------

Genus Rugubivesiculites Pierce, 1961 --------

Genus Vitreisporites Leschik, 1955 sensu 

Potonie, 1958 --------------------------- Miscellany __________________________________ _ 

Genus Taxodiaceaepollenites Kremp, 1949 ex 

Potonie, 1958 --------------------------- Genus Sequoiapollenites Thiergart, 1937 ____ _ 

Genus Classopollis Pflug, 1953 emend. Pocock 

and Jansonius, 1961 -------------------- Genus Equisetosporites Daugherty, 1941 

emend. Singh, 1964 ---------------------

Monosulcates --------------------------------- Genus Arecipites Wodehouse, 1933 ----------

Genus Liliacidites Couper, 1953 -------------

Genus Ovoidites Potonie ex Krutzsch, 1959a 

emend. Potonie, 1966 --------------------

~ Tricolpates ----------------------------------

Genus Aesculiidites Elsik, 1968b -----------

Genus Grewipollenites Srivastava, 1969d ___ _ 

Genus Pulcheripollenites Srivastava, 1969d __ 

Genus Tricolpites Cookson ex Couper, 1953 

emend. Potonie, 1960 --------------------

Genus Tricolpopollenites· Pflug and Thomson 

in Thomson and Pflug, 1953 --,------------

Genus Cranwellia Srivastava, 1966 ---------

Genus Fibulapollis Chlonova, 1961 ----------

Genus Aquilapollenites Rouse, 1957 ---------

Genus Mancicorpus Mtchedlishvili, pro parte, 

Krutzsch, 1970 -------------------------

Polycolpates ---------------------------------- Genus Sindorapollis Krutzsch, 1969 ________ _ 

Tricolporates ---------------------------------

Genus Cupanieidites Cookson and Pike, 1954 _ 

Genus Tricolporites Cookson, 1947 ----------

Genus Montanapollis Tschudy, 1971 --------

Genus Siberiapollis Tschudy, 1971 ----------

Triporates ----------------------------------- Genus Triatriopollenites Pflug in Thomson 

and Pflug, 1953 ------------------------

Genus Subtriporopollenites Pflug and Thomson in Thomson and Pflug, 1953 ---------

Genus Proteacidites Cookson ex Couper, 1953_ 

m 

Page 

15 

15 

16 

16 

18 

18 

19 

19 

20 

20 

20 

20 

20 

20 

21 

21 

21 

21 

22 

22 

24 

26 

26 

26 

29 

29 

29 

30 

30 

30 

31 

31 

31 

31 

31 

32 
IV CONTENTS 

Page Page 

Systematics-Continued Systematics-Continued 

Polyporates ---------------------------------- Genus Periporopollenites Pflug and Thomson 

32 Tetrads --------------------------------------

Genus Inaperturotetradites van Hoeken32 

emend. Krutzsch, 1966 ----------------- 32 Klinkenberg, 1964 ---------------------- 32 

Genus Erdtmanipollis Krutzsch, 1962 ______ _ 32 Genus Droseridites Cookson, 194 7 ex Potonie, 

Dyads _______________ ------------------------ 32 1960 ----------------------------------- 33 

33 

39 

PLATE 

FIGURE 

Genus Dyadonapites Erdtman, 1947 ex 

Tschudy (this report) -------------------

References cited ---------------------------------- 32 Index ___________________________________________ _ 

1. 

2. 

3. 

4. 

5. 

6. 

7. 

8. 

9. 

10. 

11. 

1. 

2. 

ILLUSTRATIONS 

[Plates follow index] 

Cyathidites, Deltoidospora, Stereisporites, Leptolepidites, Baculatisporites, Osmundacidites, Kuylisporites, lschyosporites, Cicatricosisporites, Ghoshispora. 

Foraminisporis, Polycingulatisporites, Trisolissporites, and Dulhuntyispora. 

Gleicheniidites, C amarozonosporites, Laevig atospo rites, U mbosporites, Micro f oveolatosporis, and S eductisporites. 

S eductisporites. 

Zonalapollenites, Alisporites, Abietineaepollenites, Cedripites, and Pityosporites. 

Pityosporites. 

Podocarpidites, Pristinuspollenites, and Rugubivesiculites. 

Vitreisporites, Taxodiaceaepollenites, Sequoiapollenites, Classopollis, Equisetosporites, Arecipites, Liliacidites, 

Ovoidites, Aesculiidites Gre1vipollenites, Pulcheripollenites, and Tricolpites.· 

Tricolpopollenites, Cranwellia, Fibulapollis, and Aquilapollenites. 

Aquilapollenites, Mancicorpus, Sindorapollis, Cupanieidites, and Tricolporites. 

Montanapollis, Siberiapollis, Triatriopollenites, Subtriporopollenites, Proteacidites, Periporopollenites, Erdtmanipollis, Dyadonapites, lnaperturotetradites, and Droseridites .. 

Page 

Composite section showing stratigraphic position of pollen-spore collections from Judith River Formation 2 

Index map showing sample localities ----------- --------------------------------------------------- 3 

TABLES 

Page 

TABLE 1. Stratigraphic distribution of pollen and spores _ --------------------------------------------------- 4 

2. Pollen-spore zones, based on selected taxa from table 1 -------------------------------------------- 6 

3. Abundance ratios between two pairs of pollen s.pecies ---------------------------------------------- 7 
PALYNOLOGY OF THE UPPER CAMPANIAN (CRETACEOUS) 

JUDITH RIVER FORMATION, NORTH-CENTRAL MONTANA 

By BERNADINE D. TscHUDY 

ABSTRACT 

A large assemblage of well-preserved fossil pollen and 

spores was found to be incorporated in 13 samples from 

upper Campanian rocks from the type area of the Judith 

River Formation near the Missouri River in north-central 

Montana. Three fairly distinct zones and one transition zone, 

on the basis of the extracted palynomorphs, were recognized 

within the composite section studied. Most of the microfossils 

morphologically worthy of taxonomic designations were assignable to previously known species and (or) genera; but 

two new genera, 10 new species, and one new variety were 

named and described to accommodate new taxa. The palynomorph assemblages recovered from the 13 samples of the 

section are listed in stratigraphic s,equence, and the taxa 

are illustrated. 

INTRODUCTION 

Preliminary palynological work on 18 rock samples furnished by J. R. Gill and L. G. Schultz from 

a composite section from the type areas of the 

Claggett Shale and Judith River Formation in 

north-central Montana disclosed a large assemblage 

of well-preserved palynomorphs. This report incorporates the results of extensive work on 13 samples from five collection localities from the Judith 

River Formation part of the composite section. The 

purpose of the study was the identification of the 

taxa that make up the Judith River Formation 

pollen and spore assemblage. 

THE JUDITH RIVER FORMATION 

Hayden (1859, p. 123-124) dramatically described 

the area near the mouth of the Judith River which 

he called the "Bad Lands of the Judith," and he 

later (1871, p. 97) formally designated the strata 

of the Judith basin as the "Judith Group." Stanton 

and Hatcher (1905, p. 11-66) summarized the history of early exploration in the region along the 

Missouri River near the mouth of the Judith River 

and described the lithologic and paleontologic features of the Judith River Formation. On the bases 

of early reports by Hayden and of their own observations, Stanton and Hatcher (1905, p. 32-33) defined "the original area of the Judith River formation" as lying between the mouths of the Judith 

River on the west and "Little Rocky Mountain 

Creek" (Rock Creek) on the east and as extending 

both north and south of the Missouri River for about 

20 miles. 

The composite section (fig. 1) of the Judith River 

Formation in areas from which J. R. Gill and L. G. 

Schultz collected samples is about 775 feet thick. 

The lower marine part of the section was included 

in the Claggett Formation by Stanton and Hatcher 

( 1905, p. 32, 35-36) . 

SAMPLE LOCALITIES 

The 13 samples used in this study are the pollenspore collections which J. R. Gill and L. G. Schultz 

made within their composite section of the Judith 

River Formation in its type area. Eleven of the 

samples were collected near the west end of the 

area, at sites near the Judith River and Dog Creek 

south of the Missouri River and near Birch Creek 

north of the Missouri River. The other two samples 

were collected toward the east end of the area from 

sites slightly north of the Missouri River. The geographic localities from which the samples were collected are shown on the index map (fig. 2). Section, 

township, and range localities of the individual sEctions and the relative positions of the samples within 

the composite stratigraphic section are shown in 

figure 1. 

PROCESSING OF SAMPLES AND PHOTOGRAPHY 

The samples were processed and the slides prepared by standard laboratory procedures, as described by Robert H. Tschudy (1970, p. F2). 

A Zeiss photoscope was used for the photographs 

of all specimens. Neofluar X 54 and X 100 and 

apochromatic x 40 oil-immersion objectives were 

used on the photoscope for all except one of the 

photographs; a neofluar 10 x objective was used 

for the photograph of an extremely large spore, 

Ghoshispora sp. (pl. 1, fig. 17). Adox KB-14, 35-mm 

(millimeter) film was used for all the photographs. 

1 
2 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

Bearpaw 

Shale 

NW¥<1 NE¥<1 sec. 31, 

T. 22 N., R. 24 E., 

Phillips County 

Gill and Schultz 

section G61-13E 

NWl/4 sec. 26, 

T. 24 N., R. 17E., 

Blaine County 

----,.---+--------ll-04067-----+-----

z 

0 

t-

<( 

:2: 

a:: 

0 

LL. 

a:: 

LJ.J 

> 

a:: 

J: 

tt 

ro 

c. 

Q) 

c 

-~ 

E 

c 

0 

c 

Q) 

c. 

c. 

::J Gill and Schultz 

section G61-13C 

About center of 

N El;4 sec. 31, 

- 03725-A 

~ 03725-B 

1- 03725;-C 

r-- 03725-0 

- 03725-E 

T. 23 N., R. 17 E., 

Fergus County 

SWlj4 SW1;4 SWI;4 sec. 15., 

T. 23 N., R. 22 E., 

Phillips County I- 03726-B 

o 

::J 

...., ----+04069----i-03726-A-----

·Ill 

c. 

Q) 

c 

·;:: 

ro 

E 

Q) 

:;: 

0 

. ...J 

Claggett 

Shale 

SWlj4 SW1;4 sec. 36, 

T. 23 N., R. 16 E., 

1- 0 3726-E Fergus County 

1-03726-0 I 

j04070 

1-03726-C 

l Feet [0 

FIGURE 1.-Composite section showing stratigraphic position of pollen-spore collections from Judith River Formation. Collections made by J. R. Gill and L. G. Schultz 

in 1961 and by J. R. Gill in 1967. Data from unpublished composite section by J. R. Gill and L. G. Schultz. 

PROCEDURE OF STUDY 

Two hundred palynomorphs, consisting of pollen 

and . spores, were counted under high-power (X 

430) magnification for each of the 13 samples used 

in this study (table 1). An individual count for any 

taxon may be divided by two to determine percentage. At least two (and for some samples as many 

as six) slides were scanned in search for taxa not 

found within the 200-count procedure; these additional taxa are indicated by the symbol X. The taxa 

and groups of taxa in table 1 are numbered consecutively, and these numbers are used in the section "S~stematics" of this report and in tables 2 

and 3. 

Dinoflagellates and acritarchs were found in a 

few of the samples throughout the section. These 

were not included in the counts sho\\ n in table 1. 

In samples where present, these made up 2 percent 

or less 6f the totar palynomorph counts. 

Most of the taxa that were morphologically 

worthy of taxonomic designations were assignable 

to previously known and described species and (or) 

genera. A few of these genera and species are redescribed on the basis of the additional data obtained from the Judith River Formation specimens, 

but new descriptions are furnished only if contributions to previous descriptions can be made. 

Two new genera, 10 new species, and one new 

variety were named and described to accommodate 

new taxa. Several of the new species occurred in 

the samples in vecy low frequency but were numerically well represented on the slides and were 

conspicuous elements of the microfossil assemblage. 

Some of these new species apparently have limited 

stratigraphic ranges. All descriptions of new species 

except one were based on at least 15 specimens. Two 

additional new genera and their respective type 

species were named and described from samples of 

this series of Judith River Formation samples in 

an earlier report (Tschudy, 1971). 

During the literature search involved in the systematic part of this study several taxonomic problems were evident. The problems are discussed, and 

attempts are made to contribute to their resolutions. 

DEPOSITORY OF TYPES AND OTHER 

ILLUSTRATED SPECIMENS 

All specimens found in samples of the Judith 

River Formation that are illustrated in this report 

are deposited in the slide files of the paleobotanical 

collections of the U.S. Geological Survey, Denver, 

Colo. The tYiPe specimens are within the ink-circled 

areas on the slides but may also be located by the 

mechanical stage coordinates which are given in 

the plate explanations, this report. These coordinates may be converted to those of another mechanical stage as explained by Tschudy ( 1969, p. A1) or 

Tschudy (1971, p. B53). 
POLLEN-SPORE ZONES 3 

T. 

24 

N. 

R. 16 E. R. 19 E. R. 20 E. 

T. ~~~--+---~~~------r-~~~~ 23 

N. 

T. 

22 

R.23 E. R. 24 E. R.25 E. R. 26 E. 

! I 

N. FERGUS COUNTY 

T. 

21 

N. 

0 5 MILES 

FIGURE 2.-lndex map showing sample localities. 

ACKNOWLEDGMENTS 

Many people contributed to this study and their 

help is appreciated. Dr. B. E. Balme furnished a 

slide containing specimens of Dulhuntyispora which 

were useful for comparison with species of the new 

genus Trisolissporites that is described in this report. J. R. Gill and L. G. Schultz furnished the rock 

samples and the stratigraphic information pert'aining to them. Sharon Van Loenen did the laboratory 

preparations of the sample material. John W. Hall 

loaned me some of Richard L. Pierce's type specimens from the pale~botanical collection of the University of Minnesota's Botany Department. 

POLLEN-SPORE ZONES 

Pollen-spore zones within the section studied are 

not easily discernible from the total data shown in 

table 1; however, three zones and one transitional 

zone-based on 22 selected taxa from table 1-are 

recognizable and are indicated in table 2. The highest sample in the section ( D4067) which was believed by J. R. Gill to be Bearpaw Shale (basal) 

or Judith River Formation (uppermost) is palynologically similar to the next lower sample, D3725-A. 

These two samples make up the uppermost zone, 

here designated as zone 1. The next four samples, 

lower in the section, contain several taxa that are 

not present in the uppermost zone; these four samples constitute zone 2. The lowermost four samples 

of the section are palynologically distinguishable 

from the samples higher in the section. These contain several taxa not present in the samples above, 

and lack several taxa that are more or less consistently present in the upper samples. The lowermost four samples make up zone 3. Between zones 

2 and 3 are three samples that have pollen and spore 

characteristics (based on the selected taxa) partly 

resembling those of the upper zone (zone 2) and 

partly resembling those of the lower zone (zone 3) ; 

these make up the transitional zone. 

Many of the more abundant species and groups 

of taxa of the Judith River Formation assemblage 

range throughout the section studied, with erratic 

percentage occurrences from sample to sample and 

with no apparent trend toward greater or less 

abundance from the top to the bottom of the section; these taxa thus do not appear to have any 

stratigraphic significance within the Judith River 

Formation section. A few of the more abundant 

species, however, show trends toward higher percentage occurrence in either the lower or the upper 

part of the section-for example, Tricolpites reticulatus (taxon 24) and Tricolpites hians (taxon 25) 

are more abundant in the lower part of the s·ection, 

and Tricolpopollenites levitas (taxon 29) and Tricolpopollenites tessellatus (taxon 30) are more 

abundant in the upper part. Ratios in table 3 clearly 

show this reversal in abundance of the two pairs 
4 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

TABLE !.-Stratigraphic distribution of pollen and spores 

[Counts divided by 2=percentages. X denotes occurrence; asterisk (*) indicates questionable determination] 

Judith River Formation 

Taxon or group 

Upper nonmarine part Lower marine part 

No. Name 

1 Cyathidites minor --------------------- 2 

2 Deltoidospora sp ---------------------- 1 

3 Stereisporites spp --------------------- 1 

4 Leptolepidites major and 

cf. L. major X 

5 Foraminisporis wonthaggiensis --------- X 

6 Kuylisporites spp --------------------- X 

7 Cicatricosisporites radiatus ------------ X 

8 Ghoshispora sp. ----------------------- X 

9 Gleicheniidites spp -------------------- 11 

10 Camarozonosporites spp --------------- 1 

11 Unclassified triletes ------------------- 9 

12 Laevigatosporites spp ------------------ 19 

13 Umbosporites callosus ----------------- X 

14 Microfoveolatosporis canaliculatus ______ 2 

15 Rugubivesiculites sp ------------------- X 

16 Unclassified bisaccates ----------------- 1 

17 Unclas·sified inaperturates -------------- 1 

18 Taxodiaceaepollenites hiatus ------------ 37 

19 Equisetosporites spp ------------------- X 

20 Liliacidites complexus ----------------- 9 

21 A recipites spp. and unclassified 

monosulcates ----------------------- 4 

22 Aesculiidites dubius ------------------- 2 

23 Pulcheripollenites krempii -------------- 1 

24 Tricolpites reticulatus ----------------- 1 

25 Tricolpites hians ---------------------- 5 

26 Tricolpites spp ------------------------ 1 

27 Unclassified tricolpates (polar 

axis le·ss than 20p) ------------------ 28 

28 Miscellaneous unclassified tricolpates ___ 5 

29 Tricolpopollenites levitas --------------- 2 

30 Tricolpopollenites tesselatus ----------- 22 

31 Cranwellia bacata -------------------- X 

32 Aquilapollenites quadrilobus ------------ 14 

33 trialatus var. uniformis _ X 

34 Aquilapollenites sp -------------------- X 

35 Aquilapollenites attenuatus ------------- X 

36 Aquilapollenites spp ------------------- 1 

37 Mancicorpus calvus -------------------- 2 

38 Tricolporites rhomboides --------------- 2 

39 Montanapollis endannulatus ------------ X 

40 Unclassified tricolporates -------------- 4 

41 Subtriporopollenites alpinus ------------ 5 

42 Proteacidites spp ---------------------- X 

43 Erdtmanipollis pachysandroides -------- X 

44 Single cells of Dyadonapites reticulatus?_ 7 

45 Cicatricosisporites spp ----------------- 46 Alisporites grarulis -------------------- 47 Alisporites thomasii ------------------- 48 Cedripites parvus ---------------------

3 

X 

2 

4 

27 

X 

8 

2 

14 

2 

4 

6 

* 1 

14 

16 

7 

5 

21 

9 

X 

6 

1 

3 

1 

11 

1 

1 

2 

1 

2 

2 

1 

1 

1 

X 

6 

X 

3 

6 

X 

47 

3 

1 

3 

3 

23 

7 

5 

1 

1 

2 

1 

30 

1 

X 

30 

3 

X 

2 

20 

X 

1 

X 

X 

1 

2 

10 

X 

1 

1 

X 

13 

2 

43 

23 

2 

X 

X 

3 

4 

32 

11 

X 

10 

1 

X 

USGS paleobotany locality No. 

2 

2 

1 

1 

12 

3 

55 

1 

4 

X 

4 

28 

13 

5 

2 

X 

2 

5 

7 

1 

3 

X 

6 

7 

2 

* 1 

1 

3 

11 

23 

2 

68 

3 

X 

2 

18 

7 

1 

X 

6 

4 

1 

32 

X 

1 

X 

X 

1 

1 

2 

4 

11 

84 

3 

1 

46 

8 

18 

4 

3 

2 

2 

1 

2 

X 

2 

1 

4 

8 

33 

X 

70 

X 

4 

X 

2 

X 

42 

7 

6 

1 

X 

5 

1 

1 

1 

2 

1 

6 

6 

30 

4 

8 

4 

10 

1 

4 

6 

2 

1 

24 

21 

4 

7 

11 

6 

2 

2 

1 

10 

X 

1 

X 

X 

1 

1 

2 

X 

2 

X 

6 

3 

X 

X 

16 

5 

44 

X 

11 

3 

22 

28 

X 

7 

7 

1 

X 

1 

X 

2 

X 

X 

4 

6 

X 

2 

1 

7 

2 

4 

16 

X 

7 

14 

3 

3 

7 

2 

35 

5 

1 

2 

2 

6 

25 

6 

7 

X 

1 

6 

3 

1 

1 

1 

1 

X 

7 

7 

6 

2 

1 

4 

25 

7 

1 

1 

7 

X 

45 

20 

1 

2 

1 

1 

10 

10 

11 

3 

1 

X 

3 

2 

2 

2 

10 

6 

22 

4 

5 

5 

6 

20 

3 

* 1 

11 

17 

* 1 

10 

5 

1 

7 

1 

17 

4 

5 

2 

1 

1 

1 

1 
POLLEN-SPORE ZONES 5 

TABLE !.-Stratigraphic distribution of pollen and spores-Continued 

;§~ '0 ... 

=~ Judith River Formation 

~~ ocu Po .-.p, 

i.e 

ell 

..c= Taxon or group .... 0 

cu~ 

]~ rno 

I!: I&! 

Cllt Upper nonmarine part Lower marine part e.:: 

8!~ 

~ 

USGS paleobotany locality No. 

No. Name 

< ~ 0 ~ f";a;l 

~ 1 f";a;l 9 0 

too I I I .b I I I It:> It:> It:> It:> "' a> "' "' "' 0 "' "' C'l C'l C'l C'l C'l C'l "' C'l C'l c<l t- C'l 0 t- t- too too too t- 0 t- too t- 0 too 

""" <'.) <'.) <'.) <'.) <'.) <'.) 

""" <'.) <'.) <'.) """ <'.) 

~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 

49 Pityosporites cf. P. c.onstrictus --------- 3 X 3 

50 Pityosporites elongatus var. elongatus -- 3 2 1 7 1 

51 Pityosporites elongatus var. grand is ---- 2 X 1 4 1 

52 Podocarpidites cf. P. ornatus ---------- X 2 

53 Vitreisporites pallid us ----------------- 1 1 X 1 1 4 4 4 

54 Classopollis spp ---------------------- 1 X 1 1 2 

55 Liliacidites 'Uariegatus ----------------- 9 1 3 1 1 

56 Liliacidites spp ----------------------- 1 1 2 X 1 2 1 2 2 1 

57 Cranwellia rumseyensis ---------------- X 3 x. 2 3 

58 Cf. Cranwellia ------------------------ X 

59 Aquilapollenites turbidus --------------- X 1 X X X X X X 5 

60 M ancicorpus tripod if ormis ------------- X X 1 

61 Triatriopollenites co status ------------- 1 X 

62 Unclassified triporates ----------------- 2 1 1 2 3 1 1 4 X 5 5 3 

63 Inaperturotetradites scabratus ---------- 1 1 X X 1 1 2 1 

64 Baculatisporites sp -------------------- X 

65 Aquilapollenites senonicus -------------- 1 1 X 4 X 

66 Dyadonapites reticulatus --------------- 15 7 38 2 1 1 

67 Seductisporites emtnens ---------------- X * 1 X X * 1 

68 Cedripites cretaceus ------------------- X 2 3 

69 Ovoidites ligneolus -------------------- X 1 1 

70 Abietineaepollenites varius _____________ 1 1 1 

71 Aquilapollenites clarireticulatus _________ X X *2 

72 Siberiapollis montanensis ---------·----- X X 2 

73 Osmundac.idites sp --------------------- 1 2 1 X 

74 Ischyosporites cf. I. punctatus ---------- X 

75 Fibulapollis scabratus ----------------- X X X X 

76 Aquilapollenites trialatus -------------- X X 1 

77 A quilapollen ites reticulatus ------------ X 3 1 

78 Sindorapollis granulatus --------------- 10 

79 Trisolis1~porites montanensis ------------ X X 

80 S eductispori tes gemmiferus ------------ X X 

81 Pityosporites alatipollcnites ------------ X 

82 Pristinuspollenites microsaccus --------- X 3 1 

83 Cyathidites sp ------------------------ X 

84 Cicatricosisporites stoveri -------------- X 

85 Polycingulatisporites spp --------------- X X X 

86 Trisolissporites chlonova.e -------------- X 

87 S eductisporites barbatus --------------- 2 

88 Zonalapollenites sp -------------------- X 1 1 

89 Podocarpidites multesimus ------------- X 

90 Sequoiapollenites spp ------------------ X 1 

91 Cupanieidites sp ---------------------- X 

92 P eriporopollenites sp ------------------ 2 1 X 

93 Droseridites cf. D. spinulosus ----------- 1 

94 Grewipollenites radiatus --------------- 7 1 

95 Liliac.idites cf. L. morrinensis ---------- 1 

96 Tricolporites sp. ----------------- 8 1 

Total ----------------------------- 200 200 200 200 200 200 200 200 200 200 200 200 200 

494-146 0 - 73 - 2 
6 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

TABLE 2.-Pollen-spore zones, based on selected taxa from table 1 

[Taxon or group numbers and the counts correspond to those in table 1. Counts divided by 2 =percentages. X denotes occurrence; asterisk ( *) 

indicates questionable determination] 

Taxon or group 

No. Name 

Zone 1 

20 Liliacidites complexus ----------------- 9 

31 Cranwellia bacata --------------------- X 

32 Aquilapollenites quadrilobus ------------ 14 

35 Aquilapollenites attenuatus ------------- X 

39 Montanapollis endannulatus ------------ X 

57 Cranwellia rumseyensis ---------------- 65 Aquilapollenitcs senonicus -------------

4 

9 

X 

X 

1 

1 

1 

3 

1 

Upper nonmarine 

part 

Judith River Formation 

USGS paleobotany locality No. 

0 A IO:l ~ 

I I I I It:> It:> It:> "' 0> C'l C'l C'l C'-1 

t- t- t- "' t- 0 

"'-' "'-' "'-' "'-' ...,. A A A A A 

Transitional Zone 2 zone 

2 1 X 

2 X 1 

2 

3 X X 

X 2 

1 X 

1 

"' C'l 

t-

"'-' A 

6 

6 

3 

4 

Lower marine 

part 

IO:l A I I 

"' C'l "' C'l t- t- "'-' "'-' A A 

0 

t0 

...,. 

A 

Zone 3 

X 

0 

I 

"' C'l 

t-

"'-' A 

66 Dyadonapites reticulatus --------------- 15 7 38 2 1 1 

67 Seductisporites eminens ---------------- 72 Siberiapollis montanensis -------------- 75 Fibulapollis scabratus ----------------- 76 Aquilapollenites trialatus -------------- 77 Aquilapollenites reticulatus ------------ 78 Sindorapollis granulatus --------------- 79 Trisolissporites montanensis ------------ 80 Seductisporites gemmiferus ------------ 82 Pristinuspollenites microsaccus --------- 85 Polyc,ingulatisporites spp --------------

86 Trisolissporites chlonovae -------------- 87 Seductisporites barbatus --------------- 92 Periporopollenites sp ------------------ 94 Grewipollenites radiatus ---------------

of species. in the upper and lower parts of the section. 

SYSTEMATICS 

In this section, taxa of species rank are arranged 

under their appropriate genera. Spore genera are 

followed by those of pollen. All genera are regarded 

as form genera. Previously known and described 

genera and species are redescribed only when the 

author believes that contributions to the int~;rpretation of the taxa, based on data from the Judith 

River Formation specimens, can be made. 

TRILETES 

Genus CY ATHIDITES Couper, 1953 

Cyathidites Couper, 1953, p. 27. 

Cyathidites Couper, 1953. In Dettmann, 1963, p. 22. 

Type species.- Cyathidites australis Couper, 

1953, p. 27, pl. 2, fig. 11 (holotype) ; New Zealand; 

Ohika beds, Jurassic. 

X * 1 X X * 1 

X X 2 

X X X X 

X X 1 

X 3 1 

10 

X X 

X X 

X 3 1 

X X X 

X 

2 

2 1 X 

7 1 

Cyatbidi.tes minor Couper, 1953 

Plate 1, figure 1; table 1, No. 1 

Cyathidites minor Couper, 1953, p. 28. 

Holotype.-ln Couper, 1953, p. 28, pl. 2, fig. 13; 

Ohika beds; New Zealand; Jurassic. 

Cyatbidites sp. 

Plate 1, figure 2; table 1, No. 83 

Genus DEL TOIDOSPORA Miner, 1935 em£nd. Potonie, 1956 

Deltoidospora Miner, 1935, p. 618. 

Deltoidospora Miner, 1935 emend. Potoniee, 1956, p. 13. 

Type species.-(designated by Potonie, 1956, p. 

13) Deltoidospora hallii Miner, 1935, p. 618, pl. 24, 

fig. 7 (lectotype) ; Cascade County, Mont. Kootenai 

Formation ; Early Cretaceous. 

Deltoidospora sp. 

Plate 1, figure 3; table 1, No. 2 

Genus STEREISPORITES Pflug in Thomson and Pflug, 1953 

Sphagnum-sporites Raatz, 1937, p. 9, (No generic description.) 
SYSTEMATICS 7 

TABLE·3.-Abundance ratios between two pairs of pollen species in the upper and lower parts of the composite section 

[Ratios be~ween total counts of Tricolpites r(!ticulatus plus Tricolpites hians and total counts of Tricolpopollenites levitas plus Tricolpopollenites tes- seUatus m the tl_lree zones and transitional zone (see table. 2)---showing relative abundance of the two pairs of species in the upper and lower 

parts of the sectwn Taxa and counts from table 1. Counts divided by 2 =percentages] 

Taxon or group 

< ill I I 

No. Name 1:- 10 10 <0 ~ ~ 

0 1:- 1:- '<I' ~ ~ 

A A A 

Zone 1 

24 Tricolpites reticulatus ----------------- 1 14 --- 25 Tricolpites hians ---------------------- 5 16 3 

Total -------------------------- 36 

29 Tricolpopollenites levitas --------------- 2 21 5 

30 Tricolpopollenites tessellatus ----------- 22 --- ---

Total -------------------------- 45 

Ratio: 

Total T. reticulatus + T. hians 36 

--- -=0.80 

Total T. levitas + T. tessellatus 45 

Stereisporites Pflug in Thomson and Pflug, 1953 (March), 

p. 53. 

Sphagnites Cookson, 1953b (October), p. 463. 

Sphagnumsporites Raatz, 1937. In Potonie, 1956, p. 17. 

Stereisporites Thomson and Pflug, 1953 [March] s. 1. In 

Krutzsch, 1963b, p. 9-20. 

