Normapolles Pollen from the 

Mississippi Embayment

GEOLOGICAL SURVEY PROFESSIONAL PAPER 865

Work done in cooperation with the 

Kentucky Geological Survey
Normapolles Pollen from the 

Mississippi Embayment

By ROBERT H. TSCHUDY

GEOLOGICAL SURVEY PROFESSIONAL PAPER 865

Work done in cooperation with the 

Kentucky Geological Survey

Taxonomy and stratigraphic ranges of 

Normapolles genera and species found 

in Mississippi embayment rocks

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1975
UNITED STATES DEPARTMENT OF THE INTERIOR

GEOLOGICAL SURVEY 

V. E. McKelvey, Director

Library of Congress Cataloging in Publication Data

Tschudy, Robert H.

Normapolles pollen from the Mississippi embayment.

(Geological Survey Professional Paper 865)

Bibliography: p.

Includes index.

Supt. of Docs. No.: 119.16:865

1. Normapolles. 2. Paleobotany-Cretaceous. 3. Paleobotany-Tertiary. 4. Paleobotany-Southern States.

I. Title. II. Series: United States Geological Survey Professional Paper 865.

QE993.2.T76 56l'.13 75-619037

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402 

Stock Number 024-001-02680-5
CONTENTS

Abstract ____________________

Introduction _________________

Previous reports, North American __ 

Taxonomic reports ________

Nontaxonomic references _____

Acknowledgments ___________

Methods ___________________

Fossil pollen sample localities _______

Type specimens _______________

Systematic descriptions ___________

Genus Basopollis ____________

Basopollis obscurocostatus, n. sp

Genus Choanopollenites ________

cf. Choanopollenites sp ______

Genus Complexiopollis _________

Genus Extremipollis __________

Extremipollis vivus, n. sp ____

Extremipollis versatilis, n. sp __ 

Extremipollis caminus, n. sp __

Genus Interpollis ____________

Interpollis cf. /. supplingensis __ 

Interpollis microsupplingensis __ 

Interpollis intranodus, n. sp ___

Genus Interporopollenites _______

__________ 9

-----____ 9

__________ 9

	10 

________ 10

__________ 11

__________ 11

	12 

_________ 12

__________ 12

Interporopollenites turgidus, n. sp __________ 13

Genus Kyandopollenites ___________________ 13

Kyandopollenites anneratus ______________ 13

Genus Megatriopollis ____________________ 13

Megatriopollisl sp ____________________ 14

Genus Minorpollis ______________________ 14

Minorpollis minimus __________________ 14

Minorpollis aff. M. minimus _____________ 14

Genus Nudopollis ______________________ 15

Nudopollis endangulatus ________________ 15

Nudopollis aff. N. endangulatus ___________ 15

Nudopollis terminalis __________________ 15

Nudopollis terminalis subsp. hastaformis ______ 15

Nudopollis terminalis subsp. hebeformis _______ 15

Nudopollis aff. N. terminalis _____________ 16

Nudopollis thiergartii ___________________ 16

Nudopollis thiergartii subsp. rneridium _______ 16

Nudopollis thiergartii subsp. conspicuus _______ 16

Nudopollis thiergartii subsp. minutus ________ 16

Nudopollis aff. N. thiergartii _____________ 16

Genus Pecakipollis ______________________ 16

cf. Pecakipollis sp ____________________ 16

Genus Plicapollis _______________________ 17

Plicapollis usitatus, n. sp _______________ 17

Plicapollis retusus, n. sp ________________ 18

Plicapollis vacuus, n. sp ________________ 18

Page

Systematic descriptions Continued

Genus Plicapollis Continued

Plicapollis rusticus, n. sp _______________ 18

Genus Pompeckjoidaepollenites ______________ 19

Pompeckjoidaepollenites aff. P. subhercynicus ___ 19

Genus Oculopollis ______________________ 20

Genus Pseudoculopollis ___________________ 20

Pseudoculopollis admirabilis, n. sp __________ 20

Pseudoculopollis sp. A _________________ 20

Genus Semioculopollis ____________________ 21

Semioculopollis sp. A __________________ 21

Genus Pseudoplicapollis ___________________ 21

Pseudoplicapollis endocuspis, n. sp __________ 22

Pseudoplicapollis serenus, n. sp ____________ 22

Pseudoplicapollis sp. A_________________ 22

Genus Pseudovacuopollis __________________ 23

Pseudovacuopollis involutus, n. sp __________ 23

Genus Thomsonipollis ____________________ 24

Thomsonipollis magnificus _______________ 24

Thomsonipollis magnificoides _____________ 24

Thomsonipollis aff. T. magnificoides _________ 24

Thomsonipollis expolitus, n. sp ____________ 25

Genus Trudopollis ______________________ 25

Trudopollis variabilis, n. sp _____________ 25

Trudopollis plenus, n. sp ________________ 26

Trudopollis cf. T. artifex ________________ 26

Trudopollis sp. A ____________________ 27

Trudopollis sp. B ____________________ 27

Trudopollis sp. C ____________________ 28

Genus Vacuopollis ______________________ 28

Vacuopollis munitus, n. sp _______________ 28

Genus Osculapollis, n. gen _________________ 29

Osculapollis aequalis, n. sp ______________ 29

Osculapollis perspectus, n. sp _____________ 29

Osculapollis sp. A ____________________ 30

Genus Praecursipollis, n. gen ________________ 30

Praecursipollis plebius, n. sp _____________ 30

Genus Pseudatlantopollis, n. gen ______________ 30

Pseudatlantopollis simulatus, n. sp __________ 31

Genus Endoinfundibulapollis, n. gen ___________ 31

Endoinfundibulapollis distinctus, n. sp _______ 31

New genus A _________________________ 32

New genus B _________________________ 32

New genus C _________________________ 32

New genus D _________________________ 33

Key to Mississippi embayment Normapolles genera _____ 33

Stratigraphic distribution of Normapolles genera _______ 35

Stratigraphic distribution of Normapolles species ______ 37

References _____________________________ 39

Index ______ ____ ____ _____ ____ ___ 41

III
IV CONTENTS

ILLUSTRATIONS

[Plates follow index]

PLATE 1. Basopollis obscurocostatus and cf. Choanopollenites sp.

2. Extremipollis vivus and E. versatilis.

3. Extremipollis caminus.

4. Interpollis cf. /. supplingensis and /. microsupplingensis.

5. Interpollis intranodus.

6. Interporopollenites turgidus.

7. Kyandopollenites anneratus and Megatriopollisf sp.

8. Minorpollis aff. M. minimus, Nudopollis aff. AT. endangulatus, N. aff. N. thiergartii, and AT. aff. AT. terminalis.

9. cf. Pecakipollis sp., Plicapollis usitatus, and P. retusus.

10. Plicapollis vacuus and P. rusticus.

11. Pompeckjoidaepollenites aff. P. subhercynicus and Pseudoculopollis admirabilis.

12. Pseudoculopollis sp. A. and Semioculopollis sp. A.

13. Pseudoplicapollis endocuspis, P. serenus, and P. sp. A.

14. Pseudovacuopollis involutus and Thomsonipollis magnificus.

15. Thomsonipollis magnificoides and T. aff. T. magnifieaides.

16. Thomsonipollis expolitus and Trudopollis uariabilis.

17. Trudopollis plenus, T. cf. T. arti/e*, and T. sp. A, B, and C.

18. Vacuopollis munitus, Osculapollis aequalis, 0. perspectus, and 0. sp. A.

19. Praecursipollis plebius and Pseudatlantopollis simulatus.

20. Endoinfundibulapollis distinctus and New Genera A, B, C, and D.

FIGURES 1-42. Sketches showing: Page

1. Basopollis obscurocostatus -------------------------------------------------- 8

2. cf. Choanopollenites sp _________________________--------------------------- 8

3. Extremipollis vivus ------------------------------------------------------ 9

4. Extremipollis versatilis ---------------------------------------------------- 10

5. Extremipollis caminus ---------------------------------------------------- 10

6. Interpollis cf. /. supplingensis ------------------------------------------------ 11

7. Interpollis intranodus ----------------------------------------------------- 12

8. Interporopollenites turgidus ------------------------------------------------- 13

9. Kyandopollenites anneratus ------------------------------------------------- 14

10. Megatriopollis1? sp. _________________________---__------------------------ 14

11. Minorpollis aff. M. minimus ------------------------------------------------- 15

12. cf. Pecakipollis sp. ____________________________---_---------------------- 17

13. Plicapollis usitatus ------------------------------------------------------ 17

14. Plicapollis retusus ------------------------------------------------------- 18

15. Plicapollis vacuus ------------------------------------------------------- 18

16. Plicapollis rusticus ------------------------------------------------------ 19

17. Pseudoculopollis admirabilis ------------------------------------------------- 20

18. Pseudoculopollis sp. A ___________________--------------------------------- 21

19. Semioculopollis sp. A _____________________________------------------------ 21

20. Pseudoplicapollis endocuspis ------------------------------------------------- 22

21. Pseudoplicapollis serenus ___________________________________________________ 22

22. Pseudoplicapollis sp. A ____________________________________________________ 23

23. Pseudovacuopollis involutus _________________________________________________ 23

24. Thomsonipollis aff. T. magnificoides ____________________________________________ 24

25. Thomsonipollis expolitus ___________________________________________________ 25

26. Trudopollis variabilis _____________________________________________________ 26

27. Trudopollis plenus _______________________________________________________ 26

28. Trudopollis cf. T. arti/ex ______________________________________________ 27

29. Trudopollis sp. A. ______________________________________________ 27

30. Trudopollis sp. B. _______________________________________________________ 27

31. Trudopollis sp. C. _______________________________________________________ 28

32. Vacuopollis munitus ______________________________________________________ 28

33. Osculapollis aequalis _____________________________________________________ 29

34. Osculapollis perspectus ____________________________________________________ 29

35. Osculapollis sp. A. ______________________________________________________ 30

36. Praecursipollis plebius ____________________________________________________ 30

37. Pseudatlantopollis simulatus _________________________________________________ 31

38. Endoinfundibulapollis distinctus ______________________________________________ 31
CONTENTS v

FIGURES 1-42. Sketches Continued Page

39. New Genus A ______________________________________________________ 32

40. New Genus B ______________________________________________________ 32

41. New Genus C ______________________________________________________ 33

42. New Genus D ______________________________________________________ 33

43. Chart showing comparison of stratigraphic distribution of Normapolles genera in Europe and Mississippi

embayment ______________________________________________________ ___ 36

44. Chart showing stratigraphic ranges of Normapolles species in Mississippi embayment __________________ 38
NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

By ROBERT H. TSCHUDY

ABSTRACT

The Normapolles group of fossil pollen genera is extinct. Pollen types 

of the included genera are triaperturate, morphologically complex, and 

distinctly different from types produced by extant pollen genera. 

Genera included in this somewhat heterogeneous group were first 

described from the Late Cretaceous and early Tertiary of Europe. In recent years increasing mention has been made of Normapolles genera 

observed in pollen preparations from North American rocks, and 

several new genera, not known from Europe, have been described. Taxonomic evaluations of North American Normapolles genera and 

species, however, have been minimal. This paper lists and evaluates 

the Normapolles genera and species found in rocks from the Mississippi embayment province.

At the present time, 57 Normapolles genera have been reported from 

Europe, 47 from boreal Middle Europe and 10 from western Hungary. 

Only four Normapolles genera are common to the two European 

regions. In Mississippi embayment rocks I have found pollen 

assignable to 18 of the European Normapolles genera, to 2 additional 

genera previously described from the United States, and to 4 new 

genera that I have proposed. I have found and illustrated specimens of 

four additional potentially new genera, but have not named them 

because of insufficient numbers of specimens for diagnosis.

The development of Normapolles genera in the Mississippi embayment began with their first appearance in the Cenomanian. They increased in numbers, reaching a peak in the Campanian Maestrichtian interval, then declined in numbers during the early Tertiary and 

finally disappeared at the end of middle Eocene time. This development approximately parallels the generic occurrence of the Normapolles group in Europe.

Some of the genera common to Europe and the Mississippi embayment show remarkably similar stratigraphic ranges, but the ranges of 

others are distinctly different in the two regions. Several genera apparently appear earlier in Mississippi embayment rocks than in European rocks. This phenomenon is at odds with the commonly held 

hypothesis that Europe is the center of development and dispersion of 

the Normapolles group as a whole. If a migration route from Europe to 

North America allowed some Normapolles genera to move from Europe 

to North America, an alternate hypothesis would allow other Normapolles genera originating independently in North America to 

migrate to Europe and, thus, appear there later in the stratigraphic 

record.

A total of 59 species assignable to Normapolles genera have been 

recognized from the Mississippi embayment. Of this number, 43 have 

been found in sufficient numbers that specific names could be 

assigned. Thirty-three of these species are confined to North America. 

This fact suggests that the North American genera were relatively 

isolated from their European counterparts.

Normapolles species by themselves, or in combination with pollen 

assemblages, have proved to be excellent indicators of the several 

segments of the Late Cretaceous-early Tertiary part of the 

stratigraphic column. The known range zone of each Normapolles 

species discussed is presented in a stratigraphic chart.

INTRODUCTION

The term "Normapolles" was proposed by Pflug 

(1953) to embrace a group of pollen form genera that 

possessed morphological characteristics distinctly 

different from the morphological characteristics of most 

modern pollen types. Pollen grains assigned to genera of 

the Normapolles group are commonly triangular and 

shallow lens-shaped (short axial), with apertures at the 

angles, although a few are spherical or nearly so. The 

apertures are complex and morphologically highly 

differentiated, sometimes to a degree that they have 

been referred to as having a "bizarre" construction. The 

morphology of the apertures and the associated wall 

structure, although vastly different within the group, is 

sufficiently distinctive that none of the included genera 

can be conveniently included in the porate, colpate, or 

colporate morphotypes. Plants that produced pollen 

assigned to these genera are, so far as is known, extinct, 

and their relationships to modern genera, and indeed 

families, are still in the realm of conjecture. The Normapolles group is a group of convenience and neither 

genetic nor morphologic relationship among all included 

genera is implied.

Pollen genera assignable to the Normapolles group 

had their origin during the Cenomanian and became virtually extinct at the end of the Eocene. Their postulated 

center of origin and dispersal was in central Europe 

(Goczan and others, 1967, p. 435). In North America this 

group of genera is virtually limited to the Mississippi 

embayment and the eastern coast, although specimens 

assignable to Normapolles genera have been reported 

from rocks deposited to the west of the Late Cretaceous 

North American midcontinental epeiric sea. In North 

America also, pollen of Normapolles genera has been 

found only in rocks of Late Cretaceous and early Tertiary 

age.

Krutzsch (1959) enlarged the normapolles group by 

the addition of several new form genera from the European Cretaceous and early Tertiary. Goczan, Groot, 

Krutzsch, and Pacltova (1967), in the light of new 

knowledge of the fine morphology of these pollen grain 

types, in an excellent monographic work, reevaluated all 

the then-known Normapolles genera, erected several 

new genera and included more complete and accurate
NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

diagnoses and diagrams, thus contributing to the 

clarification of many morphological details that serve to 

characterize the several genera. It is this European work 

by Goczan and his coworkers that provides the basis for 

an evaluation and listing of Normapolles genera and 

species from the Mississippi embayment area of the 

United States.

PREVIOUS REPORTS, NORTH AMERICAN

Previous reports of Normapolles genera from North 

America are largely confined to the eastern part of the 

United States; but four, the reports of Newman (1965), 

Drugg (1967), B. D. Tschudy (1971), and Chmura (1973), 

record Normapolles genera from the western part. 

Although new information has altered somewhat the circumscriptions of some genera, other genera are no longer 

valid, and some determinations were made on the basis 

of few specimens or poorly preserved material, the reader 

should note that the several authors at the times of 

publication assigned specimens to what they then consedered to be appropriate Normapolles genera and 

species.

These reports can be divided into two categories: (1) 

reports that include taxonomic diagnoses or descriptions, and (2) reports that mention Normapolles genera 

but do not provide descriptions or taxonomic details.

TAXONOMIC REPORTS

The first mention of a Normapolles genus from the 

United States appeared in a publication by Krutzsch 

(1960a). In that report Krutzsch established the genus 

Thomsonipollis, basing his diagnosis upon specimens 

from the early Eocene of Europe and from lower Eocene 

rocks near Austin, Tex. Two other reports proposed new 

Normapolles genera from the United States. Stover (in 

Stover and others, 1966) erected the new monotypic 

genera Choanopollenites and Kyandopollenites from 

specimens found in the early Eocene of Texas. These 

genera, though not formally included in the Normapolles 

group, undoubtedly belong there because of their complex morphological characteristics. B. D. Tschudy (1971) 

found and described two new Normapolles genera from 

upper Campanian rocks of Montana. These new genera 

are Siberiapollis and Montanapollis.

The first North American descriptions of species 

belonging to heretofore strictly European Normapolles 

genera are found in a publication by Groot, Penny, and 

Groot (1961). They described one species each of 

Turonipollis (now Complexiopollis), Sporopollis (now 

Pseudoplicapollis) , Latipollis (now Complexiopollis), 

Quedlinburgipollis and Trudopollis, two species of 

Vacuopollis and three species of Plicapollis from Upper 

Cretaceous rocks of the Eastern United States, thus 

firmly establishing the presence of Normapolles genera 

other than Thomsonipollis on the North American continent. Groot and Groot (1962) mentioned or described one

species each of the genera Latipollis (now Complexiopollis), Plicapollis, and Extratriporopollenites from 

the Brightseat Formation (Paleocene) of Maryland; and 

Fredericksen (1973) described a species of Plicapollis 

from the Tertiary of Mississippi. From the Cretaceous of 

Colorado, Newman (1965) described one species each of 

Conclauipollis (now Vacuopollis) and Trudopollis and 

mentioned a species of Sporopollis (now 

Pseudoplicapollis). Chmura (1973) described one species 

of Trudopollis and mentioned one species of Plicapollis 

from the Cretaceous of California. The reports by Newman and Chmura are the only ones describing European 

Normapolles genera and species from pollen specimens 

obtained from Western United States. Two reports on 

Late Cretaceous palynomorphs from Delaware and New 

Jersey appeared in 1966. Gray and Groot (1966) provided 

descriptions of two species of Plicapollis and two species 

of Trudopollis, and Kimyai (1966) described a single 

species of Conclauipollis (now Vacuopollis). Elsik (1968), 

in discussing the palynomorph flora of the Paleocene of 

Texas, emended the genus Thomsonipollis and listed 

one species each of Trudopollis, Plicapollis, Extratriporopollenites, Choanopollenites and two species of 

Nudopollis. More recently, R. H. Tschudy (1973b), in 

tracing the evolutionary development and stratigraphic 

distribution of the Complexiopollis lineage, described 

four new species of Complexiopollis and five new species 

of Choanopollenites from Cretaceous and lower Tertiary 

rocks of the Mississippi embayment region.

NONTAXONOMIC REFERENCES

Mention of the Genera Extratriporopollenites and 

Trudopollis in Paleocene and lower Eocene rocks of 

Arkansas was made by Jones (1962), and in the same 

year Brown and Pierce (1962) figured specimens from the 

Turonian of Texas that probably belong to genera of the 

Normapolles group under the name of Punctatricolporites. Leopold and Pakiser (1964) listed the 

genera Basopollis, Sporopollis (now Pseudoplicapollis), 

Complexiopollis, Latipollis (now Complexiopollis), 

Nudopollis, Minorpollis and Conclauipollis (now 

Vacuopollis) from the Late Cretaceous of western 

Alabama. Engelhardt (1964) listed and illustrated two 

species of the genus Extratriporopollenites from the 

Eocene Cockfield Formation of Mississippi. Drugg 

(1967) mentioned single species of the genera Nudopollis 

and Vacuopollis from the Late Cretaceous-early Tertiary transition of California. Doyle (1969) and Wolfe 

and Pakiser (1971) examined Upper Cretaceous rocks 

from the Atlantic Coastal Plain area and both recorded 

the genera Atlantopollis, Complexiopollis, 

Praebasopollis, Pecakipollis, Trudopollis, Plicapollis 

and Pseudoplicapollis. Doyle recorded Vacuopollis in addition, and Wolfe and Pakiser recorded in addition 

Semioculopollis, Heidelbergipollis, Minorpollis, 

Primipollis, Neotriangulipollis, and 14 new unnamed
FOSSIL POLLEN SAMPLE LOCALITIES

genera. They made no attempt to record the 

morphological characteristics of any of these genera. 

Fairchild and Elsik (1969), in discussing the 

characteristic palynomorphs of the Gulf lower Tertiary, 

listed two species of Nudopollis and one species each of 

Thomsonipollis and Choanopollenites. R. H. Tschudy 

(1973a) recorded one species of Kyandopollenites, two 

species of Nudopollis, and the genus Thomsonipollis 

from the Eocene of the Mississippi embayment. Goczan, 

Groot, Krutzsch, Pacltova (1967, p. 511) reported that 

types similar to Thomsonipollis magnificoides are known 

from "the Laramie of Raton Pass, Alabama/U.S.A." 

(Undoubtedly the Upper Cretaceous Laramie Formation 

from Raton Pass, Colo.).

ACKNOWLEDGMENTS

I thank Sharon Van Loenen and Kathryn Dieterich for 

their assistance in the preparation of the samples and 

Kathryn Dieterich for photography of the specimens.

METHODS

The methods employed for processing samples and 

preparing slides are standard for this laboratory and 

have been reported previously (Tschudy, R. H., 1970). A 

minor modification in procedures is the employment of 

the synthetic resin "Histoclad" as a mounting medium 

for some of the slides, replacing the previously used 

medium "Permount."

FOSSIL POLLEN SAMPLE LOCALITIES

The samples from Cretaceous and Tertiary rocks used 

in this study are from the numbered USGS Paleobotany 

localities listed below. The locations and detailed 

descriptions of the Cretaceous and Paleocene localities 

have been presented previously (Tschudy, R. H., 1973b). 

The localities and detailed information from Eocene 

samples have also been given in a previous publication 

(Tschudy, R. H., 1973a). This information is repeated 

here because of the desirability of having accurate 

locality data available in the same publication that 

pioposes new taxa.

usr-s

Paleobot mv 

loc.

Stratigraphic 

unit

D3415 _ _ _ Outcrop at an altitude of 510 

ft on the south side of U.S. 

Highway 78, NW 1 4 sec. 34, 

T. 3 S., R. 15 W., Colbert 

Courty, Ala. Collected by 

W. \V. Olive, U.S. 

Geological Survey, 

"probably from unit 4 

described by Stephenson 

and Monroe, 1940, p. 46" 

(W. W. Olive, written 

commun., 1964).

D3265 _ _ _ E. W. Berry field designation 

6474. Collected at Broken

Tuscaloosa Formation, 

Cenomanian.

Eutaw Formation, 

Coniacian.

USGS

Paleobotany 

loc.

Locality

Stratigraphic 

unit

Arrow Bend of the Chattahoochee River about 13 

miles below Columbus, 

Chattahoochee County, 

Ga. Matrix material from 

leaf locality collected by E. 

W. Berry, U.S. Geological 

Survey, 1909. Matrix submitted by Jack Wolfe, 

U.S. Geological Survey, 

1963.

D3260 _ _ _ E. W. Berry field designation 

6471. Collected at the 

junction of Cowickee 

Creek and Chattahoochee 

River, Barbour County, 

Ala. Matrix material from 

leaf locality collected by E. 

W. Berry, U.S. Geological 

Survey, 1909. Matrix submitted by Jack Wolfe, 

U.S. Geological Survey,

1963.

D3412___ Type locality, near middle. 

Elevation 440 ft along bluff 

on left bank Tennessee 

River 0.3 mile downstream 

from river mile 185 and 0.9 

mile east of crossroads 

north of Coffee Landing, 

Hardin County, Tenn. 

Collected by W. W. Olive, 

U.S. Geological Survey,

1964.

D3413 _ _ _ From roadcut along east side 

Tennessee Route 22, 0.8 

mile north of intersection 

with Tennessee Route 100, 

Henderson County, Tenn. 

Collected by W. W. Olive, 

U.S. Geological Survey, 

1964.

D1967 ___ Type locality of McNairy 

Sand Member. West end 

of big cut in Southern RR. 

east of Dismal Swamp and 

Tuscumbia River. Base is 

approximately same elevation as track and is 40 ft 

northwest of railroad 

crossing. South center 

NW'-iSW^, Chewala 

quadrangle, McNairy 

County, Tenn.

D1967-A_ _ 0-14 in. above bottom of sequence. 

B_ _ 14-26 in. above bottom of

sequence. 

C_ _ 26-38 in. above bottom of

sequence.

Collected by E. M. Gushing, 

U.S. Geological Survey, 

1962.

Cusseta Sand Member, 

Ripley Formation, 

Campanian.

Coffee Sand, Campanian.

Coon Creek Tongue of 

Ripley Formation, 

Maestrichtian.

McNairy Sand Member of Ripley Formation Maestrichtian.

Do. 

Do. 

Do.
NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

USiJS

 'aleobotany

loc.

Locality

Stratigraphic 

unit

D3416 _ _ _ 6 ft below unconformable 

contact with Owl Creek 

Formation; NW^NWW 

sec. 10, T. 27 N., R. 11 E., 

Advance 7 l/i -minute 

quadrangle, Stoddard 

County, Mo. Collected by 

W. W. Olive, U.S. 

Geological Survey, 1964.

D3410 _ _ _ Type locality. From right 

bank of Owl Creek, 2.5 

miles northeast of Ripley, 

Miss.; E''2 sec. 7, T. 4 S., 

R. 4 E., Tippah County, 

Miss. Collected by W. W. 

Olive, U.S. Geological 

Survey, 1964.

D1966 ___ 1.5 miles south-southwest of 

Cotton Plant, Miss., 

southwest wall of tributary 

to Guyton Creek along 

country road. SVzNW 1^ 

sec. 8, T. 6S.,R. 3E., Tippah County, Miss.

D1966-A_ _ 6 ft above basal limestone. 

B_ _ 9 ft above basal limestone. 

C_ _ 14 ft above basal limestone. 

D_ _ 15 ft above basal limestone. 

E_ _ 17 ft above basal limestone. 

F__20-23 ft above basal

limestone.

Collected by E. M. Gushing, 

U.S. Geological Survey, 

1962.

