Middle Devonian Rugose Corals 

of the Central Great Basin

GEOLOGICAL SURVEY PROFESSIONAL PAPER 799
Middle Devonian Rugose Corals 

of the Central Great Basin

ByC.W.MERRIAM

GEOLOGICAL SURVEY PROFESSIONAL PAPER 799

A stratigraphic-paleontologic study of 

middle Middle Devonian rugose corals 

and their use in geologic correlation

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

ROGERS C. B. MORTON, Secretary

GEOLOGICAL SURVEY 

V. E. McKelvey, Director

Library of Congress catalog-card No. 73-600112

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

Washington, B.C. 20402 - Price $1.85 (paper cover)

Stock Number 2401-02374
CONTENTS

Page

Abstract .........................................-.............. -........... ..........-.. 1

Introduction.............................................. .....-.......................... 1

Purpose and scope .............................................................. 1

History of investigation........................................ ........... 2

Acknowledgments................................................................ 2

Great Basin Devonian coral zones .......................................... 3

Facies belts of the central Great Basin Devonian .......... 3

Rugose corals and stratigraphy of Middle Devonian coral 

zones F and G in the central Great Basin .......................... 7

Rugose corals and stratigraphy of Middle Devonian

zones F and G in the Antelope-Roberts Mountains 

facies belt.................................................................... ..... 7

Reference section and coral zone F at Lone Mounts in 7 

Strata of Middle Devonian coral zone F and possibL

zone G in the Roberts Mountains ................................ 10

Middle Devonian rocks of coral zone F in the

northern Antelope Range .............................................. 11

Middle Devonian strata of coral zone F in the

southern Fish Creek Range ................................... 11

Cockalorum Wash Formation .................................. 14

Middle Devonian strata of the Hot Creek Range ........ 15

Middle Devonian rugose corals of the Diamond

Mountains belt .................................................. ............ 15

Great Basin Devonian coral zone G .................................. 17

Middle Devonian strata of Bare Mountain, southern

Nye County, Nev. .............. N ................................................... 17

Great Basin Middle Devonian Rugosa and their bearing

upon geologic correlation with distant regions........... 17

Distribution and correlation value of Rugosa ................ 18

Correlation with rugose coral deposits of western

Canada ...................................................................... 18

Hume Formation ........................................................ 18

Horn Plateau Formation .......................................... 19

Rugose corals and correlation with Middle Devonian 

strata of southeastern Alaska ........................................ 19

Page

Great Basin Middle Devonian Rugosa and their bearing

upon geologic correlation with distant regions Con.

Correlation with rugose coral deposits of the Rhine

Valley, Germany ............................................................ 20

Middle Devonian rugose corals of eastern North

America compared with those of the Great Basin .... 21

Classification of Great Basin Middle Devonian rugose

corals in coral zones F and G ................................................ 22

Systematic and descriptive paleontology .............................. 23

Subfamily Siphonophrentinae, new subfamily of

Streptelasmatidae ....._............................. ............. 23

Genus Siphonophrentis O'Connell .................................. 23

Genus Cystiphylloides Chapman ...................................... 24

Genus Zonophyllum Wedekind ........................................ 25

Genus Digonophyllum Wedekind .................................... 26

Subgenus Mochlophyllum Wedekind....................... 27

Genus Mesophyllum Schliiter .......................................... 28

Subgenus Lekanophyllum Wedekind ............ ........ 29

Genus Acanthophyllum Dybowski .................................. 29

Genus Paracanthus, new genus ........................................ 32

Genus Cyathophyllum Goldfuss ...................................... 32

Subgenus Morauophyllum Kettnerova................... 33

Subgenus Orthocyathus, new subgenus................... 34

Genus Keriophyllum Wedekind ...................................... 35

Genus Sociophyllum Birenheide ...................................... 36

Genus Utaratuia Crickmay .............................................. 37

Genus Tabulophyllum Fenton and Fenton .................... 38

Genus Australophyllum Stumrn ...................................... 39

Genus Hexagonaria Giirich .............................................. 39

Genus Taimyrophyllum Chernychev .............................. 44

Genus Lyrielasma Hill .............................................^.......^ 45

Locality register .......................................................................... 45

Selected references .................................................................... 47

Index .............................................................................................. 51

ILLUSTRATIONS

PLATE 1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

[Plates follow index]

Cystiphylloides, Zonophyllum, Mesophyllum, and Siphonophrentis.

Acanthophyllum and Digonophyllum.

Digonophyllum and Mesophyllum.

Cyathophyllum, Keriophyllum?, Acanthophyllum, and Paracanthus.

Keriophyllum and Cyathophyllum.

Cyathophyllum, Paracanthus, and Tabulophyllum.

Cyathophyllum and Lyrielasma.

Utaratuia, Australophyllum, and Sociophyllum.

Taimyrophyllum.

Taimyrophyllum.

Hexagonaria.

Hexagonaria.

Hexagonaria.

Hexagonaria.

Ill
IV CONTENTS

Page 

FIGURE 1. Map of Nevada showing distribution of Middle Devonian rocks containing Rugosa of coral zones F and G .......... 2

2. Generalized correlation diagram showing Great Basin Devonian coral zones A through I in relation to

Devonian stratigraphic units of the central Great Basin .............................................................................................. 4

3. Diagram showing characteristic rugose coral genera and their stratigraphic ranges in Great Basin

Devonian coral zones ......................................................................................................................................................... 5

4. Map showing Devonian facies belts and Middle Devonian rugose coral-collection sites in the central Great Basin 6

5. Geologic map of northeastern part of Lone Mountain, Eureka County, Nev., showing principal collection

sites for coral zone F in unit 4 of the Nevada Formation .............................................................................................. 9

6. Geologic sketch map of the southernmost part of the Fish Creek Range, Cockalorum Wash quadrangle,

showing distribution and structural relations of Cockalorum Wash Formation, which contains faunas

of Middle Devonian coral zone F ............................................................................. ............. ................................... 12

7. Columnar correlation diagram showing relations of Devonian rocks of the southern Diamond Mountains to

those of the Sulphur Spring Range, Lone Mountain, and the Cockalorum Wash area ........................................ 16

TABLE

TABLE 1. Rugose corals of Middle Devonian coral zones F and G and their areas of occurrence

Page
MIDDLE DEVONIAN RUGOSE CORALS OF THE 

CENTRAL GREAT BASIN

By C. W. MERRIAM

ABSTRACT

Great Basin Middle Devonian rugose corals here described 

and illustrated comprise 20 genera and subgenera of coral zones 

F and G. The entire Devonian column of this province is subdivided on the basis of Rugosa into nine zones, A through I in 

ascending stratigraphic order. Of these genera and subgenera, 

13 have not been found in the lower Middle Devonian below 

coral zone F; none of them are recognized above coral zone G 

and the Stringocephalus zone. A coral evolutionary burst in 

Great Basin coral zone D is separated from that of zone F by 

a barren zone (coral zone E) with few Rugosa. The final 

Devonian burst of Rugosa, that of the Phillipsastraeidae in 

Upper Devonian coral zone I, is separated from zone G by 

another interval of sparse Rugosa in coral zone H, wherein the 

stromatoporoids predominate.

Most of the described Rugosa, those of middle Devonian 

coral zone F (unit 4 of the Nevada Formation at Lone Mountain), occur in the Antelope-Roberts Mountains facies belt. 

This belt is distinguished on the basis of lithology and faunas 

from both the Monitor-Simpson Park facies belt on the west 

and the Diamond Mountains facies belt on the east.

Rugosa of coral zone F characterize the Cockalorum Wash 

Formation in the southernmost Fish Creek Range. This new 

geologic unit includes more siltstone and gritty sandstone than 

either unit 4 of the Nevada Formation at Lone Mountain or the 

Woodpecker Limestone Member of the Nevada Formation in 

the Diamond Range, both of which are approximately correlative with the upper beds of zone F. The Cockalorum Wash 

Formation lies in a highly complex structural relation to contiguous rocks and is without exposed stratigraphic top or bottom.

Digonophyllidae and Ptenophyllidae related to species of 

western Canada and the Rhine Valley of Germany are the distinctive solitary rugose corals of Great Basin coral zones F and 

G. Among the digonophyllids are Mesophyllum (Lekanophyllum) and Digonophyllum (Mochlophyllum); the Ptenophyllidae are represented by Acanthophyllurn. Of the abundant 

colonial Rugosa, yardarm-carinate Hexagonaria of the H. 

fisherae group is common throughout the Antelope-Roberts 

Mountains facies belt and is especially abundant locally in the 

Cockalorum Wash Formation as an incipient patch reef builder.

Of special stratigraphic and correlation value are the colonial 

genera Sociophyllum, Utaratuia, and Taimyrophyllum in coral 

zone F, occurring also in the correlative Hume and Nahanni 

Formations of western Canada.

Coral zone G is best represented in the Diamond Mountains 

facies belt, where it is typified by a 30-foot band in the Nevada 

Formation bracketing the top of the Woodpecker Limestone 

Member and the base of the Bay State Dolomite Member. In 

coral zone G occur the lowest Stringocephalus, Rensselandia, 

Heliolites, and the rugose corals Digonophyllum (Mochlophyllum) alhambraensis, Cyathophyllum (Moravophyllum) 

alhambraensis, and species of Mesophyllum, Cystiphylloides,

and Acanthophyllurn. Great Basin coral zone G lacks the distinctive colonial Rugosa of coral zone F.

Middle Devonian strata of southeastern Alaska include the 

Stringocephalus zone and contain Digonophyllidae and Ptenophyllidae similar to those of Great Basin coral zones F and G. 

The middle Middle Devonian coral-bearing formations of eastern North America have fewer paleontologic ties with the Great 

Basin than do those of southeastern Alaska, western Canada, 

and Europe.

Great Basin coral zone G at the bottom of the Stringocephalus zone is correlative with the Rhine Valley Givetian. 

Coral zone F below the Stringocephalus zone is considered later 

Eifelian, approximately correlative with the Junkerberg 

Schichten of the Eifel district, Germany.

INTRODUCTION

Middle Devonian rugose corals of the Great Basin 

are best known in a 20-mile-wide linear belt extending 

southward 125 miles from the Roberts Mountains 

through the Antelope Range to the south end of the 

Hot Creek Range, Nev. (figs 1, 4). Within the prevailingly carbonate lithofacies of this Antelope-Roberts 

Mountains belt, exposures containing abundant corals 

have a patchy distribution, suggesting that environments conducive to prolific coral growth were restricted 

to scattered and relatively small areas of Devonian sea 

bottom. Elsewhere these Middle Devonian rocks are 

either sparsely fossiliferous, or the common fossils are 

brachiopods, Mollusca, and trilobites with an occasional 

solitary rugose coral or tabulate. Tentaculites are numerous locally. Strata with abundant Rugosa may be 

either limestone or dolomite in favorable coral growth 

sites. However, in this Middle Devonian interval the 

greatest development and diversity of a potentially 

reef-forming coral biota appear to be in dolomitic limestone facies, as for example in unit 4 of the Nevada 

Formation and in Devonian coral zone F at Lone 

Mountain.

PURPOSE AND SCOPE

The main purposes of this long-term investigation 

are to describe, classify, and illustrate the rugose corals 

of Great Basin Middle Devonian coral zones F and G 

and to elucidate their stratigraphic occurrence. Conclusions regarding stratigraphic order and zonation of 

enclosing rocks are supported by detailed geologic
MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

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FIGURE 1. Distribution of Middle Devonian rocks containing 

Rugosa of coral zones F and G.

mapping at Lone Mountain and in parts of the Cockalorum Wash quadrangle by Merriam and in the Pinto 

Summit quadrangle, southern Diamond Mountains by 

Nolan, Merriam, and Blake (1973). Finally, the Middle 

Devonian coral-bearing strata here dealt with are correlated by means of Rugosa and certain key brachiopods with those of western Canada, southeastern 

Alaska, and the Rhine Valley of Germany.

HISTORY OF INVESTIGATION

This coral study is an outgrowth of stratigraphic and 

paleontologic investigations of Devonian rocks in the 

Roberts Mountains region that began in 1932 (Merriam, 1940). Rugose corals collected throughout the 

Silurian and Devonian rock of this structurally complex 

area were especially useful in identifying, mapping, and 

subdividing rocks of these systems. A long-term 

descriptive and taxonomic investigation of Great Basin 

Devonian Rugosa was initiated in subsequent years, as 

reconnaissance geologic mapping and stratigraphic

work proceeded to the south at Lone Mountain and 

Antelope Valley (Merriam, 1963).

During the 1950/s geologic mapping and stratigraphic 

investigation by Nolan and Merriam under the Kobeh 

Valley project gave added impetus to these paleontologic studies. Since 1963, the opportunity has presented 

itself to concentrate upon and bring this coral research 

to completion.

Prior to this study a number of rugose corals from 

Lone Mountain and the Eureka mining district were 

described by the late E. C. Stumm (1937, 1938) in 

reports based upon study of Smithsonian Institution 

collections. These collections were made largely by 

C. D. Walcott and associates during initial mapping of 

the Eureka district in 1880 (Hague, 1892), supplemented in later years by Edwin Kirk of the U.S. Geological Survey and by the writer. General advances in 

knowledge of Devonian Rugosa within the past 25 years 

by many contributors throughout the world made 

research on Great Basin corals far more rewarding than 

would have been possible before 1940.

A monographic study of Lower and lower Middle 

Devonian Rugosa of the central Great Basin was completed in 1969 (Merriam, 1973), a work dealing with 

the faunas of coral zones A, B, C, and D. The present 

contribution completes description of Middle Devonian 

rugose corals through coral zone G of Givetian age. 

Comparative research has been carried on concurrently 

with similar Middle Devonian Rugosa of southeastern 

Alaska. Recently published investigations of Middle 

Devonian rugose corals of western Canada by Lenz 

(1961), by McLaren (in McLaren and Norris, 1964), 

and by Pedder (1964) have facilitated meaningful 

paleontologic comparison with coral assemblages of the 

northern Cordilleran seaways.

ACKNOWLEDGMENTS

Acknowledgment is made to the Smithsonian Institution for loan of type specimens figured by the late 

E. C. Stumm. Other collections made use of in this 

study are from the Confusion Range, Utah, made by 

R. K. Hose, U.S. Geological Survey; from the Hot Creek 

Range, Nev., by F. J. Kleinhampl and H. W. Dodge of 

the U.S. Geological Survey; and from the Bare Mountain area, Nevada, by H. R. Cornwall and F. J. Kleinhampl. A collection from Red Canyon, Roberts Mountains, was loaned by J. G. Johnson.

Ellis Yochelson, of the U.S. Geological Survey, 

Washington, D.C., provided identifications of gastropod 

genera associated with the described Rugosa in the 

newly named Cockalorum Wash Formation.

Thin sections used in connection with this report 

were prepared mainly by Robert Shely of the Menlo 

Park center; those of the Red Canyon corals were made
GREAT BASIN DEVONIAN CORAL ZONES

in Washington, D.C., by William Pinckney. All photographs of corals are the work of Kenji Sakamoto.

GREAT BASIN DEVONIAN CORAL ZONES

Devonian rugose corals of the central Great Basin 

range upward from the Rabbit Hill Limestone of Early 

Devonian (Helderberg) age to Late Devonian beds in 

the upper part of the Devils Gate Limestone. No 

Rugosa have been found in the uppermost Devonian of 

this region. Figure 2 presents the Great Basin Devonian 

coral zones and illustrates the composite nature of the 

coral sequence. Unfortunately no individual stratigraphic section has been found that includes all coral 

evidence of zones in continual sequence.

The proposed Great Basin Devonian coral zones, nine 

in number (Merriam, 1973b), are designated by capital 

letters in stratigraphic order A through I. In reference 

sections where each is defined, six of the coral zones 

correspond to mappable lithologic units. Coral zones B 

and C fall within a single map division, unit 1 of the 

Nevada Formation. Coral zone G occupies a narrow 

band bracketing the topmost beds of the Woodpecker 

Limestone Member of the Nevada and the lowermost 

beds of the overlying Bay State Dolomite Member.

Coral zones A, B, and C are of Early Devonian age; 

coral zone D is Early and early Middle Devonian. Zones 

E, F, G, and probably the lower half of zone H are Middle Devonian. Coral zone I is Late Devonian; the highest Devonian of this region, in the lower part of the 

Pilot Shale or its limestone equivalent, has yielded no 

Rugosa. For convenience the lettered primary coral 

zones are further divided into subzones as DI, D2 , and 

D3 . In this instance, subzone DI has a rugose coral 

assemblage regarded as late Early Devonian, D2 is considered Early or early Middle Devonian, and D3 is 

Middle Devonian.

Following the major burst of rugose coral evolution 

(Merriam, 1973a) in Late Silurian limestone environments of the Great Basin (Silurian coral zone E), the 

Rugosa of Helderbergian coral zone A became greatly 

restricted in number of families. In most exposures of 

the Helderbergian Rabbit Hill Limestone and the correlative part of the Beacon Peak Dolomite Member of 

the Nevada Formation, coral zone A is characterized by 

Syringaxon (fig. 3); this abundant small solitary form 

is associated with Pleurodictyum and the favositids. 

Australophyllum and Billingsastraea are present but 

very uncommon in this zone. Coral zones B and C are 

characterized by solitary Halliidae. Large endemic 

unnamed members of this family in zone B probably 

gave rise to Papiliophyllum of zone C, associated here 

with other Halliidae such as Aulacophyllum and 

Odontophyllum. The siphonophrentids appear in the 

record at this time with Siphonophrentis (Breviphrentis), which becomes the most abundant coral in 

succeeding coral zone D. Papiliophyllum persists into 

Di, as the last of the Halliidae. Subzone D2 is marked 

by another major burst of rugose coral differentiation 

in which Siphonophrentis (Breviphrentis), Bethanyphyllum, Billingsastraea, an early sub genus of Hexagonaria, Cystiphylloides, and Mesophyllum are common. Digonophyllidae including Mesophyllum and 

Zonophyllum first appear at this time. The large Mesophyllum (Arcophyllum) is introduced in subzone D3 .

Rugosa are poorly represented in coral zone E by 

Disphyllum and cyathophyllids in what is called the 

"barren zone". Coral zone F introduces several new 

groups to populate an interval of great rugose coral 

proliferation, tapering off in coral zone G when the 

Stringocephalus brachiopod facies came to predominate.

Coral zone H, like zone E, has yielded few Rugosa. 

Massive stromatoporoids and Amphipora predominate 

in zone H and the lower part of the Devils Gate wherein 

they became major carbonate builders. Newly introduced Phillipsastraeidae predominate in coral zone I of 

the higher Devils Gate, following the disappearance of 

Digonophyllidae, Ptenophyllidae, and the colonial 

genera of late Middle Devonian time. This marks the 

ultimate evolutionary burst of Devonian rugose corals 

in the Great Basin province. None have been found in 

the uppermost Devonian rocks.

FACIES BELTS OF THE 

CENTRAL GREAT BASIN DEVONIAN

Distribution and geographically changing abundance 

of Devonian rugose corals in the central Great Basin 

clearly reflect east-west facies changes. On the basis 

of lithology and of biofacies, three north-south-trending 

facies belts are recognized in this region, in order as 

follows from west to east: (1) Monitor-Simpson Park 

facies belt, (2) Antelope-Roberts Mountains facies 

belt, and (3) Diamond Mountains facies belt (fig. 4). 

West of the Antelope-Roberts Mountains belt, known 

rugose corals are mainly small solitary forms of the 

genus Syringaxon in Lower Devonian limestones of the 

roughly parallel Monitor-Simpson Park belt; Middle 

Devonian Rugosa are almost unknown in this westerly 

belt. East of the Antelope-Roberts Mountains belt, the 

Middle Devonian strata of the Diamond Mountains 

belt are less rich in corals, and the underlying Lower 

Devonian dolomites and sandstones have yielded 

almost no identifiable fossils.

The locally coral-rich Antelope-Roberts Mountains 

belt probably extends southward beyond the Hot Creek 

Range, for colonial Rugosa typical of this belt occur at 

Bare Mountain 90 miles south of Warm Springs (fig. 4), 

which is the southernmost coral locality in the Hot 

Creek Range.
MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

FORMATION

AREA

Lone Mountain Devils Gate Sulphur Spring Range, 

southern part

Diamond Mountains, 

Oxyoke Canyon

Monitor Range, 

Rabbit Hill

Lower part 

of Pilot Shale

Topunexposed

Devils Gate 

Limestone

Lower Pilot 

equivalent

Devils Gate 

Limestone

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Unit 4, 

675ft

Unit 3, 

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Unit 2, 

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Sentinel 

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400ft

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Dolomite 

Member, 

470ft

Lone Mountain 

Dolomite

Top unexposed

Rabbit Hill 

Limestone

Silurian 

graptolite shale

FIGURE 2. Great Basin Devonian coral zones A through I in relation to Devonian stratigraphic units of the central Great Basin.

See figure 4 for location of sections.
GREAT BASIN DEVONIAN CORAL ZONES

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EXPLANATION 

Devonian facies belts

Diamond Mountains facies belt 

Upper Devonian, mainly limestone 

Middle Devonian, predominantly dolomite with minor quartzitic sandstone; subordinate limestone and 

siltstone

Lower Devonian, dolomite and 

quartzitic sandstone

Antelope Roberts Mountains

facies belt

Upper Devonian, mainly limestone 

Middle Devonian, argillaceous to silty 

limestone with minor quartzitic 

sandstone; arenaceous limestone, 

crinoidal limestone, and dolomite 

Lower Devonian, dolomite and argillaceous, silty, and arenaceous 

limestones

Monitor Simpson Park 

facies belt

No Upper Devonian rocks recognized

Middle Devonian unrecognized, except in Coal Canyon vicinity at 

north end of belt

Lower Devonian widespread, mainly 

limestone; no dolomite recognized 

in Lower Devonian of this belt

Middle Devonian beds 

of Devonian coral zone G

Middle Devonian beds 

of Devonian coral zone F

Middle Devonian corals

of Devonian coral zone F

In conglomerate cobbles at base of

Tertiary volcanic sequence, Coal

Canyon, Simpson Park Mountains

20 MILES

FIGURE 4. Devonian facies belts and Middle Devonian rugose coral-collection sites in the central Great Basin.
CORAL ZONES F AND G, ANTELOPE-ROBERTS MOUNTAINS FACIES BELT

In the eastern Great Basin, beyond the limits of this 

study, the prevailingly dolomitic Middle Devonian 

rocks have been little explored for rugose corals. The 

Simonson dolomite and lower part of the Guilmette 

Formation in western Utah contain solitary Rugosa of 

Middle Devonian age; however, the Middle and the 

Lower Devonian dolomites of the eastern Great Basin 

generally appear to contain few Rugosa in coral biotas 

which are probably less diverse taxonomically than are 

those in the Antelope-Roberts Mountains belt of the 

central Great Basin.

The reference stratigraphic section of the AntelopeRoberts Mountains facies belt is situated at Lone 

Mountain, Eureka County, Nev. (figs. 2, 4, 5). Here 

the Nevada Formation is subdivided lithologically as 

map units numbered 1 through 5 in ascending stratigraphic order. The coral zones here employed are those 

previously used (Merriam, 1973b). In accordance with 

this proposed scheme, the Great Basin Devonian coral 

zones, nine in number, are lettered A through I in 

ascending sequence. The rugose corals here described 

occur in coral zones F and G, which in the European 

scale are approximately upper Eifelian and Givetian. A 

majority of the described forms come from Nevada unit 

4 and Devonian coral zone F at Lone Mountain.

Reference section of the Diamond Mountains Devonian belt is in upper Oxyoke Canyon southeast of 

Eureka, Nev. Here the rocks bearing the corals of 

Devonian coral zone G are well exposed at the base of 

the Bay State Dolomite Member and the top of the 

Woodpecker Limestone Member of the Nevada Formation. At Lone Mountain, unit 5 of the Nevada occupies 

the approximate stratigraphic interval of the Bay State.

RUGOSE CORALS AND STRATIGRAPHY OF

MIDDLE DEVONIAN CORAL ZONES F AND G

IN THE CENTRAL GREAT BASIN

Rugose coral study material used in this report is 

classified in 53 taxa (table 1), of which 45 percent is 

assigned to formally named species and subspecies. The 

remainder is provisionally designated specifically by 

letter, in most instances being represented in our collections by few individuals or by incomplete specimens. 

Of these taxa, 54 percent is colonial genera, a figure 

reflecting the especially vigorous coral growth activity 

during this interval of Devonian time in the Cordilleran belt of western North America.

The formal taxonomy of Rugosa in Great Basin 

Devonian coral zones F and G is dealt with elsewhere 

in this report.

RUGOSE CORALS AND STRATIGRAPHY OF 

MIDDLE DEVONIAN CORAL ZONES F AND G 

IN THE ANTELOPE-ROBERTS MOUNTAINS 

FACIES BELT

Most of the Rugosa collected from zones F and G of 

the Middle Devonian section in the Antelope-Roberts 

Mountains facies belt occur in Great Basin Devonian 

coral zone F. A coral assemblage from Red Canyon in 

the Roberts Mountains may possibly pertain to coral 

zone G, as discussed later.

Geologic mapping provides stratigraphic control in 

several areas from whence came the rugose corals of 

the Antelope-Roberts Mountains belt. In the Roberts 

Mountains, in the Antelope Valley area, and southward, the mapping is of a reconnaissance nature (Merriam and Anderson, 1942; Merriam, 1963; Kleinhampl 

and Ziony, 1967). More detailed mapping has in recent 

years been done at Lone Mountain and the important 

Devils Gate area as special stratigraphic projects supplementary to geologic mapping of the Whistler Mountain 15-minute quadrangle. Throughout this belt, 

thrust faults are major geologic structures that 

together with later high-angle faults and rapid facies 

change, greatly complicate stratigraphic interpretation. 

Complex faulting of coral-bearing Devonian strata 

called for detailed mapping of parts of the Cockalorum 

Wash quadrangle in an attempt to clarify stratigraphic 

relations (fig. 6).

REFERENCE SECTION 

AND CORAL ZONE F AT LONE MOUNTAIN

Ordovician, Silurian, and Devonian rocks in continuous sequence at Lone Mountain form an eastdipping homocline which includes most of the exposed 

bedrock (fig. 5). The lower pediment slopes of this 

mountain are underlain in part by Devonian and 

Silurian beds and in part by graptolitic beds of the 

Ordovician Vinini Formation differing greatly in facies 

from the Ordovician Pogonip Group and the Hanson 

Creek Formation in the main homoclinal block. Associated with the Vinini are fusulinid-bearing Permian 

limestones. Although good exposures of bedrock are 

few and widely scattered on the pediments, it is fairly 

evident that the Vinini and associated Permian strata 

lie in thrust fault relation to the normal carbonate 

rocks of the homocline.

Strata in the Devonian reference section at Lone 

Mountain are the Nevada Formation overlain by the 

Devils Gate Limestone of the east-dipping homocline. 

The Nevada Formation of Early and Middle Devonian 

age rests conformably upon the Silurian Lone Moun-
MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

TABLE 1. Rugose corals of Middle Devonian coral zones F and G and their areas of occurrence

Siphonophrentis (Siphonophrentis) sp. b...... .... .... X

Cystiphylloides sp. a .......................................... .... .... ....

iddingsin. sp ................................................ X

sp., cf. C. sp. b .............................................. .... .... X

sp. c .............................................................. .... .... ....

Zonophyllum sp. r .............................................. .... .... .... .... .... X

Digonophyllum (Digonophyllum)

occidentalis n. sp ............................................ .... .... X

sp. c .............................................................. ....

(Mochlophyllum) alhambraensis n. sp.... ....

alhambraensis subsp.

robertsensis n. subsp .............................. .... .... .... .... .... X

Mesophyllum (Mesophyllum) sp. f ................ X

n. sp .............................................................. ....

(Mesophyllum)? sp. k................................ X

(Lekanophyllum) sp. 1................................ X

cf. sp. 1.......................... .... .... .... .... .... .... .... .... .... .... X

Acanthophyllum sp. a............................ ......... ....

robertsensis n. sp ........................................ .... .... X

sp. c .............................................................. ....

sp. p .............................................................. .... X

sp .................................................................. x .... .... .... .... .... .... .... .... .... ....

Acanthophyllum? sp .......................................... .... .... .... .... .... X

Paracanthus nevadensis n. gen., n. sp.............. .... .... .... .... .... X .... .... .... x

cf. P. nevadensis .......................................... .... .... .... .... .... .... .... .... .... X

Paracanthus? sp. f .............................................. X

Cyathophyllum (Moravophyllum)

alhambraensis n. sp........................................ .... .... .... .... .... .... .... .... .... ....

(Moravophyllum) sp. 1.............................. X

sp ................................ X .... .... .... .... .... .... .... .... ....

(Moravophyllum)? sp .............................. .... .... .... .... .... .... .... .... .... X

(Orthocyathus) flexum (Stumm) ............ X

Keriophyllum mclareni n. sp............................ X

? kobehense (Stumm) ................................ X

? sp., cf. K. kobehense (Stumm).............. X

Sociophyllum eurekaensis n. sp........................ X X

eurekaensis subsp. b .................................. .... .... X

? sp ................................................................ .... .... ....

Utaratuia eurekaensis n. sp .............................. X

Tabulophyllum antelopensis n. sp .................. .... .... .... .... .... .... .... X

Australophyllum prismatophylloides

(Stumm) .......................................................... X

Hexagonaria fisherae (Merriam) .................... X X .... X cf. .... cf.

fisherae subsp. cockalorumensis n. subsp .... .... .... .... .... .... .... .... X X

fisherae subsp. antelopensis n. subsp........ .... .... .... .... .... .... .... X .... X

meekin. sp.................................................... .... .... .... .... .... .... .... .... .... X

sp., cf. H. meeki n. sp.................................. .... .... X

sp. b (Merriam, 1940)................................ .... .... .... X

sp. c .............................................................. .... .... .... .... .... .... .... .... .... X

sp. f .............................................................. .... .... . .... .... .... .... .... .... X

? sp. h............................................................ .... .... .... .... .... .... .... .... .... .... X

sp. n .............................................................. .... .... .... .... .... .... .... .... .... .... .... X

sp. r .............................................................. .... .... .... .... .... X

sp. w .............................................................. .... .... .... .... .... .... .... .... .... .... .... X

Taimyrophyllum nolani n. sp............................ X X

Disphyllum sp .................................................... ....

Lyrietasma antelopensis n. sp.......................... .... .... .... .... .... .... .... X
CORAL ZONES F AND G, ANTELOPE-ROBERTS MOUNTAINS FACIES BELT

Base from U.S. Geological 

Survey 1:62,500, Bartine Ranch 

and Whistler Mountain, 1956

0

Geology by C.W. Merriam, 

1954-72

5000 FEET

Contact 

Dashed where approximately located

Fault, showing dip

Dashed where approximately located; 

dotted where concealed

Strike and dip of beds

AM1360 

Fossil locality, coral zone F

^1362 

Fossil locality

-a

13 C 

C O

EXPLANATION

Alluvium

Undifferentiated 

dolomite

Western facies rocks

Hanson Creek Formation

Dolomitized Vinini Formation 

Graptolite shale 

and limestone

FIGURE 5. Northeastern part of Lone Mountain, Eureka County, Nev., showing principal collection sites for coral zone F in

unit 4 of the Nevada Formation.

tain Dolomite and is overlain without recognized break 

by the Devils Gate. For mapping purposes the Nevada 

is subdivided as five lithologic units numbered in 

ascending order 1 through 5 and shown in figure 5. Unit 

1 and the lowermost part of unit 2 are Early Devonian. 

Middle Devonian Rugosa occur in the middle and upper 

parts of unit 2 and upward through unit 4. Unit 3 has 

yielded only poorly preserved disphyllid and cyathophyllid corals. Unit 5, a dolomite member aproximately 

equivalent to the Bay State Dolomite Member in the 

Diamond Mountains belt, includes Stringocephalus 

beds at Lone Mountain but has yielded no identifiable 

Rugosa at this locality.