Stereisporites Thomson and Pflug, 1953 (March). In Potonie, 

1966, p. 19. 

Type species.-Stereisporites (Sporites) stereoides (Potonie and Venitz) Thomson and Pflug, 

1953, p. 53 (designated by Potonie, 1966, p. 19); 

lectotype in Potonie and Venitz (1934, p. 11, pl. 1, 

fig. 4) near KOln, Germany; late Oligocene to Miocene. 

Remarks.-Pflug in Thomson and Pflug (1953) 

was the author of Stereisporites-not Thomson and 

Pflug (1953) as in Krutzsch (1963b, p. 9) and in 

Potonie (1966, p. 19). See Thomson and Pflug 

(1953, p. 53). 

Stereisporites spp. 

Plate 1, figure 4, 5; table 1, No. 3 

Genus LEPTOLEPIDITES Couper, 1953 emend. Potonie, 1966 

Leptolepidites Couper, 1953, p. 28. 

Leptolepidites Couper, 1953 emend. Potonie, 1966, p. 30. 

Type species.-Leptolepidites verrucatus Couper, 

1953, p. 28, pl. 2, fig. 14 (holotype); New Zealand; 

Ohika beds; Jurassic. 

Judith River Formation 

Under nonmarine Lower marine 

part part 

USGS paleobotany locality No. 

0 A r:il ill < r:il I I I I I I ~ 10 10 10 <0 Cl> <0 <0 <0 0 ~ ~ ~ ~ <0 ~ ~ ~ 1:- 1:- 1:- 1:- 1:- 0 1:- 1:- 1:- 0 ~ ~ ~ ~ '<I' ~ ~ ~ '<I' 

A A A A A A A A A 

Transitional Zone 2 zon·e Zone 3 

1 X X 46 --- 24 22 2 1 

1 4 2 8 2 21 28 6 7 

11 101 94 

43 --- --- --- --- 11 --- --- 1 

23 5 1 --- 6 --- 1 --- 2 

77 17 5 

11 101 94 

-=0.14 -=5.94 -=18.80 

77 17 5 

Leptolepidites major Couper, 1958 

Plate 1, figure 6; table 1, No. 4 

Leptolepidites major Couper, 1958, p. 141. 

0 

I 

<0 

~ 

1:-

~ 

A 

11 

17 

1 

---

Holotype.-In Couper, 1958, p. 141, pl. 21, fig. 7; 

Yorkshire, England; Sycarham Beds; Middle Jurassic. 

Leptolepidites cf. L. major 

Plate 1, figure 7; table 1, No. 4 

Genus BACULATISPORITES Pftug and Thomson in Thomson 

and PJiug, 1953 

Bac:ulatisporites Pflug and Thomson in Thomson and Pflug, 

1953, p. 56. 

Baculatisporites Thomson and Pflug, 1953. In Potonie, 1956, 

p. 33. 

Baculatisporites Thomson and Pflug, 1953. In Dettmann, 

1963, p. 34. 

Type species.-Baculatisporites primarius (Wolff) 

Thomson and Pflug, 1953, p. 56; holotype in Wolff, 

1934, p. 66, pl. 5, fig.· 8; near Dettingen, Germany; 

Pliocene. 

Baculatisporites sp. 

Plate 1, figure 8; table 1, No. 64 

Genus OSMUNDACIDITES Couper, 1953 

Osmundacidites Couper, 1953, p. 20. 

Osmundacidites Couper, 1953. In Potonie, 1956, p. 30-31. 

Conosmundasporites Klaus, 1960, I>· 127-129. 
8 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

Osmundacidites Couper, 1953. In Dettmann, 1963, p. 31-32. 

Osmundacidites Couper, 1953. In Potonie, 1966, p. 36. 

Type species.-Osmundacidites wellmanii Couper, 

1953, p. 20, pl. 1, fig. 5 (holotype) ; New Zealand; 

Ohika beds; Middle Jurassic. Holotype also illustrated in Couper, 1960, pl. 1, fig. 1. 

Osmundacidites sp. 

Plate 1, figure 9; table 1, No. 73 

Genus KUYLISPORITES Potonie, 1956 

Kuylisporites Potonie, 1956, p. 38. 

Type species.-Kuylisporites waterbolki Potonie, 

1956, p. 38. Illustrated in Kuyl, Muller, and Waterbolk, 1955, pl. 1, fig. 7 (holotype) ; Trinidad, West 

Indies ; late Tertiary. 

Kuylisporites spp. 

Plate 1, figures 10, 11; table 1, No. 6 

Genus ISCHYOSPORITES Bahne, 1957 

Ischyosporites Balme, 1957, p. 23. 

Ischyosporites Balme, 1957. In oPtonie, 1960, p. 46. 

Type species.-lschyosporites crateris Balme, 

1957, p. 23-24, pl. 3, fig. 45 (holotype); Western 

Australia; ?Neocomian-early. Aptian. 

lschyosporites punctatus Cookson and Dettman, 1958 

Ischyosporites punctatus Cookson and Dettmann, 1958, p. 104. 

Holotype.-In Cookson and Dettman, 1958, p. 104, 

pl. XVI, fig. 1; South Australia; Early Cretaceous. 

Ischyosporites cf. I. punctatus 

Plate 1, figure 12; table 1, No. 7 4 

Genus CICATRICOSISPROITES Potonie and Gelletich, 1933 

emend. Potonie, 1966 

Cicatricosi-sporites Potonie and Gelletich, 1933, p. 522. 

Cicatricosisporites Potonie and Gelletich, 1933. In Potonie, 

1956, p. 47-48. 

Cic,atricosisporites Potonie and Gelletich, 1933 emend. Potonie, 1966, p. 58. 

Type species.-Cicatricosisporites dorogensis Potonie and Gelletich, 1933, p. 522, pl. 1, fig. 1 (lectotype, designated by Potonie, 1956, p. 47) ; Dorog, 

Ungarn, Germany; late Paleocene to Eocene. 

Cicatricosisporites stoveri Pocock, 1964 

Plate 1, figure 13; table 1, No. 84 

Cicatricosisporites stoveri Pocock, 1964, p. 161. 

Holotype.-In Pocock, 1964, p. 161, pl. 2, figs. 

24, 25; Saskatoon area, Saskatchewan, Canada; 

Upper part, Mannville Group; middle Albian. 

Cicatricosisporites sp. 

Plate 1, figure 14; table 1, No. 45 

Cicatricosisporites radiatus Krutzsch, 1959b 

Plate 1, figures 15, 16; table 1, No. 7 

Cicatricosisporites radiatus Krutzsch, 1959b, p. 126-127. 

Anemia radiata (Krutzsch) Stanley, 1965, p. 258-259, pl. 33, 

figs. 6, 7; South Dakota; Paleocene. 

Holotype.-Illustrated in Krutzsch, 1957, pl. 1, 

figs. 41, 42; middle Europe; late Oligocene or early 

Miocene. 

Genus GHOSHISPORA Srivastava, 1966 sensu Leffingwell, 1971 

Ghoshispora Srivastava, 1966, p. 511. 

Styx Norton in Norton and Hall, 1967, p. 104. 

Ghoshispora Srivastava, 1966. In Leffingwell, 1971, p. 24. 

Type species.-Ghoshispora scollardiana Srivastava, 1966, p. 511-512, pl. II, fig. 9 (holotype) ; 

Scollard, Alberta, Canada; Edmonton Formation 

(upper part) ; late Maestrichtian. 

Ghosispora sp. 

Plate 1, figure 17 (X 500); table 1, No. 8 

Genus FORAMINISPORIS Krutzsch, 1959a 

Foraminisporis Krutzsch, 1959a, p. 130. 

Foraminisporis Krutzsch, 1959. In Dettmann, 1963, p. 71. 

Foraminisporis Krutzsch, April 1959. In Potonie, 1966, p. 35. 

Foraminisporis Krutzsch, 1959b. In Krutzsch, 1963a, p. 2. 

Type species. -Foraminisporis foraminis 

Krutzsch, 1959a, p. 130, pl. XIX, figs. 203-206 (bolotype, four levels) ; Geisel tal, Germany; Eocene. 

Foraminisporis wonthaggiensis (Cookson and Dettman, 1958) Dettman, 1963 

Plate 2, figures 1, .2; table 1, No. 5 

Apiculatisporis wonthaggiensis Cookson and Dettmann, 1958, 

p. 100. 

Foraminisporis wonthaggiensis (Cookson and Dettman) Dettmann, 1963, p. 71, pl. XIV, figs. 19-23; southeastern 

Australia; late-Mesozoic. 

Foraminisporis wonthaggiensis (Cookson and Dettman) 

Dettmann. In Norris, 1967, p. 98, pl. 13, fig. 23; central 

Alberta, Canada; lower part, Colorado Group; Albian-

?Cenomanian. 

Holotype.-In Cookson and Dettmann, 1958, p. 

100, pl. XIV, fig. 8; Wonthaggi State Coal Mine area, 

Victoria, eastern Australia; Early Cretaceous. 

Genus POLYCINGULA TIS PORITES Simoncsics and Kedves, 1961 emend. 

Playford and Dettmann, 1965 

Poly~ingulatisporites Simoncsics and Kedves, 1961, p. 34. 

Polycingulatisporites Simoncsics and Kedves, 1961 emend. 

Playford and Dettmann, 1965, p. 143-144. 

Type species.-Polycingulatisporites circulus Simoncsics and Kedves, 1961, p. 34, pl. VI, figs. 1-6; 

Jfungary; Early Cretaceous. 

Polycingulatisporites spp. 

Plate 2, figures 3-5; table 1, No. 85 

Genus TRISOLISSPORITES n. gen. 

Type species. - Trisolissporites montanensis n. 

sp., USGS paleobotany locality D3726-E, slide (3) 

at 99.3 x 11.3; plate 2, figures 6, 7 (holotype, two 
SYSTEMATICS 9 

levels), center NE1,4 sec. 31, T. 23 N., R. 17 E.; 

Fergus County, Mont.; Judith River Formation 

(lower marine part); late Campanian. 

Description.- Trilete miospores with narrow 

equatorial cingulum. Equatorial outline round, 

rounded triangular, or slightly irregular. Trilete 

mark distinct, with or without margines. Rays of 

trilete long, reaching to or almost to equator of 

spore. Proximal surface ornamented with three sunburst or daisylike configurations located in the three 

sectors partitioned off by the three rays of the trilete 

mark---each sunburst configuration completely occupying its tripartite sector of the proximal surface 

of the spore. Distal surface ornamented with thickenings-these of various sizes and shapes, randomly 

arranged or zonate. 

Rem,arks.-This new genus has been erected to 

accommodate a group of exotic trilete spores having 

in common three sunburstlike configurations on the 

proximal surfaces. Chlonova ( 1960, p. 37) remarked 

that some unique spores which she described, and 

which I have transferred to this genus, probably 

deserve separation into a special morphological 

genus. She referred to her species as the three-sun 

spores ; hence, I selected the name Tr·isolissporites 

for this genus. 

New combinations.-The following species conform to the new genus Trisolissporites and are 

here transferred to it. 

Trisolissporites (Stenozonotriletes) radiatus ( Chlonova, 1960, p. 36-37, pl. IV, figs. 22-23; 

Chulimo-Y enisei depression; Cenomanian-Turonian) n. comb. 

Trisolissporites ( Stenozonotriletes) stellatus ( Chlonova, 1960, p. 38, pl. IV, fig. 25; Chulimoy enisei depression ; Cenomanian-Turonian) n. 

comb. 

Trisolissporites ( Taurocusporites) radiatus ( Chlonova) (Krutzsch, 1963b, p. 20) n. comb. 

Trisolissporites (Taurocusporites) stellatus (Chlonova) (Krutzsch, 1963b, p. 20) n. comb. 

Trisolissporites (Dulhuntyispora? minuta) minutus 

(Jansonius, 1962, p. 48-49, pl. 11, figs. 1, 2; 

western Canada; Early Triassic) n. comb. 

Trisolissporites ( Taurocusporites) chlonovae (Doring, 1966, p. 105-106, pl. I, figs. 7-9; Brandenburg, Germany; Wealden) n. comb. 

Comparison with other genera.-Potonie ( 1966, 

p. 43) suggested that the spores (Stenozonotriletes 

radiatus and Stenozonotriletes stellatus) which 

Chlonova (1962, p. 300, pl. 1, figs. 2-4) illustrated 

could perhaps be placed in Dulhuntyispora. Trisolissporites superficially resembles the Australian Permian genus Dulhuntyispora Potonie ( 1956, p. 37-38) 

but is easily distinguishable because its interradial 

ornaments are on the proximal surface of the spore 

and are not inflations of the outer wall layer, whereas the interradial ornaments of Dulhuntyispora are 

equatorially to surbequatorially located and appear 

as domelike inflations or blisters-the wall of the 

scutula being. distinctly detached from. the lower 

wall layer or layers of the spore. (See Dulhuntyispora dulhuntyi, pl. 2, fig. 11.) 

The radiating proximal ornamentations and the 

distal ornamentation of Trisolissporites easily separate it from Stenozonotriletes N aumova ex N aumova 

emend. Potonie (1958, p. 25) whose exine was 

described by Potonie as smooth to chagranate 

("glatt his chagrinat"). 

Trisolissporites is separable from Taurocusporites 

Stover (1962 emend. Playford and Dettmann, 1965, 

p. 14.6) by its three daisylike proximal interradial 

orna~ents. Zonate concentric thickenings are present in some species of Trisolissporites but are not 

the essential features that they are for Taurocusporites. The proximal ornamentation of Trisolissporites 

extends to the equator of the spore whereas the 

equatorial area of the proximal surface of Taurocusporites segmentatus, the type species of Taurocusporites, is usually unornamented or has only a 

few scattered verrucae. 

Trisolissporites montanensis n. sp. 

Plate 2, figures 6-10; table 1, No. 79 

H olotype.-USGS paleobotany locality D3726-E, 

slide (3) at 99.3X11.3; plate 2, figures 6, 7 (two 

levels), center NE1,4 sec. 31, T, 23 N., R 17 E.; 

Fergus County, Mont.; Judith River Formation 

(lower marine part); late Campanian. 

Paratype.-USGS paleobotany locality D3726-E, 

slide (2) at 81.8 x 11.2, plate 2, figure 8; same locality data as for holotype. 

Description-(based on eight specimens) .-Cingulate, trilete miospores. Equatorial outline in polar 

view circular, rounded triangular, or slightly irregular. Some specimens having slight equatorial indentations in cingulum adjacent to ends of trilete rays. 

Equatorial diameter 35J-t-49J-t. Equatorial cingulum 

3J-t-4J-t wide. Rays of trilete mark reaching approximately to inner margin of cingulum; in some sp·ecimens rays bifurcating near their radial extremities 

and branches thence appearing to merge with inner 

margin of cingulum. Proximal surface ornamented 

with three sunburst or daisylike configurations located in the three sectors set apart by the three 

rays of the trilete mark. Centers of configurations 

located at approximate centers of the three sectors 

of the proximal surface. Rays of each configuration 
10 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

15-25 in number, consisting of thickenings of exine 

-these arranged to resemble the ray flowers of 

daisies. Center part of configurations circular to 

irregular in shape, devoid of exinal thickenings, 

approximately 5p. in diameter. Distal surface ornamented with flat, randomly arranged, oval, round, 

or irregular exinal thickenings 3p.-7 p. in diameter 

and 1p.-3p. apart. 

Comparison.-The proximal surface of T. montanensis n. sp. ~ seems to be indistinguishable from 

that of T. radiatus ( Chlonova) n. comb., but the 

dis~tal surface ofT. montanensis is ornamented with 

numerous flat randomly arranged round to slightly 

irregular thickenings, whereas the distal surface 

of S. radiatus, according to Chlonova ( 1960, p. 37), 

always has concentric solid or broken narrow ridges. 

Chlonova's diagram ( 1962, fig. 2b) of the distal 

side of her Stenozonotriletes radiatus shows a concentric arrangement of the distal thickenings ; and 

photographs of additional spores from the Turonian 

of Western Siberia that were identified as S. radiatus Chlonova also show a ringlike thickening (see 

Skuratenko, 1966, pl. II, fi.g. 3 and Samoilovitch and 

others, 1961, pl. XLII, figs. 5-7). T. radiatus and 

T. montanensis are probably from closely related 

plants, but the two species are easily distinguishable !because of the different configuration of their 

distal thickenings. 

T. montanensis n. sp. is easily distinguishable 

from T. stellatus (Chlonova) n. comb and T. chlonovae (Doring) n. comb. by its coarser proximal ornamentation. 

T. montanensis n. sp. is larger than T. minutus 

( J ansoni us) n. comb. and lacks the ringlike distal 

thickenings that distinguish I. minutus. 

Trisolissporites chlonovae (Doring, 1966) n. comb. 

Plate 2, figure 12-14; table 1, No. 86 

Taurocusporites chlonovae Doring, 1966, p. 105-106. 

Holotype.-ln Doring, 1966, p. 105-106, pl. I, figs. 

7-9; Brandenburg, Germany; Wealden, Early Cretaceous. 

Description (based on 13 specimens from USGS 

paleobotany locality D3726-E) .-Cingulate, trilete 

miospores. Equatorial outline in polar view circular 

or nearly so. Equatorial diameter 37 p.-56p.. Equatorial cingulum 2p.-5p. wide. Trilete mark distinct; 

margines 1p.-·2p. wide, in some specimens not clearly 

visible; rays extending to or almost to equator, in 

some specimens rays appearing to bifurcate near 

their radial extremities. Proximal surface of spore 

ornamented with numerous microverrucae, verrucae, 

and elongate thickenings-these alined toward a 

central point to form three starlike configurations 

whose centers are located approximately at the 

center of each of the three sectors set off by the 

three rays of the trilete mark. Individual microverrucae and verrucae of the starlike configurations 0.5p.-1p. in diameter, elongate thickenings 

0.5p.-1p. wide and as much as 5p. long. Distal surface of spore ornamented with circular to very 

irregular exinal thickenings 1p.-9p. in diameter and 

of low relief; distal thickenings numerous on some 

specimens, sparse on others, randomly arranged, 

not arranged to form concentric zones. 

Remarks.-! transferred Taurocusporites chlonovae Doring to Trisolissporites because of the three 

sunburstlike configurations on its proximal surface. 

Doring placed this species in Taurocusporites but 

in his description did not mention zonate thickenings on its distal surface, and if these thickenings 

are not present the species does not conform to 

Taurocusporites. In assigning my specimens to the 

species T. chlonovae, I am assuming that T. chlonovae does not have a zonate distal surface. 

T. chlonovae is possibly synonymous with T. stellatus (Chlonova) n. comb. Drawings rather than 

photographs ilustrated the holotype of T. stellatus 

and do not clearly show the characteristics of the 

species. 

T. chlonovae is easily differentiated from T. radiatus and from T. montanensis because of its much 

smaller and more numerous proximal exinal thickenings that make up the three sunburst configurations. 

Genus GLEICHENIIDITES Ross, 1949 emend. Skarby, 1964 

Gleicheniidites Ross, 1949, p. 31-32. (Valid name, monotypic.) 

Gleicheniidites Ross, 1949. In Delcourt and Sprumont, 1955, 

p. 26. 

Gleicheniidites (Ross, 1949) Delcourt and Sprumont, 1955. 

In Potonie, 1956, p. 14-15. 

Triremisporites Delcourt and Sprumont, 1957, 61-63. 

[in part] Gleicheniidites (Ross, 1949) Delcourt and Sprumont, 1955, emend. Delcourt and Sprumont, 1959, p. 34. 

(Included spores having continuous equatorial thickening of wall.) 

[in part] Gleicheniidites Ross, 1949 emend. Krutzsch, 1959a, 

p. 109-115. (Included ornamented spores.) 

[not] Gleicheniidites Grigorjeva in Samoilovitch and others, 

1961, p. 59-60. (Later homonym of Gleicheniidites Ross, 

1949.) 

Fasciatisporites Sato, 1961, p. 85. 

Gleicheniidites Ross ex Delcourt and Sprumont emend. Dettmann, 1963, p. 64-65. 

Gleicheniidites Ross, 1949 emend. Skarby, 1964, p. 60-65. 

Type species. - Gleicheniidites senonicus Ross, 

1949, p. 31-32, pl. I, fig. 3 (holotype) ; Asen, Scania, 

Sweden; late Santonian or early Campanian. 
SYSTEMATICS 11 

Gleicheniidites spp. 

Plate 3, figures 1-4; table 1, No. 9 

Genus CAMAROZONOSPORITES Pant ex Potonie', 1956 emend. 

Klaus, 1960 

Camarozonosporites Pant, 1954, p. 51. (Genus described but 

no species assigned.) 

Camaroz·onosporites Pant, 1954 ex Potonie, 1956, p. 65. 

Hamulatisporis Krutzsch, 19·59a, p. 157-158. 

Camerozonosporites Pant, 1954 ex Potonie, 1956. In Krutzsch, 

1959a, p. 186-187. 

Camarozonosporites Potonie, 1956 emend. Klaus, 1960, p. 

135-136. 

Camarozonosporites Pant, 1954 ex Potonie, 1956. In Krutzsch, 

1963a, p. 18, 20-23. 

Camarozonosporites (Pant ex Potonie, 1956) Klaus, 1960. In 

Potonie, 1966, p. 92. 

Camarozonosporites (Potonie, 1956) Klaus, 1960. In Hedlund, 

1966, p. 17. 

Type species.-Camarozonosporites (Rotaspora) 

cretaceus (Weyland and Krieger, 1953) Potonie, 

1956, p. 65; holotype in Weyland and Krieger, 1953, 

p. 12, pl. 3, fig. 27; Aachen, Basiston, Germany; 

middle Senonian. 

Camarozonosporites spp. 

Plate 3, figures 5, 6; table 1, No. 10 

MONOLETES 

Genus LAEVIGA TOSPORITES lhrahim, 1933 emend. Schopf, Wilson, 

and Bentall, 1944 

Laevigato-sporites Ibrahim, 1933, p. 39. 

Polypodium-sporites Raatz, 1937, p. 10. 

Polypodiaceae-sporites Thiergart, 1937, p. 297. 

Phaseolites Wilson and Coe, 1940, p. 182. 

Laevigato-sporites Ibrahim, 1933 emend. Schopf, Wilson, and 

Bentall, 1944, p. 36-37. 

Type species.-Laevigatosporites vulgaris (Ibrahim) Ibrahim, 1933, p. 39-40; holotype in Ibrahim, 

1932, p. 448, pl. XV, fig. 16; Westfal, Germany; 

Carboniferous. 

Laevigatosporites spp. 

Plate 3, figures 7, 8; table 1, No. 12 

Genus UMBOSPORITES Newman, 1965 

Umbosporites Newman, 1965, p. 10. 

Type species.-Umbosporites ca.llosus Newman, 

1965, p. 10; pl. 1, fig. 2 (holotype) ; northwestern 

Colorado ; basal Iles Formation ; Campanian. 

Umhosporites callosus Newman, 1965 

Plate 3, figures 9, 10; table 1, No. 13 

Genus MICROFOVEOLATOSPORIS Krutzsch, 1959a 

Microfoveolatosporis Krutzsch, 1959a, p. 211. 

Type species.-Microfoveolatosporis pseudodentatus Krutzsch, 1959a, p. 212, pl. 41, fig. 463 (bolotype) ; Geisel tal, Germany; Eocene. 

Microfoveolatosporis canalicalatus Dettmann, 1963 

Plate 3, figure 11; table 1, No. 1 

Microfoveolatosporis canaliculatus Dettman, 1963, p. 87. 

Holotype.-In Dettmann, 1963, p. 87, pl. XIX, 

figs. 15, 16; South Australia; Tambo Formation; 

Albian. 

ZONATES 

Genus SEDUCTISPORITES Chlonova, 1961 

Seductisporites Chlonova, 1961, p. 56-57. 

Seductisporites Chlonova, 1961. In Potonie, 1966, p. 96. 

Type species.-Seductisporites signifer Chlonova, 

1961, p. 56-57, pl. VII, fig. 44 (holotype) ; mouth of 

Vakh River, western Siberian lowlands; Maestrichtian-Danian. 

Description.-Inaperturate miospores having a 

thin outer wall layer which forms a zone at equator. 

Shape of spores in polar view circular to rounded 

triangular. Equatorial zona (flange) narrow or as 

much as 9p. wide. Outer wall layer membranous or 

as much as 1.5p. thick. Zona distinct and complete, 

or fragmentary, or almost lacking. Fragmentary or 

indistinct tetrad mark present in some specimens 

and species; when present, with or without thickened margins. Distal hemisphere of spore ornamented with simple, bifurcating, or netlike muroidal 

ridges or folds. Proximal hemisphere smooth or 

variously ornamented but differing from distal hemisphere. (Inasmuch as specimens that have fragmentary tetrad marks have their muroidal ridges 

on the distal hemisphere, it is assumed by analogy 

that specimens that lack tetrad marks also have 

their muroidal ridges on the distal side.) 

Comparison with similar genera.-Rouseisporites 

Pocock (1962, p. 52-53) differs from Seductisporites 

in that its zonate layer has invaginations in each 

radial region at the equator. (See Dettmann, 1963, 

p. 96.) 

Zlivisporis Pacltova (1961, p. 86-87) differs from 

Seductisporites in that it has a distinct trilete mark 

which reaches to the equator. 

The anastomosing folds of the perisporal layer of 

the Triassic genus Peroaletes Bharadwaj and Singh 

(1964, p. 38-39) have slight resemblance to the 

muroidal folds on the distal hemispheres of Seductisporites, Zlivisporis, and Rouseisporites; however, 

the two hemispheres of Peroaletes are apparently 

similarly ornamented and are not distinctly different from each other as they are in Seductisporites, 

Zlivisporis, and Rouseisporites. 

Seductisporites gemmiferus n. sp. 

Plate 3, figures 12-17; table 1, No. 80 

Holotype.-VSGS paleobotany locality D3726-A, 
12 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

slide (2) at 96.7X19.8; plate 3, figures 12, 13, 

center NE1;4 sec. 31, T. 23 N., R. 17 E.; Fergus 

County, Mont. ; Judith River Formation; late Campanian. 

Paratype.-USGS paleobotany locality D3726-A, 

slide (1) at 91.4 x 14.1; plate 3, figuies 14, 15; same 

locality data as for holotype. 

Paratype.-USGS paleobotany locality D3726-A, 

slide (2) at 92.4 x 18.3; plate 3, figures 16, 17; same 

locality data as for holotype. 

Description (based on 20 specimens) .-Inaperturate miospores. Shape in polar view round, rounded 

triangular, or irregular owing to folding of the delicate outer wall layer. Spores consisting of a central 

body and a thin-walled outer layer which forms a 

zone at the equator. Wall of central cell 1,u-2,u thick. 

Equatorial diameter of central body 42,u-59,u, of 

outer layer 48,u-70,u; zone extending 1,u-7 ,u beyond 

equator of central body. Width of zone in individual 

specimens nearly constant (pl. 3, fig. 16) or variable 

(pl. 3, fig. 12). Fragmentary tetrad mark present 

in some specimens; when present, raised, cordlike, 

contorted, sparingly warty. Surface of outer layer 

smooth to finely scabrate; distal surface having 

three to six radially oriented, raised, knifelike ridges 

or folds- these more or less perpendicular to surface, 2,lt'-4,u high, decreasing in height at outer ends, 

in some places bifurcating; in most specimens folds 

not extending as far as equator of central cell. 

Proximal surface of central cell ornamented with 

scattered, nearly spherical (1,u-4,u in diameter) 

wartlike projections or gemmae. 

Remarks.-This species was given the name gemmiferus (gemmifer, bearing or producing gems) 

because its proximal hemisphere is conspicuously 

adorned with small protrusions. 

Comparison with similar species.-The configuration of the ridges on the distal surface of S. gemmiferus resembles that of the muroidal ridges of 

Rouseisporites radiatus Dettmann ( 1963, p. 98, pl. 

XXIII, figs. 13-17) ; however, the two species are 

distinctly different because the zona of S. gemmiferus is wider than that of R. radiatus, S. gemmiferus does not have conical invaginations in the 

radial regions at the equator, and S. gemmiferus 

is ornamented on the proximal side, whereas R. 

radiatus is smooth. 

S. gemmiferus is separable from Seductisporites 

signifer because of the distinct wartlike ornamentation on its proximal hemisphere. 

S. gemmiferus lacks the flask-shaped equatorial 

invaginations that characterize Rouseisporites 

granospeciosus (Delcourt and Sprumont) Delcourt, 

Dettmann, and Hughes (1963, p. 291); and the wartlike ornamentation on the proximal surface of the 

central body of S. gemmiferus also distinguishes it 

from R. granospeciosus which has a smooth proximal surface. 

S. gemmiferus n. sp. differs from S. barbatus n. 

sp. because of its distinctly different proximal ornamentation and its less fragile equatorial zone. 

Seductisporites eminens n. sp. 

Plate 4, figures 1-3; table 1, No. 67 

[not] Lycopodium novomexicanum Anderson, 1960, p. 14-15, 

pl. 1, fig. 2. 

Lycopodiumsporites novomexicanum (Anderson) Drugg, 1967, 

p. 40, pl. 6, fig. 27. 

Holotype.-USGS paleobotany locality D3726-E; 

slide (5) at 99.0X10.3; plate 4, figure 1; center 

NE%, sec. 31, T. 23 N., R. 17 E.; Fergus County, 

Mont.; Judith River Formation (lower marine 

part) ; late Campanian. 

Paratype.-USGS paleobotany locality D3726-E, 

slide (9) at 84.8 x 16.1; plate 4, figure 2; same locality data as for holotype. 

Paratype.-USGS paleobotany locality D3726-E, 

slide (8) at 109.2X19.7; plate 4, figure 3; same 

locality data as for holotype. 

Description (based on 15 specimens, all in polar 

view) .-Inaperturate miospores. Shape in polar view 

rounded triangular. Equatorial diameter 52tL·-73,u. 