D3548 _ _ _ Auger hole J. 12, about 1 mile 

from southwest corner 

Joppa 7 ''2 -minute quadrangle; Kentucky coordinates, South zone: E. 

1,094,590 ft; N. 308,200 ft, 

McCracken County, Ky. 

[Depths are from top of 

auger hole]

D3548-A_ _ Depth 93-98 ft.

B_ _ Depth 77-93 ft.

C_ _ Depth 62-77 ft.

D_ _ Depth 52-62 ft.

Collected by W. W. Olive, 

U.S. Geological Survey, 

1965.

D3285 ___ Channel of McCullough 

Fork, 300 ft N. KentuckyTennessee State line. Kentucky coordinates, South 

zone: E. 1,263,500 ft; N. 

70,250 ft, Hazel quadrangle, Galloway County, 

Ky. Collected by W. W. 

Olive, U.S. Geological 

Survey, 1963.

Upper part of McNairy 

Sand, Maestrichtian.

Owl Creek Formation, 

Maestrichtian.

Clayton Formation, 

Paleocene.

Do. 

Do. 

Do. 

Do. 

Do. 

Do.

Clayton Formation,

Paleocene.

Porters Creek Clay, 

Paleocene. 

Do. 

Do.

Do.

USGS

Paleobotany 

loc.

Locality

Stratigraphic 

unit

D1849-

D1850___ NW'/iSW1 '* sec. 6, T. 11 N., 

R. 11 W., De Soto Parish, 

La. Sample provided by 

W. W. Fairchild, Standard 

Oil Company of Texas, 

1962.

D3138 _ _ _ Upper half Naheola Formation from along Alabama 

Highway 17 in SE^NW'/i 

sec. 3, T. 15 N., R. 2 W., 

Choctaw County, Ala. 

Sample provided by C. W. 

Copeland, Geological 

Survey of Alabama, 1963.

D3268 _ _ _ Type locality. From Coal 

Bluff on Alabama River, 

sec. 7, T. 11 N., R. 7 E., 

Wilcox County, Ala. Sample provided by H. T. 

Ames, Pennsylvania State 

Univ., 1963.

D3222 _ _ _ 0.2 mile east of bridge across 

Tucabum River on 

Lavaca-Pennington road 

on south side of new roadcut, sec. 22, T. 14 N., R. 1 

W., Choctaw County, Ala. 

(See MacNeil, geologic 

map, 1946.) Collected by 

W. W. Olive and R. H. 

Tschudy, U.S. Geological 

Survey, 1963.

D1854 _ _ _ About 1 mile northeast of 

Ackerman, Miss., in 

railroad cut, Illinois Central Railway, at Blantons 

Gap, sees. 28 and 21, T. 17 

N., R. 11 E., Choctaw 

County, Miss. (See Vestal, 

1943, p. 30.) Collected by 

W. W. Olive, U.S. 

Geological Survey, 1962.

D1862 _ _ _ From south wall of the valley 

of Potts Creek, about 75 

yards east of road bridge, 

NW'/4 sec. 4, T. 6 S., R. 1 

W., Marshall County, 

Miss. (See Vestal, 1954, p. 

41.) Collected by W. W. 

Olive, U.S. Geological 

Survey, 1962.

D1846___ Sample F48, NW^SE 1 * sec. 

12, T. 6 N. R. 13 W., 

Sabine Parish, La., elev 

165 ft. (See Andersen, 

1960, p. 74, 75, fig. 13.) 

Supplied through the 

courtesy of W. W. 

Fairchild, Standard Oil 

Co. of Texas, 1962.

Naborton Formation, 

Paleocene.

Naheola Formation, 

Paleocene.

Coal Bluff Marl 

Member of Naheola 

Formation, Paleocene

Nanafalia Formation. 

Wilcox Group, lower 

Eocene.

Wilcox Formation 

(formerly Ackerman 

Formation, type 

locality), lower 

Eocene.

Wilcox Formation 

(formerly Ackerman 

Formation), lower 

Eocene.

Bayou Lenann Member of Pendleton Formation (type 

locality) (Andersen, 

1960). In Wilcox 

Group, lower Eocene.
FOSSIL POLLEN SAMPLE LOCALITIES

uses

Paleobotanv Locality

Stratigraphic 

unit

D1851 _ _ _ Sample F-129 from north 

side of Louisiana Highway 

6, sec. 6, T. 6N.,R. 11 W., 

Sabine Parish, La. Ostrea 

multilirata zone. (See 

Andersen, 1960, p. 76.) 

Supplied through the 

courtesy of W. W. 

Fairchild, Standard Oil 

Co. of Texas, 1962.

D1855 ___ North side of roadcut 1.5 

miles west of corporate 

limit of Meridian, Miss., 

on U.S. Highway 80, sees. 

15, 22, T. 6 N., R. 15 E., 

Lauderdale County, Miss. 

(See Foster, 1940, p. 67.) 

Collected by W. W. Olive, 

U.S. Geological Survey, 

1962.

D1848 _ _ _ Sample F-71, from bed of 

Two Mile Creek, 1.5 miles 

due south of U.S. Highway 

79 bridge across Brazos 

River, Milam County, 

Tex. (Stop 2 of Houston 

Geol. Soc. Ann. Field Trip, 

1959. See Smith, 1959.) 

Supplied through the 

courtesy of W. W. 

Fairchild, Standard Oil 

Co. of Texas, 1962.

D1858 _ _ _ Holly Springs Brick and Tile 

Co., Holly Springs, Miss., 

pit south of Mississippi 

Highway 78, Poorhouse or 

No. 2 pit, SE 1 ^ sec. 26, T. 

3 S., R. 3 W., Marshall 

County, Miss. (See Vestal, 

1954, p. 67-90.) Collected 

by W. W. Olive, U.S. 

Geological Survey, 1962.

D1856 _ _ _ 2.9 miles east of west city 

limits of Hickory, Miss., 

on U.S. Highway 80, NE''i 

sec. 32, T. 6 N., R. 13 E., 

Newton County, Miss. 

(See Priddy, 1961, p. 37.) 

Collected by W. W. Olive, 

U.S. Geological Survey, 

1962.

D1852 ___ Sample F-130 from 

northeast side of Louisiana 

Highway 476, SE 1 * sec. 

10, T. 5N.,R. 12 W., south 

of Negreet, Sabine Parish, 

La. (See Andersen, 1960, 

p. 87 and pi. 13.) Supplied 

through the courtesy of W. 

W. Fairchild, Standard Oil 

Co. of Texas, 1962.

Pendleton Formation 

(Andersen, 1960). In 

Wilcox Group, lower 

Eocene.

Uppermost part, 

Hatchetigbee Formation. In Wilcox 

Group, lower Eocene.

Marquez Shale Member (of Stenzel, 1938) 

of Reklaw Formation. In Claiborne 

Group, middle Eocene.

Tallahatta Formation. 

In Claiborne Group, 

middle Eocene.

Zilpha Clay. In Claiborne Group, middle 

Eocene.

Sparta Sand. In 

Claiborne Group, 

middle Eocene.

l tSGS

Paleobotany 

loc.

Locality

Stratigraphic 

unit

D3421 ___ NW^SW'iSW'M sec. 23, T. 

16 S., R. 14 W., near south 

end of Gallon Lake, Union 

County, Ark. Collected by 

H. N. Halberg, U.S. 

Geological Survey, 1964.

D4524 _ _ _ Just above iron bridge over 

Techeva Creek, SE^ 

NE'/4 sec. 16, T. 13 N., R. 

1 W., Yazoo County, Miss. 

(Yegua Member of Lisbon 

Formation as used by 

Mellen, 1940, p. 14 and fig. 

1.) Collected by E. H. 

Boswell, U.S. Geological 

Survey, 1970.

D4525___ NE 1/4SW 1/4SW 1/4 sec. 11, T. 

5 N., R. 11 E., Newton 

County, Miss, (stratigraphically about 

equivalent to D4526. E. H. 

Boswell, written commun., 

1970). Collected by E. H. 

Boswell, U.S. Geological 

Survey, 1970.

D4526___ SE^SW^ sec. 32, T. 4 N., 

R. 13 E., !/2 mile southeast 

of Rose Hill, Jasper County, Miss. (See DeVries, 

1963, p. 22, fig. 6.) 

Collected by E. H. 

Boswell, U.S. Geological 

Survey, 1970.

D3224___ South bluff of Garland 

Creek, 2.6 miles east of 

steel bridge over 

Chickasawhay River, 2 

miles from house at road 

fork straight east along log 

trail. 6 ft from top of formation, NWV4 sec. 28, T. 1 

N., R. 16 E., Clarke County, Miss. (See Holden and 

others, 1948, p. 60.) 

Collected by W. W. Olive 

and R. H. Tschudy, U.S. 

Geological Survey, 1963.

D3228 _ _ _ NWViNW 1^ sec. 36, T. 6 N., 

R. 1 E., Riverside Park 

Jackson, Hinds County, 

Miss. Lignitic clay with 

borings. Collected by W. 

W. Olive and R. H. 

Tschudy, U.S. Geological 

Survey, 1963.

D3227 ___ NW'/iNW'/i sec. 36, T. 6 N., 

R. 1 E., Riverside Park, 

Jackson, Hinds County, 

Miss., 1 MJ ft below coquina 

bed. (See Moore, 1965, p. 

48.) Collected by W. W.

Cook Mountain Formation. In Claiborne 

Group, middle Eocene.

Cockfield Formation. 

In Claiborne Group, 

middle Eocene.

Middle part, Cockfield 

Formation. In Claiborne Group, middle 

Eocene.

Do.

Upper part, Cockfield 

Formation. In 

Claiborne Group, 

middle Eocene.

Do.

Moodys Branch Formation (reference 

section). In Jackson 

Group, upper Eocene.
NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

uses

I'.ileo botany

loc

Locality

Stratigraphic

unit

Olive and R. H. Tschudy, 

U.S. Geological Survey, 

1963.

D3697-B__ SE'iSW^SE 1 * sec. 32, T. 

12 N., R. 2 W., Yazoo 

County, Miss., 124 ft 

above base of formation. 

Core samples of Yazoo 

Clay supplied through the 

courtesy of the Mississippi 

Geological Survey, 1963.

D3699-A _ _ Jackson Ready-Mix Concrete Light Aggregate 

Quarry, SWUNE 1 ^ 

NW>4 sec. 36, T. 7N.,R. 1 

W., 7 miles west-southwest 

of Madison, Hinds County, Miss., 30 ft below top of 

formation. (See Priddy, 

1960, p. 40, fig. 9.) 

Collected by R. H. 

Tschudy, U.S. Geological 

Survey, 1966.

D4383-A _ _ Steep bluff on right bank of 

Arkansas River, in EVij 

sees. 19 and 30, T. 3 S., R. 

10 W., Jefferson County, 

Ark. (See Wilbert, 1953, p. 

114.) Collected by B. R. 

Haley and R. H. Tschudy, 

U.S. Geological Survey, 

1969.

D4407-B__ Center of east line NW ] 4 

sec. 36, T. 5 N., R. 3 K, 

Whitmore quadrangle, St. 

Francis County, Ark. Sample from 2 ft above marl. 

(Marine fossils. B. R. 

Haley, writen commun., 

Sept. 15, 1969.) Collected 

by B. R. Haley, U.S. 

Geological Survey, 1969.

Yazoo Clay (core hole 

at type locality). In 

Jackson Group, upper Eocene.

Yazoo Clay. In Jackson 

Group, upper Eocene.

Jackson Formation or 

base of White Bluff 

Formation of Jackson Group (Wilbert, 1953), upper 

Eocene.

Jackson Formation, 

upper Eocene.

Four additional Late Cretaceous samples not included 

in previous publications were used in this study. These 

samples are not reference samples but are samples submitted for age determination by geologists working on 

the Kentucky cooperative mapping project. These 

samples were used because they yielded particularly 

good specimens of some of the taxa discussed.

Sample Locality Formation

D3002 _ _ _ Drill hole 35 (depth 32-37 ft) 

immediately south of Kentucky Highway 444 on east 

side and adjacent to 

Beechy Creek; Kentucky

McNairy Sand Member of Ripley Formation, Maestrichtian.

Locality Formation

coordinates, South zone: 

E. 1,298,250 ft; N. 89,250 

ft, New Concord quadrangle, Galloway County, 

Ky. Collected by W. W. 

Olive, U.S. Geological 

Survey, 1962.

D3292__ From west bank of 

Grindstone Creek; Kentucky coordinates, South 

zone, E. 1,306,700 ft; N. 

95,200 ft; Hamlin quadrangle, Galloway County, 

Ky. Collected by W. W. 

Olive U.S. Geological 

Survey, 1963.

D3420___ From 1.15 miles northnortheast of Thomson 

Chapel, 550 ft north, 75 ft 

east of road junction, east 

side of U.S. Highway 60; 

0.9 mile south of north 

edge Little Cypress IVi 

minute quadrangle; Kentucky coordinates, South 

zone, E. 1,227,200 ft; N. 

292,400 ft; Livingston 

County, Ky. Collected by 

W. W. Olive, U.S. 

Geological Survey, 1964.

D4268-A _ _ Grand Rivers, Livingston 

County Highway 453 roadcut, 0.3 mile northwest of 

Grand Rivers; about middle of 10-foot-thick black 

clayey silt bed; Kentucky 

coordinates, South zone: 

E. 1,271,500 ft; N. 254,800 

ft; Livingston County, Ky. 

Collected by R. W. Swanson, U.S. Geological 

Survey, 1968.

D4268-B _ _ Same locality, immediately 

above black clayey silt.

McNairy Sand Member of Ripley Formation, Maestrichtian.

Do.

Do.

Do.

TYPE SPECIMENS

All specimens illustrated in this report are preserved 

on slides deposited in the paleobotanical collections of 

the U.S. Geological Survey, Denver, Colo. All illustrated 

specimens are within black-inked circles marked 

directly on the slides; they may also be located on the 

slides by the mechanical-stage coordinates given with 

the type designations and in the plate explanations. In 

order that others may convert their mechanical-stage 

readings to those recorded for the specimens included in 

this report, the coordinates for the center point of a 1- 

X3-inch standard microscope slide are 108.0X12.3 mm. 

The method of accurately locating the center of a stan-
SYSTEMATIC DESCRIPTIONS

dard microscope slide has been described by R. H. 

Tschudy (1966, p. D78). With the slide label to the left, 

the vertical coordinates decrease toward the near edge of 

the slide, and the horizontal coordinates decrease toward 

the right edge of the slide.

Color transparencies of type and illustrated specimens 

described in this report are available from the U.S. 

Geological Survey laboratory, Denver, Colo., on a 

limited-time loan basis.

SYSTEMATIC DESCRIPTIONS

The genera belonging to the Normapolles group are 

discussed in alphabetical order. Newly proposed genera 

are presented at the end of the report. An arbitrary 

minimum of 10 good specimens was selected as a requirement before a name could be assigned to a new 

genus or new species. As a result of this decision several 

probable new taxa are mentioned but not named. As an 

aid to future workers, provisional descriptions are 

provided for these unnamed entities, but no formal 

diagnoses are made. For those genera and species 

already established, generic diagnoses are given. For 

many genera, these diagnoses are somewhat modified 

from the original German; a conscientious attempt has 

been made to retain the meaning expressed in the 

original versions. Direct word-for-word translation would 

have provided much more stilted diagnoses.

In several descriptions Goczan and others (1967) 

provided new wordings ("Neufassungen") for diagnoses 

of already established genera. These new wordings are 

not considered to be emendations by those authors, nor 

do I consider them to be emendations. For example, Goczan and others (1967), under the heading Basopollis 

Pflug, 1953b (p. 449), say "Wie aus der Diagnose zu 

ersehen ist, wurde sie unwesentlich verandert, sie ist vor 

allem weiter prazisiert worden. Dies ist heute moglich 

durch die Vielzahl neugefundener Formtypen. Die 

Bemerkungen zur Einheit in R. Potonie 1960, Syn. Ill (S. 

126) sind uberholt und waren auch wenig zutreffend."

The complex morphology of pollen grains assigned to 

the Normapolles group of genera has prompted several 

European workers to institute new terminology for some 

of the more unusual morphological features of the grains. 

Many of these new terms, such as "interloculum" (space 

between the ektexine and endexine), are defined in the 

"Morphological Encyclopedia of Palynology" by Kremp 

(1965). All the terms used by Goczan and others (1967) 

in the new wordings of generic diagnoses are defined in 

section D of their publication.

Skarby (1968) raised some doubtless valid objections 

to the interpretations of the so-called diagnostic 

characters used by Pflug (1953) and by others for 

differentiating some of the Normapolles genera. Skarby 

(1968, p. 3) claimed that "Several of these diagnostic 

characters are derived from incorrect observations or

misinterpretations of existing properties * * *." This 

claim is to some degree true. Unfortunately, Skarby was 

not familiar with the paper by Goczan and others (1967) 

in which the Normapolles genera are more accurately 

differentiated and morphological features present are 

more accurately described. This publication removes the 

bases for most of her objections. Skarby emended the 

genus Extratriporopollenites to include 23 Normapolles 

genera and the Postnormapolles genus 

Triatriopollenites. She did not recognize characters, 

such as "atrium," "interloculum," "solution-meridian," 

"oculus," and "endanulus" as diagnostically significant. 

Certain other generic characteristics such as 

"praevestibulum" (in Basopollis), she suggested may be 

optical phenomena or misinterpretation as the result of 

poor lenses or poor observation.

I have observed the morphological features described 

in Goczan and others (1967) for certain European Normapolles genera in specimens from the Mississippi embayment. As a result of the careful observations recorded 

by Goczan and others (1967), I have experienced little 

difficulty in differentiating Normapolles genera on the 

bases established by the above publication. I do not 

believe that the extreme "lumping" of genera resorted to 

by Skarby serves any useful purpose. Furthermore, the 

inclusion by Skarby (1968) of such morphologically 

diverse genera as Triatriopollenites, Complexiopollis 

(Latipollis, Turonipollis), Basopollis, and Plicapollis in 

the genus Extratriporopollenites is, I believe, totally unjustified.

Genus BASOPOLLIS Pflug, 1953

Type species. Basopollis orthobasalis (Pf.) Pf.

Diagnosis. ("New wording" from Goczan and 

others, 1967.) Pollen grains triangular, shallow lensshaped to extreme triplane; equatorial contour rigid 

triangular to extremely concave triangular; the three 

equatorial germinals at the angles are sometimes 

strongly distorted but generally do not contribute particularly to the equatorial contour, so that normally the 

equatorial contour is not strongly interrupted. Polar axes 

of very different lengths. Wall several layered, the outer 

part thicker than the closely lying inner part; surface 

smooth or mostly with strong punctate sculpture. 

Sometimes with plicae (principally in those with a 

tendency to a triplane shape). Exopores vertical slit 

shaped, more or less symmetrical to the equator, with 

two lamella complexes in the exogerminal: (1) A 

relatively thick sculpture bearing outer layer without internal structure, and (2) a typical polyannulate complex 

that is present from the root of the exogerminal; these 

complexes are sometimes clearly structured with 

radially directed bacula. Pore canal index among the 

several form species very different, mostly above 0.3. 

Vestibulum present. Mesopore equatorial slit shaped 

with more or less gable-shaped protrusions of the upper
NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

and lower lips; lips are, however, generally without 

thickenings. "Endopore" is developed from the inner 

wall as a straight margined or weakly concave atrium. 

No oculi or polpapillae. Mostly medium sized, but 

sometimes large forms.

Remarks. Distinguished from Complexiopollis principally by the typical polyannulate exogerminals.

Basopollis obscurocostatus, n. sp.

Plate 1, figures 1-12; text figure 1

Holotype. Sample D1862, slide 3, prep. 2, coordinates 95.7X3.9. Wilcox Formation, Mississippi, lower 

Eocene.

Paratype. Sample D1862, slide 3, prep. 2, coordinates 92.6X16.1. Wilcox Formation, Mississippi, lower 

Eocene.

Diagnosis. Shape of pollen grains concave 

triangular in polar view, often distorted into the triplane 

form. Size 28jU-42|U in equatorial diameter (14 

specimens). Wall of two principal layers, about 2yu thick

FIGURE 1. Basopollis obscurocostatus, n. sp., ca. X 2,000. A, 

Aperture in polar view showing polyvestibulate characteristic of the 

exogerminal. B, Aperture in equatorial or partial triplane view showing arci and endexinal flaps. C, Surface view showing ornamentation.

in the interapertural areas. Presence of equatorial arci or 

ribs gives the impression of a thicker wall when grains 

are oriented in strict polar attitude; arci generally seen 

clearly only in triplane or equatorial view. Surface with 

scabrate to microverrucate sculpture. Ektexine about 

twice as thick as endexine. Apertural areas elongate. Exogerminals vertical slit shaped, bordered by 

characteristically polyannulate thickened ektexinal

walls. Outer layer of germinal areas thin, structureless; 

inner layer of radially projecting bacula of uneven 

lengths forming thereby one or more praevestibulae. 

Vestibulum at base of exogerminal prominent but 

narrow. Endexine at position of endogerminal separated 

into two gable-shaped flaps separated by a large 

equatorially oriented slit. Delicate atrium below flaps 

not observed in specimens examined.

Remarks. Basopollis obscurocostatus, n. sp., is distinguished from Basopollis orthobasalis by its more 

prominent concave triangular shape, thinner walls, and 

the presence of arci. The outer surfaces of Basopollis 

basdlis, B. atumescens, and B. ornatus are smooth in 

contrast to the distinctly sculptured outer surface of 

Basopollis obscurocostatus.

Stratigraphic distribution. Nanafalia, Wilcox, 

Pendleton, and Hatchetigbee Formations.

Genus CHOANOPOLLENITES Stover, 1966

Type species. Choanopollenites eximius Stover, 

1966 (in Stover and others, 1966), Kansas Univ. Paleont. 

Contr. Paper 5, p. 8, pi. 4, figs, la-lh.

The genus Choanopollenites and the stratigraphic distribution of its species have been discussed in a previous 

publication (Tschudy, R. H., 1973b). In that publication 

the following new species were proposed: 

Choanopollenites consanguineus, C. transitus, C. discipulus, C. conspicuus and C. patricius. Two other possible new species were mentioned but not named, owing to 

an inadequate number of good specimens.

cf. Choanopollenites sp.

Plate 1, figures 13-15; text figure 2

Sample D3138, slide 4, coordinates 88.5X13.2.

FIGURE 2. cf. Choanopollenites sp., ca. X 2,000. A. Aperture in 

polar view showing external prismatic layer and internal baculate 

layer. B, Same, showing the two endexinal flaps, one above the 

other. C, Surface view showing ornamentation.
SYSTEMATIC DESCRIPTIONS 9

Description. Shape of pollen grain triangular in 

polar view; apertures at the angles. Size 26^ in equatorial diameter (one specimen). Wall about 1.5^ thick in 

interapertural areas, ektexine about twice as thick as 

endexine. Surface coarsely fossulate. Exogerminals vertical slit shaped. Ektexine in germinal areas two layered, 

the outer layer columellate (prismatic) structured, 

narrow adjacent to apertures, then becoming distinctly 

thicker toward the roots of the germinals. Inner layer of 

prominent, radially directed bacula. Interior of exogerminals narrow funnel shaped, wider at base than at tip. 

Vestibulum narrow. Endogerminal equatorial slit 

shaped bordered by cresent-shaped somewhat thickened 

lips of endexinal flaps. Endexine structureless, thinner 

in interapertural areas than adjacent to the endogerminals.

Stratigraphic distribution. Naheola Formation 

only.

Genus COMPLEXIOPOLLIS Krutzsch, 1959, 

emend. Tschudy, 1973b

Type species. Complexiopollis praeatumescens 

Krutzsch, 1959, Palaeontographica, ser. B, v. 105, p. 135, 

pi. 31, figs. 39-54.

The genus Complexiopollis and the stratigraphic distribution of species from the Mississippi embayment 

assigned to the genus were presented in a previous 

publication by R. H. Tschudy (1973b). The following 

new species from Mississippi embayment rocks were 

proposed: Complexiopollis funiculus, C. patulus, C. abditus, and C. microuerrucatus. One other possible new 

species was mentioned but, because too few specimens 

were found, was not named.

Genus EXTREMIPOLLIS Krutzsch, 1967

Type species. Extremipollis maastrichtiensis 

Krutzch, 1967 (in Goczan and others, 1967), Palaontologische Abhandlungen, ser. B, v. 2, no. 3, p. 463, pi. 6, 

figs. 7-10.

Diagnosis. (From Krutzsch, in Goczan and others, 

1967). Pollen grains spherical to triangular shallow lens 

shaped; exogerminals equatorial, located at the angles. 

Exogerminals provided with three pairs of club-shaped 

to blunt elongate monannulate annuli. Inner part of annuli sometimes with structure. Exopores vertical slit 

shaped, somewhat symmetrical to the equator, mostly 

gaping. Endogerminals with deep incidence to platea 

(similar to many Trudopollis species). Wall several 

layered, ratio of the outer to the inner complex about 1:1. 

Sculpture various. Structure seldom present in the extragerminals. No oculi, endannuli, arci, plicae; only 

seldom a weak interloculum. Medium-sized forms.

Remarks. Distinguished from Extratriporopollenites by the presence of a strong incidence or 

cuneus, and ektexine and endexine of virtually the same 

thickness. Distinguished from Extrapollis by the

absence of a collarlike structure at the root of the germinals.

Extremipollis vivus, n. sp.

Plate 2, figures 1-10; text figure 3

Holotype. Sample D3420, slide 2, coordinates 

95.2X21.0, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Paratype. Sample D3420, slide 4, coordinates 

94.4X21.0, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Diagnosis. Shape of pollen grains triangular to 

slightly concave triangular in polar view. Apertural areas 

prominently protruding; exclusive of apertural areas, 

pollen grains almost circular in equatorial outline. Size 

20ju-29ji in equatorial diameter (18 specimens). Wall of 

two principal layers about 1.5/i thick in the interapertural areas. Surface shallowly fossulate, almost smooth. 