Great Basin Devonian coral zone D brackets unit 2 

of the Nevada Formation and in this area is further 

subdivided as subzones Dl5 D2 , and D3 . Of these subzones, Di, containing advanced Papiliophyllum, is 

viewed as late Early Devonian. D2 , with Breviphrentis,

Bethanyphyllum, Mesophyllum, Zonophyllum, Billingsastraea, and Hexagonaria, is considered to be late 

Early or early Middle Devonian, and D3 , with Arcophyllum, is assigned to the Middle Devonian (early 

Eifelian). Rugosa of coral zones A through D are 

treated in a separate paper on Lower and lower Middle 

Devonian rugose corals of the central Great Basin 

(Merriam, 1973b).

Unit 4 at Lone Mountain rests conformably on the 

flaggy and heavy-bedded gray limestones of unit 3, 

which includes thick lenses of rather fine grained 

crinoidal limestone. Corals and other shell fossils are 

fragmentary and poorly preserved in these partly bioclastic limestones. In the Diamond Mountains belt the 

rocks of this interval have been named the Sentinel 

Mountain Dolomite Member, are dolomitized, and 

have yielded no identifiable fossils, but otherwise have 

somewhat the gross lithologic appearance of unit 3.
10 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

Lithology and fossils of unit 4 of the Nevada Formation make possible a twofold division of the unit at 

Lone Mountain. Of the 675 feet of this unit (fig. 7), the 

upper 250 feet are distinguished by thicker bedding 

and more prominent massive weathering in places and 

are in considerable part dolomitic. Corals, especially 

the colonial Rugosa of coral zone F, are abundant in the 

dolomites of local growth sites and are generally silicified. The lower 425 feet of unit 4 shows less tendency to 

become dolomitized at Lone Mountain and comprises 

thinly bedded and flaggy limestones with occasional 

beds more than 1 foot thick. Much of this limestone is 

very fine textured and medium dark to light gray on 

fresh surface, weathering light gray to buff gray with 

pinkish streaks. There are some argillaceous partings, 

but these are less numerous than in the shaly members 

of unit 2. Silicified brachiopods, tentaculites, and other 

fossils are numerous and excellently preserved in many 

beds. Rugose corals are less common than in the upper 

250-foot division and are mainly the smaller solitary 

forms. Locally these lower beds are loaded with tentaculites of the Noivakia type. The "Martinia kirki zone" 

of Merriam (1940, p. 56) is essentially the lower 425 

feet of unit 4 at Lone Mountain. Rugose corals of coral 

zone F are most numerous in the upper 250-foot, partly 

dolomitic division of unit 4.

No identifiable rugose corals have been collected at 

Lone Mountain above unit 4 of the Nevada Formation. 

Except for an occasional thick Stringocephalus bed, the 

overlying dark- and light-gray saccharoidal dolomites 

of unit 5 are sparsely fossiliferous. These resistant dolomites form most of the main ridge of Lone Mountain 

and are approximately equivalent lithologically and 

faunally to the Bay State Dolomite Member in the 

Diamond Mountains belt. Above the dolomites of unit 

5, the thick-bedded limestones of the Devils Gate Limestone contain a great abundance of stromatoporoids of 

both the massive types and of Amphipora, apparently 

to the exclusion of most Rugosa.

Rugose corals of zone F collected during the course 

of this study from the upper 250-foot interval of unit 4 

of the Nevada Formation at Lone Mountain are as 

follows:

Cystiphylloides iddingsi n. sp.

Mesophyllum sp. f

Mesophyllum (Lekanophyllum) sp. 1

Keriophyllum mclareni n. sp. "

Keriophyllum? cf K.? kobehense (Stumm)

Acanthophyllum sp.

Paracanthus? sp. f

Cyatkophyllum (Moravophyllum) sp.

Cyathophyllum (Moravophyllum) sp. 1

Cyathophyllum (Orthocyathus) flexum (Stumm)

Sociophyllum eurekaensis n. sp.

Utaratuia nevadensis n. sp.

Taimyrophyllum nolani n. sp.

Among rugose corals collected by C. D. Walcott at 

Lone Mountain in 1880 and later described by Stumm 

(1937; 1938) are the following forms that probably 

came from unit 4 and coral zone F: 

Mesophyllum? sp. k 

Keriophyllum? kobehense (Stumm) 

Cyathophyllum (Orthocyathus) flexum (Stumm) 

Australophyllum prismatophylloides (Stumm) 

Hexagonaria fisherae (Merriam)

STRATA OF MIDDLE DEVONIAN CORAL ZONE F AND 

POSSIBLE ZONE G IN THE ROBERTS MOUNTAINS

East-dipping structural blocks of Nevada Formation 

with abundant Middle Devonian rugose corals are well 

exposed in the Roberts Mountains (fig. 4) at Devonian 

Peak, Bobcat Peak, and Pyramid Hill, where these beds 

were overridden by upper plate rocks of the Roberts 

Mountains thrust (Merriam and Anderson, 1942). 

Except for Pyramid Hill, no detailed stratigraphic 

work has been done here since the initial reconnaissance studies (Merriam, 1940; Merriam and Anderson, 

1942). Strata in the middle and upper parts of the 

Nevada Formation extending northward in the Roberts 

Mountains from Pyramid Hill to Devonian Peak are 

similar lithologically and faunally to those at Lone 

Mountain. Dolomitization is, however, less evident in 

the uppermost Nevada, within unit 5 the approximate 

equivalent of the Bay State Dolomite Member. Presence of Devonian coral zone G has not been established 

with certainty in the Roberts Mountains.

Less adequately investigated are coral-bearing 

Middle Devonian limestone outliers in overthrust terrane at Jackass Hill in the western Roberts Mountains, 

exposures at Dry Creek in the northern Roberts 

Mountains, and in Red Canyon west of Devonian Peak. 

These isolated exposures have yielded Hexagonaria. 

The Red Canyon coral assemblage may belong to coral 

zone G.

Devonian Peak. Within this large east-dipping fault 

block 2.5 miles west of the top of Roberts Creek Mountain, the beds near the summit of Devonian Peak contain Hexagonaria fisherae and the large brachiopod 

Warrenella occidentalis (Merriam). This horizon probably represents coral zone F. Near this horizon, but 

possibly somewhat higher stratigraphically, the limestones contain Heliolites and Hexagonaria sp. b (Merriam, 1940). The Bay State Dolomite Member (correlative with Nevada unit 5) and the overlying basal Devils 

Gate Limestone were not recognized in this block.

Bobcat Peak. The thick east-dipping Devonian section on the east slopes of this mountain passes upward 

through Nevada Formation unit 4 and includes unit 5 

as well as the lower part of the Devils Gate Limestone. 

Coral-rich beds of unit 4 and coral zone F are well
CORAL ZONES F AND G, ANTELOPE-ROBERTS MOUNTAINS FACIES BELT 11

exposed at locality M1342, where the following corals 

occur:

Siphonophrentis sp. b

Digonophyllum occidentalis n. sp.

Acanthophyllum robertsensis n. sp.

Cystiphylloides sp., cf. C. sp. b

Sociophyllum eurekaensis subsp. b

Heliolites sp.

Thamnopora sp. (coarse, thick form)

Higher in the Bobcat Peak section at locality M1365 

and below the massive upper beds, Hexagonaria cf. H. 

meeki n. sp. occurs in association with large Warrenella similar to W. occidentalis. Thamnopora is abundant near this occurrence.

Pyramid Hill. In this isolated hill northeast of the 

Roberts Creek Ranch house an east-dipping, nearly 

complete section of the Nevada Formation is especially 

well exposed. Some 300 feet of thick-bedded, massive 

Devils Gate Limestone that caps this hill rests conformably upon Nevada unit 5. As at Lone Mountain these 

lower limestones in the Devils Gate contain abundant 

Amphipora together with massive stromatoporoids. 

Coral zone F assemblages have been collected here from 

three localities (M1345, M1348, M1350) in Nevada 

unit 4. The rugose corals are as follows:

Acanthophyllum sp. p

Sociophyllum eurekaensis n. sp.

Taimyrophyllum nolani n. sp.

Hexagonaria fisherae (Merriam)

Jackass Hill. In this limestone outlier 2.5 miles 

northeast of Three Bar Ranch, the Nevada Limestone 

is surrounded by the overthrust Ordovician Vinini Formation. Coral zone F is represented here by carinate 

Hexagonaria similar to H. fisherae.

Dry Creek. In the northernmost Roberts Mountains, carinate Hexagonaria of the fisherae type representing coral zone F occurs 1.3 miles southeast of Willow Creek Ranch on the west side of Dry Creek.

Red Canyon. At locality M1339 in Red Canyon 1.9 

miles west of the summit of Devonian Peak, limestones 

of the Nevada Formation have yielded the following:

Paracanthus nevadensis n. gen., n. sp.

Acanthophyllum? sp.

Zonophyllum sp. r

Digonophyllum (Mochlophyllum) alhambraensis subsp. 

robertsensis n. subsp.

Hexagonaria sp. r

Thamnopora sp. (coarse, thick form)

stromatoporoids

The stratigraphic horizon of this assemblage may be 

that of Digonophyllum (Mochlophyllum) alhambraensis in the Diamond Mountains, that is coral zone G.

MIDDLE DEVONIAN ROCKS OF CORAL ZONE F 

IN THE NORTHERN ANTELOPE RANGE

Middle Devonian limestones of unit 4 and coral zone 

F, both in the Nevada Formation, are well exposed in 

the northernmost part of the Antelope Range (fig. 4)

which lies in the Cockalorum Wash and Horse Heaven 

Mountain quadrangles. Here between Antelope summit (7,829) and the mouth of Ninemile Canyon, Devonian rocks rest in thrust fault contact with Upper 

Cambrian and Ordovician beds. Thick-bedded limestones just east of the Ninemile Canyon fault (Merriam, 1963, pi. 2) contain Hexagonaria and probably 

represent unit 4 and coral zone F.

At locality M1347 on the west side of Antelope summit, low-dipping coralline limestones contain the following:

Favosites sp. (massive form)

Tabulophyllum antelopensis n. sp.

Lyrielasma antelopensis n. sp.

Hexagonaria fisherae subsp. antelopensis n. subsp.

Warrenella sp.

Atrypa sp.

Three-fourths of a mile north of Antelope summit at 

locality M30 on the west side of this ridge, platy limestones of unit 4 contain a brachiopod fauna, together 

with abundant Receptaculites, in what is possibly a 

lower stratigraphic horizon than locality M1347. 

Among the common fossils here are:

Receptaculites sp. (large platelike form)

Thamnopora sp. (coarse, thick form)

Heliolites sp.

acanthophyllid solitary coral

Sociophyllum? sp.

stromatoporoids (massive)

Schizophoria sp.

Leiorhynchus n. sp., cf. L. castanea (large form)

Atrypa sp. (large spiny form)

Warrenella kirki (Merriam)

Dechenella? (pygidium)

MIDDLE DEVONIAN STRATA OF CORAL ZONE F 

IN THE SOUTHERN FISH CREEK RANGE

A discontinuous outcrop belt of Middle Devonian 

strata containing zone F corals extends southward 6 

miles from Cockalorum Wash to the vicinity of Willow 

Creek at the south tip of the Fish Creek Range (figs. 

4, 6). This new unit, whose formal description as the 

Cockalorum Wash Formation is given later, is made up 

of limestones, siltstones, and sandstones. This coralbearing sequence lies in complexly faulted relation with 

rocks of other systems and reveals no normal stratigraphic boundary with older or younger Devonian beds. 

To elucidate the stratigraphy and structure of the new 

formation, certain parts of the Cockalorum Wash quadrangle have been mapped geologically in some detail 

(fig. 6). The type section passes through Reef Hill in 

the northern part of the type area. At Reef Hill large 

silicified Hexagonaria colonies are very numerous in a 

biohermal limestone near the top of the formation. At 

the south end of the Devonian outcrop in Coral Ridge 

near Willow Creek several comparable loci of abundant 

coral growth occur in these dislocated Cockalorum beds.
12 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

R.51 E. 116°10' R.52E.

Base 

1:62

from U.S. Geological Survey 

,500, Cockalorum Wash, 1956

Geology by C.W. Merriam, 1954-72

2 MILES

FIGURE 6. Southernmost part of the Fish Creek Range, Cockalorum Wash 

quadrangle, showing distribution and structural relations of Cockalorum Wash 

Formation, which contains faunas of Middle Devonian coral zone F.
CORAL ZONES F AND G, ANTELOPE-ROBERTS MOUNTAINS FACIES BELT

EXPLANATION

13

Include plant- and bone-bearing

tuffaceous deposits

Conglomerate, brown sandstone,

Include fresh-water molluscan fauna

Conglomerate, sandstone, siltstone,

and pebbly limestone

Include fusulinids and brachiopods

Brown quartzitic sandstone

Includes plant remains

Devils Gate Limestone and unnamed

Include Amphipora

Cockalorum Wash Formation

Siltstone, shale, sandstone, and

coral-bearing limestone

Quartzite and dolomite 

Possibly correlative with Oxyoke 

Canyon Sandstone Member of the 

Nevada Formation in Diamond 

Mountains. Occur in upper thfust 

plate

DC

o \-

Siltstone and shale with chert beds 

Age uncertain. Possibly correlative with Mississippian part of Pilot Shale. May include 

Cockalorum Wash Formation on Willow 

Creek. Occur in lower thrust plate

Hornfels and quartzite of unknown age 

Possibly Devonian. Occur in upper thrust plate

Contact 

Dashed where approximately located

Fault

Dashed where approximately located; 

dotted where concealed

M1367

Thrust fault

Dotted where concealed. Sawteeth 

on upper plate

Fossil locality Cockalorum Wash Formation 

with Hexagonaria

FIGURE 6. Continued.
14 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

COCKALORUM WASH FORMATION

The name Cockalorum Wash Formation (fig. 7) is 

here applied to some 1,200 feet of limestone, shale, siltstone, and sandstone; it crops out discontinuously in 

thrust fault slices and high-angle fault blocks in the 

southernmost Fish Creek Range between Cockalorum 

Wash and Willow Creek. The type area lies south of 

Nyeka Hill and west of Conglomerate Ridge at the 

center of the Cockalorum Wash quadrangle. Beds in the 

type section dip steeply west, passing upsection from 

the west base of Conglomerate Ridge westward through 

Reef Hill (fig. 6). Because no physical, stratigraphic, 

or paleontologic evidence of overturning exists, these 

strata are interpreted as becoming younger toward the 

west, the rich coral beds of Reef Hill therefore being 

uppermost.

Far less deformed conglomerate and sandstone underlying Conglomerate Ridge are in contact with the Devonian on the west; the relationship is apparently a 

combination of faulting and stratigraphic overlap. A 

fragmentary productid brachiopod in a conglomerate 

cobble suggests that these east-dipping beds are postearly Carboniferous and are probably post-Paleozoic. 

Six miles north at Fenstermaker Mountain, similar 

east-dipping conglomerates and coarse sandstones contain abundant poorly preserved plants and fossil wood. 

Three miles north on the Nye-Eureka County line are 

£7mo-bearing land-laid sandstones of Late Cretaceous 

or Paleocene age (W. A. Cobban and L. S. Russell, written communs., 1968). In general these less-deformed 

and less-indurated siliceous clastic rocks are distinguishable lithologically from denser plant-bearing 

coarse gritty brown sandstones of the Carboniferous, 

also present in fault blocks of this area.

North of the Cockalorum Wash Formation type section, dense white quartzites of uncertain age and derivation are thrust over the Middle Devonian Cockalorum Wash, probably from the west. Whereas these 

quartzites resemble the Ordovician Eureka Quartzite, 

it appears more probable they pertain to a sequence of 

white quartzite and dolomite which crops out 2 miles 

north-northeast near the county line in a thrust sheet 

of Devonian rocks correlated with the Oxyoke Canyon 

Sandstone Member of the Nevada Formation near 

Eureka.

The lower 600 feet of the Cockalorum Wash Formation comprises light-tan-weathering shales, platy siltstones, and sandstones, with lenses and interbeds of 

limonite-stained arenaceous pebbly to conglomeratic 

somewhat platy limestone and light-gray calcarenite. 

Brachiopod fragments are present in the lower clastic 

limestones. Abundant small calcitic plates in the calcarenites show a round median canal and have more

the appearance of dasycladacean algal debris than of 

Amphipora or crinoidal matter. Many of the carbonate 

clasts also appear to be of algal origin. Other spherical 

bodies are possible calcispheres. Minute poorly preserved spherical structures with a suggestion of radial 

spines may be Radiolaria.

Near the middle of the Cockalorum Wash type section, the light-gray limestone outcrops become thicker 

bedded in places and finer textured. Calcarenites in 

this part of the section lack the siliceous pebbles of the 

lower limestones. Tentaculites are present, together 

with small gastropods and other fragmentary shell fossils. Calcispheres and calcitic plates with a median 

canal are less conspicuous. Limestone lenses some 40 

feet thick extend laterally at least 300 feet. Separating 

the limestone beds are noncalcareous tan-weathering 

platy siltstones and pebbly siliceous sandstones. Gritty 

sandstone beds contain angular clasts of reworked rock. 

Limestones predominate within the upper 500 feet. One 

lenticular limestone body is about 175 feet thick and 

comprises platy to flaggy weathering medium- to lightgray fine-textured limestone. These beds contain fragmentary brachiopods, gastropods, and tentaculites. 

Calcispheres are common in the calcarenites. Tentaculites of the Nowakia type are abundant in some layers.

The important coral-bearing limestone underlying 

Reef Hill at the top of the type section is about 150 feet 

thick; this unit consists of flaggy to platy medium- and 

light-gray limestones, mostly fine textured, ranging 

from aphanitic to fine-grained calcarenite. The abundant corals are silicified. Where the coral heads are 

most numerous, at locality M1351, there is no evidence 

of inclined bedding to suggest moundlike organic 

growth with bottom relief, as in the case of a more 

typical bioherm.

On the west side of Cobble Hill, 0.5 mile south of 

Reef Hill, the interbedded fine-textured gray limestones, tan siltstones, and cherty shales are strongly 

deformed. No corals or other shell fossils were found 

here; poorly preserved probable Radiolaria are present 

in the aphanitic limestones. An exploration trench at 

locality M1366 exposes contorted tan and light-gray 

weathering siltstones containing eurypterid claws 

nearly identical with those from the Morey Peak area 

35 miles to the south, where they are associated with 

other well-preserved eurypterid body plates. The 

eurypterid claw beds and associated limestones of the 

Cockalorum Wash quadrangle are considered part of 

the Middle Devonian Cockalorum Wash Formation.

In Coral Ridge east of Willow Creek Ranch, the 

coral-bearing Cockalorum Wash Formation has been 

mapped along the ridge north and south of Willow 

Creek for 1.5 miles. Eurypterid claws and tentaculites 

occur here also in the siltstones and platy limestones.
DIAMOND MOUNTAINS BELT 15

The limestones and clastic beds dip steeply or are in 

places nearly vertical, and they appear on Willow Creek 

to have been thrust over dark shales and siltstones 

with chert interbeds. The shales and siltstones of the 

lower plate have yielded no fossils, but they may represent part of the Upper Devonian and Lower Mississippian Pilot Shale like that in a comparable structural 

setting 9 miles north at Bluff Canyon in the Fish Creek 

Range (fig. 4).

The Reef Hill fauna consists mainly of a single form 

Hexagonaria fisherae subsp. cockalorumensis, which 

occurs in great abundance, together with a large massive Favosites. Interbeds with rather sparse brachiopods yield Leiorhynchus sp. and a large species of 

Warrenella resembling W. occidentalis (Merriam).

The following is a combined faunal list including fossils from several localities of the Cockalorum Wash 

Formation in the northern part of Coral Ridge:

Favosites sp. (massive form in large cylindrical colonies)

Heliolites sp. (massive)

Thamnopora sp. (coarse variety)

chaetetids

Cyathophyllum (Moravophyllum) ? sp. (large solitary 

form)

Paracanthus nevadensis n. gen., n. sp.

Paracanthus cf. P. nevadensis n. gen., n. sp.

Hexagonaria fisherae subsp. antelopensis n. subsp.

Hexagonaria meeki n. sp.

Hexagonaria sp. c

Hexagonaria sp. f

massive stromatoporoids

Leiorhynchus (several types)

Atrypa sp. (fine-ribbed, no spines, fairly large)

Warrenella? sp.

Dechenella sp. (pygidium)

Ptomatis sp., or Knightites (Retispira) sp.

Porcellia sp., cf. P. nais (Hall)

Pleurotomaria-like genus (lacks slit band of

Pleurotomaria; surficially resembles P. capillaria)

MIDDLE DEVONIAN STRATA OF THE HOT CREEK RANGE

Carbonate facies of the Antelope-Roberts Mountains 

Devonian belt reappear 30 miles south of Cockalorum 

Wash in the Hot Creek Range. Near Warm Springs 

(fig. 4) at the south end of this range, faunas of Great 

Basin Devonian coral zone F are well represented. In 

the Morey Peak area of the middle Hot Creek Range, 

small coral collections suggest either coral zone F or G. 

Dark-gray to black carbonaceous limestones at locality M1354 just southwest of Warm Springs have yielded 

abundant silicified fossils, among which are the following:

Favosites sp. (massive form)

Thamnopora sp. (coarse, thick form)

chaetetids

Heliolites sp. (massive form)

Paracanthus sp., cf. P. nevadensis n. gen., n. sp.

Hexagonaria sp. w

Hexagonaria sp. n

Warrenella sp., cf. W. occidentalis (Merriam)

Atrypa sp. (large, no spines)

Leiorhynchus sp.

tentaculites

This assemblage pertains to coral zone F. Although the 

dark-gray and black limestones here differ considerably 

in appearance from unit 4 of the Nevada Formation at 

Lone Mountain, they are probably correlative. Nearly 

identical black limestones occur in the same interval at 

Bare Mountain, 90 miles south of Warm Springs.

Gray limestones north of Morey Peak at locality 

M1338 contain Mesophyllum (Lekanophyllum) cf. M. 

(L.) sp. 1 and Cystiphylloides sp. c. This and other 

types of Mesophyllum from limestones and dolomites 

of the Morey Peak area probably represent coral zone 

F. Hexagonaria? sp. h occurs at locality M1358, about 

2 miles north of the Morey mine. In this vicinity the 

Devonian rocks contain eurypterid claws and body 

plates. Detailed mapping and stratigraphy may demonstrate that some of these beds are a southward continuation of the Cockalorum Wash Formation.

MIDDLE DEVONIAN RUGOSE CORALS 

OF THE DIAMOND MOUNTAINS BELT

Geologic mapping of the Pinto Summit quadrangle by 

T. B. Nolan (Nolan and others, 1973) and stratigraphic 

investigation under the Kobeh Valley project demonstrate that the depositional facies in the lower part of the 

Devonian in the Diamond Mountains belt (fig. 4) differ 

widely from those of the Antelope-Roberts Mountains 

belt on the west (Nolan and others, 1956). Nearly 

unfossiliferous partly sandy dolomites and orthoquartzite having a combined thickness of about 850 feet constitute the Beacon Peak Dolomite Member and Oxyoke 

Canyon Sandstone Member in the Oxyoke Canyon 

area (fig. 7). These eastern facies seemingly occupy 

the approximate time-rock interval of units 1 and 2 of 

the Nevada Formation in the Lone Mountain section 

(figs. 2, 7), where they are rich in fossils of Early and 

early Middle Devonian ages (Merriam, 1973b). West 

to east lithofacies changes are less profound within the 

overlying intervals of units 4 and 5. There are, however, 

significant faunal differences, as the more diagnostic 

rugose corals of unit 4 and coral zone F at Lone Mountain have not been recognized in the presumably 

equivalent Woodpecker Limestone Member of the Diamond Mountains belt. Moreover, the characteristic 

Rugosa of coral zone G that occupy basal beds of the 

Bay State Dolomite Member and topmost beds of the 

Woodpecker Limestone Member in the Diamond 

Mountains belt do not appear where they would be 

expected near the boundary of units 4 and 5 of the 

Nevada Formation in the Antelope-Roberts Mountains 

belt.
DEVONIAN SILURIAN

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GEOLOGIC CORRELATIONS WITH DISTANT REGIONS 17

GREAT BASIN DEVONIAN CORAL ZONE G

The Great Basin Devonian coral zone G (fig. 7) 

includes the lowest strata with Stringocephalus and 

occupies a 30-foot silty dolomite and limestone interval 

bracketing the top of the Woodpecker Limestone Member and the base of the Bay State Dolomite Member 

of the Nevada Formation (Nolan and others, 1956, p. 

47). The following is a combined list of fossils in Great 

Basin Devonian coral zone G:

Thamnopora sp. (coarse, thick form)

Heliolites sp. (massive form)

Syringopora sp.

chaetetids

Cystiphylloides sp. a

Digonophyllum (Mochlophyllum) alhambraensis n. sp.

Mesophyllum n. sp.

Acanthophyllum sp. c

Cyathophyllum (Moravophyllum) alhambraensis n. sp.

Disphyllum sp.

Stringocephalus sp. a

Stringocephalus sp. b

Rensselandia n. sp.

Amphiporal sp.

stromatoporoids (massive)

tentaculites

Notably lacking are the colonial Rugosa of coral zone 

F, among which are Hexagonaria, Taimyrophyllum, 

Sociophyllum, and Utaratuia.

Strata of Great Basin coral zone G are well exposed 

in the Pinto Summit quadrangle on the southwest side 

of Sentinel Mountain, in the upper Oxyoke Canyon, 

and at the south end of the Alhambra Hills on the east 

side. The lowermost Bay State in Oxyoke Canyon is a 

light-gray silty dolomite containing abundant large 

Stringocephalus sp. b with which are associated Thamnopora sp., Acanthophyllum sp. c, Mesophyllum n. sp., 

Cyathophyllum (Moravophyllum) sp., Disphyllum sp., 

and stromatoporoids. Solitary acanthophyllids are the 

only Rugosa found in the underlying Woodpecker 

Limestone Member.

The lowermost bed in the Bay State Dolomite Member of the Alhambra Hills is a medium-gray silty dolomite bed probably correlative with a light-gray silty 

dolomite exposed in Oxyoke Canyon; beneath this bed 

is dark-gray limestone containing the very large cylindrical Digonophyllum (Mochlophyllum) alhambraensis, Cyathophyllum (Moravophyllum) alhambraensis, 

Cystiphylloides sp. a, Heliolites sp., Syringopora sp., 

Stringocephalus sp. a, and Rensselandia n. sp. Mapped 

with the uppermost Woodpecker Limestone Member, 

these limestones are of special importance because they 

locally contain abundant diagnostic Rensselandia, 

known elsewhere at the bottom of the Stringocephalus 

zone. Here, as in Oxyoke Canyon, only the solitary 

acanthophyllids have been found in the Woodpecker 

beneath the 30-foot interval of coral zone G. .

MIDDLE DEVONIAN STRATA OF

BARE MOUNTAIN, SOUTHERN NYE COUNTY,

NEVADA

The southernmost known exposures carrying faunas 

of coral zone F in the Antelope-Roberts Mountains belt 

are at Bare Mountain near Beatty, Nye County, Nev., 

90 miles south of Warm Springs (Cornwall and Kleinhampl, 1960). Here in Tarantula Canyon (locality 

M1359) black, carbonaceous limestones containing 

these fossils resemble the limestones with a correlative 

assemblage at Warm Springs. The fossils are as follows:

Favosites sp. (massive form)

Thamnopora sp.

Alveolites sp.

Hexagonaria fisherae subsp. antelopensis

Atrypa sp.

Crurithyris sp. (large form)

Cyrtina sp.

Schizophoria sp. (large form)

stromatoporoids (massive)

GREAT BASIN MIDDLE DEVONIAN RUGOSA 

AND THEIR BEARING UPON GEOLOGIC 

CORRELATION WITH DISTANT REGIONS

Great Basin Middle Devonian Rugosa are especially 

valuable for correlation of their enclosing rocks with 

Devonian rocks in western Canada, southeastern 

Alaska, and the more remote Eifel district of Germany. 

As might be anticipated paleogeographically, fairly 

close paleontologic ties exist to the north with coralbearing strata of the central Mackenzie River region 

west of Great Bear Lake and northwest of Great Slave 

Lake. Less evident are genetic affinities to rugose corals 

of the continental interior and eastern North America. 

In fact, these studies demonstrate that the overall 

generic makeup of eastern Middle Devonian rugose 

coral assemblages differs appreciably from that of the 

western or Cordilleran faunas. Vagaries of Devonian 

faunal spread and provincial biogeography may be 

invoked to explain closer similarity of some Cordilleran 

Rugosa to species of the more distant Rhine Valley 

and Eifel district.

The Stringocephalus zone. Intercontinental and 

provincial correlation of Great Basin Middle Devonian 

rugose coral-bearing strata gains support through supplementary evidence from study of two distinctive 

brachiopod groups: the widely distributed Stringocephalus and the more localized Leiorhynchus castanea 

(Meek) faunas of the Cordilleran belt.

Since recognition by Kirk (1927) of the importance 

of Stringocephalus as a Great Basin zone fossil, this 

large terebratuloid has been found to be largely confined to the Bay State Dolomite Member of the Nevada 

Formation in the Eureka district and the approximately correlative unit 5 of nearby areas. At Lone
18 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

Mountain this genus ranges upward into lowermost 

beds of the overlying Devils Gate Limestone. Stringocephalus seems to be similarly restricted vertically elsewhere in the Great Basin, in Alaska, and in western 

Canada (McLaren, 1962). As in western Canada, the 

other large and distinctive terebratuloid, Rensselandia, 

occurs also in the Eureka district at the bottom of the 

Stringocephalus zone in coral zone G. In connection 

with proposed correlation with the European Devonian, 

the Great Basin Stringocephalus zone is provisionally 

regarded as essentially equivalent in time-stratigraphic 

sense to the Rhine Valley Givetian. It is, however, recognized that some authorities have suggested that 

Stringocephalus may be confined to the middle and 

upper parts of the standard Givetian. The lowest beds 

of coral zone G are perhaps the most important paleontologic datum in the Great Basin Devonian column.

Leiorhynchus castanea. The key fossil Leiorhynchus castanea (Meek) is a common brachiopod in the 

Woodpecker Limestone Member of the Nevada Formation of the Diamond Mountains belt, the member 

which underlies the Bay State Dolomite Member and 

the Stringocephalus zone. Warrenella kirki is sparsely 

represented here with abundant L. castanea. Leiorhynchus castanea has not been found even though expected, 

in unit 4 of the Nevada Formation west of the Diamond 

Mountains belt at Lone Mountain, where unit 4 

occupies the Woodpecker interval below Stringocephalus and contains the abundant Rugosa of coral zone F. 

Warrenella kirki is a common brachiopod here, unaccompanied by L. castanea. Facies may explain these 

east-to-west faunal changes, for the bulk of the zone F 

coral assemblage and especially the colonial genera 

have not been found in the easterly Woodpecker. As 

shown by McLaren (1962, fig. 1), the vertical range of 

Leiorhynchus castanea is greater in western Canada, 

this species overlapping the range of Stringocephalus 

and probably passing upward into younger beds.

DISTRIBUTION AND CORRELATION VALUE OF RUGOSA

Internally complex, specialized solitary corals of the 

families Digonophyllidae and Ptenophyllidae differentiated rapidly in both Europe and western North 

America, reaching evolutionary peaks during the intervals of Great Basin Middle Devonian coral zones F and 

G. Although of wide value for geologic correlation, these 

families are little known if indeed represented in eastern North America. Cystiphylloides, a diverse cosmopolitan genus well represented both east and west, has 

by some workers (Stumm, 1961) been classified with 

Digonophyllidae; however, Cystiphylloides lacks many 

true digonophyllid features and is here, by preference, 

reassigned to the Family Cystiphylloidae.

The distinctive colonial genera Utaratuia, Taimyrophyllum, and Sociophyllum remain undocumented thus 

far in eastern North America but are otherwise significant in this connection. Sociophyllum and Taimyrophyllum are typically Old World genera.