Tetrad mark when present contorted, in most specimens fragmentary, in some branching, with thickened margins approximately 1/L wide. Spore having 

two conspicuous wall layers ; inner layer 1tL-2fL thick, 

scabrate, confining inner cell; outer layer 0.5fL-1.5fL 

thick, enveloping inner cell and forming zona at 

equator. Equatorial zona narrow (1tL-3,u wide) in 

inter angular areas, wider (as much as 7 fL wide) at 

angles. Distal surface ornamented with muroidal 

folds which form a coarse reticulum; muri approximately 1/L in height, undulating; lumina irregular 

in shape, 8,u-15f-t in diameter; reticulum not extending beyond equator of central cell. Proximal surface 

smooth to scabrate. 

Remarks.-This species was given the name 

eminens (standing out, eminent) because of the 

conspicuous muroidal folds on its distal surface. 

Comparison with similar species.-A specimen 

from the Maestrichtian of Escarpado Canyon, Calif., 

which Drugg (1967, p. 40, pl. 6, fig. 27) identified 

as Lycopodiumsporites novomexicanum (Anderson) 

Drugg, seems to be the same as the new species 

described above. However, Seductisporites eminens 

seems to be different from Lycopodium novomexicanum Anderson (1960, p. 14-15, pl. 1, fig. 2). 

Anderson stated that the body of L. novomexicanum 
SYSTEMATICS 13 

is bordered equatorially by a thin irregular flange, 

with thickened extensions of the faint laesurae projecting into the flange on many specimens ; whereas, 

the equatorial flange of S. eminens n. sp. is narrow 

but not very thin walled or irregular, and laesurae 

have not been seen to extend into the flange. 

S. eminens, in common with Rouseisporites laevigatus Pocock (1962, p. 53-54, pl. 7, fig. 106), has a 

flange that is wider at the apices (angles) , but it 

differs from R. laevigatus in that it has a very distinct distal reticulum and does not have structures 

resembling pores at the apices. 

Its smooth to scabrate proximal surface and its 

lack of porelike apical structur·es easily distinguishes 

S. eminens from Rouseisporites reticulatus Pocock 

(1962, p. 53, pl. 7, fig. 101). 

The distal reticulum of S. eminens resembles that 

of Zlivisporis blanensis Pacltova (1961, p. 87, pl. 

II, figs. 1-4) but S. eminens is separable from Z. 

blanensis because of its thicker wall, its zona which 

is wider at the angles, and by its lack of a distinct 

Y -mark which extends to the equator. 

Seductisporites barbatus n. sp. 

Plate 4, figures 4-7; table 1, No. 87 

Holotype.-USGS paleobotany locality D3726-E, 

slide (9) at 111.3 X 3.2; plate 4, figure 4; center 

NE14 sec. 31, T. 23 N., R. 17 E.; Fergus County, 

Mont.; Judith River Formation (lower marine 

part); late Campanian. 

Paratype.-USGS paleobotany locality D3726-E, 

slide (7) at 82.0 x 12.5; plate 4, figure 5; same locality data as for holotype. 

Paratype.-USGS paleobotany locality D3726-E, 

slide (10) at 94.5X7.7; plate 4, figures 6, 7; same 

locality data as for holotype. 

Description (based on 35 specimens) .-Alete miospores having an outer wall layer which forms a 

zone at the equator. Outer wall membranous, delicate, slightly larger than central body, forming a 

narrow zone (1t-t-3t-t wide) at equator. In some 

specimens, zone fragmentary or completely lacking. 

Outline of spore in polar view round, rounded triangular, or irregular owing to folding or distortion. 

Diameter of central cell 49fL-76t-t; polar axis apparently shorter than equatorial diameter (judged 

from two specimens seen in equatorial view). Wall 

of central cell 1.5fL-2t-t thick, outer wall layer thin. 

No trilete mark evident in any of the 35 specimens 

examined. One hemisphere (distal?) of spore ornamented with muroidal ridges-these apparently 

composed of folds of the membranous outer wall 

layer; configuration of muroidal ridges varying 

from (1) a few ridges with no definite pattern, to 

494-146 0- 73- 3 

(2) a pseudo Y-mark with bifurcating rays, to (3) 

a more complex pattern of bifurcating ridges that 

join to form a very coarse more or less complete 

reticulum. Muri 1t-t-2t-t high and lumina (where present) 15t-t-30t-t in greatest diameter. Opposite hemispher·e (proximal?) ornamented with short (1t-t-3f-t) 

spines or hairlike projections. 

Observations have not yet proved whether the 

outer wall layer completely envelops the spinose 

hemisphere of the spore. 

Remarks.-This species was. given the name barbatus ( barbatus, bearded) because of its bristly or 

bearded proximal ( ?) ornamentation. 

Comparison with similar species.-The narrow 

membranous equatorial zone and the configuration 

of the muroidal ridges on one hemisphere (of some 

specimens) of S. barbatus n. sp. resemble those of 

Zlivisporis blanensis Pacltova (1961, p. 87, pl. II, 

figs. 1-4), but S. barbatus differs by not having a 

Y-mark and by having fine spines or hairlike projections on one hemisphere. 

The narrow equatorial zone and the configuration. 

of the muroidal ridges of S. barbatus also resemble 

those of Rouseisporites reticulatus Pocock ( 1962, p. 

53, pl. 7, fig. 101) and Rouseisporites radiatus 

Dettmann (1963, p. 98, pl. XXIII, figs. 13-17), but 

S. barbatus lacks the conical invaginations in the 

radial regions at the equator that characterize the 

other two species. The fine hairlike projections on 

one hemisphere of S. barbatus also distinguish it 

from R. reticulatus and R. radiatus. 

S. barbatus n. sp. differs from S. gemmiferus n. 

sp. by its distinctly different ornamentation and by 

its more fragile equatorial zone. 

Genus ZONALAPOLLENITES Pftug in Thomson and Pftug, 1953 

Tsugae-pollenites. In Potonie and Venitz, 1934, p. 17. 

Zonalapollenites Pflug in Thomson and Pflug, 1953, p. 66. 

Tsugaepollenites Potonie and Venitz, 1934. In Potonie, 1958, 

p. 48. 

Tsugaepollenites Potonie and Venitz, 1934. In J ansonius, 1962, 

p. 51. 

Tsugaepollenites Potonie and Venitz emend. Potonie. In Manum, 1962, p. 45. 

Tsugaepollenites Potonie and Venitz emend. Potonie, 1958. In 

Dettmann, 1963, p. 99. 

Tsugaepollenites (Potonie and Venitz, 1934) Potonie, 1958. 

In Potonie, 1966, p. 117. 

Zonalapollenites Pflug (in Thomson and Pflug). In Pocock, 

1968, p. 640. 

Tsugaepollenites Potonie and Venitz, 1934. In Potonie, 1970, 

p. 72. 

Type species.-Zona.lapollenites (Sporonites) igniculus (Potonie) Thomson and Pflug 1953; bolotype in Potonie, 1931d, p. 556, fig. 2; near KOln, 

Germany; Miooene. 
14 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

Remarks.-Potonie ( 1931d, p. 556) described, by 

means of symbols, and illustrated the new species 

Sporonites igniculus. Sporonites Potonie had earlier 

been introduced (Potonie, 1931c, p. 1) as an artificial genus (Gattung) for the provisional grouping 

of spores. 

In Potonie and Venitz (1934, p. 17), "Tsugaepollenites igniculus R. Pot." was introduced and 

described. The name Tsugae-pollenites was not 

there validated under the provision of a combined 

generic and specific description (see Lanjouw and 

others, 1966, art. 42) because the species name 

igniculus was not new. 

Pflug (in Thomson and Pflug, 1953, p. 66) described the new genus Zonalapollenites. In the same 

report, Sporonites igniculus Potonie was transferred 

to Zonalapollenites, and Z. igniculns was described. 

Pflug did not designate a type species for Zonalapollenites but according to artice PB. 4 (Lanjouw 

and others, 1952) which was in effect at that time 

"The type of a genus of fossil plants is the first 

described species which shows such characters as 

are necessary for distinguishing the genus from 

other taxa." Z onalapollenites igniculus met the requirements of article PB. 4 and hence became the 

type species for the genus Zonalapollenites. Also, 

according to article 37 (Lanjouw and others, 1966), 

the indication of a type species for a new genus 

was not a requirement for validation until J anuary 1, 1958. 

Potonie (1958, p. 48) considered Tsugaepollenites 

Potonie and Venitz, 1934 a valid name. He designated Tsugaepollenites ( al. Sporonites) igniculus 

(Potonie, 1931) Potonie and Venitz, 1934 as the 

"Genotypus" for Tsugaepollenites. He described the 

genotypus hut did not describe the genus Tsugaepollenites. 

Tsugaepollenites was described by Jansonius 

(1962, p. 51) and the name was then validated; 

however, it was a junior synonym of the earlier 

validated name Z onalapollenites. 

Pocock (1968, p. 639-640) recognized Zonalapollenites as a validly published name and considered Tsugaepollenites its junior synonym. He formally assigned Zonalapollenites igniculus as its type 

species. 

Zonalapollenites sp. 

Plate 5, figure 1; table 1, No. 88 

BISACCATES 

Genus ALISPORITES Daugherty, 1941 

Alisporites Daugherty, 1941, p. 98. 

Alisporites Daugherty, 1941. In Potonie and Kremp, 1956, p. 

176-177. 

Alisporites Daugherty, 1941. In Potonie, 1958, p. 60. 

Alisporites Daugherty, 1941. In Potonie, 1966, p. 119. 

Type species.-Alisporites opii Daugherty, 1941, 

p. 98, pl. 34, fig. 2; Arizona; Chinle Formation; 

Late Triassic. 

Alisporites grandis (Cookson, 1953b) Dettmann, 1963 

Plate 5, figures 2, 3; table 1, No. 46 

Disaccites grandis Cookson, 1953b, p. 471-472. 

Pityosporites grandis (Cookson) Balme, 1957, p. 36, pl. 10, 

figs. 110, 111; Western Australia; Early Cretaceous. 

Alisporites rotundus Rouse, 1959, p. 316, pl. 1, figs. 15, 16; 

British Columbia, Canada; Late Jurassic (?). 

Alisporites rotundus Rouse. In Pocock, 1962, p. 61-62, pl. 9, 

figs. 140, 141 western Canada; Early Cretaceous. 

Alisporites grandis (Cookson) Dettmann, 1963, p. 102, pl. 

XXV, figs. 1-4; southeastern Australia; late Mesozoic. 

Alisporites grandis (Cookson) Dettmann, 1963. In Srivastava, 

1966, p. 524, pl. VI, figs. 5, 7, 10; Alberta, Canada; 

Maestrichtian, Late Cretaceous. 

Alisporites grandis (Cookson) Dettmann. In Norris, 1967, 

p. 101, pl. 15, fig. 1; central Alberta, Canada; lower part 

of Colorado Group; Albian- ?Cenomanian. 

Holotype.-In Cookson, 1953b, p. 471-472, pl. 2, 

fig. 41; Comaum, South Australia; Mesozoic. 

Alisporites thomasii (Couper, 1958) Nilsson, 1958 

Plate 5, figures 4, 5; table 1, No. 4 7 

Pteruchipollenites thomasii Couper, 1958 (March), p. 150-

151. 

Alisporites thomasii (Couper) Nilsson, 1958 (November), p. 

83-84, pl. 8, fig. 1; Hoganas, southern Sweden: Lias. 

Alisporites bilateralis Rouse, 1959, p. 316, pl. 1, figs. 10, 11; 

southeastern British Columbia, Canada; Kootenay Formation; Early Cretaceous. 

Alisporites thomasii (Couper) Pocock, 1962, p. 62-63, pl. 9, 

figs. 142, 143; western Canada; Vanguard Formation; 

Late Jurassic; and quartz sand member of Mannville 

Group; Early Cretaceous. 

Alisporites thomasii (Couper) Pocock, 1962. In Singh, 1964, 

p. 109, pl. 14, figs. 11, 12; east-central Alberta, Canada; 

Mannville Group; Early Cretaceous. 

Alisporites bilateralis Rouse, 1959. In Hedlund, 1966, p. 23, 

pl. 7, fig. 3; Bryan County, Okla.; Woodbine Formation; 

Cenomanian. 

Alisporites bilateralis Rouse. In Norris, 1967, p. 101, pl. 14, 

figs. 18, 19; central Alberta, Canada; lower part, Colorado Group; Albian- ?Cenomanian. 

Holotype.-In Couper, 1958, p. 150, pl. 26, fig. 

10; northeastern Scotland; Middle Jurassic. 

Genus ABIETINEAEPOLLENITES Potonie, 1951 ex Delcourt and 

Sprumont, 1955 

Abietineae-pollenites Potonie, 1951, p, 144-145. (Not valid, 

genus not described.) 

Abietineaepollenites Potonie, 1951 ex Delcourt and Sprumont, 

1955, p. 51. 

Type species.-(designated by Del court and Sprumont, 1955, p. 51) Abietineaepollenites (Piceaepollenites) microalatus (Potonie. 1931c) Delcourt 
SYSTEMATICS 15 

and Sprumont, 1955. Holotype in Potonie, 1931c, p. 

5, fig. 34; Lausitz, Germany; Miocene. 

Abietineaepollenites varius Norton in Norton and Hall, 1969 

Plate 5, figure 6; table 1, No. 70 

Abietineaepollenites varius Norton in Norton and Hall, 1969, 

p. 29. 

H olotype.-In Norton and Hall, 1969, p. 29, pl. 4, 

fig. 5; Montana; Hell Creek Formation; Late Cretaceous. 

Genus CEDRIPITES Wodehouse, 1933 

Cedripites W odehouse, 1933, p. 490. 

Cedripites Wodehouse, 1933. In Singh, 1964, p. 111. 

Cedripites Wodehouse, 1933. In Pocock, 1962, p. 63. 

Type species.-Cedripites eocenicus Wodehouse, 

1933, p. 490-491, fig. 13; Colorado or Utah; Green 

River Formation; middle Eocene. 

Cedripites parvus Norton in Norton and Hall, 1969 

Plate 5, figure 7; table 1, No. 48 

Cedripites parvus Norton in Norton and Hall, 1969, p. 29. 

H olotype.-In Norton and Hall, 1969, p. 29; pl. 

4, fig. 7; Montana; Tullock Member of Fort Union 

Formation; Paleocene. 

Cedripites cretaceus Pocock, 1962 

Plate 5, figure 8; table 1, No. 68 

Cedripites cretaceus P~ock, 1962, p. 63. 

Cedripites cretac.eus P~ock, 1962. In Singh, 1964, p. 111, 

pl. 15, figs. 3-5; east-central Alberta, Canada; Mannville Group; Early Cretaceous. 

Cedripites cretaceus Pocock. In Norris, 1967, p. 101, pl. 14, 

figs. 20, 21 and pl. 15, fig. 5; central Alberta, Canada; 

lower part of Colorado Group; Albian- ?Cenomanian. 

Holotype.-In Pocock, 1962, p. 63, pl. 9, fig. 145; 

western Canada ; Quartz Sand Member, Mannville 

Group; Barremian. 

Genus PITYOSPORITES Seward, 1914 emend. Manum, 1960 

[in part] Pityosporites Seward, 1914, p. 23. 

Pityosporites Seward, 1914 emend. Potonie and Klaus, 1954, 

p. 534-536. 

Pinuspollenites Raatz, 1937 ex Potonie, 1958, p. 62. 

Pityosporites Seward, 1914 emend. Manum, 1960, p. 14. 

Pityosporites Seward, 1914 emend. Manum, 1960. In Singh, 

1964, p. 121-122. 

Type species.-Pityosporites antarcticus Seward, 

1914. In Manum, 1960, p. 14, figs. 1-4 (type specimen). 

Pityosporites constrictus Singh, 1964 

Pityosporites constrictus Singh, 1964, p. 122-123, pl. 16, figs. 

8, 9. 

Pityosporites constrictus Singh, 1964. In Srivastava, 1966, p. 

524, pl. VI, fig. 2; Scollard, Alberta, Canada; Edmonton 

Formation; Maestrichtian, Late Cretaceous. 

Holotype.-In Singh, 1964, p. 122-123, pl. 16, fig. 

8; Fort Augustus No. 1 well, Aberta, Canada; EllersHe Member of McMurray Formation, Mannville 

Group; Early Cretaceous. 

Pityosporites cf. P. constrictus 

Plate 5, figure 9; table 1, No. 49 . 

Remarks.-Except for their slightly smaller size 

(total breadth of grains 49,u-70,u) these grains ap:-

pear very similar to those of P. constrictus that 

were described and illustrated by Singh. 

Pityosporites alatipollenites (Rouse, 1959) Singh, 1964 

Plate 6, figure 1; table· 1, No. 81 

Pinus alatipollenites Rouse, 1959, p. 314, pl. 1, fig. 7; southeastern British Columbia, Canada; Kootenay Formation; 

Late Jurassic (?) . 

Pityosporites alatipollenites (Rouse) Singh, 1964, p. 123, 

pl. 16, fig. 10; east-central Alberta, Canada; Mannville 

Group; Early Cretaceous. 

Holotype.-In Radforth and Rouse, 1954, p. 195, 

pl. I, fig. 15; Entrance, Alberta, Canada; Brazeau 

Formation; Late Cretaceous. 

Pityosporites elongatus Norton in Norton and Hall, 1969 n. comb. 

Pinuspollenites elongatus Norton in Norton and Hall, 1969, 

p. 27. 

H olotype.-In Norton and Hall, 1969, p. 27, pl. 4~ 

fig. 1; Garfield County, Mont. ; Tullock· Member of 

Fort Union Formation; Paleocene. 

Pityosporites elongatus (Norton) Tschudy var. elongatus 

Plate 6, figures 2-4; table 1, No. 50 

H olotype.--"-Same as for Pityosporites elonga.tus 

(see above) . 

Pityosporites elongatus var. gran dis n. var. 

Plate 6, figures 5, 6; table 1, No. 51 

H olotype.-USGS paleobotany locality D3726-E, 

slide (5) at 116.0 X4.9; plate 6, figure 6; center 

NE14 sec. 31, T. 23 N., R. 17 E.; Fergus County, 

Mont. ; Judith River Formation (lower marine 

part); late Campanian. 

Paratype.;_USGS paleobotany locality D3726-E, 

slide (6) at 96.9 X 13.2; plate 6, figure 5; same locality data as for holotype. 

Description (based on 28 specimens) .-Bisaccate 

pollen grains. In polar view central body circular 

to slightly oval-in some specimens body breadth 

greater than length, in others length greater than 

breadth ; sacci distinctly to slightly smaller than 

body, slightly more than semicircular to almost circular in outline, narrowing toward roots. Dorsal 

attachment of sacci at equator or slightly beyond; 

sacci distally pendant, clearly set off from body, a 

distinct reentrant angle present where proximal 

roots of sacci join . the central body. Marginal ridge 

not present. Distal tenuitas 17 ,u-21,u wide. Wall of 
16 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

proximal cap l,u-2,u thick; columnar layer visible at 

X 1,000 magnification, tectate; surface of tectum 

smooth; infrareticulum very fine. Sacci irregularly 

reticulate, lumina 0.5,u-3,u in diameter. 

Observations on many specimens indicate the body 

of this species probably was originally nearly 

spherical. 

Size range (in microns) (terminology based on 

Pocock (1962, p. 59-60) .-

Length of bladders --------------------------- 42-71 

Length of central body ----------------------- 56-80 

Breadth of bladders -------------------------- 31-59 

Total breadth of grain _______________________ 100-153 

Height of central body ----------------------- 58-70 

Breadth of central body ---------------------- 52-104 

Comparison with similar species.-P. elongatus 

var. grandis is larger than Pityosporites alatipollenites (Rouse) Singh (1964, p. 123), and the ornamentation of the sacci of P. elongatus var. grand is 

is very distinct from that of the body, whereas 

Rouse (1959, p. 314) described the ornamentation 

of P. alatipollenites as more or lEss uniform on body 

and bladders. Except for its larger size, P. elongatus 

var. grand is resembles Pityosporites elongatus Norton n. comb. 

Genus PODOCARPIDITES Cookson ex Couper, 1953 emend. 

Potoni~, 1958 

Disaccites (Podocarpidites) Cookson, 1947, p. 131. 

[in part] Podocarpidites Cookson, 1947 ex Couper, 1953, p. 

35. 

Podocarpidites Cookson, 1947 ex Couper, 1953 emend. Potonie, 

1958, p. 68. 

Type species.-(designated by Couper, 1953, p. 

35) Podocarpidites ellipticus Cookson, 1947; lectotype (designated by Potonie, 1958, p. 68) in Cookson, 1947, p. 131, pl. XIII, fig. 6; Waterfall Gorge, 

Kerguelen Archipelago; Tertiary. 

Podocarpidites multesimus (Bolkbovitina, 1956) Pocock, 1962 

Plate 7, figures 1, 2 ; table 1, No. 89 

Podocarpus multesimus Bolkhovitina, 1956, p. 127. 

Podocarpidites biformis Rouse, 1957, p. 367, pl. II, fig. 13; 

south-central Alberta, Canada; Oldman Formation; Late 

Cretaceous. 

Podocarpidites multesimus (Bolkhovitina, 1956) Pocock, 1962, 

p. 67; western Canada; Mannville Formation; Early 

Cretaceous. 

Podoc,arpidites multesimus (Bolkhovitina) Pocock, 1962. In 

Singh, 1964, p. 116, pl. 15, figs. 12, 13; east-central 

Alberta, Canada; Mannville Group; Early Cretaceous. 

Holotype.-ln Bolkhovitina, 1956, p. 127, pl. 

XXIV, fig. 235; Vilyui Basin, U.S.S.R.; Early 

Jurassic. 

Remarks.-The specimens from the Judith River 

Formation samples which are here identified as P. 

multesimus also resemble Podocarpidites maximus 

(Stanley, 1965) Norton (in Norton and Hall, 1969, 

p. 31). P. maximus resembles P. multesimus and 

may be synonymous with it. 

Podocarpidites ornatus Pocock, 1962 

Podocarpidites ornatus Pocock, 1962, p. 67-68. 

Holotype.-In Pocock, 1962, p. 67-68, pl. 11, fig. 

164; western Canada; Quartz Sand Member, Mannville Group; Barremian. 

Podocarpidites cf. P. ornatus 

Plate 7, figure 3; table 1, No. 52 

Genus PRISTINUSPOLLENITES n. gen. 

Type species.-Pristinuspollenites (Alisporites) 

microsaccus (Coupe·r) (Pocock, 1962) n. comb. 

(Seep. 17.) 

Description.-Bisaccate pollen grains. In polar 

view outline of grain ellipsoidal to circular. Length 

of body equal to, slightly greater than, or slightly 

less than its breadth. Sacci distally attached (entirely on distal side of equator), clearly set off from 

body, very small in comparison to body, in fossilized 

state variously inclined-either leaning toward or 

away from the tenuitas; sacci on some specimens 

and in some species wrinkled or slightly ruffled. 

Tuberclelike thickenings never present at base of 

sacci (as in Dacrydiumites Cookson, 1953a). Distal 

sulcus clearly defined in some specimens. Body variously ornamented, sacci finely or indistinctly reticulate but never coarsely r_eticulate. 

Remarks.-Pristinuspollenites was erected to accommodate a group of distinctive bisaccate pollen 

species which have greatly reduced distally attached 

sacci and which do not have thickenings at the base 

of the sacci (as in Dacrydiumites) . 

Pristinuspollenites (Pteruchipollenites) microsaccus Couper, 1958 n. comb. was designated as the 

type species for the genus because it was an adequately described and illustrated species and has 

the characteristics that represent Pristinuspollenites. 

New combinations and new names: 

Pristinuspollenites (Bacubivesiculites) inchoatus 

(Pierce, 1961) n. comb. (Holotype in Pierce, 

1961, p. 34, pl. II, fig. 35; Minnesota; early 

Late Cretaceous. Also illustrated in this report: 

pl. 7, fig. 7.) 

Pristinuspollenites ( Clavabivesiculites inchoatus 

Pierce, 1961) crassus n. name. (Holotype in 

Pierce, 1961, p. 34-35, pl. II, fig. 36; Minnesota; 

early Late Cretaceous. Also illustrated in this 

report: pl. 7, figs. 8, 9.) The new species name 

crassus was here introduced because Pristinuspollenites (Clavabivesiculites) inchoatus 

(Pierce, 1961) n. comb. would have been a 
SYSTEMATICS 17 

later homonym of Pristinuspollenites (Bacubivesiculites) inchoatus. (See above.) 

Pristinuspollenites ( Clavabivesiculites) pannosus 

(Pierce, 1961) n. comb. (Holotype in Pierce, 

1961, p. 35, pl. II, fig. 37; Minnesota; early 

Late Cretaceous. Also illustrated in this report: 

pl. 7' fig. 10.) 

Pristinuspollenites ( Granabivesiculites inchoatus 

Pierce, 1961) sulcatus n. name. (Holotype in 

Pierce, 1961, p. 35, pl. II, fig. 38; Minnesota; 

early Late Cretaceous. Also illustrated in this 

report: pl. 7, fig. 11.) The new species name 

sulcatus was here introduced because Pristinuspollenites ( Granabiv esiculites) inchoa tus 

(Pierce, 1961) n. comb. would have been a 

later homonym of Pristinuspollenites (Bacubivesiculites) inchoatus (see above). 

Pristinuspollenites (Phyllocladidites) microreticulatus (Brenner, 1963) n. comb. (Holotype in 

Brenner, 1963, p. 79-80, pl. 30, fig. 3a-b; Maryland; Early Cretaceous.) 

Pristinuspollenites ( Parvisaccites) gra.nisaccus 

(Hedlund, 1966) n. comb. (Holotype in Hedlund, 1966, p. 22-23, pl. 6, fig. 6a; Oklahoma; 

Cenomanian.) This species appears to be very 

similar to, and is probably synonymous with, 

Pristinuspollenites sulcatus. (See above.) 

Remarks.-Norris (1967, p. 103) transferred 

Granabivesiculites inchoatus Pierce to Phyllocladidites Cookson ex Couper. This is not an acceptable 

transfer because Granabivesiculites incho·atus Pierce 

(holotype illustration, this report, pl. 7, fig. 11) 

does not have proximal tubercles and thus does not 

fit into Dacrydiumites Cookson, 1953 (synonym 

Phyllocladidites). In his description of the species, 

Norris did not mention if his specimens from central Alberta have proximal tubercles. There is no 

indication of tubercles at the base of the sacci in 

his specimens illustrated in his plate 15, figures 8, 9. 

Those specimens closely resemble and may be assignable to Pristinuspollenites 'microsaccus. 

Pocock (1962, p. 61) had transferred the species 

Pteruchipollenites microsaccus Couper to the genus 

Alisporites, a genus to which that small winged 

taxon does not conform. 

Comparison.-Pristinuspollenites n. gen. is distinguishable from Dacrydiumites by its lack of 

thickenings ("proximal tubercles" of Cookson, 1947, 

p. 133) at the base of the sacci. 

Pristinuspollenites, in common with Parcisporites 

Leschik (1956, p. 55), has strongly reduced sacci 

but differs in that its sacci are distinctly delineated 

from the body whereas in Parcisporites the zone or 

line of attachment of the sacci is irregular or indefinite. The sacci of Parcisporites are apparently 

attached much closer to the equator than are those 

of Pristinuspollenites. 

The sacci of Pristinuspollenites are not veillike 

as are those of Indusiisporites Leschik (1956, p. 55). 

Pristinuspollenites does not have radially arranged thickenings on the sacci as does Parvisaccites 

Couper (1958, p. 154). 

Distribution.-In addition to the species localities 

mentioned above (under new combinations and new 

names), forms assignable to this genus have been 

reported from several other parts of the Northern 

Hemisphere. Specimens that seem to conform to 

Pristinuspollenites were reported from the Cretaceous of the U.S.S.R by Bolkhovitina (1953, p. 74, 

pl. XI, figs. 5, 6) and by Chlonova (1960, p. 60-61, 

pl. IX, figs. 6, 7). Chlonova (1961, p. 67, 

pl. XI, fig. 70) assigned a specimen from the 

Cenomanian-Turonian of the Chulymn-Enisei depression of the U.S.S.R. to the species Pteruchipollenites microsaccus Couper. Dacrydium? transiens Sato (1961, p. 89-90, pl. I, fig. 23) from the 

Upper Cretaceous of Japan appears to be assignable 

to the genus Pristinuspollenites. 

Robert Tschudy (oral commun., March 1971) 

reported several forms assignable to Pristinuspollenites from Upper Cretaceous rocks from the 

Appalachian Mountains region of the United States. 

Pristinuspollenites microsaccus (Couper, 1958) n. comb. 

Plate 7, figures 4-6; table 1, No. 82 

Pteruchipollenites microsaccus Couper, 1958, p. 151, pl. 26, 

figs. 13, 14. 

Alisporites (Pteruchipollenites) microsaccus (Couper) Pocock, 1962, p. 61. 

Alisporites microsaccus (Couper, 1958) Pocock, 1962. In 

Burger, 1966, p. 259, pl. 35, figs. 1a, b; eastern Netherlands; Lias, Jurassic. 

Holotype.-In Couper, 1958, p. 151, pl. 26, fig. 13; 

Cambridge Observatory Borehole, southeastern England; Upper Estuarine Series; Middle Jurassic. 

Description (based on Couper's description and 

illustrations and on 11 specimens from Judith 

River Formation samples-all specimens in polar 

and oblique polar views) .-Bisaccate pollen grains. 

In polar view, outline of grain ellipsoidal to circular. 

Length of body equal to, slightly greater than, or 

slightly less than its breadth. Sacci distally attached, 

clearly set off from body, much smaller than body ; 

sacci arc-shaped, in some specimens one or both 

sacci wrinkled or distorted ; maximum length of 

sacci at their base, length of sacci greater than their 

breadth; sacci variously inclined-either leaning 

toward or away from the tenuitas, appearing flattened or slightly inflated. Proximal tubercles not 

present at base of sacci. Distal sulcus not clearly 
18 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

defined, in most specimens not distinguishable. 

Proximal cap 1,u-1.5fL thick, sculpture scabrate. Body 

wall slightly finer textured and appearing thinner 

in area between bladders than on proximal cap. 

Sculpture of sacci finely and indistinctly reticulate. 

Sacci in reclined position covering as much as half 

the distal surface of grain; in true polar position 

of grains sacci not contributing to outline of grain. 

