Complexiopollis Pollen Lineage r*/<j 

Mississippi Embayment Rocks

GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C

Work done in cooperation with the 

Kentucky Geological Survey
Complexiopollis Pollen Lineage in 

Mississippi Embayment Rocks

By ROBERT H. TSCHUDY

CONTRIBUTIONS TO PALEONTOLOGY

GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C

Work done in cooperation with the 

Kentucky Geological Survey

Stratigraphic ranges of new species belonging to the 

genera Complexiopollis and Choanopollenites

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

ROGERS C. B. MORTON, Secretary

GEOLOGICAL SURVEY 

V. E. McKelvey, Director

Library of Congress catalog-card No. 73-600117

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

Washington, D.C. 20402 - Price 75 cents Stock Number 2401-00345
CONTENTS

Page

Abstract ___________________________ Cl

Introduction __________________ __ 1

Acknowledgments _______-___,_____ 2

Methods ______________________________ 2

Type specimens _______________________ 2

Fossil pollen localities _________________ 2

Systematic descriptions _______ ______ 4

Genus Complexiopollis Krutzsch, 1959, emend __ 4

Complexiopollis funiculus n. sp ___ 4

Complexiopollis microverrucosus n. sp __- 5

Complexiopollis patulus n. sp _____ 6

Complexiopollis abditus n. sp _______ 7

Complexiopollis sp. A _______________ 7

Genus Choanopollenites Stover, 1966 ______ 7

Systematic descriptions Continued

Genus Choanopollenites Stover, 1966 Continued

Choanopollenites eximius Stover, 1966 .

Choanopollenites consanguineus n. sp _.

Choanopollenites transitus n. sp __ .

Choanopollenites discipuhis n. sp .

Choanopollenites sp. A _____ .

Choanopollenites sp. B _ .

Choanopollenites conspicuus (Grootand 

Groot) Tschudy n. comb ___ _____ .

Choanopollenites patricius n. sp _ . 

Suggested relationships of species included in the 

Complexiopollis lineage _______ __ .

References cited _____________ _ 

Index .

ILLUSTRATIONS

[Plates follow index]

PLATE 1. Complexiopollis funiculus n. sp.

2. Complexiopollis microverrucosus n. sp.

3. Complexiopollis patulus n. sp.

4. Complexiopollis abditus n. sp. and Complexiopollis sp. A.

5. Choanopollenites consanguineus n. sp.

6. Choanopollenites transitus n. sp. and Choanopollenites sp. B.

7. Choanopollenites discipuhis n. sp. and Choanopollenites sp. A.

8. Choanopollenites conspicuus (Groot and Groot 1962) Tschudy n. comb.

9. Choanopollenites patricius n. sp.

Page

FIGURE 1. Map of Mississippi Embayment showing fossil localities _____ C3 

2-11. Sketches of:

2. Complexiopollis funiculus _______ __ _ ____ 5

3. Complexiopollis microverrucosus ______________ 6

4. Complexiopollis patulus _____-_ -_____ 6

5. Complexiopollis abditus _________ _____ 7

6. Choanopollenites eximius _ _ 8

7. Choanopollenites consanguineus _ ____ 9

8. Choanopollenites transitus ______ __________ 9

9. Choanopollenites discipulus ___________________ 10

10. Choanopollenites conspicuus ____ ____ _ 11

11. Choanopollenites patricius _____ __ 12 

12. Diagram showing suggested relationships of species of Complexiopollis and Choanopollenites in Mississippi Embayment __-__ 13

Page

C8

8

9

9

10

10

10

12

12

14

15

III

CONTRIBUTIONS TO PALEONTOLOGY

COMPLEXIOPOLLIS POLLEN LINEAGE IN 

MISSISSIPPI EMBAYMENT ROCKS

By ROBERT H. TSCHUDY

ABSTRACT

Complexiopollis Krutzsch 1959, a genus proposed for fossil 

pollen grains and originally described from Europe, is represented in the Mississippi Embayment by four new species 

herein named and described. Choanopollenites Stover 1966 is 

a fossil pollen genus somewhat similar morphologically to 

Complexiopollis and by evolution is considered to be derived 

from it. Four new species of Choanopollenites are named 

and described and one species, reported previously, is included as a new combination.

The range zones of the species of Complexiopollis and 

Choanopollenites discussed in this paper are delineated, and 

the relationships between the species are suggested.

INTRODUCTION

The genus Complexiopollis, originally described 

from Europe by Krutzsch (1959), is a member of 

a complex of genera composing the group Normapolles of Pflug (1953). Pollen grains assigned to 

genera of the Normapolles group are commonly triangular, short axial, and morphologically highly 

differentiated, and they possess complex, and sometimes bizarre, apertures. So far as is known, these 

genera are extinct, and their relationship to modern 

families and genera is as yet unknown. Normapolles 

is a group of convenience and no genetic relationships among all the included genera are implied. 

Complexiopollis is one of the earliest, if not the 

earliest, genus of the Normapolles group to appear 

in the fossil record. In Europe it appears in the 

basal Cenomanian and persists at least into the 

basal Campanian.

Krutzsch (in Goczan and others, 1967) restudied 

genera belonging to the Normapolles group, erected 

several new genera, but of more importance, 

clarified the morphological details of wall and aperture structures of the several genera. He redescribed 

Complexiopollis and placed his genera Latipollis 

Krutzsch 1959 and Turonipollis Krutzsch 1959 in 

synonymy with Complexiopollis.

Several species were found in reference material 

from formations in the Mississippi Embayment and 

tentatively assigned to the genus Complexiopollis. 

The present work is the outcome of more careful 

examination of these species in preparation for providing descriptions and names for the undescribed 

new species.

The genus Complexiopollis (including Latipollis) 

has been reported from North American rocks by 

Leopold and Pakiser (1964) from the pre-Selma 

Cretaceous of Alabama, by Groot and Groot (1962) 

from the Paleocene of Maryland, by Gray and Groot 

(1966) from the Upper Cretaceous of Delaware and 

New Jersey, by Doyle (1969) from the Raritan and 

Magothy Formations of the Atlantic Coastal Plain, 

and by Wolfe and Pakiser (1971) from the Magothy 

and Raritan Formations of New Jersey and Maryland. Except for the papers by Groot and Groot 

(1962) and Gray and Groot (1966) none of these 

contributions provides species names or descriptions.

The material on hand from some Mississippi Embayment Upper Cretaceous and Tertiary formations 

has been ample for the description of nine species, 

eight new and one new combination, assignable to 

the Complexiopollis lineage. Furthermore, when the 

ranges of the several species are considered in stratigraphic sequence, evolutionary development of the 

pollen Complexiopollis to Choanopollenites with the 

retention of the basic morphologic features of Complexiopollis becomes evident.

The stratigraphic distribution of the several species and the evolutionary lineage that developed in 

North America is presented. The morphological features that characterize the genus Complexiopollis 

and that are present in modified form in the genus 

Choanopollenites Stover 1966 are demonstrated. 

Choanopollenites is here considered as belonging to 

the Complexiopollis lineage.

Cl
C2 CONTRIBUTIONS TO PALEONTOLOGY

ACKNOWLEDGMENTS

I thank the following people for providing some 

of the reference samples: Jack Wolfe of the U.S. 

Geological Survey for matrix material from collections of E. W. Berry in the U.S. National Museum, 

E. M. Gushing and W. W. Olive of the U.S. Geological Survey for the Clayton Formation and McNairy Sand samples studied, W. W. Fairchild of 

Standard Oil Company of Texas for the sample from 

the Naborton Formation, C. W. Copeland of the 

Geological Survey of Alabama for a sample from 

the Naheola Formation, and H. T. Ames of Pennsylvania State University for the sample from the 

Coal Bluff Marl Member of the Naheola Formation. 

Kathryn Dieterich and Sharon Van Loenen assisted 

in the preparation of the samples and the photography of specimens.

METHODS

The methods employed for processing rock samples and preparing slides are standard for this 

laboratory and have been reported previously 

(Tschudy, 1970).

I set an arbitrary minimum of 10 good specimens 

as the basis for assigning a name to a new species. 

Several potentially new species are mentioned but 

because of insufficient specimens are not named.

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 in the plate explanations. In order 

that other workers may convert their mechanicalstage readings to those recorded for specimens included in this report, the coordinates for the center 

point of a 1- by 3-inch standard microscope slide 

are 108.0 and 12.3 mm (designated as 108.0x12.3). 

The method of accurately locating the center of a 

standard microscope slide is described by Tschudy 

(1966, p. D78). If the slide label is placed 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.

