Larger Foraminifera of 

Late Eocene Ag;e From 

Eua, Tonga

GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-B
Larger Foraminifera of 

Late Eocene Age From 

Eua, Tonga

By W. STORKS COLE

LATE EOCENE FOSSILS FROM EUA, TONGA

GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-B

Study of eight species 

characteristic of upper Eocene 

(Tertiary b) stage

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1970
UNITED STATES DEPARTMENT OF THE INTERIOR 

WALTER J. HICKEL, Secretary

GEOLOGICAL SURVEY 

William T. Pecora, Director

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

Washington, D.C. 20402 - Price 55 cents (paper cover)
LATE EOCENE FOSSILS FROM EUA, TONGA FOREWORD

One of the most widespread units of the Cenozoic 

section in the islands of the open Pacific is a series of 

limestones assigned to the upper Eocene (Tertiary b). 

Such limestones, containing diagnostic larger Foraminifera, have been reported in many parts of an area 

spreading 4,000 miles across the tropical Pacific (fig. 1), 

from Palau and the Mariana Islands on the northwest 

through the Marshall Islands (Eniwetok) to Fiji and 

Tonga on the southeast (Whipple, in Hoffmeister, 1932, 

p. 79-86; Asano, 1939; Cole, 1950, 1957a, 1957b, 1960. 

In almost all the islands the limestones are dense and 

crystalline. Foraminifera and algae are abundant locally, but in most places fossils cannot be extracted and 

must therefore be studied in random thin sections. On 

the little island of Eua, Tonga, a locality was recently 

found where the Eocene limestone is tuffaceous, considerably weathered, and richly fossiliferous. Abundant 

fossils that represent a dozen organic groups were found. 

Such abundance and diversity signaled the find as a 

remarkable one that would add greatly to our knowledge of life in the western Pacific during the Eocene.

The island of Eua measures only 12 by 5 miles but 

it rises 1,000 feet above sea level. It occupies an interesting position tectonically, as its steep eastern side 

faces the Tonga Trench. In addition, Eua is the oldest 

island in the Tonga group that has a plutonic core 

(Guest, 1959) and a series of associated volcanic rocks, 

which are partly blanketed by thick limestones of late 

Eocene age. Younger volcanic rocks and sediments of 

late Tertiary age are also present (Hoffmeister, 1932).

This series of reports is concerned with one fades of 

the upper Eocene limestone. After the limestone series 

was deposited, Eua was uplifted periodically and a 

sequence of six terraces was cut in the limestones on 

the windward (eastern) side. Hoffmeister was the 

first to recognize the Eocene age of the main limestone 

of the terraces, three of which have veneers of Pliocene 

reef corals. He made a planetable map of the terraced 

eastern ridge and recorded the average altitudes of the 

terraces as 100, 200, 340, 400, 550, and 760 feet, The 

east-facing "rocky backbone" of Eua thus looks in 

profile like a giant staircase facing the Tonga Trench. 

The Eocene limestone may once have covered all of Eua

but is now largely limited to the eastern ridge (Hoffmeister, 1932; the Eocene Foraminifera were described 

by Whipple in this same report, p. 79-86).

The fossils described in this series of reports vere 

obtained from an outcrop on the 400-foot terrace 

about a quarter of a mile north of Vaingana (fig. 2). 

At this locality, the limestone lies close to the underlying volcanic rock and is tuffaceous and partly weathered; almost everywhere else on Eua the limestone is 

pure, hard, and crystalline.

In 1943, Harold T. Stearns, then of the U.S. Geological Survey, also served as a consultant to the 

Armed Forces at Pacific bases and made a brief visit 

to Eua. He collected a sample that contained half 

a dozen fossil brachiopods from the 400-foot terrace 

on the eastern side of the island. Stearns recorded the 

locality as: "Tele-a-hiva at elevation of 400 feet about 

% mile north of army lookout tower, at the second 

stream north of Vaigana [sic]." The brachiopods vere 

examined by G. A. Cooper of the U.S. National Museum. Some years later when I was studying other 

island fossils collected by Stearns, Cooper showed me 

the brachiopods and expressed a desire for additional 

specimens so that he could continue his study of their 

internal structures.

In 1966,1 learned that Yoshio Kondoof the Bernice P. 

Bishop Museum in Honolulu intended to visit Eua 

in connection with his studies of living Pacific island 

land snails (under National Science Foundation grant 

GB-3974). I sent Stearns' locality data and marked 

copy of Hoffmeister's Eua map to Kondo, and I informed Stearns of the plan to collect additional 

material.

Late in August 1967, Kondo reached Eua and, aided 

by a Tongan guide, Tomiki, and an interpreter, Mo^ese 

Vea, spent 2 days searching for the fossil locality. The 

lookout tower mentioned by Stearns 110 longer exists 

and Kondo found that Tele-a-hiva translates to "Nine 

Gulches." Traveling northward from "Vaigana" (Otu 

Vaingana) through heavy brush on exceedingly rugged 

karst topography for about 1,000 feet, he reached the 

first of the gulches. There he found a soft fossiliferous 

layer between two harder limestones and collected a

in
40-pound sample of the material. This gulch locality 

is probably not the exact spot visited by Stearns. The 

two collections have minor differences in nature of 

preservation, but they obviously came from the same 

formation.

The exact extent of the richly fossiliferous bed is

not known but it is probably limited both horizontally 

and vertically. In 1926, when Hoffmeister made his 

map of the terraces, he did not come upon this facies, 

and in 1928 when I spent 2 weeks on Eua with Hoffmeister, reviewing his mapping, no exposures of this 

zone were seen although we visited Vaingana. AddiFIGURE 1. Location of Tonga and other island groups in the southwest Pacific where upper Eocene limestone has been identified.

Dashed line marks structural boundary of the Pacific Basin (andesite line).
FIGURE 2. Map of Eua, Tonga, showing the location of. the 

recently discovered fossil outcrop (X) and the main mass of 

Eocene limestone (patterned area) on the east side of the 

island, as mapped by Hoffmeister (1932).

tional fieldwork in the area of the rugged "l rine 

Gulches" would be worthwhile.

William Melson of the Smithsonian Institution examined hand specimens and this section of the tuffaceous limestone and noted that the volcanic constituents are highly altered, making it difficult to determine 

their original nature. The rock is composed of 50 

percent or more of volcanoclastic debris, much of which 

has been replaced by calcite. The predominant volcanic 

fragments are of porphyritic pumiceous glassy material; 

most of the phenocrysts are plagioclase, now largely 

replaced by calcite. The original groundmass of pumiceous glass is now devitrifled and dark brown. 

Fragments of tuff are rare. There appears to be a 

large and varied assemblage of secondary minerals. 

The volcanic garments are mainly porphyritic andesitic rocks, or possibly plagioclase-bearing dacites. The 

presence of abundant fossils suggests that the volcanic 

material has been reworked.

The soft tuffaceous limestone collected by Kondo 

was treated with a wetting agent and penetrant ir the 

laboratory. The material broke down easily, revealing 

a variety of fossil remains: Foraminifera, discoasters, 

corals, hydrozoans, brachiopods, bryozoans, anneUds, 

crinoids, echinoids, ostracodes, barnacles, decapod crustaceans, mollusks, shark teeth, otoliths, and spores and 

other plant microfossils.

W. Storrs Cole has described the larger Forairinifera; these fossils suggest to him a depth of depos: tion 

of about 200 feet, but other groups notably the 

smaller Foraminifera, the corals, brachiopods, bryozoans, mollusks, ostracodes, and barnacles point to a 

considerably greater depth of deposition.

Material representing a total of 17 organic groups 

was distributed to paleontologists for study and report. 

Seven of these collections were small or were made up 

of incomplete specimens leading only to summary 

reports, but the others, except for the larger Foraminifera, contained much new material. The brachiopod, 

bryozoan, ostracode, barnacle, and mollusk collections 

contained the first identifiable Eocene species fron: the 

island of the open Pacific, an area extending 4,000 

miles from Palau to Tonga.

HARRY S. LADD

CONTENTS

Foreword-________________

Abstract..- _ ______________

Introduction. _____________

The fauna.__________--__-

Paleoecological implications _ 

Description of species. _

Family Camerinidae___

Genus Camerina,..

Gemis Pellatispira.

Page

ill

Bl

1

1

2

3

3

3

5

Description of species Continued 

Family Camerinidae Continued 

Genus Heterostegina- ________

Genus Spiroclypeus, ________

Family Discocyclinidae__________

Genus Discocyclina, _________

Genus Asterocyclina- ________

References cited.___________________

Index____-_______-_______--_--___-

Page

B 9 

10 

10

10

11

14

16

ILLUSTRATIONS

[Plates 1-5 follow index]

PLATE 1. Pellatispira.

2. Camerina, Spiroclypeus, Asterocyclina, Heterostegina, Discocyclina, and Pellatispira.

3. Asterocyclina, Heterostegina, and Camerina.

4, 5. Asterocyclina.

TABLE

Page

FIGURE 1. Map showing the location of Tonga and other island groups in the southwest Pacific where upper Eocene limestone has been identified-____--_-__________________________________-__----__--____-__-___--__-_-__ iv

2. Map of Eua, Tonga, showing the location of the recently discovered fossil outcrop and the main mass of Eocene

limestone.__-_--__,-_____________________._________________________-___---_-_-_--___--_-_____-_- v

Page 

TABLE 1. Distribution of larger Foraminifera in the Tertiary b (upper Eocene) sample from Eua, Tonga, and elsewhere. __ B2

VII

LATE EOCENE FOSSILS FROM EUA, TONGA

LARGER FORAMINIFERA OF LATE EOCENE AGE FROM EUA, TONGA

By W. STORES COLE

ABSTRACT

Eight species of larger Foraminifera characteristic of the 

Tertiary b (upper Eocene) stage of the Indo-Pacific region 

were recovered from a sample collected at an elevation of 400 

feet about one-quarter mile north of Vaingana, Eua, Tonga. The 

most abundant specimens in this fauna are Pellatispira. Heterostegina and Asterocyclina were found in moderate numbers, 

but Camerina and Spiroclypeus were uncommon. This fauna 

developed in warm, shallow waters (about 200 feet in depth) 

in a protected ecologic niche.

The genus Biplanispira Umbgrove, 1938, is demonstrated 

to be a synonym of Pellatispira Boussac, 1906. Specimens previously identified as Pellatispira fulgeria Whipple (synonym: 

Biplanispira absurda Umbgrove) are considered to be aberrant 

forms of Pellatispira madaraszi (Hantken).

INTRODUCTION

In 1932, Hoffmeister published a report on the geology of Eua, Tonga, one section of which was an analysis 

of larger Foraminifera prepared by G. Leslie Whipple. 

Species of larger Foraminifera from Tertiary b (upper 

Eocene) were described and illustrated, and "Recent 

species In the younger limestones" (Whipple, in Hoffmeister, 1932, table 5) were listed without discussion 

or illustrations.

Cole (1957a, p. 333, 334) examined the types of certain of the species described by Whipple and referred 

specimens from Saipan Island to these species. Later, 

Cole (1960a, p. A3) wrote:

Whipple (1932, in Hoffmeister, p. 79) described several species 

from the Tertiary b (upper Eocene) of Eua, Tonga. Certain of 

these species were identified incorrectly and others were inadequately described. A revised list of certain of these species follows :

Names used by Whipple Names used in this report

Camerina pengaronensis (Verbeek)__ C. pengaronensis

(Verbeek). 

Pellatispira fulgeria Whipple. ______ Biplanispira fulgeria

(Whipple).

Discocyclina (Discocyclina) fritschi D. (D.) omphala 

(H. DouvillS). (Fritsch).

euaensis Whipple,________ D. (D.) omphala

(Fritsch).

(Asterocyclina) stellata A. matanzensis Cole. 

(d'Archiac).

Harry S. Ladd of the U.S. Geological Survey sent the 

author abundant, excellently preserved, matrix-free 

specimens of larger Foraminifera collected by Yoshio 

Kondo of the Bernice P. Bishop Museum at an elevation

of 400 feet about one-quarter mile north of Vaingana, 

Eua, Tonga (fig. 2). This locality would be at the eastern 

edge of the Eocene limestone on the areal geologic map 

prepared by Hoffmeister (1932, p. 19, fig. 3) and 

associated with the 400-foot terrace (Hoffmeister, 1932, 

p. 14, fig. 2).

The most abundant specimens in the sample were 

Pellatispira madaraszi (Hantken). Whipple (in Hoffmeister, 1932, p. 81) identified specimens in the samples 

he studied as P. rutteni Umbgrove, a species which is 

considered to be a junior synonym of P. madc,raszi. 

Whipple wrote: "This species occurs abundantly at an 

altitude of 840 feet (E-390), associated with Camerina 

pengaronensis on the highest ridge of the island." In 

addition, specimens of Heterostegina sp. (= H. saipanensis Cole) were recorded from this locality (E-390).