Size range (in microns) .- Combination of 

specimens 

(Couper and 

Couper T.~chudy Tschudy) 

Length of central body _ 55-78 37-56 37-78 

Breadth of central body_ 30-58 30-46 30-58 

Length of sacci -------- 53-75 25-46 25-75 

Breadth of sacci _______ 18-30 7-13 7-30 

Total breadth of grain _ 35-63 30-46 30-63 

Rema.rks.-My specimens from the Judith River 

Formation of Montana, except that many of them 

are smaller, closely resemble those of P. microsaccus 

as it was described and illustrated by Couper. 

Comparison.-Pristinuspollenites sulcatus n. name 

(see p. 17) resembles Pristinuspollenites microsaccus n. comb. but is distinguishable because of its 

broad and clearly defined sulcus. P. sulcatus also 

tends to have a thicker proximal cap than P. 'microsaccus and its sacci tend to be narrower at their 

roots. 

Genus RUGUBIVESICULITES Pierce, 1961 

Rugubivesiculites Pierce, 1961, p. 39. 

Type species.-Rugubivesiculites convolutus 

Pierce, 1961, p. 39-40, pl. II, fig. 57; Minnesota; 

early Late Cretaceous. 

Rugubivesiculites sp. 

Plate 7, figur·es 12, 13; table 1, No. 15 

Remarks.-These specimens resemble photographs 

of Pinus ruginosa Stanley (1965, p. 277, pl. 39, figs. 

1-7) but apparently are different because Stanley 

described P. ruginosa as having a finely to moderately reticulate cap and my specimens from the 

Judith River Formation from Montana have a finely 

rugulate cap. 

My specimens also resemble Rugubivesiculites 

minimus Burger (1966, p. 257, pl. 33, fig. 3), except 

that their proximal cap seems to be more finely 

ornamented. 

Genus VITREISPORITES Leschik, 1955 sensu Potonie, 1958 

Vitreisporites Leschik, 1955, p. 53. 

Vitreisporites Leschik, 1955. In Potonie, 1958 (January), p. 

56. 

Caytonipollenites Couper, 1958 (March), p. 149. 

Caytoniales-Pollenites Pautsch, 1958 (July), p. 323. 

Vitreisporites Leschik. In Nilsson, 1958 (November), p. 77-

78. 

Vitreisporites Leschik, 1955 emend. Jansonius, 1962 (April), 

p. 55. 

Vitreisporites (Leschik) Jansonius. In Pocock, 1962 (October), p. 58. 

Vitreisporites (Le.schik) Jansonius emend. De Jersey, 1964, 

p. 9. 

Vitreisporites Leschik, 1955, emend. Jansonius, 1962. In 

Singh, 1964, p. 102. 

Type species.-Vitreisporites signatus Leschik, 

1955, p. 53-54; pl. 8, fig. 10; Neuewelt near Basel, 

Switzerland; Keuper, middle Late Triassic. 

Rema;rks.-Vitreisporites was described by Leschik (1955, p. 53) as having a w•eak Y-mark on 

the proximal pole. Potonie (1958, p. 56) and Nilsson 

(1958, p. 77-78) in their descriptions of Vitreisporites did not omit the Y -mark distinction but included 

within the genus those forms not having, as well 

as those having, a Y-mark. Jansonius (1962, p. 55) 

in his emendation of Vitreisporites omitted any 

designation of a Y-mark because he was of the 

opinion that a Y -mark had not been demonstrated 

convincingly. Pocock (1962, p. 58) and Singh (1964, 

p. 102) also omitted recognition of the Y -mark from 

their descriptions of Vitreisporites. De Jersey (1964, 

p. 9) reemended Vitreisporites (Leschik) Jansonius, merely expanding the genus to allow for the 

inclusion of grains with little, if any, ornamentation on the bladders. I accept Vitreisporites (Leschik) sensu Potonie in preference to Vitreisporites 

(Leschik) Jansonius because I do not believe the 

Y -mark designation should be excluded from the 

genus description when it was an important characteristic of the genus and of its type species as 

they were originally described by Leschik. 

Vitreisporites pallidus (Reissinger} Nilsson, 1958 

Plate 8, figures 1, 2; table 1, No. 53 

Pityosporites pallidus Reissinger, 1939, p. 14. 

Pityopollenites pallidus (Reissinger) Reissinger, 1950, p. 

109, pl. XV, figs. 1-5. (Pityopollenites used as a new 

genus but not described.) 

Pityosporites pallidus Reissinger. In Balme, 1957, p. 36-37, 

pl. 10, figs. 112, 113; Western Australia; Mesozoic. 

Caytonipollenites pallidus (Reissinger) Couper, 1958, p. 150, 

pl. 26, figs. 7, 8; Yorkshire, England; Middle Jurassic. 

Isle of Wight, England; Aptian. 

Vitreisporites pallidus (Reissinger) Nilsson, 1958, p. 78-79, 

pl. 7, figs. 12-14; Sweden; Lias. 

Caytonipollenites pallidus (Reissinger) Couper. In Lantz, 

1958, p. 926, figs. 46, 4 7; Dorset, Great Britain; Jurassic. 

Vitreisporites pallidus (Reissinger) Nilsson. In Pocock, 1962, 

p. 58-59, pl. 9, figs. 134, 135; westeTn Canada; Deville 

Member and quartz sand member of Mannville Group; 

Early Cretaceous. 

Vitreisporites pallidus (Reissinger) Nilsson, 1958. In Balme, 

1963, p. 24-25, pl. 6, fig. 7; western Australia; Kockatea 

Shale; Early Triassic. 

Vitreisporites pallidus (R·eissinger) Brenner, 1963, p. 74, pl. 

25, figs. 1, 2; Maryland; Patapsco and Patuxent Formations of Potomac Group; Early Cretaceous. 
SYSTEMATICS 19 

Vitreisporites pallidus (Reissinger) Nilsson, 1958. In Singh, 

1964, p. 102-103, pl. 14, fig. 1; east-central Alberta, 

Canada; Grand Rapids Formation of Mannville Group; 

Albian. 

Caytonipollenites pallidus (Reissinger) Couper. In Leop'old 

and Pakiser, 1964, p. 82, 88, pl. 6, figs. 33, 34; western 

Alabama; Eutaw Formation; Late Cretaceous. 

Vitreisporites pallidus (Reissinger) Nilsson, 1958. In Playford and Dettmann, 1965, p. 156, pl. 17, fig. 58; South 

Australia; Rhaetic-Liassic. 

Vitreisporites pallidus (Reissinger) Brenner. In Habib, 1969, 

pl. 2, fig. 11; deep-sea core, northeast of Cat Island, the 

Bahamas, West Indies; middle Cretaceous. 

Lectotype.-(here assigned) In Reissinger (1950, 

pl. XV, fig. 5; Teufelsgraben near Altdorf, east 

from N tirnberg, Germany; Lias; Early Jurassic. 

Remarks.-Grains resembling Vitreisporites p.a.Zlidus have been reported from Cretaceous deposits of 

Russia by Bolkhovitina (1953, p. 72-73, pl. XI, fig. 

1) and by Chlonova (1961, p. 65, pl. XI, figs. 66, 67). 

Description (based on 30 specimens) .-Bisaccate 

pollen grains, shape oval in polar view; total breadth 

of grain 25t-t-38t-t, length of bladders 11rt-21t-t, 

breadth of bladders 7.5p:-16t-t, length of central 

body 14p:-21t-t, breadth of central body 7 t-t-16t-t; sulcus not discernible in specimens examined ; exine 

of bladders approximately 0.5t-t thick, sculpture 

finely and irregularly reticulate; exine of body 

thinner than that of bladders, sculpture smooth to 

granulate. 

Similar species.-Vitreisporites pallidus resembles Vitreisporites signatus Leschik but differs in 

that it does not have a Y -mark on the proximal pole. 

MISCELLANY 

Genus T AXODIACEAEPOLLENITES Kremp, 1949 ex Potonie, 1958 

Taxodiaceaepollenites Kremp, 1949 ex Potonie, 1958, p. 78-79. 

Type species.-Taxodiacea.epollenites (Pollenites) 

hiatus (Potonh~) Kremp, 1949 ex Potonie, 1958. 

Taxodiaceaepollenites hiatus (PotonHi) Kremp, 1949 ex Potonie, 1958 

Plate 8, figures 3, 4; table 1, No. 18 

Pollenites hiatus Potonie, 1931c, p. 5. 

Taxodium hiatipites Wodehouse, 1933, p. 493-494, fig. 19 (not 

fig. 17 as stated). (Wodehous.e compared this species 

with species of Taxodiwn, Juniperus, Libocedrus, Thuja, 

and Zamia). 

Taxodieae-Poll. hiatus Potonie. In Thiergart, 1940, p. 30. 

Taxodiaceae-Poll. hiatus Potonie. In Kremp, 1949, p. 59. 

Taxodiaceae-Poll. hiatus Potonie. In Kremp, 1949, p. 91. 

Taxodoidites hiatus Potonie. In Potonie, Thomson, and Thiergart, 1950, p. 49. (In this :veport also spelled Taxodioidites.) 

Taxodioipoll. hiatus Potonie. In Potonie, 1951, p. 140, 149. 

Inaperturopollenites hiatus (Potonie) Thomson and Pflug, 

1953, p. 65. 

Taxodiaceaepollenites kiatus (Potonie, 1931) Kremp, 1949. 

In Potonie, 1958, p. 79. 

Taxodieaepollenites hiatus (Potonie, 1931). In Kremp and 

Gerhard, 1961, p. 265. 

Thuja? hiatus (Potonie) Stanley, 1965, p. 273-274. 

Holotype.-In Potonie, 1931c, p. 5, fig. 27; Ville 

bei Koln, Beisselsgrube; (Oligocene) Miocene. 

Desc1·iption.-Pollen grains, probably originally 

spherical but characteristically split from one-half 

to nearly total diameter of grain; the split varying 

from narrow V-shaped to broadly gaping. Exine 

approximately 0.5t-t thick, wall layers not distinguishable, surface smooth to scabrate. 

Remarks.-The species first described as Pollenites hiatus Potonie ( 1931c, p. 5) has since been 

included in combination with a succession of valid 

and invalid names. 

Thiergart (1940, p. 30) used the combination 

Taxodieae-Poll. hiatus but obviously did not intend 

that as a formal binomial name, because in his illustrations he referred to the grains as "Taxodium," 

"cf. Taxodiu·m," and "Taxodieae" rather than as 

Taxodieae-Poll. hiatus. Furthermore, Taxodieae-Poll. 

Thiergart, 1940, could not be considered a valid name 

under the provision of Article 42 of the "International Code of Botanical Nomenclature" (Lanjouw, chm., 

and others, 1966) because the taxon hiatus was not a 

new species and because Ta.xodie.ae-Poll. Thiergart 

was not monotypic-Thiergart in the same report 

also described the new form "Poll. hiatus R. Pot. 

paleocaenicus n.f." 

Kremp ( 1949, p. 59) used the combination Taxodiaceae-Poll. hiatus but it is not clear whether he 

intended Taxodiaceae-Pollenites as a generic name 

because he (p. 91) merely placed Pollenites hiatus 

in the family Taxodiaceae. At any rate, TaxodiaceaePoll. Kremp, 1949, was not valid. The species hiatus 

was not new and consequently Taxodiaceae-Pollenites could not have been valid under the provision of 

Article 42 (Lanjouw, chm., and others, 1966). 

Potonie, Thomson, and Thiergart (1950, p. 49) 

used the combination Taxodoidites hiatus (probably 

a misprint, as this was under the heading Taxodioidites). They did not describe Taxodoidites (or 

Taxioidites) or designate it as a new genus. 

Potonie (1951, p. 140, 149) used the combination 

Taxodioipoll. hiatus but did not describe it. 

The first valid new combination was that in 

Thomson and Pflug (1953, p. 64, 65)-where the 

new genus Inaperturopollenites Pflug and Thomson 

was described and the species hiatus Potonie was 

formally transferred to it. 

Taxodiaceaepollenites became a valid name when 

Potonie (1958, p. 78-79) used it as a generic name, 

described it, and designated Taxodiaceaepollenites 
20 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

(al. Pollenites) hiatus (Potonie, 1931) Kremp, 1949 

as its type species ( Genotypus) . 

Kremp and Gerhard (1961, p. 265) considered 

Taxodiaceae-pollenites hiatus (Potonie, 1931) 

Kremp, 1949, a synonym of Ta.xodieae-pollenites 

hiatus (Potonie, 1931) Theirgart, 1940. It is true, 

"Taxodieae-Poll." was the first name other than 

Pollenites to be used in combination with the species 

hiatus but it was not a valid name. 

Stanley (1965, p. 273-274) transferred the species 

hiatus to Thuja? This combination is not acceptable 

because "Thuja ?" is not a proper generic name. 

In the present report, Ta.xodiaceaepollenites is 

considered to be a form genus because I believe 

that it cannot be assigned with certainty to the 

family Taxodiaceae. Taxodiaceaepollenites as defined by Potonie ( 1958, p. 79), is clearly distinguishaJble from lnaperturopollenites Pflug and Thomson 

as emended by Potonie (1958, p. 77-78). 

Genus SEQUOIAPOLLENITES Thiergart, 1937 

Sequoia-pollenites Thiergart, 1937, p. 301-302. 

Sequoiapollenites Thiergart, 1937. In PotoniE~, 1958, p. 79-80. 

Sequoiapollenites Thiergart. In Manum, 1962, p. 43-44. 

Type species.- Sequoiapollenites polyformosus 

Thiergart, 1937, p. 301-302, pl. 23, fig. 6 (lectotype, 

selected by Potonie, 1958, p. 79) ; Senftenberg, Germany; Miocene. 

Sequoiapollenites spp. 

Plate 8, figures 5, 6; table 1, No. 90 

Genus CLASSOPOLLIS Pflug, 1953 emend. Pocock and Jansonius, 1961 

Classopollis Pflug, 1953, p. 91. 

Classopollis Pflug, 1953 emend. Pocock and J ansonius, 1961, 

p. 443. 

Classopollis (Pflug, 1953) Pocock and Janson ius, 1961. In 

Potonie, 1966, p. 130-132. 

Type species. - Classopollis classoides Pflug 

emend. Pocock and Jansonius, 1961, p. 443-444; 

holotype in Pflug, 1953, p. 91, pl. 16, figs. 29-31 

(three levels) ; Siegelsum, Germany; Lias, Early 

Jurassic. 

Classopollis spp. 

Plate 8, figures 7, 8; table 1, No. 54 

Genus EQUISETOSPORITES Daugherty, 1941 emend. Singh. 1964 

Equisetosporites Daugherty, 1941, p. 63. 

Equisetosporites Daugherty, 1941 emend. Singh. 1964, p. 129. 

Equisetosporites Daugherty emend. Singh, 1964. In Srivastava, 1968b, p. 216. 

Type species.-Equisetosporites chinleana Daugherty, 1941, p. 63, pl. 24, fig. 4; Arizona; Late Triassic. 

Equisetosporites spp. 

Plate 8, figures 9, 10; table 1, No. 19 

MONOSULCATES 

Genus ARECIPITES Wodehouse, 1933 

Arecipites Wodehouse, 1933, p. 497. 

Arecipites Wodehouse, 1933. In Potonie, 1958, p. 97. 

Type species.-(selected by Potonie, 1958, p. 97) 

Arecipites punctatus Wodehouse, 1933, p. 497, fig. 

22; Colorado or Utah; Green River Formation; 

Eocene. 

Arecipites sp. 

Plate 8, figure 11; table 1, No. 21 

Genus LILIACIDITES Couper, 1953 

Liliacidites Couper, 1953, p. 56. 

Retimonoc,olpites Pierce, 1961, p. 47. 

Type species.-Liliacidites kaitangataensis Couper, 1953, p. 56, pl. 7, fig. 97 (holotype) ; Kaitangata 

Coalfield, New Zealand; sub-W angaloan; Late Cretaceous. 

Liliacidites variegatus Couper, 1953 

Plate 8, figure 12; table 1, No. 55 

Liliacidites variegatus Couper, 1953, p. 56. 

Liliacidites variegatus Couper, 1953. In Rouse, 1957, p. 368, 

pl. II, figs. 29, 30; south-central Alberta, Canada; Oldman Formation; Santonian. 

Liliacidites variegatus Couper, 1953. In Srivastava, 1966, p. 

525, pl. IV, figs. 15, 16, Scollard, Alberta, Canada; Edmonton Formation; Maestrichtian. 

Liliacidites variegatus Couper, 1953. In Elsik, 1968a, p. 312, 

pl. XV, fig. 1; Milam County, Tex.; Rockdale Formation; 

Paleocene. 

Liliacidites variegatus Couper, 1953. In Srivastava, 1969b, p. 

977, pl. I, fig. 10; Alberta, Canada; Edmonton Formation; Maestrichtian. 

Holotype.-In Couper, 1953, p. 56, pl. 7, fig. 99; 

Mataura Valley, Southland, New Zealand; Landonian, early to middle Oligocene. ( Holoty,pe also 

illustrated in Couper, 1960, pl. 9, fig. 9.) 

Liliacidites morrinensis Srivastava, 1969b 

Liliacidites nwrrinensis Srivastava, 1969b, p. 977. 

Holotype.-In Srivastava, 1969b, p. 977, pl. I, fig. 

9; near Morrin, Alberta, Canada; Edmonton Formation ; Maestrichtian. 

Liliacidites cf. L. morrinensis 

Plate 8, figure 13; table 1, No. 95 

Liliacidites complexus (Stanley, 1965) Leffingwell, 1971 

Plate 8, figures 14, 15; table 1, No. 20 

Schizosporis complexus Stanley, 1965, p. 267-268, pl. 36, figs. 

7-17. 

Schizosporis complexus Stanley. In Norton and Hall, 1969, 

p. 58, pl. 3, fig. 9; Montana; Fox Hills Sandstone; 

Maestrichtian. 

Liliac,idites complexus (Stanley, 1965) Leffingwell, 1971, p. 

41, pl. 7, fig. 3; Wyoming; Lance Formation; Late Cretaceous. 

Holotype.-In Stanley, 1965, p. 267-268, pl. 36, 
SYSTEMATICS 21 

figs. 7-9 (three levels) ; Crow Butte, Harding 

County, S. Dak.; Hell Creek Formation; Maestrichtian. 

Description (based on 30 specimens from USGS 

paleobotany locality samples from the Judith River 

Formation of Montana) .-Monosulcate, bilateral 

pollen grains; shape oval to almost circular; size 

221-"-33.51-" X 27.5f-t-42f-t. Aperture: Sulcus slitlike; 

as seen in distal view. of grain sulcus extends approximately three-fourths the total length of grain· 

in some views of specimens the sulcus is not clearl; 

visible; in some specimens a narrow smooth margo 

is present adjacent to sulcus. Pollen wall : Endexine 

thin; ektexine 11-"·-1.51-" thick, surface sculpture 

reticulate; lumina of reticulum 1v-3fL in diameter, 

lumina smaller on distal surface, very small near 

sulcus; muri as much as 0.5f-t wide, duplibaculate. 

Remarks.-The specimens described above vary 

considerably in size, shape, and coarseness of surface ornamentation; however, none of my specimens 

from the Judith River Formation were as large or 

as coarsely reticulate as one illustrated by Stanley 

(1965, pl. 36, figs. 13, 14). Some of my specimens 

are more finely reticulate than Stanley's illustrated 

specimens, but most of th€m are very similar to 

his holotype specimen (Stanley, 1965, pl. 36, figs. 

7-9). 

Liliacidites spp. 

Plate 8, figures 16-18; table 1, No. 56 

Genus OVOIDITES Potoni~ ex Krutzsch, 1959a emend. Potonie, 1966 

Ovoidites Potonie, 1956, ex Krutzsch, 1959a, p. 249. 

Ovoidites Potonie, 1951 emend. Potonie, 1966, p. 201-202. 

Type species.-Ovoidites ligneolus (Pontonh~) 

1951 ex Krutzsch, 1959a. 

Ovoidites ligneolus (Potonie, 1931b) Potonie, 1951 ex Krutzsch, 1959a 

Plate 8, figure 19; table 1, No. 69 

Pollenites ( ?) Ugneolus Potonie, 1931b, pl. 2, fig. V25a. 

Sporites ligneolus ( Potonie) Potonie and V enitz, 1934, p. 15, 

Ovoidites ligneolns (Potonie) Potonie, 1951, pl. XXI, fig. 185. 

(Not a valid name, species not new and genus not described.) 

Ovoidites ligneolus, Potonie. In Thomson and Pflug, 1953 p. 

113. ' 

Ovoidites ligneolus ( Potonie) Potonie, 1951 ex Krutzsch, 

1959a, p. 249. (Valid name because genus Ovoidites here 

described.) 

Holotype.-ln Potonie, 1931b, pl. 2, fig. V25a; 

Niederrheim-Bucht; Miocene. 

TRICOLPA TES 

Genus AESCULIIDITES Elsik, 1968b 

Aesculiidites Elsik, 1968b, p. 632. 

Type species.-Aesculiidites (T1·icolpites) circumst1·iatus (Fairchild) Elsik, 1968b. In Stover, Elsik, 

494-146 0- 73 - 4 

and Fairchild, 1966, p. 5, pl. 2, figs. 3-7 (holotype 

fig. 3a-c); Milam County, Tex.; Wilcox Group; 

early Eocene. 

Aesculiidites dubius (Jardine and Magloire, 1965) n. comb. 

Plate 8, figures 20, 21; table 1, No. 22 

Striopollenites dubius Jardine and Magloire, 1965, p. 215. 

Holotype.-In Jardine and Magloire, 1965, p. 215, 

pl. X, fig. 50; Senegal, northwestern Africa; late 

Albian. 

Remarks.-The genus Striopollenites Rouse (1962, 

p. 212), as already pointed out by Potonie ( 1966, 

p. 164), is synonymous with Striatopollis Krutzsch 

(1959b, p. 142-143). However, the holotype of 

Striopollenites dubius as shown in the illustration 

(Jardine and Magloire, 1965, pl. X, fig. 50) has 

striations approximately parallel with the equator 

of the grain and so pertains to the genus Aesculiidites Elsik ( 1968b) which has circumstriate ornamentation, rather than to Striatopollis which has 

longitudinal striations. 

Some of my specimens found in samples of the 

Judith River Formation are slightly smaller 

(range 16~-"-21!-L) than Jardine and Magloire's 

specimens (range 181-L-25!-L) but otherwise they 

closely resemble their holotype specimen. Their 

plate X, figures 48 and 49, illustrate specimens assigned to Striopollenites dubius that appear to have 

longitudinal striations; these specimens probably are 

a different species and belong in the genus Striatopollis Krutzsch (1959b). 

Genus GREWIPOLLENITES Srivastava, 1969d 

Grewipollenites Srivastava, 1969d, p. 60-61. 

Type species.-Grewipollenites canadensis Srivastava, 1969d, p. 61, pl. 2, fig. 27; Alberta, Canada; 

Edmonton Formation; Maestrichtian. 

Grewipollenites radiatus n. sp. 

Plate 8, figures 22-25; table 1, No. 94 

Holotype.-USGS paleobotany locality D3726-D, 

slide (2) at 104.7 X 18.8; plate 8, figure 22; center 

NE1;4 sec. 31, T. 23 N., R. 17 E.; Fergus County, 

Mont.; Judith River Formation (lower marine part) 

late Campanian. 

Paratype.-USGS paleobotany locality D3726-D, 

slide (2) at 86.0 X 15.3; plate, 8, figure 23; same 

locality data as for holotype. 

Paratype.-USGS paleobotany locality D3726-D, 

slide (2) at 88.1 X 4.1; plate 8, figure 24; same locality data as for holotype. 

Description (based on 28 specimens) .-Tricolpate 

pollen grains; outline circular to rounded triangular; diameter 14t-L-20fL. Apertures: Colpi extending 

one-half to two-thirds the distance to poles, slitlike 
22 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

or gaping slightly, usually visible only in specimens 

that are oriented in polar view; pores in addition 

to colpi are suggested in some specimens, but definite 

pores have not been observed. Pollen wall: Walls 

less than 0.5/L thick, ornamented with ridges that 

diverge from several foci on surface of grain; 

ridges 1.5/£-3/L long, as much as 1/L high, less 

than 1/L wide at widest part, and tapered to sharp 

or blunt ends. 

C01nparison.-Specimens that I have described 

under the new species name Grewipollenites racliatus 

are smaller and apparently have slightly coarser 

surface ornamentation than Grewipollenites canadensis Srivastava; however, G. radiatus may later 

be found to merge with G. canadensis. 

Genus PULCHERIPOLLENTES Srivastava, 1969d 

Pulcher1'pollenites Srivastava, 1969d, p. 61. 

Ty]Je species.-Pulcheripollenites narcissus Srivastava, 1969d, p. 61-62, pl. 2, figs. 28, 29; Alberta, 

Canada; Edmonton Formation; Maestrichtian. 

Pulcheripollenites krempii Srivastava, 1969d 

Plate 8, figures 26-28; table 1, No. 23 

Pulcheripolleni.tes k1·empii Srivastava, 1969d, p. 62. 

Holotype.-In Srivastava, 1969d, p. 62, pl. 2, figs. 

30, 31; Alberta, Canada; Edn1onton Formation; 

Maestrichtian. 

Description (based on 26 specimens in polar and 

oblique views and four in equatorial view) .-Tricolpate pollen grains, radially symmetrical, angulaperturate, isopolar; outline in polar view nearly circular, to triangular with rounded corners; outline in 

equatorial view elliptical to rounded rhomboidal; 

equatorial diameter 27 tL-491-L, polar axis 27 /L-421t, 

polar axis 27 /L-42/L. Apertures (as seen in polar view 

of grains) : Colpi short, simple, slitlike to gaping, 

reaching one-fourth to one-third the distance to 

poles. Pollen wall : Endexine in most specimens less 

than 0.5/L thick, in some specimens as much as 

0.5/L thick near colpi where endexine is slightly 

thicker than on remainder of grain. Ektexine 0.5/L1.5/L thick, thickening to as much as 2/L near apertures. Endexinal and ektexinal thickening of wall 

begins approximately 10/L from colpi and continuEs 

to colpal slits-in this area tops of muri sheathed 

by a tectum as much as 0.5fL thick. Ektexinal surface ornamentation striate-reticulate, fingerprintlike, consisting of elongate, somewhat parallel and 

variously oriented muri which are connected at 

right angles by short muri that have slightly lower 

relief; the elongate muri are less than 0.51t to 11-L 

high, approximately 1,4/L wide, with rounded to 

pointed tops, and approximately as far apart as 

their width. 

Remarks.-The size range (equatorial diameter 27 tL-49/L) of my specimens of P. krempii from 

the Judith River Formation samples is slightly 

broader than that of P. krempii (equatorial diameter 35/L-42/L) as described by Srivastava. Pulche1·ipollenites narcissus Srivastava (1969d, p. 61, pl. 2, 

figs. 28, 29), except for its larger size, appears to 

be very similar to P. kren~pii; and possibly, when 

more specimens are found, these two species will 

merge. 

Genus TRICOLPITES Cookson ex Couper, 1953 emend. Potoni~, 1960 

[In part] Tricolpites Cookson ex Couper, 1953 (August), p. 

61. 

Tricolpites (Erdtman, 1947, Cookson, 1947, Ross, 1949) 

Couper, 1953 emend. Potonie, 1960, p. 95. 

[In partl Tricolpites (Cookson, 1947 ex Couper, 1953) 

emend. Belsky, Boltenhagen, and Potonie, 1965, p. 75. 

[In part] Tricolpites (Couper, 1953) Belsky, Boltenhagen, 

and Potonie, 1965 emend. Boltenhagen, 1967, p. 344, 346. 

Tricolpites Cookson ex Couper emend, Potonie, 1960. In 

Srivastava, 1969d, p. 55-56. 

Type Species.- (designated by Couper, 1953, p. 

61) Tricolpites Teticulatus Cookson, 1947, p. 134, 

pl. XV, fig. 45; Kerguelen Archipelago, Indian 

Ocean; early Tertiary. 

Description (not intended as an emendation).-

Isopolar, tricolpate pollen grains. Shape in polar 

view three lobed, rounded angular, or nearly circular. Colpi slitlike to broadly gaping, with or without 

distinct margines. Surface of grains finely reticulate 

(or finely areolate); lumina of reticulum less than 

1/L in diameter. (See Srivastava, 1969d, p. 55-56.) 

Remarks.-Tricolpites, though validly published 

by Couper in August 1953, was virtually synonymous with the broadly circumscribed genus Tricolpopollenites Pflug and Thomson (in Thomson 

and Pflug, 1953, p. 95) which was validly published 

in March 1953. The genus Tricolpite.s was reinstated 

by Potonie in 1960, when it was restricted to 

spherical to ovoid isopolar tricolpate forms having 

bulged (convex) mesocolpia, wide furrows with 

thickened margins, a more or less three-lobed equator, and finely reticulate exine. Belsky, Boltenhagen, 

and Potonie (1965, p. 75) reemended and expanded 

Tricolpites to include strongly reticulate forms 

without thickened colpal margins and with spheroidal to ball shape. Boltenhagen (1967, p. 346) recommended emending Tricolpites to take in striate 

grains as well as those of other sculptures. Tricolpites as emended by Belsky, Boltenhagen, and Potonie 

and as recommended by, Boltenhagen is not clearly 

defined and would overlap other more exactly defined 

tricolpate pollen genera. In this report Tricolpites 

sensu Potonie, 1960 is accepted. 
SYSTEMATICS 23 

Tricolpites reticulatus C1>okson, 1947 

Plate 8, figures 29-31; table 1, No. 24 

Tricolpites reticulata Cookson, 1947, p. 134. 

Tricolpites reticulatus Cookson, 1947. In Couper, 1953, p. 61. 

[not] Retitricolpites microreticulatus (Van der Hammen) 

Van der Hammen and Wymstra, 1964, p. 234, pl. II, fig. 

1. 

Tricolpites microreticulatus Belsky, Boltenhagen, and Potonie, 1965, p. 75-76, pl. 12, figs. 8, 9; Mandji-Halhinsel, 

Gabun, Equatorial Africa, Late Cretaceous. (According 

to a written communication from the publisher, the date 

of issue of the "PaHiontologieal Zeitschrift" in which 

this species was published wa::; July 9, 1965). 

Tricolpites interangulus Newman, 1965, p. 10-11, pl. 1, fig. 

