?

Lower and Lower Middle Devonian 

Rugose Corals of the 

Central Great Basin

GEOLOGICAL SURVEY PROFESSIONAL PAPER 805
Lower and Lower Middle Devonian 

Rugose Corals of the 

Central Great Basin

By C.W. MERRIAM

GEOLOGICAL SURVEY PROFESSIONAL PAPER 805

A stratigraphic-paleontologic study of 

rugose corals as aids in age 

determination and correlation of 

Great Basin Devonian rocks with 

those of other regions

UNITED STATES GOVERNMENT PRINTING OFFICE, WASHINGTON : 1974
UNITED STATES DEPARTMENT OF THE INTERIOR 

ROGERS C. B. MORTON, Secretary

GEOLOGICAL SURVEY 

V. E. McKelvey, Director

Library of Congress catalog-card No. 73-600011

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

Washington, D.C. 20402 - Price $2.85 (paper cover)

Stock Number 2401-02488
CONTENTS

±"age

Abstract ........................................................................................ 1

Introduction ................................................................................ 1

Purpose and scope of investigation.................................. 3

History of investigation...................................................... 4

Methods of investigation.................................................... 5

Acknowledgments .............................................................. 6

Stratigraphy of central Great Basin Devonian rocks........ . 6

Great Basin Devonian f acies belts and coral

reference sections................................................... 6

Antelope-Roberts Mountains facies belt.................. 6

Lone Mountain reference section...................... 8

Nevada Formation, unit 1.......................... 8

Nevada Formation, unit 2.......................... 8

Nevada Formation, unit 3.................. .. 8

Nevada Formation, unit 4.......................... 8

Nevada Formation, unit 5.......................... 9

Devils Gate Limestone................................ 9

Southern Sulphur Spring Range

reference section...................................... 9

Beacon Peak Dolomite Member............... 11

Nevada Formation, unit 1........................... 11

Nevada Formation, unit 2...................... .. 13

Sentinel Mountain Dolomite Member.... 13

Woodpecker Limestone Member.............. 13

Bay State Dolomite Member.................... 13

Devils Gate Limestone................................ 15

Diamond Mountains facies belt................................ 15

Beacon Peak Dolomite Member ...................... 15

Oxyoke Canyon Sandstone Member................ 15

Sentinel Mountain Dolomite Member ............ 15

Woodpecker Limestone Member.................. . 16

Bay State Dolomite Member ............................ 16

Devils Gate Limestone........................................ 17

Monitor-Simpson Park facies belt....................... .. 17

Rabbit Hill Limestone........................................ 17

Coral biostratigraphy of the central

Great Basin Devonian .................................................... 17

Early and early Middle Devonian coral zones.......... . 19

Coral zone A ................................................................ 19

Coral zone B .......................................................... ... 20

Coral zone C ................................................................ 20

Coral zone D ................................................................ 20

Middle Middle Devonian coral zones.............................. 21

Coral zone E ................................................................ 21

Coral zone F ................................................................ 21

Coral zone G ................................................................ 21

Late Middle and Late Devonian coral zones.................. 21

Coral zone H................................................................ 22

Coral zone I.................................................................. 22

Biof acies aspects of Devonian coral distribution

and zonation .................................................................... 22

Syringaxon biofacies .......................................................... 23

Kobeha biofacies ................................................................ 23

Papiliophyllum biofacies .................................................. 24

Breviphrentis biofacies ...................................................... 24

Hexagonaria-Sociophyllum biofacies .............................. 25

Phillipsastraea biofacies .................................................... 25

Page 

Coral succession and evolution as related to geologic

change and faunal migration ........................................ 25

Coral changes at close of Silurian.................................... 27

Late Silurian and Devonian bursts of rugose

coral evolution ......................................................... 28

Zone D coral burst...................................................... 28

Zone F coral burst........................................................ 28

Zone I coral burst........................................................ 29

Age and correlation of Great Basin Early and early

Middle Devonian coral zones........................................ 29

Age and correlation with distant regions. . . . . 29

Coral zone A ..................... ........................ .......... 29

Coral zone B ................................................................ 29

Coral zone C ................................................................ 29

Coral zone D . ...........-...........................'............. 30

Correlation with other areas in the Cordilleran Belt.... 30

North-central Great Basin ........................................ 31

Northern Sulphur Spring Range...................... 31

Southern Tuscarora Mountains........................ 31

Cortez Mountains .............................................. 31

West-central Great Basin.......................................... 31

Toiyabe Range .................................................... 31

South-central Great Basin ........................................ 32

Hot Creek Range ................................................ 32

Southern Great Basin ................................................ 32

Ranger Mountains .............................................. 32

Desert Range ...................................................... 32

Northern Panamint Mountains. .. 32

Funeral Mountains ............................................ 32

Northern Inyo Mountains.......... ..................... 33

Systematic and descriptive paleontology................................ 33

Classification of Great Basin Early and early

Middle Devonian Rugosa .............................................. 33

Taxonomic interpretation of rugose coral structure...... 34

Exterior corallum features ........................................ 34

Taxonomic evaluation of corallum interior

structures ........................................................ 35

Septa .................................................................... 35

Tabulae ................................................................ 36

Dissepiments ...................................................... 36

Stereoplasmic deposits ...................................... 37

Rugose coral symmetry and taxonomy.................... 37

Growth changes and reproductive features............ 37

Variation and dimorphism........................................ 38

Descriptive terms ................................................................ 38

Family Laccophyllidae ...................................................... 38

Genus Syringaxon ...................................................... 39

Family Streptelasmatidae ................................................ 40

Subfamily Siphonophrentinae ........:....................... 41

Genus Siphonophrentis .................................... 41

Subgenus Breviphrentis ............................ 42

Genus Nevadaphyllum ...................................... 44

Family Kodonophyllidae .................................................. 44

Family Stauriidae .............................................................. 45

Genus Dendrostella .................................................... 45

Family Halliidae ................................................................ 46

Subfamily Halliinae .................................................. 46

in
IV CONTENTS

Page

Systematic and descriptive paleontology Continued 

Family Halliidae Continued

Subfamily Halliinae Continued

Genus Aulacophyllum ............................... 46

Genus Odontophyllum ............................... 47

Subfamily Papiliophyllinae .............................. 47

Genus Kobeha ............................................ 47

Genus Papiliophyllum .............................. 50

Genus Eurekaphyllum .............................. 52

Family Bethanyphyllidae ................................................ 53

Genus Bethanyphyllum ............................................ 53

Family Chonophyllidae .................................................... 55

Genus Sinospongophyllum ........................................ 55

Family Endophyllidae ...................................................... 56

Genus Australophyllum ............................................ 57

Family Disphyllidae .......................................................... 58

Genus Disphyllum ...................................................... 58

Page

Systematic and descriptive paleontology Continued 

Family Disphyllidae Continued

Genus Hexagonaria .................................................... 60

Subgenus Pinyonastraea .................................. 61

Genus Billingsastraea ................................................ 62

Family Cystiphylloidae .................................................... 64

Genus Cystiphylloides .............................................. 65

Family Digonophyllidae .................................................. 67

Subfamily Digonophyllinae ...................................... 67

Genus Mesophyllum .......................................... 67

Subgenus Mesophyllum sensu stricto...... 68

Subgenus Arcophyllum ............................ 68

Subfamily Zonophyllinae .......................................... 70

Genus Zonophyllum .......................................... 70

Locality register .................................................................. ..... 71

Selected references .................................................................... 74

Index ............................................................................................. 79

PLATE 1.

2.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

FIGURE 1.

2.

3.

4.

5.

6.

7.

ILLUSTRATIONS

[Plates follow index]

Syringaxon.

Kobeha and siphonophrentoid coral.

Kobeha.

Kobeha.

Kobeha.

Kobeha.

Papiliophyllum.

Papiliophyllum and Eurekaphyllum.

Odontophyllum, Aulacophyllum and Aulacophyllum-lilie rugose corals.

Bethanyphyllum.

Be thanyphyllum.

Bethanyphyllum.

(?) Kodonophyllum, Nevadaphyllum, Nevadaphyllum-like rugose coral, and Bethanyphyllum.

Siphonophrentis (Breviphrentis).

Siphonophrentis (Breviphrentis).

Siphonophrentis (Breviphrentis).

Sinospongophyllum and Siphonophrentis (Breviphrentis).

Dendrostella and Disphyllum.

Cystiphylloides.

Cystiphylloides and Zonophyllum.

Mesophyllum and Mesophyllum (Arcophyllum).

Mesophyllum (Arcophyllum).

Cystiphylloides and Hexagonaria (Pinyonastraea).

Billingsastraea.

Australophyllum, Billingsastraea?, and (?) Hexagonaria.

Page

Index map showing occurrences of Lower and lower Middle Devonian rugose corals in the central and

southern Great Basin.......................................................................................................... ..... ... . .. ........ 2

Map showing distribution of Lower and lower Middle Devonian marine facies belts in the central Great Basin... 7 

Geologic map and cross section of part of Lone Mountain showing lithologic units in the Nevada Formation......... 10

Geologic sketch map showing Silurian and Devonian units in southern Sulphur Spring Range.................................. 12

Geologic cross section showing Devonian stratigraphic units and coral zones in the southern

Sulphur Spring Range ......,........................................-................ .... .. ...................... ......-.--..-....... ...................... 13

Columnar diagram showing relationships of the Devonian reference sections of Lone Mountain and

Sulphur Spring Range to Devonian sections of the Monitor Range......................................................................... 14

Columnar diagram showing relationship of the Devonian reference sections of Lone Mountain and

Sulphur Spring Range to that of the Diamond Mountains facies belt at Oxyoke Canyon................................ 16
CONTENTS V

Page

FIGURE 8. Diagram showing the Great Basin Devonian coral zones, equivalent rock units, coral biofacies, and

previously employed biostratigraphic designations........................................................................................................ 18

9. Characteristic rugose coral genera of central Great Basin Late Silurian, Early Devonian, Middle Devonian, and

Late Devonian coral zones, showing known and inferred vertical ranges ............................................ ........ ... 26

TABLE

TABLE 1. Evolutionary coral bursts in the Great Basin during the Silurian and Devonian Periods.

Page

28
LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS 

OF THE CENTRAL GREAT BASIN

By C. W. MERRIAM

ABSTRACT

Rugose corals of the Great Basin Devonian seaways underwent three major bursts of evolutionary activity: (1) in late 

Early to early Middle Devonian (early Eifelian) time; (2) in 

the medial Middle Devonian (late Eifelian); (3) in Late 

Devonian (Frasnian) time. No Rugosa are known in the 

highest Devonian (Famennian) of this province. The product 

of the first of these bursts of evolution is part of the subject of 

this paper.

Devonian rocks of the central Great Basin occur in three 

north-south facies belts of which the Antelope-Roberts Mountains belt is the more important in terms of Early and Middle 

Devonian coral distribution. No Rugosa were found in Early 

and early Middle Devonian rocks of this age of the Diamond 

Mountains facies belt on the east, where strata of this age are 

barren dolomite and siliceous sandstone. Within the MonitorSimpson Park facies belt on the west, only the earliest Devonian Rabbit Hill Limestone coral fauna is known.

Nine successive coral zones are defined in the Nevada Formation and overlying Devils Gate Limestone of the AntelopeRoberts Mountains belt. These zones have been designated A 

through I in ascending stratigraphic order. Described and illustrated herein are Rugosa and Early Devonian coral zones A, 

B, and C and those of coral zone D, which is Early and early 

Middle Devonian. Rugosa of coral zone A are best known in 

the westerly Monitor-Simpson Park belt, where they characterize the Syringaxon facies of the Helderbergian Rabbit Hill 

Limestone.

The first great burst of Devonian coral activity is that of 

coral zone D, followed by an even greater differentiation of 

certain families in coral zone F, which is marked by spread of 

the Disphyllidae and Digonophyllidae and disappearance of 

Halliidae and Bethanyphyllidae of the older coral zones. Late 

Devonian strata of coral zone I have few surviving rugose coral 

families. However, the newly arrived and very distinctive 

Phillipsastraeidae evolved rapidly and became abundant as 

colonial and solitary forms here as in most of the Late Devonian throughout the world. No Rugosa have been found in the 

latest Devonian of the Great Basin.

Of 13 families of Rugosa in the Great Basin Silurian, 8 do not 

continue on in the Devonian. Eleven Devonian rugose coral 

families are described in the Great Basin province, of which 

only the Laccophyllidae, Streptelasmatidae, and Chonophyllidae carry over from the Silurian as major Devonian groups. 

By early Middle Devonian time five new families have 

appeared: Halliidae, Bethanyphyllidae, Disphyllidae, Cystiphylloidae, and Digonophyllidae. These may have evolved 

from such Silurian stocks as the Lykophyllidae, Kyphophyllidae, and Cystiphyllidae.

Halliidae are especially characteristic of Early Devonian 

beds in coral zones B and C. The small solitary Syringaxon of 

the Laccophyllidae is the only rugose coral in most deposits of 

coral zone A. Locally, however, the compound genus Australophyllum of the Endophyllidae carries over from Late Silurian 

coral zone E to the Early Devonian Rabbit Hill of Devonian 

coral zone A.

Of Early Devonian Halliidae, the endemic Kobeha and 

Papiliophyllum are known only in the Great Basin province. 

Corals of the subfamily Siphonophrentinae of the Streptelasmatidae are abundant in coral zones C and D where they 

paralleled the Onondaga and older members of this subfamily 

in eastern North America. Billingsastraea nevadensis of coral 

zone D is similar to species in somewhat older beds of eastern 

North America. Coral zone D Digonophyllidae of the genera 

Mesophyllum and Zonophyllum are significant ties with the 

Rhine Valley Middle Devonian of western Europe.

INTRODUCTION

Corals are among the common fossils in Great Basin 

Devonian rocks. Of the two extinct Paleozoic Orders, 

Tabulata and Rugosa, the first is more abundantly 

represented, but the Rugosa by reason of their morphologic diversity, more complex internal features, 

and greater evolutionary plasticity prove to be a more 

rewarding subject for stratigraphic paleontology and 

provide better material for taxonomic study and investigation of evolutionary trends through the Devonian 

strata.

Like present-day Zoantharia, the extinct Rugosa 

were beyond doubt highly sensitive to environmental 

vicissitudes, and their geographic-stratigraphic discontinuities within the Great Basin Devonian clearly 

reflect environmental control. Within the Great Basin 

geologic province (fig. 1), the central Great Basin 

Devonian includes a great diversity of marine rock 

types or lithofacies and illustrates better than other 

parts of the province a wide range of coral biofacies. 

For example, the sandstones and dolomites of Early 

and early Middle Devonian age in the Diamond 

Mountains facies belt have yielded no rugose corals. 

To the west, however, in the impure limestones of the 

Antelope-Roberts Mountains facies belt (fig. 2), corals 

become abundant and taxonomically diverse within
LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

120° 118° 116° 114°

\ Hawthorne 

Bridgeport 0 \

\

I M 'I I ! I I I I I 

25 0 25 50 75 100 125 150 175 KILOMETERS

38

36°

34°

FIGURE 1. Index map showing occurrences of Lower and lower Middle Devonian rugose corals in 

the central and southern Great Basin. Symbols indicate Devonian coral zone.
INTRODUCTION

EXPLANATION

LOWER AND LOWER MIDDLE DEVONIAN 

RUGOSE CORAL OCCURRENCES

 Coral zone F, Middle Devonian (Eifelian)

E Coral zone E, Middle Devonian

A Coral zone D, late Early and Middle Devonian

 Coral zone C, late Early Devonian

C Coral zone B, Early Devonian (Oriskany)

A Coral zone A, Early Devonian (Helderberg)

NUMBERED LOCALITIES

1. Lone Mountain map area (see fig. 3)

2. Southern Sulphur Spring Range map area (see fig. 4)

3. Roberts Creek Ranch (M1042)

4. Roberts Mts., Cooper Peak area (M1073)

5. Roberts Mts., Niel Creek area (M1072)

6. Simpson Park Mts., Walti Ranch (M1074)

7. Simpson Park Mts., Coal Canyon (M1075)

8. Cortez Mts., (M1083)

9. Mineral Hill area; McColley Canyon

10. Modoc Peak (M1071)

11. Combs Peak (M27)

12. Grays Canyon (M3; M51)

13. Northern Antelope Range (M1035)

14. Northern Antelope Range (M1053)

15. Fish Creek Range (M1033)

16. Cockalorum Wash area, southeast of Eightmile Well

17. Rabbit Hill (M48; M187)

18. Northern Toquima Range, Petes Canyon area (M1150)

19. Toiyabe Range, west side south of Reeds Canyon (M1151)

20. Dobbin Summit (M1067; M1068; M1069)

21. Hot Creek Canyon (M1066)

22. Northern Panamint Range (M184; M1065)

23.' Funeral Mts. (M1059; M1060; M1061; M1062; M1063; M1064)

24. Ranger Mts. (M1034; M1058)

25. Desert Range (M1057)

Note: Numbers preceded by M are

USGS (Menlo Park) fossil localities

about the same stratigraphic interval. Yet, farther 

west, rugose coral faunas of limestones characterizing 

the Monitor-Simpson Park facies belt are extremely 

restricted taxonomically; most sections contain but a 

single genus of small solitary coral.

In a time-rock sense a great part of the Devonian is 

doubtless recorded by the 3,500-4,000 feet of marine 

strata which make up the system in this province 

(Merriam, 1940). Dolomite and limestone of shallowwater shelf environments predominate. The Devonian 

dolomite, which is mainly of diagenetic origin, especially characterizes the eastern Great Basin; for the 

province as a whole, diagenetic dolomite occupies a 

greater outcrop area than limestone. Where dolomite 

and limestone facies come together in the central Great 

Basin, the vertical and lateral changes are complex; 

within this belt of unstable, fluctuating environment 

the vertical range and geographic spread of coral 

species were greatly affected by facies restrictions.

Corals are more conspicuous on the outcrop and are 

seemingly more numerous in Devonian argillaceous 

and silty limestones than in the cleaner carbonate 

rocks. To some extent this seeming fossil abundance 

may be misleading. The collector soon recognizes that 

fossils weather free and are more easily obtainable on 

the shaly, rubbly erosion surfaces formed by argillaceous limestones than on the denser, less impure, 

albeit fossil-rich carbonate rocks. In the past most 

Great Basin Devonian fossil collecting by geological 

exploring parties was from loose weathered material 

of this nature.

Systematic fossil collecting during the 1930's in 

conjunction with geologic mapping of central Nevada 

Devonian rocks demonstrated the stratigraphic utility 

of rugose corals and was the initial inspiration for 

this project. More recently the Cordilleran Rugosa of 

Silurian and Devonian ages and those of eastern 

North America became the subject of detailed research by several coral specialists. Progress since 1940 

in the light of world advances in the knowledge of 

rugose coral structure, classification, stratigraphic distribution, and ecology makes possible a more meaningful approach to continued study of western Rugosa 

than would earlier have been possible.

PURPOSE AND SCOPE OF INVESTIGATION

The primary objectives of this study are the description, classification, and illustration of Great Basin 

Lower and lower Middle Devonian Rugosa in accordance with modern standards and techniques, together 

with the interpretation of their evolutionary history 

in the light of new knowledge of their stratigraphic 

ranges and geographic occurrences. A proposed zonation based on the corals is the chief product of these 

paleontologic researches; in addition, geologic mapping and structural interpretation of the areas in 

which the Devonian reference sections are situated 

have been accomplished.

Study of the Devonian rugose corals and their ecologic associates has an important bearing upon geologic correlation of the Great Basin strata with strata 

elsewhere in the Cordilleran belt and with standard 

Devonian sections in eastern North America and 

Europe, especially in the Rhine Valley region. These 

studies point up the endemic character of some Great 

Basin coral genera and the more cosmopolitan nature 

of others.

In this report the descriptive and taxonomic paleontology includes only the rugose corals of the Rabbit 

Hill Limestone and the lower half of the Nevada 

Formation; the "barren zone" or coral zone E is an 

appropriate cutoff horizon above which entirely different coral faunas were introduced.
LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

HISTORY OF INVESTIGATION

Study of Great Basin Devonian rugose corals began 

with Meek's (1877, p. 25-33, pi. II) description of 

Upper Devonian species collected in the White Pine 

mining district, Nevada, by the 40th Parallel party. 

Meek was guided in this modest contribution by the 

memoirs of Edwards and Haime (1851; 1850-54), 

whose works set high standards for coral research and 

remained the authoritative references on this subject 

for more than a half century.

Coral-rich Devonian strata, the subject of this report, were discovered by pioneering geologists who 

made the first topographic and geologic map of the 

Eureka mining district, Nevada, in the 1880's (Hague, 

1892). Paleozoic coral research was at that time in a 

static phase, largely because of technical difficulties in 

thin-section preparation. Although large collections of 

loose corals were zealously made by Walcott and his 

field associates, it is not surprising to find that "Paleontology of the Eureka District" (Walcott, 1884, p. 

104-106), published an amazingly short time after 

fieldwork, gave scant attention to the rugose corals. 

None were illustrated.

The Eureka district Devonian coral material deposited in the U.S. National Museum was supplemented considerably during the 1920's by the efforts 

of Edwin Kirk of the U.S. Geological Survey, who 

revisited many of Walcott's localities and discovered 

new localities in outlying areas. At Kirk's suggestion, 

the writer in 1933 began a program of sectioning and 

study of Lower and lower Middle Devonian Rugosa 

collected in the course of a geologic reconnaissance of 

the Roberts Mountains. Comparisons were made with 

the Eureka material and with Great Basin material 

collected by earlier Great Basin exploring parties. 

(The earlier material was also on deposit in the Smithsonian.) At the outset of this program it became evident that the older Eureka district collections had 

limited stratigraphic value, because of the manner in 

which collecting and reconnaissance geology were 

done in the 1880's. Some of the collections rather 

certainly represent fossils taken from 100 feet to 

several hundreds of feet of strata, but they are included 

in a single lot; hence, cautious evaluation and sorting is required for purposes of detailed stratigraphy 

and ecologic association by present standards.1

Systematic thin-section study of American

to Edwin Kirk (oral commun., 1935; written commun., 1954), 

Walcott had described to him the manner in which some of the Devonian 

fossil collecting was done at Lone Mountain and at the Eureka vicinity. Several U.S. Naval Academy midshipmen attached to the Hague party for summer engineering experience were assigned to fossil collecting under Walcott's 

supervision. Large piles of corals and other fossils, found at central locations 

on the outcrop about 1928, were believed by Kirk to be unretrieved caches of 

the Annapolis trainees.

ian corals was little advanced at the inception of this 

investigation in 1933. Few published descriptions were 

accompanied by adequate illustration of internal 

corallum structure, thus preventing meaningful comparison. Progress in Europe was more rapid. A milestone in Paleozoic coral research was the burst of 

activity at Marburg, Germany, by Rudolf Wedekind 

and his colleagues during the 1920-36 period. Detailed descriptions and thin-section illustrations of 

Rhine Valley Devonian Rugosa by these workers have 

had special value in connection with identification and 

classification of Great Basin and other American farwestern forms. Although the coral taxa of Wedekind 

and his associates are keyed to stratigraphic units and 

have value as zonal indices, their excessive generic 

splitting is a controversial feature. Doubtless many 

Wedekind genera will eventually be reduced in rank 

or combined in synonymy by more conservative 

workers (Birenheide, 1962a; 1964).

Stumm's (1937; 1938; 1940) descriptive work on 

Devonian rugose corals of the Eureka district and 

adjacent areas dealt mainly with the Smithsonian collections mentioned above. These contributions placed 

on record previously unknown Early and Middle 

Devonian genera and provided a foundation for continued taxonomic work upon these partly endemic 

Great Basin forms.

Since 1940 general progress in the understanding of 

the Rugosa has been great, with a rapidly expanding 

world literature, some of which is referred to in the 

section "Systematic and Descriptive Paleontology." 

Among indispensable compendia are:

(1) "Index of Paleozoic Coral Genera" by Lang, 

Smith, and Thomas (1940).

(2) "Revisions of the Families and Genera of the 

Devonian Tetracorals," by Stumm (1949).

(3) "Rugosa," by Hill (1956) in "Treatise on 

Invertebrate Paleontology."

To a considerable extent these Great Basin Silurian 

and Devonian coral studies were a product of geologic 

mapping and stratigraphy which were stimulated by 

economic and resource factors. Search for petroleum 

and metals in the Great Basin and successful oil exploration in Devonian rocks of western Canada within the 

past 25 years have encouraged study of all aspects of 

the stratigraphy and paleontology. For the first time 

meaningful comparison of Great Basin faunas with 

those of the northern Cordilleran Belt has become 

possible. Thus far close paleontologic similarities are 

evident mainly among higher Middle and Upper 

Devonian species, rather than among fossils of the 

lower zones here dealt with.

Most of the corals described and figured herein
INTRODUCTION

were collected after 1947, during the course of geologic mapping and stratigraphic investigation in the 

Eureka mining district, Nevada, and adjacent areas 

by the writer in conjunction with the detailed geologic mapping program of T. B. Nolan. Central Great 

Basin Devonian collections were supplemented from 

time to time by coral material collected by U.S. Geological Survey mapping parties in other parts of the 

province. In 1959 the mapping and stratigraphic 

effort involving Silurian and Devonian rocks of six 

15-minute quadrangles west of the Eureka mining district was revitalized under the Kobeh Valley project 

of Nolan and Merriam.

METHODS OF INVESTIGATION

Understanding the fossil distribution in the greatly 

dislocated strata of the Great Basin Paleozoic requires 

fairly detailed geologic mapping in order to set up 

reliable reference sections. At best the standard 

paleontologic column is a composite of stratigraphic 

segments in disconnected structural blocks. Further 

complexities are introduced by profound facies 

changes, as for example the almost complete disappearance of siliceous sandstone in the Lower Devonian between the Diamond Mountains and Lone 

Mountain. Dependable key beds with great lateral 

continuity are uncommon in the Devonian rocks of 

this vicinity. Attempts have been made to relate fossil 

occurrences to structure sections prepared in the 

advanced stages of the mapping procedure.

Stratigraphic data applicable to Devonian rugose 

coral zonation have accrued from geologic mapping 

since 1959 in the Whistler Mountain, Bartine Ranch, 

and parts of the Garden Valley and Cockalorum Wash 

quadrangles. Much light is shed upon the profound 

Lower Devonian facies changes by the mapping of 

T. B. Nolan near Eureka, especially in the southernmost Diamond Mountains in the Pinto Summit quadrangle.

The more continuous measurable Devonian sections 

within this region of thrust faulting occupy carbonate autochthons. Like the central mass of Lone 

Mountain (fig. 3), these more rigid strata are little 

deformed by thrusting, but they are cut by innumerable high-angle faults which break stratigraphic continuity. In this connection Lone Mountain, including 

the primary Devonian coral reference section, has 

been separately mapped.

Lone Mountain provides an excellent example of 

highly complex Great Basin geologic structure wherein stratigraphic sections unaffected by pervasive thrust 

and normal faulting are indeed exceptional. The central part of the mountain is a fairly straightforward

homocline of marine beds, mostly carbonate, ranging 

in age from early Middle Ordovician to late Middle 

Devonian. However, the lower or pediment slopes 

reveal an entirely different and highly complex pattern 

of thrusting which involves Ordovician graptolitic 

beds, Permian limestones, and Devonian limestones. 

Lone Mountain discloses to advantage the characteristic structure pattern of the region thrusting 

followed by high-angle, mostly normal, faulting.

To the south beyond the limits of this study in the 

Fish Creek Range occur Middle Devonian rocks of 

coral zone F, lithologically quite different from the 

normal autochthonous carbonates of this interval; 

these highly deformed southern partly argillaceous 

facies occupy what appear to be major thrust plates.

Coral collecting by the writer and associates during 

the initial phase of this study (Merriam, 1940) was 

in considerable part from loose weathered material, 

precise stratigraphic horizon not being rigorously 

determined. In later years more of the collecting was 

done bed-by-bed on measured sections to secure in 

situ associations, a procedure greatly facilitated by 

new and accurate topographic maps. Whereas sections 

were measured by tape and Brunton for collecting 

purposes within the mapped areas, large-scale map 

location of collecting sites was found to be more reliable because of pervasive faulting, which was difficult 

to recognize or interpret where stratigraphic observations are confined to individual traverse courses.

Little progress has been made in biofacies study of 

the Great Basin rugose corals and their ecologic associates. Whereas Rugosa are dominant megafossils in 

some lithologic units, in others they are numerically 

less important elements of varied assemblages containing corals, brachiopods, mollusks, and trilobites. 

A good biofacies indicator is Syringaxon of the Early 

Devonian Rabbit Hill Limestone, in which this abundant small solitary genus is normally the only rugose 

coral. By contrast coral zone F of the higher Middle 

Devonian contains locally abundant large compound 

genera in a potentially reef-forming biota.

With regard to relative age and correlation values 

of the several phyla, it may be said that specialist 

conclusions based upon independent study of separate fossil categories have not always agreed with 

those founded upon rugose coral research. Converging 

and adjusted conclusions from research upon all the 

major fossil categories are needed. Some corals are 

perhaps better indicators of specific environments 

than precise geologic age. In this work the associated 

brachiopods have been utilized largely to support the 

coral evidence for correlation and age.
LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

ACKNOWLEDGMENTS

Among Great Basin workers whose rugose coral 

collections have added materially to this study are 

the following: J. F. McAUister, Panamint Range and 

Funeral Mountains; M. S. Johnson and D. E. Hibbard, Ranger Mountains; F. G. Poole, Ranger Mountains; C. R. Longwell, Desert Range; F. J. Kleinhampl, 

Monitor Range; H. E. Cook, Hot Creek Range; R. H. 

Washburn, Toiyabe Range; J. H. Stewart and E. H. 

McKee, Toiyabe and Toquima Ranges; T. B. Nolan, 

Eureka mining district; and A. J. Boucot and J. G. 

Johnson, northern Sulphur Spring Range and Cortez 

Mountains.

An earlier draft of this report was read by the late 

E. C. Stumm of the University of Michigan, whose 

constructive suggestions regarding taxonomic matters 

are appreciated. D. J. McLaren of the Geological 

Survey of Canada has been most helpful in providing 

information on the type species of Cystiphylloides and 

other genera of Devonian rugose corals.

All photographic illustrations are the work of Kenji 

Sakamoto of the U.S. Geological Survey, Menlo Park, 

Calif.

STRATIGRAPHY OF CENTRAL 

GREAT BASIN DEVONIAN ROCKS

Coral-bearing Devonian formations of the central 

Great Basin are in ascending stratigraphic order as 

follows: (1) Rabbit Hill Limestone of Early Devonian 

(Helderberg) age; (2) Nevada Formation of Early and 

Middle Devonian age; and (3) Devils Gate Limestone 

of late Middle and Late Devonian age. Physical characteristics of these formations have been described 

in other reports (Merriam, 1940, 1963; Nolan and 

others, 1956) together with details of their areal distribution and geologic structure. The Lone Mountain 

and Sulphur Spring Range reference columns are 

adopted as joint primary yardsticks for these rocks, 

supplemented by partial sections in the Devils Gate 

area and in the northern Monitor Range.

With reference to Early and early Middle Devonian 

coral-bearing strata dealt with in this report, approximately correlative noncoralline rocks in the Diamond 

Mountains to the east comprise entirely different 

dolomite and sandstone facies. To the west in the 

Monitor and Simpson Park Ranges, the Early Devonian Rabbit Hill Limestone represents yet another 

facies belt.

