Eocene Corals from 

Eua, Tonga

GEOLOGICAL SURVEY PROFESSIONAL PAPER 6 4 0-G
Eocene Corals from 

Eua, Tonga

By JOHN W. WELLS

With a statement on EOCENE FISH FAUNA OF EUA, TONGA, 

BASED UPON ADDITIONAL OTOLITHS

By JOHN E. FITCH

LATE EOCENE FOSSILS FROM EUA, TONGA

GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-G

Descriptions and paleoenvironmental 

analysis of 23 ahermatypic corals 

from off-reef tuffaceous limestone

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1976
UNITED STATES DEPARTMENT OF THE INTERIOR 

THOMAS S. KLEPPE, Secretary

GEOLOGICAL SURVEY

V. E. McKelvey, Director

Library of Congress Cataloging in Publication Data

Wells, John West, 1907- 

Eocene corals from Eua, Tonga.

(Late Eocene fossils from Eua, Tonga) (Geological Survey professional paper ; 640-G)

Bibliography: p.

Includes index.

Supt. of Docs, no.: I 19.16:640-G

I. Corals, Fossil. 2. Fishes, Fossil. 3. Paleontology Eocene. 4. Paleontology Tonga Eua (Island) 5. Otol: ths.

I. Title. II. Series. III. Series: United States. Geological Survey. Professional paper ; 640-G. 

QE75.P9 no. 640-G [QE778] 557.3'08s 76-608144 [563'.6'099612]

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

Washington, D.C. 20402 

Stock Number 024-001-02903-1
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 facies 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 were 

obtained from an outcrop on the 400-foot tem.ce 

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 YZ mile north of army lookout tower, 

at the second stream north of Vaigana [sic]." The 

brachiopods were examined by G. A. Cooper of the 

U.S. National Museum. Some years later wher 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 Kondo of the Berrice 

P. Bishop Museum in Honolulu intended to visit Eur, in 

connection with his studies of living Pacific island Ir.nd 

snails (under National Science Foundation gr-mt 

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

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, Mosese 

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

lookout tower mentioned by Stearns no 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
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.

In 1969 Wilfred Bryan of the Carnegie Institution 

of Washington collected additional material from the 

locality sampled by Kondo. Bryan's material was 

taken from soft calcareous tuffs 2-3 feet in thickness 

that dipped 30°-40° E. These calcareous tuffs were

MAR.ANJ / MARSHALL

FIGURE 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). Islands shown include the surrounding reefs.

IV
directly underlain by several feet of harder fossiliferous material that, in turn, lay above agglomerate 

with truncated dikes. The horizon sampled was at an 

altitude of about 355 feet in a gully notched into the 

400-foot terrace.

The exact extent of the richly fossiliferous bed is 

not known. In 1926, when Hoffmeister made his

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).

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

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. Additional fieldwork in the area of the rugged "Nine 

Gulches" would be worthwhile.

William Melson of the Smithsonian Institution 

examined hand specimens and thin sections 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 

devitrified and dark brown. Fragments of tuff are 

rare. There appears to be a large and varied assemblage of secondary minerals. The volcanic fragments 

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 limestones collected on Eua 

were treated with a wetting agent and penetrant in 

the laboratory. The material broke down easily, revealing a variety of fossil remains: Foraminifera, 

discoasters, corals, hydrozoans, brachiopods, bryozoans, annelids, crinoids, echinoids, ostracodes, 

barnacles, decapod crustaceans, mollusks, shark 

teeth, otoliths, and spores and other plant microfossils.

W. Storrs Cole has described the larger Foraminifera; these fossils suggest to him a depth of deposition of about 200 feet, but other groups notably the 

smaller Foraminifera, the corals, brachiopods, br^ozoans, 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 from the islands of the open Pacific, 

an area extending 4,000 miles from Palau to Tonga.

HARRY S. LADD
SUPPLEMENTARY NOTES

In 1971, Harold Stearns published a detailed account of the 5 days he spent on Eua in 1943, during 

the course of which he discovered the Eocene fossil 

deposit. In 1972, Bryan and others (Bryan, Stice, 

and Ewart; Ewart and Bryan) published articles 

dealing with the volcanic rocks of Eua.

In 1971, I visited Eua with Frank I. Coulson of 

the Fiji Geological Survey. Our purposes were to 

check on the stratigraphic relations of the fossiliferous horizon in the gully visited by Kondo and by 

Bryan, to search for other exposures of the horizon 

in gullies to the north, and to check all elevations in 

the immediate area. We were aided by aerial photographs and were able to determine key elevations 

with the help of controlled surveying aneroids.

Though the age of the Eua deposit seemed clearly 

determined by Foraminifera (U.S. Geol. Survey 

Prof. Papers 640-A, B) as late Eocene, we recognized that a mixture of much younger shallow-water 

forms might have been carried seaward to mingle 

with still-unburied late Eocene fossils. To test this 

possibility, five specimens of a small shallow-water 

brachiopod having tightly closed valves were opened, 

and the material filling them was examined for calcareous nannofossils. Mr. David Bukry of the U.S. 

Geological Survey recognized 10 species suggesting 

early Oligocene or latest Eocene age (written commun., 1972). The examination confirmed the earlier 

interpretation that the assemblage, though a mixture of shallow and deep forms, is not a mixture of 

young and old forms.

Most of the fossils described in the published chapters of Professional Paper 640 are assigned to one of 

three U.S. Geological Survey Cenozoic locality numbers. Actually, all three represent the same locality: 

24686, collected by Yoshio Kondo in 1967; 25745, collected by Wilfred Bryan in 1969; and 25147, collected by H. S. Ladd and F. I. Coulson in 1971. All 

the corals described by John Wells in chapter G were 

assigned number 24686.

As mentioned in the"Foreword," fossils represents 

ing more than a dozen organic groups were distributed to paleontologists for study and report. 

Eight of these groups contained sufficiently numerous and varied fossils to warrant .separate chapters, 

and work on seven of these has now been completed 

(U.S. Geol. Survey Prof. Papers 640A-G) ; study of 

the barnacles is continuing. The stud^T of seven 

smaller groups has led to summary statements that 

have been attached to larger reports. Chapter G, 

dealing with the corals, is accompanied by a supplementary report on the otoliths.

REFERENCES CITED

A,sano, Kiyoshi, 1939, Limestones of the Soutl Sea Islands 

under the Japanese Mandate: Yabe Jubile? Pub., v. 1, 

p. 537-550, 2 figs., 2 pis.

Bryan, W. B., Stice, G. D., and Ewart, A., 1972, Geology, 

petrography and geochemistry of the volcanic islands of 

Tonga: Jour. Geophys. Research, v. 77, no. 8, p. 1566- 

1585.

Cole, W. S., 1950, Larger Foraminifera fro^n the Palau 

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

 1957a, Larger Foraminifera, in Pt. 3. Paleontology, 

of Geology of Saipan, Mariana Islands: U.S. Geol. Survey Prof. Paper 280-1, p. 321-360, pis. 94-118.

 1957b, Larger Foraminifera from Eriwetok Atoll 

drill holes: U.S. Geol. Survey Prof. Paper 2-0-V, p. 743- 

784, pis. 230-249, 1 text-fig. [1959].

 1960, Upper Eocene and Oligocene larger Foraminifera from Viti Levu, Fiji: U.S. Geol. Surve7 Prof. Paper 

374-A, p. A1-A7, pis. 1-3, 1 text-fig.

Ewart, A., and Bryan, W. B., 1972. Petroerarahv and geochemistry of igneous rocks from Eua, Tongan Islands: 

Geol. Soc. America Bull., v. 83, no. 11, p. 37.81-3298.

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

Survey Dept., Ann. Kept. 1958, Council Pawer 17, p. 3.

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

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

Stearns, H. T., 1971, Geologic setting of an Eocene fossil 

deposit of Eua Island, Tonga: Geol. Soc. America Bull, 

v. 82, no. 9, p. 2541-2551.

Whipple, G. L., 1932, Eocene Foraminifera, m Hoffmeister, 

J. E., Geology of Eua, Tonga: Bernice P. Bishop Mus. 

Bull. 96, p. 79-86.

HAPRY S. LADD

VI
CONTENTS

Foreword _________. 

Supplementary notes . 

Abstract _ _ . 

Introduction ___ _ 

Acknowledgments 

Material and preservation

Paleoenvironment . 

Previous work _ . 

The coral fauna .

Undetermined corals 

Systematic paleontology

Hydrozoa _ _

Stylasteridae _

Page

III

VI

Gl

1

1

1

1

1

2

Pa-re 

Systematic paleontology Continued

Anthozoa - G7

Octocorallia - 7 

Scleractinia - 7 

Fungiidae - 7 

Oculinidae _ - 7 

Anthemiphylliidae . 7 

Caryophylliidae ___ . 8 

Flabellidae _________ . 11

Guyniidae _ - 11

Dendrophylliidae . 11

References cited __-_ _ - 12

Eocene fish fauna of Eua, Tonga, based upon additional

otoliths, by John E. Fitch _____ . 13

Index ______ __ . 17

ILLUSTRATIONS

TABLES

[Plates follow index]

PLATE 1. Stylaster, Stenohelia, Crypthelia, Astya, Corallium, Fungiacyathus, and Parisis.

2. Madrepora, Anthemiphyllia, Caryophyllia, Cyathoceras, Deltocyathvs, Crispatotrochus, Steiwcyathus.

3. Notocyathus, Desmophyllum, Anomocora, Goniocorella,Javania, Enallopsammia.

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

3. Photograph showing concentrate of Eocene corals from Eua, Tonga _ _ __ -__. G2

4. Photograph showing sample of Holocene corals from off-reef slope east of Enyu Island, Bikini Atoll, at

depth of 720 m __ _________________.___________ ___ __ _ ___. 3

TABLE 1. Geologic ranges of genera and subgenera of corals from Eua, Tonga _ _ _ 

2. Bathymetric ranges of genera and subgenera of Eocene corals from Eua, Tonga 

3. Thermometric distribution (°C) of genera and subgenera of Eocene corals from Eua, Tonga

4. Number of teleost otoliths in each collection from Eua, Tonga ___ __ _.

Pave

G4

5

6

13

VII

LATE EOCENE FOSSILS FROM EUA, TONGA

EOCENE CORALS FROM EUA, TONGA

By JOHN W. WELLS

ABSTRACT

Twenty-three ahermatypic corals are described from tuffaceous limestone of late Eocene age. Four are stylasterine 

hydrozoans, 3 of which are new species; 2 are octocorals; 17 are scleractinians, 13 of which are described as 

new. The assemblage indicates a cool bathyal thicket assemblage at depths greater than 200 m (650 ft).

INTRODUCTION

Ahermatypic corals of Eocene age have not heretofore been found in the Pacific, except in New Zealand, and their discovery in Eua adds a significant 

new dimension to the history of corals in this vast 

area.

Compared with Eocene ahermatypic coral faunas 

elsewhere, the Eua corals do not have the "Eocene 

Look," an aspect noted by Ladd (1970) for the mollusks of Eua. However, the Eua corals are a deepwater assemblage, whereas most, if not all, other 

Eocene ahermatypic faunas are from the neritic 

zone. Further, deep-water faunas in general are cosmopolitan on the generic level, both geographically 

and stratigraphically (table 1). None of the genera 

from Eua are extinct, and all are widespread in deep 

water in present seas. Comparisons and correlations 

with other Eocene ahermatypes are not productive 

at present. The only possibly similar fauna is that 

described by Squires (1958, 1962) from the Eocene 

Arnold Group of New Zealand. This fauna is a cool 

(10°-16°C.), shallow-water (20-40 m) assemblage 

of different generic composition.