3; Routt County, Colo.; basal. part of Williams Fork 

Formation; Campanian. (According to a written communication from the publisher, the date of release of 

the publication in which this species was published was 

July 22, 1965.) 

Tricolpites microreticulatus Belsky, Boltenhagen, and Potonie, 1965. In Srivastava, 1966, p. 548, pl. VII, fig. 24, 

Scollard, Alberta, Canada; Maestrichtian. 

[not] Tricolpites reticulatus Cookson, 194 7. In Srivastava, 

1966, p. 547, pl. VII, figs. 17-19, 26; Scollard, Alberta, 

Canada; Maestrichtian. 

Retitricolpites microreticulatus (Belsky, Boltenhagen, and 

Potonie) Brenner, 1968, p. 36·6, pl. VIII, fig. 5; northeastern Peru; Turonian, middle Cretaceous. 

Tricolpites reticulatus Cookson. In Norton and Hall, 1969, p. 

43-44, pl. 7, fig. 1; Garfield County, Mont.; Hell Creek 

Formation; Late Cretaceous. 

Tricolpites reticulatus Cookson, 1947. In Oltz, 1969, p. 141; pl. 

41, fig. 103; east-central Montana; Fox Hills Sandstone and Hell Creek Formation; Maestrichtian; Tullock Member of Fort Union Formation; Paleocene. 

Gunnera mic.roreticulata (Belsky, Boltenhagen, and Potonie, 

1965) Leffingwell, 1971, p. 37, pl. 6 figs. 7, 8; eastcentral Wyoming; Lance Formation, Maestrichtian. 

Holotype.-ln Cookson, 1947, p. 134, pl. XV, fig. 

45; Kerguelen Archipelago, Indian Ocean; early 

Tertiary. 

Remarks.-Pollen grains from the Oldman Formation (Upper Cretaceous) of western Canada called 

Tricolpopollenites liblarensis (Thomson) Thomson 

and Pflug by Rouse (1957, p. 367, pl. III, figs. 62, 

63) are also probably equal to the species T. reticulatus. 

T. reticulatus Cookson (1H47) ; T. microreticulatus Belsky, Boltenhagen, and Potonie ( 1965) ; and 

T. interangulus Newman (1965) are characterized 

by deeply cut interangular colpi, round to rounded 

triangular shape (in polar view), and finely reticulate sculpture-the original descriptions and illustrations of these species indicate that they are not 

clearly distinguishable from each other. I believe 

they should be considered synonymous species until 

or unless they can be shown by electron microscopy, 

or by other means, to be different from each other. 

Leffingwell's (1971, p. 37, pl. 6, figs. 7, 8) specimens from the Lance Formation of east-central 

Wyoming which he assigned to Gunnera r;nicroreticulata (Belsky, Boltenhagen, and Potonie, 1965) 

Leffingwell appear to be extremely fine textured and 

this appearance in part at least is probably due to 

the photographic method used. For example, in our 

laboratory, specimens of T. reticulatus that have 

been photographed under high dry magnification 

appear to be more finely sculptured and have less 

distinctly defined texture than the same specimens 

photographed under high-magnification oil immersion. 

Description.-Tricolpate isopolar pollen grains; 

shape in polar view three lobed, rounded triangular, 

or nearly circular; outline in equatorial view ellipsoidal to nearly circular; equatorial diameter 21t-L32fL, polar axis 23ft-28f-t. Apertures (as seen in polar 

view of grains) : Col pi extending approximately 

two-thirds the distance to poles; slitlike, V -shaped, 

or broadly gaping; bordered by narrow margines 

(in some specimens margines are indistinct) ; col pi 

inter angular, or (on nearly circular specimens) 

symmetrically arranged around equator. Pollen wall 

(as observed under x 1000 oil immersion magnification) : Wall 0.5/-L-1/L thick, slightly thicker in 

polar areas (as seen in equatorial view of grains) . 

Wall at equator of most specimens slightly thicker 

near col pi (as seen in polar view of grains) . Ektexine columnar, tectate, two to three times thicker 

than endexine. Ektexinal sculpture finely reticulate, 

lumina less than 0.51-L in diameter. (It would be 

desirable to examine this species with the aid of a 

scanning electron microscope. The surface sculpture 

appears to be finely reticulate at high focal level, 

but it is possible that this is a negative reticulum.) 

Surface of grains under high dry magnification appearing smooth to slightly rough. Some specimens 

(in polar view) having distinct and variously 

shaped, or indistinct darkened polar areas. 

Remarks.-T. reticulatus is a variable species and, 

on the basis of differences in size, shape of grains, 

coarseness of ornamentation, and presence or absence of polar thickenings (dark areas), it might 

conceivably be subdivided into several different 

species or varieties. However, observations of many 

specimens from a wide range of localities and ages 

(Trinidad Sandstone of northeastern New Mexico, 

Mitten Black Shale Member of Pierre Shale and 

Lance Formation of east-central Wyoming, Hell 

Creek Formation of northeastern Montana, Judith 

River Formation of north-central Montana, and 

basal part of Williams Fork Formation of northwestern Colorado (Newman's type locality of T. 

interangulus)) have shown that the morphological 
24 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MON'f. 

characteristics intergrade to such an extent that a 

subdivision does not seem to be feasible. I have also 

been unable to correlate any of the morphological 

characteristics with geographic locality or geologic 

age. 

Tricolpites hians Stanley, 1965 

Plate 8, figures 32-34; table 1, No. 25 

Tricolpites hians Stanley, 1965, p. 321-322. 

Tricolpites parvus Stanley, 1965, p. 322, pl. 4 7, figs 28-31. 

Holotype.-In Stanley, 1965, p. 321-322, pl. 47, 

fig. 24; Harding County, S. Dak.; Ludlow Member 

of Fort Union Formation; Paleocene. 

Description.-Tricolpate isopolar pollen grains; 

outline in polar view nearly circular, with three 

evenly spaced V -shaped to widely gaping col pi ; outline in equatorial view ellipsoidal; equatorial diameter 15fL-22fL, polar axis 18/L-24/L. Apertures: Colpi 

simple, long, without margines; apocolpia small. 

Pollen wall: Wall 1/L-1.51--t thick. Endexine comparatively thick; in intercolpal areas (as seen in 

polar views of grains) endexine approximately as 

thick as ektexine but thins to very thin near colpi. 

Ektexinal elements visible at X 1000 oil-immersion 

magnification, consisting of microclavae. Surface 

sculpture at high focal level microreticulate under 

oil immersion (X 1000), surface of grains appearing nearly smooth at x 450 magnification. 

Remarks.-The specimens described above appear 

to be the species Tricolpites hians (and Tricolpites 

parvus) as illustrated by Stanley (1965, pl. 47, figs. 

24-26, 28-31). Many of the specimens illustrated 

by Elsik (1968b, pl. 23, figs. 13-16, 18 and pl. 24, 

figs. 1-4, 8-14) under the name Tricolpopollenites 

hians (Stanley) Elsik do not appear to be equal to 

Stanley's Tricolpites hians or Tricolpites parvus. 

Tricolpites varius Norton in Norton and Hall 

(1969, p. 45, pl. 6, fig. 15) from the Fort Union 

Formation of Montana appears to be similar and 

may be equal to T. hians Stanley. 

Tricolpites spp. 

Plate 8, figures 35-37; table 1, No. 26 

Genus TRICOLPOPOLLENITES Pftug and Thomson in Thomson 

Pflug, 1953 

Cornaceoipollenites Potonie, 1951, p. 151. (Not a valid name, 

genus not described until 1960 (in Potonie, 1960, p. 93) .) 

Tr,icolpopollenites Pflug and Thomson in Thomson and Pflug, 

1953 (March), p. 95. 

Tricolpites Cookson, 1947 ex Couper, 1953 (August), p. 61. 

[not] Tricolpites (Erdtman, 1947, Cookson, 1947, Ross, 

1949) Couper, 1953 emend. Potonie, 1960, p. 95. 

Tricolpites (Couper, 1953) Belsky, Boltenhagen, and Potonie, 1965 emend. Boltenhagen, 1967, p. 344, 346. 

Type species. - Tricolpopollenites (Pollenites) 

parmularius (Potonh~) Thomson and Pflug, 1953, p. 

97; holotype in Potonie, 1934, p. 52, pl. 2, fig. 7 and 

pl. 6, fig. 11; Geiseltals, Germany; Eocene. 

Description.-Radially symmetrical, isopolar, tricolpate pollen grains. Colpi comparatively simple, 

not accompanied by distinctly differentiated pores 

but geniculi (Thomson and Pflug, 1953, p. 41) may 

be present. Polar axis equal to or greater than equatorial diameter. Size of grains and sculpture various. A form genus intended to accommodate tricolpate pollen that are not assignable to more precisely 

circumscribed genera. 

Remarks.-The form genera Tricolpopollenites 

Pflug and Thomson and Tricolpites Cookson ex 

Couper were published almost simultaneously, and 

both were apparently intended to embrace a wide 

range of tricolpate pollen species having uncertain 

botanical affinity. Pflug and Thomson, in March 

1953 (in Thomson and Pflug, 1953, p. 95), diagnosed Tricolpopollenites as follows: (translation) 

"Three germinals are present. These are symmetrical to each other. Apertures are developed only as 

meridional colpi. The polar axis is a three numbered 

symmetrical axis." Pflug and Thomson divided the 

genus into two sections. These are: section (a) 

species with smooth, baculate, fossulate, or other 

sculptures (except echinat~), and section (b) species with echinate sculpture. Tricolpopollenites was 

placed (see Thomson and Pflug, 1953, p. 95) in the 

division "Longaxones" (polar axis longer than or 

equal to the equatorial axis). Couper, in August 

1953 (p. 61), validated the genus Tricolpites. He 

described it as follows: "Free, isopolar, tricolpate. 

Exine variable in thickness and sculpture. Size variable." Tricolpites was indistinguishable from Tricolpopollenites on the basis of shape, inasmuch as 

Couper included in the genus subprolate, subspheroidal, prolate-spheroidal, prolate, and spheroidal 

species. The two genera were not distinguishable 

on the basis of sculpture because both, as described, 

had a wide variety of surface ornamentations. 

Potonie (1960, p. 95) reinstated Tricolpites, limiting it to finely reticulate forms having broad, margined furrows, and three-lobed shape. Subsequently, 

Belsky, Boltenhagen, and Potonie (1965, p. 75) 

reemended and expanded Tricolpites to include 

strongly reticulate forms without colpal margines 

and with oblate to spherical shape. Boltenhagen 

( 1967, p. 344, 346) recommended further expansion 

of Tricolpites that would have made it again 

synonymous with Tricolpopollenites (see "Remarks,'' 

this report, p. 22). 

Krutzsch (1959b, p. 143) stated that he examined Tricolpopollenites parmularius and it is tri-
SYSTEMATICS 25 

colporate. In view of Krutzsch's statement, Potonh~ 

(1966, p. 168) was of the opinion that Tricolpopollenites would not apply to tricolpate forms. Pflug 

and Thomson (in Thomson and Pflug, 1953, p. 41) 

already indicated, at the time that they described 

Tricolpopollenites, that a geniculus was characteristic of some species which they placed in the genus. 

They believed that the geniculus or equatorial nick 

in the colpus and in the caverna probably displays 

a morphological stage in the formation of a pore. 

I do not believe that pores in the type species of 

Tricolpopollenites in any way alter the usefulness 

of Tricolpopollenites as a fonn genus for tricolpate 

pollen, as well as for tricolporate pollen which have 

indistinct, underdeveloped, or uncertain pores. I believe that this was the original intended scope of 

the genus. 

Tricolpopollenites leviitas n. sp. 

Plate 9, figures 1-6; table 1, No. 29 

Holotype.-USGS paleobotany locality D3725-C, 

slide (2) at 113.2 x 3.8; plate 9, figure 1; NW%~ 

sec. 26, T. 24 N., R. 17 E.; Blaine County, Mont.; 

Judith River Formation (upper nonmarine part) ; 

late Campanian. 

Paratype.-USGS paleobotany locality D3725-C, 

slide ( 4) at 80.9 X 6.8; plate 9, figures 2, 3; same 

locality data as for holotype. 

Paratype.-USGS paleobotany locality D3725-C, 

slide ( 4) at 84.4 X 11.3, plate ~), figure 4; same locality data as for holotype. 

Description (based on more than 100 specimens, 

of which 35 were measured} .-Tricolpate (rarely 

four-col pate) pollen grains ; outline in polar view 

circular to rounded triangular, in equatorial view 

circular to broadly elliptical; equatorial diameter 

16tL-24fL, polar axis 16fL·-26fL. Apertures: As seen in 

polar views of grains, colpi reaching approximately 

two-thirds the distance to poles, bordered by narrow margines (0.5/L or less wide) ; in some specimens colpal slits jagged or irregular, appearing 

eroded. As seen in equatorial views, some specimens 

appear to have colpi of difrerent lengths (as in 

Eucommiidites) ; however, grains in true polar 

view seem to have colpi of virtually equal lengths. 

In both polar and equatorial views, a few specimens 

have been seen which appear to have equatorial 

pores accompanying the colpi (pl. 9, fig. 5). Pollen 

wall : Wall (except near col pi 1/L -2rt thick, in some 

specimens thickened at poles. Ektexine two to four 

times thicker than endexine, in some specimens 

wall layers almost indistinguishable ; as seen on 

equator (in polar views of grains), the wall gradually thins to 0.5fL or less near colpi; ektexine not 

ornamented, surface smooth. 

Remarks.-The species name levitas (evenness 

of surface) was chosen for this taxon because of 

its smooth surface texture. 

Comparison.-In general appearance, some specimens of Tricolpopollenites levitas resemble Eucommiidites troedssonii Erdtman ( 1948, p. 266, figs. 

13-15) and Eucommiidites minor Groot and Penny 

(1960) as illustrated by Hughes (1961, pl. 37, figs. 

17, 18). Consideration was given to placing this 

new species in the genus Eucommiidites because it 

does have some characteristics in common with 

members of that genus; for example, its jagged 

edged colpi (jagged slits were mentioned by Couper, 

1958, p. 160, for E. troedssonii) and the thinning 

of its equatorial exine near the colpi (mentioned 

for E. troedssonii by Erdtman, 1948, p. 268 and 

shown in his diagrams, p. 266, figs. 4, 8) . However, 

T. levitas does not conform to the genus Eucommiidites as it was interpreted by Couper (1958, p. 

160) or by Hughes (1961, p. 292). Couper described 

Eucommiidites as having one colpus generally better 

developed than the other two; and Hughes, in his 

description, referred to one flat face having a long 

furrow and the opposite more convex face bearing 

a ring-furrow. On the contrary, T. levitas has three 

symmetrically arranged equal colpi. 

Some specimens of T. levitas resemble Hedlund's 

illustrations of Tricolpites erugatus (Hedlund, 1966, 

pl. 9, figs. 2a, 2b) ; however, the shorter colpi and 

distinctive colpal margines of T. levitas distinguish 

the species from T. erugatus. 

T ricolpopollenites tessellatus n. sp. 

Plate 9, figures 7, 8; table 1, No. 30 

Holotype.-USGS paleobotany locality D3725-E, 

slide (2) at 107.6X8.9; plate 9, figure 8; NW!Jt. sec. 

26, T. 24 N., R. 17 E.; Blaine County, Mont.; Judith 

River Formation (upper nonmarine part) ; late 

Campanian. 

Description (based on many specimens, 28 of 

which were measured) .-Isopolar, tricolpate pollen 

grains; outline in polar view circular, with three 

evenly spaced colpal incisions; outline in equatorial 

view narrowly to broadly ellipsoidal; equatorial diameter 8fL-21fL, polar axis 13.5fL-21fL. Apertures: 

Colpi siltlike to gaping, reaching two-thirds to threefourths the distance to poles (as seen in polar view 

of grains) . Pollen wall : Wall approximately 0.5p. 

thick, ektexine two to three times thicker than 

endexine, in many specimens wall layers not discernible. Ektexinal ornamentation consisting of 

helter-skelter groups of parallel muri; muri typi-
26 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

cally 1.5/L-3/L long, less than 0.5fL wide, and less 

than 0.5/L high; muri in groups of three or four, 

individual muri parallel to each other and spaced 

approximately as far apart as their width; groups 

of parallel muri oriented in various directions over 

surface of grains to form a mosaiclike pattern. 

Remarks.-This taxon was given the species name 

tessellatus because of its tessellated, checkered, or 

mosaic surface ornamentation. It was assigned to 

the heterogeneous form genus Tricolpopollenites 

because I do not believe that this single species warrants the erection of a new genus. Perhaps at a later 

date a more appropriate niche will be found for this 

new species. It does not pertain to the striate genera 

Striatopollis Krutzsch (1959b, p. 142-143) which 

has longitudinal striations (parallel to polar axis 

or to Aesculiidites Elsik (1968b, p. 632) which has 

circum.striate ornamentation. 

Comparison with other species.-! have not seen 

any fossil grains or found in the literature any 

pollen with ornamentation comparable to that of 

T. tessellatus. This species with its variously oriented groups of parallel muri is distinctly different 

from Aesculiidites (Tricolpites) circumstria.tus 

(Fairchild in Stover, Elsik, and Fairchild) Elsik 

(1968b, p. 632, 634) which has circumstriate ornamentation, from Striatopollis swrstedtensis Krutzsch 

(1959b, p. 143-144) which has striations parallel 

to its polar axis, and from Grewipollenites canadensis Srivastava (1969d, p. 61) which has striae dispersed radially from centers. 

Genus CRANWELLIA Srivastava, 1966 

Cranwellia Srivastava, 1966, p. 537. 

Type species.-Cranwellia (Elytranthe) striat,a 

(Couper, 1953) Srivastava, 1966, p. 537-538; bolotype illustrated in Couper, (1960, pl. 8, fig. 15; 

Mataura Valley, New Zealand; Waitakian Stage; 

middle Oligocene. 

Remarks.-Srivastava proposed the genus Cranwellia to accommodate fossil pollen grains which 

show a similarity or affinity with the pollen of the 

extant genus Elytranthe. He assigned the genus to 

the family Loranthaceae. CTanwellia is used here 

a.s a form genus because, even though the species 

assignable to it show some resemblance to members 

of the genus Elytranthe, I am not sure that they 

are assignable to the family Loranthaceae. 

Cranwellia rumseyensis Srivastava, 1966 

Plate 9, figure 9; table 1, No. 57 

Cranwellia rumseyensis Srivastava, 1966, p. 538. 

Holotype.-ln Srivastava, 1966, p. 538, pl. XI, 

figs. 3, 7; Scollard area, Alberta, Canada; upper 

part of Edmonton Formation ; Maestrichtian. 

Cranwellia bacata Srivastava, 1969d 

Plate 9, figures 10-12; table 1, No. 31 

Cranwellia bacata Srivastava, 1969d, p. 51. 

Holotype.-In Srivastava, 1969d, p. 51, pl. 1, fig. 

2; Drumheller locality, Alberta, Canada; lower 

member of Edmonton Formation; Maestrichtian. 

cf. Cranwellia 

Plate 9, figure 13; table 1, No. 58 

Genus FIBULAPOLLIS Chlonova, 1961 

Fibulapollis Chlonova, 1961, p. 87. 

Fibulapollis Chlonova, 1961. In Tschudy, 1969, p. A13. 

Type species.-Fibulapollis (Triporina) mirificus 

(Chlonova, 1957) Chlonova, 1961, p. 87. 

Fibulapollis scabratus Tschudy, 1969 

Plate 9, figures 14-16; table 1, No. 75 

Fibulapollis scabratus Tschudy, 1969, p. A13-A14, pl. 16, 

figs. 11-20. 

Holotype.-In Tschudy, 1969, p. Al3, pl. 16, fig. 

11; Colville River, Alaska; Colville Group; Late 

Cretaceous. 

Genus AQUILAPOLLENITES Rouse, 1957 

Aquilapollenites Rouse, 1957, p. 370. 

Type species.-A quilapollenites quadrilobus 

Rouse, 1957, p. 371; holotype in Radforth and 

Rouse, 1954, pl. I, fig. 14; Alberta, Canada; Brazeau 

Formation; Late Cretaceous. 

Remarks.-Aquilapollenites as used here encompasses only those species having gross morphological 

characteristics (three equatorial protrusions ; and 

two unequal polar protrusions such as those of A. 

quadrilobus, or two more or less equal polar protrusions such as those of A. trialatus) which correspond to those of the two original species which 

were assigned to the genus by Rouse. Subsequent 

to its inception Aquilapollenites has been emended, 

subdivided, and variously interpreted by many 

authors, including Funkhouser (1961, p. 193-194), 

Stanley (1961, p. 338, 340), Chlonova (1961, p. 82-

83), Mtchedlishvili (in Samoilovitch and others, 

1961, p. 209), Belsky, Boltenhagen, and Potonie 

(1965, p. 80_.:_81), Srivastava (1968a, p. 668-670), 

Krutzsch (1970, p. 104, 109), Srivastava and Rouse 

(1970, p. 1592-1593), and Tschudy and Leopold 

(1971, p. 120-124). The emendations and subdivisions in part have merit but I do not believe any of 

them are entirely satisfactory. 

In this report, M ancicorpus, as restricted by 

Krutzsch (1970, p. 104-106), is recognized as a 

useful genus because it is distinct from Aquilapollenites Rouse; however, Mancicorpus, as introducEd 

by Mtchedlishvili (in Samoilovitch and others, 1961, 

p. 218-219) or as emended by Srivastava (1968a, 
SYSTEMATICS 27 

p. 695-697), is not acceptable because some of the 

species included in it are extremely similar to the 

type species of Aquilapollenites. As discussed earlier 

(Tschudy and Leopold, 1971!, p. 120-121), several 

genera proposed by Mtchedlishvili that included 

species earlier described under Aquilapollenites are 

not acceptable because they are not clearly distinct 

from each other or from Aquita.pollenites. Krutzsch's 

new genus, Hemic01·pus Krutzsch (1970, p. 104, 

107), is not acceptable because it is not distinct 

from Aquilapollenites; in fact, I believe its type 

species, H emicorpus (Aquilapollenites) pule her 

(Funkhouser) Krutzsch (1970, p. 107), is synonymous with A. quadrilobus, the type species of Aquilapollenites. Krutzsch's limitation of Aquilapollenites to isopolar and subisopolar species is probably 

based on his misunderstanding of the type species 

of the genus-Krutzsch (1970, p. 109) cited Rouse 

(1957, pl. II, fig. 9) as the type of A. quadrilobus 

but that figured specimen is not the holotype specimen designated by Rouse. 

Aquilapollenites quadrilobus Rouse, 1957 

Plate 9, figures 17, 18; table 1, No. 32 

Aquilapollenites quadrilobus Rous·e, 1957, p. 371. 

Aquilapollenites quadrilobus Rouse, 1957. In Srivastava and 

Rouse, 1970, p. 1597-1598, pl. I, figs. 3-7. 

Aquilapollenites quadn"lobvs Rouse, 1957. In Tschudy and 

Leopold, 1971, p. 125-127, 130--131, pl. I, figs. 5A, B and 

pl. 2, figs. lA, B ( holotype, two levels), 2-6. 

Holotype.-In Radforth and Rouse, 1954, pl. I, 

fig. 14; Alberta, Canada; Brazeau Formation; Late 

Cretaceous. 

Aquilapollenites senonicus (Mtchedlishvili) Tschudy and Leopold, 1971 

Plate 9, figure 19; table 1, No. 65 

Manc.icorpus senonicum Mtchedlishvili in Samoilovitch and 

others, 1961, p. 224-225. 

Aquilapollenites senonicus (Mtchedlishvili) Tschudy and 

Leopold, 1971, p. 131-132, pl. 1, fig. 2. 

Holotype.-In Samoilovitch and others, 1961, p. 

224-225, pl. 72, figs. 2a-c; western Siberian lowlands; lower Symian subsuite; Senonian. 

Aquilapollenites trialatus Rouse var. uniformis Tschudy and Leopold, 1971 

Plate 9, figure 20; talble 1, No. 33 

Aquilapollenites trialatus Rouse var. uniformis Tschudy and 

Leopold, 1971, p. 145, 148. 

Holotype.-In Tschudy and Leopold, 1971, p. 145, 

pl. 4, figs. 8A, B; Niobrara County, Wyo.; Mitten 

Black Shale Member of Pierre Shale; middle or 

early Campanian. 

Aquilapollenites clarireticulatus (Samoilovitch) Tschudy, 1969 

Plate 9, figure 21; table 1, No. 71 

Integricorpus clarireticulatus Samoilovitch, 1965, p. 123-124. 

Aquilapollenites clarireticulatus ( Samoilovitch) Tschudy, 

1969, p. A6, A 7. 

Holotype.-In Samoilovitch, 1965, p. 123-124, figs. 

2:a, b; Linde River, Yakutia, U.S.S.R.; top of Chirimyian suite; Maestrichtian-Danian. 

Aquilapollenites trialatus Rouse, 1957 

Plate 9, figures 22-24; table 1, No. 76 

Aquilapollenites tr·ialatus Rous.e, 1957, p. 371. 

Aquilapollenites trialatus Rouse, 1957. In Tschudy, 1969, p. 

A 7, pl. 4, figs. 1-5; northern Alaska; Colville Group; 

Late Cretac.eous. 

Aquilapollenites trialatus Rouse, 1957. In Srivastava and 

Rouse, 1970, p. 1598-1599, figs. 8-17 (specimens from 

type locality) . 

Aquilapollenites trialatus Rouse var. trialatus. In Tschudy 

and Leopold, 1971, p. 144. 

Holotype.-In Rouse, 1957, p. 371, pl. II, fig. 14; 

south-central Alberta, Canada; Oldman Formation; 

Santonian, Late Cretaceous. 

Aquilapollenites reticulatus (Mtchedlishvili) Tschudy and Leopold, 1971 

Plate 9, figure 25; table 1, No. 77 

Pa.rviprojectus reticulatus Mtchedlishvili in Samoilovitch 

and others, 1961, p. 226-227. 

Aquilapollenites reticulatus (Mtchedlishvili) Tschudy and 

Leopold, 1971, p. 149-151. 

Holotype.-ln Samoilovitch and others, 1961, p. 

226-227, pl. 73, figs. 2a-c; Sym River, western 

Siberian lowlands; upper Symian subsuite; Maestrichtian-Danian. 

Aquilapollenites sp. 

Plate 9, figures 26; table No. 34 

Aquilapollenites turbidus Tschudy and Leopold, 1971 

Plate 10, figures 1, 2; table 1, No. 59 

Aquilapollenites turbidus Tschudy and Leopold, 1971, p. 157-

159, pl. 1, fig. 6, and pl. 2, figs. 7 A-10. 

Holotype.-ln Tschudy and Leopold, 1971, p. 157-

159, pl. 2, figs. 7 A, B; Black Hills area, Niobrara 

County, Wyo.; Red Bird Silty Member of Pierre 

Shale; middle Campanian. 

Aquilapollenites attenuatus Funkhouser, 1961 

Plate 10, figures 3-6; table 1, No. 35 

Aquilapollenites attenuatus Funkhous·er, 1961, p. 194, 196. 

Triprojectus dispositus Mtchedlishvili in Samoilovitch and 

others, 1961, p. 205-206, pl. 65, figs. la-c; Sym River, 

western Siberian lowlands; Maestrichtian-Danian. 

Aquilapollenites asper Mtchedlishvili in Samoilo-vitch and 

others, 1961, p. 213, pl. 68, figs. 2a-c; Sym River, western 

Siberian lowlands; Maestrichtian-Danian. 

Aqunapollenites granulatns Mtchedlishvili in Samoilovitch 

and others, 1961, p. 212, pl. 68, figs. la-c; Vakh River, 

western Siberian lowlands; early Maestrichtian. 

Aquilapollenites a.mplus Stanley, 1961, p. 342, 344, 346, pl. 

1, figs. 1-6; Harding County, S. Dak.; Hell Creek 

Formation; Maestrichtian. 
28 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

Aquilapollenites matsumotoi Takahashi, 1964, p. 252-253, pl. 

38, figs. 1a, b; Hokkaido, Japan; lower Hokobuchi Formation; Campanian. 

Aquilapollenites accipiteris Srivastava, 1969a, p. 133-134, pl. 

I, figs. 3-5; Alberta, Canada; Edmonton Formation; 

Maestrichtian. 

Aquilapollenites ascriptivus Srivastava, 1969a, p. 136, pl. II, 

figs. 12-15; Alberta, Canada; Edmonton Formation; 

Maestrichtian. 

Aquilapollenites augustus Srivastava, 1969a, p. 137, pl. III, 

figs. 18-20; Alberta, Canada; Edmonton Formation; 

Maestrichtian. 

Aquilapollenites h1'spidus Srivastava, 1969a, p. 139-140, pl. 

V, figs. 31, 32; Alberta, Canada; Edmonton Formation; 

Maestrichtian. 

Aquilapollenites regaUs Srivastava, 1969a, p. 142, pl. VI, 

figs. 42, 43; Alberta, Canada; Edmonton Formation; 

Maestrichtian. 

Holotype.-In Funkhouser, 1961, p. 194, 196, pl. 

2, figs. la-c; west flank of Rawlins uplift, Rawlins, 

Wyo.; Lance Formation; Maestrichtian. 