Ektexine and endexine of about equal thickness in interapertural areas. Ektexine thickened, annulate and 

made up of two layers in the apertural areas; outer layer 

structureless, inner layer consisting of tightly packed, 

radially directed elongate bacula; presence of bacula difficult to distinguish in most specimens except at the inner part of the exogerminals. Exogerminals vertical slit 

shaped, slightly eccentric to the equator, sometimes gaping. Vestibulum narrow and almost absent. Endexine 

thin, less than Iji thick, separated from ektexine by a 

narrow interloculum. Endopores about 4/u wide bordered 

by small endannuli. A prominent semicircular cuneus is 

present below each endopore.

Remarks. This species is distinguished from Extremipollis maastrichtiensis by its concave rather than 

convex triangular equatorial outline and by its more 

elongate exogerminals. Many specimens were observed 

in sample D3420.

Stratigraphic distribution. McNairy Sand 

Member, Ripley Formation.

FIGURE 3. Extremipollis vivus, ca. X 2,000. A, Section view of 

germinal. B, Slit-shaped exogerminal showing short slit on one side 

of grain. Compare with C. C, Long slit on obverse side of grain. Surface ornamentation shown.
10 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

Extremipollis versatilis, n. sp.

Plate 2, figures 11-22; text figure 4

Holotype. Sample D3420, slide 3, coordinates 

97.6X11.5, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Paratype. Sample D3420, slide 3, coordinates 

108.2X5.3, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Diagnosis. Shape of pollen grains triangular in 

polar view; apertural areas sometimes protruding and 

interrupting the triangular outline. Size 23/u-31/u in 

equatorial diameter (25 specimens). Wall of two principal layers, about 2/u thick in interapertural areas. Surface coarsely fossulate. Ektexine of about same thickness 

as endexine or slightly thicker in interapertural areas. 

Ektexine prominently annulate and made up of two 

layers adjacent to the apertures; outer layer structureless, inner layer of prominent radially directed 

bacula. Exogerminals vertical slit shaped, slightly 

eccentric to the equator, seldom gaping. Vestibulum 

narrow, almost absent, present only at the extreme root 

of the exogerminals. Endexine thin, commonly less than 

1/u thick and separated from the ektexine by an interloculum. Endopores 5/u-6/u wide; a prominent 

semicircular to somewhat V -shaped cuneus present 

below each endopore. Inner lamella of endexine forming 

an inconspicuous additional atrium in some specimens.

Remarks. Distinguished from Extremipollis uiuus 

by the absence of endannuli, less protruding exogerminals, and coarser sculpture. Many specimens present 

in sample D4268-A. Several apparently somewhat transitional forms were observed. This variation is 

demonstrated by apparently thinner endexine and 

slightly more narrow and protruding exogerminals (pi. 2, 

figs. 20-22).

Stratigraphic distribution. Coffee Sand, Coon 

Creek Tongue of Ripley Formation, and McNairy Sand 

Member of Ripley Formation.

FIGURE 4. Extremipollis versatilis, ca. X 2,000. A, Sectional view of 

germinal. B, Short slit on one side of grain. C, Elongate slit on 

obverse side of grain. Surface ornamentation shown.

Extremipollis caminus, n. sp.

Plate 3, figures 1-18; text figure 5

Holotype. Sample D3420, slide 4, coordinates 

79.7X4.5, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Paratype. Sample D3420, slide 2, coordinates 

84.3X14.8, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Diagnosis. Shape of pollen grains triangular to 

slightly concave triangular in polar view with prominent 

chimneylike protrusions (germinals) at the angles. Size 

25/U-41/U in equatorial diameter (30 specimens). Wall of 

two principal layers, about 1.5/u thick in the interapertural areas. Surface microverrucate with low relief.

FIGURE 5. Extremipollis caminus, ca. X 2,000. A and A', Sectional 

view of aperture showing variation in development of the visible 

baculate layer and the splitting of the endexine at the margins of the 

endogerminals. B and B', Elongate apertural slit present on one side 

of grain is in contrast with length of slit on obverse side. C and C", 

Short apertural slit of exogerminal on obverse side; variation of surface sculpture of small and large specimens is also shown.
SYSTEMATIC DESCRIPTIONS 11

Ektexine and endexine of about equal thickness in interapertural areas. In germinal areas the ektexine is 

greatly thickened and structureless in the outer part; the 

inner part appears lined with short radially projecting 

bacula. Exogerminals vertical slit shaped, distinctly 

eccentric to the equator (one side of slit extends a much 

greater distance toward the pole than does the other 

side). Neck of exogerminal area with a prominent 

biconvex chamber widens toward the base. Vestibulum 

narrow and present only at the base of the exogerminal. 

Endexine thin, about 0.5/it thick and separated from the 

ektexine by a narrow interloculum. Endopore 6/it-7/it wide 

subtended by a prominent U- or V-shaped cuneus. 

Endexine commonly splitting into two or more lamellae 

adjacent to the endopore.

Remarks. This species is distinguished from all 

other species of Extremipollis by the long chambered 

neck of the exogerminal. The two drawings in text figure 

5 show variation in the development of the ektexinal 

baculate layer and the finer surface ornamentation present on the smaller specimens.

Stratigraphic distribution. McNairy Sand 

Member, Ripley Formation.

Genus INTERPOLLIS Krutzsch, 1960

Intratriporopollenites supplingensis PL, 1953 (in Thomson and Pflug, 

1953), Palaeontographica, v. 94, ser. B, p. 89, pi. 10, fig. 26. 

Type species. Interpollis supplingensis (Pf.) 

Krutzsch, 1960, Geol. Gesell DDK. Berichte, v. 5, no. 4, 

p. 305.

Diagnosis. ("New wording" from Goczan and 

others, 1967.) Pollen grains triangular double-lens 

shaped, owing to an equatorial constriction, seldom 

single lens shaped because of a reduced constriction. 

Subequatorial contour mostly weakly convex triangular, 

sometimes also weakly concave triangular, seldom 

rounded triangular. Equatorial contour primarily 

strongly concave triangular with projecting rounded sectors at the angles. The polar regions often somewhat 

depressed and limited by band-shaped arcilike angle-toangle differentiation of both hemispheres. Wall two 

layered, ratio of ektexine to endexine about 1:1; walls 

sometimes separated by weak local interloculi. Surface 

smooth or weakly sculptured; inner layer often intrapunctate. Germinals at the angles without annuli, 

with two small, somewhat round subequatorial proximal 

and distal exopores at each angle. The endopores are at 

the equator. The structured inner wall layer sometimes 

forms a distinct visible atrium. Forms small to maximum middle sized.

Interpollis cf. I. supplingensis (Pf.) Krutzsch, 1960b

Plate 4, figures 1-21; text figure 6

Type specimen. (Lectotype designation, Pflug, 

1953, pi. 24, fig. 39).

FIGURE 6. Interpollis cf. /. supplingensis, ca. X 2,000. A, Large 

specimen in which baculate interloculum shows clearly. B, Smaller 

specimen. Triangular equatorial contour shown on this figure. C, 

Bacula in interloculum visible only under oil immersion lens.

Synonymy. Intratriporopollenites supplingensis 

Pf., in Thomson and Pflug, 1953.

Interporopollenites supplingensis Pflug, 1953. 

Interpollis supplingensis (Pf.) Krutzsch, 1960b.

Remarks. Interpollis supplingensis was first 

described by Pflug (in Thomson and Pflug, 1953) as Intratriporopollenites supplingensis. In a succeeding 

volume, Pflug (1953) transferred the species to Interporopollenites supplingensis without adding to the 

original diagnosis. Later, Krutzsch (1960b), after noting 

the presence of six exopores and an equatorial constriction, erected the new genus Interpollis and chose the 

species Interpollis supplingensis as the type species. 

Krutzsch did not provide a new diagnosis for the species, 

even though the original diagnosis by Pflug (in Thomson 

and Pflug, 1953) was extremely brief and inadequate. I 

am therefore including here a description of the 

Mississippi embayment forms that are similar to or conspecific with Interpollis supplingensis. This is not intended as an emendation.

Description. Shape of pollen grains triangular to 

almost circular, equatorial constriction prominent in 

some specimens, in others scarcely evident. Size 16
12 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

(30 specimens) in equatorial diameter, most grains in the 

20ju-25/u size range. Wall two layered, thin, about 

1.5/u-2/u thick; ektexine and endexine of about the same 

thickness separated by a very narrow baculate layer. Extexine smooth, appearing granulate except in polar 

areas; granulate appearance due to presence of short 

bacula between ektexine and endexine. Appearance of 

polar scar or thin area is due to absence of bacula. 

Baculate layer of ektexine distinctly thicker in atria than 

in interapertural areas. Exogerminals six, two at each 

angle of grains, subequatorial or nearly so, one on each 

hemisphere; apertures circular or slightly elongate 

equatorially. Endogerminals three, equatorial, 

somewhat hourglass shaped in equatorial view, without 

annuli; endexine separated from ektexine in apertural 

areas by a vestibulum.

Remarks. The original diagnosis lists the size of this 

species as 18^-40^. I found only three specimens as large 

as 28/u, and two specimens in the 16/u range. The 

equatorial constriction is shown particularly well on 

plate 4, figures 6-13 and 16-18.

Stratigraphic distribution. Cusseta Sand Member 

of Ripley Formation, Coon Creek Tongue of Ripley Formation, McNairy Sand Member of Ripley Formation, 

upper part of McNairy Sand, and Owl Creek Formation. 

One poor specimen (redeposited?) was observed in the 

Naborton Formation.

Interpollis microsupplingensis Krutzsch, 1960b 

Plate 4, Figures 22-25

Diagnosis. (From Krutzsch, 1960b.) A very small 

form-species of Interpollis. Size lOju-20/u, in section 

14,u-17,u. Double-lens shaped. Equatorial contour concave triangular, subequatorial contour strongly 

triangular to weakly convex triangular. Surface 

shagrenate to finely extrapunctate.

Remarks. Exopores very small, circular, about 1/u in 

diameter. Mississippi embayment specimens conform 

very well to the circumscription and illustrations of 

Interpollis microsupplingensis.

Stratigraphic distribution. Hatchetigbee and 

Reklaw Formations.

Interpollis intranodus, n. sp.

Plate 5, figures 1-15; text figure 7

Holotype. Sample D3285, slide 1, coordinates 

90.3X15.3, Porters Creek Clay, Kentucky, Paleocene.

Paratype. Sample D3285, slide 2, coordinates 

97.7X21.7, Porters Creek Clay, Kentucky, Paleocene.

Diagnosis. Shape of pollen grain convex-triangular, 

equatorial constriction not evident on most specimens. 

Size 20/u-36/u in equatorial diameter (11 specimens). 

Wall two layered, about 2/u thick. Ektexine and endexine 

of about equal thickness. Ektexine undulate to 

microrugulate, two layered; the outer layer structureless, 

the inner consisting of very short fine bacula. Bacula

FIGURE 7. Interpollis intranodus, ca. X 2,000. A, Peglike bacula 

visible in interloculum and in surface view. B, Wall layers with large 

peglike bacula and also small dense bacula attached to ektexine.

present except in polar areas. Interloculum prominent; 

scattered interlaminar short peglike bacula or gemmae 

separate endexine from ektexine. Exopores equatorially 

elongate, small, six in subequatorial pairs at the angles, 

not clearly evident on all specimens. Endopores in 

endexine, and atria almost identical to those found in 

Interpollis c.f. I. supplingensis.

Remarks. Distinguished from other species of Interpollis by the presence of scattered coarse bacula or gemmae separating the endexine and ektexine. Surface 

views of peglike bacula show particularly well on plate 

5, figures 1, and 13-15.

Stratigraphic distribution. Owl Creek and Clayton 

Formations, Porters Creek Clay, and Naheola Formation.

Genus INTERPOROPOLLENITES Weyland and Krieger, 1953

Type species. Interporopollenites proporus 

Weyland and Krieger, 1953, Palaeontographica, ser. B. 

v. 95, p. 20, pi. 1, fig. 34.

Diagnosis. ("New wording" from Goczan and 

others, 1967.) Compact triangular lens shaped; those 

with a weak equatorial constriction slightly double lens 

shaped. Equatorial contour from concave triangular to 

nearly circular. Polar axis always shorter than the 

equatorial axis. Wall always at least two layered; if 

more, two principal wall complexes recognizable; ratio of 

ektexine to endexine 1:1 to 1:5 and more. Sometimes 

without arci but mostly with arci, these very distinct in 

form and distribution (breadth, thickness, course); 

sometimes only plicalike folds present. Surface smooth 

to weakly sculptured; wall sometimes with structure 

(punctate, baculate). Seldom with an interloculum, in 

most species, however, with an endannulus. Endopores 

equatorial, at angles, mostly round to oval, never with 

strong incidence, cuneus, and so forth; inner lamella 

layer tightly attached. Six exopores present, at the
SYSTEMATIC DESCRIPTIONS 13

angles, one of each pair subequatorial on the distal and 

one on proximal side; exopores round, oval or polygonal, 

never very large; these may also be located somewhat 

laterally. The exogerminal is closed at the equator, and 

in some form-species this area may be thickened annuluslike to pointed. Forms small to medium-sized, very 

seldom larger representatives.

Interporopollenites turgidus, n. sp.

Plate 6, figures 1-18; text figure 8

Holotype. Sample D3548-A, slide 1, coordinates 

77.5X11.8, Clayton Formation, Kentucky, Paleocene.

Paratype. Sample D3548-A, slide 2, coordinates 

78.4X6.5, Clayton Formation, Kentucky, Paleocene.

Diagnosis. Pollen grains triangular in polar view 

with very prominent hemispherical protrusions at the 

angles. Size 24ju-29ju in equatorial diameter (10 

specimens); hemispherical corner protrusions about onethird the diameter of the grains. Wall of two principal 

layers; the ektexine in the interapertural areas thin, 

about 0.5ju thick, in the apertural areas much thicker, 

owing to the presence of an additional internal baculate 

layer. Endexine much thicker than ektexine, in the interapertural areas 2ju-3ju thick. Ektexine usually smooth, 

faint granular arcuslike sculpture extending from aperture to aperture present on some specimens. Two small 

exogerminals present in each of the three germinal areas, 

one on each side subequatorially; germinal pores may be 

elongate equatorially or in the polar direction. Exogerminal wall 2yu-3/u thick, thinner toward the root of the 

prominent vestibulum. Bacula in the interior of the exogerminal areas very abundant and tightly packed, often 

somewhat difficult to observe. Endogerminal in the form 

of a circular equatorial pore, much larger than the exogerminal pores. Endexine adjacent to the pores with endannuli as much as 5jw thick; endexine in pore area may 

be split into several distinct lamellae.

FIGURE 8. Interporopollenites turgidus, ca. X 2,000. A, B, Sectional 

views of apertures showing baculate areas, thick endexine, large 

equatorial endexinal pores and small nonequatorial exopores. C, 

Surface view showing nonequatorial exopore and faint arci.

Remarks. Distinguished from Interporopollenites 

proporus by the faint arcuslike exterior sculpture and by 

the thickened ektexine in the exogerminal area.

Stratigraphic distribution. Clayton Formation and 

Porters Creek Clay.

Genus KYANDOPOLLENITES Stover, 1966

Type species. Kyandopollenites anneratus Stover 

(in Stover and others, 1966), Kansas Univ. Paleont. 

Contr. Paper 5, p. 6, pi. 3, figs. 1, 2.

Diagnosis. (From Stover, 1966.) "Pollen isopolar, 

triaperturate; apertures aspidate, equatorial, compound. Outer aperture circular or nearly so, inner aperture elliptical, elongate. Grains circular in polar view, 

oblate in equatorial view. Exine relatively thick (5^-8^), 

sexine thicker than nexine; endosexine lacking or very 

thin around apertures. Surface granulate on type 

species. Monotypic."

Kyandopollenites anneratus'Stover, 1966

Plate 7, figures 1-5; text figure 9

Remarks. Although Stover (in Stover and others, 

1966) did not definitely assign Kyandopollenites to the 

Normapolles group, he did compare the genus with two 

Normapolles genera, Emscheripollis and Papillopollis. 

The size range of Kyandopollenites given by Stover is 

47.5/U-54M- Specimens measured from our material increase the range slightly to 41jw-58/u 

Stratigraphic distribution. Wilcox and Pendleton 

(Andersen, 1960) Formations.

Genus MEGATRIOPOLLIS Goczan and Krutzsch, 1967

Type species. Megatriopollis santonius Groot and 

Krutzsch, 1967 (in Goczan and others, 1967), Palaontologische Abhandlungen, ser. B, v. 2, no. 3, p. 477, pi. 

10, figs. 17-22.

Diagnosis. (From Goczan and Krutzsch, in Goczan 

and others, 1967.) Pollen grains triangular shallow lens 

shaped, equatorial contour triangular to triangular convex. The equatorial contour of the inner part of the 

pollen grains is circular (rund bis uberrund). Wall of two 

principal layers, the outer thicker than the inner, the 

latter sometimes very thin. Surface variously strongly 

sculptured; sometimes with a structured layer in the inner part of the wall. Tendency to a "Schichtfuge" or interloculum between the two wall layers. Exogerminal 

with annulus or tumescens, it is, however, monannulate, 

although sometimes showing fine structure in the inner 

part. Exopores vertical slit shaped, lying weakly 

asymmetrical to the equator. With vestibulum. Endogerminal without essential thickening, with a fairly 

wide endopore that is generally bounded by a shallow 

atrium. The opening of the innermost wall layer is 

similarly formed. No oculi, arci, plicae, and so forth. 

Forms medium sized.
14 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

FKIUHK 9. Kyandopoltenites anneratus Stover, 1966. A, Pollen grain, 

ca. \ 1,000, showing sectional view of aperture and surface sculpture. B, Sectional view of pore, X 2,000, showing three prominent 

wall layers.

Megatriopollis? sp.

Plate 7, figures 6-10; text figure 10

Description. Shape of pollen grains triangular to 

slightly convex-triangular in polar view. Size 25^-29^ in 

equatorial diameter (two specimens). Wall thin, two 

layered, l/u-1.5/u thick in interapertural areas. Surface 

smooth or finely punctate, owing to the structure of the 

outer wall. Exogerminals vertical slit shaped, ektexine 

adjacent to exogerminals in the form of labra, outer part 

of labra structureless, inner part fine structured, in our 

specimens radially directed bacula observed. Endexine 

very thin, separated from ektexine by a narrow interloculum. No discernible vestibulum. Atrium large, 

with plane or slightly concave inner boundary.

Remarks. This species was not named, because too 

few specimens were available. The specimens observed 

differ from the circumscription of the genus by the 

absence of a recognizable vestibulum. However, the 

presence of a vestibulum is not clearly represented in the 

photographs of the type species.

Stratigraphic distribution. Coffee Sand and Porters 

Creek Clay.

Genus MINORPOLLIS Krutzsch, 1959

Type species. Minorpollis minimus Krutzsch, 1959, 

Palaeontographica, ser. B, v. 105, p. 141, pi. 32, figs. 

10-14.

Diagnosis. ("New wording" from Goczan and 

others, 1967.) Pollen triangular shallow lens shaped, 

seldom somewhat triplanoid because of slightly longer 

polar axis; equatorial contour round to strongly 

triangular with sometimes a crooked side contour. Wall 

two layered, presence distinguishable. Surface smooth to 

finely sculptured. With three equatorial germinals at the 

angles; exogerminal with oval to vertical slit-shaped outer opening, with annulus, tumescence, or labrum. 

Without oculus. Endogerminal variously structured, 

sometimes with endannulus; endopores displayed as 

equatorial elongated oval to vertical oval. Sometimes 

with indications of plicae. Often secondary folding but 

without solutions. Size: 6^ to about 15/t; only very small 

forms assembled here.

Minorpollis minimus Krutzsch, 1959

Diagnosis. (From Krutzsch, 1959.) A lens-shaped 

form-species of Minorpollis with a weak tendency to 

sinuose folding, somewhat rigid equatorial contour, 

sometimes with a weak constriction of the Turonipollis 

type. Exine two layered, very thin, surface shagreenate, 

exogerminal with long club-shaped annulus. Endexine 

relatively thick, sometimes with weak endannulus. Endopores equatorially elongate, no typical atrium present. 

Size S

Minorpollis aff. M. minimus

Plate 8, figures 1-9; text figure 11

Description. Shape of pollen grains triangular with 

rounded angles in polar view, apertural areas slightly 

swollen, interrupting the outline. Size 13/u-16/u in 

equatorial diameter (27 specimens). Wall no more than 

IM thick in interapertural areas, ektexine and endexine 

of about equal thickness. Ektexine thin, smooth. Exogerminal oval to short vertical slit shaped; ektexine adjaFIGURE 10. Megatriopollis^ sp., ca. X 2,000. Sectional view of 

aperture (A), showing baculate area adjacent to exogerminal, deep 

atrium, wide endogerminal, and interloculum.
SYSTEMATIC DESCRIPTIONS 15

FIGURE 11. Minorpollis aff. M. minimus, ca. X 2,000. A, Sectional 

view of aperture showing baculate area and triangular extension of 

the endexine below the exogerminal.

cent to the exogerminal two layered, the outer layer 

structureless, the inner layer of short radially projecting 

bacula. Baculate area readily visible in some specimens, 

obscure in others; ektexinal walls adjacent to exogerminal annulate to labrate. Endexine thin and with 

irregular margin below the exogerminal, commonly with 

a pointed or triangular outward projecting extension. 

This extension subtends two oval vestibulumlike areas 

immediately below the baculate layer of ektexine. Endogerminal probably equatorial slit shaped; exogerminal 

slit smaller.

Remarks. The specimens found are all larger than 

the size mentioned by Krutzsch (1959) for Minorpollis 

minimus (8/u-ll/i). Otherwise our specimens appear to 

conform to the circumscription of the species. The pollen 

grains are similar to grains of Pseudoplicapollis serenus 

except that they do not possess plicae. Furthermore, 

Minorpollis aff. M. minimus has a different 

stratigraphic range.

Stratigraphic distribution. Clayton Formation and 

Porters Creek Clay.

Genus NUDOPOLLIS Pflug, 1953

Extratriporopollenites endangulatus Pflug, 1953 (in Thomson and 

Pflug, 1953), Palaeontographica, ser. B, v. 94, p. 72, pi. 6, fig. 37.

Nudopollis endangulatus (Pflug) Pflug, 1953, Palaeontographica, ser. 

B, v. 95, p. 108, pi. 25, fig. 24.

Type species. Nudopollis endangulatus (Pf.) Pflug.

Diagnosis. ("New wording" from Goczan and 

others, 1967.) Pollen grains triangular shallow lens 

shaped, the three equatorial germinals located at the 

angles, sometimes very prominent; germinals monannulate, seldom poly annulate. Equatorial contour from 

slender concave triangular (with sometimes slightly 

irregular sides) to triangular concave extending to forms 

with only slightly elongate exogerminals. Central body 

round to triangular convex. Without or with weak plicae. 

Wall variously thick, the outer part mostly thicker than 

the sometimes thinner inner part. Intermediate development is observed in some form species; for example, in 

"endangulatus" a fairly pronounced structure layer is 

developed and enlarged in the germinal region to form 

the mesopore. Surface smooth to occasionally weakly 

sculptured. Exopores vertical slit shaped, and weakly

asymmetrical to the equator, opening outward, toward 

the inner part oval-round, that is with a strongly constricted canal ending. Annuli sometimes with distinct 

structure in the outer wall (punctate-baculate). The 

mesopore is developed as a basketlike filter. Endogerminal without thickening; endopores formed as large, 

variously deep, half circular atria. No arci, oculi only as 

a tendency, interloculum seldom (probably only as a 

secondary development). Medium-sized forms.

Nudopollis endangulatus (Pflug) Pflug, 1953

Diagnosis. (From Pflug, in Thomson and Pflug, 

1953.) 25/U-35/U. Contour strongly triangular to weakly 

concave. Inner contour convex triangular, not circular. 

Long pore canal. Annulus with pointed tip. Endexine 

thinner than ektexine, mostly very thin, the ektexine 

closely appressed at all points. No solution meridian, exine shagreenate.

Nudopollis aff. N. endangulatus

Plate 8, figures 10-11

Remarks. Few specimens found. Both the ektexine 

and the endexine are very thin in interapertural areas. 

"Filter apparatus" very poorly developed in N. endangulatus in contrast to its development in N. terminalis.

Stratigraphic distribution. Nanafalia Formation.

Nudopollis terminalis (Thomson and Pflug) Pflug, 1953

Extratriporopollenites terminalis Thomson and Pflug, 1953, Palaeontographica, ser. B, v. 94, p. 71, pi. 6, fig. 31.

Nudopollis terminalis (Thomson and Pflug) Pflug, 1953, Palaeontographica, ser. B, v. 95, p. 161. pi. 22, figs. 1-6.

Diagnosis. (From Pflug and Thomson, 1953.) 

20/u-24ju. Contour convex. Inner contour more triangular 

than round. Pore canal not more than 8/u long, centripetally conically expanded. Without vestibulum but 

with atrium. Endexine at best half as thick as ektexine. 

Exine smooth to shagreenate, at the atrium intrabaculate.

Nudopollis terminalis subsp. hastaformis

Diagnosis. Annulus with pointed tips.

Nudopollis terminalis subsp. hebeformis

Diagnosis. Annulus with blunt tips.

Remarks. Extratriporopollenites terminalis, including the two subspecies, was transferred to 

Nudopollis terminalis by Pflug (1953) on the explanation 

of plate 22, p. 161. The subspecies N. hastaformis according to the International Rules of Botanic 

Nomenclature must become Nudopollis terminalis terminalis. However, the illustrations of the two subspecies 

do not clearly distinguish the two. Variation within the 

species is such that no good purpose is served by attempting to recognize the subspecies. They are mentioned here as a means of including the complete original 

diagnosis.
16 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

Nudopollis aff. N. terminalis (Thomson and Pflug) Pflug, 1953

Plate 8, figures 16-25

Remarks. The specimens illustrated most closely 

approximate Nudopollis terminalis terminalis. There is 

a recognizable stratigraphic size difference observed in 

this species. The Paleocene specimens range from 22/u to 

about 30/u in equatorial diameter, whereas those from the 

Eocene range from 22^ to about 38^ in equatorial 

diameter. No specimens larger than 30jU were observed in 

Paleocene or older samples. The size range of N. terminalis given by Thomson and Pflug is 20jU-40/u.

Stratigraphic distribution. All formations from 

Porters Creek Clay through the Cockfield Formation.