ACKNOWLEDGMENTS

The writer is indebted to H. S. Ladd for the opportunity of studying this unusual coral fauna, to 

those who painstakingly picked out the several 

thousand bits and pieces of coral, and to F. M. Bayer 

for advice on the octocorals.

MATERIAL AND PRESERVATION

The Eua tuffaceous deposit and the method of recovering the embedded fossils have been described 

by Ladd (1970). The samples were broken up, releasing the fossils, which were sorted into the various taxa. Thousands of remains of corals, mostly 

small fragments (fig. 3), were turned over to the 

writer, and from these, several hundred specimens 

were selected for study and are the subject of this 

report. All material bears U.S. Geological Survey 

Cenozoic locality No. 24686.

All specimens are more or less worn and fragmentary, and on many of them, detail and structures 

are obscured by remnants of tightly cemented matrix; sufficient morphologic features, however, can 

be distinguished to substantiate generic assignments. 

All original aragonite has been altered to calcite, obscuring very fine detail. Many specimens are stained 

or coated with iron-manganese compounds, the significance of which is uncertain though such coatings 

are common on Holocene corals from the deep bathyal and abyssal zones.

PALEOENVIRONMENT

PREVIOUS WORK

Studies of other fossil taxa from Eua have provided somewhat conflicting interpretations of the 

environment of these fossils although there is no 

doubt that it was quite different from the contemporary reef biocoenose.

Todd (1970), on the basis of the depth ranges of 

the Holocene genera of Foraminifera found in the 

Eua beds, concluded that the depth was not less than 

about 200 m. Cole (1970), who studied the larger 

Foraminifera, noted the lack of abrasion of these 

forms which suggested to him that they were not 

subjected to extensive wave or current action, either 

because of water depth or because of dense mats of 

algae; Cole (p. B3) opted for "warm, shallow water

Gl
G2 LATE EOCENE FOSSILS FROM EUA, TONGA

(about 200 ft in depth) in a lagoon or protected situation on the outermost part of a reef." Kier (1970) 

thought that the fragmentary echinoderm material 

indicated water deeper than 80 m. Roberts (1970) 

thought that the decapod crustacean fragments accumulated in either the littoral or sublittoral zone. 

Ladd (1970, p. C2), from the evidence of the molluscan fauna, favored "deposition at a depth of about 

100 fathoms (180 m)."The ostracodes were analyzed 

by Hazel and Holden (1971), who recognized two 

assemblages, one autochthonous, indicative of the 

upper bathyal zone, and the other allochthonous, 

from the sublittoral environment. The small brachiopod fauna, as deduced from the meager data on living brachiopods, was thought by Cooper (1971) to 

be a mixture of shallow- and deep-water forms. 

Fitch (1971) thought that the teleostean otoliths 

indicated depths of more than 182 fathoms (325 m). 

The Eua cheilostome bryozoans studied by Cheetham 

(1972) included both allochthonous shallow-water 

and autochthonous deep-water genera, the latter 

much more abundant and interpreted as having lived 

at depths exceeding 200 m and in water cooler than 

13°C. The coral fauna considered in the present 

paper favors the bathyal zone below 200 m (650 ft).

Thus, the Eua thanatocoenose as a whole seems to 

contain a mixture of neritic and bathyal elements. 

The coral thicket environment, now almost wholly 

represented by worn, poorly sorted fragments, may 

have shoaled to its demolition, permitting an admixture of neritic types, many of which are less worn, 

but in fact, forms of corals characteristic of the 

neritic zone are lacking.

THE CORAL FAUNA

The Eua Eocene ahermatypic coral fauna is clearly a thanatocoenose that probably accumulated close 

to its life habit. The very high proportion of fragments of dendroid colonial forms (Madrepora, Enallopsammia, and stylasterids, fig. 3) to the varied but 

relatively small number of solitary types is indicative of derivation from banks or thickets (Squires, 

1964). The banks or thickets developed on a muddy 

or silty bottom, as suggested by the presence of occasional specimens of such eleutherozoic genera as 

Fungiacyathits, Anthemiphyllia, Deltocyathus, and 

Paradeltocyathus. The environment of the banks or 

thickets was similar to that of the Holocene banks 

on the Blake Plateau at depths of around 750 m, as 

shown in photographs by Stetson, Squires, and Pratt 

(1962) . Solitary corals that adhered to the branches 

of the thicket-formers, such as Caryophyllia, DesmoFlGURE 3. Concentrate of Eocene corals from Eua, Tonga: 

mainly worn fragments of Madrepora sp., a few of Enallopsammia eocenica, n. sp., and one of Desmophyllum lirioides, n. sp. X 2.

phyllum, Crispatotrochus, Cyathoceras, and Javania, 

are more common than the free-living forms.

A Holocene Pacific assemblage comparable with 

the Eua coral fauna was found in a dredge sample 

obtained during the Mid-Pacific Expedition in 1950 

from a depth of 720 m on the outer slope of Bikini 

Atoll east of Enyu Island. It consists of a handful of 

worn fragments of Enallopsammia, Madrepora, Desmophyllum, Stenocyathus, and the stylasterines Distichopora and Errina (Lepidopora) (fig. 4).

The relative abundance and variety of stylasterine 

hydrozoans (four of the 18 recognized genera) is 

significant. The only other comparable hydrocoral 

assemblage is that from the Paleocene (younger 

Danian) of Denmark described by Nielsen (1919), 

which also contains the only other fossil occurrence 

of the rare genus Astya. On the other hand, the cosmopolitan genus Flabellum, species of which are 

common throughout the Cenozoic, is notably absent. 

Eocene species of Flabellum are neritic types having 

stout coralla, as contrasted with the more delicate 

structures and generally larger size of modern 

bathyal forms. These forms may have been present 

in the Eua community but failed to survive the wear 

and tear that affected the corals; more likely, the 

thicket environment was unfavorable for such 

bathyal types, as at the present time.
CORALS G3

FIGURE 4. Sample of Holocene ahermatypic corals from offreef slope east of Enyu Island, Bikini Atoll, depth 720 m, 

dredged by Mid-Pacific Expedition, 1950: mostly worn but 

a few fresh fragments of Madrepora sp. and a few of Enallopsammia amphelioides (Alcock) and Desmophyllum sp. 

X2.

Table 1 shows the nonneritic character of the Eua 

coral fauna, in which 13 of the 18 genera are recorded for the first time from the Eocene. Bathymetric ranges of the corals are summarized in table 

2, where again the deep-water aspect is evident. 

With one or two exceptions, specimens of the genera 

from less than 100 m deep are very few, and most of 

the genera, especially Enallopsammia and Madrepora, the thicket formers, are commonest below 200 

m. Table 3 shows the temperature ranges of the 

same genera. In general, the genera that are eurybathic are eurythermic, and the thicket formers are 

found only at or below 16°C and commonly below 

this maximum.

The data in tables 2 and 3 seem to show that the 

paleoenvironment of the Eua fauna was that of 

thickets, coppices, or banks in the bathyal zone below 200 m, in temperatures of less than 16°C. The 

paleoenvironment could have been much deeper, for 

with the exception of three genera having very few 

records, all the genera today occur in depths of more 

than 900 m in temperatures as low as 4°C.

UNDETERMINED CORALS

In addition to the corals described below, many 

fragments are too poorly preserved to be identified

with any certainty, although several genera are 

represented: several very small, bowl-shaped turbinoliids, possibly Peponocyathus; many small turbinate coralla, probably caryophylliids, in the form 

of internal molds; a single specimen of a small solitary dendrophylliid; and several small, cylindricalscolecoid types.

SYSTEMATIC PALEONTOLOGY

Class HYDROZOA

Order STYLASTERINA

Family STYLASTERIDAE

Genus STYLASTER Gray 1831

Stylaster sp. 

Plate 1, figures 1, 2

Some fragments of branchlets are referred to this 

genus, which was previously unrecorded earlier than 

Oligocene (S. antiquus Sismonda, Oligocene, Italy). 

All are worn, and most structures are poorly preserved. The branchlets range in diameter from 1 to 

2.5 mm, and have fine longitudinal striae alternating 

with fine perforations. Cyclosystems more or less 

regularly alternating on right and left sides, 0.75 

mm in diameter, slightly elevated. Gastrostyles not 

observed. Dactylopores about 11 in number, regularly enclosing gastropore. Many fragments with scattered hemispherical ampullae, commonly broken and 

appearing as rimmed pits.

Material. Figured specimens, USNM 208294; 

and 32 other fragments.

Remarks. S. mooraboolensis (Hall), Miocene of 

Australia, redescribed by Boschma (1953), has more 

slender branchlets and cyclosystems of the same size 

with an average of 12.5 dactylopores, but the latter 

do not completely encircle the gastropores. S. antiquus is as yet unfigured and known only from a brief 

description.

Genus STENOHELIA Kent 1870

Stenohelia boschmai Wells, n. sp.

Plate 1, figures 3-6

Branchlets slender, 1-1.5 mm in diameter, bearing 

large expanded cyclosystems on anterior surface, 

closely spaced about 1-1.5 mm apart. Coenosteum 

coarsely striate, with one or two rows of perforations between the subacute striae. Cyclosystems 

slightly compressed transversely, 1.5x2-3 mm in 

diameter. Gastropores deep, sides gently .sloping, 1 

mm in diameter. Gastrostyles not observed. Dactylopores 15-19 in number, wider than intervening "septa," prominent with minute, rarely preserved dactylostyles. Margins of one or two lower dactylopores 

in some cyclosystems elevated and inclined inward. 

Ampullae large, hemispherical, with irregularly
G4 LATE EOCENE FOSSILS FROM EUA, TONGA 

TABLE 1. Geologic ranges of genera and subgenera of corals from Eua, Tonga

[X, first record from the Eocene]

Caryophyllia 

Desmophyllum 

Deltocyathus 

Fungiacyathus 

Astya 

Stenocyathus 

Madrepora 

Anthemiphyllia 

Cyathoceras

Crispatotrochus

Notocyathus 

(Paradeltocyathus)

Stylaster

Enallopsammia 

Javania 

Goniocorella 

Anomocora 

Stenohelia 

Crypthelia

Cretaceous

?

Paleocene

?

Eocene

X 

X

X 

X 

X 

X 

X 

X 

X 

X

X 

X

X

0)

c

0) 

0

o .00

6

Miocene Pliocene

Pleistocene Holocene

reticulate coenosteal striae, strongly porous when 

worn, commonly one or two borne on undersides of 

cyclosystems.

Material Holotype, USNM 208295; paratypes, 

USNM 208296; and 13 other fragments.

Remarks. This is the first record of fossil Stenohelia. All specimens are fragments, and none show

the mode of branching. This species is distinguished 

from the 10 Holocene species by the very coarse coenosteal striae, the large ampullae on the underside 

of the cyclosystems, and large open dactylopores.

The species is named for Dr. H. Boschma of Leiden, the leading authority on the stylasterine 

hydrocorals.
CORALS

TABLE 2. Bathymetric ranges (in meters) of genera and subgenera of Eocene corals from Eua, Tonga

G5

Numerals indicate number of records below 100 m 

Neritic Bathyal 

0 50 100 200 300 400 500 600 700 800 900 1000

Anomocora

Anthemiphyllia 

Astya*

Caryophyllia 

Crispatotrochus 

Crypthelia* 

Cyathoceras 

Deltocyathus

Desmophyllum 

Enallopsammia 

Fungiacyathus 

Goniocorella 

Jaoania

Madrepora

Notocyathus 

(Paradeltocyathus)

Stenocyathus 

Stenohelia *

Stylaster*

1 

3

1

1

1

1 

2 

2

1

1 

1

X

> °475

^ ^700

 >1080

 -* 4275 

 >2035

> T>Q*>

 > 5870

 >3000

 ^1220

 ^ 1901

^ 1 /- on

*,Stylasterid

x,One record between 100 and 200 m

Genus CRYPTHELIA Milue Edwards and Haime 1849 

Crypthelia vetusta Wells, n. sp.