Species of the generalized, long-ranging Disphyllum 

are common both east and west in North America but 

at present do not have great correlation significance. 

Somewhat more useful is Tabulophyllum, typically 

Upper Devonian but present in the Cordilleran Middle 

Devonian and in that of eastern North America.

The ubiquitous, worldwide Hexagonaria, a structurally and taxonomically very diverse genus with multitudes of local subgenera and species, has at present 

rather limited use for definitive geologic correlation.

The complex Family Phillipsastraeidae, as here construed, has no recognized ancestors in the Great Basin 

Middle Devonian. Earlier Old World representatives of 

this group are reported, especially in Australia. 

Distinctive Phillipsastraeidae are the predominating 

Rugosa of Late Devonian coral zone I in the Great 

Basin, presumably having been introduced in Frasnian 

time from other faunal provinces.

CORRELATION WITH RUGOSE CORAL DEPOSITS 

OF WESTERN CANADA

Digonophyllidae similar to species of Great Basin 

Devonian coral zones F and G are known from the 

Hume Formation of the central Mackenzie River, from 

correlative beds on the arctic Anderson River, and from 

the Horn Plateau Formation northwest of Great Slave 

Lake. Acanthophyllum of the contemporaneously 

evolving Ptenophyllidae is recorded from Hart River, 

Yukon Territory, and probably occurs also in the Hume 

Formation. The distinctive colonial genera Sociophyllum, Utaratuia, and Taimyrophyllum characterize 

the Hume as well as the beds of Great Basin Devonian 

coral zone F. Sociophyllum occurs also on Hart River, 

Yukon Territory.

The Nahanni Formation west of Fort Simpson and 

Great Slave Lake also yields Sociophyllum, Utaratuia, 

and Taimyrophyllum related to the Hume species, and 

the two formations are doubtless in considerable part 

correlative (Pedder, 1964). Also present in the Hume is 

Australophyllum resembling A. prismatophylloides of 

Great Basin coral zone F. Of the few described Hexagonarias from northwest Canada, none appear closely 

related to carinate Hexagonaria of Great Basin coral 

zone F.

HUME FORMATION

The stratigraphy of the widely distributed coralbearing Hume Formation in the central Mackenzie 

River area has been elucidated by Bassett (1961), and 

its coral faunas studied by Lenz (1961) and more
GEOLOGIC CORRELATIONS WITH DISTANT REGIONS 19

recently by Pedder (1964). Bassett's (1961, fig. 3) 

comparative stratigraphic columns between Fort Good 

Hope and Norman Wells show the Hume overlain conformably by the Hare Indian Formation and underlain 

disconformably by the Bear Rock Formation. Resting 

conformably upon the Hare Indian is the Kee Scarp 

Limestone which includes the Stringocephalus zone 

and is thus correlative with Stringocephalus-bearing 

Bay State Dolomite Member and coral zone G of the 

Great Basin. At the bottom of the Devonian column, 

the dolomitic beds of Bear Rock are sparsely fossiliferous and have not been dated with certainty. Early 

Middle Devonian is doubtlessly present, but there is 

evidently no paleontologic basis for correlation with 

Lower Devonian horizons in the Nevada Formation of 

the Great Basin.

Leiorhynchus castanea is recorded by Bassett above 

the Hume in lower beds of the Hare Indian Formation. 

According to McLaren (1962, fig. 1), this key brachiopod ranges upward in areas of Hume exposure well into 

the Kee Scarp Limestone, thus overlapping the range 

of Stringocephalus.

As described and illustrated by Pedder (1964), the 

Hume corals Sociophyllum glomerulatum, Utaratuia 

laevigata, and Taimyrophyllum triadorum appear to 

have counterparts in the Great Basin coral zone F 

species Sociophyllum eurekaensis, Utaratuia eurekaensis, and Taimyrophyllum nolani.

The Hume Formation contains species of Mesophyllum illustrated by McLaren, Norris, and McGregor 

(1962) and by Lenz (1961); these appear analogous to 

Mesophyllum (Lekanophyllum) sp. 1 and Mesophyllum 

sp. f of Great Basin coral zone F. Billingsastraea verrilli (Meek) figured by Pedder (1964) from the Hume 

and Nahanni has no matching form in Great Basin 

coral zone F. The most closely related Great Basin 

corals, Billingsastraea nevadensis (Stumm) and its 

subspecies arachne (Stumm), are not conspecific with 

the northern verrilli. These Great Basin corals occur 

together in unit 2 (coral zone D2 ) considerably below 

coral zone F. Dendrostella trigemme of the Hume and 

Nahanni Formations (Pedder, 1964, p. 434) also has 

no counterpart in coral zone F. In the Great Basin, 

Dendrostella has thus far been found only in unit 2 and 

coral zone D2 .

The presence of the Stringocephalus zone within the 

Kee Scarp Limestone and the similarity of the Great 

Basin coral zone F rugose coral fauna to those of the 

Hume and Nahanni indicate a probable fairly close 

correlation and suggest a late Eif elian rather than early 

Givetian age.

HORN PLATEAU FORMATION 

Outcrops of the Middle Devonian Horn Plateau Formation 90 miles northwest of Slave Point, Great Slave 

Lake (Norris, 1965, p. 78-83, fig. 9), yield rugose corals 

with affinity to Great Basin species. Of the 12 rugose 

coral genera present (McLaren and Norris, 1964), five 

are represented in the Great Basin by different but 

comparable species. These are Sociophyllum redactum 

(McLaren), Mesophyllum (Lekanophyllum) cf. punctatum (Wedekind), M. (Lekanophyllum) nebracis 

(McLaren), Cystiphylloides spinosum McLaren, and 

Disphyllum salicis McLaren. Stringocephalus, Leiorhynchus castanea, and Warrenella are not reported. 

The Horn Plateau Sociophyllum and the digonophyllid 

species may be compared with Sociophyllum eurekaensis and with Mesophyllum (Lekanophyllum) sp. 1 

of Great Basin coral zone F. Lacking in the Horn 

Plateau fauna are the compound Rugosa Utaratuia and 

Taimyrophyllum, which characterize the Hume, Nahanni, and Great Basin coral zone F.

McLaren (in McLaren and Norris, 1964, p. 5) concluded that the Horn Plateau fauna suggests a middle 

to late but not latest Givetian age. The Horn Plateau 

fauna shows no very close resemblance to the Great 

Basin coral zone G of Givetian age at the bottom of the 

Stringocephalus zone.

RUGOSE CORALS AND CORRELATION

WITH MIDDLE DEVONIAN STRATA OF

SOUTHEASTERN ALASKA

Middle Devonian stratigraphy and faunas of southeastern Alaska are best known at San Alberto Bay near 

Craig on the west side of Prince of Wales Island (Eberlein and Churkin, 1970). Here the presence of Stringocephalus was recorded by Kirk (1927, p. 219), and the 

beds have yielded rugose corals of the families Digonophyllidae and Ptenophyllidae, as well as the colonial 

Hexagonaria and Australophyllum. Within the Alberto 

Islands, Stringocephalus occurs fairly high stratigraphically in the east-dipping Devonian section, which 

passes up section to the east through these islands and 

across southern Wadleigh Island, where it continues 

above into the Upper Devonian. Below Stringocephalus, 

these rocks contain Digonophyllidae of the three genera 

Digonophyllum, Mesophyllum, and Arcophyllum, and 

Acanthophyllum of the Ptenophyllidae. Also present 

are colonial Rugosa of the genera Australophyllum, 

Xystriphyllum, and Loyolophyllum.

The abundant Rugosa in certain parts of the San 

Alberto Bay vicinity are large maturely subcylindrical 

members of the Paracanthus group. Those from Fern 

Point, San Fernando Island, 4 miles southwest of the 

Alberto Islands, resemble Paracanthus nevadensis n. 

sp. of Great Basin coral zone F and beds possibly pertaining to coral zone G.

Whereas none of these southeastern Alaska corals 

appears conspecific or very closely related to Hume or
20 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

Great Basin species, it is probable they occupy the 

time-stratigraphic interval of Great Basin coral zones 

F and G. Australophyllum from the Alberto Islands on 

the west occurs low in this section and resembles Australophyllum prismatophylloides of Great Basin coral 

zone F. From the same locality and horizon comes a 

large complex Arcophyllum quite similar to A. dachsbergi (Vollbrecht) of middle Middle Devonian age in 

the Eifel district of Germany. It is probable these 

Alberto Islands Middle Devonian beds range upward 

from the Eifelian Stage into the Stringocephalus-beaiing Givetian. Sociophyllum occurs high in this column, 

presumably below the top of the Middle Devonian.

Many scattered island exposures of gently dipping 

Middle Devonian coral-bearing strata occur at Karheen 

Passage, 20 miles north of the Alberto Islands. The 

Stringocephalus datum has not been found in this 

vicinity; the stratigraphic sequence is somewhat 

obscure and can not at present be correlated in detail 

with that of San Alberto Bay. Among the rugose corals 

are Digonophyllum, Acanthophyllum, Hexagonaria, 

and Dendrostella. The Karheen Hexagonaria is noncarinate, unlike those most characteristic of Great 

Basin coral zone F. Dendrostella cf. D. rhenana 

resembles the Rhine Valley Givetian (Glinski, 1957) 

typical rhenana and is similar also to Dendrostella 

trigemme from the Hume and Nahanni Formations of 

western Canada (Pedder, 1964).

Of more diagnostic colonial Rugosa in the Hume, 

Nahanni, and Great Basin coral zone F, the genus 

Utaratuia has not been found in southeastern Alaska, 

and Sociophyllum is known only from scrappy material 

at a single locality. A thamnastraeoid to aphroid 

colonial genus collected by Eberlein and Churkin 

(1970) at Sunny Hay Mountain, 4.5 miles east of Craig, 

reveals features of both Billingsastraea and Taimyrophyllum. This interesting coral does not appear to be 

related either to the Hume and Nahanni Billingsastraea 

verrilli or to Taimyrophyllum in Great Basin coral zone 

F, the Hume, or the Nahanni.

In summary, the southeastern Alaska Middle Devonian rugose coral-bearing rocks of Eifelian and Givetian 

age have yielded stratigraphically significant Digonophyllidae as well as Acanthophyllum of the Ptenophyllidae. However, the more diagnostic colonial genera of 

Great Basin coral zone F and the Hume and the 

Nahanni of western Canada remain unrecognized or 

poorly represented. Stringocephalus is known locally, 

but the key brachiopod, Leiorhynchus castanea, has 

not been identified in these beds of the Pacific Border 

province, where rocks and faunas generally of the 

Silurian and Devonian Systems differ somewhat from 

those of the main Cordilleran seaways. -

CORRELATION WITH RUGOSE CORAL DEPOSITS 

OF THE RHINE VALLEY, GERMANY

Paleontologic comparison of Great Basin Middle 

Devonian rugose corals with those of the Rhine Valley 

is especially meaningful in terms of intercontinental 

geologic correlation. Coral research by Schliiter, Wedekind, Vollbrecht, Birenheide, Glinski, and others since 

the 1880's makes possible fairly detailed comparisons 

of internal structure with certain specialized Great 

Basin solitary genera. In the Eifel district especially, 

the stratigraphic occurrence of Middle Devonian 

Rugosa has been intensively investigated by stratigraphers and coral specialists. The Belgian Couvinian 

Rugosa, although recently elucidated by Tsien (1969), 

are less understood by modern standards than are those 

of the more easterly Eifelian.

Among Rhine Valley Middle Devonian Rugosa, the 

closest affinity to Great Basin forms is in the larger 

specialized Digonophyllidae and Ptenophyllidae of the 

Eifel district. Both families appear to have peaked in 

the standard Eifelian, within the stratigraphic interval 

Nohner Schichten to Junkerberg Schichten. Although 

continuing on into the Rhine Valley Givetian, the 

Ptenophyllidae are seemingly less important than the 

digonophyllids towards the end of Middle Devonian 

time.

Among Digonophyllidae in the Great Basin coral 

zone F, Mesophyllum (Lekanophyllum) sp. 1, although 

not conspecific with either, may be compared with 

Mesophyllum (Lekanophyllum) punctatum (Wedekind) and lissingenense (Schliiter) of the Gerolstein 

area. Digonophyllum (Mochlophyllum) alhambraensis 

of Great Basin coral zone G resembles maximum 

(Schliiter) of the Eifelian Junkerberg Schichten. Of 

the Ptenophyllidae, species of Acanthophyllum in coral 

zone F reveal similarities to Acanthophyllum heterophyllum (Birenheide, 1961) of the Eifelian Junkerberg 

Schichten.

Keriophyllum, represented by the solitary K. mclareni 

in Great Basin coral zone F, has a counterpart in Keriophyllum heiligensteini Wedekind, possibly of the Eifelian Junkerberg (Birenheide, 1962a, p. 108). According 

to Birenheide (1962a, p. 108; 1963, p. 390) the type 

species of this complex genus is a synonym of Cyathophyllum (Peripaedium) turbinatum Goldfuss, also of 

the Eifelian Junkerberg Schichten.

Colonial Rugosa of the Rhine Valley have less in 

common with those of western North America, although 

Eifel district Sociophyllum and carinate Hexagonaria 

are worthy of comparison with Great Basin species. 

Sociophyllum eurekaensis n. sp. of coral zone F is comparable with, but not conspecific with, S. elongatum
GEOLOGIC CORRELATIONS WITH DISTANT REGIONS 21

(Schliiter) of the Eifel district Givetian in the range 

from Ahbach Schichten to Ciirten Schichten. According to Birenheide (1962b, p. 53), Sociophyllum is 

present also in the upper Eifelian of Europe, which is 

the favored age of Great Basin coral zone F.

Great Basin carinate Hexagonaria of coral zone F is 

comparable with, but not conspecific with, carinate 

members of this genus in the Middle Devonian of the 

Eifel district. Among Eifel species is the Paffrath 

Hexagonaria hexagona (Goldfuss), representing the 

type species as interpreted by Stumm (1948a, p. 14, 

pi. VI, figs. 1, 2). Stratigraphic range of this form is 

unknown.

In summary, Great Basin Devonian coral zone G at 

the bottom of the Stringocephalus zone must accordingly be interpreted as Givetian in terms of the Rhine 

Valley standard. Coral zone F lies below the Great 

Basin Stringocephalus beds. With its digonophyllids of 

Eifelian stamp, a possible Eifelian Keriophyllum (or 

Peripaedium), archaic Siphonophrentis, and carinate 

Hexagonaria, Great Basin coral zone F is believed to 

harmonize better with later Eifelian than with early 

Givetian of the Eifel district. Coral zone F may be 

approximately correlative with the German Junkerberg 

Schichten.

MIDDLE DEVONIAN RUGOSE CORALS OF EASTERN

NORTH AMERICA COMPARED WITH THOSE

OF THE GREAT BASIN

Great Basin Middle Devonian rugose coral faunas of 

zones F and G show few convincing resemblances to 

coral biotas of inferred equivalent age in eastern North 

America and the continental interior. It appears to be 

true, however, that older Middle Devonian beds in the 

interval of Great Basin coral subzones D2 and D3 contain Rugosa with somewhat closer gross resemblances 

to eastern species. Such, for example, are the abundant 

Siphonophrentis-like corals and the Bethanyphyllidae 

of Do that exhibit a degree of convergent morphologic 

similarity and possible evolutionary parallelism suggesting age equivalence. Yet among these analogs, none 

are conspecific or very closely related genetically. As 

noted under the discussion "Great Basin Devonian 

Coral Zones," the Great Basin D2 corals constitute both 

Early and early Middle Devonian elements, of which 

the Digonophyllidae appearing first in this subzone 

have their genetic ties, not with eastern North America, 

but with the Eifelian of Europe. Subzone D3 of the 

Great Basin is by itself considered Eifelian without 

reservation.

A considerable number of Great Basin Middle Devonian rugose coral groups are thus far unrecorded in

eastern North America. Among these are the European 

specialized Digonophyllidae and Ptenophyllidae, as 

well as the Old World colonial Sociophyllum and 

Taimyrophyllum and the thus-far exclusively western 

North America Utaratuia of coral zone F.

Rugose coral genera common to Great Basin coral 

zone F and the Middle Devonian of the East are Tabulophyllum, Cystiphylloides, Disphyllum, and Hexagonaria. On the other hand, several strongly yardarm 

carinate colonial genera of the eastern Middle Devonian, such as Cylindrophyllum, Eridophyllum, and certain species doubtfully placed inBillingsastraea (Ehlers 

and Stumm, 1951) remain unidentified in the Great 

Basin. To these may be added the profusely yardarm 

carinate, usually solitary Heliophyllum so prominent in 

the eastern coralline Middle Devonian. Less-carinate 

species in the Horn Plateau Formation of western 

Canada have been assigned to Heliophyllum by 

McLaren in McLaren and Norris, (1964).

Tabulophyllum, more often a Late Devonian genus, 

appears first in Great Basin coral zone F. In the Michigan column, Tabulophyllum occurs in the Middle Devonian Bell Shale and Alpena Limestone (Stumm, 1962) 

that fall approximately within the interval of Great 

Basin coral zones F and G. The long-ranging and generalized Disphyllum has at present little correlation 

value.

Cystiphylloides, abundant in the eastern Middle 

Devonian, and by some workers classified with the 

Digonophyllidae (Stumm, 1961), has numerous analogs in the western Devonian. Unidentified in the west 

are the specialized cystimorphs that are allied to Cystiphylloides but differ in the presence of heavy stereoplasmic deposits; among these are Cladionophyllum 

Stumm and Edaphophyllum Simpson.

Great Basin carinate Hexagonaria bears among 

western Rugosa the closest structural resemblance to 

eastern Middle Devonian species. In general the 

strongly yardarm carinate Hexagonarias appear to outnumber those with smooth septa in the Lower Devonian, and especially in the lower Middle Devonian of 

the east. Although carinate Hexagonaria persists here 

and there into younger Middle Devonian of the east, 

those with smoother septa become more evident upward. In the Late Devonian (Frasnian) of Europe, 

Hexagonarias with smooth septa appear to predominate 

(Sorauf, 1967).

Carinate Hexagonaria resembling that of Great Basin 

coral zone F is abundantly represented in the central 

lowland Devonian west of the Appalachians from the 

Middle Devonian Jeffersonville Limestone of Kentucky 

to the Middle Devonian beds of Michigan (Stumm, 

1948a;1964;1970).
22 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

CLASSIFICATION OF GREAT BASIN

MIDDLE DEVONIAN RUGOSE CORALS

IN CORAL ZONES F AND G

Described rugose corals of the Great Basin Lower 

and Middle Devonian (coral zones A through G) are 

classified in 14 families. Those of the Lower and lower 

Middle Devonian, or coral zones A through D of Merriam (1973b), constitute 11 families, the three families 

unrepresented (Ptenophyllidae, Cyathophyllidae, and 

Spongophyllidae) being first recognized in coral zones 

F and G. The eight families of Rugosa that characterize 

zones F and G do not include Laccophyllidae, Kodonophyllidae, Stauriidae, Halliidae, Bethanyphyllidae, and 

Chonophyllidae present in the lower zones A through 

D. Especially important stratigraphically are the abundant Halliidae and Bethanyphyllidae of zones B 

through D; above these zones they are unknown in this 

western province. Digonophyllidae, appearing in the 

higher beds of zone D, assume increasing importance 

upward stratigraphically to peak in zones F and G, 

together with the Ptenophyllidae. Disphyllidae, well 

established in early Middle Devonian beds of coral zone 

D, continue to diversify upward as the genera Hexagonaria and Taimyrophyllum of coral zone F in which 

they peak.

The classification adopted (Merriam, 1973b) is for 

the greater part in accord with that of Hill (1956), the 

most authoritative thus far published. Departures from 

Hill's arrangement include proposal of a new subfamily, 

Siphonophrentinae, under the Streptelasmatidae, restoration of Stumm's Family Cystiphylloidae, use of 

the Family Endophyllidae in a more inclusive sense, 

and restoration of the Family Cyathophyllidae of Dana. 

For the very complex Digonophyllidae, Hill's scheme 

(1956, p. F314-F320) would appear to be the most 

satisfactory proposed thus far, except for inclusion of 

Cystiphylloides as a subgenus of Mesophyllum. 

Order Rugosa Edwards and Haime 1850

Family Streptelasmatidae Nicholson 1889 (in Nicholson and

Lydekker 1889 as Streptelasmidae) 

Subfamily Siphonophrentinae, new subfamily 

Genus Siphonophrentis O'Connell 1914 

Subgenus Siphonophrentis O'Connell 1914 

Siphonophrentis (Siphonophrentis) sp. b 

Family Cystiphylloidae Stumm 1949

Genus Cystiphylloides Chapman 1893 

Cystiphylloides sp. a 

Cystiphylloides iddingsi n. sp. 

Cystiphylloides sp. c 

Family Digonophyllidae Wedekind 1924 

Subfamily Zonophyllinae Wedekind 1924 

Genus Zonophyllum Wedekind 1924

Zonophyllum sp. r

Subfamily Digonophyllinae Wedekind 1924 

Genus Digonophyllum Wedekind 1923 

Subgenus Digonophyllum Wedekind 1923

Digonophyllum (Digonophyllum) occidentalis n. sp. 

Digonophyllum (Digonophyllum)? sp. c 

Subgenus Mochlophyllum Wedekind 1923 (as a genus) 

Digonophyllum (Mochlophyllum) alhambraensis

n. sp. 

Digonophyllum (Mochlophyllum) alhambraensis

subsp. robertsensis n. subsp. 

Genus Mesophyllum Schliiter 1889 

Subgenus Mesophyllum Schliiter 1889 

Mesophyllum (Mesophyllum) sp. f 

Mesophyllum (Mesophyllum)? sp. k 

Subgenus Lekanophyllum Wedekind 1924 (as a genus)

Mesophyllum (Lekanophyllum) sp. 1 

Family Ptenophyllidae Wedekind 1923

Genus Acanthophyllum Dybowski 1873 

Acanthophyllum sp. a 

Acanthophyllum robertsensis n. sp. 

Acanthophyllum sp. c 

Acanthophyllum sp. p 

Acanthophyllum? sp. 

Genus Paracanthus new genus

Paracanthus neuadensis n. gen., n. sp. 

Paracanthus? sp. f 

Family Cyathophyllidae Dana 1846

Genus Cyathophyllum Goldfuss 1826

Subgenus Morauophyllum Kettnerova 1932 (as a

genus) 

Cyathophyllum (Moravophyllum) alhambraensis

n. sp.

Cyathophyllum (Morauophyllum) sp. 1 

Subgenus Orthocyathus n. subgen.

Cyathophyllum (Orthocyathus) flexum (Stumm) 

Genus Keriophyllum Wedekind 1923 

Keriophyllum mclareni n. sp. 

Keriophyllum? kobehense (Stumm) 

Family Spongophyllidae Dybowski 1873

Genus Sociophyllum Birenheide 1926b (as subgenus of

S t ringophyllum) 

Sociophyllum eurekaensis n. sp. 

Sociophyllum eurekaensis subsp. b 

Genus Utaratuia Crickmay 1960

Utaratuia eurekaensis n. sp. 

Family Endophyllidae Torley 1933

Genus Tabulophyllum Fenton and Fenton 1924

Tabulophyllum antelopensis n. sp. 

Genus Australophyllum Stumm 1949

Australophyllum prismatophylloides (Stumm) 

Family Disphyllidae Hill 1939

Genus Hexagonaria Giirich 1896 

Hexagonaria fisherae (Merriam)

Hexagonaria fisherae subsp. cockalorumensis n. subsp. 

Hexagonaria fisherae subsp. antelopensis n. subsp. 

Hexagonaria meeki n. sp. 

Hexagonaria sp. c 

Hexagonaria sp. f 

Hexagonaria? sp. h 

Hexagonaria sp. n 

Hexagonaria sp. r 

Hexagonaria sp. w 

Genus Taimyrophyllum Chernychev 1941

Taimyrophyllum nolani n. sp. 

No family assignment

Genus Lyrielasma Hill 1939 

Lyrielasma antelopensis n. sp.
MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN 23

SYSTEMATIC AND DESCRIPTIVE 

PALEONTOLOGY

Descriptive terms applicable to rugose coral structure 

have been adequately defined and stabilized by Hill 

(1935,1956). Hill's terminology is widely accepted and, 

with minor modifications, is adopted for purposes of 

this report.

In general, the diagnoses and descriptions here presented take into account only gross features of the 

corallum. Fine skeletal structure (Kato, 1963; Wang, 

1950) was observed in thin sections of a few well-preserved corals; the study of minute structural features 

did not, however, contribute materially to the taxonomic results. More often than not, silicification has 

destroyed original trabecular details which might otherwise be useful.

Among the described corals are several that are 

designated provisionally by letter in preference to a 

formal species name. These lettered taxa have undoubted stratigraphic value, but study material is at 

present considered inadequate for specific naming.

Family STREPTELASMATIDAE Nicholson 1889

(in Nicholson and Lydekker 1889 as Streptelasmidae)

Subfamily SIPHONOPHRENTINAE, new subfamily

Reference form. Siphonophrentis gigantea (Lesueur) 1821. Middle Devonian, New York.

Diagnosis. Streptelasmatidae with narrow to wide 

septal stereozone, amplexoid septa and a weak to prominent fossula; tabulae mostly complete, straight or 

domed, with axial and peripheral sag. Normally no 

dissepiments.

Remarks. Nondissepimented solitary corals of this 

subfamily are especially characteristic of the North 

American Early and early Middle Devonian and include 

Siphonophrentis in the strict sense, Siphonophrentis 

(Breviphrentis) of Great Basin Devonian coral zones C 

and D and Heterophrentis and Compressiphyllum of 

the Onondaga Limestone of eastern North America. 

Nevadaphyllum and Homalophyllum may also belong 

in this family. Members of this group are quite uncommon above Great Basin coral zone D and are unknown 

above zone F.

Genus Siphonophrentis O'Connell 1914

1914. Siphonophrentis O'Connell, p. 190.

1949. Siphonophrentis O'Connell. Stumm, p. 12-13.

1950. Siphonophrentis O'Connell. Wang, p. 214. 

1956. Siphonophrentis O'Connell. Hill, p. F270. 

1960. Siphonophrentis O'Connell. Oliver, p. 87.

Type species Caryophyllia gigantea Lesueur 1821; 

by original designation. Onondaga Limestone, New 

York.

Diagnosis. Oliver's diagnosis (1960, p. 87) is as 

follows:

Simple ceratoid to cylindrical corals with amplexoid septa withdrawn from axis except on and just above upper surfaces of 

tabulae. Tabulae dome-shaped, flat, or concave axially, 

depressed at the fossula. No axial structure or dissepiments.

Remarks. Siphonophrentis in the strict sense commonly develops very elongate solitary coralla of large 

size. Breviphrentis Stumm (1949, p. 13) of the Great 

Basin Early and early Middle Devonian (coral zones 

C and D) is regarded by the writer (Merriam, 1973b) 

as a subgenus of Siphonophrentis; it differs from 

Siphonophrentis (Siphonophrentis) by having narrow, 

very elongate coralla of markedly segmented appearance because of the shelving indentations and rejuvenescence flanges.

Siphonophrentis (Siphonophrentis) sp. b 

Plate 1, figures 16, 17

Figured material USNM 166423. Roberts Mountains; Bobcat Peak locality M1342. Unit 4 of the 

Nevada Formation.

Diagnosis. Large Siphonophrentis having septal 

stereozone of medium width and closely spaced, mostly 

complete tabulae which bend downward sharply at 

periphery.

Transverse thin section. Major septa 44, the longest 

extending about one-half the distance to the axis; minor 

septa short, one-third to one-half length of major septa. 

Stereozone about 3 mm wide, which is great for this 

genus; some septa thickened stereoplasmically through 

most of their length. No fossula observed. One segment 

of a transverse thin section reveals what appear to be 

three small, closely spaced, peripherally concave dissepiments in sequence near the stereozone.

Longitudinal thin sections. Some closely spaced 

tabulae complete, but most extend only part of the 

width; a few short tabellae near periphery. Abrupt 

peripheral sag adjoining the stereozone. Tabulae 

thickened stereoplasmically on distal surfaces. Several 

very small dissepimentlike structures in a small segment of one thin section near stereozone.

Fine structure. Trabecular structure poorly defined; 

peripheral stereozone reveals fine lamellar texture.

Comparison with related forms. This large species 

resembles Siphonophrentis (Siphonophrentis) gigantea 

(Lesueur), the type species, more closely than Siphonophrentis (Breviphrentis) invaginatus (Stumm) of the 

lower part of the Nevada Formation. Siphonophrentis 

(Siphonophrentis) sp. b differs from S. gigantea by 

having a peripheral tabular sag, longer minor septa, and 

a wider stereozone. S. gigantea possesses a fossula not 

recognized in S. sp. b.

Occurrence. Middle Devonian coral zone F. Roberts 

Mountains, Bobcat Peak locality M1342. Unit 4 of the 

Nevada Formation. Study material consists of one 

incomplete corallum.
24 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

Family CYSTIPHYLLOIDAE Stumm 1949 

Genus Cystiphylloides Chapman 1893

1859. Cystiphyllum aggregatum Billings, p. 137, text fig. 28. 

1893. Cystiphylloides Chapman, p. 46. 

1937. not Cystiphylloides Yoh, p. 50-54, pi. V, figs. 1-4. 

1940. Cystiphylloides Chapman (in part). Stumm, p. 39 (in

part); not pi. 19, figs. 1-7; not pi. 20, figs. 14-15. 

1956. Mesophyllum (Cystiphylloides) Chapman, 1893. Hill, 

p. F319 (in part), fig. 219-6.

Type species. Cystiphyllum aggregatum Billings, 

1859. Onondaga Limestone, Devonian, near Simcoe, 

Ontario, Canada. The taxonomic problems relating to 

this genus have been dealt with elsewhere (Merriam, 

1973b).

Diagnosis. Dendroid to phaceloid and solitary 

cystiphylloid Rugosa in which single corallites range 

from trochoid to maturely cylindrical. Mature dissepimentarium of medium width, composed mostly of small 

to medium-sized, partly subglobose dissepiments in the 

type species. Border between tabularium and dissepimentarium fairly well defined but not sharp, with 

change from small steep dissepiments to large axially 

inclined tabellae. Tabularium medium wide to wide. 

Septa and septal spines or crests commonly absent in 

mature coralla of some species.

Remarks. A dendroid colonial Cystiphylloides from 

the Onodaga Limestone of Ontario, Canada, placed in 

Billings' type species aggregatum reveals almost no 

septal spines or crests. Great Basin forms here 

classified as Cystiphylloides may have weak to stout 

peripheral crests and scattered spines internally. Cystiphylloides of the Devonian parallels structurally the 

Silurian genus Cystiphyllum, which commonly, but not 

always, has radially alined trabecular septal spines; 

these are usually more slender and more numerous than 

the stronger crests and spines which characterize some 

species of Cystiphylloides. Other Silurian cystiphyllids 

like Hedstroemophyllum have strong internal spines. 

Peripheral wall crests do not appear to be a feature of 

true Cystiphyllum. As there are no recognized fine 

structure distinctions, the Silurian forms are in some 

instances difficult to separate. Many of the Devonian 

forms of Cystiphylloides, as that genus is here construed, are axially quite excentric, as in "Nardophyllum" Wedekind 1925 and "Wedekindophyllum" 

Stumm 1949, the generic validity of which is doubtful. 

Such marked excentricity has not been observed among 

the Silurian cystiphyllids. None of the Silurian forms 

are known to be colonial, like some species of Devonian 

Cystiphylloides.

Stumm's Family Cystiphylloidae seems to be at present a more appropriate receptacle for these Devonian 

cystiphyllids than either Cystiphyllidae or Digonophyllidae (Hill, 1956); assignment to the Digonophyllidae 

as a subgenus of Mesophyllum is open to question.

Cystiphylloides, as here construed, rather conclusively 

does not belong in synonymy with Plasmophyllum, 

where the genus has recently been placed by Birenheide 

(1964, p. 17).