Nudopollis thiergartii 1 (Thomson and Pflug, 1953 

[non R. Pot., 1951]) Pflug, 1953

Extratriporopollenites thiergarti Thomson and Pflug,

1953. Palaeontographica, v. 94, ser. B., p. 71, pi. 6, fig. 43 

Nudopollis thiergarti (Thomson and Pflug) Pflug, 1953, Palaeontographica, v. 95, ser. B., p. 109. 

Nudopollis thiergarti (Thomson and Pflug, 1953 [non R. Pot. 1951])

Pflug, 1953, Berichte Geol. Gesel DDR., v. 5, no. 4, p. 326 (Lectotype

designation by Krutzsch, 1960b).

Diagnosis. (From Thomson and Pflug, 1953.) 

12/U-35/U. Contour convex to weakly concave, inner contour always circular. Endexine tightly appressed to 

ektexine, scarcely recognizable, without solution meridian or with it only intimated. Rodlet layer not evident. 

Exine shagreenate to faded intrarugulate. Pore canal 

sometimes gaping centrifugally. Praevestibulum 

sometimes suggested.

Nudopollis thiergartii subsp. meridium Pflug, 1953, 

in Thomson and Pflug, 1953

Diagnosis. 25^-50^. Small canal-shaped solution 

meridian, exine smooth to shagreenate. Ektexine in interapertural areas less than 2^u thick. Contour concave.

Nudopollis thiergartii subsp. conspicuus Pflug, 1953, 

in Thomson and Pflug, 1953

Diagnosis. 25/u-35ju. Contour mostly concave to interrupted concave. No solution meridian. Shallow 

fossulate sculpture. Ektexine in interapertural areas 

more than 2/u thick.

Nudopollis thiergartii subsp. minutus Pflug, 1953, 

in Thomson and Pflug, 1953

Diagnosis. 12^-25^- Contour mostly weakly convex. 

Minute semicircular solution meridian. Shallow 

fossulate sculpture. Ektexine in interapertural areas less 

than 2jU thick.

Remarks. The species Nudopollis thiergartii is discussed in detail by Krutzsch (1960b, p. 326). Krutzsch 

designated the specimen plate 6, figure 43, of Thomson 

and Pflug (1953) as the lectotype. This specimen was 

originally designated as Nudopollis thiergartii subsp. 

conspicuus. Krutzsch called attention to the fact that 

according to International Rules of Botanic

'Originally spelled thiergarti. International Code of Botanical Nomenclature, Article 73 

recommends the change from "i" to "ii" when the name of a person ends in a consonant.

Nomenclature this subspecies must be designated as 

Nudopollis thiergartii subsp. thiergartii. Both Pflug 

(1953, p. 109) and Krutzsch (1960b, p. 326) indicated 

that this species is variable and complex. Diagnoses of 

subspecies included here for completeness.

Nudopollis aff. N. thiergartii 

Plate 8, figures 12-15

Remarks. Fairchild and Elsik (1969) indicated that 

this species is present in the Paleocene and early Eocene 

and becomes virtually extinct at the end of early Eocene 

time.

Stratigraphic distribution. McNairy Sand through 

the Tallahatta Formation in our reference samples.

Genus PECAKIPOLLIS Kr. and Paclt., 1967

Type species. Pecakipollis bohemiensis Kr. and 

Paclt., 1967, Palaontologische Abhandlungen, ser. 8, v. 

2, no. 3, p. 486, pi. 12, figs. 14-20.

Diagnosis. (From Kr. and Paclt., in Goczan and 

others, 1967.) Pollen grains triangular shallow lens 

shaped, equatorial contour interrupted triangular to circular. Equatorial contour of the central body circular, 

the three equatorial germinals at the angles affect the 

contour strongly. Wall of two principal layers, ratio of 

about 1:1, not thick, without structure. Surface with 

weak sculpture, punctate, shagreenate to weak 

irregularly wrinkled. No interloculum. Exogerminals 

with slightly thickened plump labra, thereby forming a 

large vestibulum. Exopore weakly vertical slit shaped to 

elongate oval, outer opening arranged weakly 

asymmetric to the equator. On inspection the exogerminal appears as a shallow to equilateral triangle, or correspondingly a blunt-tipped trapezoid. The outer opening of the endogerminal is constructed of an equatorial 

elongate slit in the outer lamella with a straight or 

weakly convex outer margin. The inner opening is constructed by one or more broad fringed to jaggedappearing atria formed by the divided ends of the 

lamellae on both sides of the inner opening. No (or in any 

case, no typical) plicae, arci, oculi, and so forth, are 

developed. Forms small to at most medium sized.

cf. Pecakipollis sp.

Plate 9, figures 1-5; text figure 12

Remarks. One specimen, 23/u in equatorial 

diameter, was found. This specimen appears to possess 

most of the characteristics of Pecakipollis. The ektexine 

of this specimen is 1.5/u-2jU thick, very much thicker than 

the endexine. A "Schichtfuge" appears to be present 

between the ektexine and endexine. The ends of short 

bacula can be seen in the large vestibula. Insufficient 

specimens were found to provide an adequate evaluation 

of this taxon.

Stratigraphic distribution. Coon Creek Tongue of 

Ripley Formation.
SYSTEMATIC DESCRIPTIONS 17

Genus PLICAPOLLIS Pflug, 1953, emend. Tschudy

Type species. Plicapollis serta Pflug, 1953, Palaeontographica, v. 95, ser. B, p. 97, pi. 19, figs. 7-9.

Diagnosis. Pollen grains triangular flat lens shaped 

to triplanoid. Equatorial contour varies from triangular 

convex to triangular concave. Wall of two principal 

layers, ratio about 1:1, surface sculpture smooth to 

verrucate. With plicae, these generally prominent. Three 

germinals equatorially arranged at the angles. Exogerminals usually with somewhat strong annuli; annuli 

sculptured on the inner face with radial bacula, 

sometimes strongly, sometimes less so. Pore canal index 

less than 0.3. Exopores shallowly vertical slit shaped to 

circular, commonly symmetrical with the equator. With 

vestibula. Endogerminals slightly to strongly thickened 

adjacent to the somewhat equatorial slit shaped to oval 

or circular endogerminals (seen in equatorial view); in 

polar view, they possess a somewhat straight to weakly 

convex or concave boundary. No arci, no oculi; a 

"Schichtfuge" or narrow interloculum may be present. 

Mostly small to medium-sized forms.

Remarks. In the original short diagnosis, Pflug 

(1953) referred to Plicapollis as tricolporate with pores 

all equatorial. Goczan and others (1967) enlarged upon 

the diagnosis without emending it, as follows:

Diagnosis. ("New wording" from Goczan and others, 1967.) Pollen 

grains triangular flat lens shaped to triplanoid. Equatorial contour 

varies from triangular convex to triangular concave. Wall with two 

principal layers, ratio about 1:1, surface mostly smooth. With plicae, 

these more or less prominent, sometimes reduced. Three germinals 

equatorially arranged at the angles. Exogerminals with somewhat 

strong annuli; annuli sculptured on the inner face with radial bacula, 

sometimes strongly, sometimes less so. Pore canal index less than 0.3. 

Exopores vertical slit shaped, terminating outward, mostly approximately symmetrical with the equator. With vestibulum. Endogerminals not thickened, with somewhat equatorial slit shaped to 

elongated oval pores (seen in equatorial view); in polar view they 

possess a more or less straight to weakly convex or concave boundary. 

No arci, no oculi, no interloculum. Mostly small to medium sized 

forms.

Goczan and others (1967) categorized the exogerminals as being vertical slit shaped and stated that the

FIGURE 12. cf. Pecakipollis sp., ca. X 2,000. A. Sectional view of

aperture.

endogerminals were not bordered by thickenings. The 

photographs provided by these workers do not show 

these details with sufficient clarity to rule out the 

possibility that some of the species included in the genus 

Plicapollis may possess circular pores and a thickened 

endexine adjacent to the endogerminals. Specimens 

from Mississippi embayment rocks otherwise conforming to the circumscription of Plicapollis all possess 

definite unequivocal pores. (See pi. 9, figs. 6-24; pi. 10, 

figs. 1-21.) Most of the specimens from the Mississippi 

embayment assigned to the genus Plicapollis also show a 

somewhat pronounced thickening of the endexine adjacent to the endogerminals. Rather than creating a new 

genus to accommodate these specimens, I have emended 

the somewhat inadequate original diagnosis to include 

them.

Plicapollis usitatus, n. sp.

Plate 9, figures 6-14; text figure 13

Holotype. Sample D3260, slide 2, coordinates 

89.8X10.3, Cusseta Sand Member, Ripley Formation, 

Alabama, Campanian.

Paratype. Sample D3260, slide 1, coordinates 

110.8X17.7, Cusseta Sand Member, Ripley Formation, 

Alabama, Campanian.

Diagnosis. Shape of pollen grains triangular, with 

rounded rather than blunt angles. Plicae present, extending from base of germinals to adjacent germinals 

over the poles. Size 16ju-22M in equatorial diameter (16 

specimens). Wall thin, about 1.5/i thick in interapertural 

areas. Ektexine and endexine of about equal thickness 

and separated by a very narrow interloculum or 

"Schichtfuge"; ektexine smooth. Exogerminals circular 

pores, surrounded by ektexinal annuli. Ektexine in apertural areas two layered; outer layer thin and structureless, inner consisting of an elongate baculate layer; 

bacula longer near exopore, decreasing in length toward 

the root of the germinal. Vestibulum prominent, roomy, 

4/u-S/u deep. Endexine at base of germinals, somewhat 

annulate, sometimes splitting into several lamellae. Endogerminals larger than exogerminals, circular to oval

FIGURE 13. Plicapollis usitatus, ca. X 2,000. A, B, Sectional views of 

germinals, showing dense baculate layer, narrow interloculum, and 

annulate endogerminals. C, Surface view showing pore and plica.
18 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

with irregular margins.

Stratigraphic distribution. Cusseta Sand Member 

of Ripley Formation, Coffee Sand, Coon Creek Tongue of 

Ripley Formation, and McNairy Sand Member of Ripley 

Formation.

Plicapollis retusus, n. sp.

Plate 9, figures 15-24; text figure 14

Holotype. Sample D3412, slide 3, coordinates 

112.5X2.4, Coffee Sand, Tennessee, Campanian.

Paratype. Sample D3412, slide 4, coordinates 

106.0X19.9, Coffee Sand, Tennessee, Campanian.

Diagnosis. Pollen grains triangular to slightly concave triangular, with blunt, slightly swollen angles 

(apertural regions). Plicae prominent, extending from 

the base of the germinal region to adjacent germinals 

over the poles. Size 20/u-28/u, mostly about 23/u, in 

equatorial diameter (14 specimens). Wall thin, 1.5/u-2/u 

thick in interapertural areas; ektexine and endexine of 

about equal thickness, separated in some specimens by a 

very narrow interloculum; ektexine microverrucate. Exogerminals circular pores surrounded by ektexinal annuli. Ektexine in apertural areas two layered; the outer 

layer thin and structureless, the inner consisting of 

somewhat robust radially directed bacula. Vestibulum 

prominent, roomy, about 5/j. deep. Endexine at base of 

germinal enlarged, somewhat annulate, commonly split 

into several lamellae. Endogerminal larger than exogerminal, oval in equatorial direction and with irregular 

margins.

Remarks. Pollen of Plicapollis retusus can be distinguished from that of Plicapollis usitatus by its more 

blunt termination of the germinals, by its slightly larger 

size, and by its microverrucate, rather than smooth, surface sculpture.

Stratigraphic distribution. Coffee Sand and Coon 

Creek Tongue of Ripley Formation.

Plicapollis vacuus, n. sp.

Plate 10, figures 1-9; text figure 15

Holotype. Sample D3412, slide 4, coordinates 

87.0X3.1, Coffee Sand, Tennessee, Campanian.

Paratype. Sample D3412, slide 1, coordinates 

78.9X17.7, Coffee Sand, Tennessee, Campanian.

Diagnosis. Shape of pollen grains triangular with 

rounded angles (apertural areas); usually straight sided, 

sometimes slightly concave or convex. Delicate plicae 

extend from the base of each aperture and join over the 

poles. Size 16/u-23/u in equatorial diameter, most 

specimens about 18/u (based on 16 specimens). Wall 

about l/j. thick in interapertural areas, ektexine and 

endexine of about equal thickness; sometimes a narrow 

interloculum or "Schichtfuge" can be distinguished 

between the two wall layers; ektexine smooth to 

microverrucate. Exogerminal a circular pore in the 

ektexine. Ektexine in germinal areas generally one 

layered, structureless, and without thickening. In the inFKJITRE 14. Plicapollis retusus, ca. X 2,000. A, B, Sectional views of 

germinals showing baculate layer, narrow interloculum and annulate 

endogerminals. C, Surface view showing surface sculpture, pore, and 

plica.

terior of the wall near the pore, scattered short bacula 

are visible, more abundant near the pore than at the 

base of the germinals. In occasional specimens bacula 

are aggregated near the exopore and form a labrum or 

slight annulus. Vestibulum prominent, about 3yu deep. 

Endexine thin, terminating at base of germinal with little or no thickening. Endogerminal oval in equatorial 

direction with irregular margins.

Remarks. Pollen assignable to Plicapollis vacuus 

can be distinguished from that of Plicapollis usitatus 

and Plicapollis retusus by the virtual absence of 

aggregates of bacula in the exogerminal, by the possession of thinner walls, and by its generally smaller size.

Stratigraphic distribution. Coffee Sand and 

McNairy Sand Member of Ripley Formation.

Plicapollis rusticus, n. sp.

Plate 10, figures 10-21; text figure 16

Holotype. Sample D3412, slide 4, coordinates 

110.9X9.4, Coffee Sand, Tennessee, Campanian.

Paratype. Sample D3412, slide 1, coordinates 

108.0X11.8, Coffee Sand, Tennessee, Campanian.

B

FIGURE 15. Plicapollis vacuus, ca. X 2,000. A, B, Sectional view of 

apertures, showing only scattered bacula, circular pores, and narrow 

interloculum. C, Surface view showing plica.
SYSTEMATIC DESCRIPTIONS 19

Diagnosis. Shape of pollen grains triangular to 

slightly concave triangular in polar view, with blunt corners (apertural areas). Plicae extend from the base of the 

germinals over the poles to adjacent germinals. Size 

22/U-29/U in equatorial diameter (14 specimens). Wall 

1.5/u-2;u thick in interapertural areas, ektexine slightly 

thicker than endexine and separated from it by a narrow 

interloculum, extexine verrucate with low relief. Exogerminals circular pores surrounded by annulate, twolayered ektexine; outer layer structureless, inner layer of 

tightly packed, radially directed bacula. Vestibulum 

prominent, about 4/u deep. Endexine enlarged, 

somewhat annulate at the endogerminals, separating 

into indistinct lamellae. Endogerminal oval in 

equatorial direction with irregular margins.

Remarks. Pollen of Plicapollis rusticus is distinguished from that of other species by its prominent 

verrucate sculpture and its somewhat larger size.

Stratigraphic distribution. Coffee Sand.

FIGURE 16. Plicapollis rusticus, ca. X 2,000. A, B, Sectional views of 

apertures, showing circular pores, baculate areas, interloculum, and 

annulate endexine. C, Surface view showing sculpture and plica.

Genus POMPECKJOIDAEPOLLENITES Krutzsch, 1967

Type species. Pompeckjoidaepollenites subhercynicus (Krutzsch, 1954) Krutzsch, 1967 (in Goczan and 

others, 1967), Palaontologische Abhandlungen, ser. 8, v. 

2, no. 3, p. 491, pi. 13, fig. 25.

Synonymy. A guide to the synonymy of the species 

P. subhercynicus is provided by Krutzsch (in Goczan 

and others, 1967, p. 492).

Diagnosis. (From Krutzsch, in Goczan and others, 

1967) Pollen shallow triangular to oval lens shaped, 

equatorial contour triangular convex to round. Wall with 

two distinct principal layers separated by an interloculum. Surface fine to distinctly punctate (LOpattern) with a columella layer as the interior limit of 

the outer-wall section. Outer wall, in addition, 

sometimes with structure (baculate, tubulate, and so 

forth). With three equatorial germinals at the angles.

Exogerminals with annuli, these mostly radial baculate 

structured. Exopores not vertical slit shaped, but, at 

most, only vertical oval or, in general, only circular. Endopori connected over the poles by the development of 

platea, whereby the pole regions sometimes can have 

endexinous islands. With or without endannuli in the 

equatorial region (when viewed in optical section). Small 

to medium sized, rarely large forms.

Remarks. This genus is listed by Goczan and others 

(1967) as Pompeckjoidaepollenites (Pflug, 1953) n. 

emend. W. Kr. However, Pompeckjoidae-pollenites was 

not listed as a genus by Pflug (1953) but only as a section 

of the genus Trudopollis. Pollen assignable to this new 

genus can be distinguished from that now included in 

Trudopollis by the presence of round to oval exopores 

rather than vertical slit-shaped germinals and by the 

presence of fully developed platea.

Pompeckjoidaepollenites aff. P. subhercynicus (Kr., 1954) Kr., 1967 

Plate 11, figures 1-8

Remarks. The lectotype for Trudopollis subhercynicus (now Pompeckjoidaepollenites subhercynicus) is 

the specimen-plate 4, figure 12, of Potonie (1934). (See 

Krutzsch, 1961.) The original description of this taxon, 

in the light of more recent observations and interpretations of the fine structure of Normapolles genera, is 

somewhat misleading, I have therefore included below a 

description of the Mississippi embayment specimens 

that I believe are closely related to Pompeckjoidaepollenites subhercynicus.

Description. Shape of pollen grains convex 

triangular to almost circular in polar view, apertural 

areas slightly protruding. Size 21^-26^ in equatorial 

diameter (10 specimens). Wall of two principal layers, 

about 2/u thick in the interapertural areas, surface 

smooth, appearing scabrate to microverrucate, owing to 

the presence of bacula in the interloculum. Ektexine and 

endexine of about equal thickness in interapertural 

areas; ektexine thickened and annulate and consisting of 

two distinct layers at the apertures. Outer ektexine 

layer structureless, inner layer of radially projecting 

bacula that appear to fill the vestibula. Exogerminals 

consist of circular pores opening into a parallel-walled 

channel. Endexine in the form of three platea separated 

by a solution meridian over the poles. Individual platea 

cup shaped and separated from the ektexine by a prominent interloculum, inner layer of ektexine provided with 

short bacula that project into the interloculum. Platea 

may become disattached from the ektexine and lie at 

random in the cell lurnen. Outer margins of the endexine 

thickened, club shaped (in optical section) adjacent to 

the apertures; the thickened part may extend part way 

poleward along the inner margins of the platea. Endexine appears to separate into additional lamellae at the 

apertures.

Stratigraphic distribution. Reklaw Formation.
20 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

Genus OCULOPOLLIS Pflug, 1953

Type species Oculopollis concentus Pflug, 1953, 

Palaeontographica, v. 95, ser. B, p. 110, pi. 19, figs. 

45-49.

Diagnosis. ("New wording" from Goczan and 

others, 1967.) Central part of pollen grains shallow lens 

shaped, one hemisphere with a concave and one with a 

convex surface. The three equatorial germinals located 

at the angles possess typical, sharply limited oculi in 

both hemispheres, these oculi strongly influence the 

shape and contour of the grains, although more so on one 

hemisphere than on the other. Equatorial contour 

generally is triangular convex to circular, mostly with interrupted outline. Surface smooth to strongly sculptured, wall sometimes also strongly structured. Exogerminals strong, monannulate, commonly with structure. 

Exopores in the form of vertical slits, mostly 

asymmetrical to the equator, pore canal opening outward, strongly constricted toward the inner part. 

Vestibulum mostly shallow slit shaped; sometimes with 

interloculum. Endogerminals with only weak 

thickenings of both outer lamellae of the endexine; endopores small, forming shallow atria with concave 

borders. The innermost wall lamella forms a wide atrium 

at the germinal. Small, mostly medium sized, seldom 

large forms.

Remarks. The diagnosis of the genus Oculopollis is 

included here even though specimens of Oculopollis were 

not observed, because the diagnosis of Pseudoculopollis 

(Goczan and others, 1967) refers directly to the diagnosis 

of Oculopollis. Specimens of Pseudoculopollis were 

found in the Mississippi embayment samples.

The Oculopollis complex includes the genera 

Bohemiapollis, Oculopollis, Pseudoculopollis and 

Semioculopollis. Goczan and others (1967) suggested 

that the genera Oculopollis, Bohemiapollis, and 

Pseudoculopollis could be combined into one genus with 

three subgenera and that Semioculopollis species represent a transition group between Oculopollis and 

Trudopollis.

Genus PSEUDOCULOPOLLIS Goczan, Kr. and Paclt., 1967

Type species. Pseudoculopollis principalis (Weyl. 

and Krg.) Krutzsch, 1967 (in Goczan and others, 1967), 

Palaontologische Abhandlungen, ser. B, v. 2, no. 3, p. 

495, pi. 14, figs. 7-13.

Diagnosis. (From Goczan and others, 1967). As in 

Oculopollis, but the swellings of the three oculi join over 

the pole in one hemisphere.

Pseudoculopollis admirabllis, n. sp.

Plate 11, figure 9-18; text figure 17

Holotype. Sample D3420, slide 5, coordinates 

81.5X9.8, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Paratype. Sample D3420, slide 1, coordinates

102.3X14.4, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Diagnosis. Shape of pollen grains circular in polar 

view with three equally spaced strongly protruding germinals with well-developed oculi. Extensions of ocular 

swellings extend over one pole, oculi not present or 

poorly developed on other hemisphere. Size 27^-32^ in 

equatorial diameter (36 specimens). Surface granulate to 

finely rugulate. Wall consisting of two principal layers, 

about 1.5^-2^ thick in the interapertural areas; ektexine 

about half as thick as endexine. Interloculum absent or 

very narrow. Exogerminals in the form of vertical slits, 

slightly subequatorial, slit extending farther toward the 

pole on the hemisphere bearing fused oculi. Prominent 

annulate thickenings of the ektexine present adjacent to 

the exogerminals; annulate area structureless. A prominent, though somewhat narrow, vestibulum is present 

between the endexine and ektexine in the germinal 

areas. Endexine with labra or small annuli adjacent to 

the mesogerminal pore. Mesopores small, somewhat circular and about 4^ in diameter. An interior lamella of 

the endexine forms a shallow submesoporal atrium.

Remarks. Pseudoculopollis admirabilis can be distinguished from P. principalis by the finer, less strong 

rugulate sculpture and by the much better development 

of the fused oculi on one hemisphere. The species P. concentricoides is smooth.

Stratigraphic distribution. McNairy Sand 

Member, Ripley Formation.

Pseudoculopollis sp. A

Plate 12, figures 1-6; text figure 18

Remarks. Only one specimen of this species was 

observed. It differs from other species of 

Pseudoculopollis by the presence of arci extending from

B

FIGURE 17. Pseudoculopollis admirabilis, ca. X 2,000. A, Sectional 

view of aperture. B, Elongate slit-shaped exogerminal and oculus 

fused over the pole on one side of grain. C, Short slit-shaped exogerminal and absence of oculus on obverse side of grain. Surface sculpture shown on B and C.
SYSTEMATIC DESCRIPTIONS 21

B

FIGURE 18. Pseudoculopollis sp. A, ca. X 2,000. A, Sectional view of 

aperture. B, Oculus on one hemisphere. C, Absence of oculus in 

obverse hemisphere. Arci present in this hemisphere only.

the oculi over one hemisphere to adjacent oculi. The 

specimen observed is 86/1 in equatorial diameter. A very 

small submesoporal atrium was observed. I consider one 

specimen inadequate for the erection of a new genus or 

species and am including a drawing and illustrations to 

demonstrate an additional variation in the Oculopollis 

complex.

Stratigraphic distribution. McNairy Sand Member 

of Ripley Formation.

Genus SEMIOCULOPOLLIS Gdczan, W. Kr. and Paclt., 1967

Type species. Semioculopollis minutus W. Kr. and 

Paclt., 1967 (in Goczan and others, 1967), Palaontologische Abhandlungen, ser. B, v. 2, no. 3, p. 503, pi. 

16, figs. 13-15.

Diagnosis. (From Goczan and others, 1967.) Pollen 

grains shallow lens shaped, equatorial contour weakly 

convex triangular, whereby the three equatorial germinals enter strongly in the contour. Wall of several 

layers, with two chief segments, occurrence of these distinctive, often the inner somewhat thicker than the outer. Surface smooth to finely sculptured. Germinal opening assembled somewhat as in Oculopollis, only the 

enlargement of the oculi is not typical, mostly somewhat 

strongly developed only on one side; the oculi can, 

however, be almost absent. Only small to medium-sized 

forms.

Semioculopollis sp. A

Plate 12, figures 7-13; text figure 19

Description. Pollen grains circular in polar view 

with prominent protruding oculi surrounding the germinal apertures. Oculi present on one hemisphere only 

and do not extend to the pole. Size 23/i-25/i in equatorial

B C

FIGURE 19. Semioculopollis sp. A., ca. X 2,000. A, Sectional view of 

aperture. B, Short slit-shaped exogerminal on one side of grain. C, 

Oculus and elongate slit-shaped exogerminal on obverse side of 

grain. Granulate surface sculpture shown on B and C.

diameter (three specimens), surface granulate. Wall two 

layered, ektexine about half the thickness of the endexine in interapertural areas; no interloculum. Exogerminals in the form of vertical slits, slightly subequatorial, slit extending farther toward pole on the 

hemisphere with oculi. Large annulate thickenings adjacent to exogerminals, about 4/1-5/1 high and 4/1-5/1 broad 

on each side of exogerminal, annulate area structureless. 

A prominent roomy vestibulum is present between the 

endexine and ektexine. Endexine with slight annuli adjacent to the mesopores. Mesopores small, 2/i-3/i in 

diameter. Interior lamella of endexine forms a shallow 

submesoporal atrium.

Remarks. This species appears to differ from other 

species of Semioculopollis by the presence of larger, 

more protruding oculi. Too few specimens were observed 

to justify the diagnosis and typification of a new species.

Stratigraphic distribution. Cusseta Sand Member 

of Ripley Formation and McNairy Sand Member of 

Ripley Formation.