Plate 1, figures 7, 8

Branchlets 0.7-1.15 mm in diameter, finely and 

evenly striated longitudinally as in Astya nielseni, 

bearing cyclosystems on the anterior surface commonly at points of origin of new branchlets. Cyclosystems slightly elevated, 1-1.75 mm in diameter, 

short lids inclined upwards at about 60°. Cyclopores 

deep, sides nearly vertical. Dactylopores 10-17 in

number, commonly 16-17. Ampullae large, domed, 

developed on lids, leaving an irregular pit where 

broken.

Maternal. Holotype, USNM 208297; paratypes 

(4), USNM 208298; and 35 other fragments.

Remarks. Crypthelia has previously been reported only from the Holocene, 12 species having 

been listed by Boschma (1957). C. vetusta is closest 

to such Pacific species as C. pudica Milne EdwaHs 

and Haime and C. japonica Milne Edwards and 

Haime.
G6 LATE EOCENE FOSSILS FROM EUA, TONGA 

TABLE 3. Therniometric distribution, (°C) of genera, and subgenera of Eocene corals from Eua, Tong",

Caryophyllia

Deltocyathus 

Stylaster *

Anthemiphyllia

Desmophyllum

Notocyathus 

(Pa radeltocyath us)

Cyathoceras

Fungiacyathus 

Crypthelia *

Gkmiocorella 

Javania

Anomocora 

Stenohelia * 

Enallopsammia

Madrepora 

Stenocyathus

Crispatotroch u s 

Astya

28 27 26 25 24 23 22 21 20 19 18 17 16 15 14 13 12 11 10

1

2

*> Stylasterids

1, One record

2,One record (7°C)

Genus ASTYA Stechow 1921 

Astya nielseni Wells, n. sp.

Plate 1, figure 9

Branchlets 0.5-1.25 mm in diameter, thickest near 

base of attachment of corallum, strongly striated 

longitudinally, bearing cyclosystems on anterior surface, most commonly at points of origin of new 

branchlets. Cyclosystems slightly elevated above surface of branchlets, 1.5-4 mm apart, scarcely compressed, 0.75-1.75 mm in diameter, the large ones

bearing ampullae. Gastropores circular with nearly 

vertical sides. No gastrostyles seen. Dactylopores 17- 

19 in number. Ampullae encircling cyclosystems; 

when broken, they roughen the exteriors of the 

cyclosystems.

Material Holotype, USNM 208299; paratypes 

(3), USNM 208300; and 53 other fragments.

Remarks. The rings of ampullae around the cyclosystems and apparent absence of gastrostyles are 

typical of the sole living species, A. subviridis (Mose-
CORALS G7

ley), to which the present species is closely related; 

A. nielseni differs mainly in having smaller cyclosystems and more delicate branchlets. In A. crassa 

(Nielsen) from the Paleocene of Denmark, the cyclosystems do not appear to be confined to the 

anterior surface of the branchlets.

This species is named for K. Briinnich Nielsen 

(1872-1942), who studied the coral faunas of the 

Danish Paleocene.

Class ANTHOZOA 

Subclass OCTOCORALLIA

Order GORGONACEA 

Suborder SCLERAXONIA

Undetermined octocorals 

Plate 1, figure 10

Some fragments of octocoral axes are referable to 

the suborder Scleraxonia, but identification below 

this level is uncertain, as these supporting structures 

rank low among the criteria of classification. A few 

specimens are illustrated (USNM 208301). Solid, 

smooth or thorny unjointed axes are probably CoralHum, and ridged internodal segments appear to be 

Parisis.

Material About 50 fragments. USNM 208301.

Subclass ZOANTHARIA

Order SCLERACTINIA

Family FUNGIIDAE

Genus FUNGIACYATHUS G. 0. Sars 1872

Fungiacyathus euaensis Wells, n. sp.

Plate 1, figures 11-14

Corallum small, cupolate, diameter four to five 

times the height. Base flat, imperforate, with low, 

thin, acute, equal costae extending to the center. 

Costae of fifth cycle in some specimens tending to 

break up into elongate granules near the center. Septa (96) in five complete cycles, thin, laterally with 

thin alternating trabecular ridges giving a zigzag 

aspect to the septa in plan. Margins of the 12 firstand second-cycle septa elevated midway between 

margin and axis, dissected into several tabulations. 

Three or four concentric rings of synapticulae. Columella not observed but apparently in a small axial 

depression.

Measurements. Holotype, USNM 208302: height 

3 mm, diameter (restored) 14 mm; paratypes, 

USNM 208303, 208304: height 3 mm, diameter 10.5 

mm.

Remarks. For practical purposes, the species of 

Fungiacyathus ("Bathyactis") can be arranged in 

two groups: (1) those having four cycles of septa 

(48), including most of the fossil species, and (2) 

those having five cycles of septa (96). The latter 

group includes F. hawaiiensis (Vaughan), Holocene,

Hawaii; F. Stephana (Alcock), Holocene, Ind'an 

Ocean; F. palifera (Alcock), Holocene, Malaysia, 

Japan, South Australia; F. sibogae (Alcock), Holocene, Malaysia; and F. kikaiensis (Yabe and Eguchi), Pleistocene (Ryukyu Limestone), Ryukyu Islands. Bathyactis eocenica Gerth, Eocene, Java, is 

probably not Fungiacyathus. F. euaensis differs 

from all of these by having the equal, low, ac'ite 

costae.

Material. In addition to the holotype and paratypes, nine fragments show some details of septa r.nd 

costae.

Family OCULINIDAE

Subfamily OCULININAE

Genus MADREPORA Linnaeus 1758

Madrepora sp.

Plate 2, figures 1-3

Many small fragments, mostly representing small 

distal branchlets 1.5-4 mm in diameter, are characteristic of this genus. Corallites are budded alternately in one plane, walls faintly costate in young 

corallites, smooth and thickened by external ste.reome on larger branchlets. Calices deep, 2 mm in 

diameter. Septa thin, laterally granulated, in three 

cycles arranged 6/6/12, the last cycle very short, 

free on their inner edges. Upper margins of senta 

thickened at calicular margin. First two cycles urite 

deep in the calice to form a weak columella. Corallites on larger branches filled with stereome.

Material Figured branchlet, USNM 208305; 

figured section, USNM 208306, and about 100 small 

fragments.

Remarks. At least 30 Tertiary "species" of 

Madrepora ("Amphelia") have been described, six 

of them from the Eocene, but species discrimination 

is unresolved, and the Tongan form is not named 

here. It could be related to M. granulata (TenisonWoods) of the Eocene and Oligocene of New Zealrnd 

(Squires, 1958), but that form is based on larger 

fragments.

Family ANTHEMIPHYLLIIDAE 

Genus ANTHEMIPHYLLIA Pourtale* 1878

Anthemiphyllia catinata Wells, no. sp.

Plate 2, figures 4, 5

Corallum bowl-shaped, small, diameter 7-8 mm, 

height 4.5-5 mm. Wall thick, costate. Costae equal, 

thick, rounded, finely granulated, extending p^rt 

way to center of base. Calice shallow. Three cy^es 

of septa with a variable number of fourth-cycle s^pta (32-40). Septa thick, slightly exsert, those of 

first three cycles extending to columella, upper margins bearing 5 or 6 thick granulated dentations. Columella large (2 mm), papillate.
G8 LATE EOCENE FOSSILS FROM EUA, TONGA

Material Holotype, USNM 208307; paratype, 

USNM 208339; and two smaller, worn coralla.

Remarks. Coralla of this species are about the 

size and shape of the type species, A. patera of the 

Caribbean, but have fewer septa, fewer and coarser 

septal dentations, and a papillary rather than spongy 

columella, The living Indo-Pacific species, A. dentata 

(Alcock), is larger, more patellate, and has more 

septa. The latter species has been reported previously (Yabe and Eguchi, 1937) as fossil from the Ryukyu Limestone (Pleistocene) of Kikai-Zima. A. patella Gerth (1921) from the late Oligocene of Java, 

like A. dentata, is larger and more patellate. A. verbeeki Gerth (1921), is a large turbinate epithecate 

form, possibly a Cynarina.

Family CARYOPHYLLIIDAE

Subfamily CARYOPHYLLINAE

Genus CARYOPHYLLIA Lamarck 1801

Caryophyllia denaria Wells, n. sp. 

Plate 2, figures 6-10

Corallum cornute, fixed by a small base. Exterior 

nearly smooth, with very faint low equal broad costae corresponding to all septa, commonly obliterated 

by stereome. In very small corallites (1.5 mm) the 

costae are thin with single rows of granulations. 

Calice round or slightly ovate, shallow. Septa slightand nearly equally exsert, laterally granulate, very 

thick at calicular margin, thinning axially. Septa 40 

in number, arranged 10/10/20, the third group fused 

to the second. Ten prominent thick solid pali before 

the second group of 10 septa. Columella composed of 

one or two thick elongate laths.

Measurements. 

Diameter of calice (mm)

Specimen

Height 

(mm) Internal External

1.

2.

3.

4.

5.

6.

Holotype,

Paratype,

Paratype,

USNM

USNM

USNM

208308

208309

208310

__ 19

__ 14

__ 23

15

14

5

7.5

7

5.5

5.5

5

3 X 3.5

10

10

8.

8

6.

3.

5

5

5 X4

The decameral arrangement of the septa and pali 

and obsolescence of the costae by stereome accumulation distinguish this species from Pacific species of 

Caryophyllia. The closest relationship appears to be 

with such Holocene species as C. octopali Vaughan.

Material. Represented by 23 specimens, all more 

or less worn and basal attachment broken.

Caryophyllia sp. cf. C. japonica von Marenzeller 

Plate 2, figures 11, 12

Caryophyllia japonica Squires 1958. New Zealand Geol. Survey, Palaeontology Bull. 29, p. 46, pi. 7, figs. 8-14, text 

fig. 15.

Four specimens may represent this species, reported from the Miocene and Pliocene of New Zealand by Squires, and known from Holocene localities 

in the Indian Ocean, East Indies, and Japan. They 

are larger than C. denaria and have more septa (44), 

which are more exsert and have thinrer corallite 

walls. The calicular structures are poorly preserved 

but there appear to be 11 or 12 thin pali and a crispate columella.

Measurements. Figured specimen, USNM 

208312: height 22 mm, diameters 12X13; another 

specimen: height 19 mm, diameter 10 mm.

Genus CYATHOCERAS Moseley 1881

Cyathoceras kondoi Wells, n. sp.

Plate 2, figures 13, 14

Corallum trochoid-conical, attached by a small 

base. Wall thick, smooth, costate near calice. Costae 

low, rounded, faintly and minutely granulated. Septa 

(48) in four complete cycles, thin internally, thickened at calicular margin, those of the firrt two cycles 

equal and exsert, those of the third and fourth cycles 

less exsert. Calice shallow, with prominent chicoraceous columella about a third the diameter of the 

calice.

Measurements. Holotype, USNM 20PH13: height 

18 mm, diameter 14 mm; paratype US1 T M 208314: 

16 mm, diameters 13 X 14 mm; paratype USNM 

208340: height 15 mm, diameter 11 mm.

The earliest previously known occurrence of Cyathoceras is C. periallus Squires (1962) from the lower Oligocene of New Zealand, from whicl x the present 

species differs by having fewer septa and broader 

rounded calice.

Material. Three slightly worn specimens with 

bases broken at diameter 2.5 mm.