In the Sulphur Spring Range the Early and early 

Middle Devonian beds most closely resemble, both 

lithologically and faunally, those of the Lone Mountain yardstick, but fortunately include westerlyreaching quartzitic sandstone tongues from the 

Diamond Mountains belt as well as easterly extending

beds which carry elements of the Rabbit Hill Helderbergian fauna of the Monitor-Simpson Park belt. 

Hence in this key reference column of the Sulphur 

Spring Range, lithologic and paleontologic features of 

all these facies belts are rather satisfactorily united. 

Most of the stratigraphic units and coral zones 

treated in this report occur within the Nevada Formation of Early and Middle Devonian age, as earlier 

redefined by Nolan, Merriam, and Williams (1956, p. 

40-48). On the basis of lithology and fossils, this 

formation has been subdivided vertically and laterally 

in considerable detail. Its more characteristic facies 

lie within the Antelope-Roberts Mountains belt, but 

the easterly Diamond Mountains dolomite-arenaceous 

facies are also included in the Nevada Formation. The 

westerly Helderbergian Rabbit Hill Limestone is geographically separate and, except for its presence in 

th northernmost Simpson Park Mountains, is not 

known to be present in sections containing the Nevada 

Formation. Beds with Rabbit Hill fossils earlier mentioned in the Sulphur Spring Range are more closely 

allied lithologically to the Beacon Peak Dolomite 

Member of the Nevada Formation as defined within 

the Diamond Mountains facies belt.

GREAT BASIN DEVONIAN FACIES BELTS 

AND CORAL REFERENCE SECTIONS

For purposes of this discussion, Devonian strata of 

the central Great Basin may be treated in accordance 

with its distribution in three subparallel facies belts 

(fig. 2) as follows:

1. Antelope-Roberts Mountains facies belt,

2. Diamond Mountains facies belt,

3. Monitor-Simpson Park facies belt.

Each facies belt possesses distinctive lithologic, structural, and paleontologic characteristics. Whereas 

north-south subparallel alinement of the belts is 

today geomorphic-orographic, it may to some extent 

reflect the predeformation distribution pattern of sedimentation in shelf seaways of Devonian time.

ANTELOPE-ROBERTS MOUNTAINS FACIES BELT

The Antelope-Roberts Mountains belt stretches 

south for about 90 miles from the Sulphur Spring 

Range and Roberts Mountains, through the Fish 

Creek Range and the Antelope Range. The strata of 

this belt probably extend southward in the littleexplored Hot Creek Range. In a time-rock sense nearly 

all the Devonian System as known in the Great Basin 

is represented within the Antelope-Roberts Mountains 

facies belt. A possible exception is the latest Devonian 

of the upper Pilot Shale. Characteristic are the fossilrich silty and argillaceous limestones of Nevada
STRATIGRAPHY OF CENTRAL GREAT BASIN DEVONIAN ROCKS

117° 116°

EXPLANATION

Diamond Mountains facies belt

Dolomite and siliceous sandstone

Antelope-Roberts Mountains facies belt 

Argillaceous limestone with subo 

dolomitic limestone and calcarenite; sniceou 

sandstone interbeds on east side, 

abundant

Monitor-Simpson Park facies belt 

Limestone and calcareous shale; no dolom 

Syringaxon coral facies in Rabbit Hill 

Limestone

LU

Whistler Mtn 

Lone Mtn

Devils Gate

MAHOGANY / .

39° -

FIGURE 2. Distribution of Lower and lower Middle Devonian marine facies belts in the central Great Basin.
8 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Formation, unit 2, which have not been recognized in 

the other belts. Present along its eastern margin are 

quartzitic sandstone and dolomite tongues interpreted 

as westerly facies extensions from the Diamond Mountains belt. Situated within this belt are the primary 

coral reference sections for the Lower and lower Middle 

Devonian at Lone Mountain and in the southern 

Sulphur Spring Range.

LONE MOUNTAIN REFERENCE SECTION

Five lithologic units have been used in the detailed 

geologic mapping of the Nevada Formation at Lone 

Mountain (fig. 3), each characterized by distinctive 

rugose corals and other fossils. The stratigraphy and 

paleontology are better understood here than at 

Modoc Peak, where the Nevada Formation type section was earlier designated (Merriam, 1940, p. 15). 

However, the Modoc section is in other ways somewhat more representative of the formation as a whole, 

for it includes a quartzitic sandstone member of the 

easterly Diamond Mountains facies, a tongue which 

feathers out completely before reaching Lone Mountain. The map divisions of the Nevada Formation at 

Lone Mountain are designated units 1 through 5 in 

ascending order.

The basal dolomite of unit 1 rests with a seemingly 

gradational boundary upon the upper beds of the 

Lone Mountain Dolomite. These lower Nevada dolomites contain the Papiliophyllum fauna of coral zone 

C, and the Acrospirifer kobehana fauna of about 

Emsian age. No depositional record of the Rabbit Hill 

Limestone or of the Beacon Peak Dolomite Member, 

each with a Helderbergian fauna and coral zone A, 

has been recognized here at Lone Mountain where 

they would be expected just above the Lone Mountain 

Dolomite. Likewise, coral zone B with Kobeha is not 

represented by conclusively identifiable fossils. No 

physical evidence of disconformity was found here 

at the Lone Mountain Dolomite-Nevada Formation

boundary.

NEVADA FORMATION, UNIT 1

The lower unit of the Nevada Formation (fig. 3) is 

about 260 feet thick in Coral Gulch on the south side 

of Lone Mountain, where it consists of thick-bedded 

blocky dolomite, dolomitic limestone, and limestone 

of light-gray to mouse-gray color. Lower beds of unit 

1 have much the same appearance as the underlying 

Lone Mountain Dolomite, with which it appears to be 

gradational at this locality. A disconformity separates 

these two formations in the Mahogany Hills and in 

the Diamond Mountains belt. Argillaceous matter is 

present in the upper beds of this unit where it grades 

into unit 2.

Unit 1 becomes less dolomitic traced northwest

toward Contact Gulch. Gray chert nodules flattened 

parallel to the bedding are characteristic, and the 

fossils are generally silicified. Chert and fossils weather 

limonite brown.

The lowest identifiable fossils collected from unit 1 

at Lone Mountain represent coral zone C. Poorly preserved large horn corals at the base may be in coral 

zone B. In the Sulphur Spring Range calcarenite with 

scattered rounded quartz grains occurs at the bottom 

of unit 1 and contains the Kobeha fauna of coral zone 

B with Costispirifer arenosus.

NEVADA FORMATION, UNIT 2

About 720 feet thick, unit 2 is well exposed in upper 

Coral Gulch and upper Fossil Gulch. Because of its 

partly argillaceous, thin-bedded, and banded character, the appearance of unit 2 contrasts sharply in 

outcrop with underlying and overlying thicker bedded 

units, being more resistant to weathering. The shaly 

bands yield loose fossils in large numbers; most of the 

collections first made from the Nevada Formation are 

from outcrops of this nature. Corals, especially favositid heads, are abundant in many beds; some are in 

their original upright growth position with their base 

in a limestone layer, the upper part not uncommonly 

having been buried by argillaceous sediment. Unit 2 

contains the assemblages of coral zone D in which the 

Rugosa reach a peak of abundance and diversity.

No siliceous sandstone was found in unit 2 at Lone 

Mountain, but it was found at Combs Peak and Modoc 

Peak. The eastward increase of siliceous matter, well 

shown in the southern Sulphur Spring Range, reaches 

a maximum in the Diamond Mountains southeast of 

Eureka where the Oxyoke Canyon Sandstone Member 

probably correlates with some part of unit 2.

NEVADA FORMATION, UNIT 3

About 285 feet thick, the dark-gray, well-bedded, 

and relatively nonargillaceous limestone of unit 3 is 

well exposed on the south side of Lone Mountain, 

2,000 feet northeast of West Peak. Many beds and 

lenses of crinoidal debris in it tend to be massive; 

otherwise fossils are scarce in unit 3. Unit 3 has been 

called the "barren zone" compared with the abundantly fossiliferous units below and above. In the 

Diamond Mountains at Oxyoke Canyon, the Sentinel 

Mountain Dolomite Member, has yielded no fossils. 

It is probably a facies equivalent of unit 3, but it is 

completely dolomitized. No dolomite was recognized 

in unit 3 at Lone Mountain.

NEVADA FORMATION, UNIT 4

A well-bedded division, unit 4, is about 675 feet 

thick at Lone Mountain and crops out along the west
STRATIGRAPHY OF CENTRAL GREAT BASIN DEVONIAN ROCKS

side of the main ridge between Reef Point and North 

Peak. Unit 4 includes dense, resistant, fine-grained 

limestone, dolomitic limestone, and dolomite. The unit 

ranges in color from dull mouse gray to medium and 

dark gray, and weathers light gray. These strata 

include thin-bedded almost shaly intervals and thickbedded massive dolomite lenses. Rugose corals and 

stromatoporoids are abundant locally within the thickbedded dolomitic bodies. Chert nodules weathering 

limonite brown are fairly numerous, and fossils are 

commonly silicified. Unit 4, with numerous colonial 

and solitary rugose coral genera, represents Devonian 

coral zone F, the interval of greatest differentiation of 

the colonial Rugosa in Middle Devonian time.

In the Fish Creek Range south of Eureka, Middle 

Devonian beds correlative with the Nevada Formation, unit 4 include black shale and siltstone with 

light-gray limestone bodies locally rich in rugose 

corals like those of unit 4 at Lone Mountain. In both 

places these loci of coralline proliferation suggest 

incipient bioherms or patch reefs.

The Woodpecker Limestone Member of Nolan, 

Merriam, and Williams (1956, p. 44) is the equivalent 

of unit 4 in the Diamond Mountains, where it includes 

more platy and shaly limestone, some of which is 

pinkish in color. The thick-bedded dolomitic bodies 

with a coral biota were not found in the Diamond 

Mountains f acies belt.

NEVADA FORMATION, UNIT 5

Unit 5, the uppermost unit of the Nevada Formation, is about 570 feet thick at Lone Mountain and 

consists of thick-bedded and massive saccharoidal 

dolomite and dolomitic limestone of dark-, medium-, 

and light-gray color. Alternation of very dark gray 

and light-gray thick dolomite beds is characteristic. 

This resistant unit forms the summit ridge of Lone 

Mountain between Flag Point and North Peak. It is 

overlain by massive cliff-forming Devils Gate Limestone, the lowest beds of which are crowded with 

Amphipora, as is well shown on the first main spur 

east of Section Ridge (fig. 3).

Unit 5 is sparsely fossiliferous at Lone Mountain. 

There are scattered deposits of the large brachiopod 

Stringocephalus. However, the organic structure of 

brachiopods and stromatoporoids is largely destroyed 

by dolomitization. The Bay State Dolomite Member 

of the Diamond Mountains belt is the approximate 

equivalent of the Nevada Formation, unit 5. Thick 

beds with Stringocephalus are best shown in that belt. 

Limestone beds near the base of the Bay State Dolomite Member contain rugose corals of coral zone G 

and the brachiopod Rensselandia.

DEVILS GATE LIMESTONE

The upper Middle and Upper Devonian Devils Gate 

Limestone (Merriam, 1940, p. 16-17; 1963, p. 49-56) 

is about 1,200 feet thick and rests conformably upon 

the dolomites of Nevada Formation, unit 5 at Lone 

Mountain. It consists mainly of limestone in most 

localities; locally the lower part has been subjected to 

patchy, irregular dolomitization (Merriam, 1963, p.

51). At Lone Mountain the Devils Gate is the highest 

unit of the central autochthon and consists of mediumdark to dark-gray well-bedded limestone with scattered massive, cliff-forming bodies. The topmost strata 

are not exposed here; these abundantly fossiliferous 

beds are best seen at Devils Gate or in the Newark 

Mountain vicinity of the Diamond Mountains, according to Nolan, Merriam, and Williams (1956, p. 48-

52), where both top and bottom are well exposed.

Clear-cut lithologic subunits are more difficult to 

define and trace within the Devils Gate Limestone 

than in the Nevada Formation because of greater rock 

uniformity in the former. However, the upper 250-300 

feet are quite readily separable by somewhat thinner 

bedding, the presence of dark-gray chert, large lenses 

and beds of depositional mud-breccia, and the great 

local abundance of shell fossils. The lower half of the 

Devils Gate everywhere includes a great deal of limestone formed by massive concentric stromatoporoids 

and Amphipora as at Lone Mountain, but in general 

corals and shell fossils are abundant only in widely 

scattered pockets. It has recently been found that the 

large brachiopod Stringocephalus, thought previously 

to be confined to the Nevada Formation, unit 5 (Bay 

State Dolomite Member), ranges up into thick-bedded 

massive limestones which characterize the lower third 

of the Devils Gate at Lone Mountain.

SOUTHERN SULPHUR SPRING RANGE REFERENCE SECTION

Devonian rocks of the Nevada Formation underlie 

most of the southern Sulphur Spring Range east of 

the Mulligan Gap fault (fig. 4). The five Nevada 

Formation units or their time-stratigraphic equivalents described at Lone Mountain are present, but 

the Devonian column is more inclusive in the Sulphur 

Spring Range; stratigraphically beneath the Nevada 

Formation, unit 1 are dolomites and dolomitic limestones containing Helderbergian Rabbit Hill fossils. 

These earliest Devonian beds (for this region) are 

not found at Lone Mountain and are assigned to the 

Beacon Peak Dolomite Member. In the Diamond 

Mountains facies belt where the Beacon Peak is 

described, it is the lowermost member of the Nevada 

Formation. The overall lithologic makeup of the reference section of the Sulphur Spring Range Devonian
10 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

116°15'

39°35'

Base from U.S. Geological Survey, 1.62 500 1QQO 500 0 

Whistler Mountain and Bartme Ranch, 1956 I____.____I

1000 2000

I I

3000 

I

4000 5000 FEET Geology by C. W. Mernam, 1959-66 

I________I

1000 METERS

Lone Mountain 

Dolomite

FIGURE 3. Part of Lone Mountain showingiithologic units in the Nevada Formation of this reference section. Geologic cross 

section A-A', of Lone Mountain reference section. Letters indicate Devonian coral zones.
STRATIGRAPHY OF CENTRAL GREAT BASIN DEVONIAN ROCKS 11

EXPLANATION

Devils Gate Limestone

Unit 5 

(Bay State Dolomite Member)

Undifferentiated Middle 

Devonian dolomite

Unit3

Roberts Mountains Formation (dolomitized)

Hanson Creek Formation (dolomitized)

WESTERN FAC1ES 

PALEOZOIC ROCKS

Antelope Valley Formation

Vinini Formation

Graptolite-bearing shale

and limestone

_r

Contact

90

Fault, showing dip 

Dashed where approximately located; 

dotted where concealed 

37

Strike and dip of beds

RUGOSE CORAL LOCALITIES

Coral zone F Coral zone E

.M29 

Coral zone D

0M1044 

Coral zone C

Numbers preceded by M are USGS 

(Menlo Park) fossil locality numbers. 

Devonian coral zones A and B not 

recognized at Lone Mountain

S

Fragmentary Silurian rugose corals

is somewhat intermediate between that of the Diamond Mountains belt and the Antelope-Roberts

Mountains belt. In terms of Lower and lower Middle 

Devonian stratigraphic paleontology, the Sulphur 

Spring record is the most complete of the region, 

comprising coral zones A through D in continuous 

order. At Lone Mountain only zones C and D of this 

interval were recognized.

BEACON PEAK DOLOMITE MEMBER

Thick-bedded, rather finely saccharoidal, prevailingly light-gray dolomites of the Beacon Peak Dolomite 

Member underlie the isolated Twin Hills near Tyrone 

Gap, and crop out over a broader area at the south 

end of the range between the Mulligan Gap and 

Romano faults (fig. 4). Other outcrops occur to the 

north in the vicinity of Wales Peak and the Prince of 

Wales mine. Some 400 feet thick, the Beacon Peak of 

this area passes upward without known break into 

the arenaceous magnesian limestones and calcarenite 

at the bottom of Nevada Formation, unit 1. As in its 

type section in the southern Diamond Mountains at 

Oxyoke Canyon, the Beacon Peak Dolomite Member 

of the Sulphur Spring Range contains small lenses 

of quartzitic sandstone. Here and there it changes 

toward the top into dolomitic limestone.

Because of geomorphic and structural conditions in 

gently dipping Silurian and Devonian strata of this 

range the Lone Mountain Beacon Peak boundary has 

not been revealed by erosion (fig. 5). However, typical 

coarsely saccharoidal dolomite of the Silurian Lone 

Mountain makes up East Ridge on the east side of 

the Romano fault. In this footwall fault block the 

Lone Mountain contains abundant Halysites and 

pycnostylid corals of Silurian age, and may safely be 

inferred to underlie the Beacon Peak in the west or 

hanging-wall block of the high-angle Romano fault.

Fossils are scarce in most of the Beacon Peak 

Dolomite Member of the Sulphur Spring Range, 

especially at the south end and in the isolated Twin 

Hills. Scattered deposits of well-preserved silicified 

corals and brachiopods represent coral zone A and the 

Rabbit Hill Helderbergian fauna. Because no identifiable fossils were found in the Beacon Peak Dolomite 

of the type area in the Diamond Mountains, the correlation is based upon lithologic and stratigraphic 

criteria.

NEVADA FORMATION, UNIT 1

Unit 1, which is defined at Lone Mountain, is represented by quite similar strata in the Sulphur Spring 

Range (fig. 6). At Sulphur Spring Range, however, it 

includes basal arenaceous magnesian limestones and 

calcarenite unrecognized at Lone Mountain; these 

lowermost beds contain in considerable abundance 

the Costispirijer arenosus brachiopod assemblage of
12 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

^;>*;:;:s:aes^

\ f lilil \ 1 ^^yf^y T.'.V. TOtS-.-!.-. . .-. .-. 

--

EXPLANATION

39

45' 116° 10'

Base from U.S. Geological Survey, 1:62 500

Garden Valley, 1949

we 5'

Geology by C. W. Merriam and 

T. B. Nolan, 1952-68 

1MILE

1 .5 2 KILOMETERS

Mount Hope igneous

Devils Gate Limestone

Bay State Dolomite

Woodpecker Limestone

Sentinel Mountain

Dolomite Member

Undifferentiated 

Nevada Formation 

including all units 

Nevada unit 2 and members 

quartz sandstone 

tongues in undifferentiated Nevada 

Formation

Nevada unit 1 not

mapped separately

Shown in section

Beacon Peak Dolomite Quartzite and dolomite of unknown age;

possibly Lower and Middle Devonian

Lone Mountain Dolomite

Contains Halysites and

pycnostylids

WESTERN FACIES 

PALEOZOIC ROCKS

Contact 

Dashed where approximately located

Fault

Dashed where approximately located; 

dotted where concealed

,50 

Strike and dip of beds

Vinini Formation 

Graptolite-bearing shale

O

RUGOSE CORAL 

LOCALITIES

A

Coral zone D 

Late Early and Middle Devonian

Coral zone C 

Late Early Devonian

 

Coral zone B

Early Devonian (Oriskany)

A

Coral zone A 

Early Devonian (Helderberg)

S 

Silurian coral faunas

FIGURE 4. Geologic sketch map showing Silurian and Devonian units in southern Sulphur Spring Range. Symbols indicate

Devonian coral zones. See figure 5, geologic cross section D-D'.
STRATIGRAPHY OF CENTRAL GREAT BASIN DEVONIAN ROCKS 13

ORDOVICIAN PERMIAN SILURIAN

Note: Some Quaternary 

deposits not shown 

on geologic map (fig. 4)

1000 500

1000

I I 

3000 

1000 FEET 

J

i r

6000 METERS

FIGURE 5. Devonian stratigraphic units and coral zones in the southern Sulphur Spring Range.

See figure 4 for location of section.

Oriskany age, together with the large new rugose coral 

Kobeha which characterizes coral zone B. The basal 

sandy beds with Costispirifer and Kobeha are exposed 

at Wales Peak, Oriskany Peak, and Coral Ridge (fig.

4) where they are overlain stratigraphically by higher 

beds of unit 1 containing the coral zone C assemblage 

with-Papiliophyllum and Acrospirifer kobehana as in 

the Lone Mountain section.

NEVADA FORMATION, UNIT 2

Unit 2 is lithologically and faunally quite similar to 

the type exposures at Lone Mountain, consisting of 

argillaceous to silty limestones and siltstones with 

abundant fossils of coral zone D, which everywhere 

characterize the Antelope-Roberts Mountains facies 

belt. However, unlike the exposures at Lone Mountain, unit 2 contains in the Sulphur Spring Range 

numerous tongues and lenses of light-gray to white 

siliceous sandstone resembling the Oxyoke Canyon 

Sandstone Member of the Diamond Mountains belt. 

These sandstone bodies range in thickness from an 

inch to more than 40 feet. Rounded quartz granules 

also occur in some of the limestones. Sandstone beds 

are present throughout unit 2 in this area, but the 

thicker and laterally more persistent of these siliceous 

sandstones are in the upper part of the unit. Buttes 

like North Sand Peak and West Sand Peak (figs. 4,

5) are capped by these resistant sandstones, which 

are underlain by the fossil-rich argillaceous limestones 

characteristic of Nevada unit 2.

SENTINEL MOUNTAIN DOLOMITE MEMBER

The approximate stratigraphic equivalent of the 

Nevada Formation, unit 3 at Lone Mountain, the beds 

exposed between Oriskany Peak and Terrace Butte

(fig. 4) are largely unfossiliferous dolomite. At Lone 

Mountain, unit 3 is limestone, much of it fine-grained 

encrinite. Because it has few identifiable fossils, it is 

referred to as the barren zone. Whereas the gross 

bedding and textural features of these beds suggest 

unit 3 at Lone Mountain, the dolomitic nature and 

especially the alternation of thick light-gray and darkgray dolomite bands is more indicative of typical 

Sentinel Mountain Dolomite in the Diamond Mountains belt. The stratigraphic relationship to underlying Nevada Formation, unit 2 appears to be normal 

southeast of Oriskany Peak, but at Terrace Butte the 

Sentinel Mountain Dolomite is in fault contact with 

the overlying Woodpecker Limestone Member or 

Nevada Formation, unit 4.

WOODPECKER LIMESTONE MEMBER

The limestones of the Woodpecker Member occupy 

the interval of Nevada Formation, unit 4 at Lone 

Mountain. However, they are lithologically more like 

the typical Woodpecker Limestone of the Diamond 

Mountains and, like the Woodpecker there, yield a 

brachiopod fauna with abundant Leiorhynchus castanea. The prolific rugose coral assemblages of Nevada 

Formation, unit 4 and coral zone F at Lone Mountain 

are lacking, as is the case also in the typical Woodpecker Limestone Member.

BAY STATE DOLOMITE MEMBER

Thick-bedded light-gray and dark-gray dolomites 

which form the upper part of Terrace Butte and rest 

upon the Woodpecker Limestone Member are mapped 

as the Bay State Dolomite Member. This unit in the 

Diamond Mountains belt is the equivalent of Nevada 

Formation, unit 5 at Lone Mountain. Its characteristic
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M
STRATIGRAPHY OF CENTRAL GREAT BASIN DEVONIAN ROCKS 15

fossil is the large brachiopod Stringocephalus. In the 

Diamond Mountains, coral zone G brackets the basal 

beds of the Bay State and the topmost beds of the 

Woodpecker Limestone Member. No other outcrops 

of the Bay State were recognized in the southern 

Sulphur Spring Range.

DEVILS GATE LIMESTONE

The Devils Gate Limestone is cut out by faulting in 

the Terrace Butte vicinity. A fault sliver southwest 

of Bailey Pass includes the only Devils Gate exposures 

found in the Southern Sulphur Springs Range. Where 

present along the Mulligan Gap fault, it lies between 

conglomerate of the Permian Garden Valley Formation and lower beds of the Nevada Formation, 

and consists partly of massive stromatoporoid and 

Amphipora facies.

DIAMOND MOUNTAINS FACIES BELT

The geographic limits of the easterly Diamond 

Mountains facies belt (fig. 2) are those of the Diamond 

Mountains themselves. Between this belt and the 

Antelope-Roberts Mountains belt to the west, great 

lithologic and paleontologic changes are observed. 

Facies changes are most impressive in the lower part 

of the Devonian column. Within the 20 miles separating Lone Mountain from the Diamond Mountains, 

the normal Nevada Formation facies of units 1 and 2 

disappear and their places are seemingly occupied by 

dolomite and quartzitic sandstone, with consequent 

eastward loss of faunas (fig. 7). The possibility of 

major thrust displacement beneath the intervening 

Diamond Valley introduces tectonic factors which 

must be considered in this connection by the structural geologist. The greater part of the eastern Great 

Basin Devonian is likewise dolomite, and nowhere 

east of the Diamond Mountains have the lower faunas 

of Nevada Formation, units 1 and 2 been found thus 

far. No rugose corals of Early and early Middle 

Devonian ages are known in the Beacon Peak or 

Oxyoke Canyon Member of the Diamond Mountains 

facies belt or, in fact, from Devonian dolomites anywhere to the east within the Great Basin province.

Passing east from the Antelope-Roberts Mountains 

belt to the Diamond Mountains, the lithologic changes 

are of lesser magnitude within the upper divisions 

of the Nevada Formation. However, the faunas of 

the Woodpecker Limestone Member corresponding to 

Nevada Formation, unit 4 at Lone Mountain and 

those of Nevada Formation, unit 5 (which is essentially the Bay State Dolomite Member) reveal significant eastward biofacies changes. For example, the 

prolific coral assemblages of unit 4 (coral zone F) at 

Lone Mountain do not appear in the approximately

correlative Woodpecker Member of the Diamond 

Mountains.

BEACON PEAK DOLOMITE MEMBER

The Beacon Peak Dolomite Member is composed 

of sparsely fossiliferous fine-textured dolomites with 

subordinate siliceous sandstone intercalations and 

lenses which rest disconformably upon more coarsely 

saccharoidal Lone Mountain Dolomite, according to 

Nolan, Merriam, and Williams (1959, p. 42). The 

Beacon Peak Member is in turn conformably overlain 

by the even more barren Oxyoke Canyon Sandstone 

Member. Part of the Beacon Peak Dolomite Member, 

ranging in thickness from 470 feet to about 1,000 feet, 

is considered to be correlative with the Helderbergian 

Rabbit Hill Limestone, although its higher beds may 

be equivalent to arenaceous strata in the lower part 

of Nevada Formation, unit 1 in the Sulphur Spring 

Range. Poorly preserved ghosts of large high-spired 

gastropods are the only fossils known in this member.

OXYOKE CANYON SANDSTONE MEMBER

The Oxyoke Canyon Sandstone Member consists 

of thick-bedded quartz sandstone and orthoquartzite 

with dolomite interbeds; the orthoquartzite resembles 

white Eureka Quartzite but is less dense and locally 

has carbonate cement, uncommon in the Eureka. In 

places the Oxyoke Canyon Member is more than 400 

feet thick. Fossils are very uncommon; only at the 

Phillipsburg mine, 16 miles north of Eureka, were 

poorly preserved brachiopods found in quartzitic sandstones of this unit. As noted above, the presence of 

similar quartzitic sandstone as tongues and interbeds 

within the Nevada Formation, unit 2 rather strongly 

indicates that the Oxyoke Canyon Member is equivalent to some part of that westerly unit in the AntelopeRoberts Mountains facies belt. Westerly sandstone 

tongues of Oxyoke character in unit 2 are numerous 

in the Sulphur Spring Range, at Combs Peak, and at 

Modoc Peak in the Mahogany Hills, and at Grays 

Canyon and Eightmile Well in the Fish Creek Range.

SENTINEL MOUNTAIN DOLOMITE MEMBER

Thick-bedded generally unfossiliferous saccharoidal 

dolomites of the Sentinel Mountain Dolomite Member 

exhibit alternation of light-gray and dark-gray dolomite bands to a greater extent than the finer textured 

Beacon Peak. The Sentinel Mountain Member occupies the approximate interval of Nevada Formation, 

unit 3 at Lone Mountain, which is limestone, a good 

part being encrinite. Because of the scarcity of identifiable fossils in both members, they are referred to in 

paleontologic terms as the barren zone of the Nevada 

Formation.
16 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

SYSTEM

DEVONIAN

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SULPHUR SPRING RANGE DIAMOND MOUNTAINS 

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Bay State Dolomite 

Member 738 ft

Woodpecker Limestone 

Member 387 ft

Sentinel Mountain Dolomite 

Member 610 ft

Oxyoke Canyon Sandstone 

Member 400 ft

Beacon Peak Dolomite 

Member 470 ft

- Disconformity

Lone Mountain 

Dolomite

FIGURE 7. Relationship of the Devonian reference sections of Lone Mountain and Sulphur Spring Range to that of the

Diamond Mountains facies belt at Oxyoke Canyon.

WOODPECKER LIMESTONE MEMBER

Occupying the interval of Nevada Formation, unit 

4 at Lone Mountain, the Woodpecker Limestone 

Member consists of thin- to medium-bedded limestone 

and calcareous shale or siltstone, some of which is of 

slightly pinkish to purplish color. In certain beds, 

fossils are abundant, especially the brachiopods, of 

which the most diagnostic is Leiorhynchus castanea. 

The rich coral assemblages of the dolomitic limestones 

in Nevada Formation, unit 4 and coral zone F at Lone 

Mountain do not occur in the Woodpecker.

BAY STATE DOLOMITE MEMBER

The uppermost member of the Nevada Formation, 

the Bay State Dolomite Member, in this belt is thickbedded prevailingly dark gray saccharoidal dolomite. 

Some beds are loaded with large Stringocephalus 

shells, other beds contain much favositid coralline 

material. Rugose corals are common only in the lowermost beds, where they are locally associated with 

Stringocephalus. Nevada Formation, unit 5 of the 

Lone Mountain reference section is the approximate 

equivalent of the Bay State. Great Basin Devonian
CORAL, BIOSTRATIGRAPHY OF THE CENTRAL GREAT BASIN DEVONIAN 17

coral zone G brackets the lowermost strata of the Bay 

State and the topmost beds of the Woodpecker Limestone Member in the Diamond Mountains facies belt.

DEVILS GATE LIMESTONE

The Devils Gate Limestone, some 1,200 feet thick, 

rests conformably upon the Bay State Dolomite 

Member of the Nevada Formation and is overlain 

without discordance by the Pilot Shale. As noted 

under the Lone Mountain reference section, the lower 

part of the Devils Gate includes much massive stromatoporoid and Amphipora limestone including coral 

zone H. Thinner bedded upper limestones comprise 

coral zone I of Late Devonian (Frasnian) age. Coral 

facies are not well represented in the Diamond Mountains exposures of this limestone.

MONLTOR-SIMPSON PARK FACIES BELT

Lower Devonian limestones of the westerly MonitorSimpson Park facies belt have been traced southward 

from the Tuscarora and Cortez Mountains (fig. 2) 

through the Simpson Park, Toquima, and Monitor 

Ranges to the Dobbin Summit area, a distance of 

130 miles. Unlike the two subparallel facies belts on 

the east, no dolomite is known in either the Devonian 

or Silurian of the Monitor-Simpson Park belt. Over 

most of this belt the Helderbergian Rabbit Hill Limestone is the only Devonian unit identified by fossils. 