The present sample seemingly represents the same or 

nearly the same ecological situation as the one in which 

Whipple found abundant Pellatispira and some specimens of Camerina and Heterostegina. However, the 

sample on which this report is based contained, in 

addition, many specimens of Asterocyclina and rare 

specimens of Spiroclypeus.

The specimens in the sample from Eua, Tonga, were 

sorted into groups based on external appearance. Selected specimens from each group were sectioned. Whereever possible, pairs of specimens that were identical in 

external appearance were selected from each group 

one of which was made into an equatorial sectior, and 

the other into a vertical section. A sufficient number of 

pairs of specimens were sectioned from each group to 

establish with certainty that the internal structures 

were similar. In total, over 150 oriented thin sections 

were made and photographed by the writer.

An attempt was made to obtain the specimens upon 

which Whipple (in Hoffmeister, 1932) based his report. 

However, these specimens have been misplaced, as they 

could not be located either at the University of Rochester or in the collection of the U.S. National Museum.

THE FAUNA

On first analysis, nine supposedly valid species 

(table 1), all of which had been reported from other 

Indo-Pacific localities assigned to the Tertiary b (upper

Bl
B2 LARGER FORAMINIFERA

Eocene) stage, were recognized. In addition one small 

specimen (pi. 2, fig. 18) of Discocyclina (Discocyclina), 

which could not be identified specifically, was found. 

Detailed study resulted in the reduction of the nine 

species to eight, as specimens identified as Pellatispira 

fulgeria Whipple were combined with P. madaraszi 

(Hantken). For clarity, however, in this analysis of the 

fauna, the name P. fulgeria (synonym: Biplanispira 

absurda Umbgrove) will be used, as this form has been 

reported as a valid species in many articles.

TABLE 1. Distribution of larger Foraminifera in the Tertiary b 

(upper Eocene) sample from Eua, Tonga, and elsewhere

Species found in the sample

praecipua Cole... ...

Pellatispira fulgeria l Whipple---.-- _

Occurrence elsewhere

Saipan

X 

X

X 

X 

X 

X 

X 

X

Eniwetok

X 

X 

X 

X 

X 

X 

X 

X 

X

Viti Levu, 

Fiji

X

X 

X

X 

X 

X

Indonesia

X

X

X 

X 

X

1 In the following section an attempt will be made to demonstrate that specimens 

previously identified as Pellatispira fulgeria (synonym: Biplanispira absurda) are 

aberrant forms of Pellatispira madaraszi. The specific name P. fulgeria is used in this 

section for comparative purposes only,

In the Tertiary b (upper Eocene) sediments of 

Saipan Island (Cole, 1957a, p. 322), 20 species of 

larger Foraminifera were identified in the examination 

of samples from 98 localities. In nine cores from Eniwetok Atoll drill hole F-l, 18 species of larger Foraminifera were recovered (Cole, 1957b, p. 749). Core 12 

(4,316-4,341 ft.) in this drill hole contained 10 of the 

18 species, which was the largest number recovered 

in any of the cores examined. This same core contained Pellatispira, madaraszi (Hantken) identified as 

P. orbitoidea (Provale) hi abundance.

In addition to P. madaraszi, core 12 of Eniwetok 

Atoll drill hole F-l had five other species which occur 

in the sample from Eua, Tonga. These species are: 

Asterocyclina matanzensis Cole, A. penuria Cole, A. 

praecipua Cole, Camerina pacifica (Whipple) (identified in core 12 as Operculinoides saipanensis Cole), and 

Heterostegina saipanensis Cole. Three species, Camerina pengaronensis (Verbeek), Pellatispira fulgeria 

Whipple, and Spiroclypeus vermicularis, which occur in 

the sample from Eua, Tonga, were not recovered from 

core 12; of these, however, C. pengaronensis and S. 

vermicularis occurred in core 11 (4,197-4,222 ft) of 

Eniwetok Atoll drill hole F-l.

Thus, in the 144-foot interval from the top of core 11 

(4,197 ft) to the bottom of core 12 (4,341 ft) in Eniwetok Atoll drill hole F-l, the nine species in the sample 

from Eua, Tonga, occurred. These data suggest that 

the ecological conditions under which the fauna of the 

sample from Eua, Tonga, formed were similar to those

of the interval represented by cores 11 and 12 of 

Eniwetok Atoll drill hole F-l.

Although the sample from Eua, Tonga, may be most 

satisfactorily correlated with this interval in Eniwetok 

Atoll drill hole F-l because of the number of species 

which are common, it is not implied that this fauna 

represents a distinct zone in the Tertiary b (upper 

Eocene) stage. For example, Whipple (in Hoffmeister, 

1932, table 5) recorded Discocyclina s. s. at four localities on Eua, Tonga, and Cole (1960a, p. A3) identified 

Discocyclina (Discocyclina) omphala (Fritsch) at one 

locality on Viti Levu, Fiji. As Pellatispira was not 

found in association with Discocyclina at th°<se localities, the assumption might be made that t-vo stratigraphically separate zones formed in the Tertiary b 

(upper Eocene) stage.

In the 98 samples from Saipan Island (Cc^e, 1957a, 

p. 322), Pellatispira and Discocyclina were both reported from 44 localities; however, Pellatispira and 

Discocyclina occurred in association at only 16 of these 

localities. These data suggest that the stn.tigraphic 

range of Pellatispira is the same as that of Discocyclina, 

but that Pellatispira favors one ecological situ ation and 

Discocyclina another.

Although there are sufficient data available to correlate the Tertiary b (upper Eocene) faunas tl roughout 

the Indo-Pacific region, our present knowledge does not 

indicate the possibility of subdivision of this stage into 

distinct and stratigraphically separate paleontologic 

zones characterized by one species or group of species.

As noted previously, only one small specimen that 

could be referred to the genus Discocyclina vras found 

in the Eua, Tonga, material. Hoffmeister (1932, p. 25) 

wrote that he collected Discocyclina along a small stream 

which had eroded a ravine in the 100-foot terrace, at 

which place "* * * the genus Discocyclina was found in 

the foraminiferal limestone beneath the thin coral reef 

of the lowest terrace." In discussing a high p; nnacle of 

limestone Hoffmeister (1932, p. 26) wrote: "Tv e rock at 

the base of the pinnacle is hard and compact, showing 

mainly foraminifera of the genus Discocyclina." The 

genus Discocyclina without question occurs in the Tertiary b (upper Eocene) limestones of Eua, Tonga, but it 

seemingly has a limited distribution which may be due 

to ecological controls.

Another large distinctive species which did not occur 

in the present sample is Pellatispira hqfmeisteri Whippie. This species, however, was reported by Cole (1957a, 

p. 334) from Saipan Island and is, in the writer's 

present evaluation of it, a valid species.

PALEOECOLOGICAL IMPLICATIONS

This fauna is dominated by abundant specimens of 

Pellatispira, which have an extensive variety of shapes
LATE EOCENE FOSSILS FROM EUA, TONGA B3

and sizes. Specimens of Heterostegina and Asterocyclina 

are present in moderate numbers, but representatives 

of the genera Camerina and Spiroclypeus are rare. Of 

these genera, only Camerina and Heterostegina have 

species still living.

Few, if any, of the tests of the larger Foraminifera 

in this fauna show any evidence of abrasion. The thin 

rims and elongate delicate rays of the Asterocyclina (pi. 

4, fig. 3) are perfectly preserved. The fragile margins (pi. 

1, figs. 2, 4) oiPellatispira are intact. Thus, the situation 

in which these large Foraminifera lived and their tests 

accumulated must have been protected from extensive 

wave or current action. Such protection might have 

been provided by the water depth, by a dense mat of 

seaweed, or by some favorable combination of these.

R. D. Norton (1930, p. 347) wrote: "Heterostegina 

seems to thrive in temperatures from about 22°C to 

27°C and in depths down to perhaps 40 fathoms." 

Cole (1957b, p. 750) analyzed the occurrence of certain 

Holocene genera of larger Foraminifera in the vicinity 

of the Philippine Islands and Bikini Atoll. He wrote: 

"The abundant or common occurrence of Heterostegina 

at average depths of 25-32 fathoms agrees with the 40 

fathoms given by R. D. Norton."

Cushman, Todd, and Post (1954, p. 320) wrote concerning the faunas of the lagoons of certain of the 

Marshall Islands: "In the lagoon fauna as a whole, the 

dominant species is Amphistegina madagascariensis 

D'Orbigny. Next in abundance in the lagoons is 

Heterostegina suborbicularis D'Orbigny * * *." Myers 

(1943, p. 29) observed that Heterostegina is "* * * 

found on firm sandy mud bottoms and are most numerous near patches of weeds where the juveniles may be 

taken in considerable numbers."

Genera such as Camerina seem to have a greater 

depth range than Heterostegina (Cole, 1957b, p. 750), 

as abundant specimens of Camerina have been reported 

at depths of 10 fathoms or less to 70 fathoms or more. 

Myers (1943, p. 29) found, moreover, that Camerina 

(= Operculina) flourished best on "soft mud bottoms."

The composition of this fauna in comparison with 

the ecological conditions under which living specie^1 of 

Heterostegina and Camerina exist suggests that it formed 

in warm shallow water (about 200 ft in depth) that had a 

moderately firm substratum upon which extensive 

patches of weeds grew. These conditions could be 

satisfied best in a lagoon but possibly could prevail in 

a protected situation on the outermost part of a reef.

DESCRIPTION OF SPECIES

Family CAMERINIDAE 

Genus CAMERINA Bruguiere, 1792

Camerina pacifica (Whipple) 

Plate 2, figures 1-5

1932. Operculina pacifica Whipple, in Hoffmeister, Bernice P.

Bishop Mus. Bull. 96, p. 83, pi. 20, figs. 1, 8. 

1953. Operculina sp., in Cole and Bridge, U.S. Geol. Survey

Prof. Paper 253, p. 21, pi. 4, fig. 15. 

1957. Operculinoides saipanensis Cole, U.S. Geol. Survey

Prof. Paper 280-1, p. 331, pi. 102, figs. 15, 16. 

1957. Operculinoides saipanensis Cole. Cole, U.S. Geol. Survey

Prof. Paper 260-V, p. 755, pi. 232, figs. 7-14; pi. 233, figs.

31, 32.

1959. Operculina eniwetokensis Cole, U.S. Geol. Survey Prof. 

Paper 260-V, p. 756, pi. 232, figs. 15-23 [imprint date 

1957].

1960. Operculina saipanensis (Cole). Cole, U.S. Geol. Survey

Prof. Paper 374-A, p. A4. 

1963. Operculina saipanensis (Cole). U.S. Geol. Survey Prof.

Paper 403-E, p. E3. 

1963. Operculina eniwetokensis Cole. Cole, U.S. Geol. Survey

Prof. Paper 403-E, p. E3, E16, pi. 5, figs. 11, 12- 25.

The test is small and has an inflated subcentral area 

over the embryonic chambers surrounded by a rim 

which progressively widens distally. The sutures of the 

chamber walls show on the surface of the flat rim as 

slightly raised, gently recurved lines.

Measurements of thin sections of Tongan specimens, 

to which have been subjoined measurements made from 

the median section of the type published by Whipple 

(in Hoffmeister, 1932, pi. 20, fig. 1), follow.

Measurements of sections of Camerina pacifica (Whipple)

Height __ ....................... mm.. 

Width. .............._........... .mm..

Embryonic chambers:

Number of chambers in final volution...

Fig. 1

2.18 

1.83+

55x60

40x90

100

3

10

22

50

Specimen shown on pi. 2 

Fig. 2 Fig. 3 Fig. 4 '

2. 2 2. 1 2. 1 

1.9 1.55 1.7+ .

50x50 .........................

40x70 .........................

100 ........... ..............

3.75 ............ 5 .

7 .........................

10 25 31 .

31 ..--...-...- 90 .

Type specimen

Fig. 5 Hoflmeister, 1932, 

pi. 20, fig. 1)

1. 9+ 4. 0 

........... 3.2

.5 ...,_____.._.--.....

125(?)

........... 2.5

..-_.-..--. 7

........... 15

......._... 34

220 ....................

1 Microspheric specimen.
B4 LARGER FORAMINIFERA

Remarks. Since 1932 three specific names, Operculina pacifica Whipple (in Hoffmeister, 1932, p. 83), 

Operculinoides saipanensis Cole (1957a, p. 331), and 

Operculina eniwetokensis Cole (1957b, p. 756), have 

been given to small operculine camerinids from Tertiary 

b (upper Eocene) sediments. All these specimens seemingly should be referred to one species, Camerina pacifica (Whipple).

In part, the separation of the specimens into three 

species resulted from an attempt to use an artificial 

generic classification in which such inexactly defined 

genera as Operculina and Operculinoides were recognized 

(Cole, 1966). In addition, minor variations in size, number of chambers, and similar features were given exaggerated importance.