The species is named for Dr. Yoshio Fondo of the 

Bernice P. Bishop Museum, Honolulu, Hawaii, who 

collected much of the Tongan material.

Genus DELTOCYATHUS Milne Edwards and Hai<ne 1848 

Deltocyathus sp.

Plate 2, figures 15, 16

A single worn patellate corallum is referred to this 

genus. Septa in four complete cycles, tr°. first cycle 

extending to the columella, the remaining cycles 

regularly uniting. Pali and columella not discernible. 

Costae equal, with traces of granulations, arranged 

following the septal plan.

Measurements. Figured specimen, USNM 

208315: height 2 mm, diameter 12 mm.

Remarks. Deltocyathus is fairly common from 

the Miocene and is widespread in modern seas, but 

pre-Miocene records are sparse and dubious: D.
CORALS G9

fontinalis Dennant 1904 from the Australian Miocene; D. sp. cf. D. italicus Hegedus 1962 from the 

Oligocene of Hungary; D. sp. Kolosvary 1964, Cretaceous of Hungary; D.I complectus Squires 1958, 

Cretaceous of New Zealand; D.I whitei Durham 

1943, Paleocene of California. The Eua specimen is 

the first record from the Eocene.

Genus CRISPATOTROCHUS Tenison-Woods 1878

Crispatotrochus tenison-woodsi Wells, n. sp.

Plate 2, figures 17-19

Corallum cornute, attached in early stages by a 

small base. Exterior faintly granularly costate in 

small specimens, thickened and smoothed by stereome in larger coralla. Epitheca lacking. Calice circular or slightly ovate, deep. Septa 32, arranged 

8/8/16, correspondingly exsert, very thick and 

coarsely granulated at the wall, thin axially, only the 

16 septa of the first two groups reaching the small, 

deep, coarsely trabecular columella.

Measurements. Holotype, USNM 208316: height 

27 mm, diameter 12 X 14 mm; paratype, USNM 

208317: height 20 mm, diameter 8 mm.

Remarks. Crispatotrochus groups with Ceratotrochus and Conotrochus; it resembles the latter but 

lacks the stout epitheca as well as the pali that occur 

in some species. C. tenison-woodsi differs from the 

type species, C. inornatus Tenison-Woods from the 

Holocene of Australia, by the extensive development 

of external stereome with obliteration of costae and 

much smaller columella. Ceratotrochus exiguus 

Squires 1958 from the New Zealand Oligocene appears to be a Crispatotrochus with flangelike major 

costae near the upper margin of the calice.

Material. Represented by nine specimens. This 

species is named for the Reverend J. E. TenisonWoods (1832-1889), pioneer student of Australian 

Tertiary and Holocene corals.

Subfamily TURBINOLIINAE

Genus NOTOCYATHUS Tenison-Woods 1880

Subgenus PARADELTOCYATHUS Yabe & Eguchi 1937

Notocyathus (Paradeltocyathus) orientalis (Duncan)

Plate 3, figures 1-3 

Deltocyathus orientalis Duncan, 1876, Zool. Soc. London Proc.,

p. 431, pi. 38, figs. 4-7.

D. (Paradeltocyathus) orientalis Yabe and Eguchi, 1937, 

Tohoku Univ. Sci. Bepts., ser. 2, v. 19, p. 131, pi. 20, 

figs. 1-10.

N. (Paradeltocyathus) orientalis Squires, 1958, New Zealand 

Geol. Survey Palaeontology Bull. 29, p. 55, pi. 9, figs. 

19-22.

Squires and Keyes, 1967, New Zealand Oceanog. Inst. 

Mem. 43, p. 24, pi. 3, figs. 1-7.

Ten small specimens have the characteristic bowl 

shape with sharply incised grooves between the 

thick, finely granulated costae of this widespread 

species, known from the Oligocene and Miocene of 

New Zealand; Miocene and Pleistocene of Japan; 

Pliocene of Ceram; and Holocene of Japan, Forme sa, 

New Zealand, eastern Indian Ocean, Atlantic, ?.nd 

Caribbean.

N. (P.) australiensis (Duncan) from the Miocene 

of Australia, originally described as Deltocyathus 

italicus var. australiensis, is very similar to N. (P.) 

orientalis and may prove to be the same, in which 

case Duncan's earlier name will take precedence.

Measurements. Figured specimens, US1 T M 

208323: height 2 mm, diameter 4 mm.

Subfamily DESMOPHYLLINAE 

Genus DESMOPHYLLUM Ehrenberg 1834

Desmophyllum decuplum Wells, n. sp.

Plate 3, figures 4, 5

Corallum trochoid, straight or cornute, fixed by 

expanded base. Wall moderately thick, covered by 

densely distributed fine granulations. Costae corresponding to septa developed only at or near calicrlar 

margins. Calice circular or slightly compressed. S epta 40, decamerally arranged 10/10/20, those of the 

first group of 10 more exsert than the rest and extending nearly to the axis. Septa of the second and 

third groups weakly developed, extending less than 

one-fourth the distance to the axis. Columella absent 

or a very deep, very weak tangle.

Measurements. 

Specimen

Height Diameter 

(mm) (mm)

Septa 

arrangement

Holotype, USNM 208324 __ 15

Paratype, USNM 208325 __ 11

Another specimen __ ___ 6.5

5X7 

6.5 

5

10/10/20 

10/10/20 

10/10/20

Most species of Desmophyllum have the normal 

hexameral septal arrangement.1 D. decuplum is thus 

distinguished from the other Eua species, D. lirioides and D. coulsoni, and is allied to D. taurinense 

(Michelin) from the Miocene of Italy, which has 

septa arranged 10/10/20/40, according to Chevalier 

(1961).

Desmophyllum Hrioides Wells, u. sp. 

Plate 3, figures 6, 7

Corallum tall, trochoid, straight or slightly cornute, attached by a small base. Wall thin, granular 

below, costate near calicular margin. Calice circular 

to slightly ovate, very deep. Septa thin, regularly 

slightly exsert according to cycle, upper marjrns

1 D. quinarium Tenison-Woods 1879, Holocene of Fiji, is not Degmoihyllum, but a juvenile Plerogyra.
G10 LATE EOCENE FOSSILS FROM EUA, TONGA

falling steeply towards axis, internal margins extending not more than two-thirds of the distance to 

the axis. Septa in five cycles, arranged 12/12/24/48 

in largest calices. Columella absent, the axial space 

being empty nearly to the base. 

Measurements. 

]

Specimen

Holotype, USNM 208326..

Paratype, USNM 208341 __

Height 

(mm)

25

14

30

15

13

6

Diameter 

(mm)

14 X 23 (crushed)

13 X 15

8 at base

10 X 12

9 X 11

7

3X4 (crushed)

S^pta 

arrangement

12/12/24/48

12/12/24/20

12/12/24,

all very short

12/12/24

12/12/24

12/12/24

6/6/12

Remarks. This species is distinguished by the 

relatively fragile corallum and by the empty axial 

space. The holotype is compressed and crushed, as 

are several other equally large but poorly preserved 

specimens. In most species of Desmophyllum, at 

least the first, or first and second, cycle septa extend 

to the axis. In the only other Pacific Eocene Desmophyllum, D. sp. Squires 1958 from the Bortonian of 

New Zealand, which is otherwise much like D. lirioides, the 12 septa of the first two cycles reach the 

axis.

Material. Represented by 28 specimens.

Desmophyllum coulsoni Wells, n. sp. 

Plate 3, figures 8, 9

Corallum elongate-conical, slightly curved. Wall 

costate near calice, finely granular below, thickened 

by stereome. Calice circular or slightly compressed, 

very deep axially. Septa arranged 12/12/24, those of 

the first two cycles equally slightly exsert, thickened 

at the wall, and extending nearly to the axis deep in 

the calice but not uniting. Third cycle shorter, nonuniting, and the fourth cycle very short. Columella 

absent.

Measurements. Holotype, USNM 208327: height 

27 mm (incomplete), diameter 8.5 mm; paratype, 

USNM 208342: height 42 mm, diameter 10 mm; another specimen: height 12 mm (incomplete), diameter 6 X 8 mm.

This species is distinguished from D. decuplum by 

the hexameral septal arrangement, and from D. lirioides by the elongate-conical, thick-walled corallum.

Material. Represented by seven specimens.

Named for Frank I. Coulson of the Fiji Geological 

Survey, who assisted in the investigation of the Eua 

deposit.

Subfamily PARASMILIINAE 

Genus GONIOCORELLA Yabe and Eguchi 19 "2

Goniocorella eguchii Wells, n. sp. 

Plate 3, figures 12, 13

Corallum subdendroid. Corallites cylindrical or 

slightly expanding, diameter 2-4.5 mm, new corallites branching at right angles. Wall faintly costate, 

internally thickened in older corallites. Calice deep. 

Septa (24) thin, in three cycles, regular^ arranged 

with only those of the first cycle extending to or nearly to the axis.

Measurements. Holotype, USNM 208330: height 

20 mm, diameter of main corallite 4 mm, of lateral 

buds (incomplete) 1.75-3 mm; paratype, USNM 

208331: height 17 mm, diameter of main corallite 

4.5 mm, of lateral bud 3 mm.

Remarks. This species is distinguished from the 

type- and only known species, G. dumosa (Alcock) 

from the Holocene of Indonesia, Japan, and New 

Zealand, by its feeble growth form anc1 more completely developed septation.

Occurrence. Represented by 10 specimens, all 

fragmentary and more or less worn.

This species is named for Dr. Motoki Eguchi, long 

associated with study of Pacific corals.

Genus ANOMOCORA Sruder 1878

Anomocora exilis Wells, n. sp.

Plate 3, figures 10, 11

Growth-form quasicolonial, consisting of elongate 

conical corallites with small offsets at right angles 

from the edge-zone below the calices. Wall thick, 

with rounded, lightly granulated subemial costae 

corresponding to all septa. Septa in holotype 32 in 

number, in two groups, arranged 6/6/12 in offsets. 

Larger septa thick at the wall, thinning axially, inner edges dropping steeply into the calice and extending a little more than halfway to the axis. Septa 

of the second group short, thin, not uniting to larger 

septa. Deep in the calice, the larger septa barely meet 

to form a weak trabecular columella. Endotheca very 

sparse.

Measurements. Holotype, USNM 208328: height 

(incomplete) 27 mm, diameter (just below calice) 

9 mm; paratype, USNM 208329 (an offset with two 

minor offsets) : height 17 mm, diameter of calice 3.5 

mm, septa arranged 6/6/12.

Remarks. Anomocora, although widespread in 

present seas, has not previously been recorded as 

fossil. A. exilis differs from the commor A. fecunda 

(Pourtales) in having thicker walls, fewer septa, 

and low rounded rather than thin acute costae.

Material. Represented by seven specimens.
CORALS Gil

Family FLABELLIDAE 

Genus JAVANIA Duncan 

Duncan, 1876. Zool. Soc. London Proc., p. 435.

Type (by monotypy) : J. insignis Duncan. Holocene, Japan.

Javania, by its primarily epithecal and secondarily 

thickened wall, appears to be closely related to Flabellum rather than to Desmophyllum, in which the 

costate wall is septothecal.

Javania duncani Wells, n. sp. 

Plate 3, figures 14-16

Corallum elongate conical, cornute basally in some 

cases. Wall thickened by stereome but in some specimens shows traces of external epithecal festoons 

between outer ends of the septa. Calice ovate, deep. 

Septa (12) of first two cycles equally exsert, very 

thick at the wall, thinning inwardly. Septa of third 

cycle free, very short and thin. Columella absent, but 

near the base, six stout primary septa fuse axially.