Genus PSEUDOPLICAPOLLIS Krutzsch, 1967

Type species. Pseudoplicapollis palaeocaenicus 

Krutzsch, 1967 (in Goczan and others, 1967), Palaontologische Abhandlungen, ser. B, v. 2, no. 3, p. 496, pi. 

14, figs 26-31.

Diagnosis. (From Krutzsch, in Goczan and others, 

1967.) Pollen grains triangular shallow lens shaped, 

somewhat thickened in the region of the always present 

plicae. Equatorial contour triangular (with sometimes 

weakly interrupted sides) to triangular convex with 

slightly blunted angles. Wall thin, composed of two principal layers, with a ratio of about 1:1. Surface smooth to 

weakly microsculptured. With three equatorial germinals at the angles; exogerminals with annuli, inner 

with only weak structure, sometimes also appearing 

smooth but slightly thickened and not elongate. Exopores small, canal index much under 0.3, vertically
22 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

elongate and often weakly asymmetrical to the equator, 

opening outward. Sometimes with a roomy vestibulum. 

Endogerminals not thickened, with equatorial slitshaped opening; the plicae terminating internally with 

small points that is, the tips triangular to obtuse 

triangular in the endopores. No oculi, no interloculum, 

and so forth. Throughout small to sometimes mediumsized forms.

Remarks. Krutzsch (in Goczan and others, 1967) 

erected the genus Pseudoplicapollis to accommodate 

those pollen grains previously included in the genus 

Sporopollis. A new generic designation was required 

because the holotype of the genus Sporopollis is a 

gleichenoid spore and distinctly different from the pollen 

grains previously included in the genus. Krutzsch also 

transferred previously designated Sporopollis species to 

Pseudoplicapollis, including Pseudoplicapollis 

pseudosporites (Pflug) Krutzsch, n. comb., and P. 

peneserta (Pflug) Krutzsch, n. comb. He transferred 

Trudopollis proparuus Pflug, 1953, to Pseudoplicapollis 

proparuus (Pflug) Krutzsch, n. comb., and named one 

new species, Pseudoplicapollis palaeocaenicus, and 

designated it as the type species.

Pseudoplicapollis endocuspis, n. sp.

Plate 13, figures 1-7; text figure 20

Holotype. Sample D3265, slide 1, coordinates 

101.2X6.7, Eutaw Formation, Georgia, Coniacian.

Paratype. Sample D3265, slide 3, coordinates 

107.5X9.2, Eutaw Formation, Georgia, Coniacian.

Diagnosis. Shape of pollen grains slightly convex 

triangular with somewhat acute angles. Size 16^-22^ in 

equatorial diameter, (11 specimens) slightly inflated 

delicate plicae extend from each angle (germinal) over 

the poles to adjacent germinals. Wall two layered, thin, 

less than 1/u thick in interapertural areas, about 1.5/u 

thick at the apertures. Ektexine smooth, enlarged at 

apertures in the form of a slight labrum, two layered at 

aperture, outer layer structureless, inner layer consisting 

of very short radially directed bacula. Exogerminals 

short, vertical slit shaped. Vestibulum narrow, inverted 

V-shaped. Endexine thin, provided with acute angled 

points or flaps in the germinal area as extensions of the 

plicae. Endogerminal elongate equatorially between the 

pointed endexinal extensions.

Remarks. In general appearance this species is 

similar to Pseudoplicapollis palaeocaenicus. The latter 

species, however has a finely punctate surface sculpture 

and coarser plicae.

Stratigraphic distribution. Eutaw Formation, 

Coffee Sand, Coon Creek Tongue of Ripley Formation, 

McNairy Sand Member of Ripley Formation, McNairy 

Sand, Clayton Formation, and Porters Creek Clay. 

Single grains were observed in the Wilcox and Tallahatta Formations, but these may have been redeposited.

Pseudoplicapollis serenus, n. sp. 

Plate 13, figures 8-12; text figure 21

Holotype. Sample D3412, slide 2, coordinates 

84.7X13.0, Coffee Sand, Tennessee, Campanian.

Paratype. Sample D3412, slide 4, coordinates 

87.7X6.5, Coffee Sand, Tennessee, Campanian.

Diagnosis. Shape of pollen grains slightly convex 

triangular with blunt or obtuse angles. Germinals 

slightly inflated, interrupting the equatorial contour. 

Size 13.5ju-19ju in equatorial diameter (17 specimens). 

Delicate, slightly inflated plicae extend from apertural 

regions over the poles. Wall two layered, about l/j. thick 

in interapertural areas and about 2n thick at apertures. 

Ektexine and endexine of about equal thickness. Ektexine smooth, inflated at apertures in the form of a 

thickened labrum or annulus; two layered near apertures; outer layer structureless, inner layer of short, 

radially directed bacula. Exogerminals short, vertical 

slit shaped. Endexine provided with pointed obtuse 

angled flaps in the germinal areas that are extensions of 

the endexinal plicae. Vestibulum prominent, in polar 

view appearing as two somewhat oval chambers, one on 

each side of the plica extension. Endogerminals wide, 

horizontal slit shaped, located between the pointed extremities of the flaps.

Remarks. This species is distinguishable from other 

species of Pseudoplicapollis by its obtuse, blunt germinal regions and by the double oval shape of its 

vestibulum. It is distinguished from Minorpollis by the 

presence of distinct plicae.

Stratigraphic distribution. Coffee Sand.

Pseudoplicapollis sp. A

Plate 13, figures 13-17; text figure 22 

Description. Pollen grains triangular in polar view

FIGURE 20. Pseudoplicapollis endocuspis, ca. X 2,000. FIGURE 21. Pseudoplicapollis serenus, ca. X 2,000.
SYSTEMATIC DESCRIPTIONS 23

FIGURE 22. Pseudoplicapollis sp. A., ca. X 2,000. A, Sectional view 

of aperture showing scattered robust bacula, and endogerminal. B, 

Surface view showing delicate plica and microverrucate surface 

sculpture.

with almost straight sides. Size 25jU-36jLi in equatorial 

diameter (eight specimens). Very slightly inflated 

delicate narrow plicae extend from each angle over the 

poles. Wall two layered, about Ijii thick in interapertural 

areas, 2ju to 3ju thick at the apertures. Surface 

microverrucate to verrucate with low relief. Ektexine inflated at apertures to form a slight labrum; outer layer 

thin, structureless, inner layer of short robust loosely 

arranged bacula. Exogerminals equatorial, vertical slit 

shaped. Vestibulum narrow, poorly defined. Endexine 

thin, with somewhat rounded slightly elevated terminations in the vestibulum as extensions of the plicae. 

Endogerminals elongate equatorially with irregular 

margins.

Remarks. This species can be distinguished from 

other species of Pseudoplicapollis by its larger size. Too 

few specimens were found to allow for naming this taxon.

Stratigraphic distribution. Nanafalia and Wilcox 

Formations.

Genus PSEUDOVACUOPOLLIS Krutzsch and Pacltova, 1967

Type species. Pseudovacuopollis intraconcauus 

Krutzsch (in Goczan and others, 1967), Palaontologische 

Abhandlungen, ser. B, v. 2, no. 3, p. 499, pi. 15, figs. 1-5.

Diagnosis. (From Krutzsch and Pacltova, in Goczan and others, 1967.) Pollen grains triangular shallow 

lens shaped to triangular concave, with a tendency to a 

constriction of the equatorial contour in the interapertural areas. Germinals equatorial at the angles. Wall of 

several layers, variously thick, the proportion between 

the several layers varies among the different 

formspecies. The outer principal wall layer [ektexine] 

sometimes has an inner thickening in the interapertural

areas. Surface smooth to weakly and shallowly 

verrucate; sometimes with columella layer as an inner 

limit to the outer wall complex. With or without an interloculum. Inner wall complex possessing several 

smooth lamellae; often very thin. Exogerminals with 

structured annuli; monannulate or polyannulate. Exopores vertical slit shaped to oval. Endogerminals with 

several stepped, sometimes deeply sunken, atria, the 

almost platealike solution areas sometimes reach near 

the polar regions. Irregular endannuli or exfoliation of 

the lamellae sometimes weakly developed at the sides of 

the atria. Inner wall layers only seldom clearly 

developed, most often absent. No plicae, oculi, arci, and 

so forth. Forms mostly small to medium sized.

Remarks. The characteristics of this genus are very 

similar to those of Vacuopollis; indeed, Krutzsch and 

Pacltova (in Goczan and others, 1967) suggested that 

both could rank as subformgenera of the same genus. 

They distinguish the two genera principally by the 

tendency toward development of a constriction in the interapertural areas and a greater differentiation of the 

endexine in Pseudovacuopollis.

Pseudovacuopollis involutus, n. sp.

Plate 14, figures 1-12; text figure 23

Holotype. Sample D3416, slide 2, coordinates 

92.9X6.4, McNairy Sand, Missouri, Maestrichtian.

Paratype. Sample D3416, slide 2, coordinates 

104.7X10.7, McNairy Sand, Missouri, Maestrichtian.

Diagnosis. Shape of pollen grains slightly concave 

triangular to extreme concave triangular in polar view; a 

constriction in the interapertural areas commonly present. Size 15ju-23ju in equatorial diameter (17 

specimens). Wall of two principal layers, lij.-l.5n thick 

in the interapertural areas. Surface smooth. Ektexine in 

interapertural areas sometimes conspicuously thickened 

internally, becoming less so at bases of germinal areas. 

Ektexine thickened and annulate, consisting of two 

layers adjacent to the vertical slit-shaped exogerminal; 

outer layer structureless, inner layer consisting of comFIGURE 23. Pseudovacuopollis involutus, ca. X 2,000. A, B, 

Sectional views of apertures showing baculate layer, V-shaped endogerminals, and interapertural ektexinal thickening. C, Surface
24 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

pact, radially directed bacula. Endexine thin, less than 

IM thick with weakly developed endannuli adjacent to 

the atria. Inner margins of atria V-shaped in polar view. 

Inner layers of endexine poorly developed, secondary 

atria only occasionally visible.

Remarks. This species differs from 

Pseudouacuopollis intraconcauus Krutzsch, 1967, and P. 

triangulatus (Ross) Krutzsch, 1967, by possessing a 

smooth surface and poorly developed accessory atria.

Stratigraphic distribution. Coffee Sand, Coon 

Creek Tongue of Ripley Formation, McNairy Sand 

Member of Ripley Formation, and McNairy Sand.

Genus THOMSONIPOLLIS Krutzsch, I960, non Elsik, 1968

Type species. Thomsonipolis magnificus (Thorns, 

and Pflug) Krutzsch, 1960a, Freiberger Forschungshefte, 

v. C86, p. 58. Holotype, Th. and Pf., 1953, pi. 9, fig. 112.

Diagnosis. (From Krutzsch, 1960a.) Pollen grains 

25,u-50;u in diameter possessing three germinals that are 

complex, nonprotruding, and equatorially or slightly 

subequatorially arranged. Equatorial contour round to 

convex triangular and (in uncrumpled condition) oval to 

spherical shaped. The polar axis sometimes can be considerably shorter than the equatorial axis producing a 

lens-shaped pollen grain. Wall always many layered with 

three lamella complexes. The outermost bears mostly a 

strong baculalike to mushroom-shaped sculpturing that 

only seldom reveals the underlying exolamella. The middle wall complex is developed as many layered, unstructured, hyaline smooth lamellae oppressae (primarily in 

the intergerminal areas) and as lamellae conspicuae (at 

the germinals). Innermost complex unstructured, thin, 

yet with a refractive index different from the middle 

layer, and thereby easily separable from it. No interlocula, only seldom are "Schichtfuge" developed 

between the three wall complexes (these as a result of 

corrosion?). Germinal opening: "Exopore" equatorial 

slit shaped, oval or circular, about 5n in diameter. 

"Mesopore" arises from the outer lamellae of the middle 

wall complex, these are reflexed inward, and undergo 

differentiation in the form of thickening, annulus formation, exfoliation, and so forth. The innermost lamelae of 

the middle wall complex are likewise variously strongly 

differentiated and form several roomy oval to circular 

"atria," just as do only with a regular boundary the 

third inner wall complex ( = "endopores" Atrium).

Thomsonipollis magnificus (Th. and Pf. 1953) Krutzsch, 1960a

Plate 14, figures 13-20

Synonymy. See Krutzsch, 1960a.

Diagnosis. ("New wording" from Krutzsch, 1960a.) 

A form species of Thomsonipollis with thick to very thick 

wall, inclusive of the structure layers of the extragerminal about 2^-4^ thick; considerable lamellae differentiation in the region of the middle wall complex, whereby 

often three lamella groups are recognizable, their 

thickenings, exfoliations, and so forth, swell toward the

germinals. Size 25^-50^, mostly 30/i-35)U.

Stratigraphic distribution. Naborton, Naheola, 

Nanafalia, Wilcox, Pendleton (Andersen, 1960), 

Hatchetigbee, and Reklaw Formations.

Thomsonipollis mag-nificoides Krutzsch, 1960a

Plate 15, figures 1-8

Diagnosis. (From Krutzsch, 1960a.) A form species 

of Thomsonipollis with a thinner and finer wall, inclusive of the structure layers of the extragerminals, only 

1.5)U-2)Lt thick. Innermost lamellae of the middle wall 

layer interrupted only in form of steps; exfoliation, 

thickening, and so forth, almost completely lacking. 

"Mesopore" relatively broad. Size 27)it-4(V, mostly

Remarks. At the extremes, Thomsonipollis 

magnificus and Thomsonipollis magnificoides can be 

distinguished readily. However, specimens with intermediate wall thicknesses are difficult to place in the 

appropriate species.

Stratigraphic distribution. Same as for Thomsonipollis magnificus.

Thomsonipollis aff. T. magnificoides 

Plate 15, figures 9-20; text figure 24

Description. Shape of pollen grains almost circular 

in polar view, inflated lens shaped, almost spherical. 

Size 16n~21n in equatorial diameter, average 22/i (15 

specimens). Wall of two principal layers about l.S/i thick 

in interapertural areas, ektexine and endexine of about 

equal thickness. Ektexine with coarsely clavate-baculate 

sculpture, clavae not tightly spaced, endexine hyaline, 

structureless. Triporate, pores equatorial or nearly so. 

Endexine (middle layer of Krutzsch) thinning adjacent 

to exopore, then the upper part of the endexine turns inward forming a short channel. Inner layer (of Krutzsch) 

visible only as a hyaline refractive line below the germinals.

Remarks. These specimens differ from Thomsonipollis magnificus by their smaller size, more delicate

FIGURE 24. Thomsonipollis aff. T. magnificoides, ca. X 2,000. A, 

Sectional view of aperture showing reflexed endexine and thin wall. 

B, Surface view showing circular exorgerminal, and scattered ektexinal bacula and clavae.
SYSTEMATIC DESCRIPTIONS 25

and thin walls, more widely spaced sculpturing, and less 

structured germinals. They differ from T. magnificoides 

principally by their smaller size and more widely spaced 

sculpturing. Even though no observed specimen was as 

large as the average size T. magnificoides and the bacula 

and clavae are more scattered, I am reluctant to erect a 

new species on the basis of these minor characteristics. 

Stratigraphic distribution. Owl Creek Formation, 

Clayton Formation, one specimen observed in a sample 

from the Porters Creek Clay. The Stratigraphic range of 

this form is significantly different from that of Thomsonipollis magnificoides.

Thomsonipollis expolitus, n. sp. 

Plate 16, figures 1-12; text figure 25

Holotype. Sample D3410-C, slide 4, coordinates 

74.5X11.5, Owl Creek Formation, Mississippi, 

Maestrichtian.

Paratype. Sample D3410-C, slide 1, coordinates 

104.6X21.4, Owl Creek Formation, Mississippi, 

Maestrichtian.

Diagnosis. Pollen grains almost circular in polar 

view; germinals usually not protruding. Size 16/u-24/u in 

equatorial diameter (18 specimens). Wall of two principal layers about 1.5/u thick. Ektexine and endexine of 

about equal thickness. Surface smooth to weakly 

microscabrate. Exogerminals circular pores. Endexine 

hyaline, structureless, thinning adjacent to exopore, 

then the upper part of endexine turns inward forming a 

prominent annulus with somewhat irregular inner 

margin. Endogerminal of approximately the same size as 

exogerminal. Inner wall layer (of Krutzsch) visible only 

as a refractive line below the germinals.

Remarks. This species is somewhat similar to 

Thomsonipollis gracilis Krutzsch and Lenk, 1967 (in 

Goczan and others, 1967), except that the latter has a 

much more flexible wall and less pronounced infolding of 

the endexine adjacent to the germinal openings. 

(Holotype: plate 4 explanation, figs. 13a, b, in Kunert 

and Lenk, 1964, p. 412).

Stratigraphic distribution. Owl Creek Formation.

B

FIGURE 25. Thomsonipollis expolitus, ca. X 2,OOU. A, B, Sectional 

views of apertures showing the relatively thick wall and the reflexed 

inward enlargement of the endexine at the apertures. C, Surface

Genus TRUDOPOLLIS Pflug, 1953, emend. Krutzsch, 1967

Extratriporopollenites pertrudens Pf., 1953 (in Thorns, and Pflug, 

1953) p. 74, pi. 6, fig. 74 (and other figures).

Trudopollis pertrudens (Pf.) Pflug, 1953, p. 98, pi. 22, figs. 12, 18.

Trudopollis pertrudens (Pf.) Pflug, 1953, Lectotype designated by 

Krutzsch, 1967 (in Goczan and others, 1967), same specimen as in

Thorns, and Pf., 1953, pi. 6, fig. 74.

Type species. Trudopollis pertrudens (Pflug) Pflug, 

1953.

Diagnosis. (From Krutzsch, in Goczan and others, 

1967.) Pollen grains shallow to oval lens shaped; 

equatorial contour triangular to convex triangular or circular. Wall several layered, always two principal layers 

recognizable, often with "Schichtfuge" or interloculum. 

Surface always more or less clearly fine sculptured. Wall 

also sometimes with columella layer or other structure. 

With three equatorial germinals located at the angles. 

Exogerminals with tumescens, annulus, or similar 

thickening that sometimes strongly affects the contour; 

the inner part almost always with structure (radial 

baculate, punctate, and so forth), structured part either 

free or compact and terminates in a variously large 

vestibulum, not covered by a wall layer. Pore canal index 

less than 0.3. Outer part of exopores vertical slit shaped, 

opening nearly symmetrical with the equator. Endopores 

sometimes with weak thickenings around the atrium; 

atria of different shapes and variously deep.

No solution channels over the poles; sometimes also 

the endopores lie asymmetrical to the equatorial oculi 

that occasionally are present as attachments (accessory 

developments).

Remarks. Krutzsch (in Goczan and others, 1967) 

emended the genus Trudopollis to exclude those species 

with an endexinal solution channel over the poles (those 

species with platea) and possessing circular or oval exogerminals (rather than vertical slit-shaped exogerminals). The excluded species were placed in the genus 

Pompeckjoidaepollenites.

Trudopollis variabilis, n. sp.

Plate 16, figures 13-22; text figure 26

Holotype. Sample D3412, slide 2, coordinates 

109.7X14.1, Coffee Sand, Tennessee, Campanian.

Paratype. Sample D3412, slide 2, coordinates 

99.5X14.8, Coffee Sand, Tennessee, Campanian.

Diagnosis. Shape of pollen grains triangular to convex triangular or circular with protruding equatorial germinals. Size 22/u-29/u in equatorial diameter (20 

specimens). Wall two layered, about 1.5/u thick, no interloculum observed; ektexine and endexine of about 

equal thickness. Ektexine verrucate usually with low 

relief; in some specimens verrucae larger in the apertural 

regions. In apertural areas, ektexine greatly thickened, 

annulate, indistinctly layered, the outer layer structureless, the inner with diffuse structure consisting of
26 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

FIGURE 26. Trudopollis variabilis, ca. X 2,000. A, Sectional view of 

aperture. B, Surface view showing verrucae. C, Surface view showing 

more robust verrucae present on some specimens.

tightly packed radially directed rodlets or bacula whose 

free ends are usually clearly visible in the vestibulum. 

Vestibulum shallow. Exogerminal vertical slit shaped. 

Mesogerminal a more or less circular pore. Endexine in 

pore area with labra or endannulus. Inner layer of endexine forms a shallow submesoporal atrium.

Remarks. Pollen grains of this species are somewhat 

similar to those described as Trudopollis acinosus by 

Gray and Groot (1966), except that in their specimens 

they observed a distinct interloculum and an ektexine 

about twice as thick as the endexine. Furthermore, they 

reported no endannulus.

Pollen grains of Trudopollis meekeri Newman (1965) 

also bear some similarity to T. variabilis. However, in T. 

meekeri an interloculum is present, and the endexine is 

distinctly thinner than the ektexine. In T. variabilis an 

interloculum is lacking, and the ektexine and endexine 

are of about the same thickness.

Specimens of Trudopollis from the Mississippi embayment are difficult to compare with European forms for 

the following reasons: About 80 species have been 

reported; many of the early descriptions are inadequate; 

the photographs do not show clearly the morphological 

details required for species identification. None of the 

species described by Pflug (1953) and Thomson and 

Pflug (1953) possess a verrucate surface. However, some 

of the species described as having a fossulate surface 

sculpture may be irregularly verrucate.

Stratigraphic distribution. Cusseta Sand Member 

of Ripley Formation, Coffee Sand, Coon Creek Tongue of 

Ripley Formation.

Trudopollis plenus, n. sp.

Plate 17, figures 1-9; text figure 27

Holotype. Sample D3138, slide 4, coordinates 

113.6X10.0, Naheola Formation, Alabama, Paleocene.

Paratype. Sample D3138, slide 1, coordinates 

91.3X16.7, Naheola Formation, Alabama, Paleocene.

Diagnosis. Shape of pollen grains convex triangular

C

FIGURE 27. Trudopollis plenus, ca. X 2,000. A, Sectional view of 

aperture showing the endogerminal and the baculate layer of exogerminal region extending into the well-developed interloculum. B, C, 

Surface views.

to almost circular in polar view, with only slightly 

protruding equatorial germinal areas. Size 22/u-31/u in 

equatorial diameter (based on 21 specimens). Wall of 

two principal layers 1.5/u thick in the interapertural 

areas. Ektexine and endexine of about equal thickness 

and separated by a prominent interloculum. Ektexine 

two layered, the outer surface smooth and structureless, 

the inner composed of short bacula which extend across 

the interloculum. Baculate layer provides the 

appearance of a scabrate or microverrucate surface, 

although the outer surface is smooth. Faint plicae visible 

on some specimens. Ektexine thickened, annulate near 

apertures; bacula much elongated in this area, tightly 

packed and projecting radially with somewhat free ends 

in the vestibulum. Exogerminals consist of short, 

meridionally directed equatorial or slightly subequatorial slits. Endexine with a labrum adjacent to the 

mesogerminal. Mesogerminal somewhat circular, larger 

than exogerminal; vestibulum very narrow. Inner 

lamella of endexine separates near mesogerminal and 

forms an atriate endogerminal whose margins are 

irregular and appear granulate.

Remarks. Some specimens have a torn or somewhat 

eroded incidence or solution notch, but most specimens 

do not show this.

Stratigraphic distribution. Naborton and Naheola 

Formations.

Trudopollis of. T. artifex Weyl. and Krg., 1953

Plate 17, figures 18-20; text figure 28

Diagnosis. (From Weyland and Krieger, 1953.) 

About 25/u. Contour rigid triangular. Free lamella complex strong. Weak oculus. Cubiculum with nearly circular cross section, almost fully closed to the annulus. 

Strong endexine. The clear annulus intimus is here 

characteristic. No solution meridian. Intrabaculate
SYSTEMATIC DESCRIPTIONS 27

B

FIGURE 28. Trudopollis cf. T. artifex, ca. X 2,000. A, Sectional view 

of aperture showing short bacula, radial structure of ektexine, interloculum, and endogerminal. B, Surface view showing sculpture.

structure in the ektexine.

Description. (Mississippi embayment specimens) 

Shape of pollen grains rigid triangular in polar view with 

rounded apertural areas. Size 24^-28^ in equatorial 

diameter (three specimens). Wall of two principal layers, 

separated by a prominent interloculum; wall 3jU-4M 

thick; ektexine and endexine of about equal thickness. 

Surface irregularly fossulate with low relief. Ektexine 

thinner in apertural areas than in interapertural areas; 

with prominent radial structure. Exogerminal 

meridionally slit shaped, the lips not tightly appressed. 

Exogerminal opening into a large cubiculum; short 

bacula project inward throughout roomy cubiculum. 

Endexine with definite labra adjacent to mesogerminal. 

Inner lamella of endexine forms a short V-shaped notch. 

Atria not prominent if present.

Remarks. Trudopollis artifex differs slightly from 

the Mississippi embayment specimens by possessing a 

slight oculus and a tightly closed cubiculum. However, 

our specimens seem to conform otherwise to the circumscription and illustration of Trudopollis artifex.

Stratigraphic distribution. Coon Creek Tongue of 

Ripley Formation.

Trudopollis sp. A

Plate 17, figures 10-12; text figure 29

Description. Shape of pollen grains triangular to 

slightly concave triangular in polar view. Size 22^-23^ in 

equatorial diameter (two specimens). Surface smooth. 

Wall consists of two principal layers, about 1.5/u thick in 

the interapertural areas. Endexine about twice as thick 

as ektexine except in apertural areas. No interloculum. 

Ektexine in apertural areas two layered, the outer layer 

structureless and externally smooth, the inner consisting 

of very tightly packed radially directed bacula whose 

free ends are visible in the prominent vestibulum. Exogerminals equatorial, meridionally slit shaped. EndexFIGURE 29. Trudopollis sp. A, ca. X 2,000. A, Sectional view of 

aperture. B, C, Surface views.

ine with slight labrum adjacent to the mesopore, 

mesopore somewhat circular, larger than exogerminal; 

inner lamella of endexine forming a very narrow submesogerminal atrium.

Remarks. Somewhat similar to Trudopollis rector 

Pflug, 1953, but without interloculum, and to 

Trudopollis nonperfectus (Pflug, in Thorns, and Pf., 

1953) but with apparently thicker endexine.

Stratigraphic destribution. Coffee Sand.