Exceptional are outcrops of Nevada Formation, unit 2 

(coral zone D) at the north end of the Simpson Park 

Range near Coal Canyon. Coral-bearing strata of 

probable Middle Devonian age are known on the west 

side of the Monitor-Simpson Park belt in the Toiyabe 

Range south of Austin. This occurrence near Reeds 

Canyon is the westernmost exposure of Devonian in 

this region. Underlying strata of possible Early 

Devonian age at the Reeds Canyon locality have thus 

far yielded no fossils within the stratigraphic interval 

just above the Silurian where Rabbit Hill Limestone 

would be expected.

RABBIT HILL LIMESTONE

In the type section of the Rabbit Hill Limestone 

at Copenhagen Canyon, Monitor Range (Merriam, 

1963, p. 42), about 250 feet of continuous section is 

measurable. The dark-gray limestones and calcareous 

shales of the type Rabbit Hill are deformed below a 

thrust and rest without recognized stratigraphic break 

upon graptolitic deposits of Silurian age. More complete Rabbit Hill sequences occur at Dobbin Summit 

(fig. 2) and at Coal Canyon, where in each case about 

800 feet of the formation is exposed. At Coal Canyon

the underlying limestones of Silurian coral zone E 

contain a large and diagnostic coral assemblage of 

Gotlandian character; overlying Nevada beds have 

yielded faunas of Nevada Formation, unit 2 and coral 

zone D, but the stratigraphic contact is not exposed. 

In the Dobbin Summit area, the subjacent Silurian 

strata have yielded graptolites and brachiopods only. 

The Rabbit Hill is overlain without recognized discordance by conodont-bearing limestones reported to 

be of Mississippian age (John W. Huddle, oral commun., 1966). Although no Rabbit Hill Limestone has 

been found in the adjacent Antelope-Roberts Mountains belt on the east, diagnostic elements of the 

Rabbit Hill fauna occur in strata assigned to the 

Beacon Peak Dolomite Member below Nevada Formation, unit 1 of the Sulphur Spring Range.

All the Rabbit Hill Limestone exposures that have 

been studied represent the Syringaxon facies and 

Devonian coral zone A. At Dobbin Summit upward 

faunal changes are, however, observed within this 

formation. The higher beds contain a large Costispirifer cf. C. arenosus similar to that of lowermost 

Nevada Formation, unit 1 and coral zone B.

CORAL BIOSTRATIGRAPHY OF THE 

CENTRAL GREAT BASIN DEVONIAN

Rugose corals of restricted vertical range make possible biostratigraphic subdivision of the entire Great 

Basin Devonian into nine successive coral zones labeled 

by capital letters A through I in stratigraphic order 

(fig. 8). Four of these time-stratigraphic divisions 

correspond to mapped rock-stratigraphic units; the 

remainder occupy rock units with two coral zones or, 

as in the case of coral zone G, overlap the contact 

separating two units (fig. 9). Field reference sections, 

wherein the rocks have been mapped and the stratigraphic section measured, are given for each coral zone.

Value of the proposed biostratigraphic zonation outside the Great Basin province remains to be demonstrated. A possible disadvantage is the use in the 

zonation scheme of endemic Halliidae like Kobeha and 

Papiliophyllum, not yet recorded in other provinces. 

Other families and genera employed are more cosmopolitan, like the Laccophyllidae, Siphonophrentinae, 

Bethanyphyllidae, Disphyllidae, and the Old World 

digonophyllids.

Initial biostratigraphic zoning of Great Basin 

Devonian rocks (Merriam, 1940, p. 9) was based 

largely on spiriferoid brachiopod ranges; the taxonomy 

and geologic range of these have recently undergone 

revision (Johnson, 1966a; 1966b, 1966c). Some earlier 

adopted brachiopod markers continue to agree with 

coral indices of the lettered zones. For example, the
18 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

SYSTEM

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COMMON RUGOSE CORALS

No rugose corals found

Pachyphyllum, Phillipsastraea, 

Macgeea

Rugose corals uncommon

Acanthophyllum, Digonophyllum

(Mochlophyllum)

Hexagonaria, Taimyrophyllu.m, 

Utaratuia, Sociophyllum,

Keriophyllum, Acanthophyllum,

Digonophyllum, Mesophyllum,

Tabulophyllum

Rugose corals uncommon

(barren zone)

Mesophyllum (Arcophyllum)

Piny ona sir aea, Billing sastraea,

Mesophyllum, Zonophyllum

Papiliophyllum, Breviphrentis

Papiliophyllum, Odontophyllum,

Breviphrentis

Kobeha

Syringaxon

RUGOSE 

CORAL 

EVOLUTIONARY

BURST*

*

*

^.

CORAL BIOFACIES

Phillipsa stra ea 

biofacies

Hexagonaria- Sociophyllum

biofacies

Breviphrentis

biofacies

Papiliophyllum

biofacies

Kobeha 

biofacies

Syringaxon 

" * biofacies

PREVIOUSLY 

USED ZONE

Cyrtospirifer zone

Phillipsastraea zone

String ocephalus zone

"Martinia" kirki zone

"Spirifer" pinyonensis

zone

Acrospiri/er kobehana

zone

Costispirifer arenosus 

zone ("Trematospira 

fauna 'V

EUROPEAN 

STAGE

Famennian

Frasnian

Givetian

Eifelian

__ _ _ _ _

Emsian

Siegenian 

(Oriskany)

Helderbergian

FIGURE 8. Great Basin Devonian coral zones, equivalent rock units, coral biofacies, and previously employed biostratigraphic

designations.

Acrospirifer kobehana zone is coral zone C, and that 

of Costispirifer arenosus, or the "Trematospira fauna," 

is coral zone B. The zone of "Spirifer" pinyonensis 

(now Eurekaspirifer pinyonensis) is essentially coral 

zone D, although subzone D3 is considered Middle 

Devonian and may be above the range of this spirifer. 

In connection with rugose coral study, the associated 

brachiopods and other diagnostic fossils remain to 

be more fully evaluated stratigraphically. Tabulate 

corals, actually the most abundant fossils in certain 

Great Basin Devonian coral-brachiopod-trilobite beds, 

are too little understood at present to have definitive 

stratigraphic value in this province. Cephalopods, 

other than straight-shelled genera, are uncommon in

these rocks. Age and correlation significance of the 

rare ammonoids collected thus far are discussed below. 

The proposed coral sequence is by nature composite, for in no individual section are all nine zones 

recognized in continuity. Lone Mountain has four in 

sequence: C, D, E, and F; strata of zones G and H 

are present, but thus far have yielded no Rugosa at 

Lone Mountain. For the purposes of this report, the 

Early and Middle Devonian section in the southern 

Sulphur Spring Range is the most inclusive; coral 

zones A, B, C, and D occur here in continuity. Devonian coral zones G and H are best represented in the 

southern Diamond Mountains. Zones H and I are 

recognized in the Devils Gate vicinity, and in the
CORAL BIOSTRATIGRAPHY OF THE CENTRAL GREAT BASIN DEVONIAN 19

adjacent northeastern part of the Mahogany Hills. 

Late Devonian rocks that include the interval of coral 

zone I are well exposed in the White Pine mining 

district, especially along the west margin near the 

Belmont mine and in the low hills west of Monte 

Cristo.

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 H are 

Middle Devonian. Coral zone I is Late Devonian, 

including the youngest Devonian rugose corals of this 

province; none have been found in the high Devils 

Gate Cyrtospirifer beds, or in the lower Pilot Shale 

interval that probably correlates with European 

Famennian.

For convenience, the lettered primary coral zones 

are further divided by subnumbers as DI and D2 . In 

this instance subzone DI has a distinctive rugose coral 

assemblage regarded as late Early Devonian, whereas 

that of D2 is considered Early or early Middle Devonian; subzone D3 is early Middle Devonian.

In summary, the mapped Devonian rock units containing the zone corals remain to be equated with corresponding biostratigraphic coral range zones. Beginning with the oldest, coral zone A is typified by the 

westerly situated Rabbit Hill limestones but is present 

also in the easterly Beacon Peak Dolomite Member 

interval of the Sulphur Spring Range, beneath Nevada 

Formation, unit 1 with coral zones B and C (fig. 9). 

In the Lone Mountain reference section, only coral 

zone C has been identified in Nevada Formation, unit 

1; coral zones D, E, and F of this reference section 

correspond to Nevada Formation, units 2, 3, and 4. 

Coral zone G is found in a narrow band which brackets 

the topmost beds of the Woodpecker Limestone Member and the basal beds of the Bay State Dolomite 

Member (Nevada Formation, unit 5) in the southern 

Diamond Mountains. Coral zone H occupies the lower 

half of the type Devils Gate Limestone wherein the 

stromatoporoid-Amp/wpora biofacies is predominant. 

At Devils Gate, coral zone I is typified by the 

Phillipsastraea biofacies in the lower part of the uppermost 250 feet of this formation below the Cyrtospirifer 

beds.

EARLY AND EARLY MIDDLE 

DEVONIAN CORAL ZONES

CORAL ZONE A

The type section of the Rabbit Hill Limestone in 

Copenhagen Canyon, northern Monitor Range, is the 

reference column for this Helderbergian Early Devonian coral zone. Syringaxon foerstei n. sp., index fossil 

of coral zone A and the Syringaxon biofacies, is common to abundant in a large brachiopod-coral fauna 

throughout this formation, usually unaccompanied by

other Rugosa. Syringaxon as a genus ranges through 

the Silurian and Devonian; it is represented by many 

species throughout the world. The genus has been 

recognized by fragmentary material in the Silurian 

and Middle Devonian of the Great Basin.

The stratigraphic relationship of the Rabbit Hill 

Formation and coral zone A to underlying and overlying zones of the Great Basin standard sequence can 

not be determined at Rabbit Hill, where the subjacent Silurian is a graptolitic facies and no younger 

Devonian beds are exposed. Therefore elucidation of 

the true position of these beds depends upon supplementary study of correlative strata in other areas. At 

Coal Canyon, northern Simpson Peak Range, limestones of the Rabbit Hill rest conformably upon Upper 

Silurian limestones of the Roberts Mountains Formation and Silurian coral zone E containing a rich 

Ludlovian coral fauna of Gotlandian aspect. Upward, 

however, the boundary with the Nevada Formation is 

obscure because of faulting and poor exposure. In the 

Dobbin Summit area of the southern Monitor Range, 

the Rabbit Hill Limestone is conformably upon 

Silurian limestone and is overlain with seeming conformity by limestones of uncertain age, dated provisionally by conodonts as Mississippian (J. W. 

Huddle, oral commun., 1966).

Light is shed upon the true stratigraphic position 

of the Rabbit Hill fauna and coral zone A as a result 

of geologic mapping in the southern Sulphur Spring 

Range. Here dolomite and dolomitic limestone, considered to be a northwesterly extension of the Beacon 

Peak Dolomite Member, contain Syringaxon foerstei 

and other elements of the Rabbit Hill fauna. In that 

area the Silurian Lone Mountain Dolomite is below 

the Beacon Peak above which lies Nevada Formation, unit 1, whose lowermost beds contain Kobeha 

walcotti and Costispirifer arenosus of coral zone B. 

As elsewhere noted, the normal Beacon Peak-Lone 

Mountain stratigraphic boundary is poorly exposed 

in the southern Sulphur Spring Range; in the Diamond 

Mountains it is a disconformity.

In the type section of the Rabbit Hill and coral 

zone A, Syringaxon foerstei is associated with the 

tabulate corals Striatopora, Pleurodictyum, and massive Favosites. Distinctive forms among associated 

brachiopods are Levenea cf. L. subcarinata, Howellella 

cf. H. cycloptera and new species of Kozlowskiellina, 

Leptocoelia, and Plethorhyncha. Leonaspis, Proetus, 

and Phacops are the common trilobites. Large Costispirifer resembling C. arenosus appears in higher Rabbit 

Hill beds at Dobbin Summit. This early occurrence 

of Costispirifer foreshadows its presence in coral zone 

B, normally to be expected just above the Rabbit Hill.
20 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Virtual absence of colonial Rugosa from the known 

Great Basin Lower Devonian below coral zone D is 

not easily explainable. Although the faunas of coral 

zone A normally lack the compound genera, there are 

two exceptions: the occurrence of Australophyllum 

landerensis n. sp. in an otherwise typical Rabbit Hill 

Syringaxon association of the northern Toquima 

Range, and a Billingsastraea-like coral in a Rabbit 

Hill assemblage on Maggie Creek northwest of Carlin, 

southern Tuscarora Mountains.

CORAL ZONE B

Reference sections for this Early Devonian coral 

zone are situated in the southern Sulphur Spring 

Range, on the ridge 0.5 mile south-southeast of the 

Prince of Wales mine and at hill 6927,1.75 miles southsoutheast of Mulligan Gap (fig. 4). Along the main 

ridge at hill 6927, the zone B strata are in places 

loaded with large solitary Rugosa, overlie beds of 

coral zone A, and are overlain by beds containing the 

fauna of coral zone C. Zones B and C are within the 

variable dolomitic limestone and limestone of Nevada 

Formation, unit 1, which rests without break upon the 

Beacon Peak.

The fauna of coral zone B and the Kobeha biofacies 

contains Kobeha walcotti, Costispirifer arenosus, and 

Gypidula cf. G. coeymanensis. The large solitary 

Kobeha is abundant in beds and pockets, either by 

itself or with the brachiopods. In places Costispirifer 

is crowded in layers without the coral. Tabulate corals 

are uncommon in the coral-rich beds. Other fossils of 

this zone are a large Strophonella cf. S. punctulifera 

and the large trilobite Dalmanites. It is significant 

that the fauna of coral zone B contains but a single 

rugose coral species and lacks colonial Rugosa.

Another species of Kobeha, here named Kobeha 

ketophylloides n. sp., occurs in beds low in the Devonian section 1.5 miles north of Roberts Creek Ranch, 

where it is associated with Gypidula cf. G. coeymanensis, Acrospirifer sp., and Pholidostrophiat sp.

CORAL ZONE C

The reference section for Early Devonian corai zone 

C is at Contact Gulch on the northwest side of Lone 

Mountain (fig. 3). A northwest-striking ridge on the 

north side of this gulch is underlain by Nevada 

Formation, unit 1, which in this area has changed 

laterally from dolomite and dolomitic limestone to 

the limestone here containing fossils of coral zone C. 

Other excellent exposures of these beds are on the 

southeast side of Lone Mountain, where Nevada 

Formation, unit 1 is dolomite which is loaded in places 

with silicified fossils. Coral zone B was not recognized

at Lone Mountain below coral zone C where it would 

be expected. In the southern Sulphur Spring Range, 

coral zone C occupies the higher beds of Nevada 

Formation, unit 1 overlying the strata of coral zone B.

The large solitary Papiliophyllum elegantulum replaces Kobeha walcotti of underlying zone B. Associated with Papiliophyllum in this taxonomically more 

diverse biota are Siphonophrentis (Breviphrentis) 

kobehensis, Odontophyllum meeki, Aulacophyllum sp. 

c, and Cystiphylloides lonense. Massive favositids are 

common. No colonial Rugosa are present. The associated brachiopod fauna is fairly large, including 

Acrospirifer kobehana, Meristella cf. M. robertsensis, 

Schuchertella cf. S. nevadaensis, Leptostrophia sp., 

and a large Cyrtina. This is the fauna of the Acrospirifer kobehana brachiopod zone of previous usage.

The genus Papiliophyllum is not restricted to coral 

zone C; it reappears as an advanced form in coral 

subzone DI, associated with a younger brachiopod 

fauna.

CORAL ZONE D

The reference section of coral zone D lies on the 

west side of Lone Mountain (fig. 3) between West 

Peak and Central Peak, where it brackets the argillaceous limestones of Nevada Formation, unit 2. Coral 

zone D corresponds to the "Spirifer" pinyonensis zone 

of Merriam (1940). The Early and early Middle 

Devonian coral and associated faunas of this zone are 

taxonomically the most diverse of the Great Basin 

Devonian, only those of Nevada Formation, unit 4 

and coral zone F being comparable in this respect. 

Because of the less compact shaly nature of much of 

this unit, the abundant fossils weather loose and are 

easily collected. Hence a large part of the previously 

described Nevada Formation fauna came from these 

beds. Coral zone D includes the lowest colonial Rugosa 

found thus far in the Nevada Formation.

Using rugose corals, it is feasible to subdivide coral 

zone D into three subzones, DI, D2 , and D3 , in ascending order. Most unit 2 exposures in the main outcrop 

belt at Lone Mountain yielded D2 collections only. 

Large collections representing DI came from pediment 

outcrops 5,500 feet south of Reef Point. Subzone D3 

collections were made in the higher part of Nevada 

Formation, unit 2 near the northwest end of Lone 

Mountain at the head of Contact Gulch.

Subzone DI is characterized by advanced Papiliophyllum elegantulum subsp. d, derived from P. 

elegantulum of coral zone C. Associated large solitary 

corals are Halliidae of a somewhat different type, 

probably allied to Papiliophyllum. Among brachiopods 

are Chonetes macrostriata, Gypidula loweryi, and 

Strophonella cf. S. pustulosa, together with Leptaena,
CORAL BIOSTRATIGRAPHY OF THE CENTRAL GREAT BASIN DEVONIAN 21

Schuchertella, and Meristella. Spirifers questionably 

assigned to Eurekaspirifer pinyonensis are uncommon 

in this assemblage. Chonetes macrostriata is abundant, as it usually the case in lower beds of coral 

zone D.

Subzone D2 includes among solitary Rugosa: 

Bethanyphyllum lonense, B. antelopensis, Siphonophrentis (Breviphrentis) invaginatus, Sinospongophyllum spp., Cystiphylloides robertsense, Mesophyllum (Mesophyllum) sp. b, and Zonophyllum haguei. 

The colonial corals are Disphyllum nevadensis, D. 

eurekaensis, Dendrostella romanensis, Billingsastraea 

nevadensis, and Hexagonaria (Pinyonastraea) kirki. 

Associated fossils are those of the "Spirifer" pinyonensis zone, as previously listed by Merriam (1940, p. 

54-55) in which the commonest forms are Eurekaspirifer pinyonensis, Atrypa nevadana, Gypidula 

loweryi, Schizophoria nevadaensis, Schuchertella 

nevadaensis, Chonetes macrostriata, Dalmanites 

meeki, and Phacops cf. P. rana.

Subzone D3 includes the uppermost beds of Nevada 

Formation, unit 2 in which the diagnostic coral is the 

large solitary and structurally complex Mesophyllum 

(Arcophyllum) kirki. It is doubtful that Eurekaspirifer pinyonensis survived into this early Middle 

Devonian time-stratigraphic interval, and most of the 

characteristic fossils of subzone D2 have not been 

collected from subzone D3 .

MIDDLE MIDDLE DEVONIAN CORAL ZONES

Profound changes in corals and all other shellbearing marine organisms took place in the interval 

separating Nevada Formation, unit 2 and coral zone 

D from Nevada Formation, unit 4 and coral zone F. 

These changes are not recorded in starta of Nevada 

Formation, unit 3, which has yielded few identifiable 

fossils, hence being referred to as the barren zone. 

None of the species cross the barren zone, and of five 

rugose coral genera that survived to coral zone F, 

three are different subgenerically.

CORAL ZONE E

Coral zone E occupies the interval of Nevada Formation, unit 3 at Lone Mountain (fig. 3). Crinoid 

debris in thick beds and lenses makes up a considerable part of the unit. Poorly preserved bushy colonial 

Rugosa from these beds are assigned questionably to 

Disphyllum, and scrappy solitary coralla have characteristics of Cyathophyllum.

CORAL ZONE F

The reference section for coral zone F is situated on 

the south side of Lone Mountain, west of Charcoal

Gulch (fig. 3) and is the time-stratigraphic equivalent 

of Nevada Formation, unit 4. The "Martinia" kirki 

zone of Merriam (1940) corresponds to this interval. 

The coral faunas of this zone at Lone Mountain are 

best represented in dolomitic limestones occupying the 

upper part of unit. 4

Coral zone F contains a large, diverse, and potentially reef forming biota including both solitary and 

colonial genera. The rugose corals of this zone are 

mostly undescribed, such that generic assignments 

are for the most part provisional. Among the more 

abundant genera are: Taimyrophyllum, Hexagonaria, 

Utaratuia, Sociophyllum, Acanthophyllum,Keriophyllum, Digonophyllum, Mesophyllum (Lekanophyllum), 

Tabulophyllum, and Crystiphylloides. For the first 

time in these Devonian rocks, the colonial Rugosa of 

coral zone F outnumber the solitary forms. Nearly 

all zone F species are unrelated to zone D species; none 

appear to be ancestral to forms occurring in the higher 

Devonian coral zones. Several of the diagnostic genera 

like Taimyrophyllum, Utaratuia, and Sociophyllum 

are represented also in correlative formations of 

western Canada.

CORAL ZONE G

The reference section for coral zone G is situated in 

the southern Diamond Mountains (fig. 2) at the head 

of Oxyoke Canyon, west of Sentinel Mountain (Pinto 

Summit quadrangle). It occupies a 30-foot interval 

bracketing the uppermost beds of the Woodpecker 

Limestone Member of the Nevada formation and the 

lowermost beds of the overlying Bay State Dolomite 

Member according to Nolan, Merriam, and Williams, 

(1956, p. 45). The lowest Bay State beds with Stringocephalus fall within this coral zone, which also contains 

the genus Rensselandia. Only in this zone are rugose 

corals known to be associated with Stringocephalus. 

Characteristic rugose coral genera of coral zone G are 

Digonophyllum (Mochlophyllum), Mesophyllum, 

Acanthophyllum, Cyathophyllum (Moravophyllum), 

and Cystiphylloides. The colonial Rugosa of coral zone 

F at Lone Mountain are not represented in the reference section for coral zone G.

At Lone Mountain the basal beds of Nevada Formation, unit 5, which is approximately correlative with 

the Bay State, and the topmost beds of Nevada Formation, unit 4, correlative with the Woodpecker Limestone Member, have not yielded the coral zone G 

assemblage.

LATE MIDDLE AND LATE DEVONIAN CORAL ZONES

The rugose corals and stratigraphy of the Devils 

Gate Limestone remain to be studied in detail and, 

except for generalities, are beyond the scope of this
22 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

report. Earlier investigation of Rugosa from the upper 

part of this formation by Stumm (1940) and by Merriam (1940) confirmed their Late Devonian age and 

demonstrated that these higher coral faunas were 

unrelated genetically to those of the Middle Devonian 

in coral zones F and G of present usage. Above the 

Stringocephalus zone the interval recorded by thickbedded limestones which make up the lower half of 

the Devils Gate was seemingly inhospitable to rugose 

corals, as was the dolomitic Stringocephalus zone in 

Nevada Formation, unit 5 and the Bay State Dolomite 

Member. When Rugosa reappear in force just below 

the uppermost Devils Gate Cyrtospirifer zone, they 

include newly introduced genera, mostly of a single 

family not yet discovered in the Great Basin below 

this horizon. For the Devils Gate time-stratigraphic 

interval, two coral zones are proposed: coral zone H 

below to include the lower coral-poor thick-bedded 

limestones, and coral zone I above for the upper coralrich beds.

CORAL ZONE H

Massive stromatoporoids and Amphipora are major 

rock builders in the interval of coral zone H and as 

such have masked and obscured the contributions of 

other lime-secreting organisms. No Rugosa were found 

in these beds at Lone Mountain. Rugose corals 

assigned to Temnophyllum and Disphyllum were collected from the lower 200 feet of the Devils Gate in 

the southern Diamond Mountains (Merriam, 1963, p. 

53). The middle Devonian-Late Devonian transition 

conceivably falls within the interval of coral zone H.

CORAL ZONE I

Reference section for Late Devonian coral zone I lies 

in the Devils Gate Limestone type area on the north 

side of Devils Gate Pass. The dark-gray coralline limestones below the Cyrtospirifer beds contain the locally 

abundant Phillipsastraeidae of this zone with only a 

few solitary Rugosa of other surviving families. Phillipsastraeidae present are the colonial Phillipsastraea, 

PachyphyHum, and Phacellophyllum together with the 

solitary MacGeea.

Other localities where the coral faunas of zone I are 

represented in the Devils Gate vicinity are on the ridge 

east of Yahoo Canyon, Whistler Mountain quadrangle. 

In the White Pine mining district, these faunas occur 

abundantly near the Belmont mine mill and on the 

ridge southwest of Monte Cristo, Green Springs quadrangle (Merriam, 1963, p. 54).

Limestones probably equivalent to the lower or Late 

Devonian part of the Pilot Shale crop out at Devils 

Gate above the Cyrtospirifer zone; like the Cyrtospirifer beds, these highest Devonian rocks have yielded no 

Rugosa.

BIOFACIES ASPECTS OF DEVONIAN CORAL 

DISTRIBUTION AND ZONATION

Ideally zone fossils may be environmental indicators 

as well as stratigraphic guides. Patterns of coral distribution in the Great Basin Devonian seaways and their 

facies belts (fig. 2) give suggestive evidence of environmental control. Rugose coral populations in these seas 

formed only a part of much larger ecosystems whose 

spread and survival as a whole were governed by interplay of ecologic factors, physical, chemical, and biologic. Within the same time-stratigraphic interval, 

entombing sediments of a specific kind may be viewed 

as criteria of a fairly specific and stable bottom environment where they contain about the same coral biota 

in widely separated localities. In the section on "Great 

Basin Devonian Facies Belts and Coral Reference Sections," the broader aspects of rock facies distribution 

were considered above in connection with physical 

stratigraphy. There remains the ecologic evaluation of 

coral zone indicators and their ecologic associates as 

evidence of a particular biofacies.

From the viewpoint of the biological environment, 

discontinuous lateral distribution of coral-rich Devonian deposits indicates that optimum ecologic conditions for vigorous coral growth were confined to rather 

small patches of sea bottom separated by larger areas 

of scarce shell-bearing benthonic life. Nearly all these 

coral deposits circumscribed coral loci and thus were 

not sites of true reef or bioherm building. No biohermal 

or patch reef complexes of persistent upward growth, 

with initial sea bottom relief, altered reef cores, and 

inclined flank beds like those of the Niagaran and the 

Gotland Silurian were recognized in the Great Basin 

Devonian. Only in the highest Silurian and in middle 

Devonian coral zone F do loci of coral proliferation 

suggest patch reef potential.

Carbonate rocks are generally the favored hosts of 

rugose corals, especially the colonial forms. Moreover, 

these Great Basin studies suggest an environmental 

relationship between abundance and diversity of 

Rugosa and certain petrologic characteristics of the 

entombing carbonate. Taxonomically the most diverse 

Devonian coral faunas of this province occur in the 

Nevada Formation, unit 2 and coral zone D; the faunas 

are found in impure nonmagnesian argillaceous limestones with shale and siltstone interbeds. To these 

varied coral faunas may be compared the more vigorously proliferating coral associations of Nevada Formation, unit 4 (coral zone F) which occur in combinations 

of limestone, dolomitic limestone, and dolomite having 

a lower percentage of argillaceous matter. Coral faunas 

of Nevada Formation, unit 4 consist of slightly fewer 

species, but reveal a greater numerical concentration
BIOFACIES ASPECTS OF DEVONIAN CORAL DISTRIBUTION AND ZONATION 23

of individuals, especially colonial forms, and are thus 

more important in rock building. On the contrary, the 

pure or saturated coarsely saccharoidal Devonian dolomites of this region, as distinguished from unsaturated 

dolomitic limestones, seem to have been a diagenetic 

facies singularly inhospitable to Rugosa. Among these 

are the Sentinel Mountain and Bay State Dolomite 

Members in the upper part of the Nevada Formation; 

these have yielded few rugose corals. Rugosa are likewise scarce in diagenetic dolomites making up much 

of the Devonian System of the eastern Great Basin, 

that is, east of the Diamond Mountains. Easterly 

rugose coral scarcity would appear to support these 

paleoecological observations in the Nevada Formation 

and suggests that Devonian shelf sea conditions favoring extensive rugose coral growth like that of coral 

zone F did not prevail in the eastern part of the Great 

Basin province. To be eliminated in this connection 

is the factor of possible coral destruction by dolomitic 

recrystallization. However, in these Cordilleran pure 

dolomites, where fossils were present in the initial sediments, they commonly escaped destruction locally by 

early siliceous replacement.

Six biofacies of the Great Basin Devonian (fig. 9) 

wherein rugose corals are important faunal constituents occur in the following stratigraphic order: 

6. Phillipsastraea biofacies of coral zone I 

5. Hexagonaria-Sociophyllum biofacies of coral

zone F

4. Breviphrentis biofacies of coral zone D 

3. Papiliophyllum biofacies of coral zone C 

2. Kobeha biofacies of coral zone B 

1. Syringaxon biofacies of coral zone A

SYRINGAXON BIOFACIES

Coral-bearing medium- to dark-gray organic limestones of the Rabbit Hill typify the Syringaxon biofacies and coral zone A. Separated by calcareous shaly 

intercalations, these limestones commonly include a 

high percentage of crinoid debris. Thick coarse-textured bioclastic lenses contain the remains of a large 

brachiopod population, together with tabulate corals 

of the genera Favosites, Striatopora, and Pleurodictyum and the rugose coral biofacies indicator Syringaxon; this order indicates more or less their decreasing 

order of abundance. Less numerous are the bryozoans, 

trilobites, and the Mollusca other than small orthoceratid cephalopods which are rather common. There 

are no evidences of long transportation of the fossil 

material; unbroken shells are present in the bioclastic limestones, many brachiopods having articulated 

valves. Stromatoporoids are a minor constituent. The 

trilobites Leonaspis and Phacops are locally numerous

in platy limestone and calcareous shale interbeds 

which are a subsidiary biofacies. Throughout the Rabbit Hill limestone the small, solitary Syringaxon is 

common to abundant in most fossil beds, wherein it is 

usually the only representative of the Rugosa.

No bioherms or patch reefs were found in the Rabbit 

Hill Limestone of the Syringaxon biofacies, and the 

corals had a minor role in rock building. No evidence 

of dolomitization was recognized in the Rabbit Hill 

Limestone of this biofacies. Complete silicification of 

fossils is, however, widespread, a phenomenon which 

in other formations is commonly associated with or 

preceded dolomitic replacement. In the Rabbit Hill the 

process of fossil silicification was clearly selective, 

affecting brachiopods and corals; hard parts of other 

organic groups were evidently less susceptible to this 

kind of silicic additive replacement.

Colonial Rugosa are exceedingly uncommon in the 

Rabbit Hill and Syringaxon biofacies, there being but 

two known occurrences: that of Australophyllum 

landerensis in the Toquima Range and a single Billingsastraea-like form in the southern Tuscarora 

Mountains of northern Eureka County. Absent are 

thick-walled solitary rugose coral genera of a possible 

reef environment like those described from the roughly 

correlative Coeymans Limestone of New York by 

Oliver (1960a). Possibly significant environmentally 

is the small amount of stromatoporoid material in this 

biofacies, a characteristic also of the Breviphrentis biofacies and coral zone D. Its scarcity may be related to 

frequent turbidity of the marine waters.

The Syringaxon biofacies is present also in the Beacon Peak Dolomite Member of the Nevada Formation 

in the Sulphur Spring Range; here the fossils occur 

in limestone beds which are somewhat dolomitic.