Measurements. Holotype, USNM 208332: height 

(incomplete) 17.5 mm, diameter 7X8 mm; paratype, USNM 208333: height (incomplete) 17 mm, 

diameter 7 X 8.5 mm.

Remarks. At present, pending the revision of 

Javania and related flabellid genera by H. Zibrowius, 

the only positively known species is the type, from 

which J. duncani differs by the proportionally taller 

but smaller corallum and by the development of only 

three cycles of septa.

Material. Represented by seven specimens.

The species is named for Peter Martin Duncan 

(1824-1891), who described many fossil and living 

ahermatypic corals.

Family GUYNIIDAE 

Genus STENOCYATHUS Pourtales 1871

Stenocyathus hoffmeisteri Wells, n. sp.

Plate 2, figures 20-24

Corallum elongate-cylindrical to scolecoid. Wall 

thin, epithecal but internally thickened by stereome. 

External surface not preserved but with single rows 

of small pores, about 4 per millimeter, in the thickened wall. Septa 14-16, in two groups arranged 8/8 

or 7/7, the 7 or 8 larger septa thick, extending to the 

axis to weakly join a small columella. Septa of the 

second group very short and thin.

Measurements. Holotype, USNM 208318: length 

11 mm, diameter 1.8 mm, septa arranged 8/8; paratype a, USNM 208319: length 9 mm, diameter 2 mm, 

septa arranged 8/8; paratype c, USNM 208321: 

length 5.5 mm, diameter 1.7 mm, septa arranged 7/7.

All specimens are worn, and calicular structures 

are not preserved. Basal or early stages are lacking,

and the living attitude is uncertain. Several fragments are internal molds that show clearly the deep 

pits extending into the internally thickened vail between septa, toward the site of the thin spots in the 

epitheca. These molds are markedly like the decalcified specimens "showing the complete cast of the 

inside cavity" of S. vermiformis figured by Pourtales (1880, p. 101, pi. 1, fig. 15).

Stenocyathus is marked not only by the pseudoperforate epithecal wall but more particularly by a 

single crispate columellar lath surrounded by five 

or 6 pali. The apparent absence of pali in these 

specimens, however, is expectable, as they are observable only in intact calices. Worn or broken fragments of the Holocene S. vermiformis show little or 

no trace of pali or columella.

S. hoffmeisteri differs from S. vermiformis, the 

type species which is widespread in the Caribbean, 

Atlantic, and Mediterranean, by the presence of only 

14-16 septa rather than 20-24; it differs also from 

the only other Holocene species S. decamera Ralph 

and Squires (1962) from New Zealand, the s^pta of 

which number about 30 and are decamerally 

arranged.

The only described fossil species is S. alabamiensis 

Wells (1947), reported originally from the Upper 

Cretaceous of Alabama but now known tc be of 

Paleocene (Clayton Formation) age; it has septa 

arranged 12/12, plus a few of the fourth cyc^e. Another fossil species, still undescribed, from the upper 

Oligocene (Spring Creek) of Australia, has septa 

arranged 6/6/12.

Material. Represented by 20 specimens.

The species is named for Dr. J. Edward Hoffmeister who first mapped the island of Eua.

Family DENDROPHYLLIIDAE 

Genus ENALLOPSAMMIA Sismonda and Michelotti 1871

Enallopsammia laddi Wells, n. sp. 

Plate 3, figures 17-22

Corallum dendroid, branches apparently not anastomosing. Calices on distal and terminal branches 

regularly alternating in one plane; on thick (6-8 

mm) proximal branches, calices tend to be on one 

face, united by extensive costate coenosteum. Costae 

distinct, thick, and rounded, each bearing a single 

row of low granulations, the narrow intercostal 

grooves pierced by perforations. Calices deep, moderately salient on terminal branches, slightly elevated on thick branches, 3.5 mm outside diameter, 2 

mm internally. Septa in three complete cycles (24), 

free on upper part, thickened at calicular margin, 

internally thin. The six primary septa of some calices
G12 LATE EOCENE FOSSILS FROM EUA, TONGA

more exsert at margins than those of the other two 

cycles. Deep in calice, third-cycle septa united with 

those of second. Columella very deep, composed of a 

few trabecular processes.

Measurements. Diameter of proximal branches 

5-8 mm, of terminal branchlets, 2.5-3 mm; outer 

diameter of calices 3.5 mm, internal 2 mm; depth of 

calices 2 mm; costae 12 in 5 mm.

Remarks. In addition to the five Holocene species 

of Enallopsammia recently revised by Zibrowius 

(1973), several fossil forms have been recorded, 

mostly from Europe: the type species, E. scillae (Seguenza), Miocene-lower Pliocene, E. poppelachii 

(Reuss), Miocene, E. sp. Squires, Miocene, New Zealand, and E. payricardensis Chevalier, Miocene. The 

last is better placed in Tubastraea. E. laddi is very 

similar to E. scillae, differing mainly in having less 

salient calices, weaker columella, and coarser costae.

Material Represented by several hundred small 

fragments, mostly of distal and terminal branchlets, 

all rather rolled and worn. Holotype, USNM 208334; 

paratypes, USNM 208335-208338.

The species is named for Dr. Harry S. Ladd of the 

U.S. Geological Survey.

REFERENCES CITED

Alcock, A. W., 1898, Account of the deep-sea Madrepotraria 

collected by the Royal Indian Survey Ship Investigator: 

Calcutta, 29, p., 3 pis.

 1902, Report on the deep-sea Madreporaria of the 

Siboga-Expedition: Leiden, E. J. Brill, 55 p., 5 pis. 

(Siboga-Expeditie, Mon. 16a).

Boschma, Hilbrandt, 1953, Notes on specimens of Stylaster 

mooraboolensis (Hall) in the collection of the Manchester 

Museum: Konlkl. Nederlandse Akad. Wetensch. Proc., 

Amsterdam, Ser. B., v. 56, no. 4, p. 355-363, 2 figs. 1 pi.

 1957, List of the described species of the order of 

Stylastrina: Zool. Verh., no. 33, 72 p.

Cheetham, A. H., 1972, Cheilostome Bryozoa of late Eocene 

age from Eua, Tonga: U.S. Geol. Survey Paper 640-F, 

26 p., 7 pis., 7 figs.

Chevalier, J. P., 1961, Recherches sur les madreporaires et les 

formations recifales miocenes de la Mediterranee occidental: Soc. Geol. France Mem., new ser., v. 40, 562 p., 

26 pis., 199 figs.

Cole, W. S., 1970, Larger Foraminifera of late Eocene age 

from Eua, Tonga: U.S. Geol. Survey Prof. Paper 640-B, 

16 p., 5 pis., 2 figs.

Cooper, G. A., 1971, Eocene brachiopods from Eua, Tonga: 

U.S. Geol. Survey Prof. Paper 640-F, 7 p., 1 pi.

Dennant, J., 1904, Descriptions of new species of corals from 

the Australian Tertiaries, Part VII: Royal Soc. South 

Australia Trans., v. 28, p. 52-76, pis. 22-25.

Duncan, P. M., 1876, Notices of deep-sea and littoral corals 

from the Atlantic Ocean, Caribbean, Indian, New Zealand, 

Persian Gulf and Japanese, etc., seas: Zool. Soc. London 

Proc., p. 428-442, pis. 38-41.

Durham, J. W., 1943, Pacific coast Cretaceous and Tertiary

corals: Jour. Paleontology, v. 17, no. 2, p. 19^-202, pi.

32, 2 figs. 

Fitch, J. E., 1971, Teleost otoliths, in Cooper, G. /,, Eocene

brachiopods from Eua, Tonga: U.S. Geol. Survey Prof.

Paper 640-F, p. F7-F8. 

Gerth, Heinrich, 1921, Anthozoa, in Martin, Karl, Di? Fossilen

von Java: Geol. Reichs-Mus. Leiden, Samml., new ser.,

v. 1, abt. 2, no. 3, p. 387-445, pis. 55-57.

 1933, Neue Beitrage zur Kenntnis der Korallenfauna 

des Tertiars von Java. I. Die Korallen des Eocaen und 

des alteren Neogen: Dutch East Indies, Dienst Mijnbouw 

Nederlandsch-Indie, wetenschappelijke Medede?!. 25, p. 

1-6.

Hazel, J. E. and Holden, J. C., 1971, Ostracoda of Irte Eocene 

age from Eua, Tonga: U.S. Geol. Survey Prof. Paper 

640-D, p. D1-D11, 5 pis.

Hegediis, Gyula, 1962, Magyarorszagi oligocen korallok: 

Hungary Foldtani Intezet Evi Jelentese, 1959, r. 231-261, 

pis. 1-3.

Kier, P. M., 1970, Crinoids and echinoids, in Tcdd, Ruth, 

Smaller Foraminifera of late Eocene age from Eua, 

Tonga: U.S. GeoL Survey Prof. Paper 640-A, p. A18.

Kolosvary, Gabor, 1964, Adatok erdely Mezozoos es Neozoos 

Korallfaunajanak ismeretehez [Contribution a la connaissance de la faune de coralliaires meso-et c?nozoiques 

de la Transylvanie]: Hungary Foldtani Intezet Evi 

Jelentese, 1961, pt. 2, p. 211-258, 10 pis. (Summ. in 

French and Russian.)

Ladd, H. S., 1970, Late Eocene fossils from Eua, Tonga 

Foreword: U.S. Geol. Survey Prof. Paper 640-A, p. 

iii-v, 2 figs.

 1970a, Eocene mollusks from Eua, Tonga: U.S. Geol.

Survey Prof. Paper 640-C, 12 p., 5 pis. 

Milne Edwards, H., and Haime, Jules, 1850, Rech?.rches sur

les polypes, in 5me Mem. Monographie des Oculinides:

Ann. Sci. Nat., ser. 3, v. 13, p. 62-110, pis. 3, 4. 

Nielsen, K. B., 1919, En Hydrocoralfauna fra Faxe og

Bemaerkninger om Danien et geologiske Stillirg: Dansk.

Geol. Forening Medd, ,v. 4, no. 16, 66 p., 2 pis. (French

summ.). 

PourtaleX L. F., 1880, Reports on the results of dredging

* * * in the Caribbean Sea, 1878-79, by the United States

Coast Survey Steamer "Blake" . . . VI. Report on the

corals and Antipatharia: Harvard Coll., Mus. Comp.

Zoology, Bull., v. 6, no. 4, p. 95-120, 3 pis. 

Ralph, P. M., and Squires, D. F., 1962, The extant scleractinian

corals of New Zealand: Victoria Univ. [New Zealand],

Zool. Pub. 29, 19 p., 8 pis., 1 fig. 

Reuss, A. E., 1848, Die fossilen Polyparien des Wiener Tertiarbeckens: Naturwiss. Abh. von W. Haidinger, v. 2, p.

1-109,11 pis. 

Roberts, H. B., 1970, Decapod crustaceans, in T?dd, Ruth,

Smaller Foraminifera of late Eocene age from Eua,

Tonga: U.S. Geol. Survey Prof. Paper 640-A, p. A1-A23. 

Seguenza, Giuseppe, 1864, Disquizisioni paleontobgiche intorno ai corallarii fossili delle rocce terziare del distretto

di Messina: Accad. Sci. Torino Mem., ser. 2, v. 21, p. 399-

560, pis. 6-15. 

Sismonda, Eugenic, and Michelotti, G., 1871, Mate~iaux pour

servir a la paleontologie du terrain tertiaire du Piemont.

II. Animaux. Types protozoaires et celenteres: Accad. Sci.

Torino Mem., ser. 2, v. 25, p. 257-361, pis. 1-10.
FISH FAUNA G13

Squires, D. F., 1958, The Cretaceous and Tertiary corals of 

New Zealand: New Zealand Geol. Survey Paleont. Bull. 