Trudopollis sp. B

Plate 17, figures 13-14; text figure 30

Description. Shape of pollen grains convex 

triangular to triangular in polar view. Size 25^-29^ in 

equatorial diameter (four specimens). Wall of two principal layers separated by a very narrow "Schichtfuge", 

wall 1/U-1.5M thick in interapertural areas. Endexine and 

ektexine of about equal thickness. Surface scabrate to 

microverrucate with low relief. Exogerminals equatorial, 

meridionally slit shaped. Ektexine in apertural areas 

enlarged, annulate, apparently structureless; annulus

B

FIGURE 30. Trudopollis sp. B, ca. X 2,000. A, Sectional view of 

aperture showing apparently structureless ektexinal annulus, narrow 

interloculum, and slightly enlarged endexine at margin of the endogerminal. B, Surface view showing scabrate-microverrucate sculpture.
28 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

tear shaped with rounded contour in the roof of the 

vestibulum. Vestibulurn prominent. Endexine with 

labra adjacent to the mesogerminal. Inner lamella of 

endexine does not form a recognizable inner atrium. A 

semicircular notch present below the mesogerminal.

Remarks. Trudopollis sp. B can be distinguished 

from T. uariabilis by its slightly thinner wall, the 

teardrop-shaped structureless annulus in contrast to the 

larger, more angular annulus of T. variabilis and by the 

much finer surface sculpture.

Stmtigraphic distribution. Naborton Formation.

Trudopollis sp. C 

Plate 17, figures 15-17; text figure 31

Description. Shape of pollen grains triangular to 

slightly convex triangular in polar view; germinal areas 

rounded and slightly protruding. Size 3lu in equatorial 

diameter (one specimen). Wall of two principal layers 

separated by a narrow "Schichtfuge." Wall about 2ju 

thick in interapertural areas. Endexine slightly thicker 

than ektexine. Surface scabrate to densely 

microverrucate with low relief. Exogerminals equatorial, 

meridionally narrow slit shaped. Ektexine in apertural 

areas enlarged, annulate, structureless except for innermost part. Inner part of annulus studded with short 

pointed bacula whose free ends project into the 

vestibulum. Endexine with slight labra adjacent to the 

mesogerminal. Endexine of two distinct lamellae. No obvious atrium observed. A semicircular notch is present 

below the mesogerminal.

Remarks: Very similar to sp. B except for its larger 

size; has more robust annuli, more dense and smaller 

verrucae, and a recognizable definite two-layered endexine.

Stratigraphic distribution. Pendleton Formation 

(Andersen, 1960).

B

FIGURE 31. Trudopollis sp. C, ca. X 2,000. A, Sectional view ot 

aperture showing inner part of ektexinal annulus with short bacula, 

narrow interloculum, and the two lamellae of the endexine. B, Surface view showing sculpture.

Genus VACUOPOLLIS Pflug, 1953

Type species. Vacuopollis percentus Pf., 1953, 

Palaeontographica, ser. B., v. 95, nos. 4-6, p. 104, pi. 20, 

figs. 24-26.

Diagnosis. ("New wording" from Goczan and 

others, 1967.) Pollen grains triangular shallow lens 

shaped, seldom triplanoid. Equatorial contour extends 

from weakly convex triangular with slightly interrupted 

or uninterrupted sides to strongly triangular to weakly 

triangular concave. Corners sometimes strongly 

rounded. Wall with two principal layers, thickness of the 

entire wall and thickness of the individual wall segments 

very different. Surface smooth to weakly microsculptured. Mostly without structure in the extragerminal 

region. With three equatorial germinals located at the 

angles. Exogerminals with annuli, sometime labroid, 

seldom strongly thickened, often with prominent 

baculate structure layer (-conclave pro parte, according 

to Pflug, 1953); exopore vertical slit shaped terminating 

outward, more or less symmetrical with the equator. 

Pore canal index less than 0.3. Mostly with a wide 

vestibulum. Endogerminal not thickened, at the pore 

endexine somewhat split into lamellae; therefore, two to 

three, seldom more, stepped atria in succession and with 

shallow concave, semicircular or seldom deeper margins. 

No oculi; seldom an interloculum indicated. No plicae or 

arci. Forms small to medium sized.

Vacuopollis munitus, n. sp.

Plate 18, figures 1-19; text figure 32

Holotype. Sample D3416, slide 1, coordinates 

74.1X11.8, McNairy Sand, Missouri, Maestrichtian.

Paratype. Sample D3416, slide 1 coordinates 

82.7X18.7, McNairy Sand, Missouri, Maestrichtian.

Diagnosis. Shape of pollen triangular to slightly

B

FIGURE 32. Vacuopollis munitus, ca. X 2,000. A, Sectional view 01 

aperture, showing baculate layer, internal verrucae in atrium, and 

two-layered endexine. B, Surface view showing shallow fossulate 

sculpture.
SYSTEMATIC DESCRIPTIONS 29

concave triangular in polar view. Apertural areas only 

slightly bulging at the angles. Size 23/u-28/u in equatorial 

diameter (19 specimens). Wall of two principal layers, 

2/u-3/u thick in the interapertural areas. Surface 

shallowly fossulate. Ektexine distinctly thicker than 

endexine, on some specimens two to three times as thick. 

Exogerminal vertical slit shaped, equatorial; the ektexine thickened (annulate) adjacent to the pore canal and 

consisting of two layers an outer structureless area 

and an inner layer of tightly packed radially directed 

bacula. Baculate layer not visible on most specimens. 

Endogerminal very wide, extending almost entirely 

across base of apertural area. Endexine thin, consisting 

of two distinct layers broken up at the endogerminal to 

form one or more atria with somewhat irregular margins. 

Endexine in atria broken up into somewhat prominent 

isolated internal verrucae.

Remarks. This species of Vacuopollis is somewhat 

similar to V. orthopyramis except that it has a distinctly 

thicker ektexine.

Stratigraphic Distribution. McNairy Sand Member 

of Ripley Formation, McNairy Sand.

Genus OSCULAPOLLIS, n. gen.

Type species. Osculapollis aequalis, n. sp.

Diagnosis. Shape of pollen grains triangular to concave triangular, sometimes slightly convex triangular in 

polar view. Wall variously thick, endexine always thin. 

"Schichtfuge" or interloculum present. Surface smooth 

to slightly sculptured. Exogerminals in the form of small 

circular pores surrounded by annulate ektexine; ektexine 

in germinal areas two layered, outer layer structureless, 

inner of radially directed bacula. Narrow vestibulum 

present. Endogerminals somewhat circular, larger than 

exogerminals, endexine not enlarged adjacent to endogerminals.

Remarks. The presence of porate exogerminals in 

Osculapollis distinguishes it from Trudopollis and 

Vacuopollis.

Osculapollis aequalis, n. gen., n. sp.

Plate 18, figures 10-20; text figure 33

Holotype. Sample D3412, slide 4, coordinates 

102.6X3.2, Coffee Sand, Tennessee, Campanian.

Paratype. Sample D3412, slide 3, coordinates 

98.5X2.5, Coffee Sand, Tennessee, Campanian.

Diagnosis. Shape of pollen grains triangular, mostly 

concave triangular in polar view, with blunt apertural 

regions at the angles. Size 20/u-25/u in equatorial 

diameter (12 specimens). Wall about 1.5/u thick in interapertural areas, ektexine and endexine of about equal 

thickness and separated by a narrow interloculum. Surface smooth. Faint plicae or plicalike folds sometimes 

discernible. Exogerminals small circular equatorial 

pores surrounded by an annulate ektexine. Ektexine in 

germinal areas two layered, outer layer structureless and 

of uniform thickness over entire surface of grains, inner

FIGURE 33. Osculapollis aequalis, ca. X 2,000. A, B, Sectional views 

of apertures showing baculate layer, narrow interloculum, and thin 

endexine. C, Surface view, psilate.

layer of radially directed bacula that become 

progressively shorter toward the root of the exogerminal; 

end view of bacula in neck of exogerminal shows a 

granular appearance. A narrow vestibulum present. Endogerminal somewhat circular, larger than exogerminal 

and with irregular margin. Endexine thin, not enlarged 

adjacent to endogerminal, sometimes split into two 

lamellae in germinal areas.

Stratigraphic distribution. Cusseta Sand Member 

of Ripley Formation and Coffee Sand.

Osculapollis perspectus, n. sp.

Plate 18, figures 21-27, text figure 34

Holotype. Sample D3412, slide 3, coordinates 

104.1X16.3, Coffee Sand, Tennessee, Campanian.

Paratype. Sample D3412, slide 3, coordinates 

105.7X13.7, Coffee Sand, Tennessee, Campanian.

Diagnosis. Shape of pollen grains triangular to convex triangular with apertures at the angles. Apertural 

areas only slightly protruding. Size 20/u-29/u in equatorial 

diameter; most grains 20/u-24/u in diameter (based on 13 

specimens). Wall 2/u-3/u thick in interapertural areas and 

smooth; ektexine about twice as thick as endexine,

FIGURE 34. Osculapollis perspectus, ca. X 2,000. A, B, Sectional 

views of germinals showing small semicircular baculate areas, thick 

ektexine, narrow interloculum, and thin endexine. C, Surface view, 

psilate.
30 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

separated from it by a narrow interloculum. Exogerminals in the form of small circular pores surrounded by 

an annulate ektexine. Ektexine in pore areas two 

layered; outer layer smooth, and slightly thicker at base 

of the exogerminal than at the tip; inner layer consisting 

of a small, somewhat semicircular area of radially 

directed bacula. Endexine thin, about O.Syu thick. A 

small, narrow vestibulum is present. Endogerminal 

larger than exogerminal, circular with distinctly 

irregular margins or lips.

Remarks. Osculapollis perspectus can be distinguished from 0. aequalis by its thicker ektexinal wall, 

its much smaller baculate areas, and the more acute 

angles at its germinals.

Stratigraphic distribution. Coffee Sand.

Osculapollis sp. A 

Plate 18, figures 28-31; text figure 35

Description. Pollen grains concave triangular in 

polar view with blunt angles. Size 29ju-34yu in equatorial 

diameter (two specimens). Wall about 2/u thick in interapertural areas, endexine thinner than ektexine, surface smooth. Surface has the appearance of being 

granulate because of the presence of short bacula in the 

interloculum which separates ektexine from endexine. 

Exogerminals equatorial, in the form of small circular 

pores in the center of blunt apertural regions. Ektexine 

in pore areas two layered, outer layer structureless, 

slightly thicker at pore margin than otherwise; inner 

layer of crowded short bacula that become progressively 

shorter toward the root of the germinals and merge with 

bacula in the interloculum. No vestibulum, atrium 

roomy. Endexine thin, terminating in a plane margin at 

the base of the atrium.

Stratigraphic distribution. Wilcox Formation.

FIGURE 35. Osculapollis sp. A., ca. X 2,000. A, B, Sectional views of 

apertures showing small equatorial exopores, baculate areas with 

bacula continuing into the interloculum, and thin endexine. C, Surface view, psilate.

Genus PRAECURSIPOLLIS, n. gen.

Type species. Praecursipollis plebius, n. sp.

Diagnosis. Pollen grains convex triangular in polar 

view with apertures at the angles. Size 16yu-21ju in 

equatorial diameter (20 specimens). Wall two layered, 

thin, ektexine about the same thickness as endexine, the 

combined layers about lyu thick. Surface smooth, occasionally wall wrinkled, but this is incidental. Exogerminals circular pores, bordered by labra or small annuli. 

Ektexine in vicinity of pores two layered, the outer layer 

structureless, the inner consisting of very short radially 

directed bacula (often difficult to see). A shallow 

vestibulum present, as well as an atrium formed by the 

splitting of the endexine into two distinct layers in the 

vicinity of the endopore. Endopores slightly larger than 

exopores.

Remarks. Distinguished from Neotriangulipollis by 

porate exogerminals and by two distinct endexinal 

lamellae adjacent to the endopores.

Praecursipollis plebius, n. gen., n. sp.

Plate 19, figures 1-10; text figure 36

Holotype. Sample D3265, slide 3, coordinates 

87.0X5.5, Eutaw Formation, Georgia, Coniacian.

Paratype. Sample D3265, slide 4, coordinates 

78.4X7.1, Eutaw Formation, Georgia, Coniacian.

Diagnosis. Monotypic. Same as for genus.

Stratigraphic distribution. Eutaw Formation.

FIGURE 36. Praecursipollis plebius, ca. X 2,000. A, B, Sectional 

views of apertures, showing pores, narrow baculate area, and split 

endexine forming two atria. C, Surface view, psilate.

Genus PSEUDATLANTOPOLLIS, n. gen.

Type species. Pseudatlantopollis simulatus, n. sp.

Diagnosis. Shape of pollen grains triangular, 

slightly concave triangular, to slightly convex triangular 

in polar view, with equatorial germinals at the angles. 

Wall of two principal layers, ektexine thicker than 

endexine in the interapertural areas. Ektexine tectate, 

reticulate to foveolate with prominent columella layer. 

Exogerminals in the form of circular pores. Endexine 

thin in interapertural areas, separated into three to four 

bands or layers at the apertures, thus forming three or 

more irregularly defined successive atria.
SYSTEMATIC DESCRIPTIONS 31

FIGURE 37. Pseudatlantopollis simulatus, ca. X 2,000. A, Sectional 

view of aperture showing tectate structure of ektexine and successive 

endexinal atria. B, Surface view showing foveolate-reticulate sculpture.

Pseudatlantopollis simulatus, n. gen., n. sp.

Plate 19, figures 11-28; text figure 37

Holotype. Sample D1967-B, slide 10, coordinates 

100.0X18.9, McNairy Sand Member, Ripley Formation, 

Tennessee, Maestrichtian.

Paratype. Sample D1967-B, slide 9, coordinates 

111.9X18.9, McNairy Sand Member, Ripley Formation, 

Tennessee, Maestrichtian.

Diagnosis. Shape of pollen grains triangular, more 

often slightly concave triangular, sometimes slightly 

convex triangular in polar view with equatorial germinals at the angles of the grains. Size 20ju-26ju in 

equatorial diameter (35 specimens). Wall of two principal layers, 3ju-4At thick in the interapertural areas. 

Ektexine thicker than endexine. Ektexine tectate with 

prominent reticulate sculpture; bacula supporting tecturn easily seen at the periphery of the grains. Lumina of 

reticulum of same size over entire surface of the grains. 

Exogerminals in the form of circular pores, ektexinal 

sculpture somewhat thinner near pores. Endexine thin in 

interapertural areas, and separated into three to four 

bands or layers at the apertures to form three or more 

irregularly defined successive atria. Depth of apertural 

region including atria 5^-7^-

Remarks. This taxon bears some resemblance to 

Proteacidites. In the latter genus, however the endexine 

s not separated at the germinals, but all the lamellae are 

iused to form an endexinal annulus. Specimens of 

Proteacidites sp. are found in some of the samples that 

yield Pseudatlantopollis simulatus. These specimens of 

Proteacidites also differ from Pseudatlantopollis by 

possessing a reticulum whose lumina in the interapertural areas are distinctly larger than those at the poles or 

near the apertures. Pseudatlantopollis simulatus also 

bears a close resemblance to Atlantopollis, specifically 

Atlantopollis reticulatus, except that Atlantopollis is 

characterized by a vertical slit-shaped exogerminal, and 

the exogerminal of the genus Pseudatlantopollis is a circular pore.

Stratigraphic distribution. Cusseta Sand Member 

of Ripley Formation, McNairy Sand Member of Ripley 

Formation, and Owl Creek Formation.

Genus ENDOINFUNDIBULAPOLLIS, n. gen.

Type species. Endoinfundibulapollis distinctus, n. 

sp.

Diagnosis. Pollen grains triangular to slightly convex triangular in polar view with the germinals at the 

angles. Size 20ju-25At in equatorial diameter (11 

specimens). Wall thin, two layered, combined thickness 

in interapertural areas about 1/u. Surface smooth or 

nearly so. Exogerminals short vertical slit shaped; ektexine adjacent to germinals labrate, two layered, the outer 

layer smooth and structureless, the inner layer of short, 

very tightly packed, radially directed bacula, baculate 

area tapering toward the root of the germinal. Structure 

in the inner layer often difficult to see. Vestibulum large, 

prominent double cone shaped. Endexine thin, reflexed 

inward in the form of a funnel at the apertures, endopore 

circular, 2ju-3M in diameter at the base of the reflexed 

endexine. No endannulus, no interloculum.

Endoinfundibulapollis distinctus, n. gen., n. sp. 

Plate 20, figures 1-11; text figure 38

Holotype. Sample D4268-B, slide 1, coordinates 

78.5X21.9, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Paratype. Sample D4268-B. slide 2, coordinates 

79.2X15.1, McNairy Sand Member, Ripley Formation, 

Kentucky, Maestrichtian.

Diagnosis. Monotypic, diagnosis as for genus.

Remarks. The reflexed endexine is somewhat 

similar to that observed in Longanulipollis, except that 

the endexine is much thinner. Furthermore, the structure of the ektexine in the region of the exogerminals is 

entirely different. One specimen of Endoinfundibulapollis distinctus was observed to have a serendipitous air bubble within one endogerminal (pi. 20, figs. 

1-5) that clearly demonstrates the size and shape of the 

endopore.

Stratigraphic distribution. Coffee Sand and 

McNairy Sand Member of Ripley Formation.

FIGURE 38. Endoifundibulapollis distinctus, ca. X 2,000. Apertures 

showing ektexinal vertical slit, baculate layer, and reflexed funnelshaped endexine and endopore.
32 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

NEW GENUS A 

Plate 20, figures 12-14, text figure 39

Description. Pollen rigid triangular to slightly concave triangular in polar view. Size 31/u in equatorial 

diameter (one specimen). Wall thin, two layered, combined thickness about 1/u. Ektexine fine verrucate to 

microverrucate. Germinals at the angles of the grain. Exogerminal a short vertical slit-shaped opening surrounded by a two-layered ektexine with an annulus. 

Outer layer structureless, inner layer of short radially 

directed bacula. Endexine forming a mesogerminal 

below a narrow vestibulum. A wide atriate endogerminal 

is developed centripetal to the mesogerminal. Base of 

atriate endogerminal marks an abrupt change in contour 

of the grains. Atrium roomy, 6/u-9/u deep. Centrifugal to 

the endogerminal, the endexine forms several internal 

partial annular ridges in the walls of the atrium.

Remarks. This genus is distinguished from 

Triangulipollis by the more prominent atrium that 

marks a change in the outer contour of the grain and by 

the presence of annular ridges in the neck of the germinal. It is distinguished from Santonipollis, a genus 

that it superficially resembles, by its much shorter, more 

compact baculate area and by the presence of a prominent mesogerminal near the outer part of its germinals.

Stratigraphic distribution. Eutaw Formation.

FIGURE 39. New genus A, ca. X 2,000. A, B, Sectional views of 

germinals showing baculate area, prominent mesogerminal, and 

deep atriate endogerminal. C, Surface view, verrucate.

NEW GENUS B

Plate 20, figures 15-17; text figure 40

Description. Pollen triangular shallow lens shaped, 

with slightly bulging angles (germinals). Size 18/u-21/u in 

equatorial diameter (three specimens). Wall thin, about 

In thick in interapertural areas; ektexine and endexine 

of about equal thickness; ektexine smooth. Exogerminal 

in the form of an elongate vertical slit. Aperture

bordered by a prominent annulus tapering toward the 

root of the germinal. Ektexine in apertural area two 

layered, the outer of uniform thickness and structureless, the inner consisting of radially projecting 

tightly packed bacula. Vestibulum prominent, roomy. 

Endexine thin, projecting inward below the exogerminals and the vestibulum forming a short, parallelwalled channel. Endogerminals at base of infolded 

endexine. Endexinal apertures small circular pores; 

margins of endexine adjacent to pore slightly thickened.

Remarks. Distinguished from Longanulipollis by 

the presence of a prominent baculate layer adjacent to 

the exogerminal and by a much thinner endexine. This 

genus is somewhat similar to Endoinfundibulapollis in 

the possession of a reflexed endexine; however, the exogerminal development is much more prominent in new 

genus B, and it possesses a slight endexinal thickening 

adjacent to the endopore.

Stratigraphic distribution. Coffee Sand.

FIGURE 40. New genus B, ca X 2,000. A, B, Sectional views showing 

slit-shaped exogerminals, baculate layer, and inward-projecting 

endexine forming channeled endopore. C, Endopore channel twisted, 

showing equatorial view of pore.

NEW GENUS C . 

Plate 20, figures 18-20; text figure 41

Description (Incomplete). Pollen grains triangular 

in polar view with large apertural regions at the angles. 

Size 28ju in equatorial diameter (one specimen). Surface 

smooth. Exogerminal vertical slit shaped, ektexine adjacent to exogerminals thin, inner part of wall studded 

with scattered, radially directed fairly coarse bacula, 

arranged somewhat in the form of successive annular 

ridges. Vestibulum prominent. Endexine thick, particularly in the apertural region. Endopores small surrounded by thick unstructured annuli. Endexine thicker 

than ektexine.

Remarks. I have been unable to determine with certainty whether what appear to be heavy endexinal arci 

are actual, or whether their purported appearance is 

caused by the irregular wrinkling of the specimen in the 

interapertural regions.

Stratigraphic distribution. Owl Creek Formation.
KEY TO MISSISSIPPI EMBAYMENT NORMAPOLLES GENERA 33

FIGURE 41. New genus C, ca. X 2,000. Sectional views of apertures 

with short bacula in outer part and prominent endexinal thickenings 

adjacent to the endopores.

NEW GENUS D

Plate 20, figures 21-23; text figure 42

Description. Pollen grains circular to convex 

triangular in polar view, with equally spaced equatorial 

apertures. Size 44;u in equatorial diameter (one specimen). Wall of two principal layers, about 2/j, thick in interapertural areas, ektexine and endexine of about equal 

thickness separated by a narrow interloculum. Surface 

smooth. Exogerminal in the form of a vertical narrow slit 

extending about half the distance to the poles. Ektexine

of uniform thickness except near the exogerminal, where 

it narrows abruptly. Endexine also narrows near the exogerminal, then thickens in the mesogerminal region, 

then thins again in the interapertural areas. Inner portion of endexine shows continuous exfoliation pattern 

within the large roomy atrium. Irregular short bacula 

form a somewhat annulate pattern more prominent in 

the middle part of atrium than above or below. Innermost layer of endexine is visible only as a lower limit to 

the atrium and at the sides as a thin, more refractive 

layer.

Remarks. This specimen is included in the Normapolles assemblage because of the vertical slit-shaped 

exogerminal and the complicated exfoliation pattern of 

the endexine present in the large atrium.

Stratigraphic distribution. Owl Creek Formation.

FIGURE 42. New genus D. One aperture, ca. X 2,000. Deep vertical 

slit, extexine, interloculum, and exfoliation pattern within the 

atrium shown.

KEY TO MISSISSIPPI EMBAYMENT NORMAPOLLES GENERA

A conspicuous feature of the pollen grains of many Normapolles genera is the 

presence of vertical slit-shaped exogerminals. Other Normapolles genera, in contrast, possess circular porate exogerminals. Pollen of some Normapolles genera or 

species may possess an exogerminal constructed of only a slit or vertically oval 

exogerminal. Those pollen grains, whose exogerminals might be misinterpreted 

because of small size or orientation of the specimens, have been keyed to both the 

slit-shaped and the porate categories below.

A. Exogerminals, 6; 3 subequatorial on each hemispheije

B. Wall thin, with an equatorial constriction, usually with interloculum _ _ _ _ Interpollis

BB. Wall thick, without equatorial constriction, no interloculum _____ Interporopollenites

AA. Exogerminals, 3; equatorial or nearly so 

C. Exogerminals vertical slit shaped

D. Endogerminals (mesogerminals) prominently equatorial slit shaped at right angle to

exogerminal opening

E. Typical polyannulate exogerminal with at least one prominent 

praevestibulum ____________________________ Basopollis

EE. No praevestibulum in exogerminal

F. Ektexine and endexine thin, both of about equal thickness.

G. Plicae prominent, terminating in a triangular point within 

vestibula _______________________ Pseudoplicapollis

GG. No typical plicae present ______________ Complexiopollis

FF. Ektexine significantly thicker than endexine _____ Choanopollenites

DD. Endogerminals somewhat circular to oval

H. Prominent oculi present on one hemisphere only

I. Oculi joined over one pole by a prominent external 

thickening __________________________ Pseudoculopolis

II. Oculi clearly isolated not connected over one pole _ _ Semioculopollis
34 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

HH. Prominent oculi not present

J. Large roomy atria or vestibula extending more than half the apertural 

radius (measured from the exterior of the exogerminals toward the 

pole to the inner limit of the atria or vestibula); openings of endogerminals much larger than those of exogerminals. 

K. Endogerminals with a deep V-shaped incidence or cuneus.

L. Ektexine conspicuously thickened and protruding adjacent to 

pore canal; endexine of only one distinct layer or lamella. 

___________________________ Extremipollis

LL. Ektexine usually not prominently thickened and protruding 

adjacent to the pore canal; endexine thin, consisting of two 

or more distinct layers at the endogerminals, forming 

thereby successive atria. _____________ Vacuopollis

KK. Endogerminal borders shallow crescent shaped 

_________________________ -Megatriopollis

JJ. Atria or vestibula much smaller, extending half or less the apertural

radius. 

M. Small forms, 16/u or less in diameter. ----------Minorpollis

MM. Larger forms, usually larger than 20^i in diameter. 

N. With a prominent vestibulum.

O. Diameter of endogerminal openings only slightly 

smaller than diameter of vestibulum; exogerminals labroid; no interloculum _ Pecakipollis 

OO. Diameter of endogerminal openings much less than 

diameter of vestibulum; exogerminals annulate, 

swollen; interloculum usually presentTrudopollis 

NN. No vestibulum, no interloculum, mesogerminal consisting of a basketlike filter. _______ Nudopollis

CC. Exogerminals circular pores, or at most slightly oval. 

P. Outline of pollen grains circular or nearly so.

Q. Platea and interloculum present -----------Pompeckjoidaepollenites

QQ. No development of platea.

R. Endexine reflexed inward around endopores ________ Thomsonipolis

RR. Endexine not reflexed inward, endexine of two prominent layers _ _ _

----------_____________ Kyandopollenites

PP. Outline of pollen grains triangular, concave triangular or convex triangular. 

S. With prominent plicae.

T. Plicae terminating in a triangular flap within the vestibulum _____

_______________________ -Pseudoplicapollis

TT. Plicae terminating at base of vestibulum, no pointed flap present _ _ _

-------_--_________________ -Plicapollis

SS. Without prominent plicae.