Cystiphylloides sp. a 

Plate 1, figures 1-3

Figured material USNM 166417. Diamond Range, 

Alhambra Hills locality M1337; topmost beds in the 

Woodpecker Limestone Member of the Nevada Formation.

Diagnosis. Cystiphylloides with excentric axis; 

peripheral stereozone with well-developed septal wall 

crests. Septa spines present on inner dissepiment and 

upper tabellar surfaces.

External features. A fairly large, maturely ceratoid 

to subcylindrical species, known only as a solitary coral 

having subdued but heavy rugae and well-defined 

longitudinal grooves. No external spines or excrescences 

observed.

Transverse thin sections. Stout septal wall crests 

in a narrow stereozone, numbering about 112 in a 

mature section of 52 mm diameter. Major and minor 

septa poorly differentiated. Medium-size dissepiments 

predominating; tabellae of about the same size range 

and configuration as dissepiments. Scattered small 

septal spines lacking radial alinement. Patches of 

stereoplasmic filling present within the tabularium and 

at periphery. Corallite axis somewhat excentric, but less 

so than in earlier growth stages.

Longitudinal thin sections. Tabularium not sharply 

set off from dissepimentarium. Tabularium width less 

than one-third of corallite diameter. Dissepiments 

mostly elongate, mostly of medium size for the genus, 

and of fairly uniform pattern; tabellae of same size 

range as dissepiments, numbering six to eight, transversely alined in plane of section, some with nearly 

horizontal bases. Tabellar sag in early ephebic stage 

revealing the excentric corallite axis. Peripheral stereozone excessively thickened in places; stereoplasmic festoons extend here and there across the tabularium. 

Sporadic septal spines, mostly slender, standing normal 

to inner dissepiment surfaces and projecting upward 

from tabellae.

Fine structure. Steroplasmic thickenings minutely 

lamellar. Septal crests and spines showing no trabecular 

structure.

Comparisons with related forms. This species differs from C. spinosum McLaren (in McLaren and Norris, 1964, pi. IX) by having a wider tabularium, stronger 

wall crests, and a more excentric axis. Cystiphylloides 

iddingsi n. sp. has larger tabellae and dissepiments and 

is externally spinose; C. sp. c has much larger tabellae 

and dissepiments and lacks the wall crests. Cystiphylloides in the Great Basin Early and early Middle
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 25

Devonian lack the septal spines and crests and are less 

prone to be axially excentric. Certain of these older 

forms are colonial.

Occurrence. Middle Devonian coral zone G. Diamond Range, Alhambra Hills locality M1337; Eureka 

mining district, Nevada. Uppermost beds of Woodpecker Limestone Member. Study material comprises 

one large nearly complete corallum and smaller fragmentary coralla.

Cystiphylloides iddingsi n. sp.

Plate 1, figures 4,15

1884. Cystiphyllum n. sp. Walcott, p. 106 (in part). 

1940. Cystilphylloid sp. a. Merriam, p. 56.

Type material. Holotype USNM 166418a; paratype USNM 166418b. Lone Mountain, Eureka County, 

Nev.; unit 4 of the Nevada Formation.

Diagnosis. Cystiphylloides having stout, elongate 

wall crests, large tabellae, marked axis excentricity, and 

external spines.

External form. Mature growth stages of 50 mm 

diameter with subcylindrical form. Rugate exterior 

with cycles of tubular, tapering spines 4 millimeters or 

more long. Weak longitudinal grooves largely masked 

by epitheca. Calice deep, with V-shaped profile and 

highly excentric axis. Broken septal ridges well defined 

on calice wall.

Transverse thin sections. In a mature growth stage 

of 31 mm diameter, large wall crests 3 mm long extend 

from a narrow stereozone. Inner dissepiments and 

tabellae very large for the genus. Corallite axis about 

halfway between center and wall.

Longitudinal thin sections. Dissepiments mostly 

elongate and having steeper axial inclination than the 

very large elongate tabellae, distribution of which 

reflects the excentric conical calice shape. Irregular 

stereoplasmic fillings in tabularium and extending 

peripherally.

Fine structure. In transverse section, the elongate 

wall crests reveal divergent fibrilar structure extending 

from a median dark line, without suggestion of trabecular fiber grouping.

Comparison with related forms. Cystiphylloides sp. 

a has smaller and much more numerous dessepiments 

and tabellae, a wider tabularium, and more numerous 

septal spines internally. Cystiphylloides sp. c lacks the 

wall crests of sp. a and of C. iddingsi, revealing no septal 

spines internally.

Occurrence. Middle Devonian coral zone F. Lone 

Mountain, Eureka County, Nev., localities M1341, 

M1362. Unit 4 of the Nevada Formation. Study material consists of 24 incomplete coralla.

Cystiphylloides sp. c 

Plate 1, figures 5, 6

Figured material. USNM 166419. Middle Devonian, Morey Peak area locality M1338; Hot Creek 

Range, Nev.

Diagnosis. Cystiphylloid, with a few large and very 

elongate dissepiments poorly differentiated from tabellae, a very narrow stereozone, and no septal crests 

or spines.

External features. Mature corallum subcylindrical 

with broad subdued rugae; no external spines or longitudinal grooves observed.

Transverse thin section. No septal spines or wall 

crests. The few large internal dissepiments with globose 

traces; outer dissepiments flattened. Axis excentric. No 

stereoplasmic fillings.

Longitudinal thin section. Disposition of the very 

large elongate dissepiments reflecting the V-shaped 

calice profile and excentric position of the axis.

Fine structure. Narrow peripheral stereozone with 

a weakly fibrous rather than lamellar structure; no 

trabecular spines.

Comparison with related forms. See Cystiphylloides 

sp. a and C. iddingsi n. sp. This form is suggestive of 

Silurian Microplasma, but lacks the distinctive trabecular wall structure of that genus.

Occurrence. Probably Middle Devonian coral zone 

F. Hot Creek Range, Morey Peak area, locality M1338, 

north of Morey Peak. Study material is one incomplete 

corallum.

Family DIGONOPHYLLIDAE Wedekind 1924

Subfamily ZONOPHYLLINAE Wedekind 1924

Genus Zonophyllum Wedekind 1924

This genus, of which the type species is Zonophyllum 

duplicatum Wedekind 1924, of the German lower Middle Devonian, includes Digonophyllidae with thick, 

commonly stumpy septal crests and longer incomplete 

stereoplasmically thickened septa. Septa may be irregularly thickened and connected laterally in stereoplasmic festoons. The taxonomy of this genus has 

recently been treated elsewhere (Merriam, 1973b).

Zonophyllum sp. r 

Plate 1, figures 7, 8

Figured material. USNM 166420. Nevada Formation; Roberts Mountains, Red Canyon locality M1339.

Diagnosis. Zonophyllum with thickened, nonuniform stumpy septa in outer part of mature dissepimentarium and a complex peripheral dissepiment pattern 

in which some septa abort short of the wall.

External features. Corallum small, ceratoid, with 

deep calice having a V-shaped profile.

Transverse thin section. Septa about 28 in a corallite 

of 26-mm diameter; septa not clearly differentiated as
26 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

major and minor, somewhat wavy and irregularly 

thickened. Longest septa extending more than onethird the distance to axis. Many peripheral dissepiments small, some terminating against a single septum 

in digonophyllid fashion. Some outer dissepiments 

thickened stereoplasmically on axial sides. Complex 

peripheral herringbone patterns where minor septa are 

suppressed.

Longitudinal thin section. Tabularium comprising 

three to seven tabellae, which are larger than dissepiments. Dissepiment columns in some coralla exceeding 

nine on each side. Stereozone of medium width. Outer 

dissepiments partly small and subglobose.

Comparison with related forms. Septa are more 

uniformly developed than in Zonophyllum duplicatum 

Wedekind the type species; likewise the Zonophyllum 

species of lower beds in unit 2 of the Nevada Formation 

and coral zone D have less uniform septa and stereoplasmic festoons not present in the available material 

of Z. sp. r.

Occurrence. Middle Devonian, possibly coral zone 

G. Roberts Mountains; locality M1339, Red Canyon. 

Study material consists of one incomplete corallum.

Subfamily DIGONOPHYLLINAE Wedekind 1924 

Genus Digonophyllum Wedekind 1923

Type species. Digonophyllum schulzi Wedekind 

1923; lower Middle Devonian Nohner Schichten, Eifel 

district, Germany.

Diagnosis. Digonophyllidae with major septa radially continuous and elongate, approaching the axis. 

Tabularium relatively narrow, not sharply differentiated. Pattern of dissepiments complex; normal dissepiment traces anguloconcentric, commonly becoming 

highly irregular peripherally, with scattered minute 

dissepiments that terminate against a single septum. 

Some minor septa discontinuous peripherally. A lonsdaleioid band and scattered strip carinae present in some 

species. Septa thickened radially, especially within the 

tabularium.

Remarks. Digonophyllum includes large maturely 

subcylindrical corals. Septal thickening extends from 

the tabularium toward the periphery in varying degree. 

Some species have no appreciable septal stereozone. 

The symmetry is entirely radial, and there is usually 

no recognizable fossula in mature growth stages.

Digonophyllum with continuous major septa probably grades into Mesophyllum, which, as here construed, has radially discontinuous septa grading into 

nearly aseptate cystimorphs. Birenheide (1964) elected 

to suppress the genus Digonophyllum as a synonym of 

Mesophyllum in a taxonomic scheme differing sharply 

from that proposed by Hill (1956, p. F314-F321) for 

the digonophyllids. Hill advocated adoption of some

digonophyllid taxa as subgenera of either Digonophyllum or Mesophyllum, a course that appears reasonable at present in preference to Birenheide's suppression of Digonophyllum and use of Mesophyllum as a 

subgenus of Plasmophyllum.

Members of the Family Ptenophyllidae Wedekind 

1923 that includes Acanthophyllum parallel structurally and are associated with the digonophyllids. Both 

may have a peripheral lonsdaleioid band, but that of 

Acanthophyllum is more regular and uniform. Ptenophyllidae differ by having a wider and better differentiated tabularium with partly straight tabulae, as well 

as an axial whorl of long major septal terminations 

commonly thickened terminally in lobose fashion. 

Ptenophyllidae have continuous septa and lack the 

peripheral strip carinae of digonophyllids.

Digonophyllum (Digonophyllum) occidentalis n. sp.

Plate 3, figures 10, 11

Type material. Holotype USNM 166431. Roberts 

Mountains, Bobcat Peak locality M1342.

Diagnosis. Digonophyllum, with excessively thickened septa, some of which are laterally in contact within 

the tabularium. Lonsdaleioid band wide, with small 

outermost dissepiments.

Transverse thin sections. Septal count about 104 

in a corallite of 40-mm diameter. A few septa extending 

to the axis and a few traceable to the periphery as 

crests. Nonuniform lonsdaleioid band 8 mm wide in 

places; within the band, the largest dissepiments in the 

middle, the outer zone 2 mm wide is made up of small 

dissepiments. Thickened septa somewhat wavy; inner 

dissepiment surfaces also coated by stereoplasm. No 

appreciable septal stereozone.

Longitudinal thin sections. Tabularium narrow, 

comprising one to three flattened tabellae. Calice deep, 

with V-shaped profile.

Fine structure. Longitudinal sections reveal trabecular septal structure, the trabeculae being inclined 

peripherally.

Comparison with related forms. This species differs 

from D. schulzi Wedekind, the type species, by having 

more thickened septa and a wide lonsdaleioid band and 

by lacking strip carinae. It differs from Digonophyllum 

sp. c by having a lonsdaleioid band, longer septa, and a 

less complex pattern of normal dissepiments. D. occidentalis n. sp. is closest to D. (Mochlophyllum) alhambraensis that has less thickened septa in the tabularium 

and thinned major septa in the lonsdaleioid band, some 

of which reach the wall.

Occurrence. Middle Devonian coral zone F. Roberts 

Mountains; Bobcat Peak locality M1342. Unit 4 of the 

Nevada Formation. Study material consists of five 

incomplete coralla.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 27

Digonophyllum sp. c

Plate 2, figures 11, 12

Figured material. USNM166428. Confusion Range, 

Utah. Locality Ml343, Simonson Dolomite.

Diagnosis. Digonophyllum with septa thickened 

throughout and considerably withdrawn from axis. Dissepiment traces in a highly irregular pattern without 

lonsdaleioid band. Tabularium poorly differentiated.

Transverse thin section. Septal count about 80 in a 

corallite of 40-mm diameter. Major septa extending 

more than one-half the distance to axis; some minor 

septa exceeding one-half the length of major septa. 

Septal stereoplasmic thickening extending laterally 

upon inner surfaces of dissepiments.

Longitudinal thin section. Flat tabellae similar in 

size and shape to larger dissepiments. Upper tabellar 

and inner dissepiment surfaces with stereoplasmic 

coating.

Fine structure. Stereoplasmic thickenings faintly 

lamellar, but without trabecular features.

Comparison with related forms. Digonophyllum sp. 

c differs from D. schulzi Wedekind, the type species, by 

having shorter major septa thickened throughout, 

whereas those of schulzi are thickened only in the 

tabularium. D. schulzi has a few strip carinae not present in D. sp. c. Digonophyllum? sp. b has longer excessively thickened septa and^a distinct lonsdaleioid band.

Occurrence. Confusion Range, Utah. Fish Spring 

quadrangle, locality M1343, northwest of Ibex. Middle 

Devonian, upper beds of Simonson Dolomite. Study 

material is one incomplete corallum.

Subgenus Mochlophyllum Wedekind 1923 (as a genus)

Type species. Mesophyllum maximum Schliiter 

1889, by monotypy. Lower Middle Devonian, Eifel district, Germany.

Diagnosis. Very large, maturely subcylindrical Digonophyllum with greatly narrowed tabularium at 

maturity, long thickened continuous major septa, 

dilated especially in the tabularium, a very complex, 

irregular pattern of normal dissepiments, and a nonuniform lonsdaleioid band.

Remarks. The largest individuals of Digonophyllum 

(Mochlophyllum) may have sporadic strip carinae 

peripherally, but not to the extent developed in Mesophyllum (Arcophyllum) of Markov (1926) and in the 

probably equivalent Cosmophyllum1 of Vollbrecht 

(1922). Very small outermost lonsdaleioid dissepiments are characteristic of the subgenus Mochlophyllum.

1 Coamophyttum Vollbrecht (1922) is a homonym and should probably be 

superseded by Arcophyllum, the name Cosmophyllum having been used in 

1851 for & member of the Orthoptera.

Digonophyllum (Mochlophyllum) alhambraensis n. sp. 

Plate 3, figures 1-5

Type material Holotype USNM 166429. Diamond 

Range, Alhambra Hills locality M1337. Topmost beds 

of Woodpecker Limestone Member of the Nevada 

Formation.

Diagnosis. Very large Mochlophyllum with septa 

thinned peripherally in lonsdaleioid band and lacking 

strip carinae.

External features. Very large coralla of this species 

with nearly cylindrical shape at maturity; deep calice 

having V-shaped profile and narrow brim. Rugae fairly 

heavy; no longitudinal grooves observed, as the holotype is largely encased by a stromatoporoid.

Transverse thin sections. Septal count 142 in a 

corallite of 67-mm diameter. Some major septa 

approaching the axis and some passing peripherally to 

the wall; minor septa usually not extending peripherally as far as major septa. Septal thickening greatest in 

major septa, especially toward the tabularium; septa 

thinned where present in lonsdaleioid band. Lonsdaleioid band as much as 7 mm wide, with very small 

dissepiments near wall. No strip carinae. Some dissepiments thickened stereoplasmically on inner surfaces. Some minor septa discontinuous peripherally. 

Stereozone very narrow.

Longitudinal thin sections. Normal dissepiments in 

as many as 26 columns of medium to small size on each 

side. Tabularium narrow and poorly differentiated, 

comprising two or three tabellae.

Fine structure. Stereoplasmic thickenings minutely 

lamellar and lacking distinctly trabecular features.

Comparison with related forms. This new species 

differs from the type species D. (Mochlophyllum) maximum (Schliiter) by having more uniformly and evenly 

thickened major septa, by lacking strip carinae, and by 

having more thinned major septa continuing peripherally to the wall.

Occurrence. Middle Devonian coral zone G. Diamond Range; Alhambra Hills locality M1337. Topmost 

beds of the Woodpecker Limestone Member. Study 

material is one very large incomplete corallum and fragments of other coralla.

Digonophyllum (Mochlophyllum) alhambraensis 

subsp. roberisensis n. subsp.

Plate 3, figures 6, 7

Type material Holotype USNM 166430. Roberts 

Mountains; Red Canyon, locality M1339. Middle Devonian, Nevada Formation.

Diagnosis. Mochlophyllum with septa less thickened than with alhambraensis s.s. and having fewer 

large lonsdaleioid dissepiments.
28 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

Transverse thin section. Septal count about 98 in a 

corallite of 46-mm diameter. Lonsdaleioid band 2-5 mm 

wide. Septa somewhat wavy and unevenly thickened; 

minor septa with peripheral discontinuities. Outer 

lonsdaleioid dissepiments mostly small. Septal stereozone narrow; many major septa traceable to wall. No 

septal grooves externally.

Longitudinal thin section. Tabularium very narrow. Dissepimentarium having as many as 26 columns 

of medium and small normal dissepiments on each side. 

Calice, deep, with V-shaped profile.

Comparison with related forms. This subspecies 

differs from alhambraensis s.s. by having thinner septa, 

fewer large dissepiments, and a narrower lonsdaleioid 

band.

Occurrence. Middle Devonian coral zone F or G. 

Roberts Mountains; Red Canyon, locality M1339. 

Nevada Formation. Study material is one incomplete 

corallum.

Genus Mesophyllum Schliiter 1889

1956. Mesophyllum Schliiter 1889. Hill, p. F318.

Type species. Mesophyllum defectum Schliiter 1889. 

Middle Devonian; Eifel district, Germany.

Diagnosis. As here interpreted, this genus includes 

Digonophyllinae in which nonuniformly discontinuous, 

mostly thinned septa become obsolete peripherally in 

a wide band of irregularly lonsdaleioid dissepiments 

with or without local groupings of strip carinae.

Remarks. Corals assigned to Mesophyllum range in 

structure from maturely subcylindrical cystimorphs 

with only a few weak internal traces of septa and 

scattered or no strip carinae to forms with numerous, 

but discontinuous or broken and some fairly strong, 

continuous, irregularly thickened septa and abundant 

strip carinae. It is probable that Mesophyllum intergrades with Digonophyllum. Cystiphylloides is not here 

classified as a subgenus of Mesophyllum, as was done 

by Hill (1956, p. 319), being assigned to the Family 

Cystiphylloidae of Stumm (1949).

In classification of Great Basin Devonian Rugosa 

(Merriam, 1973b), Mesophyllum is adopted as a genus 

with subgenera, more or less in accord with the scheme 

proposed by Hill (1956, p. F317-F320). At present this 

arrangement seems to be a reasonably good rationalization of the taxonomic dilemma in this complex coral 

group. Birenheide's (1964) exceedingly detailed systematics are, on the other hand, a considerable departure from Hill's, with the classifying of Mesophyllum as 

a subgenus of Plasmophyllum.

Of the seven subgenera of Mesophyllum listed by Hill 

(1956) only two, Mesophyllum (Mesophyllum) and 

Mesophyllum (Lekanophyllum), are pertinent here.

Mesophyllum (Mesophyllum) sp. f

Plate I, figures 13, 14

Figured material. USNM 166422. Lone Mountain, 

Eureka County, Nev.; locality M1341.

Diagnosis. Mesophyllum s.s. having an irregular, 

nonuniform lonsdaleioid band and groups of thin, 

broken, wavy septa extending in places to the wall.

Transverse thin section. The thin, broken, wavy 

septa very unevenly developed and in places difficult to 

distinguish within a complex dissepimental network. 

Larger, more even lonsdaleioid dissepiments patchy 

and separated here and there by a network comprising 

smaller uneven dissepiments and wavy septal crests. 

Some septa extending to the wall.

Longitudinal thin section. Except for scattered 

large lonsdaleioid dissepiments peripherally, the dissepiments mainly of small to medium size, generally 

smaller than the tabellae.

Comparison with related forms. Mesophyllum (Lekanophyllum) sp. 1 differs by having a more uniform 

lonsdaleioid band and better developed septa. It is, 

however, possible these are merely variants of a single 

species, a problem calling for quantitative analysis of a 

large suite of specimens from the same bed.

Occurrence. Middle Devonian coral zone F. Lone 

Mountain, locality M1341. Unit 4 of the Nevada Formation. Study material is one incomplete corallum.

Mesophyllum (Mesophyllum)? sp. k

Plate 1, figures 9-11

1884. Cystiphyllum n. sp Walcott, p. 106 (in part). 

1938. (?)"Cyathophyllum" kobehense Stumm, pi. 59, fig. la 

only.

Figured material. USNM 96213c. Lone Mountain, 

Eureka County, Nev. Nevada Formation.

Diagnosis. Small trochoid rugose coral with peripheral band of lonsdaleioid dissepiments and somewhat 

wavy, little-thickened septa, most of which extend 

more than half the corallite radius.

External features. Single known individual of this 

form showing early neanic axial rejuvenescence by 

oblique offset. A single large axial offset developed in 

mature calice. Septal grooves very weakly defined.

Transverse thin section. Somewhat wavy, littlethickened septa numbering 58; minor septa nearly as 

long as major septa. All septa considerably withdrawn 

from axis. Only a few septa discontinuous peripherally. 

The longitudinal section of this form not known.

Remarks. The possibility that this form should be 

assigned to Digonophyllum rather than Mesophyllum 

has been considered in view of the radial continuity of 

most septa. The single specimen here figured, collected 

by Walcott in 1880, is one of those initially referred by 

him to Cystiphyllum. It is among Stumm's labeled
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 29

paratypes (USNM 96213c) of "Cyathophyllum" kobehense Stumm. A thin section (pi. 1, fig. 10) cut from 

the proximal half of the broken corallite (pi. 1, fig. 9) 

does not agree in structural detail with Stumm's transverse thin section of kobehense (Stumm, 1938, pi. 59, 

fig. la), which lacks lonsdaleioid features and was cut 

from another coral. A transverse thin section (pi. 4, 

fig. 7) of Stumm's holotype of "Cyathophyllum" kobehense (USNM 96213a) recently cut does not appear 

to represent the same genus and species and is herein 

classified as Keriophyllumf kobehense (Stumm). It is 

possible that Mesophyllum (Mesophyllum)'? sp. k may 

be an immature corallite of either Mesophyllum (Lekanophyllum) sp. 1 or Mesophyllum (Mesophyllum) 

sp. f, a problem which will require study of large suites 

of associated specimens from unit 4 of the Nevada 

Formation.

Occurrence. The specimen here designated as Mesophyllum (Mesophyllum)^ sp. k was recognized by 

Stumm as among those collected at Lone Mountain by 

Walcott (1884, p. 106) and referred by him to Cystiphyllum. Preservation of the fossil indicates that it 

probably came from unit 4 and coral zone F, where it 

was probably associated with Cystiphylloides sp. b, as 

suggested by Walcott's discussion.

Subgenus Lekanophyllum Wedekind 1924 (as a genus)

1924. Lekanophyllum Wedekind, p. 29, 35; text figs. 36-48. 

1940. Lecanophyllum Wedekind. Lang, Smith, and Thomas,

p. 75. 

1949. Lecanophyllum Wedekind. Stumm, p. 47; pi. 22, figs.

14, 15. 

1956. Mesophyllum (Lekanophyllum) Wedekind. Hill, p.

F318, fig. 219-2a-c.

Type species. Lekanophyllum punctatum Wedekind 1924; by subsequent designation Lang, Smith, and 

Thomas. Lower Middle Devonian, Eifel district, 

Germany.

Diagnosis. Mesophyllum with a moderately wide to 

wide lonsdaleioid band, numerous fairly continuous and 

continuous major septa withdrawn from the axis and a 

fairly wide, but not sharply differentiated, tabularium 

in which the tabellae are mostly larger than the 

dissepiments.

Remarks. The proportion of more or less continuous septa in the radial sense is greater in Lekanophyllum than in Mesophyllum s.s., which may also have 

strip carinae not recognized in the former. Septa are 

more likely to be thickened in Lekanophyllum than in 

Mesophyllum s.s. Mesophyllum (Atelophyllum) Wedekind 1925 appears to differ little from sub genus Lekanophyllum, except possibly for thinner septa of species 

which have been classified in Atelophyllum. (See 

McLaren and Norris, 1964, pis. VIII, X-XII.)

Mesophyllum (Lekanophyllum) sp. 1

Plate 1, figure 12

Figured material. USNM 166421. Lone Mountain, 

Eureka County, Nev., locality M1340.

Diagnosis. Lekanophyllum with wide fairly uniform lonsdaleioid band and numerous wavy, partly discontinuous septa considerably withdrawn from the 

axis. Minor septa poorly differentiated.

External features. Mature corallum subcylindrical; 

epitheca with longitudinal grooving.

Transverse thin section. Septal count 62 is in section of 29 mm diameter. Septa wavy, withdrawn from 

axis, partly discontinuous radially and poorly differentiated as major and minor. Wall crests absent; septal 

spines uncommon in peripheral part of dissepimentarium. Wall thin, with epitheca revealing distinct longitudinal grooves. Many of the outermost lonsdaleioid 

dissepiments small, the largest being in the middle of 

the band. Tabularium less than one-third the corallite 

diameter. Longitudinal section unknown.

Comparison with related forms. Mesophyllum (Lekanophyllum) punctatum (Wedekind), the lectotype, 

has more numerous septa, better differentiated as 

major and minor and a narrower lonsdaleioid band, its 

septa are more broken radially. A species from the Horn 

Plateau Formation of western Canada compared with 

M. (L.) punctatum by McLaren (in McLaren and 

Norris, 1964, p. 23, pi. 8, figs. 3, 4) also has a narrower 

lonsdaleioid band and somewhat thicker septa which 

are differentiated. Similar Horn Plateau corals assigned 

to Atelophyllum nebracis McLaren (in McLaren and 

Norris, 1964, p. 24, pis. 10-12) have a lonsdaleioid band 

like that of the Nevada form here described, but have 

more thickened, poorly differentiated septa.

Occurrence. Middle Devonian coral zone F. Lone 

Mountain locality Ml340. Unit 4 of the Nevada Formation. Study material consists of one nearly complete 

corallum and fragmentary coralla.

Family PTENOPHYLLIDAE Wedekind 1923 

Genus Acanthophyllum Dybowski 1873

1873. Acanthophyllum Dybowski, p. 339.

1923. Ptenophyllum Wedekind, p. 26-34; text figs. 2, 5-6.

1924. Ptenophyllum Wedekind, p. 35-44; text figs. 49-62. 

1949. Acanthophyllum Dybowski. Stumm, p. 20; pi. 9, figs. 1-7. 

1956. Acanthophyllum Dybowski. Hill, p. F303 (in part); fig.

207-la-b, not fig. 207-lc-d. 

1961. Acanthophyllum Dybowski. Birenheide, p. 77-146; pis.

1-7.

Type species. Cyathophyllum heterophyllum Edwards and Haime 1851; by subsequent designation 

Schluter 1889.

Diagnosis. Large maturely subcylindrical rugose 

corals having a wide calice brim and many columns of 

normal and subordinate lonsdaleioid dissepiments;

506-412 O - 73 - 5
30 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

long, somewhat thickened, lamellar and continuous 

septa terminating axially in a more or less twisted 

whorl; a discrete, sharply differentiated tabularium 

comprising close-spaced tabulae, some of which are 

complete.

Remarks. The pattern of dissepiment traces is 

anguloconcentric internally, but peripherally may become complex and irregular, passing in some species 

into a lonsdaleioid band; small to minute dissepiments 

that terminate against a single septum are characteristic. Axial terminations of septa are commonly lobose, 

as in the type species A. heterophyllum. Minor septa 

are usually more than half the length of major septa 

and are thinner. Acanthophyllum includes loosely colonial species.

Certain members of the Family Digonophyllidae bear 

a superficial convergent resemblance to Acanthophyllum and other Ptenophyllidae with which they 

may be associated. Both families comprise large solitary corals with long, thickened septa and a complex 

pattern of normal and lonsdaleioid dissepiments. However, the tabularium of digonophyllids is never as well 

differentiated and rarely has complete tabulae; the 

digonophyllids do not possess the twisted, lobose axial 

whorl of the ptenophyllids. A well-defined lonsdaleioid 

band is more characteristic of the Digonophyllidae than 

of Ptenophyllidae, and the peripheral strip carinae, so 

characteristic of some digonophyllids, do not occur in 

the Ptenophyllidae. Unlike many digonophyllids, none 

of the Ptenophyllidae have broken or radially discontinuous septa.

Birenheide (1961) critically evaluated the taxonomic 

status of Acanthophyllum and other members of the 

Ptenophyllidae of Wedekind. In this light several 

Wedekind ptenophyllid genera appear to be synonyms 

of Acanthophyllum. Birenheide reclassified certain 

other Wedekind genera as subgenera under Acanthophyllum.

Ptenophyllidae like Acanthophyllum are quite characteristic of Eifelian Stage Middle Devonian in Europe. 

In the Great Basin these corals occur in Devonian coral 

zone F, continuing upward in diminishing numbers into 

coral zone G. No Ptenophyllidae are known here below 

coral zone F. Digonophyllidae appear earlier in Great 

Basin Devonian coral subzone D2 of possible earliest 

Eifelian age and are common in coral zone G of Givetian 

age.

Acanthophyllum sp. a

Plate 2, figures 3-6

Figured material. USNM 166425a, b, c. Diamond 

Range localities M6, M53, M204 and S8; Oxyoke Canyon and Alhambra Hills. Woodpecker Limestone Member of the Nevada Formation.

Diagnosis. Acanthophyllum with thickened septa, 

very numerous, mostly small dissepiments, and lacking 

a lonsdaleioid band.

Transverse thin sections. Septal count 72 in a corallite of 31-mm diameter; septal thickening of some individuals extending into axial whorl. Dissepiment pattern 

mostly simple concentric to anguloconcentric. Stereozone narrow.

Longitudinal thin sections. Normal dissepiments 

mostly small; columns on each side occasionally exceeding 18. Tabularium comprising combinations of partly 

straight, close-spaced tabulae and flat tabellae.

Comparison with related forms. Acanthophyllum 

sp. a differs from A. robertsensis n. sp. by having thickened septa and much more numerous and smaller 

normal dissepiments and by lacking the lonsdaleioid 

band of robertsensis.

Occurrence. Higher beds of coral zone F. Diamond 

Range, upper beds of the Woodpecker Limestone Member of the Nevada Formation. Oxyoke Canyon localities M6, M53; Alhambra Hills localities M204, S8. 

Study material comprises seven incomplete coralla.

Acanthophyllum robertsensis n. sp.

Plate 2, figures 1, 2

1940. Tabulophyllum nevadense Stumm. Merriam, p. 58; pi. 

14, fig. 6.

Type material. Holotype USNM 166434a; paratype USNM 166424b. Roberts Mountains; Bobcat 

Peak locality Ml342.

Diagnosis. Large Acanthophyllum with well-developed lonsdaleioid band, septa of axial whorl thinned, 

and a discrete tabularium with flat tabellae and closespaced tabulae, some of which are nearly straight. 

Calice fairly deep, with V-shaped profile.

Transverse thin sections. Septal count 72 in a corallite of 38-mm diameter. Septa thin or very little thickened, fairly straight; minor septa two-thirds the length 

of major septa. Tabular extensions of major septa very 

long and unthickened within the irregularly twisted 

axial plexus. Normal dissepiments mostly of simple 

form, concentric to anguloconcentric and straight, 

closely spaced in places. Lonsdaleioid band up to 6 mm 

wide, with three or more cycles of medium and large 

dissepiments. Peripheral stereozone well developed, but 

rather narrow.