29, 107 p., 16 pis., 28 figs.

 1962, Additional Cretaceous and Tertiary corals from 

New Zealand: Royal Soc. New Zealand Trans., Geology, 

v. 1, no. 9, p. 133-150, 4 pis.

 1964, Fossil coral thickets in Waifrarapa, New Zealand: Jour. Paleontology, v. 38, no. 5, p. 904-915, pis. 

147-148, 3 figs.

Stetson, T. R., Squires, D. F., and Pratt, R. M., 1962, Coral 

banks occurring in deep water on the Blake Plateau: 

Am. Mus. Nat. History, Am. Mus. Novitates 2114, 39 p., 

15 figs.

Tenison-Woods, J. E., 1878, On the extratropical corals of 

Australia: Linnean Soc. New South Wales Proc., v. 2, 

p. 292-341, pis. 4-6.

 1879, On a new species of Desmophyllum (D. quinarium) and a young stage of Cycloseris sinensis: Linnean

Soc. New South Wales Proc., v. 3, p. 17-20, pi. 1. 

Todd, Ruth, 1970, Smaller foraminifera of late Eocene age

from Eua, Tonga: U.S. Geol. Survey Prof. Paper 640-A,

17 p., 8 pis., 2 figs. 

Vaughan, T. W., 1907, Recent Madreporaria of the Hawaiian

Islands and Laysan: U.S. Natl. Mus., Bull. 59, 427 p.,

96 pis. 

Wells, J. W., 1947, Coral studies: III, Three new Cretaceous

corals from Texas and Alabama: Bulls. Am. Paleontology,

v. 31, p. 165-176, pis. 10, 11. 

Yabe, Hisakatsu, and Eguchi, Motoki, 1937, Notes on Deltocyathus and Discotrochus from Japan: Tohoku Imp. Univ.

[Sendai], Sci. Repts., Ser. 2 (Geology), v. 19, no. 1, p.

127-147, pi. 20.

 1942, Fossil and Recent simple corals from Japan: 

Tohoku Imp. Univ. [Sendai], Sci. Repts., Ser. 2 

(Geology) v. 22, no. 2, p. 105-178, pis. 9-12.

Zibrowius, H., 1973, Revision des especes actuelles du genre 

Enallopsammia Michelotti 1871, etc.: Beaufortia, v. 21, 

p. 37-54, 23 figs.

EOCENE FISH FAUNA OF EUA, TONGA, 

BASED UPON ADDITIONAL OTOLITHS

By JOHN E. FITCH 3

After the publication of my report on fish otoliths 

from Eua (Fitch, 1971), I examined three additional 

small lots from the same locality. These three supplementary collections (USGS 24686, 24745, 25147) 

contained 129 recrystallized otoliths and otolith 

fragments. Although many were so badly deteriorated that it was difficult to determine whether they 

actually were otoliths, 48 had retained enough characters to be placed into families or family groups. 

These included at least 12 kinds of fishes belonging 

to seven "families" (Berycidae, Congridae, Diretmidae, Macrouridae, ?Melamphaidae, Myctophidae, and 

Sternoptychidae) and one kind from an unknown 

family. Table 4 shows the number of otoliths (by 

family) in each collection.

2 California Department of Fish and Game, Long Beach, California

90802.

TABLE 4. Number of teleost otoliths (by family) in each 

collection from, Eua, Tonga

Collection

Family

USGS USGS 2

24745 24745

USGS 1 (Fitch, (this USGS 3

24686 1971) report) 25147

Berycidae __ __ __

Congridae _ _ _

Diretmidae _ _ _

Flatfishes __ __

Macrouridae _ __ _

Myctophidae _

Sternoptychidae

Unidentifiable - --

Totals __ _ _

1

3

2

9

1

15

1

4

37

73

3

5

13

5

26

52

1

9

44

54

1

1

5

7

1 1,000 ft north of Vaingana, Eua, in first gulch: collected by Yoshio 

Kondo 1967.

2 Same locality as USGS 24686; collected by Wilfred Bryan, 1969

3 North of Vaingana, Eua; altitude 400 ft; collected by H. Ledd and F. 

Coulson, 1971.

FAMILY ACCOUNTS

BERYCIDAE

One fragment, constituting approximately the 

anteriormost two-thirds of a relatively large right 

sagitta, is from a berycid. It has a very distirct ostium (name applied to the front part of the sulcus or 

groove on the inner face of an otolith), and much of 

the ventral profile is still intact.

In a recent review of the family, Woods and 

Sonoda (1973) stated that two genera, Beryx and 

Centroberyx, compose family Berycidae. Stinton 

(1967, 1968) has illustrated otoliths from both of 

these genera; his Trachichthodes affinis is Centroberyx. In making direct comparisons, the deeply 

rounded ventral profile of the fossil fragment from 

Eua is most nearly like sagittae from Beryx, but it 

is quite thick, and in this respect it is closest to 

Centroberyx. The ostium in berycid sagittae has a 

very distinctive shape and usually composes at least 

50 percent of total otolith length. Thus, as the ostium on the fossil fragment is 8.1 mm long, this otolith, if entire, probably would have been between 14 

and 17 mm long. At that size, it obviously was from 

an adult.

As adults, most members of the family attain 

lengths of 350 to 500 mm and inhabit deep water. 

Maul (1954) reported that in Madeiran waters 

Beryx usually is captured 400 to 600 m beneath the 

surface. On the other hand, Centroberyx sometimes 

is found in water as shallow as 20 m (Munro, 1938). 

The fossil record for the family is extensive, beginning as early as the Upper Cretaceous, on the 

basis of skeletal remains (Berg, 1940), and f.s early 

as the Eocene, on the basis of otoliths (Weiler, 

1968b).
G14 LATE EOCENE FOSSILS FROM EUA, TONGA

CONGRIDAE

Four of the recrystallized sagittae from Eua were 

from eels, and although characters of the sulcus are 

eroded and vague, their overall shape and thickness 

permit their assignment to this family. Although 

present-day members of this large and well-known 

family of eels are abundant in the meso- and bathypelagic realms of the world oceans, only one specimen has been captured in water as deep as 1,800 m 

(Grey, 1956). On the other hand, many congrids are 

found inside the Continental Shelf, some in water as 

shallow as 20 m. Kotthaus (1968) has illustrated 

sagittae of several deep-dwelling congrids that were 

trawled in the Indian Ocean.

The family has a long and continuous fossil record. 

Berg (1940) reported their skeletal remains from 

the Upper Cretaceous, but congrid otoliths did not 

appear until the Paleocene (J. E. Fitch, unpub. data, 

1975). Congrid otoliths are extremely abundant in 

Eocene deposits throughout the world (Frizzell and 

Lamber, 1962; Weiler, 1968b) and have been fossilized during all subsequent geologic periods.

The four sagittae from Eua are quite small (1.8- 

2.5 mm long), and judged by their lobular margins, 

they probably are from juvenile eels.

DIRETMIDAE

Three of the better preserved otoliths from Eua 

were from an unknown diretmid. The largest of these 

measured 7.8 mm long by 12.0 mm high (sagittae in 

my comparative collection from an adult D. argenteus, 251 mm standard length, which was trawled off 

the Oregon coast, are 7.5 mm long by 11.8 mm high) ; 

the other two were only slightly smaller. There is no 

previously published fossil record for the family, but 

I have found diretmid otoliths in Eocene deposits 

near San Diego, Calif., and in Miocene deposits at 

Bowden, Jamaica (Fitch, unpub. data, 1975).

In a recent review of the family, Woods and Sonoda (1973) assign to it only a single genus, Diretmus. Although they do not state how many species 

compose the genus, their synonymies indicate that 

they recognize only the two that they discuss. Speciments of Diretmus are relatively rare in collections, 

most having been netted from 200 to 2,000 m beneath the surface. However, judging by the number 

of diretmid otoliths I have seen in dolphin, swordfish, and shark stomachs, I believe that the absence 

of Diretmus specimens in museum collections indicates the fish's ability to avoid nets rather than any 

actual scarcity. The sagittae of Diretmus are quite

distinctive, and on the basis of those in my comparative collection as well as those I have seen in predators' stomachs, D. aureus from New Zealand is valid, 

and probably two or three additional species are 

extant.

MACROURIDAE

Previously, I recognized five macrourid sagittae 

among the identifiable otoliths from Eua but would 

not speculate as to their generic affiliation (Fitch, 

1971). Among the 48 identifiable otoliths thr.t I have 

subsequently examined, 18 are from macrourids, and 

these represent at least three species. One of these is 

almost square in outline but has rounded corners. 

Although it compares favorably with sagittae from 

an extant species of Corypliaenoides (ths.t is, C. 

armatus), I do not feel that I could justify assigning it to this genus. The other two kinds match almost perfectly the angular outline of sagittae from 

Coelorinchus, differing from each other primarily 

in ratio of height to length.

PMELAMPHAIDAE

Melamphaid otoliths have a very distinctive sulcus and can be recognized from that alone. In addition, sagittae of all melamphaid genera except Scopelogadus have one or more hairlike projections from 

one or both corners of the ventral margin which are 

equally distinctive. Unfortunately, in the s'ngle recrystallized otolith from Eua, these characters can 

not be observed, so its provisional assignment here 

is based mostly upon outline and intuition, neither of 

which is diagnostic nor infallible.

According to Ebeling and Weed (1973), the Melamphaidae are one of the largest and most varied of 

all bathypelagic families of fishes. Adults and halfgrown individuals usually are found belov 100 to 

200 m, although larvae and young may occur nearer 

the surface. Weiler (1968a) illustrated sagittae from 

the five genera of melamphaids and speculated upon 

their systematic position.

The fossil record of the family is not extensive, 

probably because melamphaid hard parts are delicate and fragile and do not preserve well. A skeletal 

imprint is known from the Miocene of California 

(Ebeling, 1962), and I have seen otoliths from the 

Miocene of Jamaica (Fitch, unpub. data, 1975) ; 

otherwise, otoliths of Melamphaes and Scop*,logadus 

are known from the lower Pleistocene of California 

(Fitch, 1968). The Eua specimen (2.6 mm long) is 

closest to Melamphaes among the five extent genera.
FISH FAUNA G15

MYCTOPHIDAE

Sixteen of the 48 identifiable otoliths that I examined for this report were from four kinds of lanternfish. The sagittae of several myctophid genera 

have spines along the ventral margin and are easily 

distinguished when these are present. Unfortunately, none of the recrystallized otoliths had retained 

these spines, if they ever had them, so generic assignment is impractical for many of these. One otolith, by its elongate shape, broadly rounded posterior 

end, and typical strong rostrum, resembles Ceratoscopelus, and several of the larger otoliths would be 

difficult to distinguish from Symbolophorus; however, the other two kinds were most similar to 

Diaphus and Benthosema,, both having otoliths with 

spinose ventral margins.

Although these Eua fossils are much like these 

four genera, I do not believe that the quality of the 

fossils is sufficient to justify making firm assignments. The 13 myctophid otoliths previously described from Eua (Fitch, 1971) represented three of 

the same four types noted above.

STERNOPTYCHIDAE

One otolith was unquestionably from an ancestral 

form of Ichthyococcus, a small luminescing mesoand bathypelagic fish found in most world oceans. 

Ichthyococcus has a very distinctive sagitta which is 

extremely large for the size of the fish and has a very 

thin bladelike rostrum projecting from the lower 

third of the otolith. Although Kotthaus (1967) illustrated sagittae of I. ovatus, his figures are unreliable 

because the preservative (formalin) used on the fish 

had dissolved the rostra before the otoliths were 

removed. The Ichthyococcus sagitta illustrated by 

Fitch and Brownell (1968), although taken from the 

stomach of a pygmy sperm whale, was little affected 

by digestive juices and shows a typical rostrum. The 

only difference I can find between fossil and living 

Ichthyococcus otoliths is a more deeply channeled 

sulcus in the fossil.