Megalospheric specimens (pi. 2, figs. 1,5) as well as 

microspheric specimens (pi. 2, fig. 4) in the sample from 

Eua, Tonga, are similar to the types of Operculinoides 

saipanensis Cole (1957a, pi. 102, figs. 15, 16) from Saipan Island and to specimens assigned to that species 

from Eniwetok Atoll drill hole F-l (Cole, 1957b, pi. 232 

figs. 7-14; pi. 233, figs. 31, 32). The types and the 

specimens from Eniwetok Atoll drill hole F-l were assigned to the genus Operculinoides.

Other specimens recovered from Eniwetok Atoll drill 

hole F-l (Cole, 1957b, p. 756, pi. 232, figs. 15-23), 

which had fewer chambers and a thinner subcentral 

umbonal area, were assigned to the genus Operculina 

and described as a new species, Operculina eniwetokensis 

Cole. One figured specimen (pi. 2, fig. 2) is similar to 

the types of this species.

Later detailed studies (Cole, 1960b, 1961, p. 377-383; 

1966) of large suites of specimens from many geographic areas and from various stratigraphic horizons 

have demonstrated that the assumed structural differences upon which many of the supposedly valid generic names for camerinids were based are common to 

all these specimens. Cole (1961, p. 377) wrote: "These 

genera have been defined in terms of intergradational 

features which are specific rather than generic 

differences."

Although the postulate that the specimens under 

consideration should be assigned to the genus Camerina may not be acceptable, it is evident from the 

structure and development of the test that they cannot 

be placed in two genera.

The test of the specimens which Whipple (1932, in 

Hoffmeister, p. 83) named Operculina pacifica is about 

twice as large as that of 0. eniwetokensis, but the 

number of coils, chambers per volution, and certain 

other features of 0. pacifica are similar to those of 

0. eniwetokensis. The distal chambers of 0. pacHica 

are slightly over twice as high and are twice as wide 

as those of 0. eniwetokensis. Thus, the only difference 

between these two species is the size of the chambers,

A possible explanation of this increase in size of 

the chambers in 0. pacifica is that these specimens 

developed in a more favorable ecological situation than 

did the smaller specimens named 0. eniwetokensis. 

Moreover, large and small specimens of a species often 

occur together (Cole, 1958, pi. 18, figs. 7, ?).

The only significant differences between specimens 

named Operculinoides saipanensis and Operculina eniwetokensis are the number of chambers p^r volution 

and the total number of chambers in the test. The 

chambers in 0. saipanensis have approximately the 

same height as those of 0. eniwetokensis, but the chambers in 0. saipanensis are narrower, thus more 

chambers developed per volution.

As the differences in the number of chambers and 

in their size can readily be controlled by e?ologic and 

other factors, such as the availability of food and 

rapidity of test development, all these specimens are 

considered to be ecological variants of one species.

The chamber walls of the one available median section of Camerina pacifica (Whipple) (1932, in Hoffmeister, pi. 20, fig. 1) are irregularly thickened on the 

distal side. This same kind of development occurs 

occasionally in specimens formerly referred to Operculinoides saipanensis (see Cole, 1957b, pi. 232, fig. 

12) and in specimens (pi. 2, fig. 2) in the present fauna.

Camerina pengaronensis (Verbeek) 

Plate 3, figures 10, 11, 13-16

1871. NummuUtes pengaronensis Verbeek, Neues Jahrb. Mineralogie, Geologic, u. Palaeontologie, Jahr. 1871, p. 3-6, 

pi. 1, figs. la-k.

1892. NummuUtes nanggoelani Verbeek, Natuurk. Tijdschr. Nederlandsche-Indie, v. 51, no. 2, p. 116, 118 [imprint date 

1891].

1932. Camerina pengaronensis (Verbeek). Doornink, Geol.-Mijnb. 

Genoot. Nederland en Kolonien Verb., geol. ser., v. 9, 

p. 283, 284, pi. 4, figs. 1-3; pi. 6, fig. 12 [references].

1932. Camerina semiglobula Doornink, Geol.-Mijnb. Genoot. 

Nederland en Kolonien Verb., p. 292-295, pi. 7, figs. 

1-14; text figs, d, e.

1932. Camerina pengaronensis (Verbeek). Whipj^e, in Hoffmeister, Bernice P. Bishop Mus. Bull. 96, p. 80.

1953. Camerina saipanensis Cole, in Cole and Bridge, U.S. Geol. 

Survey Prof. Paper 253, p. 20, 21, pi. 2, figs. 7-19.

1957. Camerina saipanensis Cole. Cole, U.S. Geol. Survey Prof. 

Paper 280-1, p. 330, pi. 102, fig. 20.

1957. NummuUtes striatus Hanzawa, Geol. Soc. America Mem. 

66, p. 41, 42, pi. 1, figs. 1-3, 9-11 [not Camerina striata 

Bruguiere, 1792].

1957. NummuUtes pengaronensis (Verbeek). HanzaATra, Geol. Soc. 

America Mem. 66, p. 43, pi. 1, fig. 8.

1959. Camerina pengaronensis (Verbeek). Cole, U.S. Geol. Survey 

Prof. Paper 260-V, p. 753, 754, pi. 231, figs. 1-17 [imprint date 1957].

1963. Camerina pengaronensis (Verbeek). Cole, U.S. Geol. Survey 

Prof. Paper 403-E, p. E3.
LATE EOCENE FOSSILS FROM EUA, TONGA B5

As the specimens in this collection are identical with 

those illustrated and described from Saipan Island 

(Cole, in Cole and Bridge, 1953, p. 20) and Eniwetok 

Atoll drill hole F-l, particularly from core 13 at 4,406-

4,431 feet, the specimens are illustrated, but not 

described.

However, certain measurements made from thin sections of these specimens are given.

Measurements of sections of Camerina pengaronensis (Verbeek)

Specimen shown on pi. 3 

Fig. 10 Fig. 16 Fig. 11 Fig. 15 Fig. 14 Fig. 13

Height..................................

Width..................................

Thickness...... ........

Embryonic chambers:

...mm. 2.6

2.45

110x160

70x160

190

4.75

8

24

83

3.0

2.7 ...

90x110 ...

80x170 ...

180 ...

6.125 ...

9 ...

23 ...

108 ...

2. 45 1. 85

1.4 1.2

......... 130x160 ...

.......... 80x160 ...

.......... 220 ...

1,115 1,000

2.5 3.45

1.71 2.45

.......... .......200

1,400 1,000

Occurrence elsewhere. Saipan Island (Cole, 1957a, p. 

330); Eniwetok Atoll drill hole F-l (Cole 1957b, p. 

753); Guam Island (Cole, 1963, p. E3); Viti Levu, Fiji 

(Cole, 1960a, p. A4); Java (Doornink, 1932, table 2, 

p. 306, 307); Borneo (van der Vlerk, 1929, p. 20).

Remarks. Doornink (1932, p. 283) and Cole (1957b, 

p. 753) have discussed this species in detail. Sen Gupta 

(1965, p. 91) placed Camerina pengaronensis (Verbeek) 

in the synonymy of Nummulites beaumonti d'Archiac 

and Haime. This Lutetian species is much larger than, 

and internally has only a superficial resemblance to, C. 

pengaronensis particularly in transverse sections. Because of the stratigraphic occurrence of C. beaumonti in 

the Lutetian of India, it is more logical to consider C. 

beaumonti as the ancestor of the late Eocene C. pengaronensis than to combine these species.

Nagappa (1959, pi. 10, fig. 5) has figured specimens 

from the Kopili stage, upper Eocene, Jamtia Hills, 

Assam, which are without question C. pengaronensis. 

His illustrations compare favorably in number of coils, 

size, and number of chambers with illustrations of 

specimens of C. pengaronensis from Saipan Island and 

Eniwetok Atoll drill hole F-l.

The data available suggest that C. beaumonti is not 

only a stratigraphically older species than C. pengaronensis, but also that the two species can be distinguished 

by differences in size, number of whorls, number of 

chambers, and other structures, such as the internal 

striated axial plug in C. pengaronensis.

Genus PELLATISPIRA Boussac, 1906

1906. Pellatispira Boussac, Soc. Geol. France Bull., 4th ser.,

v. 6, p. 91 [type: P. douvillei Boussac = Nummulites

madaraszi Hantken, 1876]. 

1936. Heterospira Umbgrove, Leidsche Geol. Mededel., v. 8,

no. 1, p. 156 [type: Heterospira mirabilis Umbgrove,

1936].

1937. Biplanispira Umbgrove, Leidsche Geol. Mededel., v. 8, 

no. 2, p. 309 (name substituted for Heterospira Umbgrove, 1936, not Heterospira Koken, 1896).

The structure of the test of the genus Biplanispira is 

the same as that of Pellatispira, except the primary 

revolving spire of chambers in Biplanispira becomes a 

double row of chamber-lets on each side of the marginal 

cord (see Cole, in Cole and Bridge, 1953, pi. 6, figs. 0-19; 

Cole, 1957a, pi. 99, figs. 1-6, pi. 100, figs 1-3). In 

Pellatispira the primary revolving spire of chambers is 

continuous to the periphery of the test (pi. 1, fig. 7; Cole, 

1957a, pi. 96, all figs.). Thus, in specimens of Pellatispira 

the marginal cord occupies the area between the chambers of the spire, whereas in specimens of Biplanispira 

the marginal cord is continuous and has bilaterally 

symmetrically arranged chamberlets on each side of it.

In certain specimens assigned to Pellatispira (pi. 2, 

figs. 20, 23; pi. 1, figs. 3, 8), the marginal cord penetrates deeply into the proximal side of some of the 

spiral chambers, and in a few specimens (Cole, 1057a, 

pi. 97, fig. 1) the marginal cord completely bisects 

the spiral chamber so that two chamberlets are produced. This same kind of arrangement of chamberlets 

and chambers is shown on the lower end of the specimen illustrated as figure 10, plate 1.

Some specimens assigned to Biplanispira have a 

sequence of small supplemental chambers at the 

periphery of the test (Cole, 1957a, pi. 99, fig. 2). This 

feature is found in specimens of Pellatispira (pi. 1, 

fig. 8, upper left side; fig. 10, top) and in specimens 

assigned to Biplanispira absurda (Umbgrove, 1938, 

fig. 15).

Inasmuch as the basic structures and arrangement 

of these structural elements in the test are so similar 

in specimens which have been separated previously 

into two genera, it seems best to consolidate these two
B6 LARGER FORAMINIFERA

genera. A description of the redefined genus Pellatispira 

(synonym: Biplanispira} follows: Test discoidal, or 

lenticular, often bilaterally symmetrical; embryonic 

chambers bilocular, large in megalospheric specimens; 

embryonic chambers followed by an evolute planispiral 

coil of median chambers in which the coils are separated from each other by an expanded marginal cord, 

or by an initial short coil of median chambers 

beyond which there are two layers of closely coiled 

chamberlets which are separated by an intervening 

sheetlike marginal cord; covering wall on each side of 

the median zone is composed of many pillars between 

which are outwardly radiating coarse canals.

Available data suggest that certain specimens which 

previously have been referred to Biplanispira mirabilis 

(Umbgrove) represent a valid species. However, these 

specimens should be referred to the redefined genus 

Pellatispira.

Biplanispira has the same geographic distribution 

and stratigraphic range in the Indo-Pacific region as 

Pellatispira. Although the proposed consolidation of 

these two genera expresses best the biologic relationship 

of the species, this proposal does not affect either the 

stratigraphic range or geographic distribution of the 

species, all of which would be assigned to Pellatispira.

In the Indo-Pacific region, three species of Pellatispira 

seemingly are valid. They are: P. madaraszi (Hantken), 

P. hofmeisteri Whipple, and P. mirabilis (Umbgrove). 

A detailed analysis of P. madaraszi and its synonyms 

follows.

Pellatispira madaraszi (Hantken) 1 

Plate 1, figures 1-13; plate 2, figures 19-23

1876. Nummulites madaraszi Hantken, Kgl. Ungarischen Geol.-

anst. Mitt. Jahrb., v. 4, p. 86, pi. 16, figs. 7a-c [imprint

date 1875]. 

1906. Pellatispira douvillei Boussac, Soc. Ge"ol. France Bull.,

4th ser., v. 6, p. 91, 92, pi. 2, figs. 10-13. 

1906. Pellatispira madaraszi (Hantken). Boussac, Soc. Ge"ol.

France Bull., 4th ser., v. 6, p. 92, 93, pi. 2, fig. 14. 

1909. Assilina madaraszi (Hantken). Provale, Riv. Italiana

Paleontologia, v. 14, p. 66-70, pi. 4, figs. 21-24; pi. 5,

figs. 1-4 [imprint date 1908]. 

1909. Assilina madaraszi orbitoidea Provale, Riv. Italiana

Paleontologia, v. 14, p. 71, pi. 5, fig. 5 [imprint date

1908]. 

1921. Pellatispira madaraszi von Hantken var. provalei Yabe,

Tohoku Imp. Univ. Sci. Repts., 2d ser. (geology), v. 4,

no. 4, p. 108. 

1928. Pellatispira madaraszi von Hantken var. provalei Yabe.

Umbgrove, Dienst Mijnb. in Nederlandsch-Indie,

Nederlandsche Akad. Wetensch. Mededel., no. 10,

p. 17, 18, figs. 27-33.