U. Platea and interloculum present ________ Pompeckjoidaepollenites

UU. No platea present.

V. Inner part of ektexine thickened or swollen in interapertural 

areas. ________________________ Pseudouacuopollis

VV. Ektexine almost the same thickness throughtout.

W. Interloculum present _______________ Osculapollis

WW. No interloculum.

X. Prominent reticulate tectum present __________

________________________ Pseudatlantopollis

XX. Without reticulate tectum.

Y. Endexine annulate or split into several lamellae

adjacent to endogerminal.

Z. Wall thin, vestibulum very shallow; baculate 

area of exogerminals poorly developed; 

successive atria formed by splitting of 

endexine ___________ Praecursipollis

ZZ. Wall thicker, vestibulum deep, roomy; 

baculate area of exogerminals well 

developed ___________ Pecakipollis

YY. Endexine not annulate nor split into several

lamellae adjacent to endogerminal. 

a. Endexine strongly reflexed inward below 

vestibulum, terminating in a small pore _ 

___________ --Endoinfundibulapollis

aa. Endexine not reflexed inward; small forms less 

than 16|U in diameter _____ -Minorpollis
STRATIGRAPHIC DISTRIBUTION OF NORMAPOLLES GENERA 35

STRATIGRAPHIC DISTRIBUTION OF 

NORMAPOLLES GENERA

Genera assignable to the Normapolles group are 

limited in Europe to the Late Cretaceous and early Tertiary. They first appear in the basal part of the Cenomanian, attain their greatest development during the Santonian and Maestrichtian Stages and gradually decrease 

in number during the early Tertiary. Only one genus, 

Plicapollis, persists beyond the Eocene into the early 

Oligocene, all others have become extinct by the end of 

Eocene time. A chart showing the stratigraphic distribution of the European Normapolles genera is provided by 

Goczan and others (1967, p. 540). Although a few Normapolles genera, notably Thomsonipollis, were known 

from the North American continent in 1967, data concerning these genera were largely omitted from that 

treatise.

Normapolles genera first appear in Mississippi embayment rocks during the Cenomanian. Complexiopollis 

is the lone Normapolles genus to appear in the 

Tuscaloosa Formation sample. In the Coniacian, four 

genera were recognized; one of these, however, new genus 

A, was represented by but a single specimen an insufficient number to permit assigning generic and specific 

epithets. The Campanian is represented by 15 genera 

and the Maestrichtian by 18 genera. A decline in 

numbers of genera begins in the Paleocene which is 

represented by only 8 genera. The lower Eocene Wilcox 

Group yielded specimens of 10 genera; by the middle 

Eocene (Claiborne time) this number had abruptly 

decreased to 1 (Nudopollis). The upper Eocene formations examined failed to yield any specimens 

assignable to Normapolles genera, although Fairchild 

and Elsik (1969) stated that they had seen several 

specimens of Nudopollis terminalis in the late Eocene 

Jackson Group.

In Europe, the greatest number of Normapolles genera 

is attained during the Santonian, followed by a second 

peak of abundance during the Maestrichtian and two 

minor peaks in the early Tertiary (Goczan and others, 

1967). In the Mississippi embayment the greatest 

number of genera was observed in the CampanianMaestrichtian interval. Thus, the development of Normapolles genera in the Mississippi embayment approximately parallels the development of those genera in 

Europe. As the sampling interval in the lower part of the 

Campanian and older stages was very sparse in the 

Mississippi embayment, it is pointless to try to compare 

in detail the European and Mississippi embayment 

maxima.

Fifty-seven genera are included in the European Normapolles complex by Goczan, Groot, Krutzsch, and 

Pacltova (1967). Forty-seven of these genera were from 

boreal middle Europe and 10 were confined to western 

Hungary with but four Normapolles genera common to

the two European regions. Only 18 of these 57 genera are 

recorded from Mississippi embayment samples. To this 

number must be added the two genera erected by Stover 

(in Stover and others, 1966) Kyandopollenites and 

Choanopollenites, and the genera established as a result 

of the present work, Osculapollis, Praecursipollis, 

Pseudatlantopollis, and Endoinfundibulapollis. This 

makes a total of 24 Normapolles genera from Mississippi 

embayment rocks, exclusive of the four possible new 

genera that were not named because of insufficient 

numbers of specimens.

Figure 43 compares the stratigraphic distribution of 

these genera in Europe and in the Mississippi embayment.

Of the genera common to Europe and the Mississippi 

embayment, several show remarkably similar 

stratigraphic ranges in the two areas, notably Complexiopollis, Trudopollis, Pseudoplicapollis, Nudopollis, 

Interpollis, and Thomsonipollis. The sample coverage 

from the early Campanian through the Cenomanian is 

poor in the Mississippi embayment. As more samples are 

examined from this interval, undoubtedly additional 

Normapolles genera will be identified and the 

stratigraphic ranges of others will be extended.

The ranges of several genera are distinctly different in 

Europe and in the Mississippi embayment. Plicapollis 

and Vacuopollis have ranges in Europe extending from 

the Turonian or Coniacian into the Eocene, and 

Plicapollis even into the basal Oligocene. In the 

Mississippi embayment their known ranges are limited 

to the Campanian and Maestrichtian. This discrepancy 

is difficult to explain. These genera apparently died out 

earlier in the Mississippi embayment than in Europe. A 

similar pattern is observed for the genus Minorpollis. 

However, this genus is considered by European workers 

as a collection genus for small pollen grains that may be 

difficult to diagnose. (See Goczan and others, 1967, p. 

478.) Consequently, several genera may be included in 

the European range of this form genus.

Genera such as Megatriopollis and Pecakipollis may 

be as rare in European rocks as they are in rocks from the 

Mississippi embayment; consequently, although, at 

present, their stratigraphic ranges do not coincide, 

further work may extend the ranges in one or both areas 

and so greater coincidence may be forthcoming.

Pompeckjoidaepollenites has a range in Europe extending from the Coniacian into the late Eocene. In 

Mississippi embayment samples specimens of this genus 

were observed only in the upper part of the lower Eocene. 

This apparent discrepancy may be due to the rarity of 

specimens of this genus in this area. This genus very 

clearly is derived from the Trudopollis stock. Parallel 

evolution taking place later in the Mississippi embayment than in Europe or later immigration from Europe 

could explain the later first appearance of the genus in 

the Mississippi embayment.
36 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

Z

2" w

o ^

1- LU

^ >-

ESTIMATED DUROF MILLIONS 

20

10

6

11

3

3

3

20

AGE

Eocene

Paleocene

EOUS

h

LLJ

0

Maestrichtian

Campanian

Santonian

Coniacian

Turonian

Cenomanian

Jackson Group

(V

E a.

'5 u

0

x a

O 3

0 0

§ 0

GENERA ESTABLISHED FROM EUROPEAN SAMPLES

Complexiopollis

7 

7 ?

F

Trudopollis

?

Pseudoplicapollis

7

11

Plicapollis

\m

Minorpollis

1

sl

lil

Pseudovacuopollis

rt

i

Pseudoculopollis

7

|

il

/A7 terporopollenites

m

:::: : : 

 I

i

Semioculopollis

i

1

Vacuopollis

7

CO

a

cPompeckjoidaepolle

7

Pecakipollis

--f

-iI EXPLANATION

i

i J Stratigraphic range in Europe

1

5jij Stratigraphic range in

iij Mississippi embay ment

il 1 Modified from Ku p (1961), Holmes (1965) a nd Gi and Cobban 1966).

FIGURE 43. Comparison of stratigraphic distribution of Normapolles
STRATIGRAPHIC DISTRIBUTION OF NORMAPOLLES SPECIES 37

GENERA ESTABLISHED 

FROM EUROPEAN SAMPLES

Megatriopollis

7

7 

7

pr

*L

Extremipollis

»

ft

Nudopollis

7

7

m

Interpollis

7

7

m

Basopollis

7

!

Thomsonipollis

i

I

EXCLUSIVELY MISSISSIPPI 

EMBAYMENT GENERA

Praecursipollis

m

New Agenus Osculapollis

1

1

topollis Pseuda tlan

1

Choanopollenites

Endoin fundibulapollis

1

1

00

D 

C 

tD 

O)

<D

2

1

CNew genus 

Q

3 

C 

CD 

D)

CD

z

Kyandopollenites

1

genera in Europe and Mississippi embayment.

The genera Thomsonipollis, Interpollis, and Extremipollis appear earlier in the stratigraphic column in 

the Mississippi embayment than in Europe. This fact 

would seem to contradict the suggestion of Goczan, 

Groot, Krutzsch, and Pacltova (1967) that central 

Europe is the center of development of the Normapolles 

group. Certainly, so far as genera are concerned, many 

more Normapolles genera are known from Europe than 

from the Mississippi embayment. However, certain 

genera, because of their rarity may not yet have been 

found in rocks as old in Europe as in North America. An 

alternative hypothesis is one that would allow for the 

origin of certain Normapolles genera in Europe and 

others in North America. It must be remembered that 

the Normapolles group is a group of convenience. No one 

has implied genetic relationship of all the genera included. Quite the contrary, it is evident on a 

morphologic basis that such genera as Interpollis and 

Nudopollis have little in common and probably 

originated from widely divergent plant stems. If the continents were at one time close enough to permit plant 

migration from east to west, there could as well have 

been migration from west to east. A further argument for 

independent origin in North America of genera now 

assigned to the Normapolles group is seen in the fact 

that the genera Choanopollenites, Kyandopollenites, 

Osculapollis, Praecursipollis, Pseudatlantopollis, Endoinfundibulapollis, and four new unnamed genera have 

been found in Mississippi embayment rocks. None of 

these genera has been reported from Europe.

STRATIGRAPHIC DISTRIBUTION OF 

NORMAPOLLES SPECIES

Normapolles species have proved to be excellent index 

fossils in Europe. They have the same stratigraphic 

utility in the Mississippi embayment. Individual 

specimens often can provide preliminary information 

concerning the stratigraphic placement of a sample. 

When used in conjunction with assemblages of other 

Normapolles species and other pollen and spore species, 

their stratigraphic value is enhanced remarkably.

Fifty-nine Normapolles species have been recognized 

from Mississippi embayment rocks. Of this number, 43 

species have been found in sufficient numbers for 

specific names to be assigned. The remainder are 

referred to by generic designation only. It is notable that 

the greater proportion of species recognized could not be 

equated with European species. The great proportion of 

new species suggests that the Mississippi embayment 

genera and species were at least somewhat isolated from 

their European counterparts.

Mississippi embayment species have been plotted in
38 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

 7

DURATION ESTIMATED 1

20

10

9

11

3

3

3

20

YEARS' |MILLION 

W->

0

LLJ

0

<rIII

u

AGE

Eocene

Maestrichtian

Campanian

Santonian

Coniacian

Turonian

Cenomanian

o

i

Q_ 

< W

c ^ 

o 5!

i= 2 

< <

STRATIGRAPHIC f ^ 

UNIT w 0

H 2

< 2

5 I-

§0 £^

D. 

Q_

<

Jackson (Group)

g Q White Bluff Formation

" Q Yazoo Clay /

Cockfield Formation -/

£ a Cook Mountain Formation

° £ Sparta Sand

£ 0 ZilphaClay

Reklaw Format.on

Hatchetigbee Formation

° o (Andersen, 1960) A,

5 0 W,lco« Formation /

Nanafaha Formation

Naheola Formation

Naborton Formation

Porters Creek Clay <

C,ayton Formation ^

Owl Creek Formation

| | McNairy Sand

1$ (

a g McNairy Sand Member x'

£ 0 I

Coffee Sand

CussetaSand Member 

of Ripley Formation

Eutaw Formation

Complexiopollis funiculus Complexiopollis patu/us Complexiopollis A sp. 

Complexiopollis abditus Praecursipollis plebius 

New Agenus 

1 1

GENERA

« « 1 1 i

! I I 111- slj i J< ! § « .|<^.Scc^ ££2-2 £ S E w *

SsIfifMJ I^il ll< i ^ilMii I ! ! ! IP 11 ! i i 1 1 1 111 * 1 1 T: 1 i 11! *

o.5:i-:oa.(/o-5 cjtjG'JjSa-i-i-COuji-zi-a.i-UjujCCL-^^

?

1

|

rr-T- i niil

I

I I 1 1 1 1 1 1 1

,;,

?

i 

1 Modified from Ku p (1961), Ho mes (1965) and G and Cobban 1966).

FIGURE 44 (above and lacing page). Stratigraphic ranges ot JNormapolles species in Mississippi embayment.
REFERENCES 39

GENERA

ffi

FIGURE 44. Continued.

stratigraphic sequence in figure 44. This figure portrays 

the sequential development of Normapolles species in 

Mississippi embayment rocks. Beginning in the 

Cenomanian three species were recognized, all belonging 

to the genus Complexiopollis. In succeeding younger 

samples, development continues to be expressed in the 

form of increased numbers of species. In the Coniacian, 7 

species were found; in the Campanian, 22; and in the 

Maestrichtian, 28. This species peak in the Maestrichtian is followed by a decline in the Paleocene to 16 

species, then to 12 species in the early Eocene, and to 

only 2 in the middle Eocene. In the Mississippi embayment rocks examined, no Normapolles genus or species 

originates in rocks younger than those of the upper part 

of the Wilcox. Concurrently, all Normapolles species apparently became extinct in North America by the end of 

middle Eocene time.

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DeVries, D. A., 1963, Jasper County mineral resources: Mississippi 

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Doyle, J. A., 1969, Cretaceous angiosperm pollen of the Atlantic 

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Drugg, W. S., 1967, Palynology of the upper Moreno Formation (Late 

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Fairchild, W. W., and Elsik, W. C., 1969, Characteristic palynomorphs 

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Foster, V. M., 1940, Lauderdale County mineral resources Geology: 

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v. 2, no. 3, p. 427-633.
40 NORMAPOLLES POLLEN FROM THE MISSISSIPPI EMBAYMENT

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Upper Cretaceous formations of Delaware and New Jersey: 

Palaeontographica, ser. B, v. 117, nos. 4-6, p. 114-134.

Groot, J. J., and Groot, C. R., 1962, Some plant microfossils from the 

Brightseat Formation (Paleocene) of Maryland: Palaeontographica, ser. B., v. Ill, p. 161-171.

Groot, J. J., Penny, J. S. and Groot, C. R., 1961, Plant microfossils and 

age of the Raritan, Tuscaloosa and Magothy Formations of the 

Eastern United States: Palaeontographica, ser. B., v. 108, p. 

121-140.

Holden, F. T., Hughes, U. B., Mellen, F. F., and Spooner, K. K., compilers, 1948, The stratigraphy and paleontology of the Eocene, 

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Field Trip, 1948, p. 52-74 (reprinted from Shreveport Geol. Soc. 

Guidebook, llth Ann. Field trip, 1934).

Holmes, Arthur, 1965, Principles of physical geology [revised ed.]: New 

York, Ronald Press Co., 1288 p.

Jones, E. L., 1962, Palynology of the Midway-Wilcox boundary in 

south-central Arkansas: Gulf Coast Assoc. Geol. Socs., Trans., v. 

12, p. 285-294.

Kimyai, Abbas, 1966, New Plant microfossils from the Raritan Formation (Cretaceous) in New Jersey: Micropaleontology, v. 12, no. 4, 

p. 461-476.

Kremp, G. O. W., 1965, Morphologic encyclopedia of palynology: 

Arizona Univ. Press, 185 p.

Krutzsch, Wilfried, 1954, Bemerkungen zur Benennung und 

Klassifikation fossiler (insbesondere tertiarer) Pollen und Sporen: 

Geologie, v. 3, no. 3, p. 258-311.

Krutzsch, Wilfried, 1959, Einige neue Formgattungen und -arten von 

Sporen und Pollen aus der Mitteleuropaischen Oberkreide und 

dem Tertian Palaeontographica, ser. B., v. 105, nos. 5-6, p. 

125-157.

____1960a, Uber Thomsonipollis magnificus (Th. and Pf. 1953) n. f. 

gen. n. comb, und Bemerkungen zur regionalen Verbreitung 

einiger Pollengruppen im alteren Palaogen: Freiberger 

Forschungshefte, v. C86, p. 54-65.

_____1960b, Beitrag zur Sporenpalaontologie der praoberoligozanen 

kontinentalen und marinen Tertiarablagerungen Brandenburgs: 

Geol. Gesell. DDR Berichte, v. 5, no. 4, p. 290-343. 

J.961, Zum Typus von "Pollenites pompeckji R. Pot., 1931a"

und von "Trudopollis subhercynicus Krutzsch 1954a": Geologie,

v. 10, no. 32, p. 94-96. 

Kulp, J. L., 1961, Geologic time scale: Science, v. 133, no. 3459, p.

1105-1114. 

Kunert, Reinhard, and Lenk, Gu'nter, 1964, Das Tertiar nordlich der

Halle-Hettstedter Gebirgsbruke: Geologie, v. 13, no. 4, p. 403-428. 

Leopold, E. B., and Pakiser, H. M., 1964, A preliminary report on the

pollen and spores of the pre-Selma Upper Cretaceous strata of

western Alabama: U.S. Geol. Survey Bull. 1160-E, p. 71-95. 

MacNeil, F. S., 1946, Geologic map of the Tertiary formations of

Alabama; U.S. Geol. Survey Oil and Gas Inv. (Prelim.) Map 45. 

Mellen, F. F., 1940, Geology, [Pt. 1] of Yazoo County mineral

resources: Mississippi State Geol. Survey Bull. 39, p. 9-72. 

Moore, W. H., 1965, Hinds County geology: Mississippi Geol. Econ.

and Topog. Survey Bull. 105, p. 21-145. 

Newman, K. R., 1965, Upper Cretaceous-Paleocene guide

palynomorphs from northwestern Colorado: Colorado Univ.

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Pflug, H. D., 1953, Zur Entstehung und Entwicklung des angiospermiden Pollens in der Erdgeschichte: Palaeontographica, ser. B., 

v. 95, nos. 4-6, p. 60-171.

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Geiseltals: Preuss. Geol. Landesanstalt, Inst. Palaobot. und 

Petrog. Brennsteine, Arb., v. 4, p. 25-125.

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Bull. 88, 123 p.

____1961, Geology study along Highway 80 from Alabama line to

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Skarby, Annie, 1968, Extratriporopollenites (Pflug) emend, from the

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Smith, F. E., 1959, Lower Tertiary and Upper Cretaceous of Brazos

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____1970, Two new pollen genera (Late Cretaceous and Paleocene)

with possible affinity to the Illiciaceae: U.S. Geol. Survey Prof.

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Prof. Paper, 743-B, 24 p.

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INDEX

[Italic page numbers indicate major references]

Page

abditus, Complexiopollis _________________ 9

admirabilis, Pseudoculopollis _________ 20; pi. 11

aequalis, Osculapollis ____________ 29, 30; pi. 18

anneratus, Kyandopollenites _________ 13, 14; pi. 7

artifex, Trudopollis ____________ 26, 27; pi. 17

Atlantopollis ________________________ 2

reticulatus _______________________31

atumescens, Basopollis __________________ 8

B

basalis, Basopollis _________-_____---_-- 8

Basopollis _____________________--_2, 1

atumescens ____________-_-----_-- 8

basalis _______________-_-----_-- 8

obscurocostatus ____ ____ _ _ 8; pi. 1

ornatus _________________-----_-- 8

orthobasalis _______________-----_7, 8

Bohemiapollis ______________________-20

bohemiensis, Pecakipollis _________ 16

caminus, Extremipollis _________ ----10, pi. 3

Choanopollenites _______________ 2, 3, 8, 35, 37

consanguineus __________________ _ __ 8

conspicuus _______________________ 8

discipulus ----------------------- 8

eximius ________________________ 8

patricius ________________________ 8

transitus ________________________ 8

sp. ____ __ _______________ pi. 1

Complexiopollis _______________ 2, 7, 8, 9, 35

abditus _________________________ 9

funiculus ________________________ 9

microuerrucatus ____________________ 9

patulus _________________________ 9

praeatumescens ____________________ 9

concentricoides, Pseudoculopollis ____________20

concentus. Oculopollis ________ __________20

Conclauipollis _______________________ 2

consanquineus, Choanopollenites _____________ 8

conspicuus, Choanopollenites ______________ 8

Nudopollis thiergartii ___________________76

discipulus. Choanopollenites _______________ 8

distinctus, Endoinfundibulapollis _______ 37; pi. 20

E

Emscheripollis _______________________13

endangulatus, Extratriporopollenites __________15

Nudopollis ___________________ 75; pi. 8

endocuspis, Pseudoplicapollis _________ 22; pi. 13

Endoinfundibulapollis-------- _ 37, 32, 34, 35, 37

distinctus __________________ 37; pi. 20

eximius, Choanopollenites ________________ 8

expolitus, Thomsonipollis ______________pl. 16

Extrapollis _________________________ 9

Extratriporopollenites ________________ 2, 7, 9

Page

Extratriporopollenites Con.

endangulatus _____________________15

pertrudens _______________________25

terminalis _______________________15

Extremipollis ______________ _9, 11, 34, 37

caminus ____________________ JO; pi. 3

maastrichtiensis ___________________ 9

versatilis ____________________ JO; pi. 2

uiutis ____________________ 9, 10; pi. 2

F, G, H

Fossil pollen, sample localities ______________ 3

type specimens _________________ _ _ _ 6

funiculus, Complexiopollis ________________ 9

gracilis, Thomsonipollis __________________25

hastaformis, Nudopollis terminalis ___________75

hebeformis, Nudopollis terminalis ___________ _75

Heidelbergipollis ______________________ 2

I

Interpollis _____________________ 77, 35, 37

intranodus ___________________ 72; pi. 5

microsupplingensis __________________72

supplingensis ______ _________ 77, 12; pi. 4

Interporopollenites ____________________ _72

proporus _____________________ 12, 13

turgidus -------_- ___________ 73; pi. 6

intraconcavus, Pseudovacuopollis _________ ___24

intranodus, Interpollis ______________ 7.2; pi. 5

Intratriporopollenites supplingensis ___________!!

involutus, Pseudovacuopollis __________ 23; pi. 14

K, L

Kyandopollenites ______ _________ 3, 73, 35, 37

anneratus _________________73, 14; pi. 7

Latipollis _________________________2, 7

Longanulipollis ____________________ 31, 32

M

maastrichtiensis, Extremipollis _____________ 9

magnificoides, Thomsonipollis ________3, 24; pi. 15

magnificus, Thomsonipollis __________ 24; pi. 14

meekeri, Trudopollis ___________________26

Megatriopollis __________________ 73, 34, 35

santonius ______________________ _13

sp. _______________________ 74; pi. 7

meridium. Nudopollis thiergartii ____________76

microsupplingensis, Interpollis ______________72

microuerrucatus, Complexiopollis ____________ 9

minimus, Minorpollis _____________ 74, 15; pi. 8

Minorpollis ________________2, 74, 22, 34, 35

Minorpollis minimus _____________74, 15; pi. 8

minutus, Nudopollis thiergartii _____________76

Semioculopollis ____________________21

Montanapollis _______________________ 2

munitus, Vacuopollis ______________ 28; pi. 18

N

Neotriangulipollis ___________________ 2, 30

Page 

New genus A___________________ 32; pi. 20

New genus B___________________ 32; pi. 20

New genus C ___________________ 32; pi. 20

New genus D ___________________ 33; pi. 20

nonperfectus, Trudopollis _________________27

Normapolles genera, Mississippi embayment, key __33 

stratigraphic distribution _____________ 35

Normapolles species, stratigraphic distribution ____37

Nudopollis ______________2, 3, 75, 34, 35; pi. 8

endangulatus _________________ 75; pi. 8

terminalis _____________ __15, 76, 35; pi. 8

hastaformis ____________________75

hebeformis ____________________75

thiergartii ___________________ 76; pi. 8

conspicuus ____________________76

meridium _____________________76

minutus _____________________ _76

0

obscurocostatus, Basopollis __________ ___8; pi. 1

ornatus, Basopollis ____________________ 8

orthobasalis, Basopollis _________________7, 8

orthopyramis, Vacuopollis ________________29

Osculapollis -------- ------ --- 29, 34, 35, 37

aequalis _________________ 29, 30; pi. 18

perspectus ________________ 29, 30; pi. 18

sp. A __________________________30

Oculopollis ---------------------- 20,21

concentus _______________________20

palaeocaenicus, Pseudoplicapollis _________ 21, 22

Papillopollis _______________________ _13

patricius, Choanopollenites---------------- 8

patulus, Complexiopollis _________________ 9

Pecakipollis ___________________2, 76, 34, 35

bohemiensis ______________________16

sp. ________________16; 17; pi. 9

percentus, Vacuopollis __________________28

perspectus, Osculapollis __________ 29, 30; pi. 18

pertrudens, Extratriporopollenites ____________25

Trudopollis ______________________25

plebius, Praecursipollis _____________ 30; pi. 19

plenus, Trudopollis _______________ 26; pi. 17

Plicapollis _________________ 2, 7, 77, 34, 35

retuaus _____________________ 78; pi. 9

rusticus _________________ 78, 19; pi. 10

serta __________________________!?

usitatus __________________ 77, 18; pi. 9

vacuus ____________________ 78; pi. 10

Pompeckjoidaepollenites __________ 19, 25, 34, 35

subhercynicus ________________ 79; pi. 11

praeatumescens, Complexiopollis ____________ 9

Praebasopollis _______________________ 2

Praecursipollis ________________ 30, 34, 35, 37

plebius ____________________ 30; pi. 19

Primipollis _________________________ 2

proparuus, Trudopollis ________________ __22

proporus, Interporopollenites ____________ 12, 13

Proteacidites _______________________ _31

sp. __________________________ _31

Pseudatlantopollis ______________ 30, 34, 35, 37

simulatus ________________ 30, 37; pi. 19

Pseudoculopollis _____________________ 20

admirabilis ________'_________ 20; pi. 11

41
42 INDEX

Page

Pseudoculopollis Con.

sp. A ___________________ 20, 21; pi. 12

Pseudoplicapollis _____________2, 21, 22, 34, 35

concentrictiides ____________________20

endocuspifi __________________ 22; pi. 13

palaeocaenicus _________________ 21. 22

pbcudosporites ___________________ _22

serenus __________________ 15, 22; pi. 13

sp. A ___________________ 22, 23; pi. 13

pseudnxporiten, Pseudopticapollifi ____________22

Pseudiii'dcuopollis __________________ 23, 34

intraconcavus __________________ 23, 24

invnlutu* ___________________ 23; pi. 14

Punctatricolporites ____________________ 2

Quedlinburgipollis _____________________ 2

rector, Trudopollis _____________________27

retieu/atus, Atlantopollis ___________ ______31

retusux. Plicapolli.i ________________ 78; pi. 9

rusting, Plicapollis _____________ 78, 19; pi. 10

Page

Semiaculopollis __________________ 2, 20, 21

minutus ________________________21

sp. A ____________________ 21; pi. 12

serenuK, Pseudoplicapollis ___________ 22; pi. 13

serf a, Plicapollis _____________________ _17

Siberiapoilis ________________________ 2

simulatus, Pseudatlantopollis _______ 30, 31; pi. 19

Sporopollis ______________^_____ 2, 22

Stratigraphic distribution, Normapolles genera ____35

Normapolles species _________________37

subhercynicus, Pompeckjoidaepollenites ___ 19; pi. 11 

supplingensis, Interpollis ___________!!, 72; pi. 4

Intratriporopollenites _________________!!