FOSSIL POLLEN LOCALITIES

The fossil pollen grains used in this study are 

cataloged by the appropriate U.S. Geological Survey 

localities and by slide numbers. The rock samples

used for obtaining reference material from formations within the Mississippi Embayment are from 

the localities shown in figure 1. Twenty-six samples 

from 15 collecting localities in Louisiana, Missouri, 

Kentucky, Tennessee, Alabama, Georgia, and Mississippi were examined in detail.

The following descriptions of sampling localities 

are given in approximate stratigraphic order, oldest 

to youngest.

USGS

paleobotany

loc. No.

Locality Stratigraphic unit

D3415

D3265

D3260

D3412

D3413

D1967

D1967-A __ 

B .._. 

C __

D3416 _____

D3410

Outcrop at an altitude of 510 

feet on the south side of U.S. 

Highway 78, NW 1^ sec. 34, 

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

County, Ala. Collected by W. 

W. Olive, U.S. Geological Survey, and indicated by him 

(writen commun., 1964) as 

being "probably from unit 4 

described by Stevenson and 

Monroe, 1940, p. 46."

E. W. B'arry field No. 6474. Collected at Broken 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.

E. W. Berry field No. 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.

Type locality, near middle. Elev. 

440 feet along bluff on left 

bank Tennessee River 0.3 miles 

downstream from 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.

From roadcut along east side Tennessee Highway 22, 0.8 mile 

north of intersection with Tennessee Highway 100, Henderson County, Tenn. Collected by 

W. W. Olive, U.S. Geological 

Survey, 1964.

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 40 feet 

northwest of railroad crossing. 

South center NW^SW1^, 

Chewala quad., McNairy 

County, Tenn. Collected by E. 

M. Gushing, U.S. Geological 

Survey, 1962.

0-14 in. above bottom of 

sequence.

14-26 in. above bottom of 

sequence.

26-38 in. above bottom of 

sequence.

6 feet below unconformable contact with Owl Creek Formation; NW!/4NW% sec. 10. T. 

27 N., R. 11 E., Advance 7%- 

min quad., Stoddard County, 

Mo. Collected by W. W. Olive, 

U.S. Geological Survey, 1964.

Type locality. From right bank 

of Owl Creek, 2.5 miles northeast of Rip'ey, Miss., EVa sec. 

7, T. 4 S., R. 4 E., Tippah 

County, Miss. Collected by W. 

W. Olive, U.S. Geological Survey, 1964.

Tuscaloosa Formation.

Eutaw Formation.

Cusseta Sand Member, 

Ripley Formation.

Coffee Sand.

Coon Creek Tongue 

of Ripley Formation.

McNairy Sand Member 

of Ripley Formation.

Do. 

Do. 

Do.

Upper part McNairy 

Sand.

Owl Creek Formation.
COMPLEXIOPOLLIS POLLEN LINEAGE IN MISSISSIPPI EMBAYMENT ROCKS C3

37

NORTH 

Approximate northern limit CAROLINA

of Mississippi embayment

SOUTH CAROLINApproximate northern limit 

of Mississippi embayment 9

50 100 150 200 MILES 

_J

FIGURE 1. Map of Mississippi Embayment showing fossil localities and USGS paleobotany locality numbers. Selected counties are delineated within each State.

uses

paleobotany 

loc. No.

Locality Stratigraphic unit

D1966

D1966-A 

B 

C 

D 

E 

F

D3548 

D3548-A 

B 

C 

D

1.5 miles south-southwest of 

Cotton Plant, Miss., southwest 

wall of tributary to Guyton 

Creek along country road. 

SVzNW^ sec. 8, T. 6 S., E. 

3 E., Tippah County, Miss. 

Collected by E. M. Gushing, 

U.S. Geological Survey, 1962.

6 feet above basal limestone.

9 feet above basal limestone.

14 feet above basal limestone.

15 feet above basal limestone.

17 feet above basal limestone.

20-23 feet above basal limestone.

Auger hole J. 12, about 1 mile 

from southwest corner Joppa 

7%-min quad.; Kentucky coordinates, south zone: E. 

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

McCracken County, Ky. Collected by W. W. Olive, U.S. 

Geological Survey, 1965.

Depth 93-98 feet.

Depth 77-93 feet.

Depth 62-77 feet.

Depth 52-62 feet.

Clayton Formation.

Do. 

Do. 

Do. 

Do. 

Do. 

Do. 

Do.

Do.

Porters Creek Clay. 

Do. 

Do.

USGS

paleobotany

loc. No.

Locality Stratigraphic unit

D3285 _______

D1849 and 

D1850.

D3138

D3268

Channel of McCullough Fork, 300 

feet north Kentucky-Tennessee 

State line. Kentucky coordinates, south zone: E. 

1,263,500 feet; N. 70,250 feet. 

Hazel quad., Galloway County, 

Ky. Collected by W. W. Olive, 

U.S Geological Survey, 1963.

NW&SW1^ sec. 6, T. 11 N., E. 

11 W., De Soto Parish, La. 

Sample provided by W. W. 

Fairchild, Standard Oil Company of Texas, 1962.

Upper half Naheola Formation 

from along Alabama Highway 

17 in SE^NW1^ sec. 3, T. 15 

N., R. 2 W., Choctaw County, 

Ala. Sample provided by C. W. 

Copeland, Geological Survey of 

Alabama, 1963.

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 University, 1963.

Porters Creek Clay.

Naborton Formation.

Naheola Formation.

Coal Bluff Marl Member, Naheola Formation.
C4 CONTRIBUTIONS TO PALEONTOLOGY

uses

paleobotany 

loc. No.

Locality Stratigraphic unit

D3222 0.2 mile east of bridge across 

Tucabum River on LavacaPennington Road, south side 

new roadcut; sec. 22, T. 14, 

N., R. 1 W., Choctaw County, 

Ala. (See F. S. MacNeil, 1946) 

Collected by W. W. Olive and 

R. H. Tschudy. U.S. Geological 

Survey, 1963.

Nanafalia Formation.

SYSTEMATIC DESCRIPTIONS

Genus Complexiopollis Krutzsch, 1959, emend.

Latipollis Krutzsch 1959, Palaeontographica sec. B, v. 105,

p. 128. 

Turonipollis Krutzsch 1959, Palaeontographica sec. B, v.

105, p. 133.

Type species. Complexiopollis praeatumescens 

Krutzsch 1959.

Diagnosis. Pollen grains triangular flat lens 

shaped, rarely thick lens shaped; many triplanoid, 

range to typically triplane. Equatorial contour concave to extremely concave and twisted triangular, 

rarely convex triangular with bizarre, elongated 

angles (germinals). Contour sometimes interrupted, 

sometimes almost uninterrupted. Length of the 

polar axis differing greatly among the individual 

form-species, depending upon the shape, outline and 

triplane aspect of the grains. Wall thin, multilayered; in some species layers are distinct, in others 

appearing as a wall-complex. Endexine as thick as 

ektexine or slightly thicker. Surface smooth to 

weakly sculptured. Structure of germinals is complex. Exogerminal with a vertical (poleward directed) slit-shaped outer opening, two layered, the 

outer part without structure, the inner with radially 

projecting tightly packed bacula. Toward the roots 

of the exogerminals the bacula may be less densely 

packed, appearing as separate rods or narrow spines. 

Vestibulum small, in most grains not prominent. 

Mesogerminal consists of two flaplike structures 

separated by an equatorially directed slitlike opening at right angle to the exogerminal opening. An 

annuluslike thickening often present at the exterior 

extremities of the flaplike aperture. Centripetal to 

the mesogerminal, additional wall lamellae with 

atriate openings are developed. The inner wall layers 

at the root of the germinal are almost without structure. Secondary folding is incidental, particularly 

in the delicate triplane forms. No oculus or interloculum is present. Typical regular plicae are absent; 

however, some species may have delicate endexinal 

arci. The pollen species range in size from about 

I5fi (microns) to about 40^i.

Remarks. Krutzsch (in Goczan and others, 1967) 

reviewed the morphology of the genera Latipollis 

Krutzsch 1959 and Turonipollis Krutzsch 1959 and 

concluded that these two genera should be suppressed. He placed pollen species previously assigned 

to them in the revised genus Complexiopollis 

Krutzsch. In his revised generic diagnosis of Complexiopollis, Krutzsch stated that arci are absent. 

Several of the new species from Mississippi Embayment rocks possess delicate threadlike endexinal arci. 

The morphological features of these species are 

otherwise identical to those found in other species 

of Complexiopollis. I have, therefore, emended the 

genus Complexiopollis to include arci rather than 

erecting a new genus for these new species.

Forms which have a somewhat rigid rhombic 

equatorial outline (typically triplane forms) and 

which were previously assigned to the genus Latipollis may owe their shape, in part at least, to the 

presence of a somewhat rigid thickened part of the 

endexine that extends from pore to pore (the arci). 