KOBEHA BIOFACIES

The Kobeha biofacies is in the strictest sense a 

rugose coral biofacies in which the single species K. 

walcotti predominates. This biofacies is best represented in the lowermost beds of Nevada Formation, 

unit 1 (coral zone B) in the southern Sulphur Spring 

Range. Here as at Lone Mountain, unit 1 changes 

laterally in quite irregular fashion from dolomite to 

dolomitic limestone to limestone. It is in lenses of calcarenite limestone with many rounded quartz granules 

that the large solitary Kobeha is most conspicuous and 

is accompanied by few other fossils. Dolomitic limestones are associated with these coralline facies. In 

the same stratigraphic interval other calcarenite lenses 

are loaded with the large shells of Costispirifer of the 

C. arenosus type. Horn corals and brachiopods appear 

to have been deposited in situ, but some of the large
24 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

spirifer valves may have been sorted by current action 

from the heavier corals to be concentrated in separate 

bands. The corals have been silicified in varying 

degrees.

PAPELIOPHYLLUM BIOFACIES

The large solitary Papiliophyllum elegantulum of 

the Halliidae is the most conspicuous fossil of this biofacies and in coral zone C, which occupies all but the 

lowermost beds of Nevada Formation, unit 1. At Lone 

Mountain and in the Sulphur Spring Range, unit 1 is 

laterally variable, changing in short distances from 

limestone to dolomitic limestone and to dolomite. Like 

the Kobeha biofacies and coral zone B at the base of 

unit 1, this biofacies lacks colonial Rugosa; however, 

the fauna as a whole is very much larger. The Halliidae, now at their evolutionary peak, also include Aulacophyllum and Odontophyllum; rugose corals of other 

families are Cystiphylloides and the earliest Breviphrentis. Favosites in massive heads is the common 

tabulate. As a whole, the fauna is a mixed association 

containing many brachiopods, of which the most distinctive is Acrospirifer kobehana. Crinoid debris is 

abundant. Gastropods and clams are fairly numerous. 

Papilophyllum succeeds Kobeha, from which it probably evolved, and continues to occupy the same ecologic niche. Within unit 1 there appears to be no special 

preference of this fauna for either limestone or dolomitic lithofacies. Corals and other fossils are silicified, 

especially in the more dolomitic carbonate. Relative 

abundance of stromatoporoids in the cleaner carbonates of this biofacies has not been determined; probably they are numerous and, like the corals, of some 

rock-building importance. No bioherms or patch reefs 

were found in beds of the Papiliophyllum biofacies.

BREVIPHRENTIS BIOFACIES

Following decline of the large Halliidae that characterize the Kobeha and Papiliophyllum biofacies, 

rugose corals of other families continued to increase 

in numbers of genera and species. In terms of corals 

and ecologically associated fossils, the Breviphrentis 

biofacies of coral zone D includes the largest and 

taxonomically most varied faunas of the Great Basin 

Devonian. The enclosing strata of Nevada Formation, 

unit 2, to which this biofacies is confined, are impure medium- and dark-gray argillaceous organic 

limestones separated by shale and siltstone partings 

and interbeds. Bottom faunas of this stratigraphic 

interval were subjected periodically to abnormally 

muddy water resulting from influx of current-transported clay or brought about by storm turbulence. 

Favositids, solitary Rugosa, brachiopods, trilobites, 

and Mollusca adapted well to the frequent turbid 

water episodes. For the first time in the Devonian,

colonial Rugosa appear in this biofacies, but these 

occur sporadically, are rather uncommon, and are far 

outnumbered by solitary genera. Stromatoporoids are 

quite scarce. It is suggested that stromatoporoid 

scarcity and the small population of colonial Rugosa 

may reflect inability of these organisms to withstand 

turbidity. In the case of stromatoporoids such a likelihood has been noted elsewhere by Rozkowska (1960, 

P. 7).

Unlike cleaner carbonate rocks of Nevada Formation, unit 1, the impure limestones of unit 2 and the 

Breviphrentis biofacies were not dolomitized diagenetically. Fossil silicification is, moreover, uncommon, 

localized, and usually affects only the brachiopods of 

unit 2. Beds with vast numbers of shells in a mixed 

coral-brachiopod-trilobite assemblage show minimal 

breakage of shells by wave or current action. Articulated brachiopod valves are common and favositid 

heads stand here and there in upright growth positions. The importance of the Rugosa as rock builders 

is greatly overshadowed by the crinoids, massive 

favositids, and brachiopods; the stromatoporoids play 

an insignificant role. No biohermal bodies or incipient 

patch reefs are known within this biofacies; mixed 

faunal deposits with corals appear to have accumulated 

as bands or beds in shallow water without appreciable 

bottom relief.

Subordinate molluscan biofacies occupy shale or 

siltstone interbeds of Nevada Formation, unit 2. 

Clams and large gastropods are abundant locally in 

these interbeds. Also present are the large heavyshelled benthonic tentaculites. Straight-shelled Orthoceros-like cephalopods are fairly common in the 

Breviphrentis biofacies, but the nautiloids and goniatites are quite uncommon.

Coralline beds with mixed faunas of the Breviphrentis biofacies are usually dominated by massive 

favositid heads. Numerically the most abundant 

Rugosa are Breviphrentis, followed in order by Bethanyphyllum, the cystiphylloids, and the digonophyllids. Compound genera like Billingsastraea, Pinyonastraea, and Disphyllum are widely scattered in 

relatively small pockets.

More detailed facies analysis of Nevada Formation, 

unit 2 and the Breviphrentis biofacies will doubtless 

lead to recognition of additional subsidiary biofacies 

within which coral, trilobite, and other benthonic 

associations may be discriminated. Biofacies comparison with the Onondaga Limestone of New York 

which is approximately correlative geologically is 

suggested. Unlike the Onondaga, no patch reefs or 

extensive concentrations of colonial rugose corals were 

recognized in the Breviphrentis biofacies. Oliver's
CORAL SUCCESSION AND EVOLUTION 25

(1968, p. 31) New York Onondaga biofacies studies 

call attention to five facies ranging in taxonomic 

coral diversity from a high figure in patch reefs or 

bioherms to a low figure in bedded coralline deposits. 

Ecologically and taxonomically significant is Oliver's 

(1968, p. 30) observation that environmental factors 

permitting high species diversity in Onondaga coral 

faunas also favored greater variation within the 

studied species.

HEXAGONARIA-SOCIOPHYLLUM BIOFACIES

Rugose coral assemblages of this predominantly 

coralline biofacies in coral zone F occupy dolomitic 

limestone, limestone, and dolomite of Nevada Formation, unit 4. The very great difference of coral faunas 

from those of the Breviphrentis biofacies and coral 

zone D is in harmony with the marked lithofacies 

disparity, and the two are separated stratigraphically 

by the barren zone of Nevada Formation, unit 3 and 

coral zone E. For the first time in the Devonian, 

colonial Rugosa outnumber the solitary forms in most 

localities. The massive Favosites, so abundant in zone 

D, is replaced by less numerous digitate favositids and 

by Heliolites, a tabulate not found in the Devonian 

below zone F. Patchy, discontinuous dolomitization is 

similar to that in Nevada Formation, unit 1 and coral 

zones B and C. Rugose corals and brachiopods of the 

Hexagonaria-Sociophyllum biofacies are strongly silicified within the dolomitic carbonate. Rugose corals 

together with stromatoporoids in situ are major carbonate builders in loci of thick-bedded massive and 

more or less dolomitic limestone. Although this biota 

is potentially reef forming, no true bioherms of large 

dimensions were observed. The abundant rugose coral 

genera in this biofacies at Lone Mountain and in the 

Roberts Mountains are the colonial Hexagonaria, 

Taimyrophyllum, Cyathophyllum, Utaratuia, and 

Sociophyllum together with the solitary genera Cystiphylloides, Mesophyllum, Digonophyllum, and Acanthophyllum.

PHILLIPSASTRAEA BIOFACIES

The Phillipsastraea biofacies of Late Devonian age 

occurs in relatively clean medium- to dark-gray limestones in the upper part of the Devils Gate Limestone 

and coral zone I. These upper beds have not been 

diagenetically dolomitized, although the lower part of 

the Devils Gate has been locally dolomitized in very 

irregular fashion. Scattered concentrations of abundant corals in the Devils Gate area and White Pine 

district consist mainly of the colonial Phillipsastraeidae in bedded deposits. Stromatoporoids, so 

abundant in stromatoporoid biofacies of coral zone H 

in the lower Devils Gate Limestone, become inconspicuous in the Phillipsastraea biofacies. The tabulate 

corals are almost unrepresented. Most of the diagnostic Rugosa of this biofacies are classified in the 

single family Phillipsastraeidae and include the 

colonial Pachyphyllum, Phillipsastraea, and Phacellophyllum together with the solitary MacGeea.

At Lone Mountain the upper beds of the Devils 

Gate Limestone of this biofacies are not exposed above 

the stromatoporoid-rich limestones in coral zone H. 

They are present at Devils Gate and southward, but 

they may best be observed in the White Pine district, 

especially near the Belmont mine mill and in the low 

range west of Monte Cristo.

CORAL SUCCESSION AND EVOLUTION AS

RELATED TO GEOLOGIC CHANGE AND

FAUNAL MIGRATION

Nearly 4,000 feet of geosynclinal marine strata in 

the central Great Basin records much of the Devonian 

Period, enclosing an unusually full succession of 

rugose corals (fig. 9). Stratigraphically and in terms 

of coral evolution, this succession is, however, far from 

complete. Limestone or dolomite layers with abundant 

corals and other fossils are separated by barren layers 

of similar rock; between the coral-bearing units are 

entire members hundreds of feet thick which yield 

almost no Rugosa. How these paleontologic gaps relate 

to physical and biologic events of Devonian history 

in the Great Basin province, and more generally in 

the Cordilleran belt is a matter of some interest. 

Worthy of consideration in this context is the possible 

influence of a normal sea bottom ecologic succession 

upon a coral biota. More important is the influence of 

contemporaneous physical geologic change in the 

surrounding terrane upon the accumulating layered 

sediments.

Lateral spread and migration of the faunas was 

influenced or compelled by a combination of such 

physical and biologic factors. Corals of a particular 

sedimentary facies were adapted to a sea floor environment which also affected or determined the character 

of the entombing sediment. As with benthonic marine 

organisms generally, the intricate fabric of rugose coral 

evolution entails factors of species spread and migration, adaptation to available ecologic niches, isolation 

of gene pools, mutation or gene recombination, and 

above all survival or extinction by natural selection. 

Classic paleontology sheds little light upon the underlying causative factors of evolution. Although the 

fossil record is demonstrably incomplete, the study of 

rugose corals from the more continuous sections of 

this region makes possible the outlining of the broad 

evolutionary trends and patterns, especially in the
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CORAL SUCCESSION AND EVOLUTION 27

hardier, more persistently adaptive lineages. In basins 

of fairly continuous carbonate accumulation such as 

these, coral lineages which survived for long periods 

of progressive in situ evolutionary change may theoretically be distinguished from less adaptive stocks, 

the fossil record of which is in this area brief. In the 

central Great Basin Devonian, persistent lineages of 

this kind are probably recognizable in the Laccophyllidae withSyringaxon,in the endemic Papiliophyllinae, 

in the Bethanyphyllidae, and among the Siphonophrentinae, as represented by Breviphrentis. Abundant and ranging through considerable thicknesses of 

strata, these evolutionary lineages may lend themselves eventually to bed-by-bed quantitative evolutionary investigation.

The physical and biologic environment within a 

given area of shallow Devonian sea bottom was undergoing constant fluctuating change. As recorded by 

sediments and their entombed fossils, these changes 

were to some extent repetitive and perhaps in some 

instances cyclical. Exclusive of physical geologic factors, the normal ecologic succession of organisms upon 

a growth site should be considered. Metabolic and 

secretional activities of ecologically interdependent 

colony-building organisms are capable of so modifying 

their habitat as to inhibit continued occupancy by 

the same biota. Hence upward disappearance of a coral 

assemblage from a particular bed of in situ growth is 

explainable. The succession of evolving coral assemblages in Nevada Formation, unit 2 provides good 

examples of this situation.

Physical geologic changes have acted through 

geologic time to keep the currents of life moving, 

indirectly affecting the course of organic evolution. 

Uplift, subsidence, mountain building, and erosion are 

primary driving forces of biologic change. Evidences 

of contemporaneous crustal deformation are difficult 

to recognize and interpret within the stratified 

Devonian rocks of the Great Basin. Although there 

are no indications of contemporaneous folding or tilting, it is clear that upward sedimentary rock changes 

were in some part controlled by crustal deformation 

affecting the shelving sea bottoms and the adjacent 

lands. Crustal movements brought local shifts in 

marine and atmospheric circulation, changing the 

direction and load capacity of debris-carrying currents. 

Landward debris sources were modified to affect the 

offshore character of accumulating marine deposits.

Convincing field evidences of unconformity within 

the central Great Basin Devonian System are few 

indeed, there being no recognized angular stratigraphic 

discordances, major stratigraphic gaps, or erosion 

intervals. Nonetheless there are many abrupt vertical

lithologic changes in seemingly parallel strata, accompanied by sudden faunal change. It is not, therefore, 

unlikely that significant disconformities have been 

overlooked. Within Nevada Formation, unit 2 may be 

observed many repetitive vertical lithologic changes 

with concurrent faunal change. In this unit fossil-rich 

limestones characteristically alternate with silty and 

argillaceous beds containing a somewhat different 

fauna. Some of these changes appear to be more or 

less cyclical and may be partly the result of ecological 

succession. More extreme rock changes in unit 2 

followed local introduction of the white quartzitic 

sands, bringing a facies usually barren of fossils. Sudden and permanent faunal change accompanied the 

great rock change at the gradational boundary separating unit 2 from unit 3; at this horizon the rich zone 

D faunas disappear completely. Less abrupt is the 

appearance of coral zone F assemblages above the 

contact of unit 4 with the barren zone of unit 3.

CORAL CHANGES AT CLOSE OF SILURIAN

Resumption of limestone or dolomite deposition in 

Early Devonian time was accompanied by abrupt and 

profound change in corals (fig. 9). In the Great Basin 

province, few rugose corals ancestral to Devonian 

forms are known from either dolomite or limestone 

facies of the Late Silurian. Change in the character 

of carbonate deposition at this systemic boundary is 

not great in several of the studied sections; physical 

evidences of disconformity observable on the outcrop 

are local and have been found thus far only in the 

dolomite facies.

It is reasonable to link the major paleontologic 

changes at the Silurian-Devonian boundary with the 

"Caledonian Revolution," a prolonged series of compressive distubances which deformed Silurian and 

older rocks in widely separated parts of the Northern 

Hemisphere. Although angular discordance or indication of Caledonian compressive deformation is 

unknown in the Great Basin, the mapping of the 

Silurian-Devonian contact in several areas gives evidence of disconformity. For example, in the Gold Hill 

mining district of western Utah (Nolan, 1935, p. 18), 

a disconformity is reported as separating Silurian 

Laketown Dolomite from Early Devonian Sevy Dolomite; this discordance is also recognized in the 

Confusion Range (R. K. Hose, written commun., 

1958). Similarly, in the Diamond Mountains and in 

the Mahogany Hills near Eureka, Nev., a mappable 

disconformity separates Silurian Lone Mountain Dolomite from the Lower Devonian beds, according to 

Nolan, Merriam, and Williams (1956, p. 38). The 

disconformity has not been traced laterally to the
28 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Lone Mountain reference section, where the Silurian 

dolomite is overlain with seeming concordance by 

dolomite at the base of Nevada Formation, unit 1. 

Unrecognized at this boundary zone are beds of the 

Early Devonian Beacon Peak Dolomite Member, 

which would be expected here, and the coral faunas 

of Devonian coral zones A and B. Although the Lone 

Mountain Dolomite-Nevada Formation boundary 

appears gradational, the possibility of a depositional 

gap here has not been entirely eliminated.

The beds in the Lone Mountain Dolomite Silurian 

facies are characterized by the halysitids, pycnostylid 

Rugosa, and Entelophyllum; none of these corals 

reappear in the Devonian. Better understood are 

boundary faunal changes in limestone facies of the 

Silurian Roberts Mountains Formation where it is 

overlain without recognized depositional break by the 

Early Devonian Rabbit Hill Limestone. For example, 

in Coal Canyon, northern Simpson Park Mountains, 

the limestones of Late Silurian coral zone E contain a 

large Gotlandian type coral assemblage with Mycophyllinae, Kodonophyllidae, Chonophyllidae, Pycnostylidae, Goniophyllidae, and Endophyllidae. The 

overlying Rabbit Hill has yielded only the solitary 

Syringaxon in this area. A single occurrence of the 

endophyllid Australophyllum is known elsewhere in 

the Rabbit Hill. Clearly the rather unimpressive 

lithologic change at the Silurian-Devonian boundary 

is not commensurate with the profound change in the 

coral fauna.

LATE SILURIAN AND DEVONIAN BURSTS OF 

RUGOSE CORAL EVOLUTION

From Late Silurian through Late Devonian time, 

the history of the Rugosa in the Great Basin seaways 

was characterized by bursts of proliferation and evolutionary differentiation, separated by intervals during 

which the Rugosa were restricted in taxonomic diversity or poorly represented. Such bursts of coral activity 

took place: (1) in Late Silurian coral zone E of the 

upper Roberts Mountains limestones (fig. 9); (2) in 

Early to early Middle Devonian coral zone D of 

Nevada Formation, unit 2; (3) in Middle Devonian 

coral zone F of Nevada Formation, unit 4; and (4) in 

Late Devonian coral zone I of the upper Devils Gate 

Limestone. Each burst involved taxonomically different coral groups which reached their peak in 

evolution at that time (see table 1). During the 

intervals of coral zones A and B, rugose corals were 

little differentiated although they were locally numerous; the rugose corals of zone C were becoming more 

diverse. Coral zones E and H show a poor representation of the Rugosa; the beds carrying coral zone G are

TABLE 1. Evolutionary coral bursts in the Great Basin during 

the Silurian and Devonian Periods

Great Basin 

Devonian 

coral zone

Rugose corals 

unknown in latest 

Devonian of 

Great Basin

I

H

G

F

E

D

C

B

A

Silurian 

coral zone 

E

Coral burst

I

F

D

SE

European

stage

Famennian

Frasnian

Givetian

Eifelian

Emsian

Siegenian

Ludlovian

Age

1 Middle Late -» 9 1

* 

a

1

Devonian

Silurian

introductory to an interval of dolomite facies unfavorable to the Rugosa.

Following the Late Devonian burst of coral zone I, 

the Rugosa declined and seemingly disappeared in this 

region before the close of the Devonian Period. When 

Rugosa reappear in the Mississippian Joana Limestone following an interval of extinction, they are 

represented by entirely different families which are 

unknown below this horizon.

ZONE D CORAL BURST

With deposition of impure argillaceous limestones 

in Nevada Formation, unit 2 of the Antelope-Roberts 

Mountains facies belt came the zone D burst of rugose 

coral differentiation. This episode opened rather 

abruptly after the interval of coral zones B and C in 

which solitary Halliidae were dominant and coral 

faunas were far less diverse. Siphonophrentinae, 

Cystiphylloidae, and Bethanyphyllidae peaked in 

coral zone D. Colonial genera, mainly Disphyllidae, 

appeared, but these are subordinate to the solitary 

genera. Large solitary Digonophyllidae arrived at this 

time, reaching their acme later in coral zone F, 

together with the Disphyllidae. Benthonic life was in 

general favored by the shallow sea environment of 

zone D; Tabulata were numerous and taxonomically 

diverse, as were brachiopods, trilobites, and Mollusca. 

In terms of genera and species, the zone D invertebrate faunas are overall the largest of the Great Basin 

Devonian.

Except for poorly preserved disphyllids and solitary 

coral species assigned to Cyathophyllum, the next 

higher interval of Nevada Formation, unit 3 (coral 

zone E) yields few Rugosa, and none from coral zone 

D are known to carry over into this barren zone.

ZONE F CORAL BURST 

Rugosa again became numerous and taxonomically
AGE AND CORRELATION OF CORAL ZONES 29

diverse within the interval of Nevada Formation, unit 

4. Colonial genera are abundant for the first time in 

the Devonian, being concentrated in growth sites 

where corals predominate and where the limestones 

are partly dolomitic. Away from these dolomitic limestone loci, corals are scarce or absent, as in rocks of 

this interval in the Diamond Mountains facies belt. 

Families and subfamilies are not so numerous as in 

coral zone D, but the Disphyllidae have differentiated 

generically and the large solitary digonophyllids are 

at their evolutionary peak. This was also an interval 

of great rugose coral proliferation through the northern 

part of the Cordilleran Belt in western Canada.

ZONE I CORAL BURST

Marine environments in which were deposited the 

coarsely saccharoidal dolomite and thick-bedded stromatoporoid limestones of zones G and H were evidently not conducive to rugose coral growth. When 

next the Rugosa appear in force within Late Devonian 

coral zone I, nearly all of the subfamilies and genera 

of zones D and F have disappeared, their places being 

taken by Phillipsastraeidae and one or two other subordinate families. Ancestry of the Phillipsastraeidae is 

unknown. In coral zone F the genus Taimyrophyllum 

superficially resembles Pachyphyllum, but a study of 

internal structure does not bear out a genetic connection.

In the upper part of the Devils Gate Limestone, 

Phillipsastraeidae of zone I occur in scattered concentrations ecologically like those of coral zone F and 

do not occupy laterally persistent coral beds. Similar 

scattered loci of zone I coral proliferation occur in the 

northern part of the Cordilleran Belt in Canada and 

Alaska; this family also characterizes the Frasnian of 

other continents.

AGE AND CORRELATION OF GREAT BASIN

EARLY AND EARLY MIDDLE DEVONIAN CORAL

ZONES

Early Devonian rocks were made known in the 

central Great Basin during the geologic reconnaissance of the Roberts Mountains in 1934. At that time 

beds containing Costispirifer similar to the eastern 

Oriskany C. arenosus were discovered in a badly 

faulted section. Stratigraphic studies in the MonitorSimpson Park belt during the 1950's led to the recognition of the Helderbergian age of the Rabbit Hill 

Limestone (Merriam, 1963, p. 42-44). A review by 

House (1962, p. 252) of cephalopod evidence in the 

American Devonian led to the conclusion that the 

"Spirifer pinyonensis zone" of Merriam (1940, p. 51), 

initially called Middle Devonian, may also be of Early

Devonian age. In recent years research upon corals 

and brachiopods allied to a broad program of geologic 

mapping and stratigraphy in the Sulphur Spring 

Range and areas adjoining the Eureka mining district 

has shed light upon age and correlation problems.

AGE AND CORRELATION WITH DISTANT REGIONS

CORAL ZONE A

The Rabbit Hill rugose coral genera Syringaxon and 

Australophyllum, because they occur in both Silurian 

and Devonian strata of distant regions, have at present 

no definitive correlation value. However, the Helderberg age of this formation is convincingly borne out 

by associated brachiopods among which are: Levenea 

cf. L. subcarinata (Hall), Howellella cf. H. cycloptera 

(Hall), Anastrophia cf. A. verneuili (Hall), and 

Orthostrophia cf. O. strophomenoides (Hall). Other 

fossils of Helderbergian affinity are trilobites assigned 

to Leonaspis and two species of the tabulate coral 

Pleurodictyum. The closest known relatives of the 

Rabbit Hill fauna thus occur in rocks of Helderbergian 

age in New York, Oklahoma, and Tennessee (Merriam, 1963, p. 43). Costispirifer foreshadowing the 

New York Oriskany C. arenosus appears in higher 

beds of the Rabbit Hill.

CORAL ZONE B

The large and abundant Kobeha walcotti of the 

Halliidae occurs here without other rugose coral associates; like Papiliophyllum of succeeding coral zone 

C, Kobeha seems to be endemic in the Great Basin 

Province and, as such, is of no distant correlation value 

at present. Approximate Oriskany or Siegenian age is 

indicated by the associated brachiopod fauna containing Costispirifer cf. C. arenosus and Gypidula with 

affinity to G. coeymanensis. The "Trematospira fauna" 

of Merriam (1940, p. 50), with C. arenosus and Trematospira coo peri, is that of coral zone B. First recognized within the intensely faulted terrane on the west 

side of Roberts Creek Mountain (Merriam, 1940, p. 

50-52, pi. 1), the fossils of this Oriskany interval have 

in recent years been discovered during the mapping of 

these deposits southward from the Old Whalen mine 

area in the Sulphur Spring Range. In this range 

several of the more complete stratigraphic sections 

include all Devonian coral zones from A through D 

in order.

CORAL ZONE C

The endemic halliid coral Papiliophyllum succeeds 

Kobeha of underlying coral zone B. A larger coral 

fauna accompanies Papiliophyllum, including the halliid Odontophyllum meeki and Breviphrentis kobehensis of the subfamily Siphonophrentinae. The Onondaga
30 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

and Hamilton Formations of eastern North America 

are characterized by members of this subfamily 

which appears earlier in the New York Coeymans 

(Oliver, 1960a, p. 87). Siphonophrentis (Breviphrentis) kobehensis is less similar to Siphonophrentis 

variabilis Oliver of the Coeymans than is Siphonophrentis (Breviphrentis) invaginatus of coral zone D. 

Eastern species of Odontophyllum (Stumm, 1948b) 

differ specifically from O. meeki of coral zone C and 

are reported to be of Middle Devonian age.

A preliminary appraisal of brachiopods in coral zone 

C, the Acrospirifer kobehana zone (Merriani, 1940, 

p. 50-53), indicates that a provisional correlation with 

the European Emsian appears reasonable.

CORAL ZONE D

Coral zone D brackets the interval of Nevada Formation, unit 2 in the Lone Mountain reference section 

and corresponds to the "Spirifer pinyonensis zone" of 

earlier usage (Merriam, 1940, p. 53). Vertical changes 

in the rugose corals make it possible to further subdivide this zone into subzones DI, D2 , and D3 in 

ascending order. Initial appraisal of zone D coral, 

brachiopod, and trilobite assemblages by the writer 

in the 1930's (Merriam, 1940) led to a Middle Devonian age assignment, although underlying strata of 

coral zone C, or the "Spirifer kobehana zone," were 

at that time considered Lower Devonian. The beds 

of Nevada Formation, unit 2 were later assigned by 

others to Early Devonian (Johnson, 1962a; 1962b; 

1966b). Paleontologic support of the Early Devonian 

interpretation comes in large degree at present from 

the ammonoid succession as reviewed by House (1962, 

p. 247-284). A well-preserved goniatite described by 

A. K. Miller (1938, p. 46, pi. 6, fig. 1) as Agoniatites 

nevadensis was collected by the writer in 1933 from 

a locality near M29 on the southwest side of Lone 

Mountain. The beds yielding the goniatite are Nevada 

Formation, unit 2, within the range of coral zone D2 ; 

associated fossils include Billingsastraea nevadensis. 

This fossil is the only fairly complete goniatite known 

from the lower part of the Nevada Formation; fragmentary material has been found at Bush Creek in 

the Roberts Mountains and at Combs Peak by Walcott. Restudy by House of the Miller type and the 

only specimen of A. nevadensis, in the light of Erben's 

(1960) revision of European Early Devonian goniatites, led to reassignment of this species to Teicherticeras, a genus considered indicative of the Emsian.

The conclusions of House and of Johnson regarding 

the Early Devonian assignment of the entire "Spirifer 

pinyonensis zone" or Nevada Formation, unit 2 are 

only in part borne out by results of these coral investigations. In harmony with an Early Devonian assignment, the coral fauna of subzone DI includes an 

advanced Papiliophyllum elegantulum called subspecies d. Upward in the section the preponderant 

Rugosa are Siphonophrentinae of the species Breviphrentis invaginatus, making these faunas similar to 

the Onondaga faunas. The eastern Onondaga is analogous, as it has an abundance locally of Siphonophrentinae; but the forms belonging to this family are mainly 

Heterophrentis. Among other Rugosa of subzone D2 , 

Billingsastraea nevadensis shares many features with 

the late Early Devonian Billingsastraea affinis (Billings) of the Grande Greve, as refigured by Oliver 

(1964).

Other Rugosa of subzone D2 , the Family Bethanyphyllidae, Family Digonophyllidae, and the Hexagonaria, are less in accord with an Early Devonian age; 

these Rugosa represent elements largely known from 

starta of Middle Devonian age. The Halliidae, which 

here characterized the Lower Devonian beds of coral 

zones B and C, are uncommon above subzone DI, their 

places being taken by the Bethanyphyllidae.

Digonophyllidae appear in subzone D2 with Mesophyllum and Zonophyllum. The Digonophyllidae in 

these beds are greatly outnumbered by Siphonophrentinae and Bethanyphyllidae Zonophyllum haguei is 

at least specifically distinct from Z. duplicatum 

Wedekind of the early Middle Devonian "Nohner 

Schichten" in the Eifel district, Germany. In Europe 

and the Cordilleran Belt of North America, Digonophyllidae, are especially characteristic of the Middle 

Devonian (Eifelian), peak in Great Basin coral zone 

F. Bethanyphyllidae in this area are seemingly confined to subzones DI and D2 . Hexagonaria of subzone 

D2 is here placed in a new subgenus Pinyonastraea. 

True Hexagonaria is known here only in coral zone 

F, where it is locally the abundant genus.

Passing upward at Lone Mountain into subzone D3 

and the uppermost beds of Nevada Formation, unit 

2, the Digonophyllidae are represented by the structurally complex Mesophyllum (Arcophyllum) kirki, a 

species resembling Arcophyllum in the Middle Devonian (Eifelian) in the Eifel district, Germany. Arcophyllum kirki has its nearest relatives in the zone of 

"Cosmophyllum dachsbergi" at Gerolstein, Eifel district (Wedekind, 1925, p. 71); following Wedekind's 

interpretation, this horizon is well within the Middle 

Devonian.

In summary, these data appear to support the 

ammonoid evidence for the Early Devonian age of 

some part of the "Spirifer pinyonensis zone" of prior 

usage, or Nevada Formation, unit 2. However, Middle 

Devonian rugose coral elements are present in sub-
AGE AND CORRELATION OF CORAL ZONES 31

zone D2 together with those of latest Early Devonian, 

and the digonophyllid Mesophyllum (Arcophyllum) 

kirki of subzone D3 is, in the absence of conflicting 

data, considered Middle Devonian. Accordingly, the 

rocks of Nevada Formation, unit 2 and coral zone D 

are here referred to as Early and early Middle 

Devonian. Conclusions regarding age and correlation 

of subzone D2 should await detailed description and 

analysis of the large brachiopod and trilobite faunas, 

together with the conodonts and tentaculites of this 

interval. Likewise the ammonoid evidence is meager 

at present. Rugose coral data suggest that the subjective Lower-Middle Devonian boundary lies figuratively 

within a "gray band" of coral zone D and Nevada 

Formation, unit 2.

CORRELATION WITH OTHER AREAS IN THE 

CORDILLERAN BELT

Within the Cordilleran Belt, Early and early Middle 

Devonian rugose coral-bearing rocks are almost unknown outside the Great Basin. In the Canadian 

Northwest, the oldest Devonian coral-bearing strata 

thus far elucidated are the Eifelian or Givetian Hume 

and Nahanni Formations, which contain faunas correlative with Great Basin coral zone F. Billingsastraealike colonial Rugosa in these Middle Devonian formations are similar to Nevada Formation, unit 4 (coral 

zone F) species. However, Billingsastraea verrilli 

(Meek) of the Hume (Pedder, 1964), though specifically distinct, possesses features of the older B. 

nevadensis of coral subzone D2 . Strata beneath the 

Hume have yielded goniatites of the genus Teicherticeras that are indicative of a possible Emsian age 

(House and Pedder, 1963, p. 491); these older beds 

are not known to contain Rugosa. The expected distribution of the Devonian System throughout the 

Cordilleran Belt makes it seem unlikely that strata 

correlative to Rabbit Hill and Oriskany of Lower 

Devonian age are absent from these northerly regions. 