1 Pellatispira fulgeria Whipple (synonym: Biplanispira absurda Umbgrove) is 

considered to be an aberrant form of Pellatispira madaraszi. P. fulgeria is discussed 

separately after typical specimens of P. madaraszi are analyzed.

1928. Pellalispira orbitoidea (Provale). Umbgrove, Dienst Mijnb.

in Nederlandsch-Indie, Nederlandsche Akr.d. Wetensch.

Mededel., no. 10, p. 18, 19, figs. 2, 3, 5, 7, 9, 11-26,

34-41.

192S. Pellatispira rutteni Umbgrove, Dienst Mijrb. in Nederlandsch-Indie, Nederlandsche Akad. Wetensch.

Mededel., no. 10, p. 20, 21, figs. 57-61.

1028. Pellatispira inflata Umbgrove, Dienst Mijrb. in Nederlandsch-Indie, Nederlandsche Akad. Wetensch.

Mededel., no. 10, p. 21, figs. 42-56.

192S. Pellatispira glabra Umbgrove. Dienst Mijnb. in Nederlandsch-Indie, Nederlandsche Akad. Wetensch. Mededel., no. 10, p. 22, figs. 62-68. 

1928. Pellatispira irregularis Umbgrove, Dienst Mijnb. in

Nederlandsch-Indie, Nederlandsche Akad. Wetensch.

Mededel., no. 10, p. 23, 24, figs. 69-74. 

1928. Pellatispira crassicolumnata Umbgrove, E : enst Mijnb.

in Nederlandsch-Indie, Nederlandsche Akad. Wetensch.

Mededel., no. 10, p. 24, 25, figs. 75-80. 

1932. Pellatispira rutteni Umbgrove. Whipple, in Hoffmeister,

Bernice P. Bishop Mus. Bull. 96, p. 81, pi. 21, fig. 6. 

1941. Pellatispira madaraszi (Hantken) var. indicct. Rao, Mysore

Univ. Jour., sec. B (sci.), v. 2, pt. 2, p. 13, 14, pi. 1,

figs. 2, 5; pi. 2, fig. 3. 

1941. Pellatispira orbitoidea (Provale). Rao, Mysore Univ.

Jour., sec. B (sci.), v. 2, pt. 2, p. 14, 15, pi. 1, figs. 1, 3,

4, 6, 8; pi. 2, fig. 4. 

1941. Pellatispira inflata Umbgrove var. minor Rao, Mysore

Univ. Jour., sec. B (sci.), v. 2, pt. 2, p. 15, figs. 3, 7. 

1947. Pellatispira reticularis Hanzawa, Japanese Jour. Geology

and Geography, v. 20, nos. 2-4, p. 59, 60, pi. 14, figs.

1-3; pi. 15, fig. 1. 

1957. Pellatispira orbitoidea (Provale). Cole, U.S. Geol. Survey

Prof. Paper 280-1, p. 333, pi. 96, figs. 3-5, 7-9; pi. 97,

figs. 1-12; pi. 99, figs. 7-11. 

1957. Pellatispira provaleae Yabe. Cole, U.S. Geol. Survey

Prof. Paper 280-1, p. 333, pi. 96, figs. 1, 2, 6; pi. 98,

figs. 1-12.

An attempt will be made to demonstrate that large, 

distinctive specimens which previously have been identified either as Biplanispira absurda Umbgrove, 1938, 

or as Pellatispira fulgeria Whipple (in Hoffmeister, 

1932) are aberrant specimens of Pellatispira madaraszi 

(Hantken) or, in some cases, that they may be exceptional specimens of Pellatispira mirabilis (Umbgrove). 

For convenience, P. madaraszi will be discussed first, 

followed by an analysis of P. fulgeria of which B. 

absurda is a synonym (Cole, 1957a, p. 338). Separate 

synonymies are given.

Remarks. -In 1908, Provale (p. 66, pi. 5, figs. 1-4) 

identified specimens from the upper Eocene of Borneo 

as Assilina madaraszi (Hantken), a European species. 

These specimens are compressed, highly ornamented, 

and have a distinct spiral surface depression which 

reflects the internal coil of median chambers. Associated with these specimens were others (Provale, 

1908, pi. 5, fig. 5) which are inflated and evenly biconvex and have the surface covered by distinct 

papillae. Provale (1908, p. 71) designated this kind 

"Assilina madaraszi var. orbitoidea n. f."
LATE EOCENE FOSSILS FROM EUA, TONGA B7

Yabe (1921, p. 106-108) figured specimens of Pellatispira from the upper Eocene of Hillsborough Island 

(Bonin Island) and Ishigaki-jima, Ryukyu Islands. He 

stated (Yabe, 1921, p. 108): "The Bornean form which 

Provale called P. madaraszi is distinguished from the 

type of the above named species and its variety douvillei 

by the more densely pustulated surface of the shell, 

and may conveniently be distinguished as P. madaraszi 

var. provalei." In this same article Yabe (1921, p. 108) 

referred to "Assilina" madaraszi var. orbitoidea Provale as Pellatispira orbitoideus (Provale).

Umbgrove (1928) studied specimens of Pellatispira 

from Java, Borneo, and Celebes. He accepted Yabes' 

definition of P. madaraszi provalei and P. orbitoidea 

(Provale). In addition Umbgrove (1928) distinguished 

five new species: P. rutteni, P. inflata, P. glabra, P. 

irregularis, and P. crassicolumnata. Hanzawa (1947, 

p. 59) named specimens from Nakanai, New Britain, as 

P. reticularis, a species which Cole (in Cole and Bridge, 

1953, p. 22) recognized as a synonym of P. rutteni. 

Umbgrove, an opinion with which Hanzawa (1957, 

p. 53) later agreed.

In his initial study of the larger Foraminifera of 

Saipan Island, Cole (in Cole and Bridge, 1953, p. 21, 22) 

identified two species of Pellatispira P. crassicolumnata 

Umbgrove and P. rutteni Umbgrove. As more specimens 

from Saipan Island became available, Cole (1957a, 

p. 333) revised his concept of certain species of Pellatispira. In addition to P. reticularis Hanzawa, heassigned P. rutteni Umbgrove and P. inflata Umbgrove 

to the synonymy of Pellatispira orbitoidea (Provale). 

Cole (1957a, p. 333) wrote: "The name P. orbitoidea 

has been given to compressed lenticular individuals. 

More inflated individuals have been called P. inflata, 

and larger indiv duals with an umbonate central area 

which is surrounded by a rim have been termed P. 

rutteni. All of these form an integrated series and are 

grouped together."

At the same time Cole (1957a, p. 333) recognized P. 

provaleae Yabe as a distinct species. He wrote: "P. 

promleae differs from P. orbitoidea in possessing pillars 

of two different sizes, the larger of which project irregularly above the surface of the test." However, Cole 

recognized that P. crassicolumnata Umbgrove was a 

synonym of P. provaleae Yabe.

Thus, by 1957, eight species of Pellatispira had been 

recognized in the Indo-Pacific upper Eocene. Of these, 

P. inflata, P. reticularis, and P. rutteni had been assigned 

to the synonymy of P. orbitoidea, and P. crassicolumnata 

had been placed in the synonymy of P. provaleae.

In the matrix-free larger Foraminifera fauna from 

Eua, Tonga, specimens of Pellatispira are the most 

abundant element. The majority of these specimens 

could be identified readily as P. orbitoidea (Provale).

Another group of specimens could be recognized as 

P. provaleae Yabe. However, specimens occurred which 

were intermediate in all respects between these two 

supposedly valid species.

The following comparison should be made between 

the specimens illustrated on plate 1 (this report) from 

Eua, Tonga, and Umbgrove's illustrations (1928).

Eua, Tonga, specimens, Compare with Umbgrove's Umbgrove's identification 

pi. 1, fig. (1928) illustration, fig. 

1...-__----_--_-.-_------- 33--...-._--.-___.___-..--- P. madaraszi provaleae.

2-----_--------_---_-_._---- 76, 78__................. crassicolumnata.

3, 4.._.__.____........ 57, 59___....___.__. rutteni.

6----------...-----------_ 37-._...................... orbitoidea.

9------.--.-- --- _-.- 54, 56-. .-.------. .-- inflata.

Comparison of Umbgrove's illustrations (1928, figs. 

50-56) of P. inflata with his illustrations of P. glabra 

(1928, figs. 62-65) will demonstrate that these specimens are similar, particularly when one notes that 

the illustrations of P. inflata are from oriented thin 

sections whereas most of those of P. glabra are from 

oblique thin sections.

All the supposedly valid species have similar median 

sections. The differences upon which the various- species have been based are observed in the external 

ornamentation and transverse sections. Yet, when 

many specimens and thin sections are examined, a 

completely gradational series is found in a single population. Therefore, the conclusion has been reached that 

only one species is represented. However, the question 

remains concerning the specific name which should 

be applied to this species.

Rao (1941) discussed and illustrated specimens of 

Pellatispira from near Surat and Broach, western 

India. He identified P. orbitoidea (Provale) and 

described P. madaraszi indica Rao and P. inflata 

minor Rao as new varieties. The specimens which Rao 

illustrated can be duplicated by specimens ir the 

series from Eua, Tonga. Thus, the geographic range 

of this Indo-Pacific species, as defined in this report, 

can be extended to western India through Assam, 

where Nagappa (1959, pi. 11, figs. 6-12) found rneciinens of Pellatispira which he illustrated under the 

names P. glabra Umbgrove and P. infiata Umbgrove.

Grimsdale (1952, p. 225) reported that the European 

species P. madaraszi (Hantken) occurs in the upper 

Eocene of Kirkuk, Iraq. Aubouin and Neumann (1959, 

pi. 1, figs. 1, 2) identified specimens from Greece as 

P. madaraszi indica Rao.

Although more records would be desirable, the 

localities at which Pellatispira has been found suggest 

a continuous distribution of this genus in the upper 

Eocene from southern Europe to the central Pacific 

at Eniwetok Atoll (Cole, 1957b, p. 765).
B8 LARGER FORAMINIFERA

Boussac (1906, p. 91) based the genus Pellatispira 

on a new species, P. douvillei, which he described from 

the Priabonian (upper Eocene) of Italy, and assigned 

Nummulites madaraszi Hantken to this new genus.

o

Later authors (see summary by Roveda, 1961, p. 205, 

206) considered that P. douvillei Boussac (1906) was 

a synonym of P. madaraszi (Hantken, 1876).

The illustrations of European specimens (Boussac, 

1906, pi. 2, figs. 10-14; Tobler, 1929, pi. 17, figs. 

1-4; Roveda, 1961, pi. 16, figs. 1-5; pi. 17, figs. 4-6) 

of P. madaraszi are similar to Indo-Pacific specimens 

which have been identified previously as P. provaleae 

Yabe. Detailed study has convinced me that P. provaleae is a synonym of P. madaraszi and that Pro vale 

(1908, p. 66) was correct when she assigned specimens 

from Borneo to the European species.

P. provaleae is without question a synonym of 

P. madaraszi. The series of specimens from Saipan Island (Cole, 1957a, p. 333) and Eua, Tonga (pi. 1, 

figs. 1-9), show complete gradation from the P. madaraszi kind (compressed, highly ornamented) to the inflated kind (P. rutteni, P. orbitoidea, and P. infata). 

Although the Indo-Pacific specimens from a given locality have a variety of shapes and sizes, illustrations of the 

European specimens seem to indicate that only the 

compressed, highly ornamented kind is present.

Many specimens of Pellatispira from many localities 

on Saipan Island (Cole, 1957a, table 1) were analyzed. 

P. provaleae (based on thin, compressed specimens) 

was recorded from 33 localities. P. orbitoidea (based 

on inflated specimens) occurred at 23 localities at 

each of which P. provaleae also occurred.

At most localities where abundant specimens of 

Pellatispira are present, the specimens run the gamut 

of shape and size, whereas at localities that have fewer 

specimens, there is more consistency in shape and size. 

This relation suggests that in an extremely favorable 

ecological situation maximum diversity and development occurs, whereas other situations may impose limitations on shape and size. Nevertheless, all these 

specimens form a continuous, gradational series and 

thus represent only one species, P. madaraszi.

One specimen (pi. 1, fig. 5) that externally resembles 

the "inflata" kind of Umbgrove is interesting in that 

it has "twinned" embryonic chambers. It is the first 

specimen of this kind encountered.

A discussion of P. fulgeria Whipple, 1932, a synonym 

of P. madaraszi (Hantken) (1876) follows:

Plate 1, figures 10-13

1932. Pellatispira fulgeria Whipple, in Hoffmeister, Bernice P.

Bishop Mus. Bull. 96, p. 82, figs. 2, 3, 5-7. 

1938. Biplanispira absurda Umbgrove, Leidsche Geol. Mededel.,

v. 10, p. 82-89, figs. 1-17.