A typical example of this triplane form is seen in 

Complexiopollis (formerly Latipollis) normis. The 

new species C. funlculus possesses much more prominent cordlike thickenings or arci extending from 

pore to pore. When compressed laterally this species 

assumes the typical triplane form.

The genus Atlantopollis Krutzsch 1967 is morphologically somewhat similar to Complexiopollis. 

Krutzsch (in Goczan and others, 1967, p. 446) distinguishes Atlantopollis from Complexiopollis by 

the prominent sculpture and the presence of a 

columella layer in Atlantopollis. However, Krutzsch 

does not mention in his diagnosis the presence of 

mesogerminal flaps or equatorially oriented mesogerminal openings, nor does he show these features 

in his diagrammatic representation of Atlantopollis. 

Consequently, for the present, I am excluding Atlantopollis from the Complexiopollis lineage.

Complexiopollis funiculus n. sp. 

Plate 1, figures 1-29; text figure 2

Holotype. USGS paleobotany loc. D3415, slide 3, 

coordinates 107.8x9.5, plate 1, figures 1-7.

Paratype. USGS paleobotany loc. D3415, slide 1, 

coordinates 78.9x21.4, plate 1, figures 8, 9.

Paratype. USGS paleobotany loc. D3415, slide 2, 

coordinates 108.5x16.9, plate 1, figures 11-15.

Diagnosis. Shape of pollen grains triangular to 

strongly concave in polar view, triplane in equatorial view and twisted into a partial triplanoid attitude. Size range, based on more than 100 specimens, 

equatorial diameter 23-36^, polar 17-21^. Wall made
COMPLEXIOPOLLIS POLLEN LINEAGE IN MISSISSIPPI EMBAYMENT ROCKS C5

B

FIGURE 2. Complexiopollis funiculus, ca. X 2000. A, 

Polar view showing arci. The upper cordlike endexinal thickenings have been shaded; the lower are 

shown in outline. B, Lateral or equatorial triplane 

view. Upper arci shaded. C, Equatorial view of aperture showing vertical outer slit and horizontal V-- 

shaped inner aperture bordered by flaps. D, Polar 

view of aperture showing additional lamella centripetal to the endexinal flaps.

up of at least two layers, the inner layer usually 

exhibiting evidence of several lamellae. Ektexine 

and endexine thin, their combined thickness less 

than 2/x. Surface of ektexine psilate to scabrate. 

Exogerminals 3.5-5^ wide (polar view) with a vertical slit-shaped outer opening. Exogerminal two 

layered, the outer thin and without internal structure, the inner consisting of radially directed tightly 

packed bacula. The thickened part of the ektexine 

in germinal area less than 5/* long in radial direction. Mesogerminal consisting of two fairly broad 

flaps on opposite sides of an equatorially elongated 

opening at right angle to the opening of the exogerminal. Cordlike endexinal thickenings extend 

from the sides of the mesogerminal openings to adjacent apertures. These form slight annuluslike thickenings at the outer margins of the flaps. In equatorial (or triplane) view the mesogerminal lamella 

is thickened above and below at the apertures. A

radially directed V-shaped opening separates the 

flaps. A very narrow vestibulum separates the exogerminal from the mesogerminal. Additional lamellae centripetal to the mesogerminal are distinguishable in a few specimens.

Stratigraphic distribution. Tuscaloosa and Eutaw Formations.

Remarks. Complexiopollis funiculus can be distinguished from most other species of Complexiopollis by its threadlike arci. Complexopollis microverrucosus sp. nov., although possessing arci, has a 

prominent verrucate surface sculpture; its apertures are larger in polar diameter; and its baculate 

layer is distinctly larger and more prominent. Complexiopollis abditus sp. nov. also possesses arci, but 

it is much smaller and its shape is triangular rather 

than concave triangular.

The specimen in triplane equatorial view (pi. 1, 

figs. 20-22) shows the upper aperture as it appears 

in polar view and the other two apertures in equatorial view. The specimen on plate 1, figures 23 and 

24 exhibits the arci in equatorial view, as drawn in 

text figure 2B.

Complexiopollis microverrucosus n. sp. 

Plate 2, figures 1-23; text figure 3

Holotype. USGS paleobotany loc. D3412, slide 3, 

coordinates 105.2x7.4, plate 2, figures 1-8.

Paratype. USGS paleobotany loc. D3412, slide 1, 

coordinates 82.7X16.3, plate 2, figures 9-11.

Paratype. USGS paleobotany loc. D3412, slide 1, 

coordinates 75.5x5.7, plate 2, figures 12-14.

Diagnosis. Shape of pollen grains triangular to 

strongly concave triangular in polar view, triplane 

in equatorial view, often twisted into a partial triplane attitude. Size, based on 25 specimens, 25-39^ 

in equatorial diameter. Wall thin, flexible; ektexine 

and endexine of about equal thickness, in combination less than 2/x thick. Ektexine verrucate with low 

relief. Exogerminal 8-10/x in polar diameter with a 

vertical slit-shaped outer opening; two layered, the 

outer thin without internal structure, the inner with 

radially projecting tightly packed bacula. Length of 

bacula shorter toward the root of the exogerminal. 

Mesogerminal consisting of two rounded flaps separated by a V-shaped opening. Mesogerminal opening equatorially directed and at right angle to the 

exogerminal opening. A narrow annulus is present 

at the extremities of the flaps, appearing as a loop 

or partial loop when seen in oblique view, and continuing as arci connecting the three apertures. Vestibulum between exogerminal and endogerminal 

somewhat prominent. At least one additional lamella
C6 CONTRIBUTIONS TO PALEONTOLOGY

FIGURE 3. Complexiopollis microverru&osus, ca. 

X 2000. The upper aperture shows a typical polar 

view, the one to the right an equatorial view, and 

the one to the left an intermediate view showing 

the loop-forming thickenings at the extremities 

of the mesogerminal flaps. Lamella forming atriate 

endogerminal shown in right aperture.

is present centripetal to the mesogerminal and forms 

an atriate endogerminal.

Stratigraphic distribution. Coffee Sand and 

Coon Creek Tongue of Ripley Formation.

Remarks. This species differs from Complexiopollis funiculus by its more verrucate surface sculpture, larger germinals and larger, more prominent 

baculate areas and more prominent loop-forming 

thickenings as extensions of the arci across the terminal portions of the mesogerminals. The looplike 

thickenings on the extremities of the mesogerminal 

flaps are similar to those figured for Turonipollis 

turonis (now Complexiopollis turonis) by Krutzsch 

(1959). However, Krutzsch presented photographs 

of but one specimen. In Krutzsch's photographs arci 

are not clearly evident, and the outer surface of his 

specimen has a smooth appearance.

The looplike appearance of the extremities of the 

mesogerminal flaps is shown particularly well on 

plate 2, figures 10, 12, and 14. Plate 2, figures 15-17 

and 21-23 shows this species in typical triplane 

attitude.

Complexiopollis patulus n. sp.

Plate 3, figures 1-18; text figure 4

Holotype. USGS paleobotany loc. D3265, slide 2, 

coordinates 97.0x15.5, plate 3, figures 1-3.

Paratype. USGS paleobotany loc. D3265, slide 3, 

coordinates 83.2x6.7, plate, 3, figures 6-9.

FIGURE 4. Complexiopollis patulus, ca. 

X 2000. A and B, Apertures in polar 

view showing variation in shape of 

germinals and the prominent lamellae 

forming endogerminal atria. C, Equatorial view of aperture showing Vshaped notch between flaps and the 

short, incomplete arci.

Diagnosis. Pollen shape triangular to slightly 

convex-triangular in polar view, germinal areas 

commonly protruding. In equatorial view rhomboidal 

triplane. Size 24-37^ in equatorial diameter (20 

specimens) and 22-28^ in polar diameter (eight 

specimens). Wall thin; ektexine and endexine difficult to distinguish, combined thickness about I/* in 

the interapertural areas. Surface psilate to microscabrate. Exogerminal a vertical slit-shaped opening surrounded by a two-layered thickening of the 

ektexine. Outer layer of ektexine without structure, 

inner layer made up of radially directed elongate 

tightly packed bacula. Wall thickness at exogerminal 

about 3^. Vestibulum prominent; in equatorial view 

ends of bacula appear as punctae. Endexine thin, 

except in vicinity of mesogerminal where short incomplete arcuslike thickenings are commonly present. Mesogerminal consisting of two delicate flaps. 