Longitudinal thin sections. Tabularium sharply set 

off, about one-third of corallite diameter; tabulae closespaced (two per millimeter). Tabularium comprising 

various combinations of incomplete tabulae, which are 

peripherally straight and slightly arched axially, with 

wide, flat tabellae. Innermost dissepiments generally 

small and nearly vertical; normal dissepiments in 9-14 

columns on each side.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 31

Comparison with related forms. This species lacks 

the septal thickening of A. heterophyllum, the type 

species; it also has a better developed, wider lonsdaleioid band and a less complex, more uniform pattern 

of concentric normal dissepiments. The coral figured by 

Merriam (1940, pi. 14, fig. 6) from the same area as 

"Tabulophyllum nevadense Stumm" is conspecific with 

Acanthophyllum robertsensis n. sp. and is a loosely 

colonial form with a slightly wider stereozone.

Reproductive offsets. The coral referred to above 

figured by Merriam has a lateral offset which probably 

developed from the calice rim.

Occurrence. Middle Devonian, coral zone F. Roberts Mountains, Bobcat Peak locality Ml342. Unit 4 

of the Nevada Formation. Study material consists of 

three incomplete solitary coralla and a partial colonial 

corallum.

Acanthophyllum sp. c

Plate 2, figures 7, 8

Figured material. USNM 166426. Diamond Range; 

Oxyoke Canyon, locality M6.

Diagnosis. Small Acanthophyllum lacking a lonsdaleioid band, having thickened septa and a wide tabularium for the genus.

Transverse thin section. Septal count 66 in a corallite of 25 mm diameter. Dissepiments small, with fairly 

uniform concentric pattern. Stereozone narrow. Axial 

segments of septa little thickened in whorl.

Longitudinal thin section. Tabularium wide and 

less sharply differentiated than normal for this genus, 

made up of rather flat tabellae. Dissepiments mostly 

small and fairly uniform, disposed in six or more 

columns on each side.

Fine structure. Trabeculae in thickened septa flat 

or inclined peripherally at a low angle.

Comparison with related forms. This form differs 

from Acanthophyllum sp. a by having a narrower and 

more uniform dissepimentarium, and from A. robertsensis n. sp. by having thicker septa and a wider tabularium and by lacking the lonsdaleioid band.

Occurrence. Middle Devonian coral zone G. Diamond Range; Oxyoke Canyon locality M6. Basal beds 

of the Bay State Dolomite Member of the Nevada Formation. In upper Oxyoke Canyon the light-buff-colored 

silty dolomite at locality M6 contains large valves of 

Stringocephalus and digitate favositids in association 

with this species and other solitary rugose coral genera. 

Study material is one incomplete corallum.

Acanthophyllum sp. p

Plate 4, figures 9-11

Figured material. USNM 166433a, b. Roberts 

Mountains; Pyramid Hill localities M1345, M1348.

Diagnosis. Large Acanthophyllum with septa considerably thickened in tabularium, thinning in places 

near the periphery. Tabularium sharply differentiated, 

with closely spaced, uneven tabulae, some of which are 

complete.

Transverse thin section. Septal count 128 in a 

corallite of 54 mm diameter. Most major septa passing 

into the twisted axial whorl, where they are considerably thickened in lobose fashion and laterally in contact. No septal thickening in some peripheral segments. 

Septal stereozone very narrow; longitudinal grooves 

faintly developed. Minor septa less thickened than 

major septa. Normal dissepiments concentric to anguloconcentric in a simple pattern. No lonsdaleioid band.

Longitudinal thin section. Tabularium in sharp 

boundary with dissepimentarium, narrow, about onefourth the corallite diameter; tabulae undulant, some 

nearly complete and closely spaced (3-5 per mm). Dissepimentarium wide, with as many as 26 columns on 

each side.

Fine structure. Trabecular structure with steep 

peripheral inclination shown faintly in longitudinal 

section.

Comparison with related forms. Acanthophyllum 

sp. p differs from A. heterophyllum, the type species, by 

having a narrower tabularium, more uniformly thickened septa, and a simpler dissepiment pattern. It differs 

from A. robertsensis n. sp., which has a lonsdaleioid 

band and unthickened septal ends in the tabularium. 

Acanthophyllum sp. a differs by having a wider tabularium.

Occurrence. Middle Devonian coral zone F. Roberts Mountains; Pyramid Hill localities M1345, M1348. 

Unit 4 of the Nevada Formation. Study material consists of three incomplete coralla.

Acanthophyllum? sp.

Plate 2, figures 9,10

Figured material. USNM 166427. Roberts Mountains, Red Canyon, locality M1339.

Diagnosis. Acanthophyllum-]ike solitary corals with 

many columns of small, normal dissepiments, a tabularium comprising close-set, partly straight tabulae, an 

incipient axial whorl, and no lonsdaleioid band.

Transverse thin section. Septal count 84 in a corallite of 28-mm diameter. Most minor septa not extending to edge of tabularium. Axial whorl weakly developed 

for this genus. Dissepiment traces simple, concentric, 

rather crowded peripherally. Stereozone very thin.

Longitudinal thin section. Dissepiment columns as 

many as 18 on each side; outer dissepiments very small, 

their size increasing toward the tabularium. Closely 

space tabulae, three or four per millimeter; some tabu-
32 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

lae nearly complete, others combined with wide, flat 

tabellae. Tabular sag characteristic.

Fine structure. Trabeculae well developed in 

thickened septa, either flat or with low peripheral inclination.

Comparison with related forms. This form has a 

well-differentiated tabularium with more complete tabulae than is usual for the genus, but the axial whorl is 

less developed than normal for Acanthophyllum. 

Whereas the tabulae are more like those of A. robertsensis n. sp. than other forms of Acanthophyllum here 

described, it differs from A. robertsensis n. sp. in weakness of the axial whorl and absence of a lonsdaleioid 

band. Small size of the outer dissepiments distinguishes 

the form in question from all others here dealt with.

Occurrence. Middle Devonian coral zone F or G. 

Roberts Mountains, Red Canyon, locality M1339. 

Nevada Formation.

Genus Paracanthus, new genus 

1867. Aulophyllum? Richardsoni Meek, p. 81, pi. 11, figs. 3,

3a. 

1891. (1) Cyathophyllum Richardsoni Meek. Whiteaves. p.

200, pi. 27, figs. 3, 4.

1945. Mictophyllum richardsoni (Meek). Smith, p. 34 (in 

part), pi. 5, figs. 10-12.

Type species. Aulophyllum? Richardsoni Meek 

1867. The Ramparts, MacKenzie River, Canada. 

Middle Devonian.

Diagnosis. Solitary Ptenophyllidae with septa 

thickened in outer part of dissepimentarium, thinning 

toward axis, lacking the complex axial whorl of Acanthophyllum. Tabularium wide, comprising fairly continuous straight tabulae with flattened tabellae; not 

sharply differentiated from dissepimentarium. A weak 

fossula present in some coralla.

Remarks. Paracanthus n. gen. differs from Acanthophyllum by its lack of the complex axial whorl of 

septal terminations, by having more restricted septal 

thickening, and by possessing a wider tabularium that 

is less sharply differentiated at the margin. Unlike 

Acanthophyllum, some species show a suggestion of a 

fossula. Mictophyllum Lang and Smith differs by having obsolete or very short minor septa and little thickening of septa. Sterictophyllum Pedder (1965) has a much 

wider septal stereozone.

Paracanthus nevadensis n. gen., n. sp. 

Plate 4, figures 14-19; plate 6, figures 4-7 

Type material. Holotype, NSNM 166434; paratypes, USNM 166435a, b, c. Roberts Mountains; Red 

Canyon, locality M1339.

Diagnosis. Paracanthus with fairly smooth, slightly 

undulant septa in which the greatest thickening is 

within the peripheral half. Dissepiment pattern fairly 

simple concentric-anguloconcentric, crowded peripherally with a few irregular dissepiments near the wall. 

Peripheral dissepiments small, steeply inclined.

Transverse thin sections. Septal count 72 in a corallite of 33-mm diameter (holotype). A few major septa 

reach the axis; minor septa about one-half the length 

of major septa. Where the dissepiments are crowded 

peripherally, stereoplasm also pervading dissepiments. 

Septal stereozone usually narrow, but widened moderately in places. A few modified chevron septal traces 

near the wall. Fossula defined by union of two major 

septa in the tabularium.

Longitudinal thin sections. Tabularium very wide. 

Small, vertical to steeply inclined dissepiments in six 

to 10 columns on each side, with sporadic large dissepiments. Tabularium with numerous flat tabellae that 

are larger than most dissepiments.

Fine structure. Well-defined, rather coarse trabecular structure shown in longitudinal sections; large trabeculae nearly horizontal in widened parts of septal 

stereozone.

Comparison with related forms. Paracanthus richardsoni (Meek) from the Mackenzie River Ramparts, 

Canada (Smith, 1945, p. 34, pi. 5, figs. 11-12). has 

fewer septa and more slender, subcylindrical corallites. 

Paracanthus? sp. f from Lone Mountain has shorter 

septa and a slightly narrower tabularium, with more of 

the nearly complete, straight tabulae.

Occurrence. Middle Devonian coral zone F or G. 

Roberts Mountains; Red Canyon, locality M1339. 

Nevada Formation. Study material consists of six 

incomplete coralla.

Paracanthus? sp. f 

Plate 4, figures 12,13

Figured material. USNM 166436. Lone Mountain, 

locality M1344.

Diagnosis. Small Paracanthus-like coral with very 

wide, flattened tabellae with intercalated straight, 

complete tabulae.

Remarks. This coral differs from Paracanthus 

nevadensis n. gen., n. sp. by having shorter septa and 

more of the wide straight tabulae; trabeculae at the 

periphery are more steeply inclined away from the axis.

Occurrence. Middle Devonian coral zone F. Lone 

Mountain, locality M1344. Unit 4 of the Nevada Formation. Study material consists of a partial corallum 

and fragmentary coralla.

Family CYATHOPHYLLIDAE Dana 1846

Genus Cyathophyllum Goldfuss 1826 

1826. Cyathophyllum Goldfuss, p. 54; pi. 15, fig. 13. 

1846. Cyathophyllum Dana (in part). 

1940. Cyathophyllum Goldfuss, Lang, Smith, and Thomas, p.

44 (in part).

1949. Cyathophyllum Goldfuss. Stumm, p. 22; pi. 10, fig. 9. 

1956. ^Cyathophyllum Goldfuss. Hill, p. F278. 

1963. Cyathophyllum Goldfuss. Birenheide, p. 369.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 33

Type species. Cyathophyllum dianthus Goldfuss 

1826; by subsequent designation Dana 1846. Devonian, 

Eifel district, Germany. Uncertainties persist regarding 

internal characteristics of the holotype of dianthus 

(Stumm, 1940, p. 23; Hill, 1956, p. F278). Stumm 

chose Goldfuss' (1826) pi. 15, figure 13 as lectotype, 

which is not known to have been sectioned. "Cyathophyllum dianthus" Lonsdale, a Silurian coral, is 

excluded. Birenheide (1963, p. 377, pi. 26, fig. 1, pi. 50, 

figs. 19a-b, pi. 51, fig. 22) selected and figured a neotype of C. dianthus Goldfuss from the Givetian Loogher 

Schichten of the Eifel district.

Diagnosis. Solitary and colonial rugose corals having wide mature corallites numerous long major lamellar 

septa, some of which reach the axis, a wide dissepimentarium comprising many columns of prevailingly 

small globose dissepiments, and a tabularium made up 

of combinations of tabellae with straight tabulae, of 

which some are nearly complete. A fossula present in 

some mature corallites of solitary species.

Remarks. Colonial forms like the type species 

range in growth habit from phaceloid to loosely cerioid. 

Colonial forms have no fossula or other evidence of 

bilateral symmetry. Lamellar stereoplasmic thickenings 

of septa and a peripheral stereozone are uncommon. 

Cyathophyllum lacks yardarm septal carinae. Some 

species have decidedly wavy, uneven septa with abundant minor zigzag carinae. Multiple calice platform 

reproductive offsets are characteristic, these being more 

commonly peripheral.

Cyathophyllum lends itself to subgeneric classification (Birenheide, 1963, p. 369-403), the following subgenera being recognized here:

1. Cyathophyllum (Cyathophyllum) Goldfuss 1826

2. Cyathophyllum (Peripaedium) Ehrenberg 1934

3. Cyathophyllum (Moravophyllum) Kettnerova 1932

4. Cyathophyllum (Orthocyathus) newsubgenus

Characters used in subgeneric separation include 

the tendency in certain species to develop minutely 

wavy septa peripherally with abundant zigzag carinae, 

the tendency as in Cyathophyllum sensu stricto toward 

minute and complex peripheral branching of septa, and 

the relative abundance of continuous tabulae in some 

species, as compared with the tabellae that predominate 

in other species. Only solitary forms are known in 

Moravophyllum; the other three subgenera are colonial, 

as known at present. Moravophyllum and the new 

Orthocyathus have fairly straight, smooth, thin septa 

with a minimum of zigzag carinae and no septal branching peripherally.

Heliophyllum, normally but not always a solitary 

coral, may be distinguished from Cyathophyllum by 

its abundant and prominent yardarm carinae; the solitary Keriophyllum of the Nevada Formation differs

from Heliophyllum by lacking yardarm carinae and 

having abundant zigzag carinae throughout. Large 

corallites of Keriophyllum may reveal complex peripheral branching of septa similar to Cyathophyllum sensu 

stricto.

Dana's (1946) Family Cyathophyllidae is provisionally adopted for Cyathophyllum, Keriophyllum, and 

other genera for which the family name Zaphrentidae 

has been used by other authors. The true characterization of the type species of Zaphrentis remains to be 

clarified.

Subgenus Moravophyllum Kettnerova 1932 (as a genus) 

1932. Moravophyllum Kettnerova, p. 27, 79. 

1940. Moravophyllum Kettnerova. Lang, Smith, and Thomas,

P. 86.

1949. Moravophyllum Kettnerova. Stumm, p. 21. 

1956. Moravophyllum Ketterova. Hill, p. F278.

Type species. Moravophyllum ptenophylloides Kettnerova 1932, by author designation. Late Middle 

Devonian (Givetian), Celechovice, Moravia.

Diagnosis. Solitary Cyathophyllum with numerous 

smooth, uniformly thin, fairly straight, uncomplicated 

septa and a weak fossula in mature growth stages. 

Tabularium wide, comprising numerous closely spaced 

tabellae and sporadic tabulae, some being nearly continuous.

Remarks. Kettnerova's text figure of a neanic 

transverse section of the type species reveals thickened 

major septa in two quadrants, which are probably 

cardinal. Other Cyathophyllum subgenera seemingly 

do not retain at maturity a clear indication of bilateral 

symmetry like that of mature Moravophyllum with its 

weak fossula.

Cyathophyllum (Moravophyllum) alhambraensis n. sp. 

Plate 5, figures 6-10

Type material Holotype, USNM 166440; Diamond 

Range, Alhambra Hills locality M1337. Paratype, 

USNM 166441; Sulphur Spring Range, locality M1346.

Diagnosis. Moravophyllum with uniformly simple 

concentric-anguloconcentric pattern of dissepiment 

traces and a wide, well-differentiated tabularium with 

subordinate straight, nearly complete tabulae.

External features. This species fairly large and subcylindrical with well-marked rugae and weak septal 

grooves. Calice wide, deep, and flat-bottomed.

Transverse thin sections. Septal count 80 in a corallite of 32-mm diameter; septa smooth, fairly straight, 

unthickened, the longest slightly withdrawn from the 

axis. Minor septa mostly two-thirds the major septa 

length. Stereozone very narrow; septal grooves slightly 

indented. Epitheca distinct. In places near the wall the 

anguloconcentric dissepiment pattern somewhat irregular and the septa slightly thickened. Fossula of holotype indistinct.
34 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

Longitudinal thin sections. Tabularium slightly 

less than one-half the corallite diameter; flattened 

tabellae predominate with some fairly straight, nearly 

complete tabulae. Dissepiments mostly small and globose in as many as 15 columns on each side; the peripheral dissepiments inclined axially at low angles; those 

toward the axis nearly vertical.

Fine structure. Longitudinal sections show patches 

of peripherally inclined trabecular structure.

Comparison with related forms. The type species 

M. ptenophylloides Kettnerova differs by having a 

wider peripheral zone of nearly flat dissepiments and 

fewer straight, nearly complete tabulae. The fossula 

is more clearly defiined in the type species. Moravophyllum sp. 1 has a narrower dissepimentarium.

Occurrence. Middle Devonian coral zone G. Diamond Range, Alhambra Hills locality M1337. Sulphur 

Spring Range, locality M1346. Topmost beds. Woodpecker Limestone Member of the Nevada Formation. 

Study material consists of three incomplete coralla.

Cyathophyllum (Moravophyllum) sp. 1 

Plate 4, figures 1-4

Figured material. USNM 166432a, b. Lone Mountain, locality Ml344.

Diagnosis. Small Moravophyllum with unusually 

wide tabularium and relatively narrow dissepimentarium. Fossula present.

Transverse sections. Septal count 74 in a corallite 

of 19 mm diameter. Septa smooth, thin, straight to 

slightly wavy, the longest extending to axis; minor 

septa more than one-half the length of major septa. 

Fossular depression defined by union of two major 

septa near axis.

Longitudinal sections. Tabularium about one-half 

the corallite diameter. Dissepiments in about six 

columns on each side. Tabularium comprising three to 

nine tabellae, some wide and flat, with occasional nearly 

complete, uneven tabulae. Calice deep, wide, and flatbottomed.

Comparison with other forms. This form differs 

from M. ptenophyloides, the type species, by having a 

narrower disspimentarium. Moravophyllum alhambraensis n. sp. also has the wider dissepimentarium, but 

with many more columns of small dissepiments.

Occurrence. Middle Devonian coral zone F. Lone 

Mountain locality M1344. Unit 4 of the Nevada Formation. Study material consists of four incomplete 

coralla.

Subgenus Orthocyathus, new subgenus

Type species. Cyathophyllum (Orthocyathus) flexurn (Stumm) 1938. Middle Devonian, unit 4 of the 

Nevada Formation; Lone Mountain, Eureka County,

Nev.

Diagnosis. Colonial Cyathophyllum having long, 

straight, fairly smooth septa with little waviness, and 

few or no zigzag carinae or other complications. Tabularium comprising a combination of tabellae and nearly 

straight to slightly arched tabulae. No suggestion of a 

fossula in mature corallites.

Remarks. Corallites on distal surfaces of coralla 

show a very great size range from very wide to very 

narrow by reason of the numerous interstitial offsets 

from parental calice brims. Coralla range from openly 

phaceloid to loosely cerioid, occasionally within the 

same colony. Orthocyathus lacks the peripheral septum 

complications of Cyathophyllum sensu stricto and of 

the subgenus Peripaedium. It differs from subgenus 

Moravophyllum by having the colonial growth habit, 

by lacking a fossula, and in the frequency of reproductive offsets from the calice brim.

Cyathophyllum (Orthocyathus) flexum (Stumm)

Plate 7, figures 1-8

1884. Cyathrophyllum n. sp. Walcott, p. 104. 

1938. Prismatophyllum flexum Stumm, p. 483; pi. 58, figs.

6a-d; pi. 59, figs. 6a-c.

1940. "Cyathophyllum" flexum (Stumm). Merriam, p. 56, 78; 

pi. 13, figs. 1, 2.

Type material. Holotype, USNM 96219a; paratypes, USNM 96219b, c, d. Middle Devonian, unit 4 

of the Nevada Formation; Lone Mountain, Eureka 

County, Nev.

Diagnosis. Orthocyathus with largest corallites of 

great maximum diameter in large coralla that range in 

habit from openly phaceloid to loosely cerioid. Septa 

relatively straight, smooth, and uncomplicated peripherally. Tabellae predominating over tabulae.

External form. As noted by Stumm (1938, p. 483), 

the coralla of this species vary greatly in external 

appearance, depending upon degree of openness or 

compactness of the colony. The holotype is a small 

not very compact cerioid mushroom-shaped colony 

showing distally a large number of small interstitial 

offset corallites. Mature corallites have a wide, deep, 

rather straight-sided calice with a median boss and a 

flattened or reflected brim, in some corallites beset with 

small daughter corallites. The underside of the holotype 

also reveals offsets that appear to have grown from the 

sides of the parent as noted by Walcott (1884, p. 104). 

More compact cerioid colonies (pi. 7, fig. 5) have polygonal mature corallite walls and fewer interstitial corallites. Loose phaceloid colonies (pi. 7, figs. 2, 3, 7, 8) 

show a great diameter range of the subcylindrical corallites, which expanded less rapidly during growth.

Transverse thin sections. Septal count of mature 

corallites averaging about 58, may exceed 60. Septa 

normally straight, uniformly thin and simple, less commonly undulant; minor zigzag carinae uncommon.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 35

Axial twisting and intersection of longest septa sufficient in some corallites to form a calice boss, but not 

producing a discrete axial structure as observed in 

longitudinal sections. Minor septa about two-thirds the 

length of major septa. Dissepiment traces rather uniformly concentric, with sporadic chevrons peripherally. 

No lonsdaleioid dissepiments. Wall thin, straight to 

strongly but evenly curved in cerioid coralla.

Longitudinal sections. Tabularium one-third to 

one-half the corallite diameter, comprising more or less 

flattened tabellae which are overall somewhat arched 

distally. Globose dissepiments disposed in about 12 

columns on each side of which the innermost are 

inclined axially, the middle four are flat and the outer 

ones are inclined peripherally, thus indicating reflection of the calice brim. Tabellae larger than most dissepiments. Septa showing no amplexoid axial extensions.

Fine structure. No initial fine structure is preserved 

in these silicified corals.

Reproductive offsets. Offsets probably exceeding 

seven, developing from the reflexed calice rim, as illustrated in plate 7, figure 2. Offsets also shown by the 

numerous small interstitial corallites (pi. 7, fig. 1), some 

of which appear to grow from the side of the parent, 

rather than from an intermediate point on the calice 

brim.

Comparison to related forms. No described species 

closely related to this species are known. It is possible 

that some forms previously assigned to Hexagonaria 

may belong in this new subgenus of Cyathophyllum. 

Other described species of colonial Cyathophyllum fall 

in other subgenera having structural complexities in the 

peripheral band. (See Birenheide, 1963.)

Occurrence. Middle Devonian coral zone F. Lone 

Mountain localities: M54, M1360, B1363. Unit 4 of the 

Nevada Formation. Type material collected by C. D. 

Walcott at Lone Mountain in 1880. Study material 

comprises five nearly complete coralla and 15 incomplete coralla.

Genus Keriophyllum Wedekind 1923

1923. Keriophyllum heiligensteini Wedekind, p. 27, figs. 3a-b.

1940. Ceriophyllum Lang, Smith, and Thomas, p. 35.

1949. Heliophyllum heiligensteini (Wedekind). Stumna, p. 21,

pi. 9, fig. 12. 

1956. Keriophyllum Wedekind. Hill, p. F278, fig. 190-2a-b.

Type species. Keriophyllum heiligensteini Wedekind, by author designation. Middle Devonian, Heiligenstein, Eifel district, Germany.

Diagnosis. Solitary rugose corals with numerous 

long, minutely zigzag septa with elbow carinae and a 

wide dissepimentarium. Most septa little thickened; 

the longer major septa typically reaching the axis, but 

not forming an axial whorl.

Remarks. No complete longitudinal section of topotype material of the type species K. heiligensteini is 

known to have been figured. Whereas Wedekind (1923, 

p. 35; 1924, p. 67-70) seems to have considered this 

genus affiliated with Heliophyllum, it lacks the diagnostic yardarm carinae of that genus and is not known to 

have longitudinal features of the American Heliophyllum. Lang, Smith, and Thomas (1940, p. 35) regard 

Keriophyllum as a synonym of Heliophyllum. According to Birenheide (1962a, p. 108; 1963, p. 390), on the 

other hand, Keriophyllum heiligensteini Wedekind, the 

type species, as a synonym of Cyathophyllum (Peripaedium) turbinatum Goldfuss, suggesting (McLaren, 

in McLaren and Norris, 1964, p. 18) that Keriophyllum 

be merged with Peripaedium.

Keriophyllum mclareni n. sp.

Plate 5, figures 1-4

Type material. Holotype, USNM 166437; paratypes, USNM 166438a, b. Lone Mountain, locality 

M1344.

Diagnosis. Medium and large Keriophyllum with 

mature septa withdrawn from the axis, a deep calice 

with rather narrowly V-shaped profile, and a very narrow tabularium. Pattern of small dissepiments and 

wavy septa complex peripherally.

Transverse thin sections. Septal count 96 in a corallite of 30-mm diameter. Minutely wavy septa considerably withdrawn from the axis; some not traceable to the 

wall. Minor septa generally more than one-half the 

length of major septa. Although there is no discrete 

lonsdaleioid band, scattered lonsdaleioid dissepiments 

occur at the periphery. In some mature individuals, the 

concentric pattern of normal dissepiments so modified 

as to produce a highly irregular and confused arrangement of these traces, especially towards the periphery. 

Small elbow carinae numerous along the zigzag septa; 

some minute nodular thickenings on one side only of 

the septum; others are actual spines passing outward 

diagonally. No true yardarm carinae observed. In late 

mature growth stages, some of the inner parts of septa 

slightly and uniformly thickened. Stereozone very 

narrow.

Longitudinal thin sections. Tabularium very narrow, lacking straight tabulae; tabellae mostly of 

medium to small size, flattened, numbering two to four 

across axial zone. Dissepimentarium very wide, having 

as many as 36 columns of mostly small dissepiments on 

each side. Trabeculae alined peripherally in such manner as to suggest weak strip carinae in depth.

External features. Coralla externally quite rugate 

with very weak longitudinal grooves; trochoid in early 

ephebic stages. Holotype constricted distally, having
36 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

its greatest diameter in earlier adult growth stages. 

Some large individuals with a more nearly turbinate 

configuration.

Fine structure. Longitudinal thin sections revealing well-defined trabeculae that are steeply inclined 

toward periphery. In outer part of dissepimentarium, 

trabeculae coalescing to give appearance of strip carinae 

in some places.

Comparison with related forms. Keriophyllum heiligensteini Wedekind, the type species, has longer major 

septa. Of described American corals, the nearest is 

Heliophyllum borealis McLaren (in McLaren and Norris, 1964) which has yardarm carinae (lacking in this 

form), longer septa, and a somewhat wider tabularium.

Occurrence. Middle Devonian coral zone F. Lone 

Mountain locality Ml344. Unit 4 of the Nevada Formation. Study material consists of six incomplete coralla.

Keriophyllum? kobehense (Stumm)

Plate 4, figure 7

1938. "Cyathophyllum" kobehense Stumm, p. 479; pi. 58, fig. 

la, not pi. 58, fig. Ib, not pi. 59, figs, la, Ib.

Type material. Holotype, USNM 96213a (Stumm, 

1938, pi. 58, fig. la). Stumm's paratypes, USNM 96213 

b, c, d, are probably not conspecific with the holotype. 

Middle Devonian; Lone Mountain, Eureka County, 

Nev.

Diagnosis. Small trochoid to ceratoid rugose coral 

with numerous long,wavy, thin septa, an irregular pattern of normal dissepiments, and a weakly developed 

nonuniform lonsdaleioid band.

Transverse thin section. Septal count 64 in corallite 

of 18-mm diameter. Many of the major septa reaching 

the axis. Some discontinuities in the thin, wavy irregular septa; some of the normal dissepiments more nearly 

straight than concentric. Longitudinal section of this 

coral unknown.

Remarks. The true generic relation of this species 

has not been established. In transverse section, this 

form shows thin wavy septa resembling those of Keriophyllum mclareni n. sp., which also has a few lonsdaileioid dissepiments. Other specimens among Stumm's 

paratypes of "Cyathophyllum" kobehense appear to 

represent species or genera different from the holotype, 

of which the transverse section is here figured (pi. 4, 

fig. 7). Another of Stumm's paratypes (USNM 96213c) 

here referred to as Mesophyllum? sp. 1 is doubtfully 

conspecific with kobehense. (See pi. 1, figs. 9-11.)

Occurrence. Middle Devonian, probably coral zone 

F. Lone Mountain; collected by C. D. Walcott in 1880, 

probably from unit 4 of the Nevada Formation.

Family SPONGOPHYLLIDAE Dybowski 1873

Genus Sociophyllum Birenheide 1962

(as subgenus of Stringophyllum)

1881. Spongophyllum Schliiter. 

1925. Spongophyllum. Wedekind, p. 18-19,47. 

Loipophyllum Wedekind, p. 53, 57. 

Neospongophyllum Wedekind, p. 57. 

1949. Spongophyllum semiseptatum Schliiter. Stumm, pi. 14,

figs. 12,13.

1962. Stringophyllum (Sociophyllum) Birenheide, p. 53. 

1962. Stringophyllum (Neospongophyllum?). McLaren, Norris, and McGregor, pi. 1, figs. 7, 8.

1964. Stringophyllum (Sociophyllum) Birenheide. McLaren 

and Norris, p. 20.

Type species. Spongophyllum elongatum Schliiter 

1881; by author designation. Middle Devonian, Eifel 

district, Germany.

Diagnosis. Loosely phaceloid Rugosa with maturely 

cylindrical corallites having a strong lonsdaleioid band, 

wide tabularium, and partly long, smooth lamellar 

septa that may be discontinuous radially or aborted.

Remarks. This distinctive genus so characteristic 

of the Cordilleran higher Middle Devonian is here 

viewed as an independent genus rather than a subgenus 

of Stringophyllum, as originally classified by Birenheide (1962b). Sociophyllum differs from the cerioid 

true Spongophyllum as typified by S. sedgwicki in 

growth habit, in its stronger lonsdaleioid marginarium, 

and in greater width and spacing of tabulae.

Sociophyllum eurekaensis n. sp.

Plate 8, figures 6-15 

1938. ?Disphyllum lonensis Stumm, pi. 59, fig. 4c only.

Type material. Holotype, USNM 166447; paratypes USNM 166448,166449. Lone Mountain localities 

M1340, M1344; Roberts Mountains locality M1348.

Diagnosis. Slender Sociophyllum with very large 

lonsdaleioid dissepiments; septa of most corallites not 

clearly differentiated as major and minor. Tabulae wide, 

having either peripheral or axial sag. Wall crests usually 

weakly developed.

Transverse thin sections. Septal count ranging from 

24 to 32, with an average of 27 in corallites of about 

9-mm diameter. Septa long, smooth, slightly wavy, 

either withdrawn from axis or, in some corallites, 

extending to axis. In a few corallites some very short 

minor septa present; usually no differentiation between 

major and minor septa. Some septa discontinuous 

peripherally with wall crests. Septal stereozone usually 

narrow. Largest lonsdaleioid dissepiments extending 

more than one-fourth of the corallite circumference.

Longitudinal thin sections. Tabularium usually 

exceeding one-half the corallite width. Lonsdaleioid
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 37

dissepiments in one or two columns, less commonly 

three or four; dissepiments ranging from near horizontal to steeply inclined toward the axis. Tabulae 

rather closely spaced, mostly complete or nearly complete, some with axial sag; some corallites with uniform 

sequences of complete tabulae that are axially flat with 

peripheral sag. Complete straight tabulae uncommon. 

Amplexoid septal projections uncommon. Large dissepiment distribution nonuniform, occurring here and 

there in groups.

Fine structure. Slightly thickened septa and stereozone revealing no clearly defined trabecular structure.

Comparison with related forms. Sociophyllum elongatum (Schluter), the type species, has more septa, 

including a greater number of minor septa, and all 

septa tend to be discontinuous. Sociophyllum glomerulatum (Crickmay) also has more numerous and more 

broken septa; in some corallites the septa are largely 

aborted (Pedder, 1964).

Occurrence. Middle Devonian coral zone F. Lone 

Mountain localities: M1340, M1344. Roberts Mountains, Pyramid Hill locality M1348. Unit 4 of the 

Nevada Formation. Study material comprises 10 partial coralla.

Sociophyllum eurekaensis subsp. b 

Plate 8, figures 16-18

Figured material. USNM 166448. Roberts Mountains, Bobcat Peak, locality Ml342.