Possibly the strata are represented as relatively unfossiliferous dolomites below the Hume and the Nahanni 

Formations.

In Alaska Early Devonian rocks are likewise nearly 

unknown. The well-understood Paleozoic sections of 

the southeast Alaska panhandle include a very thick 

Silurian column, and the Middle and Upper Devonian 

are well represented, but the presence of Early 

Devonian has not as yet been established.

Of interest in this connection is the occurrence of 

glacial cobbles in the Chewelah area of Stevens 

County, northeast Washington, that contain a fauna 

resembling the Rabbit Hill Helderberg (F. K. Miller, 

oral commun., 1965). Among the fossils are Syringaxon sp., Pleurodictyum cf. P. lenticularis (Hall), 

Levenea cf. L. subcarinata (Hall), and a species of 

Leptocoelia.

NORTH-CENTRAL GREAT BASIN 

NORTHERN SULPHUR SPRING RANGE

The Nevada Formation extends northward in the 

Sulphur Spring Range through the Mineral Hill district, where local names have been applied to lithologic 

subdivisions (Carlisle and others, 1957; Johnson, 

1962b). Fossil collections show that Devonian coral 

zones B, C, and D are present. Zone B strata in 

McColley Canyon contain a variety of Kobeha differing somewhat from typical K. walcotti. West of the 

Old Whalen mine, lower beds of Nevada Formation, 

unit 1 carry the Costispirifer arenosus brachiopod 

assemblage but have not yielded Kobeha; these Costisipirifer beds are calcarenites with quartz grains like 

those in zone B which contains Kobeha and Costispirifer in the southern Sulphur Spring Range. Beacon 

Peak Dolomite Member with the Rabbit Hill fauna 

and coral zone A has not been identified in the 

northern part of this range.

SOUTHERN TUSCARORA MOUNTAINS

The northernmost known occurrence of Lower 

Devonian beds in the Great Basin is at Maggie Creek, 

8 miles northwest of Carlin, Nev. in the southern 

Tuscarora Mountains. Limestones of Rabbit Hill age 

at this locality contain a coral-brachiopod fauna 

including Syringaxon, Pleurodictyum, and a Billingsastraea-like colonial coral. This is one of two occurrences of a colonial rugose coral in Devonian coral 

zone A. The Helderbergian limestones at Maggie 

Creek are underlain by the Roberts Mountains 

Formation.

CORTEZ MOUNTAINS

In the Cortez Mountains, strata carrying a Rabbit 

Hill Helderbergian fauna occur 1.5 miles southeast 

of Mount Tenabo summit (locality M1083). Pleurodictyum together with abundant Syringaxon foerstei 

indicate Devonian coral zone A.

WEST-CENTRAL GREAT BASIN 

TOIYABE RANGE

Devonian limestone and calcareous shale correlative 

with the Nevada Formation have recently been discovered by R. H. Washburn (written commun., 1968) 

on the west side of the Toiyabe Range 15 miles southwest of Austin, Nev. (Stewart and McKee, 1968). 

Exposures in the Reeds Canyon vicinity (locality 

M1151) have yielded the compound Billingsastraea'?
32 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

sp. T and a large Hexagonaria-like form. Provisionally 

compared to Billingsastraea and Hexagonaria (Pinyonastraea) of coral subzone D2 , a restudy suggests 

that the Billingsastraea-like form may be more closely 

allied to undescribed Taimyrophyllum of coral zone 

F. Accordingly, the age of this fauna remains somewhat uncertain, but it is more likely Middle than Early 

Devonian. As the Toiyabe Devonian section is thick 

and fairly continuous upward from underlying graptolitic Silurian, it is not improbable that other Devonian 

fossil horizons, including the Rabbit Hill, may eventually be discriminated here.

SOUTH-CENTRAL GREAT BASIN 

HOT CREEK RANGE

Corals suggesting zones C and D have been collected in little-known Devonian beds of the Hot Creek 

Range. Fragmentary specimens of probable Papiliophyllum come from locality M1066 in Hot Creek 

Canyon (H. E. Cook, written commun., 1966) and an 

Aulacophyllum-like coral comes from the Morey Peak 

vicinity (H. W. Dodge, Jr., written commun., 1968). 

Coral faunas of zone F are better known in this range, 

suggesting that most or all of the Nevada Formation 

interval is represented.

SOUTHERN GREAT BASIN

Southern Great Basin rugose coral-bearing beds of 

Early and early Middle Devonian ages are known in 

the Ranger Mountains, the Desert Range, the northern 

Panamint Mountains, and the Funeral Mountains. 

The occurrence of faunas at each place indicates a 

southerly extension of the seaway in which the lower 

part of the Nevada Formation of central Nevada was 

deposited. No fossils with certainty as old as the 

Rabbit Hill Helderbergian have thus far been discovered in these southern mountain ranges.

RANGER MOUNTAINS

Dolomite, limestone, and quartzite, some 3,000 feet 

thick and including beds rich in rugose corals, occur 

in the Ranger Mountains and other ranges bordering Frenchman Flat, Nye and Clark Counties, Nev. 

(Johnson and Hibbard, 1957, p. 350-356; F. G. Poole, 

written commun., 1964). A quartzite about 90 feet 

thick in the lower part of this section probably corresponds to the Oxyoke Canyon Sandstone Member of 

the Eureka district 200 miles north. The fauna occurring in limestone beds 170 feet below the quartzite 

and about 10 feet above a thick dolomite sequence 

includes Siphonophrentis (Breviphrentis) invaginatus, 

Zonophyllum sp., and Papiliophyllum elegantulum

subsp. d. Other fossils identified from this horizon 

are Chonetes macrostriata, Dalmanites meeki, and a 

Strophonella. The rugose corals suggest that this 

fauna represents the lower part of coral zone D and 

probably correlates with Di of the Lone Mountain 

column because of the presence of the advanced 

Papiliophyllum. It is not unlikely that the underlying 

dolomites include earlier Devonian as well as a Lone 

Mountain Dolomite equivalent.

DESERT RANGE

Devonian beds at the south end of the Desert 

Range, 30 miles east of Frenchman Flat (Locality 

M1057, C. R. Longwell, oral commun., 1958) include 

a fauna with Chonetes macrostriata, Strophonella, 

Acrospirifer sp. and an incomplete corallum of either 

Siphonophrentis (Breviphrentis) or Sinospongophyllum. This assemblage suggests lower coral zone D, 

possibly about the horizon of the Ranger Mountains 

occurrence.

NORTHERN PANAMINT MOUNTAINS

Fossil collections from the upper part of McAllister's 

Hidden Valley Dolomite unit 3b (McAllister, 1952, 

p. 17) in the southern Andy Hills of the Panamint 

Mountains (Marble Canyon quadrangle, Calif.) include Acrospirifer kobehana, Costispirifer arenosus, 

Papiliophyllum elegantulum, and Siphonophrentis 

(Breviphrentis) cf. invaginatus. The spirifers and 

Papiliophyllum indicate Early Devonian in the combined intervals of coral zones B, C, and possible DI. 

The genus Kobeha, which is to be expected with 

Costispirifer arenosus, was not found here. Possibly 

C. arenosus ranges upward in the Panamint Mountains into beds with Acrospirifer kobehana and Papiliophyllum. In the lower part of the Nevada Formation 

of central Nevada, the beds of coral zone B with C. 

arenosus and Kobeha underlie those of coral zone C 

with A. kobehana and Papiliophyllum elegantulum. 

The Panamint beds in question appear to be older 

than those in the Ranger Mountains and those in the 

Desert Range that are discussed above which appear 

to be correlative with the Nevada Formation, unit 2; 

the Panamint Early Devonian strata are correlative 

with Nevada Formation, unit 1 of the Lone Mountain 

section.

FUNERAL MOUNTAINS

The upper part of the Hidden Valley Dolomite in 

the vicinity of Pyramid Peak (Ryan quadrangle, 

Calif.), southern Funeral Mountains, contains coral 

and brachiopod faunas resembling in part those of 

the same dolomite unit in the northern Panamint 

Mountains. Collections from six localities made by 

J. F. McAllister include an abundance of a large
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 33

Meristella similar to M. robertsensis and abundant 

Siphonophrentis (Breviphrentis) invaginatus. Less 

common are Papiliophyllum elegantulum and Aulacophyllum-like forms. These collections do not contain 

Costispirifer arenosus or Acrospirifer kobehana as do 

collections from the upper Hidden Valley of the Andy 

Hills area, northern Panamint Mountains. Absence of 

Early Devonian spirifers suggests that the Funeral 

Mountains faunas may be somewhat younger than 

the horizons of coral zones B and C, possibly being of 

the age of coral zone Di which contains the highest 

Papiliophyllum.

NORTHERN INYO MOUNTAINS

Upper beds of the Vaughn Gulch Limestone at 

Mazourka Canyon, northern Inyo Mountains contain 

a species of Australophyllum similar to A. landerensis 

of the Rabbit Hill. The upper Vaughn Gulch beds are 

underlain by strata of Upper Silurian coral zone E 

that also underlie the Rabbit Hill Limestone at Coal 

Canyon, Simpson Park Mountains. Confirmation of 

Helderbergian age of the upper Vaughn Gulch by other 

fossils has not been achieved, and these beds are presently classified as Late Silurian or Early Devonian.

SYSTEMATIC AND DESCRIPTIVE 

PALEONTOLOGY

CLASSIFICATION OF GREAT BASIN EARLY AND 

EARLY MIDDLE DEVONIAN RUGOSA

Th classification adopted for Great Basin Rugosa 

here described is for the greater part in accord with 

that of Hill (1956), the most comprehensive and 

authoritative classification published thus far. Departures from Hill's arrangement of families and genera 

involve content of certain families, status of subfamilies, and family or subfamily assignment of several 

genera.

A new subfamily, the Siphonophrentinae, is proposed to accommodate certain advanced Streptelasmatidae. Because of uncertainties in the characterization of Zaphrentis, the Family Zaphrentidae is not 

used herein. The Disphyllidae are considered an independent family not closely related to the Phillipsastraeidae, by reason of fundamental differences of 

tissue structure. Stumm's subfamily Papiliophyllinae 

is restored for these seemingly endemic Halliidae; also 

adopted is Stumm's family Bethanyphyllidae, which 

has a wide geographic distribution. For certain Great 

Basin Silurian and Early Devonian species of Australophyllum the Family Endophyllidae is adopted, 

forms which might otherwise have been relegated to 

the ill-defined Spongophyllidae. Stumm's Family Cystiphylloidae is restored to include Cystiphylloides, in

preference to placing it under Mesophyllum of the 

Digonophyllidae.

Order RUGOSA Edwards and Haime 1850

I. Family Laccophyllidae Grabau, 1928

Genus Syringaxon Lindstrom, 1882 

Syringaxon foerstei n. sp.

II. Family Streptelasmatidae Nicholson, 1889

(as Streptelasmidae)

Subfamily Siphonophrentinae new subfamily 

Genus Siphonophrentis O'Connell, 1914 

Subgenus Breviphrentis Stumm, 1949 

(as genus) 

Siphonophrentis (Breviphrentis)

invaginatus (Stumm) 

Siphonophrentis (Breviphrentis)

kobehensis n. sp.

Genus Nevadaphyllum Stumm, 1937 

Nevadaphyllum masoni Stumm

III. Family Kodonophyllidae Wedekind, 1927

Subfamily Kodonophyllinae Wedekind, 1927

Genus Kodonophyllum Wedekind, 1927

(l)Kodonophyllum sp. a

IV. Family Stauriidae Edwards and Haime, 1850 

Genus Dendrostella Glinski, 1957

(as subgenus of Favistella) 

Dendrostella romanensis n. sp.

V. Family Halliidae Chapman, 1893

Subfamily Haliinae Chapman, 1893

Genus Aulacophyllum Edwards and Haime,

1850

Aulacophyllum sp. c 

Genus Odontophyllum Simpson, 1900

Odontophyllum meeki n. sp. 

Subfamily Papiliophyllinae Stumm, 1949 

Genus Papiliophyllum Stumm, 1937

Papiliophyllum elegantulum Stumm

Papiliophyllum elegantulum subsp.

d 

Genus Kobeha new genus

Kobeha walcotti n. gen., n. sp.

Kobeha ketophylloides n. gen., n. sp. 

Genus Eurekaphyllum Stumm, 1937

Eurekaphyllum breviseptatum

Stumm

VI. Family Bethanyphyllidae Stumm, 1949

Genus Bethanyphyllum Stumm, 1949

Bethanyphyllum lonense (Stumm) 

Bethanyphyllum antelopensis n. sp. 

Bethanyphyllum sp. d 

VII. Family Chonophyllidae Holmes, 1887

Genus Sinospongophyllum Yoh, 1937 

Sinospongophyllum sp. d 

Sinospongophyllum sp. e
34 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Sinospongophyllum sp. f 

VIII. Family Endophyllidae Torley, 1933

Genus Australophyllum Stumm, 1949

Australophyllum landerensis n. sp.

IX. Family Disphyllidae Hill, 1939

Genus Disphyllum de Fromentel, 1861 

Disphyllum nevadense (Stumm) 

Disphyllum eurekaensis n. sp. 

Genus Hexagonaria Giirich, 1896

Subgenus Pinyonastraea new subgenus 

Hexagonaria (Pinyonastraea) 

kirki (Stumm) 

Genus Billingsastraea Grabau, 1917

Billingsastraea nevadensis Stumm 

Billingsastraea nevadensis subsp. 

arachne (Stumm) 

Billingsastraea'? sp. T

X. Family Cystiphylloidae Stumm, 1949

Genus Cystiphylloides Chapman, 1893 

Cystiphylloides robertsense 

(Stumm)

Cystiphylloides lonense (Stumm) 

Cystiphylloides sp. d

XI. Family Digonophyllidae Wedekind, 1924

Subfamily Digonophyllinae Wedekind, 1924 

Genus Mesophyllum Schliiter, 1889 

Subgenus Mesophyllum Schliiter, 1889 

Mesophyllum (Mesophyllum) 

sp. b

Mesophyllum (Mesophyllum) 

sp. c

Subgenus Arcophyllum Markov, 1926 

(as genus)

Mesophyllum (Arcophyllum) 

kirki (Stumm)

Subfamily Zonophyllinae Wedekind, 1924 

Genus Zonophyllum Wedekind, 1924 

Zonophyllum haguei n. sp. 

Zonophyllum sp. a 

Zonophyllum sp. b

TAXONOMIC INTERPRETATION OF 

RUGOSE CORAL STRUCTURE

Traditional gross comparative morphology of skeletal features is the basis for revised classification of 

Silurian and Divonian Rugosa here adopted. Diagnostic characters of families and genera are visible to the 

unaided eye or under low magnification. Coral specialists have long sought recognition of minute histologic 

details as reliable taxonomic guides. Structures of this 

kind, visible under a magnification range of 8-20 diameters, have appropriately been called fine structure 

(Kato, 1963). Although of value in the recognition of

certain families, fine structure is somewhat less diagnostic than the well-known minute wall features of 

fusulinids and orbitoids in Foraminifera. Among Great 

Basin Silurian and Devonian Rugosa the trabecular 

groupings are meaningful histologic characters of this 

scale in wall tissue of Tryplasmatidae and Phillipsastraeidae (Ogilvie, 1897; Hill, 1936; Wang, 1950; Kato, 

1963). None of the Early and early Middle Devonian 

species here described disclose these minute features 

convincingly. Silicification and other forms of additive 

chemical change have in most instances destroyed 

original histologic structures or left them but vaguely 

discernible in thin section through a veil of secondary 

crystallization. Ideally a biologic classification based 

on gross and fine morphology might be supplemented 

by biochemical analysis of such organic traces as may 

have survived the fossilization processes.

At species and infraspecies levels, biometrics and 

statistical analysis of variation have a definite place 

in coral taxonomy (Oliver, 1968). In some instances 

statistical techniques may also be of value in the separation of convergent and homeomorphic genera, especially those of rather simple structure. Quantitative 

studies of this kind always require adequate random 

sampling of abundant in situ material of undoubted 

field association. In only a few species are the collections herein described of Early and early Middle 

Devonian Rugosa adequate at present to justify the 

quantitative approach. At this stage of Great Basin 

coral research, taxonomy at the genus level is of prime 

importance and species discrimination a refinement 

largely for the future.

The basic solitary rugose coral skeleton is an upsidedown conical platform strengthened internally by horizontal, vertical, and diagonal supports and provided 

externally with attaching processes. Within the 

Rugosa generally there are limitless possibilities of 

taxonomic subdivision using combinations of the more 

stable megascopic exterior and interior characters. The 

structural complexity varies from the simple Stauriidae to the very complex interior patterns of the 

Digonophyllidae.

EXTERIOR CORALLUM FEATURES

Exterior surface characteristics of the rugosan corallum have been adequately defined by Hill (1935; 1956) 

and require little elaboration in this regard. Among the 

solitary Devonian taxa here described, surface features 

of taxonomic value given special attention are: (1) 

median longitudinal profile of the modified cone, with 

or without tendency to become elongate-cylindrical at 

maturity; (2) strength and abundance of rugae; (3) 

frequency and spacing of rejuvenescence flanges; (4)
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 35

accentuation of septal grooves; and (5) development 

of connecting or attachment processes.

Among colonial Rugosa generally the exterior features of taxonomic value are growth habit or pattern, 

which range from loosely fasciculate or bushy to massive cerioid. In the compact cerioid forms, shape and 

size of the free-growing head are significant, as is the 

extent of holothecal investment compared to area 

occupied by the distal or calicular surface. The open 

bushy forms reveal externally such features as lateral 

and calice reproductive offsets and lateral connections, 

which may be tubular as in Pycnostylidae or highly 

irregular attachment excrescences like those of Entelophyllum.

Some of these gross external characteristics may be 

viewed as adaptive, or they may be influenced by environmental conditions, as, for example, the external 

shape of a coral head.

Calice features are part of the corallum exterior, but 

these also disclose much of the internal structure and 

the character of reproductive offsets. The individual 

calice profile in median longitudinal section reveals 

calice shape and depth; such profiles may be V-shaped, 

U-shaped, inverted bell-shaped, or squared. Other 

calice features are: calice pit, median boss, distal calice 

platform, and reflection of calice rim all external 

characters of taxonomic importance. Symmetry of the 

calice viewed transversely is that of an adult transverse 

section and shows to advantage the bilateral and quadrilateral features manifested by cardinal, counter, and 

alar septa. Departures from complete radial symmetry 

are clearly shown by pinnate arrangements of septa 

contiguous to a cardinal fossula, present in many genera. Septal edges projecting in from the sides and floor 

of the calice may be denticulate or entire. The calice 

rim may be narrow, acute, and unreflected as in 

Syringaxon or Kobeha; in Mucophyllum, which has a 

mushroom-shaped corallum, the calice rim is strongly 

reflected, forming a wide upwardly convex marginal 

platform. Digonophyllidae that have a wide dissepimentarium also have correspondingly wide calice brims 

or platforms with a conical median pit.

Compact thamnastraeoid and aphroid colonial genera like Billingsastraea, although lacking a corallite 

wall, commonly show false surficial walls or raised 

intercalice polygonal ridges on a calicular surface; 

other variants of this genus suppress the false wall but 

develop elevated median cones, as in PachyphyHum. 

Such exterior calicinal features do not seem to have 

more than subgeneric or species value. Characters of 

the outer corallite wall, septa, biologic symmetry, and 

reproductive offsets visible in the calice are treated 

elsewhere.

Some normally solitary Rugosa are closely related 

genetically to colonial species, calling for taxonomic 

evaluation of growth habit in this respect. Among normally solitary taxa usually classified as Zaphrentidae, 

the genus Heliophyllum, as demonstrated by Wells 

(1937), produces compound adaptive groupings of fasciculate to astraeoid form. Other families which include 

closely related taxa of solitary to colonial habit are the 

Tryplasmatidae and the Phillipsastraeidae. Among the 

last, solitary Macgeea is commonly associated with 

Phillipsastraea, P achy phy Hum, or Phacellophyllum, 

and it seems not unlikely this genus is quite polyphyletic, possibly a solitary differentiate, or even a manifestation of dimorphism. Among corals described in 

this report, there are partly phaceloid individuals of 

otherwise cerioid Pinyonastraea.

TAXONOMIC EVALUATION OF CORALLUM 

INTERIOR STRUCTURES

Chief internal structures of the rugose coral are 

septa, tabulae, dissepiments, axial structures, and 

stereoplasmic deposits. The septa and axial structures 

are vertical elements, the tabulae are horizontal, and 

the dissepiments are diagonal elements. Complex, patterns consisting of combinations of these elements lend 

themselves to taxonomic evaluation.

SEPTA

In terms of fine structure, the two principal types 

of rugosan septa are lamellar and acanthine. Somewhat 

unsatisfactory attempts have been made (Wang, 1950) 

to evaluate trabecular patterns of lamellar septa for 

taxonomic purposes (Kato, 1963). In general these histologic features are too poorly understood and the preservation of most Rugosa unsatisfactory for taxonomic 

evaluation by this character. No Tryplasmatidae with 

acanthine trabecular spines were found in the Great 

Basin Early and Middle Devonian.

Of the eleven families of Great Basin Rugosa here 

defined, nine possess lamellar septa; the remaining two, 

Cystiphylloidae and Digonophyllidae, do not have 

complete and continuous lamellar septa and are dealt 

with separately.

Septum characters of special systematic importance 

are: (1) septal count, (2) relative lengths of major 

and minor septa, (3) vertical continuity of septa, (4) 

radial continuity of septa, and (5) presence of true 

septal carinae. Septal thickening by stereoplasm is 

considered separately.

Evaluation of septal and other interior characteristics at the species level is effectively expressed quantitatively by use of ratios, of which that involving 

major septum count relative to mature corallite diam-
36 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

eter is the most widely used (Oliver, 1960a, 1968; Rozkowska, 1960).

Within the mature corallum, lamellar septa of most 

families here defined have a modicum of vertical continuity. Nondissepimented forms like Siphonophrentinae have amplexoid septa which lack this vertical 

continuity except at the periphery; inward the partial 

septa extend upward from distal surfaces of tabulae. 

All genera with lonsdaleioid dissepiments have radially 

discontinuous septa. This is true also of the Digonophyllidae in which septa tend to lose identity peripherally within a complex mesh. Septal crests or septal 

spines in these categories are radially alined segments growing toward the axis from inner dissepiment 

surfaces.

True septal carinae are distinguished by continuity 

in depth; as seen in longitudinal thin section a carina 

passes diagonally downward and outward toward the 

periphery, transacting dissepimental tissue. Carinae 

are of three kinds: (1) crossbar or yardarm carinae, 

(2) angulation or elbow carinae, and (3) strip carinae. 

True carinae are readily distinguishable from minor 

septal wiggles, unthickened angulations, and stereoplasmic lumps or bumps on septa which do not extend 

downward from a given transverse plane. Strip carinae 

are a derivative of crossbar carinae in the outer dissepimentarium of Digonophyllidae like Arcophyllum, 

wherein the septa are aborted, but the carinae occur 

as diagonal strips viewed in longitudinal thin section.

TABULAE

A tabula is a supporting structure that occupies the 

middle part of a corallum. Tabulae of more primitive 

Rugosa extend from wall to wall, as in tabulate corals. 

More advanced dissepimented rugose corals have the 

interior differentiated into a middle tabularium and an 

outer dissepimentarium. Tabulae are lamellar, simple, 

and almost always thin, unlike septa which may be 

stereoplasmically thickened. In the primitive condition 

tabulae are nearly straight and horizontal. Modification is by sag or axial inclination; in many advanced 

genera the tabulae are domed and arched with a 

peripheral inclination. Tabulae may be entire and complete, or they may comprise short blisterlike domes 

called tabellae with the appearance of flat dissepiments. Edges of tabellae abut upon other tabellae, on 

longer more nearly complete tabulae, or against innermost dissepiments.

The boundary separating tabularium from dissepimentarium is discrete in some genera; in other genera 

the two zones appear to blend. In general the inner 

dissepiments are steeply inclined axially, whereas the 

adjacent outer tabellae are inclined peripherally. With

genera having an aulos or inner ring, the tabulae may 

be differentiated into a horizontal axial set and a periaxial set peripherally inclined, as in Syringaxon.

In forms with wide flat tabulae (like Kobeha) the 

calice floor is a tabula which bends down abruptly at 

the cardinal fossula.

Spacing of tabulae is rather uniform in some genera; 

in others there is great irregularity. Sudden change 

from close spacing to wide may be a response to environmental change. Specimens have been observed 

among solitary Rugosa in which the abrupt change in 

attitude of tabulae was obviously a response to the 

vertically attached corallum having fallen on its side, 

resulting in a change in growth direction to maintain 

horizontality of tabulae.

DISSEPIMENTS

As normal dissepiments, these blisterlike strengthening and space-filling elements occupy interseptal loculi. Lonsdaleioid dissepiments are usually larger, occur 

in the peripheral zone and are concave toward the outside. A single lonsdaleioid dissepiment opposes the 

outer ends of several septa. Peripherally the septa 

never transect lonsdaleioid dissepiments continuously; 

they occur here only as radially alined crests.

Each normal dissepiment is three dimensionally a 

spheroid segment with the basal plane axially inclined. 

Transverse section traces of these dissepiments occur 

in concentric, axially concave sets between pairs of 

septa. Peripherally these traces may become anguloconcentric or straight, or they may blend into an irregular herringbone or general cribriform tissue in which 

the septa are suppressed.

In longitudinal section the overlapping dissepiments 

form crude columns the number of which has taxonomic meaning. Normal dissepiments may be globose 

or elongate; lonsdaleioid dissepiments are always elongate and, unlike the normal, dissepiments, are always 

concave toward the outside, if viewed in transverse 

section.

Average inclination of dissepiments has taxonomic 

significance. In solitary and colonial forms with wide 

corallites, the peripheral dissepiments are horizontal; 

toward the axis the inclination always increases, such 

that the innermost, usually smaller, dissepiments 

stand nearly vertical. A character of considerable systematic value is discreteness or sharpness of the 

boundary between these inner dissepiments and the 

tabularium.

Digonophyllid species with complex peripheral patterns may have small nontransverse or radial dissepiments; all the edges of the dissepiments abut on a 

single septum.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 37

Dissepiments are generally but not always absent 

in Ordovician Rugosa and accordingly are viewed as 

advanced structures in the evolutionary sense. Dissepimented species appear abruptly and in some abundance with the Early Silurian Lykophyllidae and 

Arachnophyllidae. Nondissepimented Siphonophrentinae are among the abundant forms in the Early and 

early Middle Devonian rocks of the Great Basin. Dissepiments undoubtedly arose independently in several 

unrelated rugose coral lineages. With regard to corals 

with lonsdaleioid dissepiments, this is fairly evident, 

as they characterize several families among which are 

the Spongophyllidae, Endophyllidae, Halliidae, and 

Chonophyllidae. By themselves these lonsdaleioid features are not regarded as indicative of genetic relationship.

Columns of true horseshoe dissepiments are a special 

category known in the Cordilleran Belt only in Late 

Devonian Phillipsastraeidae. These features are normally associated with diagnostic trabecular bundles.

STEREOPLASMIC DEPOSITS

Septa and other primary structures of the rugose 

coral skeleton are thickened and added to in lamellar 

fashion by secondary calcareous matter to which the 

terms stereoplasm or stereome have been given. Within 

a species certain of these stereoplasmic additions are 

fairly regular and uniform. Deposits of this kind affect 

mainly the septa; dissepiments may be so thickened 

on the inner surfaces, but tabulae are generally not 

thickened. Axial structures become a solid columella 

when pervaded by stereome.

Taxonomically the most important stereoplasmic 

deposits are the fairly uniform peripheral thickenings 

of septa producing a septal stereozone. Certain genera 

like Kodonophyllum and Nevadaphyllum always have 

a wide septal stereozone. Stereoplasmically strengthened columellas do not occur in the Devonian corals 

here described but are common in late Paleozoic and 

Silurian genera. Some genera have either a general 

stereoplasmic dilation of septa throughout their tapered length or a uniform dilation only within the 

tabularium.

Less significant taxonomically are the minor nonuniform lobose thickenings distributed irregularly along 

septa or affecting only axial segments. True carinae are 

not basically stereoplasmic, but may be supplemented 

by this secondary material. Especially meaningful for 

systematic purposes are the rather uniform lateral 

swellings of septa that form an inner stereoplasmic 

wall or aulos, as in Syringaxon.

Among rugose corals with a well-developed outer 

wall, the thin epitheca may readily be distinguished

texturally in thin section from the contiguous septal 

stereozone making most of the wall.

RUGOSE CORAL SYMMETRY AND TAXONOMY

Rugose corals have the underlying bilateral symmetry of the coelenterate. In most genera, however, the 

more obvious symmetry is radial. In many solitary 

Rugosa the primary bilateral symmetry is recognizable 

externally or in the calice by means of the pinnate 

arrangement of septa with reference to the cardinalcounter plane. Biologic orientation is facilitated in 

many solitary forms by the adult cardinal fossula 

enveloping the cardinal septum. Well-defined counter 

fossulae are less common. In the neanic stage of some 

species there is a distinct quadripartite pattern defined 

by cardinal, counter, and alar features; these become 

less distinct in the ephebic transverse section. Generally the four symmetry quadrants are identifiable in 

mature growth stages of solitary genera. With reference to the corallum exterior, the cardinal fossula may 

lie either on the axially concave or the convex side; 

whether this character is constant within a genus or 

species is not known.

Contiguous cardinal quadrant septa tend to be pinnate or to become alined at maturity more or less 

parallel to the cardinal septum and fossula in mature 

stages. These septal features of symmetry, together 

with the down-bending of tabulae, define the fossula.

Characters of bilateral symmetry are best shown by 

solitary Rugosa; most cerioid colonial genera lack a 

fossula and show little indication of anything but secondary radial symmetry in mature individual corallites.

GROWTH CHANGES AND REPRODUCTIVE FEATURES

Growth and reproductive features have an important bearing upon rugose coral classification as in all 

Orders of the Animal Kingdom. Sequences of development change recorded in the corallum have, since the 

inception of modern coral research, been made use 

of in this regard. With Early Devonian taxa here 

described, solitary Papiliophyllinae, when serially sectioned, reveal progressive ontogenetic loss of septal 

stereoplasmic thickening in more or less orderly fashion. Syringaxon of the Laccophyllidae shows loss of 

fossulae with development of the mature aulos.

Modes of asexual reproduction or so-called "increase" among Rugosa have been dealt with extensively (Langand Smith, 1927; Hill, 1935,1956) and are 

readily discernible as axial, peripheral, and lateral offsets in solitary and fasciculate taxa (Oliver, 1968, p. 

20). Reproductive offsets of compact colonial genera 

like YLexagonaria and Billingsastraea are difficult to 

deal with in this respect, calling for selection and special preparation of suitable material.
38 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Developmental study of colonial genera is intimately 

related to manner of increase, for not only must ontogenetic changes of corallites be considered, but the 

colony growth pattern as well. Such patterns are a 

three-dimensional arrangement of parts contingent 

upon ways in which reproductive offsets develop from 

parent corallites (Oliver, 1968; Rozkowska, 1960). 