Opening between flaps equatorially directed at 

right angle to the exogerminal opening. In equatorial 

orientation a V-shaped notch is visible between the 

tips of the flaps. Additional lamellae centripetal to
COMPLEXIOPOLLIS POLLEN LINEAGE IN MISSISSIPPI EMBAYMENT ROCKS C7

the mesogerminal form atriate endogerminals. Endogerminal lamellae often prominent.

Stratigraphic distribution. Tuscaloosa and Eutaw Formations. Many specimens present in Eutaw 

sample.

Remarks. Complexiopollis patulus can be distinguished from all other species of Complexiopollis 

by its incomplete or interrupted arci.

The incomplete arci are shown on plate 3, figures 

6-11. The triplane attitude of this species is shown 

on plate 3, figures 12-18.

Complexiopollis abditus n. sp. 

Plate 4, figures 1-25; text figure 5

Holotype. USGS paleobotany loc. D3412, slide 3, 

coordinates 87.1x2.9, plate 4, figures 1-6.

Paratype. USGS paleobotany loc. D3412, slide 1, 

coordinates 101.2x16.3, plate 4, figures 7-10.

Diagnosis. Pollen small, rigid triangular in polar 

view, with slight bulges at the angles (apertural 

areas) ; thick lens shaped in equatorial view, although seldom found in this attitude. Size 15-21/x 

in equatorial diameter, based on 35 specimens. Wall 

thin, about 1/x thick in interapertural areas. Ektexine 

and endexine of about equal thickness. Ektexine 

psilate, rarely slightly microverrucate. Exogerminal 

in the form of a short vertical slit. Lips of exogerminal two layered; the outer ektexinal layer slightly 

thicker in apertural areas than in interapertural 

areas, without structure, the inner layer consisting 

of very short radially projecting bacula. Paired endexinal thickenings or plicae lying one above the 

other in polar view extend from aperture to aperture. Mesogerminal consisting of two, somewhat 

rounded, thin flaps separated by a shallow V-shaped 

notch. Mesogerminal opening horizontally directed 

at right angle to exogerminal opening. Endexinal

B

FIGURE 5. Complexiopollis abditus, ca. X 2000. 

A, Polar view of pollen grain. Upper arci shown 

as solid heavy line; lower arci in outline. B, Distorted, partial triplane or equatorial view (upper 

aperture) showing relationship of flaps and lower 

and upper arci.

thickenings do not traverse extremities of the flaps 

to form an annulus. Vestibulum between exogerminal 

and mesogerminal narrow. At least one additional 

lamella centripetal to the mesogerminal, forming a 

submesogerminal atriate opening. This lamella or 

lamellar complex is usually prominent.

Stratigraphic distribution. Eutaw Formation 

and Coffee Sand. Many specimens observed in the 

Coffee Sand sample.

Remarks. Complexiopollis abditus can be distinguished from other species that possess arci by its 

rigid triangular shape and small size. The paired 

arci lying one above the other are shown on plate 4, 

figures 13-22, and are shown less clearly on plate 4, 

figures 23-25.

Complexiopollis sp. A

Plate 4, figures 26-35

Illustrated specimens. 

USGS paleobotany loc. D3415, slide 3, coordinates

112.4x13.7, plate 4, figures 26-29. 

USGS paleobotany loc. D3265, slide 4, coordinates

87.87x11-0, plate 4, figures 30-33. 

USGS paleobotany loc. D3265, slide 1, coordinates

84.3x21.4, plate 4, figures 34-35.

Discussion. This pollen species was not given a 

specific epithet because too few specimens were observed (four only). In size (18-20^) and shape, 

pollen of this species is virtually the same as Complexopollis abditus. It appears to differ from the 

latter by a continuation of the endexinal thickenings 

across the mesogerminal flaps in the form of a pronounced annular thickening. This feature is absent 

from C. abditus.

Stratigraphic distribution. Tuscaloosa and Eutaw Formations.

Genus Choanopollenites Stover, 1966

Pollen of the genera Complexiopollis Krutzsch 

1959 and Choanopollenites Stover 1966 are very 

similar in basic structure, yet the morphological 

attributes of the type species of the two genera are 

strikingly different. This similarity is perhaps accentuated by several pollen species found in Mississippi Embayment rocks that show somewhat transitional characteristics. It is my conviction that both 

genera belong to the same lineage and that Choanopollenites is an evolutionary variant of the Complexiopollis group. Pollen of both genera possess 

vertical (polar-directed) exogerminals bordered by 

distinct ektexinal thickenings. Both possess inner 

germinals with equatorially directed openings at 

right angles to the exogerminal openings and both
C8 CONTRIBUTIONS TO PALEONTOLOGY

possess flap or valvelike lips on either side of the 

inner germinal.

Choanopollenites pollen differs from that of Complexiopollis by possessing a relatively thick wall, in 

which the ektexine is distinctly thicker than the 

endexine. In Complexiopollis the walls of the pollen 

grains are thin, and the endexine is thinner than 

the ektexine or is of approximately the same thickness. In some species of Complexiopollis the wall is 

so thin that the two wall layers are difficult to recognize. In equatorial view the pollen of Choanopollenites species is oval and is in contrast to the triplane rhomboidal shape of Complexiopollis species 

in the same view. Choanopollenites pollen further 

differs from that of Complexiopollis by lacking atriate endogerminals. Complexiopollis is described as 

having, in addition to the flap-shaped mesogerminals, 

one or more endogerminals with atriate apertures 

centripetal to the mesogerminals. Owing to the small 

size and thin walls of many species, the several wall 

layers are often difficult to differentiate. If sufficient 

specimens of a species are available, the distinguishing wall layers particularly those that make up the 

endogerminals can usually be found. This type of 

atriate endogerminal has not been observed in any 

of the species assigned to Choanopollenites. The 

single species of Choanopollenites previously described is from the lower Eocene. It is probably an 

evolutionary advanced form of the genus, and 

appears near the stratigraphic level where the 

genus becomes extinct. No representatives of the 

genus have been observed in rocks younger than 

early Eocene age.

Choanopollenites eximius Stover, 1966 

Text figure 6

Type species. Choanopollenites eximius Stover 

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

Paleontological Contributions, Paper 5, p. 8, pi. 4, 

figs, la-lh.

Diagnosis. Stover (in Stover and others, 1966) 

stated "Pollen isopolar triaperturate with radial 

symmetry. Outline subtriangular in polar view; oval 

in equatorial view. Apertures apical, equatorial and 

compound. Exine stratification conspicuous, sexine 

thicker than nexine, endosexine greatly thickened 

adjacent to apertures. Surface smooth, indistinctly 

roughened or punctate. Monotypic."

Species description not repeated here.

Stratigraphic distribution. Lower Eocene Wilcox 

Group (Stover, in Stover and others, 1966).

Remarks. In the description of the species Choanopollenites eximius, Stover mentions that the outer

FIGURE 6. Choanopollenites eximius. A, Pollen grain showing 

shape, ca. X 500. B, Aperture, 

ca. X 1000.

apertures are "elongate poleward," and that the 

inner apertures are "broadly elliptical" and "elongate equatorially." Choanopollenites eximius has not 

been observed in our reference samples.

Choanopollenites consanguineus n. sp. 

Plate 5, figures 1-27; text figure 7

Holotype. USGS paleobotany loc. D3413, slide 1, 

coordinates 107.1x17.4, plate 5, figures 1-6.

Paratype. USGS paleobotany loc. D3413, slide 1, 

coordinates 90.0x19.3, plate 5, figures 7-9.

Diagnosis. Shape of pollen grains triangular, 

sometimes slightly concave triangular in polar view, 

oval in equatorial view. Size 16-24/A in equatorial 

diameter (24 specimens), 13-16/x in polar diameter 

(five specimens). Wall 2-3/x thick in interapertural 

areas, thicker adjacent to the germinals. Ektexine 

more than twice as thick as endexine in interapertural areas. Ektexine psilate, sometimes slightly 

microverrucate particularly in interapertural and 

polar areas. Pore canal index less than 0.3. Exogerminal vertical slit shaped surrounded by a thickened, 

rarely slightly bulging two-layered ektexine; outer 

layer structureless, inner layer thin, consisting of 

tightly packed radially directed bacula. Vestibulum 

prominent. Inner germinal consisting of two flaps
COMPLEXIOPOLLIS POLLEN LINEAGE IN MISSISSIPPI EMBAYMENT ROCKS C9

FIGURE 7. Choanopollenites consanguineus, ca. X 2000. The wide 

endexinal apertures bordered by 

thickened endexinal flaps subtend 

the ektexinal apertures.

with the opening between the flaps at right angle to 

the exogerminal. Extremities of the flaps semicircular and usually slightly thickened; in equatorial 

view aperture between the flaps U-shaped.