Description (based on Funkhouser's holotype 

specimen and on more than 100 specimens from 

Judith River Formation samples and from many 

additional USGS paleobotany locality samples).-

Tricolpate, isopolar to subisopolar pollen grains 

with three equatorial protrusions. In equatorial 

view: Polar axis 40fL-84rt (commonly 50fL·-55fL), 

equatorial diameter (including equatorial protrusions) 43p;-93fL (commonly 55fL-63fL), diameter of 

body 11~--t-42/L (commonly 17 fL-23 rt) , diameter of 

equatorial protrusions llft-22p, (commonly 14fL18fL), length of equatorial protrusions 15fL-34fL 

(commonly 20fL-25fL). Shape of body oval; diameter of polar protrusions usually greater than, or 

in some specimens equal to or less than, the diameter of equatorial protrusions; length of polar 

protrusions usually approximately equal to, or in 

some specimens slightly greater or less than length 

of equatorial protrusions; polar protrusions and 

equatorial protrusions broadly rounded at their 

ends. In polar view: Shape of grains triangular, 

with extended rounded corners and slightly concave sides; free ends of equatorial protrusions 

paddle shaped, flattened in plane parallel to polar 

axis of grain. Apertures: Colpi dissecting equatorial protrusions, extending a short distance onto 

body, sometimes gaping at free ends of Equatorial 

protrusions. Pollen wall : Exine two layered; endexine (except for endexinal costae) usually less 

than 0.5fL thick. Axillary endexinal costae bow 

shaped to lens shaped, extending one-half to twothirds the length of equatorial protrusions and 

extending onto base of polar protrusions, costae as 

much as 3fL thick in thickest part; in some specimens costae very distinct, in others faintly visible 

or absent (where absent probably were removed by 

chemical treatment during removal of grains from 

rock matrix). Ektexine approximately lfL thick; 

ektexinal elements columnar, their diameter approximately equal to the distance between them, fused 

at tips in partial tectum resulting in a foveolate to 

reticulate appearance of surface of grains. Conical 

spines extending 1.5fL·-2.5fL above surface of tectum, 

ornamenting surface of grain; spines mainly restricted to polar domes, tips of equatorial protrusions, equatorial areas, and colpal margins (along 

edges of equatorial protrusions in equatorial view 

of grains) . Spines on polar protrusions oriented 

perpendicular to surface, on equatorial protrusions 

usually oriented toward polar axis of grain, at colpal 

margins spines retroflexed (pointing toward polar 

axis of grain). Spines at ends of equatorial protrusions not conspicuously larger than those on 

rest of grain. 

Remarks.-Funkhouser described A. attenuatu.s 

as a tridemicolpate pollen species. I have examined 

the holotype specimen and believe that the demicolpi referred to by Funkhouser are thin areas from 

which endexinal costae have partially dissolved. 

Colpi are clearly visible in many of the specimens 

that I have seen; they are, as a rule, most apparent 

in polar and oblique views of grains. 

The new description of A. attenuatus presented 

above is not considered an emendation but is merely 

meant to clarify and expand the species in the light 

of present knowledge and greater abundance of 

specimens. A. attenuatus as here amplified consists 

of grains of a wide but gradational size range and 

grains with much variation in proportional length 

and width of polar and equatorial protrusions. In 

one USGS paleobotany locality sample (D3012 from 

the Bearpaw Shale of Montana) about 40 specimens 

were examined. In this sample there were grains 

closely resembling the holotype specimen of A. 

attenuatus and also large to small grains with proportionally larger bodies and smaller equatorial protrusions than the holotype. Some specimens in this 

sample had unequal polar protrusions (subisopolar). 

Sculpture also varied. Another USGS paleobotany 

sample (Dl331 from the Fox Hills Sandstone from 

Wyoming), though containing a smaller number 

of specimens, also provided grains having widely 

variable form. I have attempted to separate our 

large number of specimens into varieties of A. 

attenuatus but have concluded that variants intergrade and cannot be distinctly or reliably separated 

from each other. 

Similar species.-"Triprojectus'' unicus (Chlonova) Mtchedlishvili (in Samoilovitch and others, 

1961, p. 206-207, pl. 66, figs. la-c) and "Tr;·ipro-
SYSTEMATICS 29 

jectus" crassus Mtchedlishvil:l (in Samoilovitch and 

others, 1961, p. 207-208, pl. 66, figs. 2a-c, 3) are 

similar to A. attenuatus but differ by having more 

slender equatorial protrusions. 

Aquila;pollenites insignis 1v.ltchedlishvili (in Samoilovitch and others, 1961, p. 215-216, pl. 69, figs. 

la-c, 2, 3) closely resembles A. attenuatus but possibly it can be distinguished because in the illustrations (drawings) A. insignis appears to have knoblike projections instead of spines on the ends of the 

equatorial protrusions. 

Aquilapollenites subtilis 1\Hchedlishvili (in Samoilovitch and others, 1961, p. 214, pl. 68, figs. 3a-c, 

4) differs from A. attenuatus in its uniform distribution of spines over the entire surface of the pollen 

grain. 

Aquilapollenites latilobu.s JM:tchedlishvili (in Samoilovitch and others, 1961, p. 216, pl. 69, figs. 4ac) differs from A. attenuc~tus in its distinctly 

coarser reticulum on the polar areas. 

Aquilapollenites aucellatus Srivastava (1969a, p. 

136-137, pl. III, figs. 16, 17) and Aquilapollenites 

drumhellerensis Srivastava ( 1969a, p. 138, pl. IV, 

fig. 28) appear to be similar to A. attenuatus but 

can be differentiated from it because of their more 

abundant and smaller spines in the equatorial areas. 

Genus MANCICORPUS Mtchedlishvili, pro parte, Krutzsch, 1970 

[in part] Manc.ic01·pus Mtchedlishvili in Samoilovitch and 

others, 1961, p. 218-219. 

Mtchedlishvilia Srivastava, 1969c, p. 615-616. 

Mancicorpus Mtchedlishvili, pro parte Krutzsch, 1970, p. 

103, 105-106. 

Type species.-Mancicorpus anchoriforme 

Mtchedlishvili in Samoilovitch and others, 1961, p. 

219-220, pl. 70, figs. 3a-c; Sym River, western Siberian lowland; upper Symia:n subsuite; Maestrichtian-Danian. 

New combination.- Mancicorpus (Mtchedlishvilia) canadianus (Srivastava, 1969c, p. 616, 618, 

pl. I, figs. 1-8; Alberta, Canada; Edmonton Formation ; Maestrichtian) n. comb .. 

Remarks.-Mancicorpus as restricted by Krutzsch 

is distinct from AquilapollenUes and is therefore a 

useful form genus. Krutzsch included in Mancicorpus only forms that have a tripodal shape-forms 

having one concave or flat pole and one convex but 

not prominent pole. The remaining four species that 

Mtchedlishvili originally described and included in 

Mancicorpus-species that have one major polar 

protrusion and one much smaller or nodelike protrusion-were placed by Krutzsch in his new genus 

Hemicorpus. Hemicorpus Krutzsch (1970, p. 104), 

however, is here considered to be a synonym of 

Aquilapollenites because the type species of Hemicorpus (Hemicorpus (Aquilapollenites) pulcher 

(Funkhouser) Krutzsch) is extremely similar to, 

and I believe synonymous with, A. qua.drilobus, the 

type species of Aquilapollenites. 

The genus Mtchedlishvilia Srivastava as circumscribed by its author is not distinguishable from 

Mancicorpu.s as it is represented by its type species, 

M ancicorpus anchori forme. 

Mancicorpus calvus (Tschudy and Leopold, 1971) n. comb. 

Plate 10, figures 7-9; table 1, No. 37 

Aqu.ilapollenites calvus Tchudy and Leopold, 1971, p. 138-

140, pl. 1, fig. 4 and pl. 3, figs. 1-5. 

Holotype.-In Tschudy and Leopold, 1971, p. 138-

140, pl. 3, fig. 1; Black Hills area, Niobrara County, 

Wyo. ; lower unnamed shale member of Pierre Shale ; 

late Campanian. 

Mancicorpus tripodiformis (Tschudy and Leopold, 1971) n. comb. 

Plate 10, figures 10, 11; table 1, No. 60 

Aquilapollenites tripodiformis Tschudy and Leopold, 1971, 

p. 140-143, pl. 3, figs. 9A-10. 

Holotype.-In Tschudy and Leopold, 1971, p. 140-

141, pl. 3, figs. 9A, B; Black Hills area, Niobrara 

County, Wyo.; Pierre Shale; late Campanian. 

Remarks.-M. tripodiformis n. comb. is similar 

to Mancicorpu.s (Mtchedlishvilia) canadianus (Srivastava, 1969c) n. comb. but apparently is distinguishable because of the striated sculpture on its 

major pole and equatorial protrusions. It is possible 

that these two species will be found to merge and 

that M. tripodiformis will become a synonym of 

Mancicorpus canadianus. 

POLYCOLPATES 

Genus SINDORAPOLLIS Krutzsch, 1969 

Sindorapollis Krutzsch, 1969, p. 413. 

Type species. - Sindorapollis (Loranthacites) 

pilatu.s (Mtchedlishvili) Krutzsch, 1969, p. 413; 

holotype in Samoilovitch and others, 1961, p. 195, 

pl. 63, figs. la-c; western Siberian plains ; Senonian. 

Description.-Radially symmetrical, angulaperturate pollen grains having three pairs of demicolpi 

which are separated at equator by equatorial bridges 

and at poles by polar areas (not syncolpate) ; demicolpi either accompanied by or not accompanied by 

pores. Size of grains and exine sculpture various. 

New combination.-Sindorapollis (Loranthacites) 

catteralli (Srivastava, 1969b, p. 980, pl. II, figs. 

21, 22; Alberta, Canada; Edmonton Formation; 

Maestrichtian) n. comb. Loranthacites Mtchedlishvili, to which the species catteralli was originally 

assigned, is a later homonym of Loranthacites 

Conwentz, (1886). 
30 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER ,FORMATION; NORTH-CENTRAL MONT. 

Sindorapollis granulatus n. sp. 

Plate 10, figures 12-16; table 1, No. 78 

Holot'!:fpe.-USGS paleobotany locality D4069 

slide (1) at 106.5 x 10.5; plate 10, figure 12; 

~W ;4SW 4SW1,4 sec. 15, T. 23 N., R. 22 E.; Phillips County, Mont.; Judith River Formation· late 

Campanian. ' 

Paratype.-USGS paleobotany locality D4069, 

slide (2) at 104.3 X 3.6; plate 10, figure 13; same 

locality data as for holotype. 

Paratype.-USGS paleobotany locality D4069 

slide (1) at 105.7 x 18.7; plate 10, figure 14; sam~ locality data as for holotype. 

Description (based on 38 specimens in polar view 

~nd six in equatorial view) .-Radially symmetrical, 

Is~polar, angulaperturate pollen grains having three 

pairs of demicolpi ; outline of grain in polar view 

rounded triangular, in equatorial view oval to 

rounded rhomboidal; equatorial diameter 16.5p.-

28p. (commonly 20p.-26p.), polar axis 19p.-24p .. Apertures: Demicolpi slitlike, in some specimens gaping 

slightly, without margine.s, 5p.-8p. long, not joining 

at poles, reaching to within 0.5p.-1.5,u of equator; 

occasionally the narrow bridge between demicolpi 

at equator is apparently torn and demicolpa1 slits 

are joined (forming a colpus). Pollen wall: Wall 

0.5p.-1p. thick; endexine thin, ektexine two to three 

times thicker than end exine; in most specimens 

wall at equator slightly thick2r on corners than on 

sides of grain; ektexinal columellae very fine, not 

always visible even under x 1000 oil-immersion 

magnification, tectate; at x 430 high dry magnification, surface of grains appearing almost smooth ; 

at X 1,000 oil-immersion magnification, surface 

appearing granular. 

Comparison with similar species.-Sindorapollis 

granulatus n. sp. differs from Sindorapollis catteralli (Srivastava) n. comb. in its smaller size, more 

rounded outline, shorter demicolpi, and finer sculpture. 

S. granulatus differs from Sindorapollis pilatus 

(Mtchedlishvili) Krutzsch (1969, p. 413) in its 

smaller size, much finer surface sculpture, and lack 

of pores. 

S. granulatus is smaller than SindorapoUi8 triangulus Kruztsch (1969, p. 413) and has siltlike 

simple demicolpi, whereas S. triangulus has demicolpi with thickened margins. 

TRICOLPORA TES 

Genus CUPANIEIDITES Cookson and Pike, 1954 

Cupanieidites Cookson and Pike, 1954, p. 210. 

Cupanieidites Cookson and Pike, 1954. In Krutzsch, 1959b, p. 

144-145. 

Duplopollis Krutzsch, 1959b, p. 144. 

Cupanieidites Cookson and Pike, 1954. In Potonie, 1960, p. 

106. 

Cupanieidites Cookson and Pike, 1954. In Belsky, Boltenhagen, and Potonie, 1965, p. 77. 

Type species.-(selected by Krutzsch, 1959b, p. 

144) Cupanieidites major Cookson and Pike, 1954, 

p. 213-214, pl. 2, figs. 83-85; lectotype (here selected) in Cookson and Pike, 1954, pl. 2, fig. 83; 

Anglesea, Victoria, Australia; Eocene. 

Cupanieidites sp. 

Plate 10, figure 17; table 1, No. 91 

Genus TRICOLPORITES Cooksoa, 1947 

Tricolporites Cookson, 1947, p. 134. 

Type species.-Tricolporites p1·olata Cookson, 

1947, p. 134, pl. XV, fig. 46; Waterfall Gorge (near 

Port Jeanne d'Arc), Kerguelen Archipelago, Indian 

Ocean ; Tertiary. 

Description.-Isopolar, tricolporate pollen grains. 

Polar axis greater than, or equal to, equatorial diameter. Colpi meridional, arranged symmetrically, each 

provided with an equatorially located well-differentiated pore. Size of grains variable. Surface 

smooth or nearly so--dimensions of sculptural elements less than 1p.. 

Remarks.-Erdtman (1947, p. 110} used the term 

"Tricolporites" in his system of classification of 

fossil and recent pollen and spores but did not use 

it as a genus name. Cookson ( 194 7, p. 134) introduced Tricolporites in the generic sense when she 

described and illustrated the binomial Tricolporites 

prolata. Tricolporites Cookson was a monotypic new 

genus and was a valid name (see article 42 of the 

"International Code of Botanical Nomenclature" 

(Lanjouw, chm., and others, 1966). 

Tricolporites rhomh.oides Anderson. 1960 

Plate 10, figures 18-21; table 1, No. 38 

Tricolp€Jirites rhomboides Anderson, 1960, p. 26. 

Holotype.-ln Anderson, 1960, p. 26,. pl. 7, fig. 15; 

eastern side of San Juan Basin, New Mexico; N acimiento Formatio.n; Cretaceous.., Tertiary. 

Description (based on many specimens, of which 

30 were measured.}-Tricolporate pollen grains; 

outline in polar view rounded triangular to nearly 

circular; outline in equatorial view rhomboidal, 

elliptical, or nearly circular. Equatorial diameter 

l6p.-22p., polar axis 16.5p.-2461• Most specimens in 

polar view have a triradiate compression fold on 

polar area (see pl. 10, fig. 18). Apertures: Colpi 

long, extending two-thirds to three-fourth the distance to poles, bordered by narrow (approximately 

0.5p. wide) margines; pores as seen in equatorial 

view of grains 0.5p.-1p. (rarely as much as 1.5p.) in 
SYSTEMATICS 31 

diameter. Pollen wall: Wall approximately 1.5f.L 

thick; ektexine two to three times thicker than 

endexine; ektexinal columellae fused at tips in a 

partial tectum. Surface sculpture scabrate. 

Remarks.-The grains described above, except 

for their slightly larger size, correspond to T. rhomboides as described and illustrated by Anderson. 

T ricolporites sp. 

Plate 10, figure 22; table 1, No. 96 

Genus MONTANAPOLLIS Tschudy, 1971 

Montanapollis Tschudy, 1971, p. B58. 

Type species.-M ontanapollis endannulatus 

Tschudy, 1971, p. B58, figs. 3, 4; Phillips County, 

Mont. ; Bearpaw Shale (basal part) or Judith River 

Formation (uppermost part); late Campanian. 

Montanapollis endannulatus Tschudy, 1971 

Plate 11, figure 1; table 1, No. 39 

Montanapollis endannulatus Tschudy, 1971, p. B58. 

Holotype.-ln Tschudy, 1971, p. B58, fig. 3; Phillips County, Mont.; Bearpaw Shale (basal part) or 

Judith River Formation (uppermost part) ; late 

Campanian. 

Genus SIBERIAPOLLIS Tschudy, 1971 

Siberiapollis Tschudy, 1971, p. B54. 

Type species.-Siberiapollis montanensis Tschudy, 

1971, p. B55, 57-58, fig. 1, 2; Fergus County, Mont.; 

lower marine part of Judith River Formation ; late 

Campanian. 

Siberiapollis montanensis Tschudy, 1971 

Plate 11, figures 2, 3; table 1, No. 72 

Siberiapollis montanensis Tschudy, 1971, p. B55. 

H olotype.-ln Tschudy, 1971, p. B55, a of fig. 2; 

Fergus County, Mont.; lower marine part, Judith 

River Formation; late Campanian. 

TRI·PORATES 

Genus TRIATRIOPOLLENITES Pftug in Thomson and Pftug, 1953 

Triatriopollenites Pflug in Thomson and Pflug, 1953, p. 76-77. 

Type species.-Triatriopollenites rurensis Pflug 

and Thomson in Thomson and Pflug, 1953, p. 79, pl. 

7, fig. 95; Eschweiler, Germany; Miocene. 

Triatriopollenites costatus Norton in Norton and Hall, 1969 

Plate 11, figures 4, 5; table 1, No. 61 

Triatriopollenites costatus Norton in Norton and Hall, 1969, 

p. 40-41. 

Holotype.-ln Norton and Hall, 1969, p. 40-41, 

pl. 5, fig. 19; Garfield County, Mont. ; Hell Creek 

Formation ; Late Cretaceous. 

Genus SUBTRIPOROPOLLENITES Pflug and Thomson in Thomson an:I 

Pflug, 1953 

Subtriporopollenites Pflug and Thomson in Thomson and 

Pflug, 1953, p. 85. 

Subtriporopollenites Pflug and Thomson 1953. In Krutzsch, 

1960, p. 307-308. 

Type species. - Subtriporopollenites anulatus 

subsp. anulatus (not subsp. notus) Pflug and Thomson in Thomson and Pflug, 1953, p. 85-86, pl. 9, fig. 

49; middle Europe; Tertiary. 

Subtriporopollenites alpinus (Wolf, 1964) n. comb. 

Plate 11, figures 6-8; table 1, No. 41 

Triporopollenites alpinus Wolf, 1964, p. 114-115, pl. 1, figs. 

8-19. 

Holotype.-ln Wolf, 1964, p. 114-115, pl. 1, figs. 

8-10; Breitenbach on the Tegern Sea, Bavaria; 

Zementmergel Series; Campanian. 

Description (based on 20 specimens from Judith 

River Formation samples) .-Triporate pollen, shape 

in polar view convex triangular to circular, diameter 16f.L-24f.L. Apertures: Pores slightly subequatorial, circular or nearly so, 1.5f.L-2f.L in diameter 

(or rarely as much as 3f.L), not protruding or in 

some specimens very slightly protruding; pore margines approximately 2f.L wide, in most specimens 

inconspicuous, in some specimens not discernible. 

Pollen wall : Exine two layered, about 1f.L thick, as 

much as 1.5f.L thick in margo areas surrounding 

pores; endexine thin, approximately one-third as 

thick as ektexine; in many specimens wall layers 

not discernible. Surface sculpture microverrucate; 

verrucae 0.5f.L-1f.L in diameter, circular in outline 

or nearly so, 0.5f.L or less in height; distance between 

verrucae about equal to, or less than, their diameter. 

Remarks.-My specimens from the Judith River 

Formation appear to be very similar to S. alpinus 

(Wolf) n. comb. as it was described and illustrated 

by Wolf. One specimen that probably is the same 

species, but is slightly larger (24f.L x 29f.L) and has 

four pores, was found in the Judith River sample 

USGS paleobotany locality D3726-A. 

Comparison with similar species.-In shape of 

grain and size of pores S. alpinus resembles Triporopollenites rugatus Newman (1965, p. 12, pl. 

1, fig. 7) ; however, the two species differ in that 

S. alpinus is slightly smaller, and it has a microverrucate surface ornamentation whereas T. rugatus 

has a smooth to scabrate sculpture. 

The sculpture of S. alpinus resembles that of 

Ulmoideipites krempU Anderson ( 1960, p. 20, pl. 

6, fig. 2) and Ulmoideipites tricostatus Anderson 

(1960, p. 20-21, pl. 4, fig. 9), but S. alpinus lacks 

the arci which characterize those species. 
32 PALYNOLOGY, UPPER CAMPANIAN JUDITH RIVER FORMATION, NORTH-CENTRAL MONT. 

Genus PROTEACIDITES Cookson ex Couper, 1953 

Proteacidites Cookson, 1950, p. 170-175. (No description of 

genus and no type species designated.) 

Proteaddites Cooks·on ex Couper, 1953, p. 42. 

Type species.-(designated by Couper, 1953, p. 

42) Proteacidites adenanthoides Cookson, 1950, p. 

172-173, pl. 2, fig. 21; Wensleydale, Australia; Tertiary. 

Proteacidites spp. 

Plate 11, figures 9-12; table 1, No. 42 

POL YPORA TES 

Genus PERIPOROPOLLENITES Pflug and Thomson emend. Krutzsch, 1966 

Liquidambar-pollenites Raatz, 1937, p. 17. (Not valid, no 

genus description.) 

Periporopollenites Pflug and Thomson in Thomson and Pflug, 

1953, p. 111. 

Liquidambarpollenites Raatz, 1937 ex Potonie, 1960, p. 134. 

(Name was here validated by Potonie; however, it was 

a synonym of Periporopollenites which was valid in 

1953-both genera had the same type species.) 

Periporopollenites Pflug and Thomson emend. Krutzsch, 

1966, p, 39-40. 

Type species.-Periporopollenites (Pollenites) 

stigmosus (Potonie) Thomson and Pflug, 1953, p. 

111; holotype in Potonie, 1931a, p. 332, pl. II, fig. 1; 

Muskauer Braunkohle, Germany; Miocene. 

Periporopollenites sp. 

Plate 11, figure 13; table 1, No. 92 

Genus ERDTMANIPOLLIS Krutzsch, 1962 

Erdtmanipollis Krutzsch, 1962, p. 281. 

Type species.-Erdtmanipollis pachysandroides 

Krutzsch, 1962, p. 281, pl. VIII, figs. 1-8. 

Erdtmanipollis pachysandroides Krutzsch, 1962 

Plate 11, figure 14; table 1, No. 43 

Erdtmanipollis pachysandroides Krutzsch, 1962, p. 281. 

Holotype.-In Krutzsch, 1962, p. 281, pl. VIII, 

figs. 1-4; Spremberg, Germany; upper Chatt; late 

Oligocene. 

Remarks.-The diameter of the specimens found 

in the samples from the Judith River Formation 

ranges from 29,u. to 37 ,u.. These grains closely resemble specimens of E. pachysandroides as illustrated by Krutzsch (1962, pl. VIII, figs. 1-8). 

DYADS 

Genus DYADONAPITES Erdtman, 1947 ex Tschudy (this report) 

Dyado-napites Erdtman, 1947. p. 111. (Term proposed in 

system of classification but not used as a genus name.) 

Dyadites Pant, 1954, p. 54. (Name proposed but no species 

assigned.) 

Dyadites van der Hammen, 1956, p. 79. (Not a valid nametype specimen designated was a recent pollen grain.) 

Type species.-Dyadonapites Teticulatus n. sp. 

Description.-Inaperturate palynomorphs unit~d 

in dyads, greatest dimension of dyad less than 601),; 

individual cells of dyad alike or very similar, their 

shape spherical, oval, or irregular; surface ornamentation various. 

Comparison with simila.r genera.-Dyadonapites 

is smaller than the Liassic genus Circularesporites 

Danze and Laveine (1963, p. 86; diameter 60,u.-

500fL), and its individual cells do not have circular 

dehiscence areas as do those of Circularesporites. 

Dyadonapites reticulatus n. sp. 

Plate 11, figures 15-17; table 1, No. 66 

H olotype.-USGS paleobotany locality D3725-C, 

slide (4) at 85.0X6.0; plate 11, figure 15; NW14 

sec. 26, T. 24 N., R. 17 E.; Blaine County, Mont.; 

Judith River Formation (upper nonmarine part); 

late Campanian. 

Paratype.-USGS paleobotany locality D3725-C, 

slide {5) at 91.0X17.1; plate 11, figure 16; same 

locality data as for holotype. 

Description (based on many specimens, of which 

30 were measured) .-Inaperturate palynomorphs 

united in dyads; diameter of individual cells l4p.-

22,u., shape spherical, oval, rounded angular, or 

irregular owing to folding. Length of dyad 23,u.-

41p.. Individual cells of dyad similar but in some 

specimens one cell slightly larger than the other 

one. Wall: Exine probably· two layered, apparent 

thin endexine visible as a component of the wall in 

a few specimens. Ektexinal sculpture reticulate; 

lumina irregular in shape and variable in size, less 

than 0.5,u. to as much as 2,u. in diameter; muri delicate, less than 0.5fL wide and less than 0.5,u. high; in 

some specimens muri appearing rough and on some 

well-preserved specimens this roughness can be 

seen to consist of microbaculae or microverrucae 

on the muri peaks. 

Remarks.-In some preparations containing 

Dyadonapites reticulatus n. sp., many palynomorphs 

are present which appear to be individual cells of 

this species (see taxon 44); whereas, in other preparations very few free cells are present. 

TETRADS 

Genus INAPERTUROTETRADITES van Hoeken-Kiillkenberg, 1964 

Inapert?l.rotetradites van Hoeken-Klinkenberg, 1964, p. 226. 

Type species.-lnaperturotetradites lacunosus van 

Hoeken-Klinkenberg, 1964, p. 226, pl. VI, figs. 17a, 

b ; Inyi, Nigeria; Maestrichtian. 

lnaperturotetradites scabratus n. sp. 

Plate 11, figures 18, 19; table 1, No. 63 

Holotype.-USGS paleobotany loc'ality D3725-A, 
REFERENCES. CITED 33 

slide (2) at 96.3X4.1; plate 11, figure 18; NW1,4 

sec. 26, T. 24 N ., R. 17 E.; Blaine County, Mont.; 

Judith River Formation (upper nonmarine part) ; 

late Campanian. 

Description (based on 20 specimens) .-Inaperturate pollen grains, arranged in tetrahedral tetrads. 

Shape of individual grains spherical or nearly so. 

Diameter of tetrads 29t-t-45t-t, diameter of individual 

grains 18p-27p.. Exine thin, or as much as 0.75t-t 

thick; endexine very thin, in many specimens not 

clearly visible. Sculptural elements consisting of 

small projections less than, or slightly greater than, 

0.5p. high, and less than 0.5ft in diameter, spaced 

at intervals almost equal to their diameter ; height 

of surface projections (scabrae) varies on individual tetrads and also on different grains of a 

single tetrad. 

Comparison with similar species and remarks.-

Because I. scabratus n. sp. is inaperturate, it is 

easily distinguishable from Dicotetradites granulatus Norton, in Norton and Hall (1969, p. 53, pl. 

5, fig. 4) whose individual grains are monoporate. 

I. scabratus n. sp. appears similar to Et·icaceoipollenites rallus Stanley (1965, p. 296-297, pl, 44, figs. 

15-18) from the Paleocene of South Dakota, except 

that the grains of the tetrad do not have colpi. 

Stanley described the grains of E. rallus as inaperturate to weakly tricolpate, but I have not observed 

colpi in any of the specimens of I. scabratu.s that I 

have examined. Compression folds sometimes resemble colpi but I do not interpret them as colpi. 

It is possible that I. scabratus, which I have found 

in the Campanian and Maestrichtian of Montana 

and Wyoming, will be found to be synonymous with 

Stanley's E. rallus from the Paleocene of South 

Dakota. Because I do not believe that my specimens 

have colpi, I have placed this new s.pecies in the 

genus Ina;perturotetradites. 

Stanley (1965, p. 296) interpreted the genus 

Ericaceoipollenites to include inaperturate to weakly 

tricolpate pollen grains that are arranged in a tetrahedral tetrad, and consequently he placed his new 

species (rallus) in that genus. He either ignored or 

did not agree to (but did not mention) Manum's 

(1962, p. 29) earlier synonymy of Ericaceoipollenites under Ericipites. I believe, as did Stanley, that 

Ericaceoipollenites and Ericipites were separable. 

Stanley (1965, p. 296) pointed out that Ericaceoipollenites is separable from Ericipites by the absence of well-defined colpi. It seems to me that the 

two genera were easily distinguishable on the basis 

that Ericaceoipollenites Potonie (1951 ex Potonie, 

1960, p. 138) was described as having furrows relatively short and at times indistinct, whereas Ericip494-146 0 - 73 - 5 

ites Wodehouse (1933, p. 516-517) was described as 

having pores in addition to colpi. Manum did not 

emend Ericipites to include tricolpate as well as 

tricolporate forms. I tentatively accept Manum's 

synonymy because it was agreed to by Potonie 

(1966, p. 192), the author of Ericaceoipollenites, 

but I believe that Ericipites should be emended to 

include tricolpate in addition to tricolporate pollen. 

Genus DROSERIDlTES Cookson, 1947 ex Potoni~, 1960 

Tetradites (Droseridites) Cookson, 1947, p. 137. 

Droseridites Cookson, 1947 ex Potonie, 1960, p. 139. 

Type species.-Droseridites ( Tetradites (Droseridites)) spinosa (Cookson, 1947) Potonie, 1960; 

lectotype (selected by Potonie, 1960, p. 139) in 

Cookson, 1947, p. 137, pl. XVI, fig. 70; Waterfall 

Gorge, Kerguelen Archipelago; Tertiary. 

Droseridites spinulosus Manum, 1962 

Droseridites spinulosus Manum, 1962, p. 30-31, pl. V, figs. 

1-5. 

Holotype.-In Manum, 1962, p. 30-31, pl. V, fig. 

1; Spitsbergen; Tertiary. 

Droseridites cf. D. spinulosus 

Plate 11, figure 20; table 1, No. 93 

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Series of Iowa: Am. Midland Naturalist, v. 23, no. 1, 

p. 182-186,pl. 1. 

Wodehouse, R. P., 1933, Tertiary pollen-[Part] II, The oil 

shales of the Eocene Green River formation: Torrey 

Bot. Club Bull., v. 60, no. 7, p. 479-524. 

Wolf, Monika, 1964, Sporomorphen aus dem bayerischen 

Flysch: Fortschr. Geologie Rheinland u. Westfalen, v. 

12, p. 113-116, pl. 1. 