1957. Biplanispira fulgeria (Whipple). Cole, U.S. Geol. Survey

Prof. Paper 280-1, p. 333, pi. 98, figs. 13-1?, 

1960. Biplanispira fulgeria (Whipple). Cole, U.S. Geol. Survey 

Prof. Paper 374-A, p. A5, pi. 1, fig. 2; pi. 2, fig. 3.

Certain distinctive specimens (pi. l,figs. 11-13) have 

long been considered to represent a valid and easily 

recognized species. Umbgrove (1938) described similar 

specimens from the Eocene of Borneo as Biplanispira 

absurda. Earlier, Whipple (in Hoffmeister, 1932, p. 82 

pi. 20, figs. 2, 3, 5-7) had named similar specimens from, 

Eua, Tonga, Pellatispira fulgeria. Although Umbgrove 

(1938, p. 83) was familiar with Whipple's work, he did 

not correlate his specimens with the species named by 

Whipple. Cole (1957a, p. 333) wrote: "Comparison of 

Umbgrove's illustrations of B. absurda with those 

given by Whipple of P. fulgeria indicates that only one 

species is represented."

All these assignments may be incorrect in view of 

the data which have accumulated since that time. 

Specimens of this kind may represent aberrant forms 

of P. mirabilis (Umbgrove) and P. madaraszi (Hantken). 

which resemble each other.

Umbgrove (1938, p. 84) listed the species which 

occurred with the type specimens of B. absurda. The 

lists include several species of Pellatispira as recognized 

by Umbgrove at that time, but he did not record 

11 Biplanispira" mirabilis in these samples. It is significant that Umbgrove (1938, p. 89) suggested that "* * * 

a few sections of foraminifera, which I described formerly, in 1928, as Pellatispira crassicolumnata, belong 

to the present species [B. absurda]."

Whipple (in Hoffmeister, 1932, table 5) found P. 

fulgeria at three localities on Eua, Tonga. Although 

Whipple found specimens of P. "rutteni" and P. hojfmeiMeri Whipple at several localities, he did not record 

or illustrate any specimens which might be identified 

as "Biplanispira" mirabilis.

The present collection from Eua, Tonga, contains 

abundant specimens of Pellatispira and about seven 

specimens that are identical with those identified by 

Whipple as Pellatispira fulgeria. These large, compressed, strongly papillate specimens having p,n inflated 

periphery are recognized readily, even before they are 

sectioned.

One specimen (pi. 1, fig. 10) selected for sectioning 

appeared externally to be a rather large specimen of P. 

madaraszi. However, the transverse thin section of this 

specimen has internal structures which resemble both 

P. madaraszi and P. fulgeria. The central area resembles 

that of typical specimens of P. madaraszi (compare this 

area of fig. 10, pi. 1, with the specimens illustrated as 

figs. 2 and 4 of pi. 1). The peripheral zone of this specimen, however, is similar to that of P. fulgeria and
LATE EOCENE FOSSILS FROM EUA, TONGA B9

"Biplanispira" absurda (see Umbgrove, 1938, figs. 14, 

15).

The central area of P. Julgeria (pi. 1, fig. 13) resembles 

that of P. madaraszi (pi. 1, fig. 7), but beyond this central area in P. julgeria, the marginal cord is wide, and 

secondary chamberlets develop on each side of the 

marginal cord. In P. madaraszi (pi. 1, fig. 7), the primary coil of chambers extends to the periphery, and 

secondary chamberlets are developed only rarely.

The available data suggest that typical specimens of 

P. madaraszi can be interconnected with typical specimens of P. Julgeria through such specimens as the one 

illustrated as figure 10, plate 1. It should be emphasized 

that these specimens of P. Julgeria from Eua, Tonga, 

are associated with abundant specimens of P. madaraszi, and the specimens of "Biplanispira" absurda 

from Borneo are likewise associated with specimens of 

P. madaraszi. "Biplanispira" mirabilis was not recorded 

from the type locality of P. julgeria or of B. absurda.

Even if the suggestion that P.fulgeria is a synonym of 

P. madaraszi is not accepted, it should be obvious from 

the similarities in structures and from association that 

B. absurda is a synonym of P. Julgeria. Moreover, 

these specimens represent Pellatispira. Umbgrove 

(1938) incorrectly assigned B. absurda to Biplanispira 

because of the secondary chamberlets which develop.

The problem remains concerning the possibility that 

P. mirabilis may develop an aberrant form which resembles the one developed by P. madaraszi. Cole (1957b, 

p. 765; table 4, p. 749) reported the occurrence of P. 

Julgeria (rare) in association with abundant specimens 

of P. mirabilis in core 11 of Eniwetok Atoll drill hole 

F-l. P. mad>araszi was not found in this core.

Cole (1957a, table 1, p. 322) found specimens identified as "Biplanispira" Julgeria at 23 localities on 

Saipan Island. These specimens were associated at 15 

localities with "B." mirabilis, and at 10 of these 23 localities P. madaraszi occurred. "Biplanispira" Julgeria 

and "Biplanispira" mirabilis, without any specimens of 

Pellatispira, were found at nine localities. At one locality in the upper Eocene of Viti Levu, Fiji, Cole (1960a, 

p. A3) found "Biplanispira" mirabilis (common) 

associated with rare specimens of "Biplanispira" 

Julgeria.

These data prove that "Biplanispira" mirabilis and 

Pellatispira madaraszi occur together, and specimens 

identified as P. Julgeria and "Biplanispira" absurda 

occur in association with either one or both of these 

species.

The structure of the test of "Biplanispira" mirabilis 

is essentially the same as that of Pellatispira madaraszi. 

The major difference is that Pellatispira madaraszi has 

a coil of primary chambers (pi. 1, fig. 7) which is continuous to the periphery of the test. In Biplanispira 

mirabilis there is a short coil of primary chambers 

around the embryonic chambers after which two coils of 

chamberlets develop, one on either side of a central 

fibrous zone (marginal cord) (Cole, in Cole and Fridge, 

1953, pi. 6, figs. 9-19). Thus, aberrant growth in either 

P. madaraszi or "Biplanispira" mirabilis would result 

in specimens which would be so identical in internal 

structure that they could not be separated specifically. 

It is entirely possible that continued analysis of 

these forms will demonstrate that specimens assigned 

here to P. mirabilis can be intergrated into the P. 

madaraszi series. Certainly, there is evidence that the 

julgeria kind of specimen can develop either in the P. 

madaraszi or the P. mirabilis series.

Genus HETEROSTEGINA d'Orbigny, 1826

Heterostegina saipanensis Cole 

Plate 2, figure 15; plate 3, figures 4-6, 8, 9, 12

1953. Heterostegina saipanensis Cole, in Cole and Bridge, U.S.

Geol. Survey Prof. Paper 253, p. 23, 24, pi. 2, figs. 4, 6. 

1957. Heterostegina saipanensis Cole. Cole, U.S. Geol. Survey

Prof. Paper 280-1, p. 331, pi. 102, figs. 17-19. 

1959. Heterostegina saipanensis Cole. Cole, U.S. Geol. Survey

Prof. Paper 260-V, p. 760-762, pi. 234, figs. 13-24; pi.

235, figs. 1-13 [imprint date 1957].

Measurements of certain critical structures from 

thin sections of this species are given.

Measurements of sections of Heterostegina saipanensis Cole

Specimens shown on pi. 3 

Fig. 9 Fig. 12 Fig. 6 Fig. 4 Fig. 5

Height......... - ...-.....-..mm..

Width. .._ . _ _ -__mm__

Embryonic chambers:

2.7

2.1

80x70

30x150

120

2

2.5

3.5

3 2

70x70

40x120

120

1

2.5

4.0

2.7 -

70x70

40x120

130

1 .

3 .

3.3

1.0

90x95 ..

30x70 ..

130 -.

450

3.87

1.0

400

Remarks Whipple (in Hoffmeister, 1932, p. 83, pi. 20 

fig. 9) illustrated the central part of a Heterostegina 

without identifying this specimen as to species. Cole 

(in Cole and Bridge, 1953, p. 24), in naming specimens 

from Saipan Island as Heterostegina saipanensis, v^rote: 

"The median section resembles one illustrated by 

Whipple (1932, p. 83, pi. 20, fig. 9) from the Eocone of 

Eua, Tonga." Study of the specimens in the present 

collection proves that the specimens in the upper Eocene 

of Eua, Tonga, are indeed the same as the Saipan 

Island specimens.

362-463i
BIO LARGER FORAMINIFERA

Genus SPIROCLYPEUS Douville, 1905

Spiroclypeus vermicularis Tan

Plate 2, figures 6-12

1937. Spiroclypeus vermicularis Tan, De Ingenieur in Nederlandsche-Indie IV Mijnb. en Geol., v. 4, no. 10, p.

187-190, pi. 1, figs. 7, 8; pi. 2, figs. 6-10; pi. 3, figs. 13-

23; pi. 4, figs. 11-18. 

1953. Spiroclypeus sp. Cole, U.S. Geol. Survey Prof. Paper 253,

p. 18, pi. 14, fig. 7. 

1957. Spiroclypeus vermicularis Tan. Cole, U.S. Geol. Survey

Prof. Paper 280-1, p. 333, pi. 102, figs. 12-14. 

1957. Spiroclypeus vermicularis Tan. Cole, U.S. Geol. Survey

Prof. Paper 260-V, p. 764, pi. 238, figs. 1-6, 8-10, 11, 12. 

1960. Spiroclypeus vermicularis Tan. Cole, U.S. Geol. Survey

Prof. Paper 374-A, p. A4, A5, pi. 1, figs. 10-14.

1963. Spiroclypeus vermicularis Tan. Cole, U.S. Geol. Survey

Prof. Paper 403-E, p. E3, Ell. 

1965. Spiroclypeus vermicularis Tan. Adams, Geol. Soc. London

Quart. Jour., v. 121, p. 295, pi. 23, figs, u-w, aa.

The test is small and has an eccentric evenly biconvex 

umbonal area which is bordered particularly on the 

distal margin by a thin flat rim. Some specimens have 

prominent umbonal bosses (pi. 2, fig. 6) ancf small papillae scattered over the remainder of the umbonal area. 

Other specimens lack the umbonal bosses but have 

small papillae over the entire umbonal arer.

Measurements of the internal features follow.

Measurements of sections of Spiroclypeus vermicularis Tan

Specimen shown on pi. 2 

Fig. 10 Fig. 11 Fig. 12 Fig. 8 Fig. 7 Fig. 6 Fig.'

Width... .................... .mm_-

Thickness........ _ ......... mm. ...

Embryonic chambers: 

Diameters of initial chamber.. n_.

Number of operculine chambers __

Number of volutions.. __ . __ . ...

Lateral chambers: 

Length---.-.... __ . _ ... _ .it....

Height.....-........ ....... _»__._

Thickness of spiral sheet __ ... p. .

2.2

2.1

130x140

60x160

1

o

2.4

2.1

110x100

70x180

1

2.125

2.8

2.3 .

110x120 ..

70x190 ..

200

1 .-

2,5 ..

2.4

.65

130

50

10

3

50

50-200

1.45

.62

30-100

10

4

40-60

30-100

2.85

1.0

50-120

10-20

4

30-100

100-350

2.0

.7

100x120

70x80

200

50-160

10

4

50-60

60-120

Remarks. Although Spiroclypeus vermicularis n external view resembles certain specimens of Heterostegina 

saipanenis Cole, the two species can normally be separated by the numerous small papillae on the surface of 

S. vermicularis and the absence of these on the surface of 

H. saipanensis.

The types are from Tertiary b (upper Eocene) sediments at Koetai, East Borneo. This species is widely 

distributed in the Indo-Pacific area; it has been reported from Saipan Island, Guam Island, Eniwetok Atoll 

drill hole F-l, Viti Levu, Fiji, and Sarawak.

The material from Eua, Tonga, contained only a few 

specimens of this species. Although these specimens 

are typical, they are small. The infrequent occurrence 

and the small size of the specimens suggests that the 

ecological situation was not optimum for their 

development.

Family DISCOCYCLINIDAE

Genus DISCOCYCLINA Giimbel, 1870

Subgenus DISCOCYCLINA Giimbel, 1870

Discocyclina (Discocyclina) sp.

Plate 2, figure 18

Remarks. The specimen illustrated as figure 18, 

plate 2, has a diameter of 1.3 mm and so far as could

be ascertained seems to be a representative of the 

genus Discocyclina .§. s. This was the only specimen 

of this kind found.

The embryonic chambers are unusual as? there are 

three nearly globular, concentric chambers. The innermost chamber has an internal diameter of about 100 ju 

and a wall about 10 ju thick. This inner chamber is 

encircled by a slightly larger chamber that has a 

slightly thicker wall. The distance between these two 

chambers is about 10 M- These two chambers are enclosed by a larger chamber that has an internal 

diameter of about 200 ju and a wall about 20 ju thick. 

The distance between the outermost chamber wall and 

the outer wall of the middle chamber is about 30 p. 

This interval appears to be subdivided into small rectangular chambers.