Systematic descriptions __________________ 7

terminalis, Extratriporopollenites ____________15

Nudopollis ________________ 15, 76; pi. 8

hastaformis _______________ _ __ _ 75

hebeformis ____________________75

thiergartii, Nudopollis ______________ 76; pi. 8

Nudopollis conspicuus ________________16

meridium ________________ _ _ _ _ 76

minutus _________________ _ _ _ _ -76

Thomsonipollis _____________ 2, 3, 24, 34, 35, 37

expoiitus ___________________ 25; pi. 16

gracilis ________________________-25

magnificoides _______ ________3, 24; pi. 15

magnificus ___________________ 24; pi. 14

Page

transitus, Choanopollenites ________________ 8

Triangulipollis _______________________32

Triatriopollenitef: _____________________ 7

Trudopollis ___________2, 9, 19, 25, 26, 29, 34, 35

artifex _________________ 26, 27; pi. 17

meekeri ________________________26

nonperfectus ______________________27

pertrudens _______________________25

plenus ____________________ 26; pi. 17

proparvus _______________________22

rector __________________________27

subhercynicus _____________________19

variabilis ______________ 25, 26, 28; pi. 16

sp. A ____________________ 27; pi. 17

sp. B _________________ 27, 28; pi. 17

sp. C _____________________ 28; pi. 17

turgidus, Interporopollenites ___________ 73; pi. 6

Turonipollis ____________________ 2,7,14

Type specimens, fossil pollens ______________ 6

U, V 

usitatus, Plicapollis ______________77, 18; pi. 9

Vacuopotlis ______________ 2, 23, 28, 29, 34, 35

munitus __________________ 28; pi. 18

orthopyramis _____________________29

percentus ______________________ _28

vacuus, Plicapollis ________________ 78; pi, 10

variabilis, Trudopollis ____________ 26, 28; pi. 16

uersatilis, Extremipollis ______________ 70; pi. 2

uiuus, Extremipollis ______________ 9, 10; pi. 2
PLATES 1-20

[Contact photographs of the plates in this report are available, at cost, from the U.S. Geological .Survey Photographic Library,

Federal Center, Denver, Colorado 80225]
PLATE 1

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-12. Basopollis obscurocostatus, n. sp.

1-3. Holotype, sample D1862, slide 3, preparation 2, coordinates 95.7X3.9,

diameter 29/u. 

3. Holotype X 2,000. 

4-7. Paratype, sample D1862, slide 3, preparation 2, coordinates 92.6X16.1,

diameter 42/u. 

8-10. Sample D1851, slide 3, coordinates 107.0X4.4, diameter 29/u.

10. Same X 2,000. 

11-12. Sample D1862, slide 3, preparation 2, coordinates 105.6X13.3,

diameter 29/u. 

13-15. Cf. Choanopollenites sp.

13. Sample D3138, slide 4, coordinates 88.5X13.2, diameter 26/i, X 2,000. 

14-15. Same X 1,000.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 1

15

BASOPOLLIS OBSCUROCOSTA TUS AND CF. CHOANOPOLLENITES SP.
PLATE 2

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-10. Extremipollis vivus, n. sp.

I-5, 8. Holotype, sample D3420, slide 2, coordinates 95.2X21.0, diameter 23ju.

8. Holotype X 2,000.

6-7. Paratype, sample D3420, slide 4, coordinates 94.4X21.0, diameter 27/j.. 

9-10. Sample D3420, slide 2, coordinates 94.8X7.8, diameter 23ju. A 

specimen with somewhat shorter annuli and more equilateral 

triangular shape. 

11-22. Extremipollis versatilis, n. sp.

II-15. Holotype, sample D3420, slide 3, coordinates 97.6X 11.5, diameter 27ju.

15. Holotype X 2,000.

16-17. Paratype, sample D3420, slide 3, coordinates 108.2X5.3, diameter 27ju. 

18-19. Sample D3420, slide 2, coordinates 99.0X10.0, diameter 26^. 

20-22. Sample D4268-A, slide 1, coordinates 88.1X15.3, diameter 30^. May be 

a variety or different species with thinner ektexine, provisionally including it here.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 2

EXTREMIPOLLIS VIVUS AND E. VERSATILIS
PLATE 3

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-18. Extremipollis caminus, n. sp.

1-5. Holotype, sample D3420, slide 4, coordinates 79.7X4.5, diameter 36ju.

5. Holotype X 2,000.

6-7. Paratype, sample D3420, slide 2, coordinates 84.3X 14.8, diameter 32ju.

8-9. Sample D3413, slide 3, coordinates 104.0X12.0, diameter 41M .

10. Sample D3420, slide 3, coordinates 82.0X13.4, diameter 31ju.

11-12. Sample D3420, slide 5, coordinates 112.1X10.2, diameter 36ju.

13. Sample D3420, slide 3, coordinates 79.5X20.7, diameter 32ju.

14-18. Sample D3420, slide 2, coordinates 104.2X14.5, diameter 25M .
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 3

EXTREMIPOLLIS CAMINUS
PLATE 4

[Magnification X 1,000 unless otherwise indicated] 

FIGURES 1-21. IntprpnUis rf J supplinppnsis

1-3. Sample D3410-C, slide 4, coordinates 76.4X21.1, diameter 28^.

3. Same X 2,000. 

4-5. Sample D3410-C, slide 3, coordinates 101.9X4.4, diameter 26^.

4. One external pore (at top) subequatorial; another, equatorial. Bacula

below ektexine gives appearance of rough surface.

6-8. Sample D3410-C, slide 5, preparation 3, coordinates 106.5X22.6, 

diameter 25^. Equatorial constriction shows clearly, as well as 

absence of bacula in polar regions.

9-13. Sample 1967-A, slide 1, coordinates 81.9X13.5, diameter 23M . 

14-15. Sample D3413, slide 2, coordinates 93.3X3.5, diameter 28/u. A variant 

with prominent thickening around endogerminal and slightly larger 

size. 

16-18. Sample D1967-A, slide 1, coordinates 93.3x2.2, diameter 20ji.

Equatorial constriction shows well. 

19-21. Sample D3260, slide 8, coordinates 80.4X14.5, diameter 16^, a smaller

variety. 

22-25. Interpollis microsupplingenis.

22-23. Sample D1855, slide 6, preparation 2, coordinates 82.3X12.9, diameter

20M . 

24-25. Sample D1848, slide 3, coordinates 88.1X13.4, diameter 20M .
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 4

10 11

INTERPOLLIS CF. /. S UPPLINGENSIS AND/. MICROS UPPLINGENSIS
PLATE 5

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-15. Interpollis intranodus, n. sp.

1-5. Holotype, sample D3285, slide 1, coordinates 90.3X15.3, diameter 25/x. 

1. Shows nodelike gemmae between ektexine and endexine appearing as

black dots.

5. Holotype X 2,000.

6-7. Paratype, sample D3285, slide 2, coordinates 97.7X21.7, diameter 25/j.. 

8-9. Sample D1966-C, slide 2, coordinates 103.8X11.6, diameter 28ju.

10. Same X 2,000. Note circular subequatorial pore in upper aperture. 

11-12. Sample D1966-F, slide 14, coordinates 80.5X22.5, diameter 25ju. 

13-14. Sample D3413, slide 3, coordinates 94.7X7.6, diameter 36ju. A slightly

larger variety. 

15. Same X 2,000. Interloculum shows well on this specimen.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 5

INTERPOLLIS INTRANODUS
PLATE 6

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-18. Interporopollenites turgidus, n. sp.

1-6. Holotype, sample D3548-A, slide 1, coordinates 77.5X11.8, diameter

26/^. Photographs at different focal levels. 

6. X 2,000.

7-8. Sample D3285, slide 2, coordinates 92.1X3.9, diameter 24/u. 

9-12. Paratype, sample D3548-A, slide 2, coordinates 78.4X6.5, diameter

29/u. 

13-15. Sample D3285, slide 1, coordinates 83.6X14.9, diameter 25jt. Note thin

polar spots on this and succeeding specimens. 

16-18. Sample D3548-A, slide 2, coordinates 84.1X11.0, diameter 26/u.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 6

13 14 15

INTERPOROPOLLENITES TURGIDUS
PLATE 7

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-5. Kyandopollenites anneratus Stover

1. Sample D1862, slide 6, preparation 2, coordinates 101.8X7.1, diameter 

55/u.

2. Same X 2,000, showing wall layers and pores. 

3-5. Sample D1846, slide 1, coordinates 102.4X11.8, diameter 55/u.

3. Surface ornamentation.

4. Intermediate focus.

5. Showing wall layers. 

6-10. Megatriopollis1? sp.

6-9. Sample D3285, slide 2, coordinates 100.1X7.9, diameter 29/u10. Same X 2,000.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 7

KYANDOPOLLENITES ANNERATUS AND MEGATRIOPOLLIS1 SP.
PLATE 8

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-9.

10-25. 

10-11.

12-15.

16-25.

Minorpollis aff. M. minimus

1-5. Sample D3285, slide 1, coordinates 82.3X6.7, diameter 15ju. 

Same X 2,000. 

Sample D1966-F, slide 

Sample D1966-F, slide

5.

6-7. 

8-9. 

Nudopollis 

Nudopollis aff. N. endangulatus

Sample D3222, slide 1, 

Nudopollis aff. N. thiergartii

12-13. Sample D3416, slide 1, 

L4-15. Sample D3416, slide 2, 

Nudopollis aff. N. terminalis

16-17. Sample D3222, slide 1, 

18-19. Sample D3548-A, slide 

20-21. Sample D1848, slide 3, 

22-25. Sample D1848, slide 3,

6. coordinates 76.4X6.9, diameter 15ju.

7. coordinates 104.2X4.6, diameter 16ju.

coordinates 107.5X15.3, diameter 25ju.

coordinates 103.3X13.7, diameter 28ju. 

coordinates 90.3X5.0, diameter 25ju.

coordinates 79.3X21.0, diameter 29ju. 

2, coordinates 80.7X9.2, diameter 28ju. 

coordinates 83.1X13.0, diamter 33ju. 

coordinates 85.7X18.2, diameter 36ju.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 8

10 11

MINORPOLLIS AFF. M. MINIMUS, NUDOPOLLIS AFF. N. ENDANGULATUS, 

N. AFF. N. THIERGARTII, AND N. AFF. N. TERMINALIS
PLATE 9

[Magnification X 1.000 unless otherwise indicated]

FIGURES 1-5. Cf. Pecakipollis sp.

Sample D3413, slide 1, coordinates 90.7X10.1, diameter 23^. 

1-2. 4-5. Different focal levels.

3. X 2,000. 

6-14. Plicapollis usitatus, n. sp.

6-8. Holotype, sample D3260, slide 2, coordinates 89.8X10.3, diameter 20^.

8. Same X 2,000.

9-10. Paratype, sample D3260, slide 1, coordinates 110.8X17.7, diameter 

19^. Note torn lower left aperture. Although exogerminals are clearly 

pores, this tear provides the appearance of a vertical slit shaped 

aperture.

11-12. Sample D3413, slide 1, coordinates 111.2X11.4, diameter 19.5^. 

13-14. Sample D3260, slide 3, coordinates 92.7X18.0, diameter 20^. 

15-24. Plicapollis retusus, n. sp.

15-19. Holotype, sample D3412, slide 3, coordinates 112.5X2.4, diameter 26^.

17. Holotype X 2,000.

20-21. Sample D3413, slide 1, coordinates 82.4X17.7, diameter 20^. 

22-24. Paratype, sample D3412, slide 4, coordinates 106.0X19.9, diameter 23M23. Paratype X 2,000.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 9

22 24

CF. PECAKIPOLLIS SP., PLICAPOLLIS USITATUS, AND PLICAPOLLIS RETUSUS
PLATE 10

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-9. Plicapollis uacuus, n. sp.

1-5. Holotype, sample D3412, slide 4, coordinates 87.0X3.1, diameter 21/u.

3. Holotype X 2,000.

6-7. Paratype, sample D3412, slide 1, coordinates 78.9X17.7, diameter 18/u. 

8-9. Sample D3416, slide 1, coordinates 108.9X6.9, diameter 17/u. 

10-21. Plicapollis rusticus, n. sp.

10-14. Holotype, sampleD3412,slide4,coordinates 110.9X9.4, diameter 28/u.

12. Holotype X 2,000.

15-16. Paratype, sample D3412, slide 1, coordinates 108.0X11.8, diameter 29/u. 

17-18. Sample D3412, slide 3, coordinates 112.0X5.7, diameter 23/u. 

19-21. Sample D3412, slide 3, coordinates 111.7X2.5, diameter 25/u. 

20. Same X 2,000.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 10

&*.*

/'I

/S»

?^j?i*\

s.Yf'Cv.^

.~»«Lf<tfe^ if

14

PLICAPOLLIS VACUUS AND P. RUSTICUS
PLATE 11

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-8. Pompeckjoidaepollenites aff. P. subhercynicus

1-2. Sample D1848, slide 3, coordiantes 82.6X 18.4, diameter 24ju. Note pore

and shifted platea. 

3-6. Sample D1848, slide 2, coordinates 90.6X2.1, diameter 23ju.

Interloculum visible on figures 5 and 6.

7-8. Sample D1848, slide 2, coordinates 93.9X7.2, diameter 26ju. 

9-18. Pseudoculopollis admirabilis, n. sp.

9-13. Holotype, sample D3420, slide 5, coordinates 81.5X9.8, diameter 31^- 

Extension of ocular swellings over one pole are on the obverse side of 

this specimen. Surface ornamentation shown on figures 11, 13. 

12. Holotype X 2,000.

14-16. Paratype, sample D3420, slide 1, coordinates 102.3X14.4, diameter 

29/u. Extensions of ocular swelling over pole present on upper surface 

of this specimen.

17. Sample D3420, slide 1, coordinates 83.0X3.7, diameter 27ju.

18. Sample D3420, slide 4, coordinates 89.3X16.0, diameter 30/u. Extensions of ocular swellings present on obverse side of this specimen.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 11

POMPECKJOIDAEPOLLENITES AFF. P. SUBHERCYNICUS AND PSEUDOCULOPOLLIS ADMIRABILIS
PLATE 12

[Magnification x 1,000 unless otherwise indicated]

FIGURES 1-6. Pseudoculopollis sp. A

Sample D3002, slide 3, coordinates 82.6X2.8, diameter 36/u. 

2. Same X 2,000. 

7-13. Semioculopollis sp. A

7-9. Sample D3260, slide 2, coordinates 108.0X4.4, diameter 25ju.

8. Same X 2,000. 

LO-13. Sample D3420, slide 5, coordinates 82.8X11.0, diameter 23/u.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 12

PSEUDOCULOPOLLIS SP. A AND SEMIOCULOPOLLIS SP A
PLATE 13

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-7. Pseudoplicapollis endocuspis, n. sp.

1-3. Holotype, sample D3265, slide 1, coordinates 101.2X6.7, diameter 18».

3. Holotype X 2,000.

4-5. Paratype, sample D3265, slide 3, coordinates 107.5X9.2, diameter 22^. 

6-7. Sample D3412, slide 1, coordinates 104.1X17.3, diameter 16^. 

8-12. Pseudoplicapollis serenus, n. sp.

8-10. Holotype, sample D3412, slide 2, coordinates 84.7X13.0, diameter 16M.

10. Holotype X 2,000.

11-12. Paratype, sample D3412, slide 4, coordinates 87.7X6.5, diameter 16M13-17. Pseudoplicapollis sp. A

13-15. Sample D3222, slide 1, coordinates 101.0X15.7, diameter 34^.

15. Same X 2,000.

16-17. Sample D1854, slide 3, preparation 2, coordinates 96.1X8.1, diameter 

31/j..
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 13

12

PSEUDOPLICAPOLLIS ENDOCUSPIS, P. SERENUS, AND P. SP. A
PLATE 14

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-12. Pseudovacuopollis involutus, n. sp.

1-4. Holotype, sample D3416, slide 2, coordinates 92.9X6.4, diameter 21/u.

4. Holotype X 2,000.

5. Sample D4268-A, slide 1, coordinates 104.0X2.4, diameter 20/u.

6. Sample D1967-C, slide 1, coordinates 103.7X5.0, diameter 21/u.

7. Sample D4268-A, slide 2, coordinates 74.8X10.3, diameter 20/u. 

8-12. Paratype, sample D3416, slide 2, coordinates 104.7X10.7, diameter 

18/u. The granulate appearance is due to granulation of endexine, 

ektexine is psilate. 

10. Paratype X 2,000. 

FIGURES 13-20. Thomsonipollis magnificus Krutzsch

13-14. Sample D1855, slide 3, preparation 2, coordinates 109.9X5.3, diameter

31/u. 

15-16. Sample D1855, slide 6, preparation 2, coordinates 108.5X18.6,

diameter 26/u.

17-20. Sample D1851, slide 3, coordinates 92.4X18.5, diameter 26/u. Specimen 

with somewhat thinner endexine somewhat intermediate between 

ThomsonipolliK magnificus and T. magnificoides.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 14

PSEUDOVACUOPOLLIS INVOLUTUS AND THOMSONIPOLLIS MAGNIFICUS
PLATE 15

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-8. Thomsonipollis magnificoides

1-2. Sample D1854, slide 3, preparation 2, coordinates 81.5X19.5, diameter

31/u.

3-4. Sample D1849, slide 1, coordinates 114.7X14.1, diameter 28/u. 

5-6. Sample D1849, slide 3, coordinates 105.8X18.0, diameter 31/u. 

7-8. Sample D1851, slide 3, coordinates 104.8X6.7, diameter 29/u. 

9-20. Thomsonipollis aff. T. magnificoides

9-12. Sample D3548-A, slide 2, coordinates 77.6X17.8, diameter 21M . 

13-14. Sample D3548-A, slide 2, coordinates 104.5X17.3, diameter 23/u. 

15-16. Sample D3548-A, slide 1, coordinates 88.7X19.1, diameter 27/u. 

17-20. Sample D3548-A, slide 2, coordinates 75.1X17.5, diameter 23/u.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 15

THOMSONIPOLLIS MAGNIFICOIDES AND T. AFF. T. MAGNIFICOIDES
PLATE 16

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-12. Thomsonipollis expolitus, n. sp.

1-4. Holotype, sample D3410-C, slide 4, coordinates 74.5X11.5, diameter

18n.

4. Holotype X 2,000.

5-7. Sample D3410-C, slide 2, coordinates 107.4X21.0, diameter 18ju. 

8-10. Paratype, sample D3410-C, slide 1, coordinates 104.6X21.4, diameter

20M .

10. Paratype X 2,000,

11-12. Sample D3410-C, slide 4, coordinates 74.5X4.5, diameter 20/u13-22. Trudopollis uariabilis, n. sp.

13-15. Holotype, sample D3412, slide 2, coordinates 109.7X14.1, diameter

26/u.

15. Holotype X 2,000.

16-17. Sample D3412, slide 4, coordinates 107.8X11.6, diameter 22/u. 

18-19. Paratype, sample D3412, slide 2, coordinates 99.5X14.8, diameter

24.5/u20-22. Sample D3260, slide 6, coordinates 113.9X6.1, diameter 28/u.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 16

^

16

»£

17

21

THOMSONIPOLLIS EXPOLITUS AND 772 UDOPOLLIS VARIABILIS
PLATE 17

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-9. Trudopollis plenus, n. sp.

1-5. Holotype, sample D3138, slide 4, coordinates 113.6X10.0, diameter

29ju.

3. Holotype X 2,000. Interloculum shows particularly well. 

6-7. Paratype, sample D3138, slide 1, coordinates 91.3X16.7, diameter 29juFaint arci present. 

8-9. Sample D3138, slide 3, coordinates 111.8X7.1, diameter 30ju. Annuli on

this specimen larger and slightly protruding. 

10-12. Trudopollis sp. A

Sample D3412, slide 1, coordinates 111.5X5.7, diameter 23ju. 

13-14. Trudopollis sp. B

Sample D1849, slide 2, coordinates 89.3X21.5, diameter 26ju. 

15-17. Trudopollis sp. C

Sample D1846, slide 2, coordinates 106.5X7.9, diameter 31ju. 

18-20. Trudopollis cf. T. artifex Wey. and Krg., 1953

Sample D3413, slide 3, coordinates 105.6X6.6, diameter 28ju.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 17

k1^^-^>»

>^ . j: T',

I.-^V '!«!

16 17

TRUDOPOLLIS PLENUS, T. CF. T. ARTIFEX, AND T. SP. A, B, AND C
PLATE 18

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-9. Vacuopollis munitus, n. sp.

1-3. Holotype, sample D3416, slide 1, coordinates 74.1X11.8, diameter 2fyu.

3. Holotype X 2,000.

4-5. Paratype, sample D3416, slide 1, coordinates 82.7X18.7, diameter 26^. 

6-7. Sample D3416, slide 2, coordinates 82.3X5.3, diameter 26M . 

8-9. Sample D1967-B, slide 1, coordinates 97.8X6.7, diameter 26^. This

specimen displays a somewhat thicker ektexine. 

10-20. Osculapollis aequalis, n. gen., n. sp.

10-14. Holotype, sample D3412, slide 4, coordinates 102.6X3.2, diameter 23^.

12. Holotype X 2,000.

15-16. Paratype, sample D3412, slide 3, coordinates 98.5X2.5, diameter 21^. 

17-18. Sample D3260, slide 5, coordinates 108.8X3.5, diameter 21.5^. 

19-20. Sample D3412, slide 3, coordinates 100.9X21.7, diameter 25^. 

21-27. Osculapollis perspectus, n. sp.

21-23. Holotype, sample D3412, slide 3, coordinates 104.1X16.3, diameter

29^.

21. Holotype X 2,000. 

24-25. Paratype, sample D3412, slide 3, coordinates 105.7X13.7, diameter

28M -

26-27. Sample D3412, slide 3, coordinates 95.0X12.9, diameter 23^. 

28-31. Osculapollis sp. A

28- 29. Sample D1862, slide 3, preparation 2, coordinates 97.0X10.0, diameter

34^. 

30-31. Sample D1862, slide 3, preparation 2, coordinates 93.4X17.3, diameter
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 18

VACUOPOLLIS MUNITUS, OSCULAPOLLIS AEQUALIS, O. PERSPECTUS, AND O. SP. A
PLATE 19

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-10. F*raecursipollisplebius,n.ger\.n.sp.

1-3. Holotype, sample D3265, slide 3, coordinates 87.0X5.5, diameter 16^.

3. Holotype X 2,000.

4-5. Sample D3265, slide 1, coordinates 102.3X5.4, diameter 19^. 

6-8. Paratype, sample D3265, slide 4, coordinates 78.4X7.1, diameter 20(u.

8. Paratype X 2,000.

9-10. Sample D3265, slide 1, coordinates 114.4X18.1, diameter 19.5^. 

11-28. Pseudatlantopollis simulatus, n. gen., n. sp.

11-13. Holotype, sample D1967-B, slide 10, coordinates 100.0X18.9, diameter

23M .

13. Holotype X 2,000.

14-15. Sample D1967-B, slide 8, coordinates 86.6X10.0, diameter 26^. 

16-17. Sample D1967-A, preparation 2, slide 6, coordinates 106.2X7.2,

diameter 21/^.

18-19. Sample D1967-B, slide 5, coordinates 88.4X4.4, diameter 23^. 

20-23. Paratype, sample D1967-B, slide 9, coordinates 111.9X18.9, diameter

25/^. The nature of the reticulum shows well on this specimen. 

24-28. Sample D3260, slide 8, coordinates 112.8X3.4, diameter 21^. 

26. Same X 2,000.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 19

Iv,

19

24 28

PRAECURSIPOLLIS PLEBIUS AND PSEUDATLANTOPOLLIS SIMULATUS
PLATE 20

[Magnification X 1,000 unless otherwise indicated]

FIGURES 1-11. Endoinfundibulapollis distinctus, n. gen., n. sp.

1-5. Holotype, sample D4268-B, slide 1, coordinates 78.5X21.9, diameter

20M3. Holotype X 2,000. 

6-7. Paratype, sample D4268-B, slide 2, coordinates 79.2X15.1, diameter

21ju.

8-9. Sample D3412, slide 4, coordinates 87.7X18, diameter 20ju10-11. Sample D4268-A, slide 1, coordinates 103.0X1.6, diameter 25ju. 

12-14. New Genus A.

12-13. Sample D3265, slide 3, coordinates 81.9X2.1, diameter 31ju.

14. Same X 2,000. 

15-17. New Genus B.

15-16. Sample D3412, slide 1, coordinates 90.9X13.6, diameter 18.5ju.

17. Same X 2,000. 

18-20. New Genus C.

18-19. Sample D3410-C, slide 2, coordinates 102.3X19.3, diameter 28ju.

20. Same X 2,000. 

21-23. New Genus D.

21-22. Sample D3410-B, slide 2, coordinates 101.1X19.0, diameter 44ju. 

23. Same X 2,000, one aperture only.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 865 PLATE 20

ENDOINFUNDIBULAPOLLIS DISTINCTUS AND NEW GENERA A, B, C, AND D