This subspecies of S. eurekaensis differs from the 

typical form in its fairly uniform development of axially 

and periaxially straight tabulae with a peripheral tabular depression. Whereas some of the very wide tabulae 

are complete beneath the depression, most appear to 

terminate laterally at the inner margin of the peripheral 

depression that is floored by discrete tabular segments. 

Thus the tabularium may be said to comprise separate 

peripheral and axial-periaxial sequences. In transverse 

section, the very large lonsdaleioid dissepiments are 

less common in this subspecies.

Comparison with related forms. A species referred 

by McLaren (in McLaren and Norris, 1964, p. 19, pi. 

VII, figs. 2a-c) to Sinospongophyllum cf. S. planotabulatum Yoh has the tabular features of this form and 

appears otherwise closely related.

Occurrence. Middle Devonian coral zone F. Roberts 

Mountains, Bobcat Peak locality M1342. Unit 4 of the 

Nevada Formation. Study material consists of four 

partial coralla.

Genus Utaratuia Crickmay 1960 

1960. Utaratuia Crickmay, p. 5; pi. 1, figs. 6-9, pi. VIII, fig. 1.

1961. Utaratuia Crickmay. Lenz, p. 507; pi. II, figs. 1, 2.

1962. Utaratuia laevigata Crickmay. McLaren, Norris, and

McGregor, pi. II, figs. 5, 6. 

1964. Utaratuia Crickmay. Pedder, p. 443.

Type species. Utaratuia laevigata Crickmay, 1960; 

by author designation. Middle Devonian, Carcajou 

River, Northwest Territories, Canada.

Diagnosis. Cerioid rugose corals of cystimorph 

character with septa very short or absent; dissepiments 

lonsdaleioid in several columns. Tabularium usually 

sagging, nonuniform, comprising large tabellae with 

a few nearly complete tabulae.

Remarks. Utaratuia Crickmay 1960 may be a 

synonym of Tabellaephyllum Stumm 1948b, the type 

species of which is T. peculiaris Stumm from the Martin 

Limestone of the Bisbee area, Arizona, where it is considered to be of Late Devonian age. Stumm's holotype 

figures of T. peculiaris show no trace of septa or septal 

crests. In longitudinal section T. peculiaris shows a differentiation of the tabularium similar to that of Utaratuia. Study of additional material of the Arizona form 

may call for rejection of Utaratuia, especially if some 

individuals of Tabellaephyllum peculiaris are found to 

possess rudimentary septa. Some corallites of Utaratuia 

laevigata also have almost completely obsolete septa.

Homeomorphic forms are to be expected among unrelated cerioid Rugosa of this kind in which septa are 

reduced to peripheral crests or lost entirely and in 

which the dissepiments in consequence tend to become 

lonsdaleioid. Some species of Columnaria and of 

Hexagonaria like H. rohrensis Glinski (1955) have correspondingly reduced septa and few columns of dissepiments, but these are not lonsdaleioid and the 

tabularium is wider. Lonsdaleioid features favor the 

classification of Utaratuia with either Spongophyllidae 

or Endophyllidae. A better differentiated, more uniform 

tabularium and greater frequency of straight tabulae 

are more characteristic of both families than is the 

case with Utaratuia.

Assuming Utaratuia and Tabellaephyllum to be 

related, or even congeneric, the range of corals of this 

kind would be from Eifelian to Frasnian. Comparable 

morphologic trends occur in Australophyllum-^ke members of the Endophyllidae in the Late Silurian, some 

corallites of which have completely aborted septa.

Utaratuia eurekaensis n. sp.

Plate 8, figures 1-3

Type material Holotype USNM 166466 Lone 

Mountain, locality M1344.

Diagonsis. Small form of Utaratuia with rather 

thick wall, strong septal crests, broken outermost
38 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

columns of small lonsdaleioid dissepiments, and sporadic complete tabulae.

Transverse thin sections. Septal count 32 in a corallite of 7-mm diameter. Septa not differentiated as major 

and minor, mostly not stubby, varying in length from 

0.3 to 1 mm. Wall in places with zigzag trace shown by 

median dark line.

Longitudinal thin sections. Lonsdaleioid dissepiments in one to three columns on each side, of which 

the larger are quite elongate. Tabularium with sag, for 

the most part not sharply differentiated, comprising 

combinations of large, flat, axially inclined tabellae, 

horizontal tabellae, and a few complete, nearly horizontal to sagging tabulae. General appearance horizontally that of a cystimorph.

Fine structure. Thickened wall with well-defined 

median dark line, but no trabecular structure.

External features. Large rounded heads 25 cm or 

more wide in which the rather small corallites are of 

rather uniform size on the distal surface. Calices deep, 

without peripheral platform, and with V-shaped profile 

to rather straight sided.

Comparison with related forms. Utaratuia eurekaensis n. sp. may be conspecific with a Canadian form 

assigned with query by Lenz (1961, p. 507, pi. II, figs. 

1, 2) to Utaratuia stevensi (Chapman). U. laevigata 

Crickmay, the type species, has larger corallites with 

more numerous and generally weaker to obsolete septa 

and in places a thinner wall.

Occurrence. Middle Devonian coral zone F. Lone 

Mountain localities M1344, M1361. Unit 4 of the 

Nevada Formation. Study material consists of nine 

incomplete coralla.

Family ENDOPHYLLIDAE Torley 1933 

Genus Tabulophyllum Fenton and Fenton 1924

1924. Tabulophyllum Fenton and Fenton, p. 30; pi. VI, figs.

8-12. 

1928. Apolythophyllum Walther, p. 135, 144-145; text figs. 33,

34. 

1945. Tabulophyllum Fenton and Fenton. Smith, p. 58; pi. 2,

figs. 10,11, pi. 3, fig. 8. 

1949. Tabulophylum Fenton and Fenton. Stumm, p. 27; pi.

12, figs. 15,16,18,19. 

1956. Tabulophyllum Fenton and Fenton. Hill, p. 300, fig.

205-la-c.

Type species. Tabulophyllum rectum Fenton and 

Fenton 1924; by author designation. Hackberry Stage 

of Fenton and Fenton (1924), Upper Devonian, Iowa. 

Diagnosis. Ceratoid to subcylindrical solitary 

rugose corals with very wide tabularium comprising 

partly complete, fairly straight tabulae. Dissepiments 

large and mostly lonsdaleioid. Septa smooth, withdrawn from axis at maturity, discontinuous peripherally. No fossula.

Remarks. The figured original sections of the type 

species do not illustrate the distinctive transverse 

lonsdaleioid features of this genus; these characters are 

better shown in the illustrations of Smith, Stumm, and 

Hill. Genera which may be confused with Tabulophyllum are Ketophyllum of the Silurian and members of 

the Halliidae, such as Papiliophyllum and undescribed 

related forms, in the Lower Devonian. All of these have 

a well-developed cardinal fossula not possessed by 

Tabulophyllum; the Halliidae mentioned have strong 

septal thickening in cardinal quadrants persisting into 

mature growth stages. Ketophyllum is commonly of 

turbinate growth habit.

Diversophyllum (Sloss, 1939, p. 64) differs by having 

longer septa reaching the axis, more arched and irregular tabulae, and less tendency to produce lonsdaleioid 

dissepiments. Sinospongophyllum Yoh 1937 may be 

congeneric, but it appears to have a somewhat less 

developed lonsdaleioid band and perhaps more of a 

septal stereozone.

Occurrence. Tabulophyllum occurs in the Great 

Basin higher Middle Devonian beds of unit 4 of the 

Nevada Formation and coral zone F. More commonly, 

however, this genus is found in Late Devonian strata, 

as in Germany (Walther, 1928), southeastern Alaska, 

and Iowa.

Tabulophyllum antelopensis n. sp.

Plate 6, figures 8-12

Type material. Holotype, USNM 166442; paratypes, USNM 166443,166444. Antelope Range, Eureka 

County, Nev.; locality M1347.

Diagnosis. Tabulophyllum with many complete, 

fairly straight tabulae; septa considerably withdrawn 

from axis at maturity. Lonsdaleioid dissepiments partly 

large, very elongate both longitudinally and circumferentially. Wall crests strong, set in a narrow to medium 

width stereozone.

Transverse thin sections. Septal count 66 in a corallite of 26-mm diameter. Major septa extending onethird to more than one-half the distance to the axis. 

Minor septa thin and discontinuous; major septa very 

slightly thickened in tabularium. Largest lonsdaleioid 

dissepiments extending one-fourth the circumference of 

corallite. In places small normal dissepiments developed 

adjacent to tabularium. Stereozone less than 1 mm. 

wide. Septal grooves poorly defined in epitheca.

Longitudinal thin sections. Tabulae fairly even, 

mostly complete; spaced four tabulae per millimeter to 

more than 1 mm apart. Some lonsdaleioid dissepiments 

15 mm long. Tabulae ranging from horizontal to those 

with axial sag and with peripheral sag; axial arching 

very uncommon. Amplexoid septal projections not 

observed.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 39

External features. Mature corallites elongate, subcylindrical with prominent rounded rugae and weak 

longitudinal grooving. Calice deep and flat-bottomed. 

This species commonly colonial.

Fine structure. Septal stereozone minutely lamellar 

without well-defined trabeculae.

Comparison with related forms. The septa are more 

withdrawn from the axis and some of the lonsdaleioid 

dissepiments larger than in the type species Tabulophyllum rectum. Tabulophyllum mcconnelli (Whiteaves) reveals more axial arching of tabulae, somewhat 

longer septa at maturity, and amplexoid septal features 

not recognized in the new species. An undescribed 

species from the Coronados Islands of southeastern 

Alaska lacks the very elongate dissepiments of T. 

antelopensis.

Occurrence. Middle Devonian coral zone F. North 

end of the Antelope Range; locality Ml347. Unit 4 of 

the Nevada Formation. Study material consists of eight 

partial coralla.

Genus Australophyllum Stumm 1949 

1949. Australophyllum Stumm, p. 34; pi. 16, figures 1, 2.

Type species. Spongophyllum cyathophylloides 

Etheridge 1911; by author designation. Devonian, 

Queensland, Australia.

Diagnosis. Cerioid Endophyllidae with mediumwide to narrow, closely spaced, proximally sagging tabulae and a wide to very wide dissepimentarium having 

several columns comprising scattered to wholly lonsdaileioid dissepiments, many of which are large and 

elongate.

Remarks. Cerioid corals of this type with a few 

lonsdaleioid dissepiments have previously been referred to Spongophyllum or classified in the Spongophyllidae. It is recommended that the genus name Spongophyllum be restricted to forms with slender corallites, 

thickened walls, and few dissepiment columns, therein 

agreeing in general structure with S. sedgwicki, the 

type species, as originally illustrated by Edwards and 

Haime (1853, pi. 56, figs. 2, 2a-c, 2e). Close spacing 

of tabulae is more characteristic of Australophyllum 

than of Spongophyllum. It is probable that unrelated 

cerioid Rugosa developed lonsdaleioid features, which 

by themselves are not always viewed as evidence of 

close genetic affiliation.

Occurrence. Australophyllum is represented by 

undescribed subgenera in the Late Silurian. The genus 

in the strict sense is not known above the late Middle 

Devonian.

Australophyllum prismatophylloides (Stumm)

Plate 8, figures 4, 5

1884. tCyathophyllum davidsoni Milne-Edwards. Walcott, p. 

104.

1938. Spongophyllum prismatophylloides Stumm, p. 482; pi.

58, fig. 5a, pi. 59, figs. 5a-b.

1961. ?Australophyllum, cf. A. cyathophylloides (Etheridge), 

Lenz, p. 509, pi. II, figs. 4, 5.

Type material. Holotype, USNM 96217. Devonian, 

Nevada Formation; Lone Mountain, Eureka County, 

Nev.

Diagnosis. Cerioid rugose coral having long, thin, 

smooth major septa, irregularly developed and broken 

minor septa, and very unevenly distributed lonsdaleioid 

dissepiments of irregular shape.

Transverse thin section. Septal count about 32 in 

a corallite of 10-mm diameter. Thin straight or slightly 

wavy septa extending in part to the axis; minor septa 

ranging from one-half the length of major septa to discontinuous as septal crests, or entirely aborted. Pattern of dissepiment traces very uneven, ranging from 

concentric toward axis to lonsdaleioid peripherally. 

Some lonsdaleioid dissepiments irregularly concave 

toward the wall. In parts of some corallites all dissepiments irreguarly concave peripherally in lonsdaleioid 

fashion. Wall thin to slightly thickened.

Longitudinal sections. No longitudinal thin sections are available. According to Stumm (1938, p. 482) 

the tabularium is not sharply bounded, is about 5 mm 

wide, or about one-half the diameter, and is occupied 

by close-spaced incomplete tabulae.

Comparison with related forms. Australophyllum 

cyathophylloides (Etheridge), the type species, has a 

more regular pattern of lonsdaleioid dissepiments and 

possibly a thicker wall. A species from the Devonian of 

western Canada compared with cyathophylloides by 

Lenz (1961, p. 509; pi. II, figs. 4, 5) appears to be quite 

similar to prismatophylloides.

Irregularities of the dissepiment pattern make somewhat uncertain the assignment of this species to Australophyllum.

Occurrence. Probably Middle Devonian coral zone 

F. Collected at Lone Mountain by C. D. Walcott in 

1880, probably from unit 4 of the Nevada Formation.

Family DISPHYLLIDAE Hill 1939 

Genus Hexagonaria Giirich 1896

1896. Hexagonaria Giirich, p. 171.

1900. Prismatophyllum Simpson, p. 218.

1940. Hexagonaria Giirich. Lang, Smith, and Thomas, p. 69.

1945. Prismatophyllum Simpson. Smith, p. 44.

1948a. Hexagonaria Giirich. Stumm, p. 11.

1949. Hexagonaria Giirich. Stumm, p. 33.

1956. Hexagonaria Giirich. Hill, p. F280.

1967. Hexagonaria Giirich. Sorauf, p. 24.

Type species. Cyathophyllum hexagonum Goldfuss; by subsequent designation Lang, Smith, and 

Thomas, 1940, p. 69. Middle Devonian, Eifel district, 

Germany.
40 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

Diagnosis. Cerioid Disphyllidae forming large compact heads; deep calice pits of distal surface surrounded 

by inclined platform overlying the dissepimentarium. 

Tabularium medium to very wide. Dissepiments normalglobose to elongate in several columns, the outer ones 

flat or inclined at very low angles. Septa long, commonly thin and continuous from wall; zigzag and yardarm carinae common. Tabularium comprising partly 

complete nonuniform tabulae and tabellae, more commonly with slight arching than with overall sag. Uniformly distributed persistent thickenings of wall and 

septa uncommon. Lonsdaleioid dissepiments and horseshoe dissepiment columns absent.

Remarks. Cerioid colonies resembling the disphyllid Hexagonaria have arisen in other rugose coral 

families; among these are the Phillipsastraeidae, Spongophyllidae, Endophyllidae, and Kyphophyllidae. Of 

these, several of the most similar surficially are among 

the Phillipsastraeidae, which are distinguished by possessing columns of small horseshoe dissepiments with 

their associated trabecular bundles and by their more 

uniform septal thickenings. The other families mentioned have lonsdaleioid features and develop carinae 

to a lesser extent than do Hexagonaria and other members of the Disphyllidae.

Most American Middle Devonian species here interpreted as Hexagonaria in the strict sense possess carinae, commonly strong yardarm carinae, in some but not 

all corallites. These carinae may be restricted to parts 

of single corallites. Stumm's (1948a) figures of the type 

species Hexagonaria hexagona (Goldfuss) from the 

Eifel district (Germany) Middle Devonian reveal numerous yardarm carinae. On the contrary, Sorauf's 

(1967, p. 10, fig. 4) figures of Late Devonian (Frasnian) 

Hexagonaria from Frasnes, Belgium, assigned to H. 

hexagona, reveal no clear indication of yardarm carinae. 

It is assumed, although by no means established, that 

the Eifel district material is more nearly representative 

of the Goldfuss type species. (See Sourauf, 1967, p. 

33-36.)

Of uncertain family affiliation and unknown relation 

to Hexagonaria are three noncarinate Devonian groups 

of Hexagonaria-like corals: (1) Aberrant species structurally immediate between Hexagonaria and Columnaria, (2) the genus Xystriphyllum, and (3) a form 

group of cerioid corals having a few sporadic lonsdaleioid dissepiments. Group 1 is heterogeneous and 

polyphyletic, occurring in the Devonian of Europe; it 

comprises thick-walled species with very wide tabulae, 

a reduced dissepimentarium having few columns, and 

short, thick wedge-tapering septa (Glinski, 1955). 

Group 2, including Xystriphyllum, has a rather uniformly thickened wall, peripherally thickened, fairly 

uniform wedge-tapering, smooth septa, usually a narrow tabularium, and steeply inclined to vertical dissepiments peripherally. Group 3 with sporadic, mostly small 

and irregular lonsdaleioid dissepiments is represented 

by the species here called Australophyllum prismatophylloides (Stumm), possibly a Hexagonaria derivative. Lonsdaleioid supporting structures clearly arose 

in several genetically unrelated stocks. Australophyllum of the Endophyllidae differs by possessing a better 

defined and more uniform lonsdaleioid band.

Occurrence. Coral species assigned to Hexagonaria in the Great Basin have their earliest occurrence in 

unit 2 of the Nevada Formation (Devonian coral zone 

D2 ) and are not known with assurance above unit 4 

(Devonian coral zone F). The Hexagonaria of Hexagonaria kirki (Stumm) in Nevada unit 2 is considered a 

subgenus (Merriam, 1973b). The following are the 

species from Devonian coral zone F here dealt with:

Hexagonaria fisherae (Merriam)

H. fisherae subsp. cockalorumensis, n. subsp.

H. fisherae subsp. antelopensis, n. subsp.

Hexagonaria meeki n. sp.

Hexagonaria sp. c

Hexagonaria sp. f

Hexagonaria sp. n

Hexagonaria sp. w

Of uncertain stratigraphic horizon, possibly as high as 

Devonian coral zone G, are Hexagonaria sp. r and 

Hexagonaria sp. b (Merriam, 1940).

Hexagonaria has not been recognized during the 

course of this study in Great Basin rocks above coral 

zone G of Givetian (Middle Devonian) age. In coral 

zone I of Frasnian (Late Devonian) age in the upper 

part of the Devils Gate Limestone, the characteristic 

abundant colonial Rugosa are thamnastraeoid and 

aphroid Phillipsastraeidae, which rarely show traces of 

an outer wall. In western Europe the genus Hexagonaria 

ranges upward from the Belgian Givetian into the 

middle Frasnian (Sorauf, 1967). Overlying strata of 

the type area higher middle Frasnian contain the Phillipsastraea assemblages. Cerioid corals referred to 

Hexagonaria (McLaren and others, 1962), are reported 

in Frasnian rocks of western Canada.

"Prismatophyllum sp. undet." figured by Stumm 

(1940, pi. 8, fig. 12) from the White Pine district, 

Nevada, appears to be a carinate Hexagonaria. This 

material collected by an early-day exploring party is 

from an unknown stratigraphic position in an area 

where the Devils Gate Limestone of Frasnian age is 

well represented and coral faunas of Devonian coral 

zone I with Phillipsastraeidae are present.

Hexagonaria fisherae (Merriam)

Plate 11, figures 1-3

1938. Prismatophyllum profundum (Hall). Stumm, p. 482; 

pi. 59, figs. 7a-b.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 41

1940. Prismatophyllum fisherae Merriam, p. 78, pi. 13, figs. 

3,4.

Type material. Holotype, USNM 96390. Roberts 

Mountains; Pyramid Hill. Unit 4 of the Nevada Formation. Figured material, USNM 96218; Lone Mountain, Eureka County, Nev.; collected by C. D. Walcott.

Diagnosis. Large globular Hexagonaria with long 

septa reaching the axis, numerous zigzag and yardarm 

carinae, a wide to very wide tabularium, and septa 

thickened in the dissepimentarium. Corallites mostly 

slender.

Transverse thin sections. Septal count ranging 

from 32 to 40. Outer wall straight to zigzag, usually 

unthickened. Septa in many places stereoplasmically 

thickened from wall to tabularium; thinned septum 

extensions in tabularium wavy to twisted, meeting 

axially. Septa with rows of closely spaced zigzag carinae which pass into yardarm carinae; other septa lacking carinae or with yardarm carinae only.

Longitudinal thin section. Unknown for holotype. 

The Lone Mountain specimen (USNM 96218) with 

tabularium width more than one-half the corallite diameter; tabulae very nonuniform, wavy, and lacking 

straight elements, some slightly arched overall with 

minor axial sag. Dissepiments small, in two to five 

columns on each side; mostly flat or with very low 

axial inclination.

Comparison to related forms. This strongly carinate 

species differs from H. fisherae subsp. cockalorumensis 

by having more widespread septal thickening and more 

of the zigzag carinae.

Occurrence. Middle Devonian coral zone F. The 

type material collected by Merriam at Pyramid Hill, 

southern Roberts Mountains, in 1937 was not sectioned longitudinally. Although these specimens were 

float coral heads, they came from the outcrop belt of 

unit 4 of the Nevada Formation. No material of this 

species was collected in situ at Lone Mountain, although 

the Walcott specimen described by Stumm (1938) and 

here refigured probably came from unit 4 and coral 

zone F. This species also occurs in unit 4 at Devonian 

Peak, Roberts Mountains. Study material comprises six 

coralla.

Hexagonaria fisherae subsp. cockalorumensis n. subsp. 

Plate 11, figures 4-6; plate 12, figures 1-7

Type material. Holotype, USNM 166454; paratype, USNM 166455. Fish Creek Range; Cockalorum 

Wash area, locality M1351. Middle Devonian, Cockalorum Wash Formation.

Diagnosis. Hexagonaria forming very large globular 

heads of diameter in excess of 30 cm. Most corallites 

with septa having numerous yardarm carinae; longer 

septa of some corallites reaching the axis. Septa usually

thin, but in some corallites having patches of localized 

thickening. Tabularium moderately wide to very wide; 

dissepiments prevailingly small.

Transverse thin sections. Septal count ranging 

from 32 to 40. Septa normally thin, straight in dissepimentarium, commonly with strong yardarm carinae; 

wavy or twisted in tabularium, reaching the axis in 

some corallites. Minor septa exceeding one-half of 

major septal length, extending to tabularium. Zigzag 

carinae uncommon. Wall thin, usually straight. Concentric dissepiment traces commonly closely spaced in 

inner part of this band.

Longitudinal thin sections. Tabularium width ranging from less than one-half to more than one-half the 

corallite diameter. Dissepiments mostly small, in four 

to seven columns on each side; some outermost flatter 

dissepiments commonly elongate. Yardarm carinae with 

strong diagonal traces in depth. Tabularium comprising mostly very large tabellae with a few nearly straight 

complete tabulae; slight overall arching common; few 

tabular elements showing axial sag.

Fine structure. All study material is silicified and 

reveals no trabecular structure.

External features. Large globular mature coralla 

with modified mushroom shape, with constricted basal 

immature growth stage (pi. 12, fig. 1). Corallites narrow, of fairly uniform size, with very few small intermural offsets. Calice pits large, deep, fairly steep sided; 

peripheral calice platform flat or axially inclined. No 

low flattened mature platelike coralla like those of 

Taimyrophyllum. Reproductive offsets peripheral.

Comparison to related forms. This subspecies differs from typical H. fisherae by having less stereoplasmic septal thickening and fewer zigzag carinae. 

Hexagonaria meeki n. sp. has fewer and even thinner 

septa, the transverse dissepiment traces are less distinctly concentric, and the tabularium is narrower. 

Hexagonaria fisherae subsp. antelopensis differs by 

having more septa stereoplasmically thickened.

Occurrence. Middle Devonian coral zone F. Fish 

Creek Range, Cockalorum Wash area; Reef Hill, locality M1351; Coral Ridge, locality M1352. Cockalorum 

Wash Formation. Study material comprises 50 complete and partial coralla.

Hexagonaria fisherae subsp. antelopensis n. subsp.

Plate 13, figures 6, 7

Type material. Holotype, USNM 166456; paratype, USNM 166457. Antelope Range, locality M1347.

Diagnosis. Hexagonaria fisherae with numerous 

yardarm carinae, few zigzag carinae; most septa somewhat thickened stereoplasmically, long septa extending 

to axis; a wide tabularium; dissepiments mostly small.
42 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

Transverse thin sections. Septal count averaging 

about 32; septa straight and usually slightly and 

unevenly thickened in dissepimentarium. Yardarm 

carinae numerous and strong on some septa of many 

corallites. Long septa twisted in tabularium, extending 

to axis of some corallites. Concentric pattern of dissepiment traces well defined.

Longitudinal thin sections. Tabularium width 

usually less than one-half the corallite diameter. Dissepiments mostly small, with some elongate and flat 

toward periphery. Tabularium comprising large flat 

tabellae and nearly straight complete tabulae; slight 

overall arching of tabular elements common. Carinal 

traces well defined in depth.

Fine structure. Trabeculae ill defined.

Comparison with related forms. This subspecies 

differs from typical fisherae by having a narrower tabularium with more of the nearly straight, complete tabulae. Septal thickening is more characteristic of subsp. 

antelopensis than of subsp. cockalorumensis.

Occurrence. Middle Devonian coral zone F. Antelope Range, north end, locality M1347, unit 4 of the 

Nevada Formation. Fish Creek Range, Cockalorum 

Wash area, Coral Ridge locality Ml364; Cockalorum 

Wash Formation. Bare Mountain, Nye County, Nev., 

locality M1359. Simpson Park Range, north end at 

Coal Canyon, Horse Creek Valley quadrangle, Nevada; 

0.75 mile south of mouth of Coal Canyon, cobbles in 

conglomerate at base of volcanic sequence.

Study material consists of nine partial coralla.

Hexagonaria meeki n. sp.

Plate 14, figures 1-4

Type material Holotype, USNM 166458. Fish 

Creek Range; Cockalorum Wash area, locality M1352.

Diagnosis. Hexagonaria with thin wall and very 

thin septa having yardarm carinae, many elongate dissepiments, and a narrow tabularium for the genus. 

Long septa withdrawn from axis.

Transverse thin sections. Septal count averaging 

about 28. Septa unusually thin for the genus, slightly 

wavy and withdrawn from axis. Most corallites having 

yardarm carinae; some septa with zigzag carinae passing into yardarm carinae along a septum. Minor septa 

nearly as long as major septa. Concentric pattern of 

dissepiment traces well developed only on inner part 

of band; outer dissepiments highly irregular, larger and 

sublonsdaleioid in some corallites near periphery. In 

places septa discontinuous peripherally, not reaching 

the wall.

Longitudinal thin sections. Tabularium ranging 

from one-fourth to one-third the corallite diameter; 

tabularium comprising many complete or nearly complete tabulae and large flat tabellae; degree of tabellar

sag great for the genus, almost no arching. Dissepiments large for the genus, many being very elongate, in 

three to six columns on each side. Carinal traces well 

shown in depth. Waviness of septa evident in longitudinal section.

Fine structure. No trabecular features shown.

External features. The holotype is a large globular 

head 28 cm in diameter, corallites of which have a relatively small central pit and a broad, nearly flat calice 

platform.

Comparison with related forms. Hexagonaria meeki 

has the yardarm carinae of H. fisherae and its subspecies, but differs decidedly in the thinness and waviness of its septa, the narrowness of its tabularium, and 

the peripheral discontinuity of its septa, together with 

the development of irregular sublonsdaleioid features.

Occurrence. Middle Devonian coral zone F. Fish 

Creek Range; Cockalorum Wash area, Coral Ridge 

locality M1352. Cockalorum Wash Formation. Study 

material consists of five coralla.

Hexagonaria sp. c 

Plate 13, figures 8, 9

Figured material. USNM 166461. Fish Creek 

Range; Cockalorum Wash area, locality M1353.

Diagnosis. Hexagonaria with very wide tabularium 

and abbreviated, noncarinate septa; minor septa well 

developed. Dissepiments all axially inclined in several 

columns.

Transverse thin sections. Septal count about 36. 

Septa smooth, straight, slightly thickened; major septa 

one-fourth to one-third of corallite radius, minor septa 

subequal to major septa. Dissepiment trace pattern 

fairly uniform, concentric-anguloconcentric. Wall somewhat thickened.

Longitudinal thin sections. Tabularium width 

exceeding one-half of corallite diameter, in some threefourths the diameter. Tabulae straight and complete 

or nearly complete with marginal tabellae. Slight axial 

sag more common than arching. Dissepiments mostly 

elongate in three to six columns; all axially inclined.

Comparison with related forms. Hexagonaria sp. c 

differs from sp. n by having fully developed minor septa, 

whereas these are partly aborted in sp. n. Hexagonaria 

sp. r differs in having much longer, wavy septa.

Occurrence. Middle Devonian coral zone F. Fish 

Creek Range; Cockalorum Wash area, Coral Ridge 

locality M1353. Cockalorum Wash Formation. Study 

material is one corallum.

Hexagonaria sp. f

Plate 13, figures 4, 5

Figured material. USNM 166463. Fish Creek 

Range; Cockalorum Wash area, locality Ml353.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 43

Diagnosis. Hexagonaria with broad corallites having long, thin, straight, and wavy septa, some of which 

meet axially. Scattered corallites showing patches of 

septa with zigzag but not yardarm carinae. Peripheral 

dissepiments elongate and flat.

Transverse thin sections. Septal count about 46. 

Minor septa exceeding one-half the length of major 

septa. Stereoplasmic thickenings absent. Zigzag carinae numerous in parts of a few corallites only. Wall 

thin. Dissepiment traces concentric and close-spaced 

in inner part of band, less uniform in peripheral part.

Longitudinal thin sections. Tabularium about onethird corallite diameter. Dissepimentarium with six to 

nine columns of dissepiments on each side, of which the 

peripheral ones are in part very elongate and flat or 

peripherally inclined. Tabularium quite nonuniform 

and comprising combinations of wide, flat tabellae.

Comparison with related forms Hexagonaria sp. f. 

lacks the yardarm carinae of H. sp. w.

Occurrence. Middle Devonian coral zone F. Fish 

Creek Range; Cockalorum Wash area, Coral Ridge 

locality M1353. Cockalorum Wash Formation. Study 

material is one corallum.

Hexagonaria? sp. h

Plate 14, figures 13,14

Figured material. USNM166464. Hot Creek Range, 

Nev.; Morey Peak area, locality M1358.

Diagnosis. Hexagonaria-like coral with long, thickened, noncarinate septa, thickened wall, a wide tabularium, and nearly flat outer dissepiments.

Transverse thin section. Septal count 32. Wall and 

septa stereoplasmically thickened; septa straight, 

tapering smoothly in wedge fashion from wall to tabularium where the major septa are abruptly thinned and 

most continue to axis. Pattern of dissepiment traces 

concentric and fairly uniform. Minor septa about onehalf the length of major septa.

Longitudinal thin section. Tabularium about onehalf the corallite diameter; wide tabulae partly complete and slightly arched. Dissepiments in two to four 

columns on each side, outermost nearly flat or inclined 

axially at very low angles.

Fine structure. Trabecular structure well shown in 

longtitudinal section; trabeculae inclined peripherally.

Remarks. This genus has the thickened wall and 

septa of Xystriphyllum, but differs from that genus by 

having a wide tabularium and flat peripheral dissepiments.

Occurrence. Middle Devonian. Hot Creek Range, 

Nev.; Morey Peak area, locality M1358. Study material 

is one corallum.

Hexagonaria sp. n

Plate 14, figures 8-12

Figured material. USNM 166460. Hot Creek 

Range, Nye County, Nev.; Warm Springs area, locality 

M1354.

This distinctive cerioid coral from the Warm Springs 

area, Hot Creek Range, Nev., forms large heads 30 cm 

or more in diameter, the calices of which are wide and 

deep and lack a peripheral platform. Corallites have a 

very wide tabularium and a narrow dissepimentarium. 

Major septa are short, stereoplasmically thickened 

peripherally; minor septa are commonly aborted, with 

resulting complexity of chevron dissepiment traces. The 

wall is unevenly thickened stereoplasmically.