The equatorial chambers are arranged around the 

embryonic chambers in fairly regular concentric rows. 

These chambers are either rectangular or faintly hexagonal in shape.

The development of the embryonic apparatus 

indicates that this is an abnormal specimen. Generally 

it is impossible even with a single normal specimen 

to identify such a specimen specifically.
LATE EOCENE FOSSILS FROM EUA, TONGA Bll

Genus ASTEROCYCIINA Giimbel, 1870

Three species of Asterocyclina are recognized. These 

species are readily recognized in equatorial section by 

the marked differences in the embryonic apparatuses 

(pi. 2, figs. 13, 24, 25) and in vertical section by the 

arrangement of the lateral chambers. Although many 

specimens were sectioned, it is impossible to arrange 

these specimens in a continuous gradational series as 

could be done with the variable sequence of specimens 

assigned to Pellatispira madaraszi (Hantken).

Asterocyclina matanzensis Cole

Plate 2, figures 14, 24; plate 3, figures 1-3, 7; plate 4, figures 2-4, 

6, 8-10, 12

1932. Discocyclina (Asterocyclina) stellata (d'Archiac). Whipple, 

in Hoffmeister, Bernice P. Bishop Mus. Bull. 96, p. 84, 

85, pi. 21, figs. 1, 2; pi. 22, figs. 6, 7 (not Cakarina 

stellata d'Archiac, 1846).

1957. Asterocyclina matanzensis Cole, U.S. Geol. Survey Prof. 

Paper 280-1, p. 350, pi. 117, figs. 6-10; pi. 118, figs. 9-18.

1959. Asterocyclina matanzensis Cole. Cole, U.S. Geol. Survey 

Prof. Paper 260-V, p. 777, 778, pi. 249, figs. 1-17 

[imprint date 1957].

1960. Asterocyclina matanzensis Cole. Cole, U.S. Geol. Survey 

Prof. Paper 374-A, p. A6, pi. 1, figs. 4-9.

1963. Asterocyclina matanzensis Cole. Cole, U.S. Geol. Survey 

Prof. Paper 403-E, p. E3.

1965. Asterocyclina matanzensis Cole. Samanta, Cushman Found. 

Forarn. Research Contr., v. 16. pt. 1, p. 23-25, pi. 1, 

figs. 1-12.

1967. Asterocyclina matanzensis Cole. Cole, Bulls. Arn. Paleontology, v. 52, no. 233, p. 9, pi. 1, fig. 17; pi. 2, fig. 7.

External views of two specimens are illustrated (pi. 4, 

figs. 3, 4). The specimen illustrated as figure 4, plate 4, 

is similar to the specimens illustrated by Whipple (in 

Hoffmeister, 1932, pi. 21, figs. 2, 3). Whipple's specimens have diameters slightly over 3 mm. This specimen 

(pi. 4, fig. 4) has a diameter of about 3.2 mm and an 

unbonate area having a diameter of about 2.0 mm. 

One specimen (pi. 4, fig. 3) has long slender tapering 

rays, the longest of which is 1.7 mm, extending from 

a nearly globular central area that has a diameter of 

about 1.9 mm. A more general description of the shape 

of the test follows.

The test is composed of an umbonal area which may 

be nearly globular (pi. 3, fig. 2) and surrounded by a 

narrow rim or by long slender rays (pi. 4, fig. 3). Other 

specimens (pi. 4, figs. 4, 9) have a more compressed 

umbonal area which merges into four or five wide 

compressed rays. The umbonal area and the rays in 

most of the specimens are covered by small papillae.

The embryonic apparatus consists of a nearly circular 

initial chamber which is slightly embraced by the larger 

second chamber (pi. 3, fig. 1, 3; pi. 4, figs. 8, 10, 12). 

Two distinct periembryonic chambers are located at 

each end of the dividing wall between the embryonic 

chambers. The two periembryonic chambers are

separated from each other by two or more normal 

equatorial chambers which border the initial embryonic 

chamber between the principal embryonic che.mbers. 

The second embryonic chamber has on its margin 

three to five or more, distinct periembryonic chambers 

which are only slightly smaller than the two principal 

embryonic chambers (pi. 4, fig. 10). Certain of these 

periembryonic chambers are in contact; others are 

separated by one or more normal equatorial chambers. 

Measurements of equatorial sections follow.

Measurements of equatorial sections of Asterocyclin?, 

matanzensis Cole

Specimen shown on 

Diameter.... __-_--_-______.__mm_- 

Embryonic chambers: 

Diameters of initial chamber. _ju._ 

Diameters of second chamber. _ju_. 

Distance across both chambers. M--

Equatorial chambers: 

In rays:

In interrays:

PI.

Fig. 1

3.75

90x100 

100x190 

200 

10

30-60

25-30

25-30

20-25

3

Fig. 3

2.85

80x100 

90x185 

190

8

30-50

20-30

30-50

15-20

PI.

Fig. 8

2.3

90x110 

90x200 

200 

10

30-40

10-20

20-30

10-20

4

Fig. 12

3.6

100x130 

120x210 

230 

10

30-50

20-30

20-30

15-20

The lateral chambers are arranged in regular tiers. 

The openings of the lateral chambers are low, and the 

thickness of the intervening floors and roofs is about 

equal to the height of the lateral chamber cavity, lender 

pillars are irregularly scattered throughout the umbonate part of vertical sections.

Measurements of vertical sections follow.

Measurements of vertical sections of Asterocyclina matanzensis

Cole

Specimen shown on 

PI. 3 PI. 4

Fig. 2 Fig. 7 Fig. 2 Fig. 9 Fig. 6

Diameter of umbo.___ ........

Thickness...... ................

Embryonic chambers:

Length. ........_.............

Height. ......................

Thickness of outer wall _ .....

Equatorial layer:

Height at center... ............

Height at periphery. ..........

Lateral chambers:

Number. ... ...................

Length. .......................

Height......-..,-...---..-.--.

Thickness of roofs and floors.-.

Surface diameter of pillars. _.-__.

......mm..

. mm -

--.------jti--

........ -ft..

....... _-M--

.-..-_. .-/*.-

..--__--./*--

..--._---/*-.

....... ..ft..

........ -M--

1.6

.5 .

2.0

90

80

10

30

50

37

40-100

10-20

10

40-100

2.25

1.14

170

90

10

40

80

25

60-80

10-15

10

60-90

1.3

.35

.65

130

50

10

40

30

14

50-70

10

10

50

1.3

.5 .

.7o

12(1

80

10

30

90

14

40-60

10

10

30-50

2.85

1.5

80

60

15

30

50

36

70-80

10

10

50

Occurrence elsewhere. Saipan Island (Cole, 1957a, 

p. 350); Eniwetok Atoll drill holes (Cole, 1957b, p. 777); 

Guam Island (Cole, 1963, p. E3); Viti Levu, Fiji (Cole, 

1960a, p. A6); Assam (Samanta, 1965, p. 23).

Remarks. Whipple (in Hoffmeister, 1932, p. 84) 

assigned similar specimens from Eua, Tonga, to Discocyclina (Asterocyclina) stellata (d'Archiac). The illustra-
B12 LARGER FORAMINIFERA

tions, measurements, and description given by Whipple 

demonstrate conclusively that the specimens he identified as Asterocyclina stellata are the same as those 

which are identified here as A. matanzensis.

Concerning certain specimens Whipple (in Hoffmeister, 1932, p. 85) wrote: "At the proximal end of 

each ray there are two or three elongate equatorial 

chambers, this number increases toward the periphery 

producing a fan-like arrangement of the chambers within the ray." This unusual arrangement is shown excellently in the specimen illustrated as figure 6, plate 4. 

This same phenomenon is found in certain specimens 

(Cole, 1957a, pi. 118, fig. 17) from Saipan Island, the 

type locality of A. matanzensis.

Cole (1967, p. 9) stated previously that A. matanzensis has a "general resemblence" to A. stellata 

(d'Archiac) (Neumann, 1958, pi. 30, figs. 1-7) from the 

Lutetian and Bartonian of France. The lateral chambers 

of A. stellata have more open cavities than do those of 

A. matanzensis, and the ring of periembryonic chambers 

on the border of the second embryonic chamber in A. 

stellata is more complete and has more chambers than 

in A. matanzensis.

Relatively few adequate photographs of thin sections 

of A. stellata have been published. As additional illustrations become available, it may be possible to demonstrate satisfactorily that A. matanzensis is a junior 

synonym of A. stellata (d'Archiac). For the present, 

A. matanzensis is retained as a valid species, closely 

related to A. stellata.

Asterocyclina penuria Cole 

Plate 2, figures 16, 17, 25; plate 4, figures 1, 5, 7, 11, 13, 14

1957. Asterocyclina penuria Cole, U.S. Geol. Survey Prof. Paper 

280-1, p. 350, 351, pi. 116, figs. 1-10 [references and 

discussion]. [Nom. nov. for Orthophragmina pentagonalis 

Deprat, 1905, preoccupied by Asterodiscus pentagonalis 

Schafhault, 1863.]

1957. Discocyclina (Discocyclina) dispansa Hanzawa, not Lycophris dispansus Sowerby, 1840, Geol. Soc. America Mem. 

66, p. 83, 84, pi. 13, figs. 1, 3, 4; pi. 14, figs. 2, 3, 8, 9.

1958. Asterodiscus cuvillieri Neumann, Soc. Geol. France Mem. 

83 (new ser.), v. 37, pt. 2-3, p. 119-121, pi. 21, figs. 1-8; 

pi. 32, fig. 5; text fig. 40.

1959. Asterocyclina penuria Cole. Cole, U.S. Geol. Survey Prof.

Paper 260-V, p. 778-780, pi. 246, figs. 1-11; pi. 247,

figs. 1-15; pi. 248, figs. 8, 12-17 [imprint date 1957]. 

1959. Asterocyclina penuria Cole. Cole, Cushman Found. Foram.

Research Contr., v. 10, pt. 1, p. 12-14, pi. 1, figs. 1-4,

6-11; pi. 2, figs. 1-9; pi. 3, figs. 1-4. 

1963. Asterocyclina penuria Cole. Cole, U.S. Geol. Survey Prof.

Paper 403-E, p. E3.

The test has diameters of 2.3 to 7.0 mm and thicknesses of 1.2 to 2.7 mm. Specimens are usually strongly 

umbonate (pi. 4, figs. 5, 13), and the inflated central 

area is bordered by a narrow flange. Other specimens 

(pi. 4, fig. 11) are compressed and lenticular. The

central area does not show any rays. If rays are present, 

they are weakly developed on the bordering rim. Thus, 

it is often impossible to recognize that specimens are 

Asterocyclina until they are sectioned.

The embryonic apparatus consists of two large 

embryonic chambers in which the larger second chamber 

slightly embraces the initial chamber. The embryonic 

chambers are surrounded completely by large periembryonic chambers. The initial embryonic chamber is 

bordered by two elongate periembryonic chambers 

(pi. 4, figs. 7, 14; pi. 2, figs. 16, 25). In some specimens 

(pi. 4, fig. 14) the second embryonic chamber is surrounded by a ring of about nine rudely rectangular 

chambers. Other specimens (pi. 2, fig. 25) have the 

second embryonic chamber bounded by two periembryonic chambers which are similar to those bordering the initial embryonic chamber.

Measurements of equatorial sections

Measurements of equatorial sections of Asterocyclina penuria Cole

Specimen showr on

PI. 4 PI. 2

Fig. 1 Fig. 7 Fig. 14 Fig. 25 Fig. 16

Diameter._.._-_____-__._________ mm.. 4.8

Embryonic chambers: 

Diameters of initial chamber.... _M. . 230x300

Diameters of second chamber.. ___/*._ 180x440 

Distance across both chambers.. _ji.. 430 

Thickness of outer wall_ ........n.. 10

Equatorial chambers: 

In rays: 

Radial diameter.........._._...M.. 40-100

Tangential diameter............/*-- 20-30

In interrays: 

Radial diameter................/*.. 30-40

Tangential diameter..........._M.. 20-30

4.2 5.2 2.3

200x350 

230x470 

450 

10

240x340 

220x500 

470 

10

200x300 

180x390 

390 

10

230x300 

200x420 

430 

10

60-100 

20-40

40-60 

20-30

80-120 

15-20

40-50 

20-30

40-80 

20

60-100 

20

60-100 

30-40

30-50 

20-35

Normally, there are four rays developed in which 

the equatorial chambers are radially elongate. These 

rays are distinct in the zone adj acent to the embryonic 

apparatus but commonly are less distinct in the peripheral zone.

The lateral chambers are abundant, arranged in 

regular tiers, and have low openings between rather 

thick floors and roofs. Distinct pillars are irregularly 

scattered throughout the vertical sections, the greatest 

concentration being in the umbonal area.

Measurements of vertical sections follow.