Although the genus is unreported as a fossil (Andrews and others, 1967), R. J. Lavenberg of the 

Natural History Museum of Los Angeles County, 

Calif, (oral commun., 1975) has informed me that 

Xyrinius houshi, which has been considered an extinct Miocene wrasse, is actually a species of Ichthyococcus. Interestingly, in their description of this 

California fossil, Jordan and Gilbert (1919) gave 

details concerning its eye and mouth, yet an examination of the unique holotype shows that the anterior 

part of the head, to a point just behind the eye, is

entirely lacking and has been since the day it was 

collected. In addition, many photophores car be discerned along the ventral profile of the body, a feature 

unknown in family Labridae.

Ichthyococcus otoliths are abundant in an Eocene 

deposit near San Diego, Calif., and in Miocene deposits north of Auckland, New Zealand (Fitch, unpub. 

data, 1975) ; they very likely have gone unrecognized 

in other deep-water deposits throughout the world.

UNKNOWN FAMILY

Four otoliths are almost oval in outline and have 

a straight, deeply channeled sulcus along the midline 

of the horizontal axis. The outer faces of these otoliths are evenly convex. Unfortunately, none of these 

otoliths has retained any indication of an ostium, so 

it is impossible to make a familial assignment. Sagittae of several extant genera in an equal number of 

families are oval in outline and have a straight deep 

sulcus but show differences in the shape and proportions of the ostium. The largest of these four otoliths is 3.7 mm long.

UNIDENTIFIABLE SAGITTAE

Eighty-one otoliths were either too fragmentary 

or too poorly preserved to offer any clues to family 

affiliation.

DISCUSSION

The 48 identifiable otoliths reported here bring to 

14 the kinds of bony fishes from the Eocene of Eua 

that can be assigned to families or family groups. An 

additional five kinds, including four mentioned in 

my earlier report (Fitch, 1971), could not be so 

placed but were different from those that could be 

assigned. Of the eight families involved, only two 

("flatfishes" and Congridae) contain extant members which, as adults, are not inhabitants of meso- or 

bathypelagic environments.

On the basis of otoliths alone, one would have to 

conclude that deposition took place at considerable 

depth, probably even deeper than the 182 fathoms 

postulated in my earlier report (Fitch, 1971).

REFERENCES CITED

Andrews, S. M., and others, 1967, Pisces, in Harlan'l, W. B., 

and otheirs, The fossil record: London, Geol. Soc. London, 

p. 637-683.

Berg, L. S., 1940, Classification of fishes both Recent and fossil. Facsimile reproduction, 1947, J. W. Edwards, Ann 

Arbor, Mich. [English-Russian text] U.R.S.S. .A cad. Sci., 

Inst. Zool. Travaux, v. 5, no. 2, p. 87-517.

Ebeling, A. W., 1962, Scopelogadusl capistranensis a new fossil melamphaid (Pisces: Teleostei) from Capistrano
G16 LATE EOCENE FOSSILS FROM EUA, TONGA

Beach, California: Postilla, no. 71, p. 1-6.

Ebeling, A. W., and Weed, W. H., Ill, 1973, Order Xenoberyces (Stephanobearyciformes), in Cohen, D. M., ed., 

Fishes of the western North Atlantic, Part 6: Sears 

Found. Marine Research, p. 397-478.

Fitch, J. E., 1968, Otoliths and other fish remains from the 

Timms Point Silt (Early Pleistocene) at San Pedro, California: Los Angeles County Mus. Contr. Sci., no. 146, 

29 p.

 1971, Teleost otoliths, in Cooper, G. A., Eocene brachipods from Eua, Tonga: U.S. Geol. Survey Prof. Paper 

640-F, p. F7-F8.

Fitch, J. E., and Brownell, R. L., Jr., 1968, Fish otoliths in 

cetacean stomachs and their importance in interpreting 

feeding habits: Canada Fisheries Research Board Jour., 

v. 25, no. 12, p. 2561-2574.

Frizzell, D. L., and Lamber, C. K., 1962, Distinctive "congrid type" fish otoliths from the Lower Tertiary of the 

Gulf Coast (Pisces Anguilliformes) : California Acad. 

Sci. Proc., 4th Ser., v. 32, no. 5, p. 87-101.

Grey, Marion, 1956, The distribution of fishes found below a 

depth of 2,000 meters: Fieldiana Zoology, v. 36, no. 2, 

p. 73-337.

Jordan, D. S., and Gilbert, J. Z., 1919, Fossil fishes of the 

(Miocene) Monterey Formations of Southern California, 

Pt. 2 of Fossil fishes of southern California: Leland Stanford, Jr., Univ. Pubs., Univ. Ser., p. 13-60.

Kotthaus, Adolf, 1967, Fische des Indischen Ozeans. Ergebnisse der ichthyologischen Untersuchungen wahrend der 

Expedition des Forschungsschiffes "Meteor" in den Indischen Ozean, Oktober 1964 bis Mai 1965. A. Svstematischer Teil, I. Isospondyli und Giganturoidei: "Meteor" 

Forsch.-Ergebnisse, Reihe D, Heft 1, p. 7-84.

1968, Fische des Indischen Ozeans. Ergebnisse der

ichthyologischen Untersuchungen wahrend der Eypedition

des Fctrschungsschiffes "Meteor" in den Indischen Ozean,

Oktober 1964 bis Mai 1965. A. Systematischer Teil, III.

Ostariophysi und Apodes: "Meteor" Forsch.-Erirebnisse,

Reihe D, Heft 3, p. 14-56. 

Maul, G. E., 1954, Monografia dos peixes do Museo Municipal

do Funchal. Ordem Berycomorphi: Mus. Municipal Funchal, Bol., v. 7, no. 16-17, p. 5-41. 

Munro, I. S. R., 1938, Handbook of Australian fishes. No. 19.

Order Beiryciformes: Fisheries Newsletter, v. 16 no. 13,

p. 77-80. 

Stinton, F. C., 1967, The otoliths of the teleostean fish Antigonia capros and their taxonomic significance: Bccagiana,

v. 8, no. 13, p. 3-7.

 1968, On the study of Tertiary fish otoliths, in Colloque 

sur 1'Eocene, Paris, Mai 1968, Volume II: France Bur. 

Recherches Geol. et Minieres Mem. 58, p. 153-161.

Weiler, Wilhelm, 1968a, Die Otolithen der bathypebgischen 

Familie Melamphaidae und ihre systematische Bodeutung 

(Pisces) : Senckenbergiana Biologica, v. 49, no 34, p. 

223-230.

 1968b, Otolithi piscium (Neubearbeitung), Pt. 117 of 

Westphal, F., ed., Fossilium Catalogus. I. Animalia: The 

Hague, W. Junk N. V., 196 p.

Woods, L. P., and Sonoda, P. M., 1973, Order Berycomorphi 

(Beryciformes), in Cohen, D. M., ed., Fishes of tl ? Western North Atlantic, Part 6: Sears Found. Marine Research, p. 263-396.
INDEX

[Italic page numbers indicate descriptions and major references]

Page

affinia, Trachichthodes _________________ G13

alabamiensis, Stenocyathus ____________ 11

(Amphelia), Madrepora ______________ 7

Annelids ___.__-_______.____________ V

Anomocora ________________________ 4, 5, 6, 10

exilis --__-____.____.____._______ 10; pi. 3

fecunda ________________________ 10

Anthemiphyllia _______________ 2, 4, 5, 6, 7

catinata ---________.____________ 7; pi. 2

dentata ...._____________._____ 8

patella _______________________ 8

patera _______________________ 8

verbeeki _____________________ 8

Anthemiphylliidae _________________ 7

Anthozoa ___.-__________________ 7

antiguus, Stylaster __________________ 3

argenteus, Dirctmus ____________.___ 14

Astya .........._____________ 2,4,5,6

crassa ...... _________________ 7

nielseni _______________________ 5, 6; pi, 1

subviridis _______________________ 6

aureus, Diretmus ...._____________ 14

australiensis, Deltocyathus italicus _____ 9

Notocyathus (Paradeltocyathus) ___ 9

B

Barnacles ___________________________ V

Bathyactis eocenica ________________ 7

(Bathyactis) Fungicyathus __________ 7

Benthosema ________________________ 15

Berycidae __--__.--__.______________ 13

Beryx ......_________________ 13

boschmai, Stenohelia __-._-__.__ ___ 3; pi. 1

Brachiopods _______________ III, V,. VI

Bryozoans ___-_________________ V

Caryophyllia _____________ 2,4,5,6,8

denaria . ______________________ 8; pi. 2

japonica _______________________ 8; pi. 2

octopali ______________________ 8

sp __.__-__----------------__.__ 8; pi. 2

Caryophylliidae ._.__- __ ----- _._ 8

Caryophyllinae ______ __ __ __ 8

catinata, Anthemiphyllia ............. 7; pi. 2

Centroberyx ...... _____.

Ceratoscopelus ......__.

Ceratotrochus ...... ____.

exiguus ............

Coelorinchus ............

complectus, Deltocyathus

13

15

9

9

14

9

Congridae ______ _._ __ __ 13, 14

Conotrochus ... _.___---__-___________ 9

Corallium _________________________ 7

Corals __ _____ _ _____ V

Coryphaenoides .................____ 14

armatus _________________-- ___ 14

coulsoni, Desmophyllum __________ 9, 10; pi. 3

crassa, Astya ...... _____ __ 7

Page 

Crinoids _____________________________ V

Crispatotrochus ___________ G2, 4, 5, 6, 9 

inornatus ______________________ 9

tenison-woodsi ________________ 9; pi. 2

Crypthelia ........._____________ 4,5,6

japonica _________ ____________ 5

pudica __________________________ 5

vetusta ...... ...____-_-__--_ 5; pi. 1

Cyathoceras ... ________._____ 2,4,5,6,8 

kondoi .... ____________________ 8; pi. 2

periallus ...... ___________.____ 8

D

decamera, Stenocyathus _______________ 11

Decapod crustaceans _____________ ._ V

decuplum, Desmophyllum ......... 9, 10; pi. 3

Deltocyathus ....................... 2,4,5,6

complectus ______ ______-____ 9

fontinalis ______ ___ ___ ______ 9

italicus ______ ___ ___ -__ __ 9

australiensis _________________ 9

orientalis ________________________ 9

whitei __-_____-__-----_--_-_ ____ 9

(Paradeltocyathus) orientalis ...... 9

sp ___________________-.- 8; pi. 2

denaria, Caryophyllia ______________ 8; pi. 2

Dendrophylliidae ____________________ 11

dentata, Anthemiphyllia ______________ 8

Desmophyllinae ______ ___ ____ __ 9

Desmophyllum ............. 2,4,5,6,9,10,11

coulsoni _____________________ 9, 10; pi. 3

decuplum .................... 9, 10; pi. 3

lirioidfs ...--------------_-_ 9, 10; pi. 3

taurinense ______________________ 9

sp ____________________-. 10 

Diaphus __________-__ -_______ __- 15

Diretmidae _________._______ 13, 14

Diretmus ...... ___.

argenteus ___-__- 

aureus _______.