In longitudinal thin section, this coral reveals a narrow dissepimentarium having two to five columns of 

steeply inclined, partly elongate dissepiments on each 

side. The very wide tabularium comprises partly complete, nearly straight tabulae with very wide flat tabellae, which are neither arched nor sagging.

Hexagonaria sp. r from the Roberts Mountains has 

a similarly wide dissepimentarium and steeply inclined 

dissepiments, but has longer septa and lacks the tendency to abort minor septa.

Occurrence. Middle Devonian coral zone F. Hot 

Creek Range, Warm Springs area, locality M1354. 

Study material consists of three coralla.

Hexagonaria sp. r

Plate 14, figures 6, 7

Figured material. USNM 166462. Roberts Mountains; Red Canyon locality M1339.

Diagnosis. Hexagonaria having wide corallites, a 

very wide tabularium, and wavy, noncarinate, moderately long septa withdrawn from the axis.

Transverse thin section. Septal count from 42 to 

50. Major septa slightly, unevenly thickened, extending 

about two-thirds the distance to axis. Minor septa more 

than one-half the length of major septa. Pattern of dissepiment traces fairly evenly concentric. Wall thickened stereoplasmically.

Longitudinal thin section. Tabularium width exceeds one-half the corallite diameter and in places is 

three-fourths the diameter. Tabulae close-spaced, 

mostly complete, straight to undulant with slight axial 

sag. Dissepiments mostly small, some elongate, nearly 

all axially inclined.

External features. Large globular heads of this form 

with wide corallites lacking a peripheral platform. Calice wide, deep, steep sided, and with a flat bottom.

Comparison with related forms. This coral differs 

from Hexagonaria sp. c by having a higher septal count, 

longer and wavy septa, as well as more closely spaced 

tabulae more of which are complete. Hexagonaria sp. n
44 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

differs by having minor septa commonly aborted and 

has more widely spaced tabulae.

Occurrence. Middle Devonian coral zone F or G. 

Roberts Mountains; Red Canyon, locality M1339. 

Nevada Formation. Study material is one corallum.

Hexagonaria sp. w 

Plate 13, figures 1-3

Figured material. USNM166459. Hot Creek Range; 

Warm Springs area, Nye County, Nev., locality M1354.

This distinctive form with wide corallites has thin 

carinate septa, the longest reaching the axis. The septal 

count ranges from 40 to 44; septa are straight to wavy, 

the carinae partly of zigzag type, but predominantly of 

yardarm type. The tabularium is less than one-third 

the corallite diameter, the dissepimentarium wide, with 

partly small and partly quite elongate, flat dissepiments.

Hexagonaria sp. w. has the thin yardarm carinate 

septa of H. meeki n. sp., but differs by its larger septal 

count, longer septa, wider dissepimentarium and more 

regular arrangement of dissepiment traces and septa 

near the periphery.

Occurrence. Middle Devonian coral zone F. Hot 

Creek Range; Warm Springs area, locality Ml354. 

Study material is one corallum.

Genus Taimyrophyllum Chernychev 1941 

1941. Taimyrophyllum Chernychev, p. 12; pi. 1, figs. 1-3; pi.

2, figs. 1-3; pi. 5, fig. 5.

1958. Taimyrophyllum Chernychev. Bulvanker, pis. 76, 77. 

1961. Aphroidophyllum Lenz, p. 505; pi. 3, figs. 1, 2. 

1964. Taimyrophyllum Chernychev. Pedder, p. 436, pi. 62, 

figs. 12,13; pis. 63-66.

Type species. Taimyrophyllum speciosum Chernychev 1941; by author designation. Devonian, Tareia 

River, Taimyr, U.S.S.R.

Diagnosis. Astraeoid, thamnastraeoid to somewhat 

aphroid rugose corals forming hemispherical or flattened colonies 13 cm and more in greatest diameter. 

Calices with median pit surrounded by a platform in 

some corallites almost flat, reflexed slightly, or protruding as part of a prominent crateriform rim surrounding 

calice pit. Septa on distal surfaces forming prominent 

calicular ridges with entirely radial symmetry. Longest 

septa reaching the axis, where they are twisted. Dissepiment traces concentric to anguloconcentric within 

the inner part of the dissepimentarium, where they are 

usually somewhat crowded; pattern of traces becoming 

irregular and more open peripherally. Tabularium of 

medium width, the close-spaced tabulae having axial 

sag. Some large flat-lying dissepiments in the peripheral 

band of irregular elongate configuration, giving rise to 

aphroid features as seen in transverse section. Septa 

smooth, lacking zigzag carinae, and in some corallites 

having uniform, radially extended stereoplasmic thickenings.

Remarks. Characters which serve to distinguish 

Taimyrophyllum from otherwise similar Billingsastraea 

are prominent aphroid features of Taimyrophyllum, 

together with absence of zigzag carinae, lack of tabular 

arching, and tendency of septa to become radially 

thickened. Billingsastraea, as here interpreted, usually 

has arched, rather than depressed, tabulae and only 

rarely exhibits radially extended thickening of septa. 

Some forms of Billingsastraea have numerous zigzag 

carinae that continue in depth longitudinally.

Taimyrophyllum nolani n. sp.

Plate 9, figures 1-6; plate 10, figures 1-7

Type material. Holotype, USNM 166451; paratypes, USNM 166452, 166453. Lone Mountain localities M1349, M1344; Roberts Mountains, Pyramid Hill, 

locality M1350.

Diagnosis. Taimyrophyllum having septa thickened in tabularium, with crateriform calice rims; axial 

ends of septa considerably twisted. Chevron dissepiment traces relatively few.

Transverse thin sections. Septal count 38 in corallite of 17-mm diameter. In some corallites minor septa 

unrecognizable, in others these septa either much 

shorter or not extending as far into the twisted axial 

complex. Septa straight to slightly wavy and fairly 

smooth in dissepimentarium, becoming twisted in axial 

complex and having minor lateral bumps; septa commonly thickened from inner part of dissepimentarium 

into tabularium; thickenings with fairly even tapered 

radial distribution. Zigzag carinae absent. Aphroid 

lacunae of highly irregular shape and distribution, being 

little developed in some corallites. No trace of a wall.

Longitudinal thin sections. Approximately three to 

eight of the inner and smaller dissepiments very steeply 

inclined or vertical, with abrupt change outward to the 

flat-lying larger dissepiments bordering the corallite. 

Close-spaced, rather delicate tabulae mostly complete, 

unevenly wavy, and with overall sag, almost never 

straight and always without distal arching like those 

of Billingsastraea. Some coralla with roughly horizontal 

layers of stereoplasm connecting corallites. Large flatlying dissepiments 2-5 mm long, these producing the 

big aphroid lacunae visible in transverse section.

Fine structure. Thickened septa revealing weakly 

defined transverse trabecular strands normal to margins. Similar strands observed in fanlike groupings 

within the horizontal stereoplasmic layers between 

corallites.

External features. Discoidal flattened colonies of 

this species with prominent crateriform calice rims surficially resemble Pachyphyllum but have quite different 

internal structure.
LOCALITY REGISTER 45

Comparison with related forms, The type species 

T. speciosum Chernychev from Taimyr, northern 

Siberia, differs from nolani by lacking crateriform margins, by lacking the septal thickening, and by having 

more numerous chevron dissepiment traces. T. carinatum Bulvanker from the Kuznetz Basin, Russia, has 

larger corallites with a greater septal count and better 

differentiated minor septa. T. triadorumPeddei (1964) 

from the Hume Formation of western Canada has 

unthickened septa and a much greater number of chevron dissepiment traces and lacks the crateriform calice 

rims; T. vescibalteatum of the Nahanni Formation, 

also of western Canada, has more numerous, smaller 

aphroid lacunae than the new species and more of the 

chevron dissepiment traces. T. howelli (Lenz, 1961) of 

the Hume Formation is perhaps the most closely related 

species, but differs in its unthickened septa and its 

very large number of aphroid lacunae.

Occurrence. Middle Devonian coral zone F. Lone 

Mountain localities M1344, M1349. Roberts Mountains; Pyramid Hill locality M1350. Unit 4 of the 

Nevada Formation. Study material comprises 14 

nearly complete and partial coralla.

No family assignment 

Genus Lyrielasma Hill 1939

1939. Lyrielasma Hill, p. 243.

1949. Lyrielasma Hill. Stumm, p. 34; pi. 16, figs. 3-5.

1967. Lyrielasma Hill. Pedder, p. 1-29; pis. 1-2.

Type species. Cyathophyllum subcaespitosum 

Chapman, 1925; by author designation. Devonian, Australia.

Diagnosis. The diagnosis given by Pedder (1967, p. 

3) in a recent revision of this genus is as follows: 

Solitary (?) to fasciculate corallum with subcylindrical corallites; budding where observed, peripheral and nonparicidal. 

Peripheral stereozone prominent and in part composed of 

lamellar sclerenchyme; internal lamellar stereozones may also 

be present. Septa well developed, either radially or pinnately 

arranged, strongly carinate in early stages, later becoming less 

carinate or even smooth. Trabeculae parallel and more or less 

horizontal. Dissepiments elongate, steeply inclined and rarely 

lonsdaleioid; rare or absent in early stage; normally several 

deep in adult stages. Tabularium axially depressed, composed 

of predominantly closely spaced and incomplete tabulae. 

Embolophyllum Pedder 1967 differs mainly by having 

a narrower septal stereozone.

Lyrielasma antelopensis n. sp.

Plate 7, figures 9-11

Type material. Holotype, USNM 166445. Antelope 

Range, locality Ml 347.

Diagnosis. Colonial Lyrielasma with well-developed 

septal stereozone, short minor septa, some nearly complete straight tabulae, and a deep calice with narrow 

rim.

Transverse thin sections. Septal count about 56 in 

a corallite of 21-mm diameter. Longest major septa 

approaching the axis; minor septa ranging from stubby 

to one-third the length of major septa. Septa somewhat 

wavy, thinned in tabularium, thickened moderately 

toward periphery. Some major septa with parallel side 

branches in dissepimentarium. Dissepiment trace pattern tending to become complex peripherally, with 

chevrons and herringbone features. Septal stereozone 

about 1 mm wide. Septal grooves rather weakly defined 

in epitheca.

Longitudinal thin sections. Dissepiments elongate, 

steeply inclined in six to 10 columns on each side; 

tabularium one-third to one-fourth of corallite diameter, comprising complete, fairly straight tabulae and 

wide, flat tabellae. No amplexoid septal extensions.

Fine structure. Trabecular structure in septal 

stereozone horizontal to peripherally inclined at low 

angles.

External features. Subcylindrical mature corallites 

of this loosely phaceloid species with subdued and 

rounded rugae and longitudinal (septal) grooves. No 

lateral connective processes observed. Calice about 15 

mm deep, somewhat constricted at the bottom, with 

elongate tapering lip.

Comparison with related forms. Lyrielasma subcaespitosum, the type species, has longer minor septa; 

L. chapmani Pedder has longer minor septa, a wider 

stereozone, and narrower mature tabularium lacking 

the complete straight tabulae.

Occurrence. Middle Devonian coral zone F. Antelope Range, north end, locality M1347. Unit 4 of the 

Nevada Formation. Study material consists of three 

partial coralla.

LOCALITY REGISTER

Lone Mountain, Eureka County, Nev.:

M54. Bartine Ranch quadrangle, about 1,200 ft due south 

of highest peak (VABM 7936) of Lone Mountain. Coralbearing upper dolomitic beds of unit 4 of the Nevada 

Formation.

M1340. Bartine Ranch quadrangle, about 0.5 mile south 

of highest peak (VABM 7936) of Lone Mountain. Loose 

float blocks from coral-bearing upper dolomitic beds in 

unit 4 of the Nevada Formation.

M1341. Bartine Ranch quadrangle. South side of Lone 

Mountain, 3,000 ft southeast of highest peak (VABM 

7936), altitude 7,300 ft; 400 ft west of east boundary of 

quadrangle. Unit 4 of the Nevada Formation with silicified brachiopods and corals.

M1344. Bartine Ranch quadrangle. South side of Lone 

Mountain, 2,500 ft south-southeast of highest peak 

(VABM 7936), altitude 7,440 ft; across ravine 900 ft west 

of M1341. Unit 4 of the Nevada Formation with silicified 

corals.
46 MIDDLE DEVONIAN RUGOSE CORALS, CENTRAL GREAT BASIN

M1360. Bartine Ranch quadrangle. Northwest slope of 

Lone Mountain on crest of spur 0.8 mile N. 34° W. of 

highest peak (VABM 7936), altitude 6,800 ft, near Mineral Monument 6828. Unit 4 of the Nevada Formation 

with silicified corals.

M1361. Bartine Ranch quadrangle. South slope of Lone 

Mountain, 900 ft due south of highest peak (VABM 

7936), altitude 7,600 ft. Upper part of unit 4 of the 

Nevada Formation with silicified corals.

M1363. Bartine Ranch quadrangle. Northwest end of Lone 

Mountain, 2,000 ft S. 72° W. of Lone Mountain zinc 

mine, near top main ridge, altitude 6,600 ft. Upper part 

of unit 4 of the Nevada Formation with silicified colonial 

corals.

M1349. Whistler Mountain quadrangle, west edge. Southeast side Lone Mountain at Reef Point which is on spur 

1,500 ft southeast of summit 7360. North side of spur 

near top, altitude 6,880 ft. Unit 4 of the Nevada Forniar 

tion with silicified colonial corals.

M1362. Whistler Mountain quadrangle, west edge. Southeast end of Lone Mountain, west slope, about 800 ft 

southwest of Flag Point (7360) which is 4,800 ft southeast of top Lone Mountain (VABM 7936). Upper beds 

of unit 4 of the Nevada Formation with silicified fossils. 

Roberts Mountains, Nev.:

M1345. Roberts Creek Mountain quadrangle, southeast 

corner. Pyramid Hill, north end. Northeast of Roberts 

Creek Ranch house, NEVi sec. 19, T. 22 N., R. 51 E., on 

top spur 1,700 ft north-northwest of summit 7504 (Pyramid Hill), altitude 7,000 ft. Unit 4 of the Nevada Formation.

M1348. Roberts Creek Mountain quadrangle, southeast 

corner. Northwest side of Pyramid Hill (7504), NE1/^ 

sec. 19, T. 22 N., R. 51 E. 800 ft northwest of summit 

7504, altitude 7,000 ft. Unit 4 of the Nevada Formation 

with corals.

Ml350. Roberts Creek Mountain quadrangle, southeast 

corner. Pyramid Hill (7504), southwest slope, altitude 

6,750 ft; 3,500 ft northeast of Roberts Creek Ranch house, 

1,700 ft south-southwest of summit 7504. Float corals 

from unit 4 of the Nevada Formation.

M1365. Roberts Creek Mountain quadrangle. East side 

Bobcat Peak (9063), northwest of Roberts Creek Ranch;

I.1 miles N. 78° E. of summit 9063 in saddle just southwest of summit 8466. Unit 4 of the Nevada Formation 

with corals.

M1342. Roberts Creek Mountain quadrangle. East side 

Bobcat Peak (9063), 1,800 ft S. 80° W. of summit 8466, 

near top of east-west spur, altitude 8440 ft. Unit 4 of the 

Nevada Formation with corals.

M1339. Roberts Creek Mountain quadrangle. East side of 

Red Canyon, 1.4 miles S. 18° E. of hill 6897, altitude 

6,840 ft just east of creek bed. Nevada Formation with 

corals. 

Sulphur Spring Range, Nev.:

M1346. Garden Valley quadrangle. East side of range, 

about 0.8 mile northwest of Romano Ranch house in sec.

II, T. 23 N., R. 52 E., east side of butte 6523 (Terrace 

Butte), altitude about 6,200 ft. 

Antelope Range, Eureka County, Nev.:

M30. Bellevue Peak quadrangle, southwest corner. Antelope Range, north end, SEVi sec. 16, T. 16 N., R. 51 E., 

about 0.7 mile north-northwest of Antelope summit 

(7829) on west side summit 7602, altitude about 7,500 ft. 

Unit 4 of the Nevada Formation.

M1347. Cockalorum Wash quadrangle, northwest corner.

Antelope Range, north end. Sec. 21, T. 16 N., R. 51 E.

Antelope summit (7829) just below top on west side.

Unit 4 of the Nevada Formation with abundant corals.

Fish Creek Range:

M1351. Cockalorum Wash quadrangle. Reef Hill 1.5 miles 

south of Cockalorum Wash, 3,500 ft south-southwest of 

Nyeka Peak (7352) at north edge of NE1^ sec. 5, T. 14 

N., R. 52 E., altitude 6,840 ft. Low hill with silicified coral 

deposit.

M1352. Cockalorum Wash quadrangle. Coral Ridge, 1 mile 

N. 70° E. of Willow Creek Ranch house, in NE1/^ sec. 19, 

T. 14 N., R. 52 E.; altitude 6,850 ft, at west base of hills 

1,700 ft north of Willow Creek and 500 ft east of Indian 

Creek. Middle Devonian coral beds with Hexagonaria.

M1353. Cockalorum Wash quadrangle. Coral Ridge; top 

of ridge 1,800 ft east of BM 6928 on north section line 

NEVi sec. 19, T. 14 N., R. 52 E., altitude 7,000 ft. Middle 

Devonian beds with corals and brachiopods.

M1355. Cockalorum Wash quadrangle. Coral Ridge; hills 

north of Willow Creek in SEVi sec. 18, T. 14 N., R. 52 E.; 

1.4 miles N. 59° E. of Willow Creek Ranch house on 

ridge top, altitude 6,960 ft, 800 ft west of section line. 

Coral-bearing limestone of Middle Devonian age.

M13567 Cockalorum Wash quadrangle. Coral Ridge; hills 

north of Willow Creek, 1.1 miles N. 60° E. of Willow 

Creek Ranch house on west side of hills on north boundary sec. 19, T. 14 N., R. 52 E., altitude 6,950 ft. Middle 

Devonian limestone with corals.

M1364. Cockalorum Wash quadrangle. Coral Ridge; 1.2 

miles N. 68° E. of Willow Creek Ranch house, east side 

of ridge 2,300 ft north of Willow Creek, altitude 6,920 ft 

in NEVi sec. 19, T. 14 N., R. 52 E. Middle Devonian 

silicified corals in light-gray limestone. 

Hot Creek Range, Nye County, Nev.:

M1338. Morey Peak 3 quadrangle. North of Morey Peak, 

5,500 feet east of west boundary, 3,500 ft north of south 

boundary T. 10 N., R. 51 E. Middle Devonian beds with 

corals.

Ml354. Warm Springs area. Locality just southwest of 

Warm Springs. Dark-gray limestone with abundant silicified Middle Devonian fossils.

M1358. Morey Peak area. Two miles north of Morey mine, 

4.2 miles N. 87° W. of Moores Station on east side of 

range, altitude 7,400 ft. Middle Devonian limestone with 

corals. 

Bare Mountain, Nye County, Nev.:

M1359. Bare Mountain, southeast of Beatty. Tarantula 

Canyon, about 0.7 mile west of mouth of canyon on south 

side; northeast end of Bare Mountain. Devonian fossils 

in dark-gray carbonaceous limestone. 

Diamond Mountains, White Pine County, Nev.:

M6. Pinto Summit quadrangle. Southwest side of Sentinel 

Mountain at head of southeast branch of Oxyoke Canyon 

in SWV4 sec. 15, T. 18 N., R. 54 E. North of saddle, altitude about 7,900 ft. Tan-colored fine silty dolomite at 

base of Bay State Dolomite Member with corals and 

abundant large silicified Stringocephalus.

M53. Pinto Summit quadrangle. East side of southeast 

branch of Oxyoke Canyon. General collections from 

vicinity of locality M6 in basal tan silty dolomite of Bay 

State Dolomite Member and including uppermost beds 

of Woodpecker Limestone Member.

M204. Pinto Summit quadrangle. West of south end 

Alhambra Hills and southwest of Fair Play mine in
SELECTED REFERENCES 47

NEy4 sec. 36, T. 18 N., R. 54 E. About 500 ft southsouthwest of shaft at altitude 6,289 ft. Dark-gray limestone of Woodpecker Limestone Member with corals and 

large Atrypa.

M1337. Pinto Summit quadrangle. Alhambra Hills, south 

end; SEi/4 SWi/4 sec. 30, T. 18 N., R. 55 E. East side 

Alhambra Hills 0.25 mile southeast of Fair Play mine, 

altitude 6,100 ft. At top of Woodpecker limestone member and base of Bay State Dolomite Member. Dark-gray 

dolomitic limestone beds with corals, Rensselandia and 

Stringocephalus.

S8. Pinto Summit quadrangle. Alhambra Hills, south end. 

Measured section (Aug. 9, 1951) on east slope of hills 

about on north section line in NW1/^ sec. 31, T. 18 N., R. 

55 E., altitude 6,300 ft. Dark-gray limestone with corals 

and brachiopods in upper part of Woodpecker Limestone 

Member. 

Confusion Range, Utah:

M1343. Fish Spring quadrangle. Measured section (by 

R. K. Hose, 1955) in T. 21 S., R. 16 W., 5.25 miles south 

of Highway 6, 13 miles northwest of Ibex. About 175 ft 

below top Simonson dolomite in Hose's unit 30. Corals.

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INDEX

[Italic page numbers indicate both major references and descriptions]

Page

Acanthophyllum 18, 19, 20, 22. 26, 29, 30, 31, 32 

heterophyllum .................................. 20, 30, 31

robertsensis ............ 8, 11, 22, 30, 31, 32; pi. 2

sp. a ...................................... 8, 22, 30, 31; pi. 2

sp. c ...................................... 8, 17, 22, 31; pi. 2

sp. p ..................................... 8, 11, 22, 31; pi. 4

sp .......................... 8, 10, 11, 22, 31; pi. 2; pi. 4

aggregatum, Cystiphyllum ............................ 24

Alaska, correlation ........................................ 19

Alhambra Hills ............................ 17, 24, 27, 30, 33

alhambraensis, Cyathophyllum

(Moravophyllum) ...... 8, 17, 22, 33;

pi. 5 

Digonophyllum (Mochlophyllum) ........ 8, 11,

17, 20, 26, 27 

Moravophyllum ........................................ 34

robertsensis, Digonophyllum

(Mochlophyllum) 8, 11, 22, 27; pi. 3 

Alveolites sp .................................................... 17

Amphipora .............................................. 3, 9, 11, 14

sp ................................................................ 17

Antelope facies belt ........................................ 3, 7

Antelope Range ............................ 11, 38, 41, 45, 46

antelopensis, Hexagonaria fisherae ............ 8, 11,

15, 17, 22, 40, 41; pi. 13 

Lyrielasma .......................... 8, 11, 22, 45; pi. 7

Tabulophyllum ............ 8, 11, 22, 38, 39; pi. 6

Aphroidophyllum ............................................ 44

Apolythophylhf,m ............................................ 38

arachne, Billingsastraea nevadensis .......... 19

Arcophyllum ................................................ 9, 19, 20

dachsbergi ................................................ 20

(Arcophyllum), Mesophyllum ........................ 3, 27

Atelophyllum .................................................... 29

nebracis .................................................... 29

(Atelophyllum), Mesophyllum ...................... 29

Atrypa sp .................................................. 11, 15, 17

Aulacophyllum .................................................. 3, 37

Richardsoni .............................................. 32

Australophyllum .............. 3, 18, 19, 20, 22, 39, 40

cyathophyttoides ...................................... 39

prismatophylloides 8, 10, 20, 22, 39, 40; pi. 8

Bare Mountain ...................................... 3, 17,

Bay State Dolomite Member .... 3, 7, 9, 10,

Beacon Peak Dolomite Member ..................

Bethanyphyllidae ............................................

Bethanyphyllum ..............................................

Billingsastraea .................................. 3, 9, 20,

nevadensis ................................................

arachne ..............................................

verrilli ......................................................

Bluff Canyon ....................................................

Bobcat Peak ................................ 10,23,26,

borealis, Heliophyttum ..................................

Breviphrentia ..................................................

(Breviphrentis), Siphonophrentis ..............

invaginatus, Siphonophrentis ..............

Canada, correlation ..........

capillaria, Pleurotomaria

42,46

15,17

3, 15

21

3,9

21,44

19

19

19,20

15

30,37

36

9,23

3,23

23

18

15

Page 

carinatum, Taimyrophyllum ........................ 45

Caryophyllia gigantea .................................. 23

castanea, Leiorhynchus ............ 11, 17, 18, 19, 20

Ceriophyllum .................................................... 35

chapmani, Lyrielasma .................................. 45

Cladionophyllum ............................................ 21

Classification, fossil ...................................... 22

Coal Canyon .................................................... 42

Cobble Hill ........................................................ 14

Cockalorum Wash .................................... 14, 41, 42

cockalorumensis, Hexagonaria fisherae .... 8, 15,

22, 40, 41; pis. 11, 12 

Columnaria ...................................................... 37, 40

Compressiphyllum .......................................... 23

Confusion Range ............................................ 27, 47

Conglomerate Ridge ...................................... 14

Coral zones ...................................... 3, 7, 10, 11, 22

Correlation ...................................................... 2, 17

Cosmophyllum ................................................ 27

Crurithyris sp ................................................ 17

Cyathophyllidae .............................................. 22, 32

cyathophyttoides, Australophyllum ............ 39

Spongophyllum ........................................ 39

Cyathophyllum ............................ 22, 32, 33, 34, 35

davidsoni .................................................. 39

dianthus .................................................... 33

flexum ........................................................ 34

heterophyllum .......................................... 29

hexagonum .............................................. 39

kobehense .......................................... 28, 29, 36

Richardsoni .............................................. 32

subcaespitosum ...................................... 45

(Cyathophyllum) .................................... 33

(Moravophyllum) .................................. 33

alhambraensis ............ 8, 17, 22, 33; pi. 5

sp. 1 .............................. 8, 10, 22, 34; pi. 4

sp ................................. 8, 10, 15; pis. 5, 6

(Orthocyathus) ...................................... 33, 34

flexum .......................... 8, 10, 22, 34; pi. 7

(Peripaedium) ........................................ 33

turbinatum ...................................... 20, 35

(Cyathophyllum), Cyathophyllum .............. 33

Cylindrophyllum ............................................ 21

Cyrtina sp ...................................................... 17

Cystiphyllidae .................................................. 24

Cystiphylloidae ........................................ 18, 22, 24

Cystiphylloides .................. 3, 18, 21, 22, 24, 25, 28

iddingsi ................................ 8, 10, 22, 25; pi. 1

spinosum .................................................. 19, 24

sp. a .......................................... 8, 17, 22; pi. 1

sp. b ................................................ 8, 11, 24, 29

sp. c .............................. 8, 15, 22, 24, 25; pi. 1

sp ................................................................ 11, 25

(Cystiphylloides), Mesophyllum .................. 24

Cystiphyllum ............................................ 24, 28, 29

aggregatum .............................................. 24

D

dachsbergi, Archophyllum 

davidsoni, Cyathophyllum 

Dechenella ............................

sp ..................................

defectum, Mesophyllum .....

................. 20

................. 39

................. 11

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

................. 28

Dendrostella .................................................... 19, 20

rhenana .................................................... 20

trigemme .................................................. 19, 20

Page 

Descriptive paleontology .............................. 23

Devils Gate Limestone ............................ 3, 10, 18

Devonian Peak ................................................ 10

Diamond Mountains ...................................... 46

Diamond Mountains belt .......................... 3, 15, 18

Diamond Range .......................... 24, 27, 30, 31, 33

Dianthus .......................................................... 33

Cyathophyllum ........................................ 33

Digonophyllidae .......... 3, 18, 19, 20, 21, 22, 24, 25

Digonophyllinae .............................................. 22, 26

Digonophyllum ...................... 19, 20, 22, 26, 27, 28

occidentalis .................................... 11, 26; pi. 3

schulzi ...................................................... 26, 27

sp. b .......................................................... 27

sp. c ................................................ 26, 27; pi. 2

(Digonophyllum) occidentalia ........ 8, 22, 26

sp. c .................................................. 8, 22

(Mochlophyllum) .................................... 27

alhambraensis .......... 8, 11, 17, 20, 26, 27

alhambraensis robertsensis .......... 8, 11,

22, 27; pi. 3 

maximum .......................................... 27

(Digonophyllum) occidentalis,

Digonophyllum .................. 8, 22, 26

sp. c, Digonophyllum ............................ 8, 22

Disphyllidae .................................................... 22, 39

Disphyllum .................................................. 3, 18, 21

lonensis ...................................................... 36

solids ........................................................ 19

sp ................................................................ 8, 17

Diversophyllum .............................................. 38

Dry Creek ........................................................ 10

duplicatum, Zonophyllum ............................ 25, 26

E

Eastern North America, correlation ........ 21, 23

Edaphophyllum .............................................. 21

Eifel district, Germany .... 20, 27, 28, 29, 33, 35,

36,39 

elongatum, Sociophyttum ............................ 20, 37

Spongophyllum ........................................ 36

Endophyllidae .................................................. 22, 38

Eridophyllum .................................................. 21

Eureka Quartzite .......................................... 14

eurekaensis, Sociophyttum .... 8, 10, 11, 19, 20, 22,

36, 37; pi. 8

Utaratuia .................... 8, 19, 22, 37, 38; pi. 8

F

Facies belts ...................................................... S

Faults ................................................................ 7

Favosites .......................................................... 15

sp ........................................................ 11, 15, 17

Fcnstermaker Mountain .............................. 14

Fish Creek Range ........................ 11, 14, 41, 42, 46

fisherae, Hexagonaria 8, 10, 11, 22, 40, 41; pi. 11

Prismatophyttum .................................... 41

antelopensis, Hexagonaria ...... 8, 11, 15, 17,

22, 40, 41; pi. 13

cockalorumensis, Hexagonaria ...... 8, 15, 22,

40, 41; pis. 11, 12 

flexum, Cyathophyllum ................................ 34

Cyathophyllum (Orthocyathus) .... 8, 10, 22

34; pi. 7

Prismatophyttum .................................... 34

51
52 INDEX

Page

G

Germany, correlation .................................... 20

gigantea, Caryophyllia .................................. 23

Siphonophrentis ...................................... 23

Sipkonopkrentis (Siphonopkrentis) .. 23

Givetian, Germany ........................................ 20

glomerulatum, Sociophyttum ...................... 19, 37

Hallidae ............................................................ 3

Hanson Creek Formation ............................ 7

Hedstroemophyttum ...................................... 24

keiligensteini, Heliophyllum ........................ 35

Keriophyllum .................................... 20, 35, 36

Heliophyllum ............................................ 21, 33, 35

borealis ...................................................... 36

keiligensteini ............................................ 35

Heliolites .......................................................... 10

sp .......................................................... 11, 15, 17

Heterophrentis ................................................ 23

heterophyttum, Acanthopkyllum .......... 20, 30, 31

Cyathophyllum ........................................ 29

hexagona, Hexagonaria ................................ 21, 40

Hexagonaria .... 3, 9, 10, 11, 17, 18, 19, 20, 21, 22,

35, 37, 39 

fisherae .................. 8, 10, 11, 22, 40, 41; pi. 11

antelopensis ...... 8, 11, 15, 17. 22, 40, 41;

pi. 13 

cockalorumensis ............ 8, 15, 22, 40, 41;

pis. 11, 12 

hexagona .................................................. 21, 40

kirki .......................................................... 40

meeki ........ 8, 11, 15, 22, 40, 41, 42, 44; pi. 14

rohrensis .................................................. 37

sp. b ...................................................... 8, 10, 40

sp. e .................... 8, 15, 22, 40, 42, 43; pi. 13

sp. f ...................... 8, 15, 22, 40, 42. 43; pi. 13

sp. h .................................. 8, 15, 22, 43; pi. 14

sp. n ............................ 8, 15, 22, 40, 43; pi. 14

sp. r ...................... 8, 11, 22, 40, 42, 43; pi. 14

sp. w ...................... 8, 15, 22, 40, 43, 44; pi. 13

hexagonum, CyathophyUum ........................ 39

Homalophyllum .............................................. 23

Horn Plateau Formation, Canada .............. 19

Hot Creek Range .................. 3, 15, 25, 43, 44, 46

howelli, Taimyrophyttum .............................. 45

Hume Formation, Canada ............................ 18

I, J

iddingsi, Cystiphylloides .......... 8, 10, 22, 25; pi. 1

Introduction .................................................... 1

invaginatus, Siphonophrentia

(Breviphrentis) ...................... 23

Investigation .................................................... 2

Jackass Hill. 10

K

Keriophyllum .................... 20, 21, 22, 33, S5; pi. 4

heiligensteini .................................... 20, 35, 36

kobehense ...................... 8, 10, 22, 29, 36; pi. 4

mclareni ........................ 8, 20, 22, 35, 36; pi. 5

Ketophyllum .................................................... 38

kirki, Hexagonaria ........................................ 40

Wdrrenella .............................................. 11, 18

kirki zone, Martinia ...................................... 10

Knightites (Retispira) sp ......._.................... 15

kobehense, Cyathophyllum .................... 28, 39, 36

Keriophyllum .............. 8, 10, 22, 29, 36; pi. 4

laevigata, Utaratuia ................................ 19, 37, 38

Lecanophyllum ................................................ 29

Leiorhynchus .................................................. 15

castanea ................................ 11, 17, 18, 19, 20

Page 

sp ................................................................ 15

Lekanophyllum ................................................ 22, 29

punctatum ................................................ 29

(Lekanophyllum), Mesophyllum 15, 28, 29; pi. 3 

lissingenense, Mesophyllum .................. 20

nebracis, Mesophyllum .......................... 19

punctatum,, Mesophyllum .............. 19, 20, 29

sp. 1, Mesophyllum ........ 8, 10, 15, 19, 20, 22,

28, 29; pis. 1, 3 

lissingenense, Mesophyllum

(Lekanophyllum) .................... 20

Loipophyllum .................................................. 36

Lone Mountain .... 1, /, 17, 25, 28, 29, 34, 35, 36,

37, 39, 44, 45 

lonensis, Disphyllum ...................................... 36

Loyolophyllum ................................................ 19

Lyrielasma ...................................................... 22, 45

antelopensis ........................ 8, 11, 22, 45; pi. 7

chapmani .................................................... 45

subcaespitosum ...................................... 45

M

mcconnelli, Tabulophyttum .......................... 39

mclareni, Keriophyllum .... 8, 20, 22, 35, 36; pi. 5

Martinia kirki zone ........................................ 10

maximum, Digonophyllum

(Mochlophyllum) .................... 27

Mesophyllum ............................................ 27

meeki, Hexagonaria ........ 8, 11, 15, 22, 40, 41, 42,

44; pi. 14 

Mesophyllum .... 8, 9, 15, 17, 19, 22, 24, 26, 28, 29

defectum .................................................. 28

maximum .................................................. 27

sp. f .......................................................... 10, 19

sp. k .......................................................... 10

sp. 1 ............................................................ 36

(Arcophyllum) ........................................ 3, 27

(Atelophyllum) ........................................ 29

(Cystiphylloides) .................................... 24

(Lekanophyllum) .................. 15, 28, 29; pi. 3

lissingenense ......................._.......... 20

nebracis ............................................ 19

punctatum .................................. 19, 20, 29

sp. 1 8, 10, 15, 19, 20, 22, 28, 29; pis. 1, 3 

(Mesophyllum) ........................................ 28

sp. f .............................. 8, 22, 28, 29; pi. 1

sp. k ............................ 8, 22, 28, 29; pi. 1

(Mesophyttum), Mesophyllum ...................... 28

sp. f, Mesophyllum .......... 8, 22, 28, 29; pi. 1

sp. k, Mesophyllum ............ 8, 22, 28, 29; pi. 1

Microplasma .................................................... 25

Mictophyllum .................................................. 32

richardsoni .............................................. 32

Mochlophyllum, ................................................ 22, 27

(Mochlophyllum), DigonophyUum ............ 27

alhambraensis, Digonophyllum ...... 8, 11, 17,

20, 26, 27

robertsensis, Digonophyllum .. 8, 11, 22,

27; pi. 3

maximum, Digonophyllum .................... 27

Monitor facies belt ........................................ 3

MoravophyUum ........................................ 22, S3, 34

alhambraensis .......................................... 34

ptenophylloidea ........................................ 33, 34

sp. 1 ............................................................ 34

(Moravophyttum), CyathophyUum ............ 33

alhambraenais, Cyathophyttum .... 8, 17, 22,

S3; pi. 5 

sp. 1, Cyathophyllum ........ 8, 10, 22, 34; pi. 4

sp., Cyathophyllum .......... 8, 10, 15; pis. 5, 6

Morey Peak ........................................ 14. 15, 25, 43

N

Nahanni Formation, Canada .................