Wolff, Herbert, 1934, [Part] II, Mikrofossilien des pliocanen 

Humodils der Grube Freigericht bei Dettingen a. M. 

und Vergleich mit alteren Schichten des Tertiars sowie 

posttertiaren Ablagerungen : Preussische Geol. Landesanst. Abh., v. 5, p. 55-86, pl. 5. 
INDEX 

[Italic page numbers indicate major references and descriptions] 

Page 

A 

A bietineae-pollenites 14 

Abietineaepollenites ---------------- 14 

varius -------------------- 5, 15; pl. 5 

(Piceaepollenites) microalatus ____ 14 

accipiteris, Aquilapollenites ---------- 28 

Acritarchs ------------------------

adenanthoides, Proteacidites --------- 32 

Aesculiidites ---------------------- 21,26 

dubius -------------------- 4, 21; pl. 8 

(Tricolpites) circumstriatus _____ 21,26 

alatipollenites, Pinus --------------- 15 

Pityosporites ------------ 5, 15, 16; pl. 6 

Alisporitcs ------------------------ 14, 17 

bilateralis --------------------- 14 

grand~ ------------------- ~1~ pl. 5 

microsaccus ------------------- 17 

opii -------------------------- 14 

rotundus ---------------------- 14 

thomasii ------------------ 4,14; pl. 5 

(Pteruchipollenites) microsaccus __ 17 

(Alisporites) microsaccus, 

Pristinuspollenites ------- 16 

a/pinus, Subtriporopollenites ____ 4, 31; pl. 11 

amplus, Aquilapollenites ------------ 27 

anchoriforme, Mancicorpus ---------- 29 

Anemia radiata -------------------- 8 

antarcticus, Pityosporites ----------- 15 

anulatus, Subtriporopollenites 

anulatus --------------- 31 

anulatus, Subtriporopollenites ____ 31 

Apiculatisporis wonthaggiensis ------- 8 

Aquilapollenites ------------------- 26,29 

accipiteris --------------------- 28 

am,plus ----------------------- 27 

ascriptivus -------------------- 28 

asper -------------------------- 27 

attenuatus -------------- 4, 6, 27; pl. 10 

aucellatus --------------------- 29 

augustus ---------------------- 28 

calvus ------------------------ 29 

clarireticulatus ------------- 5, 27; pl. 9 

drumhellerens~ ---------------- 29 

granulatus -------------------- 27 

h~pidus ----------------------- 28 

insignis ----------------------- 29 

latilobus ---------------------- 29 

tnatsumotoi -------------------- 28 

quadrilobus -------- 4, 6, 26, 27, 29; pl. 9 

regal~ ------------------------ 28 

reticulatus -------------- 5, 6, 27; pl. 9 

senonicus ---------------- 5, 6, 27; pl. 9 

subtilis ----------------------- 29 

trialatus ------------- 5, 6, 26, 27; pl. 9 

trialatus ------------------- 27 

uniformis -------------- 4, 27; pl. 9 

tripodiformis ------------------ 29 

turbidus ----------------- 5, 27; pl. 10 

sp ----------------------- 4, 27; pl. 9 

( Aquilapollenites) pule her, 

Hemicorpus ------------ 27,29 

Arecipites ------------------------ 20 

puncbtus --------------------- 20 

sp ----------------------- 4,20; pl. 8 

Page 

ascriptivus, Aquilapollenites --------- 28 

asper, Aquilapollenites -------------- 27 

attenuatus, Aquilapollcnites ___ 4, 6, 27; pl. 10 

aucellatus, Aqui/apollenites ---------- 29 

augustus, Aquilapollenites ----------- 28 

australis, Cyathidites --------------- 6 

B 

bacata, Cranwellia ----------- 4, 6, 26; pl. 9 

( Bacubivesiculites) inchoatus, 

Pristinuspollenites _____ 16; pl. 7 

B:ICulatisporites 7 

primarius --------------------- 7 

sp ------------------------ 5, 7; pl. 1 

barbatus, Seductisporites ____ 5, 6, 12, 13; pl. 4 

biformis, Podocarpidites ------------ 16 

bilateralis, Alisporitcs -------------- 14 

Bisaccates ------------------------ 4, 14 

blancnsis, Zlivisporis --------------- 13 

c 

callosus, Umbosporites __________ 4, 11; pl. 3 

ca./vus, Aquilapollenites ------------- 29 

Mancicorpus -------------- 4, 29; pl. 10 

Camarozonosporites ---------------- 11 

(Rotaspora) cretaceus ---------- 11 

spp ---------------------- 4,11; pl. 3 

canadensis, Grewipollenites ------- 21, 22, 26 

canadianus, Mancicorpus ------------ 29 

Mancicorpus (Mtchedlishvilia) ____ 29 

canaliculatus, Microfoveolatosporis_ 4, 11; pl. 3 

catteralli, Sindorapoll~ -------------- 30 

Sindorapollis (Loranthacites) ____ 29 

Caytoniales-Pollenites --------------- 18 

Caytonipollenites ------------------- 18 

pallidus ----------------------- 18, 19 

Cedripites ------------------------- 15 

cretaceus ----------------- 5, 15; pl. 5 

eocenicus ---------------------- 15 

parvus -------------------- 4, 15; pl. 5 

chinleana, Equ~etosporites ---------- 20 

chlonovae, Taurocusporites ----------

Trisol~sporites ----------- 5, 6, 10; pl. 2 

Cicatricosi-sporites ----------------- 8 

Cicatricosisporites ------------------ 8 

dorogensis --------------------- 8 

radiatus ------------------- 4, 8; pl. 1 

stoveri --------------------- 5, 8; pl. 1 

sp ------------------------ 4,8;pl. 1 

Circularesporites ------------------- 32 

circulus, Polycingulatisporites --------

circumstriatus, Aesculiidites 

(Tricolpites) ----------- 21,26 

Claggett Shale -------------------- 1 

clarireticulatus, Aquilapollenites ___ 5, 27; pl. 9 

Integricorpus ------------------ 27 

classoides, Classopollis -------------- 20 

Classopollis ----------------------- 20 

classoides --------------------- 20 

spp ---------------------- 5,20;p1. 8 

Clavabivesiculitcs inchoatus ----------- pl. 7 

Page 

( Clavabivesiculites) pannosus, 

Pristinuspollenites 17; pl. 7 

( Clavabivesiculites inchoatus) crassus, 

Pristinuspollenites ________ pl. 7 

chlonovae, Trisolissporites 

( Taurocusporites) 9 

complexus, Liliacidites -------- 4, 6, 20; pl. 8 

Schizosporis ------------------- 20 

Conosmundasporites ---------------- 7 

constrictus, Pityosporites -------- 5, 15; pl. 5 

convolutus, Rugubivesic1dites -------- 18 

Cornacwipollenites ----------------- 24 

costatus, Triatriopollenites ------ 5, 31; pl. 11 

Cmnwellia -------------------- 5, 26; pl. 9 

bacata ------------------ 4, 6, 26; pl. 9 

rumseyensis -------------- 5, 6, 26; pl. 9 

(Elytranthe) striata ------------ 26 

crassus, Pristinuspollenites 

( Clavabivesiculites 

inchoatus) 

Triprojectus -------------------

crateris, Ischyosporites --------------

cretaceus, Camarozonosporites 

pl. 7 

29 

8 

(Rotaspora) ------------ 11 

Cedripites ----------------- 5, 15; pl. 5 

Cupanieidites ---------------------- 30 

major ------------------------ 30 

sp ---------------------- 5,30; pl. 10 

Cyathidites ----------------------- 6 

australis ---------------------- 6 

minor --------------------- 4, 6; pl. 1 

sp ------------------------ 5,6;pl. 1 

D 

Dacrydium transiens 17 

Dacrydiumites --------------------- 16, 17 

Deltoidospora ---------------------- 6 

hallU ------------------------- 6 

sp ------------------------ 4, 6; pl. 1 

Depository of types and other 

illustrated specimens _____ 3 

Dicotetradites granulatus ------------ 33 

Dinoflagellates --------------------- 2 

Disaccites grandis ------------------ 14 

(Podocarpidites) --------------- 16 

dispositus, Triprojectus -------------- 27 

dorogensis, Cicatricosisporites -------- 8 

Droseridites ----------------------- 33 

spinulosus ---------------- 5, 38; pl. 11 

( Tetradites ( Droseridites) ) spinosa_ 33 

(Droseridites), Tetradites ------------ 33 

drumhellerensis, Aquilapollenites ______ 29 

dubius, Aesculiidites ------------ 4, 21; pl. 8 

Striopollenites ----------------- 21 

dulhuntyi, Dulhuntyispora ---------- 9; pl. 2 

Dulhuntyispora -------------------- 3, 9 

dulhuntyi ---------------------- 9 

(Dulhuntyispora minuta) minutus, 

Trisol~sporites ---------- 9 

Duplopollis ------------------------ 30 

Dyadites -------------------------- 32 

Dyadonapites --------------------- 32 

reticulatus ------------ 4, 5, 6, 32; pl. 11 

Dyads ---------------------------- 32 

39 
40 

Page 

E 

ellipticus, Podocarpidites ----------- 16 

elongatus, Pinuspollenites ------------ 15 

Pityosporites -------------- 15, 16; pl. 6 

Pityosporites elongatus ----------

elongatus elongatus, Pityosporites ---- 5 

grandis, Pityosporites ------- 5, 15; pl. 6 

Elytranthe ------------------------ 26 

(Elytranthe) striata, Cranwellia ----- 26 

eminens, Seductisporites ------ 5, 6, 12; pl. 4 

endannulatus, Montanapollis ___ 4, 6, 81; pl. 11 

eocenicus, Cedripites ---------------- 15 

Equisetosporites -------------------- 20 

chinleana ---------------------- 20 

spp ---------------------- 4,20; pl. 8 

Erdtmanipollis -------------------- 32 

pachysandroides ----------- 4, 32; pl. 11 

Ericaceoipollenites ------------------ 33 

rallus ------------------------ 33 

Ericipites 

crugatus, Tricolpites ---------------

Eucommiidites ---------------------

minor ------------------------

troedssonii --------------------

F 

33 

25 

25 

25 

25 

Fasciatisporites -------------------- 10 

Fibulapollis ----------------------- 26 

scabratus ---------------- 5, 6, 26; pl. 9 

( Triporina) mirificus ----------- 26 

foraminis, Foraminisporis ----------- 8 

Foraminisporis -------------------- 8 

foraminis --------------------- 8 

wonthaggiensis -------------- 4, 8; pl. 2 

G 

gemmiferus, Seductisporites _ 5, 6, 11, 13; pl. 3 

Ghoshispora ----------------------- 8 

scollardiana ------------------- 8 

sp ---------------------- 1,~ 8;pl. 1 

Gleicheniidites --------------------- 10 

senonicus --------------------- 10 

spp ---------------------- 4,11;pl. 3 

Granabivesiculites inchoatus _______ 17; pl. 7 

( Granabivesiculites) inchoatus, 

Pristinuspollenites _____ 17; pl. 7 

grandis, Alisporites ------------ 4, H; pl. 5 

Disaccites --------------------- 14 

Pityosporites ------------------ 14 

elongatus ----------- 5, 15; pl. 6 

granisaccus, Pristinuspollenites 

(Parvisaccites) --------- 17 

granospeciosus, Rouseisporites -------- 12 

granulatus, Aquilapollenites --------- 27 

Dicotetradites ------------------ 33 

Sindorapollis ------------ 5, 6, 30; pl. 10 

Grewipollenites -------------------- 21 

canadensis ----------------- 21, 22, 26 

radiatus ----------------- 5, 6, 21; pl. 8 

Gun;,era microreticulata ------------- 23 

H 

hallii, Deltoidospora 6 

Hamulatisporis -------------------- 11 

Hemicorpus ----------------------- 27,29 

(Aquilapollenites) pulcher ------- 27, 29 

hians, Tricolpites ---------- 3, 4, 7, 24; pl. 8 

Tricolpopollenites -------------- 24 

hiatipites, Taxodium --------------- 19 

hiatus, lnaperturopollenites ---------- 19 

Pollenites --------------------- 19 

Taxodiaceaepollenites -------- 4, 19; pl. 8 

(Pollenites) ------------ 19, 20 

hiatus, Taxodiaceae-Poll. ------------ 19 

INDEX 

Page 

hiatus, Taxodiaceae-Poll. ------------ 19 

hiatus, Taxodiaceae-pollenites -------- 20 

hiatus, Taxodieaepollenites ---------- 19 

Taxodieae-Poll. ---------------- 19, 20 

hiatus, Taxodieae-pollenites ---------- 20 

Taxodioipoll. ------------------- 19 

Taxodoidites ------------------- 19 

Thuja ------------------------ 19 

hiatus paleocaenicus, Pollenites ------ 19 

hispidus, Aquilapollenites ----------- 28 

igniculus, Sporonites --------------- 14 

Tsugaepollenites --------------- 14 

(Sporonites) ----------- 14 

Zonalapollenites ---------------- 14 

(Sporonites) --------------- 13 

Inaperturates 4 

lnaperturopollenites ---------------- 19, 20 

hiatus ------------------------ 19 

lnaperturotetradites ---------------- 32, 33 

lacunosus -------------------- 32_ 

scabratus -------------- 5, 32, 33; pl. 11 

inchoatus, Clavabivesiculties ---------- pl. 7 

Granabivesiculites ------------ 17; pl. 7 

Pristinuspollenites 

( Bacubivesiculites) 16; pl. 7 

( Granabivesiculites) 17; pl. 7 

lndusiisporites --------------------- 17 

1~nsignis, Aquilapollenites ------------ 29 

lntegricorpus clarireticulatus ________ 27 

interangulus, Tricolpites ------------ 23 

lschyosporites --------------------- 8 

crateris __ --------------------- 8 

punctatus ------------------ 5, 8; pl. 1 

.J 

Judith River Formation -------------

Juniperus 

K 

1, 6 

19 

kaitangataensis, Liliacidites --------- 20 

krempii, Pulcheripollenites _______ 4, 22; pl. 8 

Ulmoideipites ---------------.--- 31 

Kuylisporites ---------------------- 8 

waterbolki -------------------- 8 

spp 4, 8; pl. 1 

L 

lacunosus, lnaperturotetradites 

Laevigato-sporites -----------------

32 

11 

Laevigatosporites ------------------ 11 

vulgaris ---------------------- 11 

spp ---------------------- 4, 11;pl. 3 

laevigatus, Rouseisporites ----------- 13 

latilobus, Aquilapollenites ------------ 29 

Leptolepidites --------------------- 7 

major --------------------- 4, 7; pl. 1 

verrucatus -------------------- 7 

levitas, Tricolpopollenites ____ 3, 4, 7, 25; pl. 9 

liblarensis, Tricolpopollenites -------- 23 

Libocedrus ------------------------ 19 

ligneolus, Ovoidites ------------- 5, 21; pl. 8 

Pollenites --------------------- 21 

Sporites ----------------------- 21 

Liliacidites ----------------------- 20 

complexus --------------- 4, 6, 20; pl. 8 

kaitangataensis ----------------- 20 

morrinensis ---------------- 5, 20; pl. 8 

variegatus ----------------- 5, 20; pl. 8 

spp ---------------------- 5,21;pl.8 

Liquidambarpollenites --------------- 32 

Liquidambar-pollenites -------------- 32 

Loranthacites 29 

Paee 

( Loranthacites) catteralli, 

Sindorapollis ------------ 29 

pilatus, Sindorapollis ------------ 29 

Lycopodium novomexicanum --------- 12 

Lycopodiumsporites novomexicanum 12 

M 

major, Cupanieidites --------------- 30 

Leptolepidites --------------- 4, 7; pl. 1 

Mancicorpus ---------------------- 26, 29 

anchoriforme ------------------ 29 

calvus ------------------- 4, 29; pl. 10 

canadianus -------------------- 29 

senonicum --------------------- 27 

tripodiformis ------------- 5, 29; pl. 10 

(Mtchedlishvilla) canadianus ----- 29 

matsumotoi, Aquilapollenites --------- 28 

maximus, Podocarpidites ------------ 16 

microalatus, Abietineaepollenites 

(Piceaepollenites) 14 

Microfoveolatosporis ---------------- 11 

canaliculatus --------------- 4, 11; pl. 3 

pseudodentatus ----------------- 11 

microreticulata, Gunnera ------------ 23 

microreticulatus, Pristinuspollenites 

(Phyllocladidites) -------- 17 

Retitricolpites ------------------ 23 

Tricolpites -------------------- 23 

microsaccus, Alisporites ------------- 17 

Alisporites (Pteruchipollenites) ___ 17 

Pristinuspollenites -------- 5, 6, 17; pl. 7 

(Alisporites) --------------- 16 

(Pteruchipollenites) --------- 16 

Pteruchipollenites --------------- 17 

minimus, Rugubivesiculites ---------- 18 

minor, Cyathidites -------------- 4, 6; pl. 1 

Eucommiidites ----------------- 25 

minutus, Trisolissporites -------------

Trisolissporites ( Dulhuntyispora 

minuta) --------------- mirificus, Fibulapollis (Triporina) __ _ 

Miscellany ------------------------

Monoletes -------------------------

10 

9 

26 

19 

11 

Monosulcates ---------------------- 4, 20 

Montanapollis --------------------- 31 

endannulatus ------------ 4, 6, 31; pl. 11 

montanensis, Siberiapollis ---- 5, 6, 31; pl. 11 

Trisolissporites ---------- 5, 6, 8, 9; pl. 2 

morrinensis, Liliacidites --------- 5, 20; pl. 8 

Mtchedlishvilia -------------------- 29 

( M tchedlishvilia) canadianus, 

Mancicorpus ------------ 29 

multesimus, Podocarpidites ------ 5, 16; pl. 7 

Podocarpus -------------------- 16 

N 

narcissus, Pulcheripollenites --------- 22 

novomexicanum, Lycopodium 

Lycopodiumsporites ------------- 12 

0 

opii, Alisporites ------------------- 14 

ornatus, Podocarpidites --------- 5, 16; pl. 7 

Osmundacidites -------------------- 7 

wellmanii --------------------- 8 

sp ------------------------ 5,8;pl. 1 

Ovoidites ------------------------- 21 

ligneolus ------------------ 5, 21; pl. 8 

p 

pachysandroides, Erdtmanipollis __ 4, 32; pl. 11 

paleocaenicus, Pollenites hiatus ------- 19 

pallidus, Caytonipollenites ----------- 18, 19 

Pityopollenites ----------------- 18 

Pityosporites ------------------ 18 

Vitreisporites -------------- 5, 18; pl. 8 
Page 

pannosus, Pristinuspollenties 

( Clavabivesiculites) 17; pl. 7 

Parcisporites ---------------------- 17 

parmularius, Tricolpopollenites _______ 24 

Tricolpopollenites (Pollenites) ____ 24 

Parviprojectus reticulatus _ _ _ _ _ _ _ _ _ _ _ 27 

( Parvisaccites) granisaccus, 

Pristinuspollenites _______ 17 

parvus, Cedripites ------------- 4, 15; pl. 5 

Tricolpites -------------------- 24 

Periporopollenites ------------------ 32 

(Pollenites) stigmosus ---------- 32 

sp -------------------- 5,6,32;pl. 11 

Peroaletes ------------------------ 11 

Phaseolites ----------------------- 11 

Phyllocladidites -------------------- 17 

( Phyllocladidites) microreticulatus, 

Pristinuspollenites ------- 17 

( Piceaepollenites) microalatus, 

Abietineaepollenites ------ 14 

pilatus, Sindorapollis --------------- 30 

Sindorapollis (Loranthacites) _____ 29 

Pinus alatipollenites ---------------- 15 

ruginosa ---------------------- 18 

Pinuspollenites -------------------- 15 

elongatus ---------------------- 15 

Pityopollenites --------------------- 18 

pallid us 18 

Pityosporites ---------------------- 15 

alatipollenites ----------- 5, 15, 16; pl. 6 

antarcticus -------------------- 15 

constrictus ----------------- 5, 15; pl. 5 

elongatus ----------------- 15, 16; pl. 6 

elongatus ------------------ 5 

grandis ---------------- 5, 15; pl. 6 

grandis ----------------------- 14 

pallidus ---------------------- 18 

Podocarpidites ---------------------

biformis 

ellipticus ----------------------

16 

16 

16 

maximus ---------------------- 16 

multesimus ---------------- 5, 16; pl. 7 

ornatus ---------------- 5, 16; pl. 7 

(Podocarpidites), Disaccites --------- 16 

Podocarpus multesimus -------------- 16 

Poll. hiatus paleocaenicus ----------- 19 

Pollen-spore zones ----------------- 3 

Pollenites ------------------------- 20 

hiatus ------------------------ 19 

paleocaenicus --------------- 19 

ligneolus ---------------------- 21 

( Pollenites) hiatus, 

Taxodiaceaepollenites _____ 19, 20 

parmularius, Tricolpopollenites ___ 24 

stigmosus, Periporopollenites _____ 32 

Polycingulatisporites ----------- _____ 8 

circulus ----------------------- 8 

spp --------------------- 5,6,8;pl.2 

Polycolpates ----------------------- 29 

polyformosus, Sequoiapollenites ______ 20 

Polypodiaceae-sporites -------------- 11 

Polypodium-sporites ---------------- 11 

Polyporates ----------------------- 32 

primarius, Baculatisporites ---------- 7 

Pristinuspollenites ----------------- 16 

microsaccus -------------- 5, 6,17; pl. 7 

sulcatus ----------------------- 17, 18 

(Alisporites) microsaccus -------- 16 

(Bacubivesiculites) inchoatus ___ 16; pl. 7 

( Clavabivesiculites) pannosus __ 17; pl. 7 

( Clavabivesiculites inchoatus) 

crassus ----------------- pl. 7 

( Granabivesiculites) inchoatus ___ 17; pl. 7 

( Granabivesiculites inchoatus) 

sulcatus 

( Parvisaccites) granisaccus _____ _ 

( Phyllocladidites) microreticulatus_ 

(Pteruchipollenites) microsaccus __ 

17 

17 

17 

16 

INDEX 

Page 

Procedure of study ----------------- 2 

Processing of samples and photography 1 

prolata, Tricolporites --------------- 30 

Proteacidites ---------------------- 32 

adenanthoides ------------------ 32 

spp --------------------- ~32; pl. 11 pseudodentatus, Microfoveolatosporis __ 11 

Pteruchipollenites microsaccus ------- 17 

thomasii ---------------------- 14 

( Pteruchipollenites) microsaccus, 

Alisporites ------------- 17 

microsaccus, Pristinuspollenites ___ 16 

pulcher, Hemicorpus (Aquilapollenites) 27, 29 

Pulcheripollenites ------------------ 22 

krempii ------------------- 4, 22; pl. 8 

narcissus --------------------- 22 

puncta.tus, Arecipites --------------- 20 

Ischyosporites --------------- 5, 8; pl. 1 

quadrilobus, Aquilapollenites 

Q 

R 

4, 6, 26, 27, 29; pl. 9 

radiata, Anemia ------------------- 8 

radiatus, Cicatricosisporites ------- 4, 8; pl. 1 

Grewipollenites ----------- 5, 6, 21; pl. 8 

Rottseisporites ----------------- 12, 13 

Stenozonotriletes --------------- 9, 10 

Trisolissporites ----------------- 10 

(Stenozonotriletcs) ---------- 9 

(Taurocusporites) ----------- 9 

rallus, Ericaceoipollenites ----------- 33 

regalis, Aquilapollenties ------------- 28 

reticulata, Tricolpites --------------- 23 

reticulatus, Aquilapollenites ____ 5, 6, 27; pl. 9 

Cyadonapites ---------- 4,5, 6, 32; pl. 11 

Rouseisporites ----------------- 13 

Tricolpites ---------- 3, 4, 7, 22, 23; pl. 8 

Retimonocolpites ------------------- 20 

Retitricolpites microreticulatus ------- 23 

retriculatus, Parviprojectus ---------- 27 

rhomboides, Tricolporites ------- 4, 30; pl. 10 

( Rotaspora) cretaceus, 

Camarozcmosporites ------ 11 

rotundus, Alisporites --------------- 14 

Rouseisporites --------------------- 11 

granospeciosus ----------------- 12 

laevigatus --------------------- 13 

radiatus ----------------------- 12, 13 

reticulatus --------------------- 13 

rugatus, Triporopollenites ----------- 31 

ruginosa, Pinus -------------------- 18 

Rugubivesiculites ------------------- 18 

convolutus --------------------- 18 

minirnus ---------------------- 18 

sp ----------------------- 4,18; pl. 7 

rumseyensis, Cranwellia ------- 5, 6, 26; pl. 9 

rurensis, Triatriopollenites ----------- 31 

s 

Sample localities ------------------- 1 

sarstedtensis, Striatopollis ----------- 26 

scabratus, Fibulapollis ________ 5, 6, 26; pl. 9 

Inaperturotetradites _____ 5, 32, 33; pl. 11 

Schizosporis complexus ------------- 20 

scollardiana, Ghoshispora ----------- 8 

Seductisporites -------------------- 11 

barbatus ------------- 5, 6, 12, 13; pl. 4 

eminens ----------------- 5, 6, 12; pl. 4 

gemmiferus ----------- 5, 6, 11, 13; pl. 3 

signifer ----------------------- 11, 12 

segmentatus, Taurocusporites -------- 9 

senonicum, Mancicorpus ------------- 27 

senonicus, Aquilapollenites _____ 5, 6, 27; pl. 9 

Gleicheniidites ----------------- 10 

41 

Page 

Sequoia-pollenites ------------------ 20 

Sequoiapollenites ------------------ 20 

polyformosus ------------------ 20 

spp -----------------------5• 20; pl. 8 

Siberiapollis ----------------------- 31 

montanensis ------------- 5, 6, 31; pl. 11 

signatus, Vitreisporites ------------- 18, 19 

signifer, Seductisporites ------------- 11, 12 

Sindorapollis ---------------------- 29 

catteralli ---------------------- 30 

granulatus ------------- 5, 6, 30; pl. 10 

pilatus ------------------------ 30 

triangulus --------------------- 30 

(Loranthacites) catteralli -------- 29 

pilatus -------------------- 29 

Sphagnites ------------------------ 7 

Sphagnum-sporites ----------------- 6 

Sphagnumsporites -----------------

spinosa, Droseridites ( Tetradites 

(Droseridites)) --------- 33 

spinulosus, Droseridites -------- 5, 33; pl. 11 

Sporites ligneolus ------------------ 21 

(Sporites) stereoides, Stereisporites --- 7 

Sporonites ------------------------ 14 

igniculus ---------------------- 14 

( S poronites) igniculus, 

Tsugaepollenites --------- 14 

(Spronites) igniculus, Zonalapollenites_ 13 

stellatus, Stenozonotriletes -----------

Trisolissporites ----------------- 10 

(Stenozonotriletes) ---------- 9 

( Taurocusporites) ----------- 9 

Stenozonotriletes ------------------- 9 

radiatus ---------------------- 9, 10 

stellatus ---------------------- 9 

(Stenozonotriletes) radiatus, 

Trisolissporites ---------- 9 

stellatus, Trisolissporites --------

Stereisporites --------------------- 6 

(Sporites) stereoides ------------ 7 

spp ----------------------- 4, 7; pl. 1 

stereoides, Stereisporites ( S porites) _ __ 7 

stigmosus, Periporopollenites 

(Pollenites) ----------- 32 

stoveri, Cicatricosisporites -------- 5, 8; pl. 1 

striata, Cranwellia (Elytranthe) ------ 26 

Striatopollis ----------------------- 21, 26 

sarstedtensis ------------------- 26 

Striopollenites --------------------- 21 

dubius ------------------------ 21 

Styx ----------------------------- 8 

subtilis, Aquilapollenites ------------ 29 

Subtriporopollenites ---------------- 31 

alpinus ------------------- 4, 31; pl. 11 

anulatus anulatus -------------- 31 

sulcatus, Pristinuspollenites ---------- 17,18 

Systematics ----------------------- 6 

T 

Taurocusporites 

chlonovae ---------------------

segmentatus -------------------

( Taurocusporites) chlonovae, 

Trisolissporites ----------

radiatus, Trisolissporites --------

9, 10 

9, 10 

9 

stellatus, Trisolissporites -------- 9 

Taxodiaceae ---------------------- 19,20 

Taxodiaceaepollenites --------------- 19 

hiatus -------------------- 4, 19; pl. 8 

(Pollenites) hiatus ------------- 19,20 

Taxodiaceae-Pollenites -------------- 19 

Taxodiaceae-Poll. hiatus ------------ 19 

Taxodiaceae-Poll. hiatus ------------

Taxodiaceae-pollenites hiatus --------

Taxodi€ae ------------------------

Taxodieacpo:lcnites hiatus -----------

19 

20 

19 

19 
42 

Page 

Taxodieae-Poll. -------------------- 19, 20 

hiatus ------------------------ 19,20 

Taxodieae-pollenites hiatus ---------- 20 

Taxonioidites ---------------------- 19 

Taxodioipoll. hiatus ---------------- 19 

Taxodium ------------------------- 19 

hiatipites 19 

Taxodoidites ---------------------- 19 

hiatus ------------------------ 19 

tessellatus, Tricolpopollenites __ 3, 4, 7, 25; pl. 9 

Tetradites (Droseridites) ------------ 33 

( Tetradites ( Droseridites) ) spinosa, 

Droseridites ------------ 33 

Tetrads --------------------------- 32 

thomasii, Alisporites ----------- 4, 14; pl. 5 

Pteurchipollenites -------------- 14 

Thuja ---------------------------- 19,20 

hiatus ------------------------ 19, 20 

transiens, Dacrydium --------------- 17 

trialatus, Aquilapollenites ____ 5, 6, 26, 27; pl. 9 

Aquilapollenites trialatus --------- 27 

trialatus, Aquila.pollenites __ ______ 27 

uniformis, Aquilapollenites ____ 4, 27; pl. 9 

triangulus, Sindorapollis ------------- 30 

Triatriopollenites ------------------ 31 

costatus ------------------ 5, 31; pl. 11 

rurensis ----------------------- 31 

Tricolpates ----------------------- 4, 21 

Tricolpites ------------------------ 22 

erugatus ---------------------- 25 hians __________________ 3, 4, 7, 24; pl. 8 

interangulus ------------------- 23 

microreticulatus ---------------- 23 

parvus ------------------------ 24 

reticulata --------------------- 23 

reticulatus ---------- 3, 4, 7, 22, 23; pl. 8 

varius ------------------------ 24 

spp ---------------------- 4,24; pl. 8 

(Tricolpites) circumstriatus, 

Aesculiidites ------------ 21, 26 

INDEX 

Page 

Tricolpopollenites --------------- 22, 24, 26 

hians ------------------------- 24 

levitas ---------------- 3, 4, 7, 25; pl. 9 

liblarensis --------------------- 23 

parmularius ------------------- 24 

tessellatus ------------- 3, 4, 7, 25; pl. 9 

(Pollenites) parmularius -------- 24 

Tricolpora tes 4 

Tricolporites prolata --------------- 30 

rhomboides --------------- 4, 30; pl. 10 

sp ---------------------- 5, 31; pl. 10 

tricostatus, Ulmoideipites ----------- 31 

Triletes -------------------------- 4, 6 

tripodiformis, Aquilapollenites -------- 29 

Mancicorpus -------------- 5, 29; pl. 10 

Triporates ------------------------ 5 

(Triporina) mirificus, Fibulapollis ___ 26 

Triporopollenites rugatus ------------ 31 

Triprojectus crassus --------------- 29 

dispositus --------------------- 27 

unicus ------------------------ 28 

Triremisporites -------------------- 10 

Trisolissporites ------------------- 3, 8, 10 

chlonovae --------------- 5, 6, 10; pl. 2 

minutus ---------------------- 10 

montanensis ------------ 5, 6, 8, 9; pl. 2 

radiatus ----------------------- 10 

stellatus ---------------------- 10 

(Dulhuntyispora minuta) ninutus__ 9 

(Stenozonotriletes) radiatus ------ 9 

stellatus ---------------------- ( Taurocusporites) chlcmovae ____ _ 

radiatus ------------------- 9 

stellatus ------------------- 9 

troedssonii, Eucommiidites ----------- 25 

Tsugae-pollenites ------------------ 13, 14 

Tsugaepollenites ------------------- 13, 14 

igniculus ---------------------- 14 

(Sporonites) igniculus ----------- 14 

turbidus, Aquilapollenites ------- 5, 27; pl. 10 

Page 

u 

Ulmoideipites krempii -------------- 31 

tricostatus 31 

Umbosporites --------------------- 11 

callosus ------------------- 4, 11; pl. 3 

Unclassified bisaccates -------------- 4 

Unclassified inaperturates ----------- 4 

Unclassified monosulcates ----------- 4 

Unclassified triletes ---------------- 4 

unicus, Triprojectus ---------------- 28 

uniformis, Aquilapollenites 

trialatus 4, 27; pl. 9 

v 

variegatus, Liliacidites ---------- 5, 20; pl. 8 

varius, Abietineaepollenites ------ 5, 15; pl. 5 

Tricolpites -------------------- 24 

11errucatus, Leptolepidites -----------

Vitreisporites --------------------- 18 

pallidys ------------------- 5, 18; pl. 8 

signatus ---------------------- 18, 19 

vulgaris, Laevigatosporites ---------- 11 

w 

waterbolki, Kyulisporites ------------ 8 

wellmani, Osmundacidites ----------- 8 

wonthaggiensis, Apiculatisporis ------ 8 

Foraminisporis -------------- 4, 8; pl. 2 

z 

Zamia 19 

Zlivisporis ------------------------ 11 

blanensis ---------------------- 13 

Zonalapollenites ------------------- 13 

igniculus ---------------------- 14 

(Sporonites) igniculus ---------- 13 

sp ----------------------- ~1~pl.5 

Zonates -------------------------- 11 

U. S. GOVERNMENT PRINTING OFFICE: 1973 0- 494-146 
PLATES 1-11 

Contact photographs of the plates in this report are available, at cost, from U.S. 