Measurements of vertical sections of Asterocyclina penuria Cole

Specimen shown on pi. 4 

Fig. 5 Fig. 11 Fig. 13

Diameter.................................mm.. 4.45 3.5+ 4.85+

Thickness.._._._____--.____.--.--.._.-.-.mm.- 2.7 1.37 2.57

Embryonic chambers: 

Length........................... ...._.._.* -- 360 440 450

Height.. . . ... _--._.--_..-._._Ai-- 260 250 260

Thickness of outer wall................... _M-- 10 10 10

Equatorial layer: 

Height at center...... ._____.___._.__.___M-- 50 60 80

Height at periphery--.......---.--..----../!-- 30 70 80

Lateral chambers: 

Number............... .........-._... - 31 19 38

Length M - 100-140 80-100 100-140 

Height....................................M-- 20-30 1C-20 10-20

Thickness of floors and roofs.............._M-- 15-20 20 10-20

Surface diameter of thickened areas between the 

lateral chambers ........................ft.. 70-150 150-200 50-180
LATE EOCENE FOSSILS FROM ETTA, TONGA B13

Occurrence elsewhere. New Caledonia (Deprat, 1905, 

p. 507); Soemba (Caudri, 1934, p. 97); Saipan Island 

(Cole, 1957a, p. 350); Eniwetok Atoll drill hole (Cole, 

1957b, p. 778); Guam (Cole, 1963, p. E3); seamount 

near Tuamotu Islands (Cole, 1959, p. 42); France 

(Neumann, 1958, p. 119, as Asterodiscus cuvillieri); 

Greece (Aubouin and Neumann, 1959, p. 48, as Asterodiscus cuvillieri).

Remarks. This species is characterized by its distinctive embryonic apparatus and by the abundant, long, 

appressed lateral chambers arranged in regular tiers 

between irregularly scattered, pronounced pillars.

This species was described by Deprat (1905, p. 507) 

as Orthophragmina pentagonalis and was based upon 

specimens from New Caledonia. As the specific name 

0. pentagonalis had been used by Schafhault in 1863 

for a different European species, Cole (1957a, p. 350) 

proposed the specific name Asterocyclina penuria.

Caudri (1934, p. 97) identified specimens from 

Soemba, which are without question A. penuria, as A. 

aff. pentagonalis Deprat. Van der Weijden (1940, p. 38) 

assigned European specimens from the upper Lutetian 

and Auversian to Discocyclina (Isodiscodina) pentagonalis (Deprat),

Neumann (1958, p. 119) described Asterodiscus 

cuvillieri as a new species from the upper Lutetian, 

Auversian, and Bartonian of Europe. She included in 

this new species the European specimens which van cler 

Weijden had assigned to Asterocyclina pentagonalis 

(Deprat) (= A. penuria Cole). Later, Aubouin and 

Neumann (1959, p. 48) identified specimens from the 

upper Eocene and the upper Lutetian of Greece as 

Asterodiscus cuvillieri Neumann (= Asterocyclina penuria Cole).

The European specimens to which Neumann (1958) 

assigned the name Asterodiscus cuvillieri are identical 

externally and internally to the Indo-Pacific specimens 

which Cole (1957a) named Asterocyclina penuria; thus, 

Asterodiscus cuvillieri is a junior synonym of Asterocyclina penuria.

Whipple (in Hoffmeister, 1932, p. 84, pi. 22, figs. 1, 2) 

briefly discussed and illustrated specimens from several 

stations in Eua, Tonga, as Discocyclina (Discocyclina} 

sp. His equatorial section (1932, pi. 22, fig. 1) shows 

rays of the land which are characteristic of Asterocyclina 

penuria, and his vertical section (1932, pi. 22, fig. 2) 

has abundant aligned lateral chambers between irregularly developed pillars similar to those of A. penuria. 

Unfortunately, the embryonic chambers in these illustrations do not show. However, the other internal 

stuctures are so similar to those of A. penuria that one 

may correlate Whipple's Discocyclina (Discocyclina) sp. 

with A. penuria.

Asterocyclina praecipua Cole 

Plate 2, figure 13; plate 5, figures 1-21

1959. Asterocyclina praecipua Cole, U.S. Geol. Survey Prof.

Paper 260-V, p. 780, pi. 245, figs. 11, 12, 16, 18-20

[imprint date 1957]. 

1963. Asterocyclina praecipua Cole. Cole, U.S. Geol. Survey

Prof. Paper 403-E, p. E24, pi. 9, figs. 12, 13, 18.

Three external shapes are exhibited by specimens 

assigned to this species. Certain specimens are subcircular and have low rays, which are observed only 

on the margin of the test. In equatorial sections, such 

specimens normally possess five rays (pi. 5, fig. 20), 

and in vertical section (pi. 5, figs. 4, 21), the tests are 

compressed lenticular. Other specimens (pi. 5, fig:?, 11, 

19) are quadrate and have four short, bluntly rounded 

rays. These specimens in vertical section (pi. 5, figs. 1, 

2, 12, 13) are inflated and lenticular. Specimens of 

another group have four elongate rays (pi. 5, figs. 5, 9). 

In vertical section (pi. 5, figs. 6-8), a small infated 

umbonal area, from which the elongate rays project, is 

observed. The surface of all these specimens is covered 

by small distinct papillae.

The small bilocular embryonic chambers in many of 

these specimens were destroyed. However, several satisfactory sections (pi. 5, figs. 17, 18) were obtained. The 

periembryonic chambers are small and abundant and 

form a complete encircling ring around the embryonic 

chambers.

Measurements of some equatorial sections follow.

Measurements of equatorial sections of Asterocyclina prae^pua

Cole

Specimen shown on pi. 5 

Embryonic chambers: 

Diameters of initial chamber. . _ . _/j- .

Distance across both chambers.. ./j._

Equatorial chambers: 

In interray areas :

In rays :

Fig. 19

2.1

60x100 

70x120 

140 

5

15-40 

15-20

50-65 

20-25

Fig. 20

1.6

55x70 

40x90 

100 

5

20-40 

20

50 

20

Fig. 11

1.75

130

20-30 

20-40

40-100 

20-30

Fig. 9

3.65+

110 

5

30-40 

20

100-120 

20-30

Tig. 5

2.0

100 

5

20-40 

20

40-100 

10-20

The lateral chambers are somewhat regularly aligned 

in tiers. The floors and roofs of these chambers are 

about equal in thickness to the chamber cavity. Heavy 

distinct pillars are irregularly scattered throughout the 

vertical sections. Measurements of some vertical sections follow.
B14 LARGER FORAMINTFERA

Measurements of vertical sections of Asterocyclina 

praecipua Cole

Specimen shown on pi. 5 

Fig. 1 Fig. 13 Fig. 12 Fig. 7 Fig. 8 Fig. 21

1.7 

.75

1.8+ 1.9+ 4. 45 2. 9+ I.67 

.9 1.0 .85 .8 .45

90+

80

5

15

30

Diameter. ___________________ .mm

Thickness....................mm

Embryonic chambers: 

Length._..__..__..._______.___,.-- 140 110 150 90+ ....

Height---..-...-.............;.-- 90 90 110 40+ _____

Thicknessof outer wall...... __/*.. 10 8 5 ______________

Equatorial layer: 

Height at center........... .__/*.- 30 25 40 20 30

Height at periphery...........M ._ 40 100 50 80 60

Lateral chambers: 

Number.......................... 17 18 18 22 17 10

Length........................M-- 50-70 60-80 50-70 60-70 40-60 40-70

Height. .......................A.-- 10-20 10 10 10 10 10

Thickness of floors and roofs...M-- 10-20 10 10 10 10 10

Surface diameter of pillars..-_...M.-100-180 50-100 80-100 40-100 60-120 50-100

Remarks, The types are from Eniwetok Atoll drill 

hole F-1 in core 12 (4,316-4,341 ft). This species is characterized by the small bilocular embryonic chambers, 

the thick roofs and floors of the lateral chambers, and 

the many heavy pillars. Specimens in the collection 

from Eua, Tonga, have a greater diversity of external 

shape than the types, but the internal structures of all 

these specimens are similar.

A few microspheric specimens (pi. 5, figs. 10, 15, 16) 

were found. These specimens closely resemble microspheric specimens from Guam (Cole, 1963, pi. 9, fig. 

18), which were assigned to this species.

REFERENCES CITED

Asano, Kiyoshi, 1939, Limestones of the South Sea Islands under 

the Japanese Mandate: Yabe Jubilee pub. v. 1, p. 537-550, 

2 figs., 2 pis.

Aubouin, Jean, and Neumann, Madeleine, 1959, Contribution & 

J'etude stratigraphique et micropaleontologique de 1'Eocene 

en Grece: Rev. Micropaleontologie, v. 2, no. 1, p. 31-49, 

pis. 1-5, 8 text figs.

Boussac, Jean, 1906, Developpement et morphologic quelques 

Foraminiferes de Priabona: Soc. G&>1. France Bull., 4th 

ser., v. 6, p. 88-100. pis. 1-3.

Caudri, C. M. B., 1934, Tertiary deposits of Soemba: Amsterdam, Leiden Natl. Mus. Geol. Diss., p. 1-224, 5 pis., 21 

text figs., 3 maps.

Cole, W. S., 1950, Larger Foraminifera from the Palau Islands: 

U.S. Geol. Survey Prof. Paper 221-B, p. 21-31.

 1957a, Larger Foraminifera [of Saipan, Mariana Islands]: 

U.S. Geol. Survey Prof. Paper 280-1, p. 321-360., pis. 94-118, 

4 tables.

 1957b, Larger Foraminifera from Eniwetok drill holes: 

U.S. Geol. Survey Prof. Paper 260-V, p. 743-784, pis. 230- 

249, 1 text fig., 6 tables [1959].

 1958, Names of and variation in certain American larger 

Foraminifera, No. 1: Bulls. Am. Paleontology, v. 38, no. 170, 

p. 179-213, pis. 18-25.

 1959, Asterocyclina from a Pacific seamount; Cushman 

Found. Foram. Research Contr., v. 10, pt. 1, p. 10-14, pis. 1-3.

 1960a, Upper Eocene and Oligocene larger Foraminifera 

from Viti Levu, Fiji: U.S. Geol. Survey Prof. Paper 374-A, 

p. A1-A7, pis. 1-3, 1 text fig., 1 table.

 1960b, The genus Camerina: Bulls. Am. Paleontology,

v. 41, no. 190, p. 189-203, pis. 23-26.

 1961, An analysis of certain taxonomic problems in the 

larger Foraminifera: Bulls. Am. Paleontology, v. 43, no. 197, 

p. 373-407, pis. 28-39.

 1963, Tertiary larger Foraminifera from Guam: U.S. 

Geol. Survey Prof. Paper 403-E, p. E1-E28, pis. 1-11, 1 

text fig., 8 tables.

 1966, Additional comments on the foraminiferal genus 

Camerina: Bulls. Am. Paleontology, v. 50, no. 228, p. 229- 

265, pis. 20-27.

 1967, Additional data on New Zealand Asterocyclina

(Foraminifera): Bulls. Am. Paleontology, v. 52, no. 233,

p. 5-18, pis. 1, 2, 1 text fig., 2 tables. 

Cole, W. S., and Bridge, Josiah, 1953, Geology and larger

Foraminifera of Saipan Island: U.S. Geol. Survey Prof.

Paper 253, 45 p., 15 pis., 5 tables. 

Cushman, J. A., Todd, Ruth, and Post, Rita, 1954, Recent

Foraminifera of the Marshall Islands: U.S. Geol. Survey

Prof. Paper 260-H, p. 319-384, pis. 82-9?, 3 text figs.,

5 tables. 

Deprat, M. J., 1905, Les depots Eocenes Neo-C^ledoniens, leur

analogic avec ceux de la region de la sonde: Soc. G6ol.

France Bull., 4th ser., v. 5, p. 485-516, pis. 16-19. 

Doornink, H. W., 1932, Tertiary Nummulitidae from Java:

Geol.-Mijnb. Genoot. Nederland en Kolonien Verh., Geol.

ser., v. 9, p. 267-315, pis. 1-10, 2 tables, text figs. a-L 

Grimsdale, T. F., 1952, Cretaceous and Tertiary Foraminifera

from the Middle East: British Mus. (Nat. History) Bull.,

Geology, v. 1, no. 8, p. 223-247, pis. 20-25, 3 text figs. 

Guest, N. J., 1959, Geological mapping, Tonga, in Fiji Geol.

Survey Dept. Ann. Rept., Council Paper 17, p. 3. 

Hanzawa, Shoshiro, 1947, Note on an Eocene foraminiferal

limestone from New Britain: Japanese Jour. Geology and

Geography, v. 20, nos. 2-4, p. 59-61, pis. 14,15.

 1957, Cenozoic Foraminifera from Micronesia: Geol.

Soc. America Mem. 66, 163 p., 41 pis., 12 text figs., 7 tables. 

Hoffmeister, J. E., 1932, Geology of Eua, Tonga: Bernice P.

Bishop Mus. Bull. 96, 93 p., 22 pis., 6 text figs., 6 tables. 

Myers, E. H., 1943, Ecologic relationships of larger Foraminifera:

Natl, Research Council, Div. Geology and Geography Ann.