Few developmental studies of cerioid rugose genera 

have been carried out. Rozkowska's (1960, p. 14-28) 

Hexagonaria investigations demonstrate calice offsets 

from the dissepimentarium. Developmental patterns 

involving reproductive offsets of more conveniently 

handled phaceloid colonies are touched upon by Oliver 

(1968, p. 26) in connection with statistical variation 

analyses. In general too little is at present known of 

these reproductive characters for them to figure significantly in the taxonomy of Devonian compound 

genera.

Small reproductive offsets, not found in most solitary rugose coral species of the Great Basin Devonian, 

have been recognized only in a single specimen of 

Kobeha and in aberrant Breviphrentis. Characteristic 

of Breviphrentis are rejuvenescence features produced 

by growth of a single large daughter offset that largely 

fills the parental calice.

Cone-in-cone growth of solitary Digonophyllidae 

and Chonophyllidae evidently inspired Wedekind's 

(1924, p. 21) concept of the "septal cone." In accordance with this view, certain solitary genera with lamellar but vertically discontinuous septa grew upward as 

stacked-up increments or growth cones secreted by 

the polyp basal ectoderm during successive growing 

stages. Septa are represented by radial calice ridges or 

radially alined septal crests separately secreted after 

a resting interval.

VARIATION AND DIMORPHISM

. Biometric analyses of individual variation are especially appropriate in connection with rugose coral species discrimination. Studies by Oliver (1960a; 1968) 

and Rozkowska (1960) effectively present these data 

graphically as ratio curves or scatter diagrams. The 

most commonly used ratio is that of septal count relative to mature corallite diameter.

Biometry with statistical analysis of data is of value 

in distinguishing continuous from discontinuous variation and, in connection with possibilities of gene or 

somatic mutation, dimorphism, and intergrowth within 

a colonial corallum complex of individuals from two or 

more protocorallites (Oliver, 1968, p. 24-26). Dimorphism has been suspected but not always convincingly 

demonstrated in numerous rugose coral species. Appropriate in this regard is reference to the "genomorph

concept" of Smith and Lang (1930) which was based 

upon Carboniferous coral genera in which aberrant 

corallites or parts of corallites have the appearance of 

another genus. Critical reviews (Rozkowska, 1960; 

Wilson, 1963; Oliver, 1968) have not encouraged use 

of genomorph in a taxonomic sense as advocated by its 

authors. Whereas observations leading to the genomorph idea with reference to Carboniferous Rugosa 

are valid, the cited examples among Devonian species 

appear to be founded upon misconceptions, as noted 

by Rozkowska (1960, p. 49).

Likelihood of dimorphism has not been eliminated 

in connection with certain Devonian species here 

described, studies of large suites of associated individuals on a population basis will be required. Siphonophrentis (Breviphrentis) invaginatus may bear such a 

relationship to Nevadaphyllum and to associated individuals with sporadic dissepiments that are questionably assigned to Sinospongophyllum. Among the 

Disphyllidae, Billingsastraea nevadensis and its subspecies arachne, initially described by Stumm as Radiastraea, would also lend themselves to quantitative 

appraisal with sufficient material. A similar situation 

is found in Hexagonaria (Pinyonastraea) kirki, whose 

range overlaps that of Billingsastraea and which could 

be interpreted as a genetically related cerioid disphyllid possessing an outer wall.

DESCRIPTIVE TERMS

Descriptive terms applicable to the structure of the 

rugose coral have been adequately defined and stabilized by Hill (1935, p. 481-519; 1956, p. F234-F245) 

and are now widely accepted. New and useful terms 

have in recent years been proposed by Birenheide, 

Glinski, Schouppe, and other European coral researchers. Hill's terminology with minor modifications, is 

adopted in this report.

Family LACCOPHYLLIDAE Grabau, 1928

Reference form. Syringaxon siluriensis (McCoy) 

1850.

Small solitary trochoid to subcylindrical rugose 

corals with very deep calice, straight smooth lamellar 

septa, a narrow septal stereozone, and tubular stereoplasmic aulos. Tabulae partly complete, very strongly 

uparched, nearly flat within aulos, steeply inclined 

peripherally. No dissepiments.

These nondissepimented but otherwise specialized 

corals are distinguished by the strong inner ring produced by swelling of septal tips and commonly rather 

heavy addition of stereoplasm. Arching of tabulae is 

pronounced. The Laccophyllidae occur in the Silurian 

and Devonian wherein they may be abundant in a 

facies with no other Rugosa. Nearly identical small
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 39

corals like Permia, which reappear in the Carboniferous, may be homeomorphic (Hudson, 1944, p. 360; 

Fliigel and Free, 1962, p. 232). Somewhat less convergent are the Permian Polycoeliidae elucidated by 

Schindewolf (1942, p. 55).

Syringaxon is the only member of this family recognized in the Great Basin, where it is the characterizing rugose coral of the Helderberg Early Devonian 

Syringaxon facies.

Genus SYHINGAXON Lindstrom, 1882

1882. Syringaxon Lindstrom, p. 20.

1900. Laccophyllum Simpson, p. 201.

1902. Nicholsonia Pocta, p. 184. Cited in plate explanations as

Alleynia (Nicholsonia).

1928. Laccophyllum Simpson. Grabau, p. 82. 

1928. Alleynia Pocta (Nicholsonia Pocta) Grabau, p. 82. 

1935. Syringaxon Lindstrom. Butler, p. 117. 

1938. Syringaxon Lindstrom. (in part). Prantl, p. 21. 

1940. Syringaxon Lindstrom. Lang, Smith, and Thomas, p.

129.

1949. Syringaxon Lindstrom. Stumm, p. 10. 

L956. Syringaxon Lindstrom. Hill, p. F258. 

L962. Syringaxon Lindstrom (in part). Fliigel and Free, p.

224.

Type species. By monotypy, Cyathaxonia siluriensis McCoy, 1850 (p. 281); Silurian, upper Ludlow, 

Underbarrow, Kendal, Westmorland, England. According to Butler (1935, p. 118) and Lang, Smith, and 

Thomas (1940, p. 129), Lindstrom, in naming the 

genus, gave no diagnosis; he merely used McCoy's 

species siluriensis under the new generic name Syringaxon in his faunal lists of the Gotland Silurian 

(Lindstrom, 1882, p. 20).

Diagnosis. Small, solitary turbinate and ceratoid 

to cylindrical rugose corals with deep calice and no 

dissepiments. Axial ends of major septa dilated and 

laterally in contact to form an aulos. Tabulae strongly 

uparched distally, the aulos dividing them into inner 

flat or slightly sagging segments and outer segments 

which descend steeply to meet the outer wall (Fliigel 

and Free, 1962, fig. 3); tabulae either continuous from 

wall to wall through aulos stereozone, terminating 

within the aulos, or abutting proximally as tabellae 

against adjacent tabulae. Stereome usually abundant 

as aulos thickening and general thickening or filling 

in brephic and neanic growth stages. Narrow fossulae 

commonly reveal quadrants in late neanic growth 

stages.

Remarks. Simpson (1900, p. 201) called attention 

to this distinctive rugose coral group in the American 

Silurian upon proposal of Laccophyllum, with L. 

acuminatum Simpson from the Niagaran of Perry 

County, Tenn., as type species. Butler's exhaustive 

study (1935) determines that McCoy's (1850, p. 

281) "Cyathaxonia siluriensis" is the type species of

Syringaxon Lindstrom; later Lang, Smith, and Thomas 

(1940, p. 129) concluded that the holotype of Laccophyllum acuminatum Simpson is correctly assignable 

to Syringaxon, thus making Laccophyllum Simpson 

a synonym of Syringaxon Lindstrom.

Alleynia Pocta or Alleynia (Nicholsonia) Pocta (see 

Lang and others, 1940, p. 15, 129) appears to be correctly regarded as a synonym of Syringaxon.

Barrandeophyllum Pocta (1902, p. 190) is similar 

to Syringaxon. According to Prantl (1938, p. 34), who 

studied Pocta's cotypes of the type species, Barrandeophyllum perplexum, this genus is virtually identical 

in habit, form, and general structure with Syringaxon, 

from which it may have been derived. Barrandeophyllum is distinguished from Syringaxon by irregularities of the aulos, which is usually elliptical in 

section, and by a sparing development of dissepiments. 

Stereoplasm is always present but is not abundant. 

The type species of Barrandeophyllum is reported 

from the Lower Devonian Branik Limestone of 

Bohemia. Syringaxon ranges from the Niagaran of 

Tennessee through strata of Lower Devonian Helderberg age in western North America and is reported 

in the Upper Devonian Independence Shale of Iowa 

(Stainbrook, 1946, p. 402).

As noted by Hudson (1944, p. 360), other small and 

rather simple corals closely resembling Syringaxon 

range into the Carboniferous, where they are represented by the genus Permia Stuckenberg. According 

to Hudson, Permia and Syringaxon are probable 

homeomorphs.

In the Great Basin small solitary corals similar to 

Syringaxon occur in the lower or Silurian part of the 

Hidden Valley Dolomite of the northern Panamint 

Mountains (McAllister, 1952, p. 15-17). Following 

the great proliferation in the Great Basin Helderbergian these corals seemingly almost disappeared 

from this region, although scarce and fragmentary 

specimens which may belong either in Syringaxon or 

Barrandeophyllum have been found at Lone Mountain 

in the Nevada Formation, unit 2.

Springaxon foerstei n. sp.

Plate 1, figures 1-16 

1963. Syringaxon acuminatum (Simpson). Merriam, p. 43.

Type material. Holotype, USNM 159243; figured 

paratypes, USNM 159245-159250 inclusive. Rabbit 

Hill Limestone, Early Devonian, Central Great Basin; 

Monitor Range, Nev.

Diagnosis. Syringaxon has a very deep calice, thick 

wall, prominent longitudinal grooves externally and 

on interior of calice wall, and a high septal count for 

this genus. Cardinal and alar septa are differentiated 

in neanic stages.
40 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

External features. Corallum, large for the genus, 

attains a length of 30 mm and maximal diameter of 

27 mm at edge of calice; average length about 19 mm 

and average diameter 16 mm. Calice very deep, in 

large individuals as much as 16 mm or more than half 

the corallum length. Average specimen is trochoid, but 

ceratoid individuals are fairly common; turbinate and 

nearly cylindrical individuals are uncommon. Septal 

grooves are sharply defined and distinctly pinnate 

with reference to the primary septa. Rugae are weakly 

defined or absent, rarely prominent. A few individuals 

have talons or attachment surface at apex. Septa 

extend distally as low subequal ridges on inside of 

calice and rarely become obsolete. Aulos is defined as 

a ring-shaped boss at the bottom of the calice in many 

specimens; in the others the aulos is ill defined, and 

narrow fossulae may outline quadrants made by the 

primary septa.

Transverse sections. Mature sections have 26 to 

more than 30 major septa extending three-fourths of 

the distance to the axis. Minor septa are normally 

present, but in some individuals are not recognizable 

at maturity; these septa range from mere stubs to 

one-third the length of the major septa; in some 

specimens the septa are buried within the peripheral 

stereoplasm. Septa are usually dilated toward tips, 

laterally in contact, and reinforced by stereoplasm to 

produce an aulos. Entire septum may be somewhat 

dilated. Individuals that have a poorly defined aulos 

have pairs of contiguous septa meeting axially like a 

tuning fork, and merged segments continue toward 

axis; some of these specimens reveal quadrate transverse symmetry. The thickness of the peripheral 

stereozone in some specimens exceeds 1 mm, but is 

usually not so thick as the aulos. There is considerable variation in the amount of stereoplasm; some 

early ephebic sections show the entire aulos filled with 

this material. The appearance of false dissepiments 

possibly results from section-cutting bulbous distal 

inflations of tabulae between the aulos and the outer 

wall.

Longitudinal sections. Outer segments of tabulae 

descend steeply, with upward inflection before 

meeting the stereozone. Some complete tabulae are 

vague within the aulos, others are traceable completely across. Some aulos segments of tabulae may 

be thickened stereoplasmically. Some specimens have 

complete tabulae with M-shaped curvature and nearvertical outer segments. Tabular spacing is nonuniform 

and usually is fairly wide.

Comparison with related forms. Butler's (1935) 

figures of Syringaxon siluriensis (McCoy) show a 

maturely cerioid to subcylindrical form with fewer 

septa and poorly developed minor septa. Syringaxon 

foerstei includes externally similar subcylindrical variants, but the norm is a trochoid corallum. Both tend 

to have heavy stereoplasmic deposits in the aulos and 

a wide septal stereozone. Of several species figured by 

Prantl (1938) from the Devonian of Czechoslovakia, 

most have a lower septal count and elongate subcylindrical coralla. Species described from the Greifensteiner Kalk (Eifelian) of Germany by Fliigel and 

Free (1962) have a weaker aulos and less arching of 

tabulae.

Occurrence. Rabbit Hill Limestone of Helderberg 

Early Devonian age; Devonian coral zone A. Northern 

Monitor Range, Rabbit Hill vicinity: localities M48, 

M49, M187. Middle part of the Monitor Range, 

Dobbin Summit area: localities M1067, M1068, 

M1069. Northern Simpson Park Mountains, Walti 

Hot Springs area: locality M1074; Coal Canyon area: 

localities M1032, M1075, M1076. Cortez Mountains, 

Mount Tenabo area: locality M1083.

Beacon Peak Dolomite Member (Helderberg Early 

Devonian age) of the Nevada Formation; Devonian 

coral zone A. Southern Sulphur Spring Range: 

localities M186, M197, M1081, M1082.

Family STREPTELASMATIDAE Nicholson, 1889 

(as STREPTELASMIDAE)

Reference form. Streptelasma corniculum Hall. 

Ordovician, New York.

These are solitary rugose corals with or without 

fossula, normally without dissepiments but with a discrete septal stereozone; tabulae are wide, flat or 

domed, some complete. Septa of primitive forms are 

thickened and laterally in contact through mid-neanic 

stage; septa thin peripherally from axis; in some 

genera septa reach axis and form with domed tabulae 

axial complexes or columellae. Tabulae, obscure in 

this genera, have a very wide stereozone.

Measurements of Syringaxon foerstei n. sp.

Calice bottom Ephebic section

USNM 

No.

159243 (holotype) ..............

159244 (paratype) ..............

159246 (paratype) ..............

159248 (paratype) ..............

159352 (pi. 1, figs. 9, 10) ....

Corallum length 

restored (mm)

32

17

Calice depth 

(mm)

16

8

Outside diameter 

(mm)

12

11.5

Number of 

major septa

28

24

Diameter 

(mm)

9.5

13

15.6

Number of 

major septa

26

24

26
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 41

This diverse long-ranging family will probably lend 

itself to further subfamily taxonomic subdivision 

eventually. Great Basin Devonian representatives are 

classified in a new subfamily, here designated as the 

Siphonophrentinae.

Subfamily SIPHONOPHRENTINAE, new subfamily

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

These Streptelasmatidae have a narrow to wide 

septal stereozone, amplexoid septa, and a weak to 

prominent fossula. Tabulae are mostly complete, 

straight or domed with axial and peripheral sag. Dissepiments are lacking.

Included are the following genera and subgenera:

Siphonophrentis (Siphonophrentis) O'Connell, 1914 

Siphonophrentis (Breviphrentis) Stumm, 1949 (as genus) 

Heterophrentis Billings, 1875 

Compressiphyllum Stumm, 1949 

(l)Nevadaphyllum Stumm, 1937 

(?)Homcdophyllum Simpson, 1900

Breviphyllum Stumm, 1949 may belong here as a 

synonym of Breviphrentis. Although defined as a genus 

with dissepiments, Stumm's holotype of the type 

species of Breviphyllum (plate 17, figs. 12, 13) does 

not reveal dissepiments. As noted by W. A. Oliver, Jr. 

(written commun., 1970), Homalophyllum has a welldeveloped stereocolumella unlike the other genera here 

included in this subfamily.

The significant generic characters are shape of 

corallum, width of septal stereozone, prominence of 

fossula, and rejuvenescence features. Mature corallum 

shape ranges from elongate-subcyclindrical in Breviphrentis to short and compressed laterally in Compressiphyllum (Oliver, 1958). The septal stereozone 

is relatively narrow in Siphonophrentis s.s. and some 

species of Heterophrentis but wide in some species of 

Breviphrentis and very wide in Nevadaphyllum. Septal 

length and the spacing and evenness of tabulae are 

characters of extreme variability within this subfamily. Fossulae are prominent with Heterophrentis, 

small or unrecognizable in mature coralla of some 

Siphonophrentis s.s. and in Breviphrentis. Prominent, 

shelving rejuvenescence flanges, and the narrow 

elongate corallum set Breviphrentis apart from true 

Siphonophren tis.

Solitary Rugosa that are here classified as Siphonophrentinae have from time to time been referred to 

in the literature as zaphrentoid corals or assigned to 

the Zaphrentidae, a family of uncertain characterization to which they doubtfully belong (Hill, 1938, p. 

23; Stumm, 1949, p. 11; Oliver, 1958, p. 816). Status 

of the Siphonophrentinae with respect to Zaphrentidae 

calls for evaluation of the type species of Zaphrentis, 

usually cited as Z. phrygia Rafinesque and Clifford,

1820. Opinions vary widely regarding the characterization of Z. phrygia, upon which the validity of the genus 

and of the Family Zaphrentidae depends. Hill (1956, 

p. 278, fig. 190, 3a-b) ascribes to Zaphrentis a narrow 

dissepimentarium and crossbar carinae illustrated by 

figures of Z. phrygia attributed to Schindewolf. Stumm 

(1964, p. 34, pi. 27, figs. 1-7), on the contrary, regards 

Z. phrygia as lacking dissepiments and notes the 

possibility of confusion with carinate Heliophyllum 

venatum Hall. Obviously, thin section study of topotype material is needed to resolve this taxonomic 

problem.

Siphonophrentinae and stratigraphic indicators 

within the early Middle Devonian beds (Onondaga 

Limestone) of the New York area that are approximately correlative strata of the beds in the Great 

Basin.

Associated with Siphonophrentinae in the Nevada 

Formation coral zone D2 are solitary corals that 

resemble Breviphrentis and Heterophrentis; they 

differ in having sporadic dissepiments. These problematic individuals are assigned to Sinospongophyllum 

of the Chonophyllidae, but for some individuals 

aberrant development of lonsdaleioid dissepiments in 

a siphonophrentid lineage cannot be entirely ruled out;

Genus SIPHONOPHRENTIS O'Connell 1914

1914. Siphonophrentis O'Connell, p. 187,190,191. 

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

1949. Breviphrentis Stumm, p. 13-14.

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

1956. Siphonophrentis O'Connell. Hill, p. F271. 

1960a. Siphonophrentis O'Connell. Oliver, p. 87-89.

Type species. By original designation Caryophyllia gigantea Lesueur, 1821, p. 296, 297. Onondaga 

Limestone, New York State.

Diagnosis. Oliver's (1960a, p. 87) diagnosis 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 commonly develops 

very elongate coralla and may attain large size.

Heterophrentis (type species Zaphrentis spatiosa 

Billings, 1858) differs in having a curved ceratoid to 

trochoid or almost turbinate mature corallum with 

larger and better defined fossula on the convex side 

and a narrow septal stereozone; the very elongate 

cylindrical growth habit of Siphonophrentis is lacking. 

Unlike Siphonophrentis, some mature individuals of 

Heterophrentis have major septa reaching the axis, 

where they may be loosely twisted and pervaded by 

stereome, but not to the ^extent of forming a discrete 

axial structure. Tabulae'of Heterophrentis are com-
42 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

monly more irregularly arched, less uniform, and more 

widely spaced than are those of Siphonophrentis. 

Immature and early mature growth stages of the two 

genera are probably indistinguishable.

The genus Siphonophrentis lends itself to separation as two subgenera: Siphonophrentis s.s. and Breviphrentis.

Subgenus BREVIPHRENTIS Stumm, 1949 (as genus)

1949. Breviphrentis Stumm, p. 13,14, pi. 5, figs. 22-24. 

1956. Breviphrentis Stumm. Hill, p. F271, fig. 183-6ab. 

1960a. Siphonophrentis variabilis Oliver, p. 87-89, pi. 13.

Type species. Amplexus invaginatus Stumm (1937, 

p. 427, pi. 53, fig. 2, pi. 54, figs. 2a-c). Early Middle 

Devonian, Nevada Formation, unit 2, coral zone D2 ; 

Atrypa Peak, Eureka district, Nevada.

Diagnosis. This narrow, elongate Siphonophrentis 

is of markedly segmented appearance when fully 

mature; the cylindrical corallum has deep lateral 

shelving indentations and rejuvenescence flanges. The 

cardinal fossula is commonly weak or unrecognizable 

in the mature calice. The septal stereozone is medium 

to wide.

Remarks. Oliver (1960a, p. 87) recognized the 

morphologic resemblance of Breviphrentis invaginatus 

to Siphonophrentis. Retention of Breviphrentis as a 

subgenus seems warranted, as this group constitutes 

a useful taxonomic unit differing from true Siphonophrentis in its more slender and markedly segmented 

growth habit. Typical Siphonophrentis is a larger coral 

when fully mature and lacks the pronounced rejuvenescence flanges; its septal stereozone is relatively 

narrower.

Breviphrentis is especially characteristic of the 

Great Basin Early and early Middle Devonian (coral 

zones C and D). Stumm's proposal of Breviphrentis 

was made before the stratigraphic importance of 

these corals was understood. In the New York Early 

Devonian, Siphonophrentis variabilis Oliver (1960a, 

p. 87, pi. 13, figs. 1-17) of the Coeymans Limestone 

resembles Great Basin Breviphrentis more closely 

than true Siphonophrentis.

Siphonophrentis (Breviphrentis) invaginatus (Stumm)

Plate 15, figures 1-11; plate 16, figures 1-13,15,16; 

plate 17, figures 12, 13.

1937. Amplexus invaginatus Stumm, p. 427, pi. 53, fig. 2, pi.

54, figs. 2a-c. 

1937. (?) Heterophrentis nevadensis Stumm, p. 426, pi. 53,

fig. 1, pi. 54, figs. la-c. 

1937. (?) Amplexus lonensis Stumm, p. 428, pi. 53, fig. 4, pi.

54, figs. 4a-b. 

1937. (?) Amplexus nevadensis Stumm, p. 53-54 (in part),

pi. 53, fig. 3, pi. 54, figs. 3a-b; not fig. 3c. 

1940. Amplexus invaginatus Stumm. Merriam, pi. 16, figs. 3-

4. 

1949. Breviphrentis invaginatus (Stumm), p. 13-14, pi. 5, figs.

22-24.

1949. (?) Breviphyllum lonensis (Stumm), p. 25-26, pi. 12, 

figs. 4-5.

Type material. Holotype, USNM 94443; paratype 

USNM 94443a. Lower beds of the Nevada Formation, 

Atrypa Peak, Eureka mining district, Nevada.

Figured specimens. -USNM 159296-159311 inclusive.

Diagnosis. This Siphonophrentis has an elongate 

subcylindrical mature growth habit with several rejuvenescence constrictions. Septal stereozone is narrow to fairly wide. The axial extensions of septa are 

amplexoid, and the narrow cardinal fossula is either 

vague or well defined in mature growth stages.

External features. Neanic and early ephebic 

growth is ceratoid like Heterophrentis and later 

ephebic growth subcylindrical, elongate, and with rejuvenescence constrictions. Calice rim flanges are commonly sharp and developed repeatedly at irregular 

intervals; these flanges are usually broken off. Septal 

grooves are well defined.

Transverse sections. Major septa, about 40, normally extend one-half to three-fourths the distance 

toward the axis; though usually withdrawn from axis 

in ephebic stages, some individuals have major septa 

reaching the axis. Minor septa, long for this genus, 

commonly exceed half the length of major septa. Individuals with a wide septal stereozone have a thickened 

septa; in some, the complete stereoplasmic filling extends inward almost to the tips of minor septa. Septa 

are fairly straight to minutely wavy. In some specimens, the cardinal fossula may be defined by the 

U-shaped trace of the tabular downwarping, but others 

reveal no evidence of this structure.

Longitudinal sections. Calice rim rejuvenescence 

flanges are sharp and project obliquely 8mm or more 

from the coral body. Tabulae, mostly complete, arch 

distally with a peripheral depression, and are either 

straight axially and periaxially or have a median sag. 

Tabellar loops or globose pockets develop near the 

peripheral edges of the tabulae. Incomplete tabulae 

are accompanied by flat peripheral tabellae. The regular specimens have more or less complete tabulae that 

are commonly close spaced. Some irregular individuals 

have widely spaced nonuniform tabulae. Aberrant individuals have closely spaced, nearly straight tabulae 

almost unarched.

Comparison with related forms. Siphonophrentis 

(Breviphrentis) kobehensis of coral zone C lacks the 

well-developed septal stereozone of most variants of 

zone D invaginatus and has a lower septal count and 

very short minor septa.

Siphonophrentis variabilis Oliver of the New York 

Coeymans Limestone (Oliver, 1960a, p. 87, pi. 13, figs. 

1-17) bears a rather close resemblance to invaginatus,
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 43

from which it may differ only subspecifically. Both 

include individuals with a wide septal stereozone; variabilis is not known to have variants with nearly 

straight tabulae as does the species invaginatus.

Remarks. A large suite of Breviphrentis invaginatus from Nevada Formation, unit 2 at Grays Canyon, 

Eureka district, illustrates the great variability of this 

species. The highly variable characters are the arching 

and evenness of tabulae, the spacing of tabulae, and 

the width of the septal stereozone. Some individuals 

have straight almost unarched tabulae. The narrow 

cardinal fossula varies from vague or unrecognizable 

to fairly well defined in the mature growth stages, similar to Heterophrentis. Local variants have a wide septal stereozone, suggesting that of Kodonophyllum or 

Pseudoblothrophyllum.

Among variants of B. invaginatus at Grays Canyon 

are individuals with the features of Stumm's Amplexus 

lonensis holotype, the type species of Breviphyllum. 

As noted elsewhere, the generic diagnosis of Breviphyllum (Stumm, 1949, p. 26) calls for a dissepimentarium, but no dissepiments were found in the thin 

sections of the holotype. Other individuals resemble 

Stumm's Heterophrentis nevadensis.

In longitudinal sections the axial extensions of 

major septa rest in amplexoid fashion upon the upper 

surfaces of tabulae.

Occurrence. Central Great Basin, Nevada Formation, unit 2, Devonian coral zone D. Lone Mountain 

(D2 ): localities M55, M1036, M1048. Southern Fish 

Creek Range (D2 ): locality M1033. Northern Fish 

Creek Range, Grays Canyon (D2 ): localities M3, M51. 

Mahogany Hills (D2 ): localities M27, M1084. Northern Antelope Range (D2 ): locality M1035. Southern 

Sulphur Spring Range (D2 ): localities M36, M198, 

M1078.

Southern Great Basin, Devonian coral zone D. 

Ranger Mountains: localities M1034, M1058. Northern Panamint Mountains: localities M184, M1065. 

Funeral Mountains: localities M1059, M1060, M1061, 

M1063, M1064. Desert Range: locality M1057.

Measurements of Siphonophrentis (Breviphrentis) invaginatus

Ephebic section Neanic section

USNM 

No.

Diameter 

(mm)

Number of 

major 

septa

Diameter 

(mm)

Number of 

major 

septe

159296 .......

159298 .......

159306 ........

159309 ........

....... 30.5

....... 29.3

....... 16

....... 26.6

44

40

32 8.3

42

22

Siphonophrentis (Breviphrentis) kobehensis n. ap.

Plate 14, figures 1-15; plate 16, figure 14. 

Type material. Holotype USNM 159284; para types 

USNM 159283, USNM 159285 to 159293 inclusive.

Lower part of the Nevada Formation, Lone Mountain, 

Eureka County, Nev.

Diagnosis. This Siphonophrentis has a ceratoid to 

cylindrical growth habit that reflects frequent rejuvenescence and elongate curved corallites. Septa are 

withdrawn from axis; minor septa are very short (or 

unrecognizable). Tabulae are complete, occasionally 

straight, but more commonly uparched distally and 

either flat medially or with a slight sag. Tabulae are 

usually rather widely spaced. The wall has a very narrow stereozone for this genus. There are no dissepiments, but there are occasional looplike or globose 

tabellae. The abrupt curvature changes on rejuvenescence all caused by the resumption of upright growth 

by the parent that had been recumbent.

External features. The segmented appearance of 

the corallum is caused by the marked constrictions 

resulting from rejuvenescence. The rim of the parental 

calice commonly remains a sharp flange distal to which 

the single daughter corallite may differ in growth direction; for some individuals the direction may diverge 

90°; but normally the change is only a few degrees. 

There are a few weak transverse folds between the 

rejuvenescence flanges. Septal grooves are well defined.

Transverse sections. Major septa, about 40 in 

ephebic stages, are withdrawn to about half the radius; 

the septa are of fairly uniform width, little thickened 

by stereoplasm, amplexoid, and resting upon the upper 

surfaces of the tabulae. The outer wall has a thin septal 

stereozone beyond which the minor septa project to a 

distance only about one-fifth the length of the major 

septa. In a late nepionic stage (diameter, 5 mm), the 

septa number 22; most meet toward the axis but are 

not otherwise in contact laterally; septa lateral to the 

cardinal septum are subparallel. At an earlier nepionic 

stage (diameter 3.5 mm) there are about 20 septa 

within which the cardinal quadrants are recognizable, 

and the two septa lateral to the cardinal bend inward 

as they approach the axis, thus defining the primitive 

fossula. The cardinal fossula, inconspicuous or unrecognizable in later ephebic stages, is probably indicated 

in transverse sections by a trace which appears to join 

two major septa in U fashion and which probably 

conforms to downwarping of a tabula. There are no 

dissepiments.

Longitudinal sections. Most tabulae are complete; 

they vary from straight to uparched distally and 

peripherally depressed; the axial part is slightly convex 

distally to flat or sagging. Some individuals have a few 

localized globose periaxial tabellae. The tabulae are 

usually spaced rather wide. There are no dissepiments 

and a very narrow marginal stereozone or none. The 

tabulae are irregularly uparched where the corallum 

makes a sharp bend at rejuvenescence.
44 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Reproductive offsets. An abnormal corallum shows 

a cylindrical, curving calice offset of slightly more than 

half the diameter of the calice rim. This development 

seems to be an abnormally small rejuvenescence offset 

terminating growth of the parent after it began secretion of the rejuvenescence shelf.

Comparison with related forms. Breviphrentis 

invaginatus possesses a wider septal stereozone and 

longer minor septa. Siphonophrentis variabilis Oliver 

of the New York Coeymans Limestone has a wide 

stereozone and develops tabulae with M-shaped configuration in longitudinal section; straight tabulae are 

more characteristic of kobehensis.

Measurements of Siphonophrentis (Breviphrentis) kobehensis

Ephebic section

Neanic section 

(early)

USNM 

No.

Diameter 

(mm)

Number of 

major

septa

Number of 

Diameter major 

(mm) septa

159283 (paratype) .......

159291 (paratype) .......

159294 (figured 

specimen) ...................

..... 13.7

..... 13.5

38

40

5.4 23

Occurrence. Lower Devonian (Emsian), coral zone 

C. West side of Lone Mountain, Eureka County, 

Nev. localities M286, M74 in lower part of Nevada 

Formation, unit 1 about 75 feet stratigraphically above 

base.

Genus NEVADAPHYLLUM Stumm, 1937

1937. Nevadaphyllum masoni Stumm, p. 429, pi. 53, fig. 6, pi. 

54, figs. 6a-b.