Geological Survey Library, Federal Center, Denve·r, Colorado 80225. 
PLATE 1 

[All figures X 1,000 exeept as otherwise indieated] 

FIGURE 1. Cyathidites minor Couper (p. 6). 

USGS paleobotany loc. D4067, slide (3) at 78.3 X 12.0. 

2. Cyathidites sp. (p. 6). 

USGS paleobotany loc. D3726-E, slide (6) at 104.8 X 17.0. 

3. Deltoidospora sp. 1 p. 6). 

USGS paleobotany loc. D4067, slide (10) at 111.1 X 4.1. 

4, 5. Stereisporites spp. (p. 7). 

4. USGS p,aleobotany loc. D3726-D, slide (2) at 99.0 X 20.2. 

5. USGS paleobotany loc. D3726-D, slide (2) at 109.5 X 7.9. 

6. Leptolepidites major Couper (p. 7). 

USGS paleobotany loc. D3726-E, slide (6) at 88.1 X 20.3. 

7. Leptolepidites cf. L. major (p. 7). 

USGS paleobotany loc. D4067, slide (4) at 96.9 X 12.2. 

8. Baculatisporites sp. (p. 7). 

USGS paleobotany loc. D3725-B, slide (2) at 75.9 X 13.5. 

9. Osmundacidites sp. (p. 8). 

USGS paleobotany loc. D3726-E, slide (2) at 110.8 X 4.4. 

10, 11. Kuylisporites spp. (p. 8). 

10. USGS paleobotany loc. D4067, slide (1) at 94.3 X 3.4. 

11. USGS paleobotany loc. D4067, slide (4) at 86.6 X 2.3. 

12. Ischyosporites cf. I. punctatus (p. 8). 

USGS paleobotany loc. D3725-E, slide (2) at 100.4 X 15.6. 

13. Cic,atricosisporites stoveri Pocock {p. 8). 

USGS paleobotany loc. D3726-E, slide (5) at 105.1 X 19.9. 

14. Cicatricosisporites sp. (p. 8). 

USGS paleobotany loc. D3726-A, slide (5) 93.2 X 17.1. 

15, 16. Cicatricosisporites radiatus Krutzsch (p. 8). 

Two levels. USGS paleobotany loc. D4067, slide (4) at 97.7 X 10.2. 

17. Ghoshispora sp. (p. 8). 

USGS paleobotany loc. D4067, slide (11) at 92.3 X 18.0. (X 500). 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE I 

14 - . l 

15 16 17 

CY ATHIDITES, DELTOIDOSPORA, STEREISPORITES, LEPTOLEPIDITES, BACULATISPORITES, 

OSMUNDACIDITES, KUYLISPORITES, ISCHYOSPORITES, CICATRICOSISPORITES, AND GHOSHISPORA 

4 
PLATE 2 

[All figures X 1,0001 

FIGURES 1, 2. Foraminisporis wonthaggiensis (Cookson and Dettmann) Dettmann 

(p. 8). 

1. USGS paleobotany loc. D3726-D, slide (2) at 78.8 X 5.6. 

2. USGS paleobotany Joe. D3725-A, slide (2) at 92.6 X 21.3. 

3-5. Polycingulatisporites spp, (p. 8). 

3. USGS paleobotany loc. D3726-E, slide (4) at 99.8 X 3.7. 

4. USGS paleobotany loc. D3726-E, slide (9) at 99.4 X 8.8. 

5. USGS paleobotany loc. D3726-C, slide (2) at 83.1 X 21.7. 

6-10. Trisolissporites montanensis n. sp. (p. 9). 

6, 7. Holotype, two focal levels. USGS paleobotany loc. D3726-E, 

slide (3) at 99.9 X 11.3. 

8. Paratype. USGS paleobotany Joe. D3726-E, slide (2) at 81.8 

X 11.2. 

9, 10. Two focal levels. USGS paleobotany Joe. D3726-E, slide, (1) 

at 118.8 X 1.8. 

11. Dulhuntyispora dulhuntyi Potonie (p. 9). 

Specimen from Upper Permian from Western Australia, furnished 

by Dr. B. E. Balme. 

12-14. Trisolissporites chlonovae (Doring) n. comb. (p. 10). 

12. USGS paleobotany Joe. D3726-E, slide (2) at 93.0 X 8.9. 

13. USGS paleobotany loc. D3726-E, slide (8) at 77.8 X 14.1. 

14. USGS paleobotany loc. D3726-E, slide (2) at 117.3 X 5.6. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE 2 

11 14 

FORAMINISPORIS, POLYCINGULATISPORITES, TRISOLISSPORITES, AND DULHUNTYISPORA 
PLATE 3 

[All figures X 1,000] 

FIGURES 1-4. Gleicheniidites spp. (p. 11). 

1. USGS paleobotany loc. D3726-E, slide (4) at 87.4 X 19.8. 

2. USGS paleobotany loc. D4067, slide (4) at 92.6 X 12.2. 

3. USGS paleobotany loc. D4067, slide (4) at 100.9 X 21.1. 

4. USGS paleobotany loc. D4067, slide (4) at 112.0 X 16.4. 

5, 6. Camarozonosporites spp. ( p. 11). 

5. USGS paleobotany loc. D3726-D, slide ( 1) at 80.9 X 8.0. 

6. USGS paleobotany loc. D4067, slide (10) at 111.1 X 4.1. 

7, 8. Laevigatosporites spp. (p. 11). 

7. USGS paleobotany loe. D3726-A, slide (2) at 109.7 X 7.3 

8. USGS paleobotany Joe. D3726-A, slide (2) at 100.8 X 4.3. 

9,10. Umbosporites callosus Newman (p. 11). 

9. USGS paleobotany Joe. D4069, slide (1) at 91.9 X 5.0. 

10. USGS paleobotany Joe. 03726-C, slide (1) at 82.7 X 3.7. 

11. Microfoveolatosporis canaliculatus Dettemann (p. 11) . 

USGS paleobotany Joe. D3726-E, slide (4) at 104.7 X 5.3. 

12-17. Seductisporites gemmiferus n. sp. (p. 11). 

12, 13. Holotype, two focal levels. USGS paleobotany Joe. D3726-

A, slide (2) at 96.7 X 19.8. 

14, 15. Paratype, two focal levels, USGS paleobotany Joe. D3726-A, 

slide ( 1) at 91.4 X 14.1. 

16, 17. Paratype, two focal levels. USGS paleobotany Joe. D3726-

A, slide (2) at 92.4 X 18.3. 
GEOLOGICAL SURVEY 

1 

I 

( 

-

• 

8 

15 

PROFESSIONAL PAPER 770 PLATE3 

4 

2 

/ 

• 

9 10 

7 

/ 

• 

13 

17 

16 

GLEICHENIIDITES, CAMAROZONOSPORITES, LAEVIGATOSPORITES, 

UNBOSPORITES, MICROFOVEOLATOSPORIS, AND SEDUCTISPORITES 

5 

6 

14 
PLATE 4 

[All figures X 1,000] 

FIGURES 1-3. Seductisporites eminens n. sp. (p. 12). 

1. Holotype, USGS paleobotany loc. D3726-E, slide (5) at 99.0 X 

10.3. 

2. Paratype, USGS paleobotany loc. D3726-E, slide (9) at 84.8 X 

16.1. 

3. Paratype. USGS paleobotany loc. D3726-E, slide (8) at 109.2 

X 19.7. 

4-7. Seductisporites barbatus n. sp. (p. 13) . 

4. Holotype. USGS paleobotany loc. D3726-E, slide (9) at 111.3 X 

3.2. 

5. Paratype. USGS paleobotany loc. D3726-E, slide (7) at 82.0 

X 12.5. 

6, 7. Paratype, two focal levels. USGS paleobotany loc. D3726-E, 

slide (10) at 94.5 X 7.7. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE 4 

1 

3 

2 

5 

4 

SEDUCTISPORITES 
PLAT'E 5 

[All figures X 1,000] 

FIGURE 1. Zonalapollenites sp. (p. 14). 

USGS paleobotany loc. D3726-E, slide (4) at 100.8 X 21.6. 

2, 3. Alisporites grandis (Cookson) Dettmann (p. 14). 

2. USGS paleobotany loc. D3726-E, slide ( 5) at 117.5 X 2.5. 

3. USGS paleobotany loc. D3726-E, slide (6) at 102.2 X 8.8. 

4,5. Alisporitesthomasii (Couper) Nilsson (p. 14). 

4. USGS paleobotany loc. D3726-E, slide (6) at 106.9 X 15.1. 

5. USGS paleobotany loc. D3726-E, slide (6) at 97.3 X 10.5. 

6. Abietineaepollenites varius Norton I p. 15). 

USGS paleobotany loc. D3726-A, slide (2) at 90.1 X 12.0. 

7. Cedripites parvus Norton (p. 15). 

USGS paleobotany loc. D3726-E, slide (8) at 107.8 X 9.9. 

8. Cedripites cretac,eus Pocock ( p. 15). 

USGS paleobotany loc. D3726-C, slide (2) at 110.3 X 7.2. 

9. Pityosporites cf. P. constrictus (p. 15). 

USGS paleobotany loc. D3726-E, slide (6) at 113.2 X 6.7. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE 5 

ZON ALAPOLLENITES, ALISPORITES, ABIETINEAEPOLLENITES, 

CEDRIPITES, AND PITYOSPORITES 
PLATE 6 

[All figures X 1,000] 

FIGURE 1. Pityosporites alatipollenites (Rouse) Singh ( p. 15). 

USGS paleobotany loc. D3726-A, slide (2) at 86.0 X 8.3. 

2- 4. Pityosporites elongatus (Norton) Tschudyvar. elongatus (p. 15). 

2, 3. Two focal levels. USGS paleobotany loc. D3726-E, slide (6) at 

107.7 X 8.6. 

4. USGS paleobotany loc. D3726-E, slide (6) at 104.1 X 2.7. 

5, 6. Pityosporites elongatus var. grandis n. var. (p. 15). 

5. Paratype. USGS paleobotany loc. D3726-E, slide (6) at 96.9 X 

13.2. 

6. Holotype. USGS paleobotany loc. D3726-E, slide (5) at 116.0 X 

4.9. 
G.EOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE 6 

2 

PITYOSPORITES 
PLATE 7 

[All figures X 1,000] 

FIGURES 1, 2. Podocarpidites multesimus (Bolkhovitina) Pocock (p. 16). 

1. USGS paleobotany loc. D3726- E, slide (6) at 99.0 X 22.2. 

2. USGS paleobotany loc. D3726-E, slide (8) at 95.2 X 11.2. 

3. Podocarpidites cf. P. o1·natus (p. 16) . 

USGS paleobotany loc. D3725- A, slide (2) at 82.0 X 19.1. 

4-6. Pristinuspollenites microsaccus (Couper) n. comb. (p. 16, 17). 

4. USGS paleobotany loc. D3726- E, slide (6) at 108.8 X 21.2. 

5. USGS paleobotany loc. D3726- E, slide (4) at 87.2 X 8.0. 

6. USGS paleobotany loc. D3726-A, slide (2) at 99.4 X 19.2. 

7. Pristinuspollenites (Bacubiv esic.ulites) inchoatus (Pierce) n. comb. 

(p. 16). 

Holotype. (Also see Pierce 1961, pl. II, fig. 35). 

8, 9. Pristinuspollenites ( Clavabivesiculites inchoatus Pierce) crass us n. 

name (p. 16). 

Holotype, two focal levels. (See Clavabivesiculites inchoatus Pierce, 

1961, pl. II, g. 36.) 

10. Pristinuspollenites (Clavabivesiculites) pannosus (Pierce) n. comb. 

(p. 17) . 

Holotype. (Also see Pierce, 1961, pl. II, fig. 37.) 

11. Pristinuspollenites (Granabiv esiculties inchoatus Pierc·e) sulcatus n. 

name (p. 17). 

Holotype. (See Granabivesiculites inchoatus Pierce, 1961, pl. II, fig. 

38.) 

12, 13. Rugubivesiculites sp. (p. 18) . 

12. USGS paleobotany loc. D3726-E, slide (6) at 92.2 X 11.9. 

13. USGS paleobotany loc. D3725-A, slide (2) at 106.9 X 3.9. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE 7 

10 11 

PODOCARPIDITES, PRISTINUSPOLLENITES, AND RUGUBIVESICULITES 
FIGURES 1, 2. 

3,4. 

5, 6. 

7, 8. 

9, 10. 

11. 

12. 

13. 

14, 15. 

16-18. 

PLATE 8 

[All figures X 1,000] 

Vitreisporites pallidus (R€issinger) Nilsson (p. 18). 

1. USGS paleobotany loc. D3726-B, slide (1) at 73.3 X 5.9. 

2. USGS paleobotany loe. D3726-D, slide (2) at 97.2 X 12.0. 

Taxodiaceaepollenites hiatus ( Potonie) Kremp ex Potonie \ p. 19). 

3. USGS paleobotany Joe. D3726-D, slide (2) at 80.9 X 4.1. 

4. USGS paleobotany Joe. D3725- D, slide (2) at 88.1 X 2.7. 

Sequoiapollenites spp. 1 p. 20). 

5. USGS paleobotany Joe. D3726-E, slide (4) at 89.2 X 18.8. 

6. USGS paleobotany Joe. D3726-D, slide (2) at 104.5 X 18.6. 

Classopollis spp, ( p 20). 

7. USGS paleobotany Joe. D3725-E, slide (2) at 84.9 X 18.2. 

8. USGS paleobotany Joe. D3726-C, slide (2) at 115.3 X 18.5. 

Equisetosporites spp. 1 p. 20). 

9. USGS paleobotany Joe. D3726-E, slide (6) at 84.9 X 5.0. 

10. USGS paleobotany Joe. D4067, slide (1) at 86.9 X 21.2. 

Arecipites sp. (p. 20). 

USGS palebotany Joe. D3725-C, slide (4) at 81.0 X 17.8. 

Liliacidites variegatus Couper ( p. 20). 

USGS paleobotany Joe. D3725-C, slide (2) at 111.8 X 11.0. 

Liliacidites cf. L. morrinensis ( p. 20). 

USGS paleobotany Joe. D4070, slide (2) at 102.8 X 5.3. 

Liliacidites complexus (Stanley) n. comb. ( p. 20). 

14. USGS paleobotany loc. D4067, slide (1) at 105.1 X 20.1. 

15. USGS paleobotany loe. D4069, slide (2) at 87.0 X 2.1. 

Liliacidites spp. ( p. 21). 

16. USGS paleobotany Joe. D3726-A, slide (2) at 92.'21 X 2.3. 

17. USGS paleobotany Joe. D3726-A, slide (2) at 90.~ X 4.2. 

18. USGS paleobotany Joe. D3725- C, slide (5) at 91.3 X 22.1. 

19. Ovoidites ligneolus ( Potonie) Potonie ex Krutzsch ( p. 21) . 

USGS paleobotany Joe. D3725-C, slide (5) at 96.0 X 18.2. 

20, 21. Aesculiidites dubius (Jardine and Magloire) n. comb. ( p. 21) . 

Two focal levels. USGS paleobotany Joe. D3726- B, slide (1) at 

105.4 X 15.7. 

22-25. Grewipollenites radiatus n. sp. ( p. 21). 

22. Holotype. USGS paleobotany Joe. D3726- D, slide (2) at 

104.7 X 18.8. 

23. Paratype. USGS paleobotany Joe. D3726-D, slide (2) at 86.0 

X 15.3. 

24. Paratype, USGS paleobotany Joe. D3726-D, slide (2) at 88.1 

X 4.1. 

25. USGS paleobotany Joe. D3726-D, slide (2) at 80.6 X 14.7. 

26-28. Pulcheripollenites krempii Srivastava (p. 22). 

26. USGS paleobotany Joe. D3725-C, slide (2) at 86.5 X 3.8. 

27. USGS paleobotany Joe. D3725- C, slide (4) at 85.9 X 10.6. 

28. USGS paleobotany Joe. D3725-C, slide (2) at 96.5 X 18.9. 

29-31. Tricolpites reticulatns Cookson ( p. 23) . 

29. USGS paleobotany Joe. D3725-D, slide (2) at 93.4 X 5.2. 

30. USGS paleobotany Joe. D3726-E, slide (6} at 104.6 X 12.6. 

31. USGS paleobotany Joe. D3726-E, slide (6) at 90.6 X 10.7. 

32-34. Tricolpites hian.s Stanley (p. 24). 

32. USGS paleobotany Joe. D3726-E, slide (3) at 104.8 X 18.1. 

33. USGS paleobotany Joe. D3726- A, slide (2) at 85.8 X 8.0. 

34. USGS paleobotany Joe. D3726-E, slide (3) at 110.0 X 7.4. 

35-37. Tricolpites spp, (p. 24). 

35. USGS paleobotany Joe. D3725-C, slide (4) at 80.4 X 2.5. 

36. USGS paleobotany loc. D4069, slide (1) at 108.0 X 5.5. 

37. USGS paleobotany loc. D4070, slide (2) at 100.1 X 3.5. 
GEOLOGI CAL SURVEY PROFESSIONAL PAPER 770 PLATE 8 

20 

14 

31 33 36 

VITREISPORITES, TAXODIACEAEPOLLENITES, SEQUOIAPOLLENITES, CLASSOPOLLIS, 

EQUISETOSPORITES, ARECIPITES, LILIACJDITES, OVOIDITES, AESCULIIDITES, 

GREWIPOLLENITES, PULCHERIPOLLENITES, AND TRICOLPITES 

37 
FIGURES 

PLATE 9 

[All figures X 1,000] 

1-6. Tri~olpopollenites levitas n. sp. ( p. 25). 

1. Ho!otype. USGS paleobotany Joe. D3725-C, slide (2) at 113.2 

X 3.8. 

2, 3. Paratype, two focal levels. USGS paleobotany Joe. D3725-C, 

slide ( 4) at 80.9 X 6.8. 

4. Paratype. USGS paleobotany Joe. D3725-C, slide (4) at 84.4 

X 11.3. 

5. USGS paleobotany Joe. D3725-C, slide (4) at 83.8 X 11.2. 

6. USGS paleobotany Joe. D3725-C, S'!ide (3) at 96.1 X 5.4. 

7, 8. Tricolpopollenites tessellatus n. sp. (p. 25). 

7. USGS paleobotany Joe. D4067, slide ( 1) at 103.4 X 16.4. 

8. Holotype. USGS paleobotany Joe. D3725-E, slide (2) at 107.6 

X 8.9. 

9. Cranwellia rumseyensis Srivastava (p. 26). 

USGS paleobotany loc. D3725- C, slide (4) at 83.1 X 14.0. 

10-12. Cranwellia bacata Srivastava (p. 26). 

10, 11. Two focal levels. USGS paleobotany Joe. D4067, slide (1) 

at 77.2 X 9.7. 

12. USGS paleobotany loc. D4067, slide (3) at 87.8 X 16.1. 

13. Cf. Cranwellia (p. 26). 

USGS paleobotany Joe. D3725- A, slide (2) at 100.7 X 2.5. 

14-16. Fibulapollis scabratus Tschudy (p. 26). 

14, 15. Two focal levels. USGS paleobotany loc. D3726--E, slide 

(4) at 85.7 X 14.3. 

16. USGS paleobotany loc. D3726- D, slide (2) at 87.3 X 6.2. 

17, 18. Aquilapollenites quadrilobus Rouse (p. 27) . 

17. USGS paleobotany loc. D3725- C, slide (4) at 110.3 X 3.7. 

18. USGS paleobotany loc. D4067, slide (12) at 88.1 X 8.0. 

19. Aquilapollenites senonicus (Mtchedlishvili) Tschudy and Leopold 

(p. 27). 

USGS paleobotany loc. D3725- C, slide (2) at 97.8 X 5.3. 

20. Aquilapollenites trialatus Rouse var. uniformis Tschudy and Leopold 

(p. 27). 

USGS paleobotany Joe. D3725-C, slide (4) at 90.4 X 8.7. 

21. Aquilapollenites clarireticulatus (Samoilovitch) Tschudy (p. 27) . 

USGS paleobotany loc. D4069, slide (3) at 112.5 X 8.5. 

22-24. Aquilapollenites trialatus Rouse (p. 27). 

22. USGS paleobotany Joe. D3726-E, slide (6) at 82.3 X 12.3. 

23. USGS paleobotany loc. D3726--B, slide (1) at 86.5 X 14.9. 

24. USGS paleobotany loc. D3726--E, slide (4) at 96.2 X 7.2. 

25. Aquilapollenites reticulatus (Mtchedlishvili) Tschudy and Leopold 

(p. 27) . 

USGS paleobotany loc. D3726--D, slide (2) at 104.6 X 12.1. 

26. Aquilapollenites sp. (p. 27). 

USGS paleobotany Joe. D3725-C, slide (5) at 83.5 X 20.2. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE 9 

2 

\I 

25 23 24 

TRICOLPOPOLLENITES, CRANWELLIA, FIBULAPOLLIS, AND AQUILAPOLLENITES 
PLATE 10 

[All figures X 1,000] 

FIGURES 1, 2. Aquilapollenites turbidus Tschudy and Leopold (p. 27). 

1. USGS paleobotany loc. D3726--E, slide (6) at 108.6 X 19.7. 

2. USGS paleobotany loc. D3726- C, slide (2) at 107.5 X 10.8. 

3-6. Aquilapollenites attenuattus Funkhouser (p. 27) . 

3. USGS paleobotany loc. D3725-C, slide (4) at 81.8 X 15.4. 

4. USGS paleobotany loc. D3725-C, slide (2) at 107.4 X 18.5. 

5. USGS paleobotany loc. D3725-C, slide (4) at 86.0 X 17.7. 

6. USGS paleobotany loc. D4067, slide (3) at 77.1 X 10.5. 

7-9. Mancic.orpus calvus (Tschudy and Leopold) n. comb. (p. 29). 

7. USGS paleobotany loc. D3725- C, slide (4) at 79.2 X 17.8. 

8. USGS paleobotany loc. D3725-C, slide (4) at 85.7 X 22.3. 

9. USGS paleobotany loc. D3726--C, slide (5) at 109.5 X 10.1. 

10, 11. Mancicorpus tripodiformis (Tschudy and Leopold) n. comb. (p. 29). 

10. USGS paleobotany loc. D4069, slide (4) at 97.5 X 10.1. 

11. USGS paleobotany loc. D3725-A, slide (2) at 89.0 X 15.9. 

12-16. Sindorapollis granulatus n. sp. (p. 30). 

12. Holotype. USGS paleobotany loc. D4069, slide ( 1) at 106.5 X 

10.5. 

13. Paratype. USGS paleobotany loc. D4069, slide (2) at 104.3 X 

3.6. 

14. Paratype. USGS paleobotany loe. D4069, slide (1) at 105.7 

X 18.7. 

15. USGS paleobotany loc. D4069, slide ( 4) at 83.8 X 16.1. 

16. USGS paleobotany loc. D4069, slide (1) at 82.5 X 22.3. 

17. Cupanieidites sp. ( p. 30). 

USGS paleobotany loc. D3726--E, slide (4) at 87.6 X 16.4. 

18-21. TricolPQrites rhomboides Anderson (p. 30). 

18. USGS paleobotany loc. D3725 -C, slide (5) at 110.3 X 2.9. 

19. USGS paleobotany loc. D3725-C, slide (2) at 112.9 X 13.6. 

20. USGS paleobotany loe. D3725-C, slide (4) at 108.3 X 20.2. 

21. USGS paleobotany Joe. D3725- C, slide (4) at 99.9 X 6.0. 

22. Tric.olporites sp. (p. 31). 

USGS paleobotany Joe. D4070, slide (2) at 101.6 X 13.5. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE 10 

9 

10 

14 15 11 

18 19 20 21 22 

AQUILAPOLLENITES, MANCICORPUS, SINDORAPOLLIS, CUPANIEIDITES, AND TRICOLPORITES 
PLATE 11 

[All figures X 1,000] 

FIGURE 1. Montanapollis endannulatus Tschudy (p. 31). 

USGS paleobotany loc. D4067, slide (10) at 83.9 X 8.0. 

2, 3. Siberiapollis montanensis Tschudy ( p. 31). 

2. USGS paleobotany loc. D3726-E, slide (5) at 91.1 X 14.0. 

3. USGS paleobotany loc. D3726-E, slide (8) at 82.5 X 6.9. 

4, 5. Triatriopollenites costatus Norton (p. 31). 

4. USGS paleobotany loc. D3726-A, slide (1) at 91.8 X 11.3. 

5. USGS paleobotany loc. D3726-E, slide (4) at 95.3 X 16.0. 

6-8. Sub triporopollenites alpin us (Wolf) n. comb. ( p. 31) . 

6. USGS paleobotany loc. D3725-C, slide ( 4) at 102.8 X 6.2. 

7. USGS paleobotany Joe. D3725-C, slide (2) at 96.7 X 14.3. 

8. USGS paleobotany loc. D4067, slide (2) at 90.1 X 18.2. 

9-12. Proteacidites spp. (p. 32). 

9. USGS paleobotany loc. D4069, slide (2) at 113.5 X 7.1. 

10. USGS paleobotany loc. D4069, slide (1) at 86.3 X 4.1. 

11. USGS paleobotany loc. D4067, slide (1) at 82.2 X 4.7. 

12. USGS paleobotany loc. D4067, slide (3) at 104.0 X 1.7. 

13. Periporopollenites sp. (p. 32). 

USGS paleobotany Joe. D3726-C, slide (1) at 101.9 X 11.9. 

14. Erdtmanipollis pachysandroides Krutzsch (p. 32). 

USGS paleobotany Joe. D4067, slide (10) at 79.2 X 3.1. 

15-17. Dyadonapites reticulatus n. sp. (p. 32). 

15. Holotype. USGS paleobotany loc. D3725-C, slide (4) at 85.0 X 

6.0. 

16. Paratype. USGS paleobotany loc. D3725-C, slide (5) at 91.0 X 

17.1. 

17. USGS paleobotany loc. D3725- D, slide (1) at 91.0 X 14.8. 

18, 19. lnaperturotetradites scabratus n. sp. (p. 32). 

18. Holotype. USGS paleobotany loc. D3725-A, slide (2) at 96.3 X 4.1. 

19. USGS paleobotany loc. D3725- E, slide (2) at 79.4 X 14.4. 

20. Droseridites cf. D. spinulosus (p. 33). 

USGS paleobotany loc. D3726-E, slide (3) 88.5 X 17.2. 
GEOLOGICAL SURVEY PROFESSIONAL PAPER 770 PLATE II 

4 

9 10 

18 

f 

5 

16 

2 

11 

19 

6 7 

12 13 

14 

17 

20 

MONTANAPOLLIS, SIBERIAPOLLIS, TRIATRIOPOLLENITES, SUBTRIPOROPOLLENITES, 

PROTEACIDITES, PERIPOROPOLLENITES, ERDTMANIPOLLIS, DYADONAPJTES, 

INAPERTUROTETRADITES, AND DROSERIDJTES 

8