Rept., 1942-1943, Rept. of the committee on marine ecology

as related to paleontology, no. 3, p. 26-30. 

Nagappa, Yedatore, 1959, Foraminiferal biostratigraphy of

the Cretaceous-Eocene succession in the India-PakistanBurma region: Micropaleontology, v. 5, no. 2, p. 145-192,

pis. 1-11, 11 text figs., 9 tables. 

Neumann, Madeleine, 1958, Revision des Orbitoid6s du Cre"tace

et de 1'Eocene en Aquitaine occidentale: Soc. G6ol. France

Mem. 83, new ser., v. 37, pt. 2-3, 174 p., 36 pi -., 54 text figs. 

Norton, R. D., 1930, Ecologic relations of some Foraminifera:

Seripps Inst. Oceanography Bull., tech. ser., v. 2, no. 9,

p. 331-388, 6 tables, 3 text figs. 

Provale, Irene, 1908, Di alcune Nummulitine e Orbitoidine

dell'isola di Borneo: Riv. Italiana Paleontologia, v. 14, p.

55-80, pis. 4-6 [1909]. 

Rao, S. R. N., 1941, The Tertiary sequence near Surat and

Broach (western India), with description of Foraminifera

of the genus Pellatispira from the upper Eocene of this

region: Mysore Univ. Jour., sec. B (sci.), v. 2, pt. 2, p.

5-17, pis. 1, 2. 

Roveda, Vittorio, 1961, Contributo allo studio di alcuni

macroforaminiferi di Priabona: Riv. Italiana Paleontologia,

v. 67, no. 2, p. 153-224, pis. 14-19, 19 text fits., 1 table. 

Samanta, B. K., 1965, Asterocyclina from the up'ier Eocene of

Assam, India: Cushman Found. Foram. Research Contr.,

v. 16, pt. 1, p. 22-26, pi. 1.
LATE EOCENE FOSSILS FROM ETJA, TONGA P15

Sen Gupta, B. K., 1965, Morphology of some key species of 

Nummulites from the Indian Eocene: Jour. Paleontology, 

v. 39, no. 1, p. 86-96, pis. 15-17, 2 text figs.

Tobler, August, 1929, Pellatispira Lm Priabonien von Lenk 

(Berner Oberland): Eclogae Geol. Helvetiae, v. 22, no. 2, 

p. 173-175, pi. 17.

Umbgrove, J. H. F., 1928, Het genus Pellatispira in het Indopacifische gebied: Dienst Mijnb. in Nederlandsch-Indie, 

Nederlandsche Akad. Wetensch. Mededel., no. 10, p. 1-60, 

80 figs., 1 table.

 1938, A second species of Biplanispira from the Eocene

of Borneo: Leidsche Geol. Mededel., v. 10, p. 82-89, figs. 

1-17.

Vlerk, I. M. van der, 1929, Groote foraminiferen van >\ O. 

Borneo: Dienst Mijnb. in Nederlandsch-Indie, Nederlandsche Akad. Wetensch. Mededel., no. 9, p. 5-44, 51 figs.

Weijden, W. J. M. van der, 1940, Het genus Discocyclina en 

Europa: Leiden Univ. Diss., N. V. Leidsche Courant, 

116 p., 12 pis.

Yabe, Hisakatsu, 1921, Notes on some Eocene Foramini*era: 

Tohoku Imp. Univ. Sci. Repts., 2d ser, (geology), v. 4, 

no. 4, p. 97-108, pis. 16-20, 1 table.

PLATES 1-5
PLATE 1

FIGURES 1-13. Pellatispira madaraszi (Hantken) (p. B6). 

1-6, 8-12. Transverse sections.

1. Thin, compressed specimen; compare with figure 13, 

plate 2 of Boussac (1906); X 20; USNM 651045.

2. Specimen having slightly inflated umbonal area and

large pillars; X 20; USNM 651046. 

3, 4. Specimens having strongly inflated central areas; "P. 

rutteni" kind; compare with figures 57 and 59 of 

Umbgrove (1928); X 20; USNM 651047, 651048.

5. Highly inflated specimen having two sets of embryonic chambers; X 20; USNM 651049.

6. Evenly biconvex specimen; "P. orbitoidea" kind; 

compare with figure 36-38 of Umbgrove (1928); 

X 20; USNM 651050.

8. Compressed, lenticular specimen, intermediate in 

shape between figures 1 and 2, this plate; X 20; 

USNM 651052.

9. Subglobular specimen; "P. inflata" kind; compare 

with figures 54 and 56 of Umbgrove (1928); 

X 20; USNM 651053.

10. Specimen which has internal structures of both 

Pellatispira madaraszi (central area) and "P. 

fulgeria" (top and bottom); X 20; USNM 651054.

11. Typical "P. fulgeria"; compare with figure 5, plate 

20, of Whipple, in Hoffmeister, 1932, and with 

Biplanispira absurda, figures 2-15, of Umbgrove 

(1938); X 20; USNM 651055.

12. Slightly thicker and larger specimen than figure 11,

this plate; X 12.5; USNM 651056. 

7, 13. Median sections.

7. Typical median section; X 20; USNM 651051.

13. Median section of the "P. fulgeria" kind; note the

expanded marginal cord and the development of

secondary chamberlets beyond the zone of the

primary spiral; X 12.5; USNM 651057.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-B PLATE 1

10
PLATE 2

FIGURES 1-5. Camerina pacifica (Whipple) (p. B3).

1, 2. Median sections, centered, X 20, of megalospheric specimens; USNM 651058, 651059.

3. Median section, not centered, X 20, of a megalospheric specimen; USNM 651060.

4. Median section, centered, X 20, of a microspheric specimen; USNM 651061.

5. Transverse section, X 20, of a megalospheric specimen; USNM 651062. 

6-12. Spiroclypeus vermicularis Tan (p. BlO).

6. Transverse section, X 20, of a probable microspheric specimen; USNM 651063. 

7-9. Transverse sections, X 20, of megalospheric specimens; USNM 651064-651066. 

10, 12. Median sections, X 20, of megalospheric specimens; USNM 651067, 651069.

11. Part of a median section, X 40, of a megalospheric specimen to illustrate the embryonic chambers; USNM

651068. 

13. Asterocyclina praecipua Cole (p. B13).

Central part of an equatorial section, X 40, of a megalospheric specimen for comparison of the embryonic apparatus with those of A. matanzensis (fig. 24, this plate) and A. penuria (fig. 25, this plate); see figure 18, plate 5, 

for enlargement; USNM 651070. 

14, 24. Asterocyclina matanzensis Cole (p. Bll).

14. Transverse section, X 20; USNM 651071.

24. Part of an equatorial section, X 40, of a megalospheric specimen; see explanation of figure 13, this plate;

USNM 651081. 

15. Heterostegina saipanensis Cole (p. B9).

Transverse section, X 20; introduced for comparison with similar sections of Spiroclypeus vermicularis, figures

7-9, this plate; USNM 651072. 

16, 17, 25. Asterocyclina penuria Cole (p. B12).

16. Equatorial section, X 20, of a small specimen similar in size and external appearance to certain specimens 

of A. matanzensis; USNM 651073.

17. Vertical section, X 20, of a companion specimen to figure 16; USNM 651074.

25. Part of an equatorial section, X 40, of a megalospheric specimen; see explanation of figure 13, this plate;

USNM 651082. 

18. Discocyclina (Discocyclina) sp. (p. BIO)

Part of an equatorial section, X 40, to illustrate the embryonic apparatus and equatorial chambers; USNM

651075. 

19-23. Pellatispira madaraszi (Hantken) (p. B6).

19. Transverse section, X 20, of an inflated, selliform specimen to be compared with specimens from Saipan 

Island (Cole, 1957a, pi. 97, fig. 10); USNM 651076.

20. Compressed specimen, X 20; USNM 651077.

21. Part of a transverse section, X 20, of a microspheric specimen to illustrate the development of the marginal 

cord; USNM 651078.

22. Part of a transverse section, X 20, of a large, thin, megalospheric specimen; USNM 651079.

23. Transverse section, X 12.5, of a probable microspheric specimen; USNM 651080.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-B PLATE 2

25 23 22 21 20

CAMERINA, SPIROCLYPEUS, ASTEROCYCLINA, HETEROSTEGINA, 

DISCOCYCLINA, AND PELLATISPIRA
PLATE 3

FIGURES 1-3,7. Asterocyclina matanzensis Cole.(p- Bll).

1. Equatorial section, X 20, of a specimen having a subglobular 

central area and elongate rays; see plate 4, figure 10, for enlargement of the embryonic apparatus; USNM 651083.

2. Vertical section, X 20, of a specimen having a narrow rim from 

which bluntly rounded rays protrude; USNM 651084.

3. Equatorial section, X 20, of a specimen having an evenly lenticular 

central area which merges with five bluntly rounded rays; USNM 

651085. 

7. Vertical section, X 20, of a specimen similar to the one from which

the equatorial section, figure 3, was made; USNM 651089. 

4-6, 8, 9, 12. Heterostegina saipanensis Cole (p. B9).

4, 5. Transverse sections, X 20; USNM 651086, 651087.

6. Part of a median section, X 40, of a specimen having one camerinid chamber after the embryonic chambers; USNM 651088. 

8. Part of a median section, X 40, of a microspheric specimen;

USNM 651090. 

9, 12. Median sections, X 20, of megalospheric specimens;USNM 651091,

651094. 

10, 11, 13-16. Camerina pengaronensis (Verbeek) (p. B4).

10, 16. Median sections, X 20; USNM 651092, 651098. 

11, 13-15. Transverse sections, X 20; USNM 651093, 651095-651097.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-B PLATE 3

13 "W *r 14 ^U.'^ 15

ASTEROCYCLINA, HETEROSTEGINA, AND CAMERINA

16
PLATE 4

FIGURES 1, 5, 7, 11, 13, 14. Asterocyclina penuria Cole (p. B12).

1. Equatorial section, X 12.5, to illustrate the rayed pattern of the equatorial chambers; USNM 651099. 

5. Vertical section, X 20, of an inflated specimen; USNM

651103.

7. Part of an equatorial section, X 20; USNM 651105. 

11. Vertical section, X 20, of a compressed specimen; 

USNM 651108.

13. Vertical section, X 12.5; USNM 651110.

14. Part of an exactly centered equatorial section, X 40, 

to illustrate the embryonic, periembryonic, and equatorial chambers; USNM 651111. 

2-4 6, 8-10, 12. Asterocyclina matanzensis Cole (p. Bll).

2, 9. Vertical sections, X 40, of small specimens; USNM 

651100, 651107.

3. External view, X 8, of a specimen having a subglobular central area and distinct, elongate rays; 

USNM 651101.

4. External view, X 8, of a compressed specimen having

short, bluntly rounded rays; USNM 651102. 

6. Vertical section, X 20, of an inflated specimen having

numerous lateral chambers; USNM 651104. 

8, 10. Parts of equatorial sections, X 40, to illustrate slight 

differences in size of the peiiembryonic chambers; 

USNM 651106, 651085. 

12. Equatorial section, X 20; USNM 651109.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-B PLATE 4

12
PLATE 5

FIGURES 1 21. Asterocyclina praedpua Cole (p. B13).

1-4, 6-8, 10, 12, 13, 16, 21. Vertical sections. Figures 1-3, 13, 21, X 40;

figures 4, 6-8, 10, 12, 16, X 20.

1, 2, 4, 12, 13. Inflated, biconvex, megalospheric specimens having four short, bluntly rounded 

rays; USNM 651112, 651113, 651115, 

651123, 651124.

3, 21. Moderately inflated megalospheric specimens having many strong pillars and 

moderately extended rays; USNM 

651114, 651129.

6-8. Megalospheric specimens having a distinct 

umbonal area and four extended rays; 

USNM 651117-651119.

10, 16. Microspheric specimens having a subglobular central area, strong pillars, and 

a narrow rim; USNM 651121, 651127. 

5, 9, 11, 15, 19, 20. Equatorial sections. All figures X 20,

except figure 20, X 40.

5, 9. Megalospheric specimens having tests similar to the specimens illustrated as figures 

6-8, this plate; USNM 651116, 651120. 

11, 19, .20. Megalospheric specimens having tests similar to the specimens illustrated as figures 

1-4, 12, 13, 21, this plate; USNM 651122,

651125. 651128.

15. Microspheric specimen having a test similar to the specimens illustrated as 

figures 10 and 16, this plate; USNM

651126.

14, 17, 18. Central parts of equatorial sections of megalospheric specimens. Figure 14, X 40; 

figures 17, 18, X 210.

14, 17. Embryonic and periembryonic chambers of 

the specimen illustrated as figure 19, this 

plate.

18. Embryonic and periembryonic chambers of 

another megalospheric specimen; see 

figure 13, plate 2.

 frU-S. GOVERNMENT PRINTING OFFICE: 1970 O-362-463
GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-B PLATE 5

21

ASTEROCYCLINA