1949. Nevadaphyllum Stumm, p. 19, pi. 8, figs. 20-21. 

1956. Nevadaphyllum Stumm. Hill, p. F271, fig. 183-5.

Type species. Nevadaphyllum masoni Stumm (by 

original designation). Nevada Formation, Lone Mountain, Eureka County, Nev.

Diagnosis. Large ceratoid to cylindrical solitary 

rugose corals. Many of the very numerous long, smooth 

major septa reach the axis, toward which they become 

twisted. There is a wide peripheral stereozone and a 

fossula. The tabularium is wide; the tabulae numerous, 

crowded and incomplete, uparched periaxially with 

axial and periaxial sag. Dissepiments are abnormal or 

false; the dissepiments are obscured peripherally by 

the stereozone.

Remarks. Pseudoblothrophyllum Oliver (1960a, p. 

91) has some features of Nevadaphyllum. This Helderbergian genus, as represented by Pseudoblothrophyllum helderbergium, has fewer major septa, a wider 

stereozone, and a more regular tabularium. As with 

Nevadaphyllum, the dissepiments are largely suppressed within the wide stereozone.

Nevadaphyllum masoni Stumm 

Plate 13, figures 3, 4

1937. Nevadaphyllum masoni Stumm, p. 429, pi. 53, fig. 6, pi. 

54, figs. 6a-b.

Type material. Holotype USM 94447; paratype 

USNM 9444 7a. Lower part of the Nevada Formation, 

Lone Mountain, Eureka County, Nev.

Diagnosis. See generic diagnosis.

Transverse section. Major septa, about 76; are 

smooth and fairly straight for two-thirds of the radius 

from the wall; they become increasingly wavy and 

twisted toward the axis which most reach. There is 

no axial thickening to form a discrete axial structure. 

Minor septa, less than one-half the length of major 

septa, become very thin axially. All septa thicken 

peripherally; some show a very minute waviness. Most 

of the wide stereozone is apparently infilled by stereoplasm between the septa. The well-defined fossula 

extends from near the axis into the stereozone and contains a shortened major septum that is assumed to be 

the cardinal. Laterally the dissepiment traces cannot 

be distinguished clearly from section plane intersections with the numerous irregular arched tabulae.

Longitudinal section. The tabularium is a complex 

of close-spaced irregular tabulae and tabellae lacking 

straight elements; tabulae are uparched periaxially 

with a pronounced axial sag. Tabulae and tabellae are 

poorly separated from elongate false dissepiments in 

the zone of peripheral sag; axially inclined false 

dissepiments are obscured peripherally by the wide 

stereozone. In the axial zone, septal ends and crowded 

tabulae form a plexus in which the structural details 

are obscured.

Remarks. The types of Nevadaphyllum masoni 

are the only known specimens with the full characterization of this genus. Probably this uncommon 

multiseptate form with crowded irregular tabulae and 

modified, obscured dissepiments represents an aberrant evolutionary development from the Breviphrentis 

stock or perhaps the stock which produced the dissepimented Nevada forms here assigned to Sinospongophyllum. Of these, Sinospongophyllum sp. f with a 

wide stereozone, obscured nonuniform lonsdaleioid 

dissepiments, and numerous axially twisted major 

septa is the closest to N. masoni. In this form, however, the tabularium is much less complex.

Occurrence. Lower part of the Nevada Formation, 

probably unit 2 and Devonian coral zone D2 , early 

Middle Devonian, Lone Mountain, Eureka County, 

Nev.

Family KODONOPHYLLIDAE Wedekind. 1927

Reference form. Kodonophyllum milne-edwardsi 

(Dybowski). Silurian, Gotland.

These solitary and colonial rugose corals have long 

lamellar septa, no fossula at maturity, no dissepiments,
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 45

and a very septal stereozone. The tabularium is composed of flat tabulae or arched tabellae which may 

combine with septal ends to produce an axial structure.

Two subfamilies are recognized: Kodonophyllinae 

and Mycophyllinae.

These corals are largely confined to the Silurian 

rocks; those rather characteristic of the Late Silurian 

are the Great Basin Silurian coral zone E.

The Kodonophyllinae resemble the Siphonophrentinae by having a wide stereozone; they differ in lacking 

a mature fossula, in having on the whole a wider, denser stereozone, and in possessing an axial structure 

supported by arching tabulae and tabellae.

(?)Kodonophyllum ap. i 

Plate 13, figures 1, 2

An incomplete Kodonophyllum-like solitary coral 

with an exceptionally wide septal stereozone was collected from the lower part of the Nevada Formation in 

the southern Fish Creek Range, where it was associated with Siphonophrentis (Breviphrentis) invaginatus. This species of Siphonophrentis also has a wide 

stereozone. Because this fragmentary specimen could 

be prepared only as a transverse section, the possibility 

of it being an extremely aberrant Breviphrentis or 

Nevadaphyllum is not eliminated.

A wide, dense stereozone exceeds one-half the corallum radius. The septal count is approximately 66. 

Major septa are withdrawn from the axis; minor septa 

greatly exceed one-half the length of major septa and 

extend far beyond the stereozone. All exposed septa 

are thickened and wavy, with numerous protuberances. Tabulae and axial structure are unknown.

The stereozone, similar to that of Pseudoblothrophyllum Oliver (1960a, p. 91, pi. 16), differs in having 

long, smooth septa, some of which meet the axis. The 

septa of Pseudoblothrophyllum differ in lacking lobes 

and by being thinner in the tabularium.

Occurrence. Nevada Formation, unit 2, Devonian 

coral zone D. Fish Creek Range, north end of Fenstermaker Mountain: locality M1033.

Kodonophyllum, usually a Silurian genus, has a distinctive axial structure with a calice boss; it is otherwise known only in Late Silurian coral zone E of the 

Great Basin Province.

Family STAURIIDAE Edwards and Haime 1850

Reference form. Stauria astreiformis Edwards and 

Haime, 1850. Silurian, Gotland.

These fasciculate and cerioid rugose corals have 

slender corallites, narrow septal stereozone and thin 

lamellar, nonamplexoid septa that commonly extend 

to the axis. Tabulae are complete, straight or arched. 

Dissepiments are commonly absent, but in some lineages arise as a single column or sporadically in a broken column.

Some species of Stauria reveal a quadripartite 

mature transverse section in which four primary septa 

are prominent and may meet at the axis. In the genus 

Dendrostella, the four primary septa may exceed the 

others in length, but usually they do not reach the 

axis of a mature corallum.

Stauriidae include many Late Ordovician species of 

Favistella Dana (or Favistina Flower), Cyathophylloides Dybowski, and Paleophyllum Billings; these 

genera carry over into the Silurian. Silurian Stauriidae 

also include undescribed genera in addition to Stauria. 

The Devonian representatives are Dendrostella, possibly Columnaria, and possibly Synaptophyllum Simpson, as revised by McLaren (1959).

Genus DENDROSTELLA Glinski, 1957 

1886. Cyathophylloides rhenanum Frech, p. 93, pi. 3 (15),

figs. 19, 19a. 

1892. Columnaria (Cyathophylloides) disjuncta Whiteaves,

p. 269, pi. 34, figs. 3,3a, 3b. 

1957. Favistella (Dendrostella) rhenana (Frech). Glinski, p.

88-90, figs. 1-4,16. 

Favistella (Dendrostella) praerhenana Glinski, p. 90-

91, figs. 5,14.

Type species. Cyathophylloides rhenanum Frech 

(by original designation, Glinski, 1957). Upper Middle 

Devonian, Givetian, "Biicheler Schichten", Paffrath 

Basin, Germany.

Diagnosis. These slender dendroid and phaceloid 

rugose corals have a thick wall, no dissepiments, 

smooth, simple, even, nonamplexoid and unthickened 

septa; minor septa are short or lacking. Tabulae are 

complete, either straight or with a slight doming or 

sag.

Remarks. Dendrostella was proposed by Glinski as 

a subgenus under Favistella; however, it appears sufficiently well characterized and distinctive to be considered a full genus. Denstrostella commonly has short 

septa which are not truly amplexoid because they are 

not developed only on the upper surfaces of tabulae. 

Glinski states that the range is from Silurian (upper 

Ludlovian) to Middle Devonian (Givetian). In the 

Great Basin the only known occurrence is in Nevada 

Formation, unit 2, possibly coral zone D3 which would 

be about early Eifelian.

Dendrostella romanensis n. sp. 

Plate 18, figures 1-4

Type material Holotype USNM 159318; Nevada 

Formation, Devonian coral zone D. Locality M1031, 

Sulphur Spring Range.

Diagnosis. Dendrostella has relatively short major 

septa and very short to completely suppressed minor 

septa. There is no indication of transversely bilateral
46 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

pattern of septa.

External features. Phaceloid colonies are many 

centimeters wide with long, slender branching fairly 

straight corallites loosely arranged. Surface has fine 

longitudinal grooves and striae; transverse rugae are 

weak. No lateral connecting processes were observed.

Transverse sections. Major septa, about 20, commonly extending less than half the distance to the 

axis. Minor septa pass only a short distance beyond 

the wall or are represented only by roots within the 

thick wall. The wall is divided circumferentially into 

about 40 equal radial parts (segments) by dark hairthin lines, one segment for each septum root. Septa 

have median dark lines.

Longitudinal sections. Thickened wall has minor 

undulations, but no sharp bends or angular rejuvenescence features. Tabulae are all complete, not widely 

separated, straight to undulant, but lack a persistent 

axial arch or depression. There are no dissepiments.

Comparison with related forms. Dendrostella 

romanensis n. sp. has shorter major septa than rhenana 

(Frech) and praerhenana Glinski; unlike romanensis, 

these west European forms have a suggestion of symmetry with primary septa distinguishable in mature 

stages. Dendrostella disjuncta (Whiteaves) also has 

long major septa and longer minor septa than 

romanensis.

Measurements. Diameter (in mm) and major 

septa, three corallites of holotype colony, USNM 

159318:

Corallite ................ 123

Diameter (mm) .... 9 8 7.5

Major septa ..........20 20 20

Occurrence. In somewhat arenaceous limestone of 

Nevada Formation, unit 2, Sulphur Spring Range, 

northwest of Romano Ranch: locality M1031. The 

coral-bearing bed lies about 150 feet stratigraphically 

above sandy limestone with a large fauna including 

Spirifer pinyonensis, Breviphrentis cf. B. invaginatus, 

and Leonaspis sp.; this fauna probably represents coral 

zone D2 . The Dendrostella occurrence may be in coral 

zone D3 . The other North American occurrence of 

Dendrostella recorded by Whiteaves (1892, p. 269- 

270) apears to be in rocks of Givetian age, as suggested 

by association with Stringocephalus at Lakes Manitoba and Winnipegosis. Glinski (1957, p. 105) records 

D. praerhenana as Eifelian in the lower Nohner beds, 

and D. rhenana as Givetian in the Biicheler beds, both 

in the Rhine Valley region. The horizon of romanensis 

is probably closer to Eifelian than Givetian.

Family HALLIIDAE Chapman, 1893

Reference form. Hallia insignis Edwards and 

Haime, 1850. Devonian, Ohio.

These medium and large solitary rugose corals have 

distinctly bilateral symmetry. Septa are numerous, 

long and smooth; cardinal septum is in a prominent 

fossula. Tabulae are wide, partly complete, normally 

domed with an axial sag. Dissepimentarium is narrow 

to wide; dissepiments are normal, globose, and lonsdaleioid. Septal thickening persists to adult growth 

stages in cardinal quadrants.

The Halliidae are the abundant Rugosa of the Great 

Basin Lower Devonian, where they characterize coral 

zones B and C; they diminish in importance in coral 

zone D.

Two subfamilies constitute the Halliidae: the Halliinae of the Early and Middle Devonian, and the 

Papiliophyllinae of the Early Devonian. The lykophyllid corals of the Silurian, which have been classified 

with the Halliidae by Hill (1956, p. F272), are here 

regarded as a separate family, the Lykophyllidae of 

Wedekind.

Subfamily HALLIINAE Chapman, 1893

Reference form. Hallia insignis Edwards and 

Haime, 1850. Devonian, Ohio.

The Halliidae have an especially well defined pinnate arrangement of septa in cardinal quadrants. The 

dissepimentarium, narrow to wide, is made up of normal dissepiments only.

Genera of this widely occurring subfamily are Hallia 

Edwards and Haime, Aulacophyllum Edwards and 

Haime, and Odontophyllum Simpson; of these, Aulacophyllum and Odontophyllum occur in the Great 

Basin Devonian coral zone C.

Subfamily HALLIINAE Chapman, 1893

Genus AULACOPHYLLUM Edwards and Haime, 1850

Aulacophyllum sp. c

Plate 9, figures 6-9

Medium and fairly large solitary corals of this kind 

occur in coral zone C at Lone Mountain in association 

with Papiliophyllum elegantulum and Odontophyllum 

meeki. Unlike Papiliophyllum they do not possess lonsdaleioid dissepiments, and accordingly they seem more 

appropriately assigned to Aulacophyllum. The strong 

thickening of cardinal quadrant septa persists into 

mature growth stages.

Occurrence. Nevada Formation, unit 1. Early 

Devonian, coral zone C. Lone Mountain, Eureka 

County, Nev.: Locality M74.

Other AulacophyHum-like Rugosa of coral zone C 

have thickened septa in contact laterally through the 

neanic stage but are largely devoid of dissepiments. 

These occur at locality M74 (pi. 9, figs. 10, 12) and in 

the southern Sulphur Spring Range at locality M36 

(pi. 9, fig. 11).
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 47

Genus ODONTOPHYLLUM Simpson, 1900 

1882. Aulacophyllum convergent Hall, p. 22. 

1833. Aulacophyllum convergent Hall, p. 281, pi. 17, figs. 1-2. 

1900. Odontophyllum Simpson, p. 210.

1938. Odontophyllum Simpson. Stewart, p. 31, pi. 5, figs. 5-6. 

1948b. Odontophyllum Simpson. Stumm, p. 51-59, pis. 1-2. 

1949. Odontophyllum Simpson. Stumm, p. 16, pi. 7, figs. 13-

14.

1956. Odontophyllum Simpson. Hill, p. F273, fig. 186-2a-c. 

1964. Odontophyllum Simpson. Stumm, p. 29-30, pi. 19, figs.

5-13.

Type species. Aulacophyllum convergent Hall by 

original designation. Middle Devonian; Beechwood 

Limestone Member of the Sellersburg Limestone, Falls 

of the Ohio, near Louisville, Kentucky (Stumm, 1964, 

P. 4).

Diagnosis. These solitary curved turbinate and 

curved trochoid to subpatellate Halliidae have a broad 

and commonly very deep calice. Cardinal septum is on 

the convex side in the long narrow fossula. Alar fossulae are defined, but no counter fossula. Major septa extend to the axis; septa are denticulate at distal edges. 

Some species have carinate septa. Tabulae are incomplete, distally arched. Dissepiments are present near 

the periphery in the late growth stages.

Remarks. Neanic stages of O. convergent (Hall) 

that are figured by Stumm (1948b, pi. 1, fig.ll) show 

their affinity with the Halliidae: having cardinal-counter bilateral symmetry, short cardinal septum, and 

septa of cardinal quadrants subparallel bordering the 

counter septum and bending inward toward the counter as the axis is approached. Longitudinal sections of 

individuals with a very broad shallower calice have 

tabularium with tabellae and a narrow dissepimentarium. Specimens that are here described from the 

lower part of the Nevada Formation have a deep calice 

extending downward to the nepionic growth so that 

transverse sections show little structure.

Odontophyllum meeki n. sp. 

Plate 9, figures 1-5

Type material. Holotype USNM 159261; paratype 

USNM 159262. Lone Mountain, locality M74; Devonian coral zone C.

Diagnosis. This Odontophyllum has a curved trochoid to curved turbinate corallum and a wide, deep 

calice extending downward to or almost to nepionic 

growth. Cardinal and alar fossulae are well developed. 

Cardinal septum are strongly developed and long in 

mature calices. Septa of counter quadrants are shorter 

and much less prominent in a mature calice than the 

septa of the cardinal quadrants. Some septa are 

slightly denticulate at the distal margins but have no 

development of true carinae. The distal margins of the 

calice are thin and sharp because there is no peripheral

platform as in some individuals of 0. patellatum 

(Holmes). The exterior has sharply defined septal 

grooves.

Remarks. This species shows extremely well the 

pinnate arrangement of septa and septal insertion with 

reference to primary septa. Associated with O. meeki 

are other nondissepimented solitary trochoid rugose 

corals having a deep calice and sharp rim, but with the 

fossulae not so well defined (plate 9, figs. 13, 14). 

These have narrowly cylindrical or ceratoid early 

growth stages which may be elongate, stemlike, and 

distorted. They differ from associated early growth 

stages of Papiliophyllum; though lacking dissepiments 

in the early growth stages, these have thickened septa, 

especially in the cardinal quadrants.

Measurements. Holotype: Diameter at calice rim 

30 mm, corallum length 34 mm, calice depth 26 mm; 

major septa at calice bottom 36.

Occurrence. Nevada Formation, Early Devonian, 

coral zone C. Lone Mountain, Eureka County, Nev.: 

locality M74.

Subfamily PAPILIOPHYLLINAE Stumm, 1949

Reference form. Papiliophyllum elegantulum 

Stumm, 1937. Lower Devonian, coral zone C, Great 

Basin.

These large Halliidae have a peripheral band of lonsdaleioid dissepiments; normal dissepiments are few or 

absent. Septal thickening is commonly strong in mature cardinal quadrants.

Genera of this endemic subfamily are Kobeha n. gen. 

of coral zone B, Papiliophyllum Stumm of coral zones 

C and Di, and Eurekaphyllum Stumm of coral zone C.

Genus KOBEHA, new genus

Type species. Kobeha walcotti n. sp., here designated.

Diagnosis. These large solitary Papiliophyllinae 

have curved trochoid and curved ceratoid to subcylindrical shape. The calice is deep, rather flat bottomed 

with nearly erect, straight walls and an acute margin 

without platform. The outer wall has a narrow septal 

stereozone. The cardinal fossula are well defined, usually on the convex side of the corallum. The major 

septa of mature individuals number about 60, some of 

which nearly reach the axis. Minor septa are short. In 

nepionic and early neanic stages, septa that are thickened by the stereome are laterally in contact, and 

largely filling the interior. In mature stages the septal 

dilation progressively decreases and becomes confined 

mostly to cardinal quadrants. Advanced stages have 

attenuate septa in counter quadrants and those of cardinal quadrants are also thinned, the attenuation is 

progressively peripherally from the axis. Tabularium is
48 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

broad, 60-80 percent of mature corallum width; tabulae are numerous, normally closely spaced. Some tabulae are continuous with the axial sag, others are as 

wide as the tabellae which are domed and discontinuous. The dissepimentarium has one to three columns of 

large steeply inclined dissepiments, some of which are 

exceedingly large wall blasts.

Because of the large size and small number of wall 

blasts and other large peripheral dissepiments, the 

transverse sections through the mature corallum do 

not show a conspicuous lonsdaleioid outer zone. The 

peripheral zone of large dissepiments is well shown 

only in longitudinal sections.

Remarks. Kobeha has thus far been recognized 

only in the central Great Basin. K. walcotti is the characterizing species of the Lower Devonian Kobeha zone, 

which is coral zone B of the central Great Basin Devonian. Kobeha ketophylloides n. sp. is placed in this 

new genus with reservation, but it seems to fit best 

here. Kobeha precedes, and is conceivably ancestral to, 

other Halliidae of this region, which include Papiliophyllum of coral zone C and Eurekaphyllum. Eurekaphyllum is regarded as a synonym of Papiliophyllum 

by some investigators (Hill, 1956, p. F273).

Kobeha walcotti n. gen., n. sp.

Plate 3, figures 1, 2, 6; plate 4, figures 1-4; 

plate 5, figures 1-7; plate 6, figures 8-10

1940. Zaphrentoid sp. cf. Papiliophyllum; Merriam, pi. 12, 

fig. 5.

Type material. Holotype USNM 159253; paratypes USNM 159251,159252,159256, 159257,159260. 

Nevada Formation, lower beds of unit 1; Devonian 

coral zone B, Early Devonian. Southern Sulphur 

Spring Range, Nev.

Diagnosis. Kobeha without continuous lonsdaleioid dissepimentarium showing as a complete peripheral band in mature transverse sections. Longitudinal 

sections show one to three columns of lonsdaleioid dissepiments, most of which are large wall blasts. All 

septa are thinned in late ephebic stages; major septa 

are discontinuous peripherally, but the outermost segments are uniform and complete at their union with 

the narrow septal stereozone. Major septa occur as 

partly discontinuous septal crests on the upper surfaces of tabulae. Tabulae are downwarped at the cardinal fossula, which is not confined in all individuals 

to the convex side of the corallum. Calice is nonconical 

with a broad bottom which is nearly flat to shallowly 

concave distally.

External features. Corallum is curved trochoid to 

ceratoid in the neanic stages; it becomes nearly cylindrical in mature stages of large individuals. Septal 

grooves are commonly separated by longitudinal 

ridges with a minor secondary groove on the crest of 

each ridge. Transverse features (rugae) vary from fine 

growth striations to fairly prominent ring-folds at irregular intervals. Regularly spaced rejuvenescence features are absent, but irregular breakage and repair 

structures are common and define the broken calice 

rim. Bell-shaped to flowerpot-shaped calice has an 

almost flat bottom and nearly erect sides below a 

sharp rim. The tabular floor of the calice is downwarped in the cardinal fossula, which may be situated 

either on the convex side or laterally.

Transverse sections. In advanced ephebic stages, 

the major septa number at least 70, all of which may 

be thinned. Stereoplasmic thickening of the septa in 

the cardinal quadrants persists into the earlier ephebic 

stages. The wall is a narrow septal stereozone. Some 

major septa reach the axis in the early ephebic stages; 

minor septa are short, usually less than one-seventh 

the length of the radius. Traces of large wall blasts are 

concave peripherally and encompass as many as six 

major septa. Between septa where the section intersects tabellae, the traces are usually concave peripherally; these features do not make a concentric pattern.

Late neanic to early ephebic transverse sections usually show the cardinal quadrants filled with stereoplasmic deposits and the septa with thickening in the 

counter quadrants. Septa are progressively thinned, 

beginning axially in the counter quadrants. Thickened 

septa in the cardinal quadrants are pinnate, but it is 

difficult to ascertain which of two or three shortened 

septa is actually the cardinal septum in some neanic 

sections. Late nepionic and early neanic transverse 

sections show stereoplasmic thickening of all septa; 

septa in contact laterally completely fill the corallum.

Measurements of Kobeha walcotti n. sp.

Calice bottom

USNM 

No.

159253 (holotype) ..............

159251 (paratype) ..............

159256 (paratype)..............

159257 (paratype) ..............

Corallum 

length 

restored 

(mm)

..................108

_

Calice 

depth 

(mm)

19 

23

22

Outside 

diameter 

(mm)

52 

45

57

Number of 

major 

septa

68 

66

Ephebic 

section

Diameter 

(mm)

53 

46

Number 

of 

major 

septa

64

58

Neanic 

section

Diameter 

(mm)

27

Number 

of 

major 

septa

50
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 49

Longitudinal sections. The tabularium, wide in 

mature longitudinal section, is as much as four-fifths 

of the total diameter. Complete or nearly complete 

tabulae commonly have a slight median sag, and wide 

tabellae. Tabulae are closely spaced to widely separated, downwarped at the tabularium margin, with a 

pronounced sag at the cardinal f ossula. Large lonsdaleioid dissepiments are normally in one or two columns, 

steeply inclined to nearly vertical. Axial extensions of 

major septa are carried on the upper surfaces of 

tabulae.

Comparison with related forms. Kobeha ketophylloides differs from K. walcotti by having larger less 

uniform wall blasts as observed in mature transverse 

sections, wherein these features are more prominent 

and more irregularly distributed. The number of dissepiment columns is greater in K. ketophylloides, and 

the mature dissepiments are less steeply inclined than 

in K. walcotti. In K. ketophylloides marked discontinuities of major septa where they meet wall blasts are 

more numerous.

Occurrence. Lower part of the Nevada Formation, 

basal beds of unit 1; Early Devonian, coral zone B with 

Costispirifer arenosus. Southern Sulphur Spring 

Range: localities M4, M56, M67, M69, M1041. Northern Roberts Mountains: locality M1072. Northern 

Sulphur Spring Range, McColley Canyon: locality 

M1018; specimens from this area differ subspecifically 

from typical K. walcotti.

Kobeha ketophylloides n. sp. 

Plate 3, figures 3-5, 8, 9; plate 6, figures 1-7

Type material. Holotype USNM 159258; paratypes USNM 159254, 159255, 159259. Lower Devonian, Nevada Formation, unit 1; coral zone B. Southern Roberts Mountains and Southern Sulphur Spring 

Range, Nev.

Diagnosis. Kobeha has very large and irregularly 

distributed wall blasts and lonsdaleioid dissepiments 

in mature transverse sections; these peripheral features are circumferentially uneven and show marked 

discontinuities of the major septa, segments of which 

thicken in wedge fashion to terminate peripherally 

against dissepiments or wall blasts. Individual wall

blasts subtend as many as 13 septa. Longitudinal sections have as many as seven columns of wall blasts and 

lonsdaleioid dissepiments which decrease distally in 

steepness of axial inclination to 40°.

External features. A large trochoid specimen with 

deep bell-shaped, rather flat-bottomed calice shows 

the cardinal fossula. Broken calice rim reveals the 

cone-in-cone structure with wide irregular separations 

making the characteristic wall blasts and lonsdaleioid 

dissepiments.

Transverse sections. In advanced ephebic stages 

major septa number 64. Minor septa are usually less 

than half the radius but are longer than those of K. 

walcotti. In advanced ephebic stages the septa are 

thicker at the periphery in counter quadrants than in 

K. walcotti and there are wide discontinuities where 

major septa meet large dissepiments at which points 

the septa thicken in wedge fashion locally. In later 

ephebic stages septa in cardinal quadrants exceed in 

thickness those of the counter quadrants; in earlier 

ephebic stages all major septa are of about the same 

thickness. Septa are thickened and laterally in contact 

in later nepionic stages, separating in early neanic 

stages; in late neanic stages septa of counter quadrants are thinner than those of cardinal quadrants. 

The outer wall has a septal stereozone which in late 

neanic stages is 2 mm thick. Large irregular dissepiments and wall blasts are concave peripherally.

Longitudinal sections. Most of the wide and rather 

closely spaced tabulae arched distally with a median 

sag, and are for the greater part nearly continuous. 

Short tabellae are uncommon. Large lonsdaleioid dissepiments are fairly even and regular in curvature longitudinally though irregular transversely; dissepiments decrease in steepness from early to late ephebic 

stages where the axial inclination is about 40°. The 

number of columns in the dissepimentarium increases 

to seven at maturity; this fact distinguishes the species 

from K. walcotti, the type species.

Reproductive offsets. A well-preserved silicified 

specimen assigned to K. ketophylloides shows four 

peripheral calice buds. The main corallite was itself 

developed as a calicinal offset from the partially preserved parent.

Measurements of Kobeha ketophylloides n. sp.

Ephebic section 

(early)

USNM 

No.

159258 (holotype) .................

159258 (holotype) .................

159259 (paratype) ................

159254 (paratype)................

Calice 

depth 

(mm)

_

............... 27

Calice 

bottom 

outside 

diameter 

(mm)

39

Diameter 

(mm)

55 

45 

32.5

Number of 

major 

septa

70 

60 

56

Neanic section

Diameter 

(nun)

30

Number of 

major 

septa

50
50 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OP THE CENTRAL GREAT BASIN

Comparison with related forms. Kobeha ketophylloides differs from K. walcotti in having a larger number of mature dissepiment columns, in its more conspicuous lonsdaleioid separations in advanced ephebic 

transverse sections, and longer minor septa. Heavy 

thickenings of septa in cardinal quadrants were not 

observed to persist ontogenetically into ephebic stages 

of K. ketophylloides as they do in some individuals of 

K. walcotti. The tendency for subcyclindrical growth 

in advanced ephebic stages is greater than in K. walcotti. Calice reproductive offsets were not recognized in 

K. walcotti.

In mature pattern of construction Kobeha ketophylloides resembles the species of Ketophyllum from 

the Silurian of Gotland, that is figured by Wedekind 

(1927) and the species from the Silurian of Czechoslovakia of Pocta (1902). The septal discontinuities 

of Ketophyllum are greater and lonsdaleioid separations more pronounced. Unlike Kobeha, the wide tabulae of Ketophyllum are usually rather flat, whereas 

those of Kobeha are strongly arched distally and have 

a median sag. Kobeha ketophylloides has thickened 

septa in the neanic and earlier growth stages, especially in the cardinal quadrants; the developmental 

thinning of the septa begins peripherally in the neanic 

growth within the counter quadrants, a characteristic 

feature of Kobeha and Papiliophyllum ontogeny. 

Wedekind's figures (1927, pi. 10, figs. 8-11) of the 

neanic stages of Ketophyllum elegantulum do not 

show thickened septa and the septa are short neanically, whereas in Kobeha ketophylloides the septa are 

thickened and approach the axis. These early developmental differences by themselves are believed sufficient to justify not assigning the Nevada forms to 

Ketophyllum.

Occurrence. Lower part of the Nevada Formation, 

lower beds of unit 1. Early Devonian, coral zone B, 

southern Roberts Mountains: type locality M1042. 

At the type locality, K. ketophylloides is associated 

with a Gypidula that resembles G. coeymanensis,Acrospirifer sp., Leptocoelia sp., and a brachipod that resembles the Pholidostrophia of the Rabbit Hill Helderberg. This horizon may be slightly older than the beds 

that contain specimens assigned to K. ketophylloides 

in the Sulphur Spring Range. Southern Sulphur 

Spring Range: localities M69, M1039.

Genus PAPILIOPHYLLUM Stumm, 1937 

1937. Papiliophyllum elegantulum Stumm, p. 430. 

1940. Papiliophyllum elegantulum Stumm, Merriam, p. 52,

53, pis. 12, 16.

1949. Papiliophyllum elegantulum Stumm, p. 16, pi. 7. 

1956. Not Papiliophyllum Hill, p. P274, fig. 186, 4c, 4d.

Type species. By original designation Papiliophyllum elegantulum Stumm (1937, p. 430, pi. 53, fig. 7,

pi. 54, fig. 7a-e). Lone Mountain, Eureka County, 

Nev.; lower part of Nevada Formation, unit 1, Lower 

Devonian (Emsian), coral zone C.

Diagnosis. Solitary trochoid and ceratoid Halliidae which grow to a large size have a deep flowerpotshaped, rather flat-bottomed calice with a cardinal 

fossula. Peripheral zone has medium to large steeply 

inclined lonsdaleioid dissepiments. Septa of cardinal 

quadrants are thickened in all the ephebic stages and 

are laterally in contact in all the early ephebic stages. 

Most septa are withdrawn from the axis and terminate 

peripherally at the inner margin of the lonsdaleioid 

dissepimentarium. Minor septa are either suppressed 

or inconspicuous as short discontinuous crests. In the 

neanic stages the cardinal septum is short within the 

fossula which is bordered by thick septa touching laterally. Tabulae are mostly complete, wide, close to 

rather widely spaced, and distally arched; in normal 

mature stages tabulae are rather flat to slightly sagging axially and periaxially and are peripherally depressed. Some individuals have tabulae that are rather 

irregular and sharply arched distally in the neanic 

stages. Tabulae are bent sharply downward, funnellike 

at the cardinal fossula. Septa of the counter quadrants become thinned and lonsdaleioid dissepiments 

develop in the early to mid-neanic stages; the cardinal 

quadrants are bordered by lonsdaleioid dissepiments 

externally until late neanic or early ephebic stages.