Stratigraphic distribution. Coffee Sand, Coon 

Creek Tongue and McNairy Sand Member of Ripley 

Formation.

Remarks. Choanopollenites consanguineus differs 

from all other species of Choanopollenites except C. 

transitus by its small size. It differs from C. transitus in having a more rigid triangular shape, a 

generally thicker wall, smaller baculate areas in the 

pore canal areas, and shorter pore canals, and by 

having a slight thickening at the extremities of the 

endexinal flaps.

Choanopollenites transitus n. sp. 

Plate 6, figures 1-24; text figure 8

Holotype. USGS paleobotany loc. D3416, slide 2, 

coordinates 95.4x19-5, plate 6, figures 1-4.

Paratype. USGS paleobotany loc. D3416, slide 2, 

coordinates 103.9x6.8, plate 6, figures 5-8.

Diagnosis. Shape of pollen grains triangular, 

mostly concave triangular in polar view, oval in 

equatorial view, germinals at apices often slightly 

bulging; size 20-28^ in equatorial diameter (20 

specimens) and 13-15^ in polar diameter (two 

specimens). Wall about 2^ thick in interapertural 

areas, much thicker adjacent to germinals. Ektexine 

at least twice as thick as endexine in interapertural 

areas and about 5/x thick adjacent to germinals. 

Ektexine psilate. Pore canal elongate; pore canal 

index about 0.3. Exogerminal in the form of a vertical slit surrounded by a distinctly thickened twolayered ektexine; outer layer structureless, inner 

layer prominent, consisting of long, radially directed 

bacula. Bacula in baculate layer becoming progresFIGURE 8. Choanopollenites transitus, ca. X 2000. A, Polar view. 

B, Equatorial, partially triplane

view.

sively shorter toward the base of the germinals, and 

very short in the interapertural areas. A prominent, 

though narrow, vestibulum is present between the 

exogerminals and endogerminals. Endogerminal 

opening equatorially oriented, at right angle to the 

exogerminal. Aperture bordered above and below 

by two semicircular endexinal flaps. Additional 

lamellae centripetal to the endogerminal (as found 

in Complexiopollis) not evident.

Stratigraphic distribution. Coffee Sand, Coon 

Creek Tongue of Ripley Formation, McNairy Sand 

Member of Ripley Formation, Clayton Formation, 

and Porters Creek Clay.

Remarks. Choanopollenites transitus can be distinguished from other species by its small size, concave triangular shape, and proportionately large 

baculate area. It can be distinguished from C. consanguineus by its shape, elongate pore canal, and 

prominent baculate areas. The endexinal flaps show 

well on plate 6, figures 5, 8, 14-17, and 21. Plate 6, 

figures 20-24, shows specimens in triplane attitude.

Choanopollenites discipulus n. sp. 

Plate 7, figures 1-14; text figure 9

Holotype. USGS paleobotany loc. D3548-A, slide 

2, coordinates 97.0x21.4, plate 7, figures 1-5.
CIO CONTRIBUTIONS TO PALEONTOLOGY

FIGURE 9. Choanopollenites discipulus, 

ca. X 2000.

Paratype. USGS paleobotany loc. D3548-A, slide 

2, coordinates 83.9X20.4, plate 7, figures 6-9.

Diagnosis. Shape of pollen grains triangular to 

slightly concave triangular and bulging at the angles 

(apertures). Size 24-34/x in equatorial diameter (13 

specimens). Wall about 2-3/x thick in interapertural 

areas, much thicker at germinals. Ektexine at least 

twice as thick as endexine in interapertural areas. 

Surface smooth at germinals, slightly scabrate to 

verrucate in interapertural and polar areas. Exogerminal vertical slit shaped, surrounded by a twolayered, often bulging ektexine. Outer layer structureless, smooth, and thicker at base of germinal 

than at apex; inner layer of tightly packed, radially 

projecting bacula. Vestibulum between exogerminals 

and endogerminals very narrow. Endogerminal consisting of two triangular flaps, with the openings 

between the flaps at right angle to the exogerminal 

opening. Endogerminal flaps thickened in the form 

of an inverted V. Endexine conspicuously thinner in 

interapertural areas than in germinal areas.

Stratigraphic distribution. Clayton Formation 

and Porters Creek Clay.

Remarks. Choanopollenites discipulus differs 

from C. transitus and C. consanguineus in its larger 

size, thickened extremities of the endexinal flaps, 

and pointed triangular shape of the flaps. It differs 

from C. eximius, C. conspicuus, and C. patricius 

principally by its much smaller size.

Choanopollenites sp. A

Plate 7, figures 15-18

Illustrated specimen. USGS paleobotany loc. 

D1966-A, slide 1, coordinates 106.4x14.0.

Discussion. Only one specimen of this pollen 

species was found, consequently no specific epithet 

was assigned. This species, 40/x in diameter, is intermediate in size between Choanopollenites discipulus 

(24-34/x) and C. eximius (71-82,*). It differs from 

C. consanguineus by possessing less robust apertural 

areas and a thinner ektexine. Morphologically, it 

resembles C. discipulus and may be conspecific with 

the latter species.

Occurrence. Clayton Formation.

Choanopollenites sp. B 

Plate 6, figures 25, 26

Illustrated specimen. USGS paleobotany loc. 

D1849, slide 1, coordinates 99.2x23.1.

Discussion. Only two specimens of this pollen 

species were observed. No specific epithet was assigned. This species is about 30/x in equatorial diameter and may be characterized by greatly elongated 

pore canals. The exterior wall in the apertural areas 

is very thin, the baculate area occupying most of 

the area of the exogerminal.

Occurrence. Naborton Formation.

Choanopollenites conspicuus (Groot and Groot 1962) 

Tschudy n. comb.

Plate 8, figures 1-16; text figure 10

Latipollis conspicuus Groot and Groot 1962, Palaeontographica, sec. B., v. Ill, p. 169, pi. 30, figs. 35, 36; pi. 31,

figs. 1-3. 

Extratriporopollenites sp. Groot and Groot 1962, Palaeontographica, sec. B, v. Ill, p. 170, pi. 31, fig. 7. 

Extratriporopollenites sp. Tschudy 1971, in Geol. Soc.

America Spec. Paper 127, p. 98, figs. 9, 10.

Holotype. Latipollis conspicuus Groot and Groot 

1962.

Description. Groot and Groot (1962, p. 169) 

state: "Triplanar pollen grains with three germinal 

apertures which are rather narrow, elongated openings without annulus; the exine consists of an endexine, which curves toward the center of the grain 

near the apertures, and an ektexine, which is about 

2/x thick and distinctly scabrate; diameter of the 

grain is about 55/x, height about 35/x."

Emended diagnosis. Shape of pollen grains rigid 

triangular to concave triangular in polar view, 

broadly oval in equatorial view. Size (based on 19 

specimens) 35-57/t, average 46/x, in equatorial diameter (17 specimens), 22-30/x in polar diameter (two 

specimens). Ektexine thicker than endexine, with 

psilate to scabrate surface sculpture. Ektexine 3-4/x 

thick in the interapertural areas, and as much as 

Sfi thick at the apertures. Ektexine adjacent to exogerminal three layered; outer layer thin (less than 

1/x thick), scabrate with low relief, maculae present
COMPLEXIOPOLLIS POLLEN LINEAGE IN MISSISSIPPI EMBAYMENT ROCKS Cll

A

FIGURE 10. Choanopollenites conspicuus, ca. 

X 2000. A, Polar view of apertural region. 

B, Equatorial view of apertural region.

over entire surface of most grains; middle layer 

structureless, 2-3/x thick; the inner layer consisting 

of densely packed, radially oriented bacula. Baculate 

area is somewhat comma shaped, bacula distinctly 

shorter toward the inner part of the aperture. A 

layer of very short bacula separates the endexine 

from the ektexine throughout all areas of the grains. 

Exogerminal aperture a vertical slit-shaped opening, 

in equatorial view funnel shaped, tapered centripetally. Vestibulum very narrow. Endogerminal consisting of two somewhat triangular flaps; opening 

between the flaps at right angle to the exogerminal 

slit. Extremities of flaps thickened to as much as 

3/i. In equatorial view mesogerminal opening prominent, U-shaped, up to 10/x deep. Lamellae centripetal 

to mesogerminal not evident.

Stratigraphic distribution. Porters Creek Clay, 

Naheola Formation, and Brightseat Formation 

(Paleocene) of Maryland (Groot and Groot, 1962).

Remarks. Choanopollenites conspicuus can be 

distinguished from C. transitus, C. consanguineus, 

and C. discipulus by its larger size. It can be distinguished from C. eximius by the rounded rather 

than angular shape of the germinals in polar view, 

and from C. patricius by the convex triangular outline and the more delicate and triangular shape of 

the endexinal flaps.