Discoasters _______

Distichopora ________

dumosa, Goniocorella

14 

14 

14 

V 

2 

10

duncani, Javania ...... .. _________ 11; pi. 3

Echinoids _________ ___ ______ _ V

eguchii, Goniocorella ................ 10; pi. 3

Enallopsammia ............ 2,3,4,5,6,11,12

laddi _________________ 11, 12; pi. 3 

payricardensis _____________ ___ 12

poppelachii _________________ __ 12

scillae ____________________ 12

eocenica, Bathyactis ...... ____ ____ 7

Errina (Lepidopora) _________________ 2

Eua deposits, geology of _____________ III

Eua deposits, relation to Tonga Trench. Ill 

euaenis, Fungiacyathus .............. 7; pi. 1

exiguus, Ceratotrochus ................ 9

exilis, Anomocora . ________________ 10; pi. 3

Page

F

fecunda, Anomocora ...._____ ______ G10

Flabellidae ____ 11 

Flabellum ..................____ , 2,11

fontinalis, Deltocyathus ______ -___ __ 9

Foraminifera ______._.__-_ III, V, VI, Gl 

Foreword, by H. S. Ladd _____-___-_ III 

Fungiacyathus ................... 2,4,5,6,7

euaenis ..............-.....--. 7; pi. 1

hawaiiensis . 

kikaiensis __ 

palifera .... 

sibogae ..... 

Stephana ... 

(Bathyactis) 

Fungiidae ------

Goniocorella .....------------------ 4, 5, 6,10

dumosa __________________________ 10

eguchii ______.__ _____ __ __ 10; pi. 3

Gorgonacea ___________ .__ _ 7

granulata, Madrepora ......_________ 7

Guyniidae ___--_ __ __. -.- -- 11

H

hcwaiiensis, Fungiacyathus ___________ 7

hoffmeisteri, Stenocyathus ---------- 11; pi. 2

houshi, Xyrinius _____ __ ___ _ 15

Hydrozoa ______ ___ ___ _____._-__ 3

Hydrozoans .__ _ ______ __ - V

Ichthyococcus __________.__.

ovatus ________________-

inornatus, Crispatotrochus _... 

insignis, Javania ______ ______

italicus Deltocyathus ___--_--_.

australiensis, Deltocyathus

15

15

9

11

9

9

japonica, Caryophyllia ___-_-___---__ 8; pi. 2 

Crypthelia ____________ .. 5

Javania ___________ __ ___ 2, 4, 5, 6, 11

duncani _______________.--__ 11; pi. 3

insignia ______________ __ _ 11

K

kikaiensis, Fungiacyathus _____ ___ 7 

kondoi, Cyathoceras _________________ 8; pi. 2

Ladd, H. S., Foreword ___._ III 

laddi, Enallopsammi ------------- 11, 12; pi. 3

(Lepidopora), Errina ______ ___ ____ 2

lirioides, Desmophyllum ____ _ 10

G17
G18 INDEX

Page

M

Macouridae ________________________ G13, H

Madrepora _______________ 2,3,4,5,6,7

granulata _________________________ 7

(Amphelia) __________________ 7

sp ________ ___ _________ 7; pi. 2 

Melamphaes ------ _-_ ____________ 14

Melamphaidae ____ ______________ 13, H

Mollusks ______________________ V 

mooraboolensis, Stylaster _____________ 3

Myctophidae _--_-_ -_-__.________ 13,15

N

nielseni, Astya __________________ 5, 6, 7, pi. 1

Notocyathus ________________________ 9

(Paradeltocyathus) _______________ 4,5,6

australiensis ________________ 9

orientalis .__-_---____________ 9; pi. 3

O

Octocorallia __________

octopali, Caryophyllia _ 

Oculinidae _____ ._

Oculininae __--____-__

orienttlis, Dcltocyathus

Deltocyathus (Paradeltocyathus) __ 9

Notocyathus (Paradeltocyathus) __ 9; pi. 3

Ostracodes ______ __ _.___-____._- V

Otoliths ___ _-________-__ V 

ovatus, Ichthyococcus _________________ 15

palifera, Fungiacyathus _______________ 7

Paradeltocyathus _____________________ 2, 9

Page 

( Paradeltocyathus) orientalis,

Deltocyathus ______________ G9

Notocyathus __.____-_--_________-_ 4,5,6

australiensis, Notocyathus _________ 9

orientalis Notocyathus ___________ 9; pi. 3

Parasmiliinae ___________ ___ ___ __ 10

Parisis ___________--___--___--______ 7

patella, Anthemiphyllia ________________ 8

patera, Anthemiphyllia ______________ 8

payricardensis, Enallopsammia ________ 12

Peponocyathus _______________________ 3

periallus, Cyathoccras ________________ 8

Plant microfossils _____________________ V

poppelachii, Enallopsammia ___________ 12

pudica, Crypthelia _________________ 5

Scleractinia ______ __.

scillae, Enallopsammia . 

Scleraxonia ____________

Scopelogadus __________

Shark teeth ___________

sibogae, Fungiacyathus 

Stearns, H. T., quoted

7 

12

7

14 

V

7 

III

Stenocyathus _____________ 2,4,5,6,11 

alabamiensis _______-____---__--_ 11

decamera ______________________ 11

hoffmeisteri ________________ 11; pi. 2

Vfrmiformis _____-_-_________----_ 11

Stenohelia _________________.. 3,4,5,6

boschmai _______________________ 3; pi. 1

Stephana, Fungiacyathus -______-__---_ 7

Sternoptychidae ___._-_--_________-_-- 13, 15

Stylaster ____________________ 3,4,5,6

antiqutis ______--___--____--___ __ 3

mooraboolensis _____--________----_ 3

sp _____----------_-------- 3; pi. 1

Stylasteridae ___. 

Stylasterids _____ 

Stylasterina 

subviridis, Astya 

Symbolophorits __

Page 

G3

2

3

6

15

taurinenss, Desmophyllum _________ __ 9

Tele-a-hiva ____.--._- _-. ------ III

tenison-woodsi, Crispatotrochus __-_-_- 9; pi. 2 

Trachichthodes affinis _________________ 13

Tubastraea _______--__----__-__ ___ 12

Turbinoliinae ______ ___ ___ ___ -- 9

U

Undetermined octocorals ___---_--_--_ 7; pi. 1

Vaingana _______.__-__--____- ------ III

verbeeki, Anthemiphyllia ______________ 8

 vcrmiformis, Stenocyathus ____________ 11

vetusta, Crypthelia ___________________ 5; pi. 1

whitci, Deltocyathus

Xyrinius hoiishi

Zoantharia

15
PLATES 1-3

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

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

FIGURES 1,2. Stylaster sp. (p. G3). 

X 8, USNM 208294. 

3-6. Stenohelia boschmai Wells, n. sp. (p. G3).

3,4. Holotype, anterior surface; posterior surface with large

ampullae, x 4, USNM 208295.

5-6. Paratypes, cyclosystems, X 4, USNM 208296. 

7, 8. Crypthelia vetusta Wells, n. sp. (p. G5).

7. Holotype, upper surface with large ampullae, X 8, USNM 

208297.

8. Holotype (lower right) and paratypes, anterior surfaces, X 4,

USNM 208297, 208298. 

9. Astya nielsem Wells, n. sp. (p. G6).

Holotype (left), X 6, USNM 208299; paratype (right) X 6, USNM

208300.

10. Corallimn (left and bottom), and Parisis (other four specimens), (p. 

G7).

X 2, USNM 208301. 

11-14. Fungiacyathus euaensis Wells, n. sp. (p. G7).

11,12. Holotype, basal aspect, X 1, X 5, USNM 208302.

13. Paratype, basal aspect, X 4, USNM 208303.

14. Paratype, lateral aspect of septum and synapticulae, X 4, 

USNM 208304.
PROFESSIONAL PAPER 640-G PLATE 1

STYLASTER, STENOHELIA, CRYPTHELIA, ASTYA, CORALLIUM, FUNGIACYATH'US, AND PARISIS
PLATE 2

FIGURES 1-3. Madrepora sp. (p. G7).

1. Branchlets, X 1, USNM 208305.

2. Calice, X 4, USNM 208305 (part).

3. Polished transverse section below calice, X 4, USNM 208306. 

4, 5. Anthemiphyllia catinata Wells, n. sp. (p. G7).

Holotype, basal and calicular aspects, X 4, USNM 208307 

6-10. Caryophyllia denaria Wells, n. sp. (p. G8).

6,7. Holotype, lateral and calicular aspects, X 1, X 2, USNM

208308.

8, 9. Paratypes, X 1, USNM 208309, 208310. 

10. Paratype, polished section at base of calice, X 4, USNM

208311.

11,12. Caryophyllia sp. cf. C. japonica von Marenzeller. (p. G8). 

Lateral and calicular aspects, X 1, X 2, USNM 208312. 

13,14. Cyathoceras kondoi Wells, n. sp. (p. G8).

13. Holotype (right), X 1, USNM 208313; 

paratype (left), X 1, USNM 208314.

14. Holotype, calice, X 2. 

15,16. Deltocyathus sp. (p. G8).

Calicular and basal aspects, X 4, USNM 208315. 

17-19. Cristpatotrochus tenlson-woodsi Wells, n. sp. (p. G9).

17,18. Holotype, lateral and calicular aspects, X 1, X 2, USNM 

208316.

19. Paratype, polished section below calice, X 4, USNM 208317. 

20-24. Stenocyathus hoffmeisteri Wells, n. sp. (p. Gil).

20. Holotype (right), x 4, USNM 208318;

paratypes a, b, c (left three), X 4, USNM 208319, 208320, 

208321.

21. Septa (8) of holotype, X 8.

22. Septa (7) of paratype c, X 8.

23. Paratype b, dark spots are sites of mural pits, X 8.

24. Paratype, internal mold, short spines are filling of mural 

pits, X 8, USNM 208322.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-G PLATE 2

MADREPORA, ANTHEMIPHYLLIA, CARYOPHYLLIA, CYATHOCERAS, DELTOCYATHUS, CRISPATOTROCHUS,

STENOCYATHUS
PLATE 3

FIGURES 1-3. Notocyathus (Paradeltocyathus) orientates (Duncan) (p. G9). 

USNM 208323.

1,2. Basal aspects, X 4.

3. Calicular aspect, X 4. 

4, 5. Desmophyttum decuplum Wells, n. sp. (p. G9).

4. Holotype (left), X 1, USNM 208324; 

paratype (right), X 1, USNM 208325.

5. Calice of holotype, X 4. 

6,7. Desmophyttum lirioides Wells, n. sp. (p. G9).

Holotype, lateral and calicular aspects, X 1, USNM 208326. 

8, 9. Desmophyllum coulsoni Wells, n. sp. (p. G10).

Holotype, lateral and calicular aspects, X 1, X 2, USNM 208327. 

10,11. Anomocora exilis Wells, n. sp. (p. G10).

10. Holotype (right), X 1, USNM 208328; 

paratype (left), X 1, USNM 208329.

11. Polished section of holotype, X 2. 

12,13. Gonwcorella eguchii Wells, n. sp. (p. G10).

12. Holotype (left), X 1, USNM 208330; 

paratype (right), USNM 208331.

13. Polished section of holotype, X 4. 

14-16. Javania duncani Wells, n. sp. (p. Gil).

14,15. Holotype, lateral and calicular aspects, X 1, X 2, USNM 

208332.

16. Paratype, calice, X 2, USNM 208333. 

17-22. Enallopsammia laddi Wells, n. sp. (p. G 11).

17. Holotype (top center), X 1, USNM 208334;

paratypes (right, left, and lower center), X 1, USNM 208335.

18. Holotype, X 4.

19. Paratype, polished section, X 4, USNM 208336.

20. Paratype, polished section, X 4, USNM 208337.

21. Paratype, calice, X 4, USNM 208338.

22. Paratype, calice, X 4, USNM 208335.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 640-G PLATE 3

NOTOCYATHUS, DESMOPHYLLUM, ANOMOCORA, GONIOCORELLA, JAVANIA, ENALLOPSAMMIA