Nardophyllum .............................................

nebracis, Atelophyllum .............................

Mesophyllum (Lekanophyllum) .....

18

24

29

19

Page 

NeospongophyUum ........................................ 36

(Neospongophyllum), Stringophyllum ...... 36

Nevada Formation ........ 1, 3, 7, 10. 15, 17. 18, 23,

24, 25, 27, 30, 34, 39 

Nevadaphyllum .............................................. 23

nevadense, Tabulophyllum .......................... 30, 31

nevadensis, Billingsastraea .......................... 19

Paracanthus .... 8, 11, 15, 19, 22, S2; pis. 4, 6

Utaratuia .................................................. 10

arachne, Billingsastraea ........................ 19

Ninemile Canyon ............................................ 11

Nolani ................................................................ 45

nolani, Taimyrophyttum ...... 8, 10, 11, 19, 22, 44;

pis. 9, 10 

Nowakia ............................................................ 10, 14

Nyeka Hill ........................................................ 14

O

occidentalis, Digonophyllum ............ 11, 26; pi. 3

Digonophyllum (Digonophyllum) .. 8, 22, 26 

Wdrrenella .......................................... 10, 11, 15

Odontophyttum ................................................ 3

Onondaga Limestone, Canada .................... 24

New York ................................................ 23

Orthocyathus ............................................ 22, 33, 34

(Orthocyathua), Cyathophyllum ................ 33, 34

flexum, Cyathophyllum .... 8, 10, 22, 34; pi. 7

Oxyoke Canyon ........................ 7, 14, 15, 17, 30, 31

Oxyoke Canyon Sandstone Member .......... 14, 15

Pachyphyllum .................................................. 44

Papiliophyllum .............................................. 3, 9, 38

Paracanthus .................................. 19, 22, 32; pi. 6

nevadensis ........ 8, 11, 15, 19, 22, 32; pis. 4, 6

richardsoni .............................................. 32

sp. f ...................................... 8, 10, 22, 32; pL 4

sp ................................................................ 15

peculiaris, Tabellaephyllum ........................ 37

Pedder, A. E. H.. cited ................................ 45

Peripaedium .............................................. 21, 34, 35

(Peripaedium), CyathophyUum .................. 33

turbinatum, Cyathophyttum ................ 20, 35

Phillipsastraea ................................................ 40

Phillipsastraeidae .......................................... 3, 18

Pilot Shale ...................................................... 3, 15

planotabulatum, Sinospongophyttutn ........ 37

PlasmophyUum ........................................ 24, 26, 28

Pleurodictyum ................................................ 3

Pleurotomaria capillaria .............................. 15

Pogonip Group ................................................ 7

Porcellia nais .................................................. 15

sp ................................................................ 15

prismatophylloides, Australophyllum .. 8, 10, 20,

22, 39, 40; pi. 8 

Spongophyllum ........................................ 39

Prismatophyllum ............................................ 39, 40

fisherae ...................................................... 41

flexum, ...................................................... 34

profundum ................................................ 40

profundum, Prismatophyllum .................... 40

Ptenophyllidae .................. 3, 18, 19, 20, 21, 22, 29

ptenophyttoides, Moravophyttum .............. 33, 34

Ptenophyllum .................................................. 29

Ptomatis sp ...................................................... 15

punctatum, Lekanophyttum ........................ 29

Mesophyllum (Lekanophyllum) .... 19, 20, 29

Pyramid Hill ............................................ 10, 31, 44

R

Rabbit Hill Limestone .................................. 3

Receptaculites ................................................ 11

sp ................................................................ 11

rectum, Tabulophyttum, ................................ 38, 39

Red Canyon ...................... 7, 10, 25, 27, 31, 32, 43

redactum, Sociophyttum ................................ 19
INDEX 53

Page 

Reef Hill .......................................................... 14, 15

Rensselandia .................................................... 17, 18

(Retispira) sp., Knightites ........................ 15

rhenana, Dendrostetta .................................. 20

Richardsoni, Aulophyllum ............................ 32

Cyathophyttum ........................................ 32

richardsoni, Mictophyllum .......................... 32

Paracanthus ............................................ 32

Roberts Mountains ...... 10, 23, 25, 26, 27, 30, 31,

32, 36, 37, 43, 44, 46 

Roberts Mountains facies belt .................... 3, 7

Roberts Mountains thrust ............................ 10

robertsensis, Acanthophyttum ........ 8, 11, 22, 30,

31, 32; pi. 2 

Digonophyttum (Mochlophyllum)

alhambraensis .... 8, 11, 22, 27; pi. 3

rohrensis, Hexagonaria ................................ 37

salicis, Disphyllum ........................................ 19

Schichten, Germany ................................ 20, 26, 33

Schizophoria sp .............................................. 11, 17

schulzi, Digonophyttum ................................ 26, 27

sedgwicki, Spongophyttum .......................... 36, 39

semiseptatum, Spongophyllum .................... 36

Sentinel Mountain .......................................... 17

Sentinel Mountain Dolomite Member ........ 9

Simonson Dolomite ........................................ 27

Simpson Park facies belt ............................ 3

Simpson Park Range .................................... 42

Sinospongophyllum ...................................... 37, 38

planotabulatum ...................................... 37

Siphonophrentinae ........................................ 22, 23

Siphonophrentis .............................................. 21, 22

gigantea .................................................... 23

sp. b .......................................................... 11

(Breviphrentis) ...................................... 3, 23

invaginatus ...................................... 23

(Siphonophrentis) .................................. 23

gigantea ............................................ 23

sp. b .................................... 8, 22, 23; pi. 1

(Siphonophrentis), Siphonophrentis ........ 23

gigantea, Siphonophrentis .................. 23

sp. b, Siphonophrentis ............ 8, 22, 23; pi. 1

Sociophyllum .................. 17, 18, 19, 20, 21, 22, 36

elongatum ................................................ 20, 37

Page 

eurekaensis .............. 8, 10, 11, 19, 20, 22, 36,

37; pi. 8 

glomerulatum .......................................... 19, 37

redactum ..................................................' 19

sp .............................................................. 8. 11

(Sociophyllum), Stringophyllum ................ 36

speciosum Taimyrophyllum ........................ 44, 45

spinosum, Cystiphylloides ............................ 19, 24

Spongophyllidae ............................................ 22, 36

Spongophyllum .............................................. 36, 39

cyathophylloides ...................................... 39

elongatum ................................................ 36

prismatophylloides .................................. 39

sedgwicki ................................................ 36, 39

semiseptatum .......................................... 36

Sterictophyttum .............................................. 32

stevensi, Utaratuia ........................................ 38

Stratigraphy .................................................... 7

Streptelasmatidae .......................................... 22, 23

Stringocephalus .... 3, 9, 10, 17, 18, 19, 20, 21, 31

sp. a .......................................................... 17

sp. b .......................................................... 17

Stringophyllum .............................................. 36

(Neospongophyllum) ............................ 36

(Sociophyllum) ........................................ 36

subcaespitosum, Cyathophyttum ................ 45

Lyrielasma ................................................ 45

Sulphur Spring Range .................................. 33, 46

Syringaxon ...................................................... 3

Syringopora sp .............................................. 17

Systematic paleontology .............................. 23

Tabellaephyllum .............................................. 37

peculiar-is .................................................. 37

Tabulophyllum .................................... 18, 21, 22, S8

antelopensis ................ 8, 11, 22, S8, 39; pi. 6

mcconnelli ................................................ 39

nevadense ................................................ 30, 31

rectum ...................................................... 38, 39

Taimyrophyttum ...... 17, 18, 19, 20, 21, 22, 41, 44

carinatum ................................................ 45

howelli ...................................................... 45

nolani .............. 8, 10. 11, 19, 22, 44; pis. 9, 10

speciosum ................................................ 44, 45

triadorum ................................................ 19, 45

Page 

vescibalteatum ........................................ 45

Tarantula Canyon .......................................... 17

Thamnopora .................................................... 11

sp ........................................................ 11, 15, 17

The Ramparts, Canada ................................ 32

triadorum, Taimyrophyllum ........................ 19, 45

trigemme, DendrosteUa ................................ 19, 20

turbinatum, Cyathophyttum

(Peripaedium) ........................ 20, 35

U

Unio .................................................................. 14

Utaratuia ........................ 17, 18, 19, 20, 21, 22, 37

eurekaensis .................. 8, 19, 22, 37, 38; pi. 8

laevigata ............................................ 19, 37, 38

nevadensis ................................................ 10

stevensi .................................................... 38

verritti, Billingsastraea ................................ 19, 20

vescibalteatum, Taimyrophyllum ................ 45

Vinini Formation .......................................... 7, 11

W

Warm Springs ................................................ 15

Warrenella ................................................ 11, 15, 19

kirki .......................................................... 11, 18

occidentalis ........................................ 10, 11, 15

sp ................................................................ 11, 15

Wedekindophyttum ........................................ 24

Willow Creek .................................................. 14

Woodpecker Limestone Member ...... 3, 7, 15, 17,

18, 24, 27, 30

X

Xystriphyllum .......................................... 19, 40, 43

Zaphrentis ...................................................... 33

Zonophyllinae ................................................ 22, 25

Zonophyllum .................................... 3, 9, 22, 25, 26

duplicatum .............................................. 25, 26

sp. r .................................... 8, 11, 22, 25; pi. 1
PLATES 1-14

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

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

FIGURES 1-3. Cystiphylloides sp. a (p. 24).

1, 2. Transverse and longitudinal thin sections ( x 1) of USNM 166417.

3. Enlargement (x 2l/z) of same longitudinal thin section as figure 2. Middle Devonian 

coral zone G; locality M1337, Alhambra Hills. Uppermost beds of Woodpecker Limestone Member. 

4,15. Cystiphylloides iddingsi n. sp. (p.25).

4. Transverse thin section ( x 2) of holotype USNM 166418a. Middle Devonian coral zone

F; locality M1341, Lone Mountain. Upper beds of unit 4 of the Nevada Formation. 

15. Lateral and calice view ( x 1) of paratype USNM 166418b. Same zone as fig. 4; locality M1362, Lone Mountain. 

5, 6. Cystiphylloides sp. c (p. 25).

Transverse and longitudinal thin sections ( x ll/z), USNM 166419.

Middle Devonian, probably coral zone F; locality M1338, Hot Creek Range, Morey Peak

area. 

7, 8. Zonophyllum sp. r (p. 25).

Transverse and longitudinal thin sections ( x ll/2 ), USNM 166420.

Middle Devonian, probably coral zone G; locality M1339, Roberts Mountains at Red

Canyon. 

9-11. Mesophyllum (Mesophyllum)? sp. k (p. 28).

Lateral view (x 1), transverse thin section (x IVk), calice view showing offset (X 1); 

USNM 96213c, specimen figured by Stumm (1938, pi. 59, fig. la) as "Cyathophyllum" 

kobehense.

Middle Devonian, probably coral zone F. Lone Mountain; collected by C. D. Walcott. 

12. Mesophyllum (Lekanophyllum) sp. 1 (p. 29).

Transverse thin section ( x 2), USNM 166421. Middle Devonian coral zone F. Lone Mountain, locality M1340. Nevada formation unit 4. 

13,14. Mesophyllum (Mesophyllum) sp. f (p. 28).

Transverse and longtiudinal thin sections ( x 2), USNM 166422.

Middle Devonian coral zone F. Lone Mountain, locality M1341. Nevada formation unit 4. 

16,17. Siphonophrentis (Siphonophrentis) sp. b (p. 23).

Transverse and longitudinal thin sections (x ll/z), USNM 166423.

Middle Devonian coral zone F. Roberts Mountains, locality M1342, Bobcat Peak.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 1

17 ^^^PW 15 14

»

CYSTIPHYLLOIDES, ZONOPHYLLUM, MESOPHYLLUM, AND SIPHONOPHRENTIS
PLATE 2

FIGURES 1,2. Acanthophyllum robertsensis n. sp. (p. 30).

Transverse and longitudinal thin sections ( x 1%). holotype USNM 166424a. 

Middle Devonian coral zone F. Roberts Mountains, locality M1342, Bobcat Peak. 

3-6. Acanthophyllum sp. a (p. 30).

3. Transverse thin section ( X 2), USNM 166425b. Midddle Devonian coral zone F. Woodpecker Limestone Member; Alhambra Hills, locality M204.

4. 5. Transverse thin section ( X 2), longitudinal thin section ( X 2), USNM 166425a.

Middle Devonian coral zone F. Woodpecker Limestone Member; Alhambra Hills, locality 

S8.

6. Transverse thin section ( x 3l/z), USNM 166425c. Middle Devonian coral zone F. Woodpecker Limestone Member; Oxyoke Canyon, locality M53. 

7, 8. Acanthophyllum sp. c (p. 31).

Transverse and longitudinal thin sections ( x 2), USNM 166426.

Middle Devonian coral zone G. Basal beds of Bay State Dolomite Member; Oxyoke

Canyon, locality M6. 

9,10. Acanthophyllum? sp. (p. 31).

Transverse and longitudinal thin sections ( x 1%), USNM 166427.

Middle Devonian coral zone F or G. Roberts Mountains, Red Canyon, locality M1339. 

11,12. Digonophyllum sp. c (p. 27).

Transverse and longitudinal thin sections ( X 2), USNM 166428.

Middle Devonian, upper part of Simonson Dolomite; Confusion Range, Utah, locality 

M1343.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 2

ACANTHOPHYLLUM AND DIGONOPHYLLUM
PLATE 3

FIGURES 1-5. Digonophyllum (Mochlophyllum) alhambraensis n. sp. (p. 27). 

Thin sections as follows:

Transverse (x 0.9), longitudinal (x 1), transverse (x 3.5), transverse (x 3.5), longitudinal (x 2); holotypeUSNM 166429. 

Middle Devonian coral zone G. Alhambra Hills, locality M1337; topmost beds of the

Wodpecker Limestone Member.

6, 7. Digonophyllum (Mochlophyllum) alhambraensis subsp. robertsensis n. subsp. (p. 27). 

Transverse and longitudinal thin sections (x 1%); USNM 166430. 

Middle Devonian coral zone F or G. Roberts Mountains, Red Canyon, locality M1339. 

8, 9. Mesophyllum (Lekanophyllum) cf. M. (L.) sp. 1 (p. 29).

Transverse and longitudinal thin sections ( x 1%), USNM 166466. 

Middle Devonian coral zone F. Hot Creek Range, Morey Peak area, locality M1338. 

10,11. Digonophyllum occidentalis n. sp. (p. 26).

Transverse and longitudinal thin sections of holotype ( x 1%), USNM 166431. 

Middle Devonian coral zone F. Roberts Mountains, Bobcat Peak, locality M1342.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 3

5 11

DIGONOPHYLLUM AND MESOPHYLLUM
PLATE 4 

FIGURES 1-4. Cyathophyllum (Moravophyllum) sp. 1 (p. 34).

1, 2. Transverse and longitudinal thin sections ( X 2), USNM 166432a. 

3, 4. Transverse and longitudinal thin sections (x 2), USNM 166432b. 

Middle Devonian coral zone F; Lone Mountain, locality M1344. 

5, 6. Keriophyllum^ cf. K.? kobehense (Stumm).

5. Oblique calice view ( x 2), USNM 166467a.

6. Calice view ( x 2), USNM 166467b.

Middle Devonian coral zone F. Lone Mountain, locality M1341.

7. Keriophyllum? kobehense (Stumm) (p. 36)-

Transverse thin section (x 1%) of holotype USNM 96213a.

Middle Devonian, probably unit 4 of the Nevada Formation and coral zone F. Lone Mountain; collected by C. D. Walcott.

8. Acanthophyllum sp.

Longitudinal thin section ( x 2), USNM 166468. 

Same horizon and locality as figs. 1-4. 

9-11. Acanthophyllum sp. p (p. 31).

9. Transverse thin section (x IV2 ), USNM 166433a.

Middle Devonian coral zone F. Roberts Mountains; Pyramid Hill, locality M1345.

10. 11. Transverse ( x 2) and longitudinal thin sections ( x 1%), USNM 166433b.

Middle Devonian coral zone F. Roberts Mountains; Pyramid Hill, locality M1348. 

12,13. Paracanthus? sp. f (p. 32).

Transverse and longitudinal thin sections (x 2), USNM 166436.

Middle Devonian coral zone F. Lone Mountain, locality M1344. 

14 19. Paracanthus nevadensis n. gen., n. sp. (p. 32).

14,15. Transverse and longitudinal thin sections ( x 1%) of holotype, USNM 166434.

16,17. Transverse and longitudinal thin sections (x 1%) of paratype, USNM 166435a.

18. Transverse thin section ( x 1%) of paratype, USNM 166435b.

19. Transverse thin section ( X iy2 ) of paratype, USNM 166435c.

Middle Devonian coral zone F or G. Roberts Mountains; Red Canyon, locality M1339.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 4

ff^ivm » X. ?* **__. f - V* H, f L.*~ > ^--* *-« - * ^jf^f'i"*"'-/, ' ,l;r --'"jt*st^Tv'

11 >-J»v Ma.v^ir>^^^. -^ ^ -^-j»^i.»»« lg

CYATHOPHYLLUM, KERIOPHYLLUM?, ACANTHOPHYLLUM, AND PARACANTHUS
PLATE 5

FIGURES 1 4. Keriophyllum mclareni n. sp. (p. 35).

1, 2. Transverse ( x 3) and longitudinal ( x 2) thin sections of holotype, USNM 166437.

3. Transverse thin section (x 2) of paratype, USNM 166438a.

4. Transverse thin section (x 2) of paratype, USNM 166438b. 

Middle Devonian coral zone F. Lone Mountain, locality M1344. 

5. Cyathophyllum (Moravophyllum) sp.

Transverse thin section ( x 2), USNM 166439. 

Same horizon and locality as figs. 1 4. 

6-10. Cyathophyllum (Moravophyllum) alhambraensis n. sp. (p. 33).

6-8. Transverse and longitudinal thin sections ( x 2) of holotype, USNM 166440. 

Middle Devonian coral zone G. Alhambra Hills, locality M1337.

9,10. Transverse and longitudinal thin sections of paratype ( x 1%), USNM 166441. 

Middle Devonian coral zone G. Sulphur Spring Range, locality M1346.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 5

KERIOPHYLLUM AND CF>1 THOPHYLLUM
PLATE 6

FIGURES 1. Cyathophyllum (Moravophyllum)? sp.

Transverse thin section ( x iy2 ), USNM 166469.

Middle Devonian. Cockalorum Wash area, locality M1356. 

2, 3. Paracanthus cf. P. nevadensis n. sp.

Transverse and longitudinal thin sections (X 2), USNM 166470.

Middle Devonian. Cockalorum Wash area, locality M1353. 

4, 5. Paracanthus nevadensis n. sp.

Transverse and longitudinal thin sections (x 2), USNM 166471.

Same horizon and locality as figs. 2,3. 

6, 7. Paracanthus cf. P. nevadensis n. sp.

Transverse and longitudinal thin sections (x 2), USNM 166472.

Middle Devonian. Cockalorum Wash area, locality M1355. 

8-12. Tabulophyllum antelopensis n. sp. (p. 38).

8, 9. Transverse and longitudinal thin sections ( x 2), holotype, USNM 166442.

10,11. Longitudinal thin sections (x 2), paratype, USNM 166444.

12. Transverse thin section ( x 2), paratype, USNM 166443.

Middle Devonian coral zone F. Northern Antelope Range, locality M1347.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 6

11 9 

CY ATROPHY LLUM, PARACANTHUS, AND TABULOPHYLLUM
PLATE 7 

FIGURES 1-8. Cyathophyllum (Orthocyathus) flexum (Stumm) (p. 34).

1. Calice view of holotype ( x 1), USNM 96219a.

2. Lateral view of paratype with offsets ( x 1V2 ), USNM 96219d.

3. Lateral view of paratype ( x l1/^), USNM96219c.

4. Longitudinal section (x l 1/^). Copy of Stumm (1938, pi. 59, fig. 6c) figure. 

Middle Devonian coral zone F. Lone Mountain; collected by C. D. Walcott.

5. 6. Calice view ( x 1) and transverse thin section ( x 1%), USNM 166473. 

Middle Devonian coral zone F. Lone Mountain, locality M1363; Nevada Formation unit 4. 

7, 8. Transverse and longitudinal thin sections ( x 2), USNM 166474. 

Middle Devonian coral zone F. Lone Mountain, locality M54; Nevada Formation unit 4. 

9-11. Lyrielasma antelopensis n. sp. (p. 45).

Transverse and longitudinal thin sections ( x 2) of holotype, USNM 166445. 

Middle Devonian coral zone F. Northern Antelope Range, locality M1347.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 7

.^mt*i ^P«%& ' -, t :</ ^^j*&&*

10 ' , .- ' 7 

CYA THOPHYLL UM AND L YRIELASMA
PLATE 8 

FIGURES 1-3. Utaratuia eurekaensis n. sp. (p. 37).

Transverse (x 4) and longitudinal thin sections (x 2, x 4) of holotype, USNM 166446. 

Middle Devonian coral zone F; Lone Mountain, locality M1344. 

4, 5. Australophyllum prismatophylloides (Stumm) (p. 39).

Transverse thin sections ( x 2) of holotype, USNM 96217. 

Middle Devonian; Lone Mountain. Collected by C. D. Walcott. 

6-15. Sociophyllum eurekaensis n. sp. (p. 36) -

6, 7. Transverse thin sections (x 2) of paratype, USNM 166449. 

Middle Devonian coral zone F; Lone Mountain, locality M1344. 

8-15. Transverse and longitudinal thin sections ( x 2) of holotype, USNM 166447. 

Middle Devonian coral zone F; Roberts Mountains, Pyramid Hill, locality M1348. 

16-18. Sociophyllum eurekaensis subsp. b. (p. 37).

Longitudinal thin sections ( x 2), USNM 166448.

Middle Devonian coral zone F; Roberts Mountains, Bobcat Peak, locality M1342.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATES

UTARATUIA, AUSTRALOPHYLLUM, AND SOCIOPHYLLUM
PLATE 9

FIGURES 1-6. Taimyrophyllum nolani n. sp. (p. 44).

1. Calice view of paratype ( x 1), USNM 166452.

Middle Devonian coral zone F; Lone Mountain, locality M1349.

2. Calice view of paratype ( x 1%), USNM 166453.

Middle Devonian coral zone F; Lone Mountain locality M1344.

3-5. Transverse thin section ( X 2), longitudinal thin section ( X 2), enlargement of part of

figure 3 ( x 4), holotype, USNM 166451.

Middle Devonian coral zone F; Lone Mountain, locality M1349. 

6. Transverse thin section ( x 8), paratype, USNM 166475. 

Middle Devonian coral zone F; Roberts Mountains, Pyramid Hill, locality M1350.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 9

5 6 

TAIMYR OPHYLL UM
PLATE 10

FIGURES 1-7. Taimyrophyllum nolani n. sp. (p. 44).

1, 2. Transverse and longitudinal thin sections (x 4), paratype, USNM 166476. 

Middle Devonian coral zone F; Lone Mountain, locality M1344.

3. Transverse thin section ( x 8), paratype, USNM 166477. 

Same horizon and locality as figs. 1,2.

4. 5. Longitudinal thin sections (x 8), paratype, USNM 166475.

Middle Devonian coral zone F; Roberts Mountains, Pyramid Hill, locality M1350.

6. Longitudinal thin section (X 8), holotype, USNM 166451. 

Middle Devonian coral zone F; Lone Mountain, locality M1349.

7. Transverse thin section ( X 2), paratype, USNM 166475. 

Same horizon and locality as figs. 4, 5.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 10

7 6

TAIMYR OPHYLL UM
PLATE 11

FIGURES 1-3. Hexagonaria fisherae (Merriam) (p. 40).

1, 2. Transverse thin sections ( X 6), USNM 96218a.

3. Longitudinal thin section (X 6), USNM 96218b. From specimen figured by Stumm

(1938, pi. 59) as Prismatophyllum profundum (Hall). 

Middle Devonian coral zone F; Lone Mountain. Collected by C. D. Walcott. 

4-6. Hexagonaria fisherae subsp. cockalorumensis n. subsp. (p. 41).

Transverse and longitudinal thin sections ( x 4), holotype, USNM 166454. 

Middle Devonian coral zone F. Cockalorum Wash Formation; Cockalorum Wash area, 

locality M1351.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 11

55KJ-3K

d\*jfe "Hi.

6 5
PLATE 12

FIGURES 1-7. Hexagonaria fisherae subsp. cockalorumensis n. subsp. (p. 41).

1. Oblique view of entire corallum ( X Vs), paratype, USNM 166455.

2. Caliceview(x 1%), paratype, USNM 166478.

Middle Devonian coral zone F. Cockalorum Wash Formation; Cockalorum Wash area, 

locality M1351.

3. 4. Transverse and longitudinal thin sections ( X 3), USNM 166479.

Middle Devonian coral zone F. Cockalorum Wash Formation; Cockalorum Wash area,

locality M1352.

5-7. Transverse and longitudinal thin sections ( x 8) of holotype, USNM 166454. 

Same horizon and locality as figs. 1, 2.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 12

HEX AGON ARIA
PLATE 13 

FIGURES 1-3. Hexagonaria sp. w (p. 44).

Transverse (x 4, x 8) and longitudinal ( x 2) thin sections, USNM 166459. 

Middle Devonian coral zone F; Hot Creek Range, Warm Springs, locality M1354. 

4, 5. Hexagonaria sp. f (p. 42).

Transverse and longitudinal thin sections (X 2), USNM 166463.

Middle Devonian coral zone F; Cockalorum Wash Formation, Cockalorum Wash area,

locality M1353. 

6, 7. Hexagonaria fisherae subsp. antelopensis n. subsp. (p. 41).

Transverse and longitudinal thin sections (x 2) of holotype, USNM 166456. 

Middle Devonian coral zone F; Antelope Range, locality M1347. 

8, 9. Hexagonaria sp. c (p: 42).

Transverse and longitudinal thin sections ( X 4), USNM 166461. 

Same horizon and locality as figs. 4, 5.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 13

St-L

8 f

HEXAGON ARIA
PLATE 14 

FIGURES 1-4. Hexagonaria meeki n. sp. (p. 42).

Transverse and longitudinal thin sections ( x ll/z), holotype, USNM 166458.

Middle Devonian coral zone F. Cockalorum Wash Formation; Cockalorum Wash area,

locality M1352. 

5. Hexagonaria sp. cf. H. meeki n. sp.

Transverse thin section ( x 3), USNM 166480.

Middle Devonian; Roberts Mountains, Bobcat Peak, locality M1365. 

6, 7. Hexagonaria sp. r (p. 43).

Transverse and longitudinal thin sections ( x IVz), USNM 166462. 

Middle Devonian coral zone F or G. Roberts Mountains; Red Canyon, locality M1339. 

8-12. Hexagonaria sp. n (p. 43).

Transverse and longitudinal thin sections ( x IVz), USNM 166460. 

Middle Devonian coral zone F. Hot Creek Range, Warm Springs, locality M1354. 

13,14. Hexagonaria? sp. h (p. 43).

Transverse and longitudinal thin sections ( x 4), USNM 166464. 

Middle Devonian. Hot Creek Range; Morey Peak area, locality M1358.

U.S. GOVERNMENT PRINTING OFFICE : 1973 O-506-412
GEOLOGICAL SURVEY PROFESSIONAL PAPER 799 PLATE 14

13 14

HEXAGON ARIA