Groot and Groot (1962) named this species Latipollis conspicuus from a specimen in equatorial view 

and figured specimens of L. conspicuus in partial 

triplane attitude (pi. 31, figs. 1-3) that demonstrate 

an intermediate view between polar and equatorial. 

These illustrations are similar to those shown on 

plate 8, figures 11-16. They figured a specimen in 

polar view as Extratriporopollenites sp. (pi. 31, fig. 

7). for which they provided the following description: "Triporate, pores equatorial and extremely 

protruding, long pore canals; exine consisting of 

endexine and ektexine, the former thickening toward 

the pore opening, and the latter of constant thickness; pore structure 18/x long and 15/x in diameter; 

equatorial diameter of grain 50/x." From an examination of specimens in polar, equatorial, and distorted 

triplane views, I am certain that the specimens mentioned above are conspecific and also are conspecific 

with my material. Krutzsch (in Goczan and others, 

1967) reexamined Latipollis Krutzsch 1959 and 

placed the type species into synonymy with Complexiopollis.

The holotype of Latipollis conspicuus was designated by Groot and Groot (1962, p. 169) as "Slide 

50311-B, grain 5, 105.6x20.8." However, they did 

not designate the holotype specimen of L. conspicuus 

on their plate description. Because they presented 

two and possibly three different specimens on their 

plates, I am assuming that the first one presented 

is the holotype (pi. 30, figs. 35-36).

Illustrated specimens. 

USGS paleobotany loc. D3285, slide 1, coordinates

91.7x17.5, plate 8, figures 1-3. 

USGS paleobotany loc. D3285, slide 1, coordinates

101.7X1.8, plate 8, figures 4-7. 

USGS paleobotany loc. D1966-F, slide 7, coordinates

89.3x13.7, plate 8, figures 8-10. 

USGS paleobotany loc. D1966-F, slide 1, coordinates

106.1x9.3, plate 8, figures 11-13. 

USGS paleobotany loc. D1966-F, slide 7, coordinates

99.4x7.4, plate 8, figures 14-16.
C12 CONTRIBUTIONS TO PALEONTOLOGY

Choanopollenites patricius n. sp.

Plate 9, figures 1-11; text figure 11

Holotype. USGS paleobotany loc. D3222, slide 3, 

coordinates 80.7x7.2, plate 9, figures 1-5.

Paratype. USGS paleobotany loc. D3222, slide 5, 

coordinates 83.1x20.9, plate 9, figures 6, 7.

Diagnosis. Pollen rounded triangular in polar 

view, enlarged at angles (apertures), oval in equatorial view. Apertural areas hemispheric at angles, 

separated by a narrow concave area along the sides 

of the grains midway between the apertural areas. 

Size 39-47/t in equatorial diameter (15 specimens) 

and 28-33/t in polar diameter (2 specimens). Wall 

about 3/t thick in interapertural areas, much thicker 

at the apertures. Ektexine about three times as 

thick as endexine. Exogerminal apertures short, 

vertical slit-shaped. Ektexine adjacent to exogerminal three layered; outer layer thin (less than I/* 

thick), scabrate to microverrucate, middle layer 

structureless, 6-8/* thick at the root of the exogerFlGURE 11. Choanopollenites patricius, ca. X 2000. 

A, Polar view of apertural area. B, Equatorial 

view of apertural area.

minal, inner layer of radially directed tightly packed 

bacula. Baculate layer difficult to see owing to the 

surface maculae. Endogerminal opening elongate 

equatorially, much larger than exogerminal opening 

and at right angle to it. Endogerminal with two 

very prominent valvelike thickened flaps. Terminal 

portion of flaps about 4/* thick in both equatorial 

and polar views. In equatorial view the extremities 

of the valvelike flaps are reflexed inward; opening 

between flaps broad U-shaped. Vestibulum narrow 

but prominent.

Stratigraphic distribution. Nanaf alia Formation.

Remarks. Choanopollenites patricius can be distinguished from all other species of Choanopollenites 

by its shape, and the thick hemispherical endexinal 

thickenings adjacent to the endogerminal. A pollen 

grain that appears to be conspecific with C. patricius 

was illustrated by Kuyl, Muller, and Waterbolk 

(1955, p. 68, pi. 3, fig. 9) from the "Eocene U.S.A."; 

and Fairchild and Elsik (1969, p. 85, fig. 55) figured 

a specimen from the Wilcox that also appears conspecific with C. patricius.

SUGGESTED RELATIONSHIPS OF SPECIES

INCLUDED IN THE COMPLEXIOPOLLIS

LINEAGE

The relationships suggested below can be only 

tentative. Too few samples from Campanian and 

older Cretaceous rocks have been examined from the 

Mississippi Embayment. Consequently it is anticipated that additional species of the Complexiopollis 

lineage will appear as more Cretaceous rocks from 

this area are examined.

In Europe the stratigraphic range of the genus 

Complexiopollis is from basal Cenomanian through 

the basal part of the Campanian, with questionable 

specimens found in the upper Maestrichtian 

(Goczan and others, 1967). This range is shown by 

the heavy bar in figure 12. The genus Choanopollenites has not been reported from Europe. The 

observed stratigraphic ranges of species of Complexiopollis and Choanopollenites found in Mississippi Embayment rocks are also shown on figure 12.

Complexiopollis funiculus, C. patulus, and C. sp. 

A were observed in samples from the Tuscaloosa and 

Eutaw Formations. The Tuscaloosa sample was 

taken from some distance above the base of the 

formation and also from some distance above the 

base of the Cenomanian. All three species may have 

been derived from a common older ancestor, or C. 

funiculus or C. patulus could have been the first 

species of this genus to appear in embayment rocks.
COMPLEXIOPOLLIS POLLEN LINEAGE IN MISSISSIPPI EMBAYMENT ROCKS C13

Estimated duration, in Approximate

millions of years stratigraphic

position of

Age Formation sample

20

10

6

11

3

3

3

20

Eocene

Paleocene

00

ZJ

O

Maestrichtian

Campanian

Santonian

Coniacian

Turonian

Cenomanian

Nanafalia Fm. D3222-*

Naheola Fm. D3138.D3268-*

Naborton Fm. 01849,01850-^

Porters Creek Clay 03285-^- 

Porters Creek Clay 03548-B-D-^-

Clayton Fm. D3548-A-*

Clayton Fm. D1966-A-F->

Owl Creek Fm. 03410-^

McNairy Sand D3416-*

McNairy Sand /-» 

Member of Ripley /

Formation D1967-A-C7 r> 

\Coon Creek Tongue D3413V

\ of Ripley Fm. /p*

/

Coffee Sand D3412

Cusseta Sand Member 03260-^ 

of Ripley Formation

Eutaw Fm. D3265->

Tuscaloosa Fm. 03415-^-

y\r--^_ /ii\ _J\

1

? < i

^ifL^-

^1

ll

I^X

saJ«S*i

^ v>

^^^as'

L '\- nCh.sp.B \ ^BV

W* f'W T // \\

- --- _ f \\

Ch. discipulus *

1 ^ - Ch. conspicuus

~~ \\Ch. sp. A

lAi ^" /J^\ y\ /jT\ n

^/ >», /^Vl J^\

Ch. transitus

C. microverruc

\

4%//\

\

\ 

\

\

\

^*k

1

nsus

=« ^^-^^^

Ch. consanguineus

' \ 

C. abditus ~^^

' \

\ 

\

\

\

\ 

\

\

C.funiculus

^-^^^

"^-^.

^.a.

C. sp. A C. patulus

.^-""'

FIGURE 12. Suggested relationships of species of Complexiopollis and Choanopollenites in Mississippi Embayment. Estimated duration, in millions of years, modified from Kulp (1961), Holmes (1965), and Gill and Cobban (1966). Heavy 

bar represents stratigraphic range of Complexiopollis in Europe; light solid bars, ranges of Complexiopollis species; 

open bars, ranges of Choanopollenites species.

The species C. abditus is morphologically so similar 

to C. sp. A. that the latter must have been derived 

from this stem. C. microverrucosus is the youngest 

species of Complexiopollis observed. It probably was 

derived from the older C. funiculus stock.

As has been mentioned previously, the genus 

Choanopollenites is similar structurally to the 

genus Complexiopollis. The oldest species observed 

in Mississippi Embayment rocks are Choanopollenites transitus and C. consanguineus. The former has

the greater stratigraphic range from upper Campanian through middle Paleocene; the range of the 

latter is limited to upper Campanian and Maestrichtian rocks. The species Choanopollenites discipulus, 

C. sp. A, C. sp. B, and C. conspicuus are progressively larger variants of the Choanopollenites transitus type. For the most part these larger species 

occur also in progressively younger rocks. The species Choanopollenites conspicuus has been reported 

previously from the Paleocene Brightseat Forma-
C14 CONTRIBUTIONS TO PALEONTOLOGY

tion of Maryland; and in Mississippi Embayment 

rocks this species is also confined to the Paleocene.

The morphologically more complex species of 

Choanopollenites, C. patricius, was observed in our 

material only in Nanafalia Formation rocks although 

many younger samples of Eocene age were examined.

The type species of Choanopollenites, C. eximius, 

was not observed in our reference material. Stover 

(in Stover and others, 1966) reported this species 

from the lower Eocene Wilcox Group from Milam 

County, Tex. On the basis of morphological criteria 

I believe that C. eximius probably was derived from 

the older C. conspicuus-discipulus complex.

REFERENCES CITED

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

Atlantic Coastal Plain and its evolutionary significance: 

Jour. Arnold Arboretum, v. 50, no. 1, p. 1-35.

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

palynomorphs of the Lower Tertiary in the Gulf Coast: 

Palaeontographica, sec. B, v. 128, pts. 3-6, p. 81-89.

Gill, J. R., and Cobban, W. A., 1966, The Red Bird section 

of the Upper Cretaceous Pierre Shale in Wyoming, 

with a section on A new echinoid from the Cretaceous 

Pierre Shale of eastern Wyoming by Porter M. Kier: 

U.S. Geol. Survey Prof. Paper 393-A, p. A1-A73.

Goczan, Ferenc, Groot, J. J., Krutzsch, Wilfried, and Pacltova, 

Blanka, 1967, Die Gattungen des "Stemma Normapolles 

Pflug 1953b" (Angiospermae). Neubeschreibungen and 

Revision europaischer Formen (Oberkreide bis Eozan): 

Palaont. Abhandlungen, sec. B, v. 2, no. 3, p. 429-539.

Gray, T. C., and Groot, J. J., 1966, Pollen and spores from 

the marine Upper Cretaceous formations of Delaware 

and New Jersey: Palaeontographica, sec. B, pts. 4-6, v. 

117, p. 114-134.

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

from the Brightseat Formation (Paleocene) of Maryland: Palaeontographica, sec. B., pts. 4-6, v. Ill, p. 

161-171.

Holmes, Arthur, 1965, Principles of physical geology, 2d ed. 

revised: New York, Ronald Press Co., 1288 p.

Krutzsch, Wilfried, 1959, Einige neue Formgattungen and 

-arten von Sporen und Pollen aus der mitteleuropaischen 

Oberkreide und dem Tertiar: Palaeontographica, sec. 

B, pts. 5-6, v. 105, p. 125-157.

Kulp, J. L., 1961, Geologic time scale: Science, v. 133, no. 

3459, p. 1105-1114.

Kuyl, O. S., Muller, J., and Waterbolk, H. Th., 1955, The 

application of palynology to oil geology with reference 

to western Venezuela: Geologie en Mijnbouw, no. 3, new 

series., v. 17, p. 49-76.

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.

Pflug, H. D., 1953, Zur Entstehung und Entwicklung des 

angiospermiden Pollens in der Erdgeschichte: Palaeontographica, sec. B., v. 95, pts. 4-6, p. 60-171.

Stevenson, L. W., and Monroe, W. H., 1940, The Upper 

Cretaceous deposits: Mississippi Geol. Survey Bull. 40, 

296 p.

Stover, L. E., Elsik, W. C., and Fairchild, W. W., 1966, New 

genera and species of early Tertiary palynomorphs from 

Gulf Coast: Kansas Univ. Paleont. Contr. Paper 5, p. 1- 

11.

Tschudy, R. H., 1966, Associated megaspores and microspores of the Cretaceous genus Ariadnaesporites Potonie, 

1956, emend., in Geological Survey research 1966: U.S. 

Geol. Survey Prof. Paper 550-D, p. D76-D82.

 1970, Two new pollen genera (Late Cretaceous and 

Paleocene) with possible affinity to the Illiciaceae: U.S. 

Geol. Survey Prof. Paper 643-F, 13 p.

1970, Palynology of the Cretaceous-Tertiary boundary

in the northern Rocky Mountain and Mississippi Embayment regions in Kosanke, R. M., and Cross, A. T., eds., 

Symposium on Palynology of the Late Cretaceous and 

Early Tertiary, San Francisco, Calif., Nov. 14, 1966: 

Geol. Soc. America Spec. Paper 127, p. 65-111. 

Wolfe, J. A., and Pakiser, H. M., 1971, Stratigraphic interpretations of some Cretaceous microfossil floras of the 

Middle Atlantic States, in Geological Survey research 

1971: U.S. Geol. Survey Prof. Paper 750- B, p. B35-B47.
INDEX

[Italic page numbers indicate major references]

Page 

abditus, ComplexiopoUia -----__________..._ C5, 7,13, pi. 4

Atlantopottis __._._,___ __.____...____________ 4

Choanopollenitea _-_.___-------- .-.-_-.______,________ 7

conaanguineua __._ . ___ ____________ 8, 9,10,11,13, pi. 5

conspicuua .-. . .__ 10, 13, pi. 8 

diacipulus _._..____. _____________________ 9, 11,13, pi. 7

eximius _. _ _ 8, 10, 11, 14 

patricius __ ___._--._ __________ 10, 11, 12, 14, pi. 9 

tranaitua __._ ____ _ .9, 10,11, 13, pi. 6

sp. A ______________-____.-_______________ 10, 13, pi. 7 

sp. B ____________ ____ ____ _____... 10, 13, pi. 6

Complexiopollis ----------_---------------___.--.__.. 1, 4, 7,12

abditus .. 5, 7, 13, pi. 4

funiculus -__ --__ -_---- 4, 6,12,13, pi. 1

microverrucoaua ___- ___ - -._ 5, 13, pi. 2

normis .... ___ ------------------- _ ____ ____ ____ _ _ 4

patulua .-_---_ - --_ 6, 12, pi. 3 

praeatumescena ------___________________ ___ _-__ - 4

turonis .-- _-- --- - -- 6 

sp. A ____ __ . 7,12,13, pi. 4

conaanguineus, Choanopollenitea - ------------------ 8, 9,10,11, 13, pi. 5

conspicuua, Choanopollenitea ..--------.--_. _-----_--... 10, 13, pi. 8

Latipottis 10,11

discipulus, Choanopollenitea --__--_-_-_--_--------_----_----0, 11, 13, pi. 7

eximius, Choanopollenitea _______________ _________________ 8,10,11,14

Page 

Extratriporopottemtea sp --_-_---.-. .-. .._.__ CIO, 11

Fossil pollen localities .-_---._. _.-._-_ 2 

funiculus, Complexiopollis --------- -------- .._ .. 4, 6,12,13, pi. 1

Latipottis -------______ - _______ ------ - - _ __ _ _ __ 1, 4

conspicuus _------------------ --_-_-__.--_ ...__ 10,11

normis __________________- ------------- ____ ______ 4

microverrucosus, Complexiopollis -------------- . ___ - 5, 13, pi. 2

Normapolles ----

normis, Complexiopollis

Latipoilis __.__-__

patricius, Choanopollenites - _-_-------____________ 10,11, 12, 14 pi. 9

patulus, Complexiopollis _ _-_- _--- 6,12, pi. 3 

praeatumescena, Complexiopollis ------ 4

Systematic descriptions

transitus, Choanopollenites . __ 9, 10,11,13, pi. 6 

Turonipollis - __________________ - - --- ________---_____- - 1, 4

turonis __-__ _.__ __________________ ------- _ ___ -___ 6

turonis, Complexiopollis __-_____-_-__-- ---------- ____ - - 6

TuronipoUis ___- -_.-. 6 

Type specimens, location ___.-- --- 2

C15

PLATES 1-9

Contact photographs of the plates in this report are available, at cost, from U.S. 

Geological Survey Library, Federal Center, Denver, Colorado 80225.
PLATE 1

[Magnification X 1000 unless otherwise indicated]

FIGURES 1-29. Complexiopollis funiculus n. sp.

1-7. Holotype, USGS paleobotany loc. D3415, slide 3, coordinates 107.8 X 9.5. 6, 7, Holotype X 2000.

8,9. Paratype, USGS paleobotany loc. D3415, slide 1, coordinates 78.9 X 21.4. 9, Paratype X 2000. 

10. USGS paleobotany loc. D3265, slide 1, coordinates 106.3

X 16.0.

11-15. Paratype, USGS paleobotany loc. D3415, slide 2, coordinates 108.5 X 16.9. 

16-19. USGS paleobotany loc. D3415, slide 2, coordinates 99.0

X 16.0. 

20-22. USGS paleobotany loc. D3415, slide 1, coordinates 89.8 X

22.5 

23-25. USGS paleobotany loc. D3415, slide 2, coordinates 96.0 X

6.4. 

26,27. USGS paleobotany loc. D3415, slide 1, coordinates 104.6

X 8.2.

28, 29. USGS paleobotany loc. D3415, slide 1, coordinates 89.9 X 

21.9.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C PLATE 1

22

26 ^ 27 

COMPLEXIOPOLLIS FUNICULUS
PLATE 2

[Magnification X 1000 unless otherwise indicated]

FIGURES 1-23. Complexiopollis microverrucosus n. sp.

1-8. Holotype, USGS paleobotany loc. D3412, slide 3, coordinates 105.2 X 7.4. 7, 8, Holotype X 2000.

9-11. Paratype, USGS paleobotany loc. D3412, slide 1, coordinates 82.7 X 16.3.

12-14. Paratype, USGS paleobotany loc. D3412, slide 1, coordinates 75.5 X 5.7. 14, Paratype X 2000. 

15-17. USGS paleobotany loc. D3412, slide 1, coordinates 93.4 X

6.4 

18-20. USGS paleobotany loc. D3412, slide 2, coordinates 97.8 X

15.6.

21-23. USGS paleobotany loc. D3412, slide 2, coordinates 78.1 X 

3.5.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C PLATE 2

22 23

COMPLEXIOPOLLIS MICROVERRUCOSUS
PLATE 3

[Magnification X 1000 unless otherwise indicated]

FIGURES 1-18. Complexiopollis patulus n. sp.

1-3. Holotype, USGS paleobotany loc. D3265, slide 2, coordinates 97.0 X 15.5. 3, Holotype X 2000. 

4, 5. USGS paleobotany loc. D3265, slide 4, coordinates 93.5 X

7.6.

6-9. Paratype, USGS paleobotany loc. D3265, slide 3, coordinates 83.2 X 6.7. 

10,11. USGS paleobotany loc. D3265, slide 2, coordinates 99.8 X

15.7. 

12-14. USGS paleobotany loc. D3265, slide 1, coordinates 97.8 X

8.8. 14, Magnification X 2000. 

15,16. USGS paleobotany loc. D3265, slide 2, coordinates 103.4

X 4.5.

17,18. USGS paleobotany loc. D3265, slide 1, coordinates 82.0 X 

6.1.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C PLATE 3

A

10

17 18

14

COMPLEXIOPOLLIS PA TUL US
PLATE 4

[Magnification X 1000 unless otherwise indicated]

FIGURES 1-25. Complexiopollis abditus n. sp.

1-6. Holotype, USGS paleobotany loc. D3412, slide 3, coordinates 87.1 X 2.9. 6, Holotype X 2000.

7-10. Paratype, USGS paleobotany loc. D3412, slide 1, coordinates 101.2 X 16.3.

11,12. USGS paleobotany loc. D3412, slide 3, coordinates 98.5 X 

5.5.

13-16. USGS paleobotany loc. D3412, slide 1, coordinates 78.7 X 

15.7.

17-19. USGS paleobotany loc. D3412, slide 1, coordinates 74.5 X 

17.9.

20-22. USGS paleobotany loc. D3412, slide 4, coordinates 79.5 X 

18.9.

23-25. USGS paleobotany loc. D3412, slide 1, coordinates 82.7

X 5.2. 

26-35. Complexiopollis sp. A.

26-29. USGS paleobotany loc. D3415, slide 3, coordinates 112.4 

X 13.7. 29, Magnification X 2000.

30-33. USGS paleobotany loc. D3265, slide 4, coordinates 87.8 

X 11.0.

34, 35. USGS paleobotany loc. D3265, slide 1, coordinates 84.3 X 

21.4.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C PLATE 4

13 14 15 16

10

11 12

17 18 19

20 21 22

23 24 25

fit

34 35

COMPLEXIOPOLLIS ABDITUS AND COMPLEXIOPOLLIS SP. A
PLATE 5

[Magnification X 1000 unless otherwise indicated]

FIGURES 1-27. Choanopollenites consanguineus n. sp.

1-6. Holotype, USGS paleobotany loc. D3413, slide 1, coordinates 107.1 X 17.4. 6, Holotype X 2000.

7-9. Paratype, USGS paleobotany loc. D3413, slide 1, coordinates 90.0 X 19.3.

10-15. USGS paleobotany loc. D1967-A, slide 6, coordinates 93.3 

X 23.3. 15, Magnification X 2000.

16,17. USGS paleobotany loc. D1969-C, slide 7, coordinates 114.0 

X 1.7.

18,19. USGS paleobotany loc. D3416, slide 1, coordinates 96.4 X 

11.7.

20,21. USGS paleobotany loc. D3416, slide 1, coordinates 75.3 X 

19.8.

22-24. USGS paleobotany loc. D3416, slide 2, coordinates 86.5 X 

14.0.

25-27. USGS paleobotany loc. D3413, slide 2, coordinates 105.5 

X 6.2.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C PLATE 5

12 13 14

16

23 24 25 26 

CHOANOPOLLENITES CONSANGUINEUS
PLATE 6

[Magnification X 1000 unless otherwise indicated]

FIGURES 1-24. Choanopollenites transitus n. sp.

1-4. Holotype, USGS paleobotany loc. D3416, slide 2, coordinates 95.4 X 19.5. 4, Holotype X 2000.

5-8. Paratype, USGS paleobotany loc. D3416, slide 2, coordinates 103.9 X 6.8. 

9-13. USGS paleobotany loc. D3548-A, slide 1, coordinates 73.0

X 9.1. 13, Magnification X 2000. 

14-17. USGS paleobotany loc. D3413, slide 2, coordinates 96.6 X

8.5. 16, 17 Magnification X 2000. 

18,19. USGS paleobotany loc. D3416, slide 1, coordinates 77.7 X

9.0. 

20,21. USGS paleobotany loc. D3416, slide 2, coordinates 75.2

X 17.3. 

22-24. USGS paleobotany loc. D3416, slide 2, coordinates 102.5

X 11.6.

25,26. Choanopollenites sp. B. USGS paleobotany loc. D1849, slide 1, coordinates 99.2 X 23.1.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C PLATE 6

22

CHOANOPOLLENITES TRANSITUS AND CHOANOPOLLENITES SP. B
PLATE 7

[Magnification X 1000 unless otherwise indicated]

FIGURES 1. 14. Choanopollenites discipulus n. sp.

1-5. Holotype, USGS paleobotany loc. D3548-A, slide 2, coordinates 97.0 X 21.4. 5, Holotype X 2000.

6-9. Paratype, USGS paleobotany loc. D3548-A, slide 2, coordinates 83.9 X 20.4. 9, Paratype X 2000. 

10-14. USGS paleobotany loc. D1966-F, slide 6, coordinates 87.7

X 15.3. 12, Magnification X 2000.

15-17. Choanopollenites sp. A. USGS paleobotany loc. D1966-A, slide 1, coordinates 106.4 X 14.0.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C PLATE 7

15 16 17 

CHOANOPOLLENITES DISCIPULUS AND CHOANOPOLLENITES SP. A
PLATE 8

[Magnification X 1000]

FIGURES 1-16. Choanopollenites conspicuus (Groot and Groot 1962) Tschudy n. comb. 

1-3. USGS paleobotany loc. D3285, slide 1, coordinates 91.7 X

17.5. 

4-7. USGS paleobotany loc. D3285, slide 1, coordinates 101.7

X 1.8. 

8-10. USGS paleobotany loc. D1966-F, slide 7, coordinates 89.3

X 13.7. 

11-13. USGS paleobotany loc. D1966-F, slide 1, coordinates 106.1

X 9.3.

14-16. USGS paleobotany loc. D1966-F, slide 7, coordinates 99.4 

X 7.4.
PROFESSIONAL PAPER 743-C PLATE 8

14

CHOANOPOLLENITES CONSPICUUS
PLATE 9

[Magnification X 1000]

FIGURES 1 11. Choanopollenites patricius n. sp.

1-5. Holotype, USGS paleobotany loc. D3222, slide 3, coordinates 80.7 X 7.2.

6,7. Paratype, USGS paleobotany loc. D3222, slide 5, coordinates 83.1 X 20.9.

8-11. USGS paleobotany loc. D3222, slide 2, coordinates 107.8 

X 13.4.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 743-C PLATE 9

9 10 

CHOANOPOLLENITES PATRICIUS

U. S. GOVERNMENT PRINTING OFFICE : 1973 O - 503-013