Remarks. The diagnosis here given is modified 

slightly from that of Stumm (1949, p. 16). Of special 

significance is the seeming absence of minor septa in 

some mature transverse sections, although in some 

specimens minor septa are recognizable as weak crests 

near the peripheral border of the tabularium. Also important taxonomically is the absence of lonsdaleioid 

dissepiments in the cardinal quadrants until the late 

neanic stage.

Papiliophyllum differs from Kobeha in having a better defined dissepimentarium with smaller and more 

numerous lonsdaleioid dissepiments, especially in the 

mature transverse sections. In Kobeha the septal 

crests carry through to the outer wall; they do not in 

Papiliophyllum. Minor septa are better defined and 

more continuous in Kobeha.

Papiliophyllum differs from Eurekaphyllum in possessing much more numerous and smaller lonsdaleioid 

dissepiments. Eurekaphyllum shows widely spaced 

tabulae with a pronounced axial-periaxial sag, whereas 

Papiliophyllum usually shows distally arched commonly close-set tabulae.

Papiliophyllum elegantulum Stumm 

Plate 7, figures 4-10; plate 8, figures 1-S
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 51

1937. Papiliophyllum elegantulum Stumm, p. 430; pi. 53, fig.

7, pi. 54, figs. 7a-e. 

1940. Papiliophyllum elegantulum Stumm. Merriam, p. 52-53,

pi. 12, fig. 3, pi. 16, figs. 1, 2. 

1949. Papiliophyllum elegantulum Stumm, p. 16; pi. 7, figs.

11, 12. 

1956. Papiliophyllum elegantulum Stumm. Hill, p. F274, fig.

186-4a-4b.

Type material. Stumm's transverse thin section of 

the holotype is similar to the transverse section of 

Merriam (1940, pi. 16, fig. 1) made by cellulose peel. 

The holotype is one of the fossils from the Nevada 

Formation presented by Merriam to the U.S. National 

Museum. Paratype, USNM 94448. Lower part of the 

Nevada Formation, unit 1, Lone Mountain, Eureka 

County, Nev.

Diagnosis. This is a large Papiliophyllum. The 

width of the mature lonsdaleioid dissepimentarium is 

less than half the radius and consists of three to seven 

columns of medium to large, steeply inclined dissepiments. Some tabulae are complete, arched distally. 

They are usually flattened axially and periaxially in 

the ephebic stages and commonly are arched more 

deeply and rather irregularly in the neanic stages. In 

advanced ephebic stages the dissepimentarium is narrowest at the cardinal fossula; the thickened septa 

bounding the fossula invade this peripheral band. 

Minor septa are weakly developed or unrecognizable. 

Septa of the cardinal quadrants are heavily to moderately thickened in the ephebic stages.

External features. These curved trochoid or curved 

ceratoid corals usually have well-defined septal grooves 

but commonly do not show very pronounced annular 

folds or rugae; these features suggest fairly continuous 

growth. The outer wall is commonly broken away in 

the places where large wall blasts occur within. The 

deep and rather flat-bottomed calice is bordered in 

mature specimens by blisterlike lonsdaleioid dissepiments; on the axial surfaces of these dissepiments lie 

septal crests that become weaker toward the rim. 

There is no calicular shelf.

Transverse sections. Mature transverse sections 

have an average of about 56 major septa, some of which 

reach or nearly reach the axis. Minor septa occur as

thin discontinuous crests toward the peripheral edge 

of the tabularium; some mature individuals show 

almost no trace of minor septa. Septal crests are rarely 

observed within the dissepimentarium and almost 

never meet the wall. In advanced ephebic stages the 

width of the dissepimentarium varies from less than 

one-third to more than one-third of the radius. The 

dissepiments lack uniformity, ranging in size from 

small to very large and they subtend 12 or more septa. 

In late ephebic stages septa of the counter quadrants 

are always thinned; those of the cardinal quadrants 

are thickened and commonly several remain laterally 

in contact. The cardinal septum is shortened or 

aborted in a fossula which encroaches peripherally 

upon the dissepimentarium.

In the mid-neanic stages there are about 38 septa. 

The septa vary from being all thickened and laterally 

in contact with one another to being considerably 

thinned in the counter quadrants, and some of those in 

the cardinal quadrants are partly separated even 

though they are thick. At the mid-neanic stage, but 

usually a bit later, lonsdaleioid wall blasts begin to 

develop in the counter quadrants only. The cardinal 

septum roughly defines a plane of symmetry; on either 

side of this plane the thickened septa are more or less 

parallel near the periphery but bend laterally toward 

this plane as they pass axially, some join near the axis 

to produce a primitive fossula. Counter and alar septa 

are not distinguishable with assurance. In early ephebic stages lonsdaleioid dissepiments are generally not 

developed or are just beginning to develop in the cardinal quadrants, and the thick septa are here largely 

or partly in contact laterally.

Longitudinal sections. In advanced ephebic stages 

the tabularium width exceeds one-half the corallite 

diameter. Sections parallel to the alar plane show 

fairly uniform, distally arched tabulae, some of which 

are complete and accompanied by wide tabellae. The 

broad axial-periaxial part of the tabularium is flat or 

has a slight sag, the peripheral part sags abruptly. In 

neanic stages the tabular uparching is more pronounced and axially convex to angular distally. Medial 

sections through the cardinal fossula reveal the exMeasurements of Papiliophyllum elegantulum Stumm

Figured 

specimen

1 .....................

2 .....................

3 .....................

4 .....................

5 .....................

Corallum 

length 

restored 

(mm)

... 51

Calice 

depth 

(mm)

22

Calice 

bottom 

outside 

diameter 

(mm)

23

Ephebic £

Diameter 

(mm)

82 

46 

45

section

Number of 

major 

septa

68 

52

58

Neanic section

Diameter 

(mm)

19.8 

21

Number of 

major 

septa

44 

38
52 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

tensive stereoplasmic septal thickening and the pronounced funnellike proximal sag of tabulae associated 

with the fossula. Columns of lonsdaleioid dissepiments 

number seven or more in the advanced ephebic stages; 

dissepiments vary greatly in size, and smaller ones are 

scattered throughout. Very large wall blasts are common, but large dissepiments seen in some specimens 

may also occur at the axial margin of this band.

Comparison with related forms. Papiliophyllum 

elegantulum seems to have evolved from the same 

stock that also gave rise to Kobeha and Eurekaphyllum, which are discussed elsewhere under their definitions of genera. Papiliophyllum elegantulum differs 

from Kobeha walcotti in having smaller and more numerous lonsdaleioid dissepiments and in not having 

septal crests within the dissepimentarium the absence of these crests is a diagnostic feature of K. walcotti. Eurekaphyllum breviseptatum differs in having 

only a few very large wall blasts and lonsdaleioid dissepiments and in having its very widely spaced tabulae 

which are depressed proximally, not arched distally as 

in P. elegantulum.

Occurrence. Lower part of the Nevada Formation, 

unit 1; Devonian coral zone C of late Early Devonian 

age, approximately Emsian. Associated with Acrospirifer kobehana. Lone Mountain: localities M74, M286, 

M1044. Southern Roberts Mountains: locality 18 of 

Merriam (1940). Northern Roberts Mountains: localities 13, 94 of Merriam (1940). Southern Sulphur 

Spring Range: locality M68. North end of Antelope 

Range: locality Ml035a. Hot Creek Range: locality 

M1066?. Northern Panamint Mountains, California: 

localities M184, M1065. Funeral Mountains, California: localities M1059, M1063.

Papiliophyllum elegantulum subsp. d 

Plate 7, figures 1, 2

This subspecies is insufficiently known to characterize. It is an evolutionally advanced and a geologically 

younger derivative of the elegantulum lineage with 

very numerous lonsdaleioid dissepiments and crowded 

fairly uniform wide tabulae that are only slightly 

arched.

Measurements. Diameter of the late ephebic section is 78 mm; the number of major septa is 62.

Occurrence. Lower part of the Nevada Formation, 

unit 2. Devonian coral zone Di of late Early Devonian 

age. Lone Mountain: locality M1037; associated with 

Sinospongophyllum sp. d, Chonetes macrostriata, 

Gypidula loweryi, Spirifer pinyonensis, and Dalmanites meeki. Ranger Mountains, Clark County, Nev.; 

a similar Papiliophyllum occurs here at locality M1034, 

associated with a fauna which may be that of coral 

zone DI.

Genus EUREKAPHYLLUM Stumm, 1937 

1937. Eurekaphyllum breviseptatum Stumm, p. 431, pi. 53,

fig. 8; pi. 54, figs. 8a-b. 

1940. Eurekaphyllum Stumm. Lang, Smith, and Thomas, p.

58.

1949. Eurekaphyllum Stumm, p. 17, pi. 7, figs. 15-17. 

1956. Papiliophyllum Stumm (in part). Hill, p. F273 (in

part), fig. 186-4c-d.

Type species. Eurekaphyllum breviseptatum 

Stumm (by original designation). Lower part of the 

Nevada Formation, Lone Montain, Eureka County, 

Nev.

Diagnosis. Large curved trochoid Papiliophyllumlike rugose coral with a peripheral zone of very large 

irregular lonsdaleioid dissepiments and widely spaced 

irregular wide tabulae which are axially depressed. 

Inner ends of the septa are withdrawn some distance 

from the axis; septa of the cardinal quadrants are 

thickened. Septal crests are present in the dissepimentarium.

Remarks. Eurekaphyllum has the lonsdaleioid dissepiments and thickened mature cardinal quadrant 

septa of Papiliophyllum and Kobeha, but differs from 

both in possessing exceedingly large irregular dissepiments and in having a small number of tabulae, most 

of which sag proximally. In transverse section, Eurekaphyllum resembles Kobeha in having septal crests 

within the dissepimentarium; Papiliophyllum commonly lacks these outer crests and differs in having 

a much greater number of dissepiment columns.

Eurekaphyllum breviseptatum Stumm 

Plate 8, figures 9, 10

1937. Eurekaphyllum breviseptatum Stumm, p. 431, pi. 53,

fig. 8, pi. 54, figs. 8a-b. 

1949. Eurekaphyllum breviseptatum Stumm, p. 17, pi. 7, figs.

15-17.

Type material. Holotype, USNM 94449. Lower 

part of the Nevada Formation; Lone Mountain, Eureka County, Nev. This is the only known specimen of 

Eurekaphyllum.

Diagnosis. See generic diagnosis.

Transverse section. Septa number about 64; all are 

considerably withdrawn from the axis; minor septa, 

if distinguishable, exceed half the length of the major 

septa. Septa of the cardinal quadrants are thickened. 

Some septa extend discontinuously to the outer wall 

as thin crests. Width of the dissepimentarium is about 

equal to the length of the major septa; most dissepiments are very large, irregular and nonuniform; the 

longest extends one-fourth of the circumference. Fossula is indistinct.

Longitudinal section. Tabulae are few; some are 

complete, the more nearly complete are widely separated and have a pronounced U-shaped sag with a
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 53

somewhat flattened axial-periaxial segment. Large dissepiments of irregular form are in one or two columns.

Occurrence. Lower part of the Nevada Formation, 

Lone Mountain, Eureka County, Nev. The horizon 

from which this form came is unknown; it probably 

came from unit 1, coral zone C, which is the zone in 

which Papiliophyllum is the abundant rugose coral.

Hill (1956, p. F273-F274) regards Eurekaphyllum 

as a junior synonym of Papiliophyllum. Study of a very 

large suite of specimens shows that the Eurekaphyllum structure is not normal or modal for the type of 

coral here placed in Papiliophyllum. As no additional 

representatives of Eurekaphyllum have been collected 

from the beds which probably yielded the type and 

only specimen, it seems likely that this form may be 

an aberrant and uncommon development from the 

Papiliophyllum stock. There are no intergradational 

forms, morphologically speaking. Without knowledge 

of the exact horizon or zone from which the type came, 

it is believed desirable to let the genus in question 

stand for the present. Like Kobeha it could well be 

found to have stratigraphic as well as morphologic 

significance.

Family BETHANYPHYLLIDAE Stumm, 1949

Reference form. Bethanyphyllum robustum (Hall) 

1876. Middle Devonian, New York.

Medium and large trochoid to ceratoid rugose corals 

with long, thin major septa and a cardinal fossula. 

Multiple columns of medium to small normal globose 

dissepiments occur in a wide dissepimentarium. Tabularium is medium or wide, consisting of combinations 

of straight, nearly complete tabulae and tabellae. 

Septa of the cardinal quadrants are unthickened beyond the nepionic stages.

Bethanyphyllidae may have sporadic zigzag carinae, 

but these are uncommon; in some species the fossula 

is weakly developed. The dissepimentarium-tabularium 

margin is usually not sharply defined. No lonsdaleioid 

dissepiments are known.

This family designation is appropriate for a stratigraphically important early Middle Devonian coral 

group possibly evolving from the ancestral stock of the 

Halliidae which precede Bethanyphyllidae in the Devonian of the Great Basin.

In a recent revision, the Family Bethanyphyllidae 

has been abandoned, Bethanyphyllum being assigned 

to the Zaphrentidae, together with Heliophyllum 

(Stumm, 1963, p. 139; 1964, p. 38). Pedder (1965, p. 

207) has emended the Bethanyphyllidae to accommodate Bethanyphyllum, Ceratophyllum, Moravophyllum, Tortophyllum, and Stathmoelasma. In these genera, septa are smooth or weakly carinate, unlike Heliophyllum which has strong yardarm carinae. The cardinal septum is relatively short. In mature stages it is 

difficult to recognize the cardinal fossula of some species; this is one of several features which distinguishes 

Bethanyphyllidae from Halliidae.

Genus BETHANYPHYLLUM Stumm, 1949

1876. Cyathophyllum robustum Hall, pi. 22, figs. 1-14.

1937. Cyathophyllum? lonense Stumm, p. 435, pi. 55, figs.

3a-b. 

1937. Grypophyllum giganteum Stumm, p. 433, pi. 54, figs.

9a-b. 

1937. Grypophyllum curviseptatum Stumm, p. 433, pi. 55,

figs. 2a-b. 

1937. Grypophyllum nevadense Stumm, p. 432, pi. 53, fig. 9,

pi. 55, figs. la-b. 

1937. Tabulophyllum nevadense Stumm, p. 434, pi. 55, figs.

4a-b.

1949. Bethanyphyllum Stumm, p. 18, pi. 8, figs. 1-4. 

1956. Bethanyphyllum Hill, p. F278, figs. 189-6a-b. 

1963. Bethanyphyllum Stumm, p. 139. 

1965. (?) Stathmoelasma Pedder, pi. 31, figs. 1-5.

Type species. Cyathophyllum robustum Hall, 

1876, by original designation. Middle Devonian; Hamilton Group of western New York State.

Diagnosis. Medium to large solitary trochoid to 

cylindrical rugose corals have numerous long, thin, 

usually fairly smooth septa. The longer septa reach the 

axis. Cardinal fossula is well defined to weak, without 

pronounced pinnate arrangement of several bordering 

septa. The dissepimentarium is wide, with multiple 

columns of steeply inclined mostly small dissepiments. 

The tabularium is wide or medium wide, consisting of 

a variable complex of partial to nearly complete, 

straight or bowed tabulae that are peripherally bordered by large tabellae passing into dissepiments laterally. Conspicuous vertical gaps commonly separate 

the tabluae. Outer wall is thin, lacking a peripherally 

continuous septal stereozone.

Remarks. Bethanyphyllum lacks the more evident 

bilateral arrangement of septa characterizing Aulacophyllum and usually has no axial whorl like Tortophyllum. Some species assigned to Bethanyphyllum 

have septal thickening in the initial cardinal quadrant 

which persists well into the neanic stages (B. antelopensis). In mature stages, however, the septa of Bethanyphyllum are normally unthickened in cardinal 

quadrants alone, one of several features which distinguish this genus from the Papiliophyllum stock. If 

there is septal thickening in the late stages of Bethanyphyllum, it affects especially the axial parts of major 

septa in any or all quadrants. Bethanyphyllum also 

lacks the large lonsdaleioid dissepiments of Papiliophyllum. Carinae are absent in most species of Bethanyphyllum', those species that do have scattered carinae, like B. antelopensis, are small and of the zigzag 

thickening type rather than the true yardarm carinae.
54 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Stathmoelasma, recently described by Pedder 

(1965, p. 207, pi. 31, figs. 1-5, pi. 32, fig. 10) from the 

Australian Devonian, is similar to Bethanyphyllum, of 

which it may be a synonym. Pedder's figures show 

indications of a weak fossula, and the longitudinal 

sections reveal nearly complete straight to slightly 

bowed tabulae like those of Bethanyphyllum, with 

conspicuous gaps. There is no septal stereozone.

Bethanyphyllum of coral zone D2 is preceded by 

Papiliophyllum of coral zone C, which ranges up into 

coral zone D^ Papiliophyllum has not been found in 

association with the more typical Bethanyphyllum, as 

exemplified by B. lonense (Stumm). However, in coral 

zone D! Papiliophyllum individuals with intermediate 

features occur with advanced; these individuals reveal 

minor mature septal thickening in the cardinal quadrant and in some specimens a few rather inconspicuous 

lonsdaleioid dissepiments are present. Whereas these 

specimens suggest development toward Bethanyphyllum, they appear on the whole to be somewhat closer 

to the Papiliophyllum stock.

Bethanyphyllum lonense (Stumm)

Plate 11, figures 1-4; plate 12, figures 3-10

1937. Cyathophyllum? lonense Stumm, p. 435, pi. 55, figs. 3a,

b. 

Tabulophyllum nevadense Stumm, p. 434, pi. 55, figs.

4a, b. 

1940. Cyathophyllum sp. Merriam, pi. 12, figs. 2, 4.

Type material. Holotype, USNM 94453; lower 

part of the Nevada Formation, Lone Mountain, 

Eureka County, Nev.

Diagnosis. Bethanyphyllum has thin, commonly 

rather wavy septa which may be slightly thickened 

toward the axis. Tabulae varying from straight, wide, 

and nearly complete to large and small tabellae. More 

continuous straight tabulae and larger tabellae bound 

the bigger longitudinal gaps. The dissepimentarium 

has several columns of rather small and steeply inclined dissepiments. The fossula is generally unrecognizable in mature growth stages.

External features. Poorly known. This form seemingly does not become very large and tends to remain 

trochoid with a very deep calice.

Transverse sections. Major septa of the advanced 

growth stages average 36, some of which approach or 

reach the axis in mature sections. Minor septa are 

usually less than half the radius. Septa are thin and 

wavy, never entirely smooth, with abrupt angular 

bends. Some septa may be slightly thickened axially, 

where they may be contorted but not twisted to form 

an axial whorl. Between septa, the dissepiment traces 

and tabellar intersections are concentric and either 

evenly concave axially or occur as chevrons. The outer 

wall is thin, without appreciable stereoplasmic thickening. Neanic and ephebic sections show no fossula 

or cardinal septum. No nepionic or early neanic sections are available, but presumably the septa of cardinal quadrants are thickened, to judge from the related species Bethanyphyllum antelopensis.

Longitudinal sections. The tabulariums of ephebic 

stages vary considerably in width within the corallum; 

they usually exceed half the diameter. The presence 

of a few straight, wide, and nearly continuous tabulae 

is characteristic, these tabulae separate longitudinal 

intervals of tabellae and shorter tabulae and define 

larger vertical gaps. Dissepiments, mostly small and 

globose, are disposed in numerous columns bounded 

axially by larger, less globose dissepiments or peripheral tabellae.

Comparison with related forms. Bethanyphyllum 

lonensis differs from B. antelopensis in having a wider 

tabularium and shorter minor septa; major septa are 

more sinuous in B. lonensis. Some mature individuals 

of B. antelopensis have a cardinal fossula not observed 

in lonensis. B. robustum (Hall) differs in having a 

weakly defined ephebic cardinal fossula, more even 

septa, and relatively larger peripheral dissepiments.

Measurements of Bethanyphyllum lonense Stumm

Ephebic section

Figured 

specimen

USNM 

No.

Diameter 

(mm)

Number of 

septa

.159272

.159273

.159277

.159278

159278

30

24

31.5

124

13

40

34

38

134

26

1Early ephebic.

Occurrence. Nevada Formation, unit 2. Devonian 

coral zone D2 ; in association with Siphonophrentis 

(Breviphrentis) invaginatus. Lone Mountain: locality 

58 (Merriam, 1940), localities M55, M1036, M1048. 

Northern Roberts Mountains: locality 11 (Merriam, 

1940). Southern Sulphur Spring Range: locality 

M1047. Northern Fish Creek Range, Grays Canyon: 

localities M3, M51.

Bethanyphyllum antelopensis n. sp.

Plate 10, figures 1-7; plate 11, figures 5, 6; 

plate 12, figure 11

Type material. Holotype, USNM 159265; paratypes, USNM 159266-159269, 159274. Lower part of 

the Nevada Formation, Combs Peak area, Eureka 

County, Nev.

Diagnosis. This Bethanyphyllum has long major 

septa, commonly about two-thirds as long as the major 

septa, usually with a correspondingly wide dissepimentarium, and with several axially-periaxially straight 

tabulae separated by wide vertical gaps. The septa are 

thin except for the axial part which may be somewhat
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 55

thickened; septa show only moderate waviness or irregularity. A fossula is present in some mature individuals.

External features. Inadequately known.

Transverse sections. Major septa average about 38 

in mature sections. Septa are thin, only slightly wavy, 

and for the greater part approach the axis toward 

which they may be slightly thickened, especially in 

the cardinal quadrants. Minor septa are long, in some 

sections two-thirds the length of major septa. In some 

specimens, a fossula may be defined by a bending of 

pairs of septa at their axial ends toward the symmetry 

plane of the cardinal septum, which, if correctly identified, remains fairly long in mature stages. Dissepiments between septa appear to be axially concave, and 

chevron features make a concentric pattern. Outer wall 

appears to be thin and without stereozone. Septa of 

cardinal quadrants are very much thickened and in 

lateral contact through to mid-neanic stages. One 

neanic section with 30 major septa shows thick cardinal quadrant septa laterally in contact except near the 

alar positions; the cardinal septa are arranged on 

either side of a somewhat shortened cardinal septum. 

Major septa are withdrawn slightly from the axis. 

Within the tabularium, the counter quadrant septa 

are also thickened slightly. There are rows of concentric dissepiments in the counter quadrants, but none 

were recognized in the cardinal quadrants. Minor septa 

are slightly less than half the length of the majors. The 

wall is unthickened by stereoplasm.

Longitudinal sections. The tabularium, usually 

rather narrow, ranges from one-fourth to nearly onehalf the corallite diameter in the mature growth stages. 

Nearly straight, almost complete tabulae define vertical intervals separated by intervals of less complete 

tabulae and peripheral tabellae. The more complete 

straight tabulae commonly occur where the vertical 

separation is greater. Dissepimentarium has numerous 

columns of steeply inclined, small to medium and large 

dissepiments, of which the smaller predominate.

Comparison with related forms. See Bethanyphyllum lonense.

Measurements of Bethanyphyllum antelopensis n. sp.

Ephebic section Neanic section

USNM 

No.

Diameter 

(mm)

Number of 

major 

septa

Diameter 

(mm)

Number of 

major 

septa

159265 (holotype) ....... 

159266 (paratype) ....... 

159269 (paratype) .......

159274 (paratype) .......

. 39 

. 35

. 29

38 

44

32

14.6 30

Occurrence. Lower part of the Nevada Formation, 

unit 2; Middle Devonian, Devonian coral zone D2 . 

Locality M27, 1.75 miles south of Combs Peak at a

horizon about 270 feet stratigraphically above the base 

of the Nevada Formation and just below a 15-foot 

quartzite bed of Oxyoke Canyon type. The associated 

fauna of the Spirifer pinyonensis zone includes abundant <(Martinia" undifera.

Similar corals provisionally assigned to B. antelopensis (pi. 11, figs. 7-9) occur at Lone Mountain 

(locality Ml045) in the upper part of unit 2, possibly 

as high as Devonian coral zone D3

Bethanyphyllum sp. d 

Plate 10, figures 8-10; plate 13, figures 6, 7, 9

Figured specimens. USNM 159270, 159271, 

159281, 159281a. Lower part of the Nevada Formation, Lone Mountain, Eureka County, Nev.

Specimens classified as Bethanyphyllum sp. d differ 

from B. lonense and B. antelopensis in features of the 

tabularium; they are wider, and the tabulae are more 

closely spaced with a tendency toward distal arching. 

The wide vertical gaps between some tabulae of lonense and antelopensis were not found in sp. d. It is 

probable that sp. d lacks the very deep calice of some 

individuals of lonense.

Occurrence. Lower part of the Nevada Formation, 

lower beds of unit 2; Devonian coral zone DI. Lone 

Mountain, locality M1037, where it occurs with the 

fa ma including the advanced Papiliophyllum elegantulum subsp. d.

Family CHONOPHYLLIDAE Holmes. 1887

Reference form. Chonophyllum Edwards and 

Haime, 1850.

These solitary rugose corals are of medium to large 

size, dissepiments partly or wholly lonsdaleioid; some 

tabulae are complete, flat or domed. Major septa are 

lamellar, elongate, discontinuous peripherally, little 

thickened. Fossula is weak or absent.

Genera included in this family are as follows:

Chonophyllum Edwards and Haime, 1850 

Ketophyllum Wedekind, 1927 

Tabulophyllum Fenton and Fenton, 1924 

Sinospongophyllum Yoh, 1937 

Diversophyllum Sloss, 1939

The Chonophyllidae are commonly quite rugate 

externally because of their numerous rejuvenescence 

flanges and cone-in-cone growth habit. The septal 

stereozone is usually not a conspicuous feature, and 

the tabularium ranges from narrow in Chonophyllum 

to wide or very wide in the other genera.

Genus SINOSPONGOPHYLLUM Yoh. 1937 

1937. Sinospongophyllum planotabulatum Yoh, p. 56, pi. vi,

figs. 2-5. 

1940. Sinospongophyllum Yoh. Lang, Smith, and Thomas, p.

119. 

1949. Tabulophyllum Stumm, p. 27 (in part); pi. 12, figs. 20,

21.
56 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

1956. Sinospongophyllum Yoh. Hill, p. F300, fig. 204, 5a-b.

Type species. Sinospongophyllum planotabulatum 

Yoh (by original designation). Upper part of the middle Devonian; Kwangsi, China.

Diagnosis. This ceratoid to subcylindrical Chonophyllidae has rather closely spaced, wide to very wide 

tabulae which are either straight or sag axially and 

peripherally. Septa are amplexoid, smooth or minutely 

wavy, noncarinate, and usually withdrawn from axis. 

Minor septa are well defined. Peripheral zone has septal crests, large irregularly distributed lonsdaleoid dissepiments, and smaller normal dissepiments. Septal 

stereozone is narrow to moderately wide. Fossula is 

weak or absent.

Remarks. Yoh's figures and the type show very 

steeply inclined dissepiments in a rather narrow band, 

very wide, flattened tabulae, and no fossula.

Diversophyllum (Sloss, 1939, p. 65) differs in having 

longer septa that reach the axis, more arched and 

irregular tabulae, and less tendency to produce lonsdaleioid dissepiments.

Tabulophyllum, as typified by the Upper Devonian 

T. rectum, differs from Middle Devonian Sinospongophyllum in having weakly developed minor septa and 

a less evident septal stereozone. However, the two 

genera have much in common, and Sinospongophyllum 

is regarded by Stumm (1949, p. 27) as a possible synonym. For the present, however, it seems desirable to 

retain Sinospongophyllum, pending more thin section 

study of topotype material of both species. Externally 

the eleven described Iowa Late Devonian species of 

Tabulophyllum (Fenton and Fenton, 1924) show a 

great variety of form, from turbinate to subcylindrical, 

and some with wide dissepimentarium.

In the Cordilleran belt, corals assignable to either 

Sinospongophyllum or Tabulophyllum are characteristic of late Middle and Late Devonian rocks in the 

Great Basin, southeast Alaska, and northwest Canada 

(Smith, 1945). They occupy Great Basin late Middle 

Devonian coral zone F in association with Hexagonaria. Those corals of lowermost Middle Devonian coral 

zone D that are placed in Sinospongophyllum have a 

very wide tabularium, and some specimens have only 

a few sporadic lonsdaleioid dissepiments. These coral 

zone D specimens occur with Siphonophrentis (Breviphrentis) invaginatus, variants of which they resemble. Stumm's (1949, p. 25) Breviphyllum that was 

founded upon Nevada Formation material was possibly inspired by examination of such associated material; the holotype is seemingly without dissepiments 

and probably represents a specimen of invaginatus. 

Possibly such sparingly dissepimented zone D corals 

may be derived from the Breviphrentis invaginatus 

stock by the aberrant development of dissepiments.

At present they are insufficiently known for specific 

description and accordingly are designated informally 

by letter.

Sinospongophyllum sp. d 

Plate 17, figures 1, 2, 5, 6

Figured specimens. USNM 159312, 159314; lower 

part of the Nevada Formation, Lone Mountain and 

northern Antelope Range, Eureka County, Nev.

The lonsdaleioid dissepiments are large, elongate, 

and irregularly distributed in one or two columns. The 

tabulae are closely spaced. The septal stereozone is 

narrow.

Occurrence. Late Early Devonian, coral zone DI. 

Locality Ml037, south pediment of Lone Mountain, 

Eureka County, Nev. Early Middle Devonian, coral 

zone D2 . Locality Ml035, north end Antelope Range, 

southern Eureka County, Nev.

Sinospongophyllum sp. e 

Plate 17, figures 3, 4,10,11

1937. (?) Amplexus nevadensis Stumm, p. 427, pi. 54, figs. 3a-c. 

1937. (?) Amplexus magnus Stumm, p. 428, pi. 54, figs. 5a-b.

Figured specimens. USNM 159313, 159317; lower 

part of the Nevada Formation, Lone Mountain and 

northern Antelope Range, Eureka County, Nev.

The lonsdaleioid dissepiments are small and irregularly distributed in one to several columns. The tabulae are closely spaced. The septal stereozone is narrow. 

Major septa are withdrawn from the axis.

Occurrence. Early Middle Devonian, coral zone D2 . 

LocalityM1038, Lone Mountain, Eureka County, Nev. 

Locality Ml035, north end Antelope Range, southern 

Eureka County, Nev.

Sinospongophyllum sp. f 

Plate 17, figures 7, 8, 9

Figured specimens. USNM 159315, 159316; lower 

part of the Nevada Formation, Grays Canyon and 

northern Antelope Range, Eureka County, Nev.

The lonsdaleioid dissepiments are small and irregularly distributed in one to several columns. The tabulae 

are closely spaced. The septal stereozone is narrow to 

fairly wide. Major septa extend to the axis where they 

are twisted together in an axial plexus.

Occurrence. Early Middle Devonian, coral zone 

Do. Locality M3, Grays Canyon, southern Eureka 

district, Nev.: Locality M1035, north end Antelope 

Range, southern Eureka County, Nev.

Family ENDOPHYLLIDAE Torley, 1933 

Reference forms. Endophyllum bowerbanki 

Edwards and Haime and E. abditum Edwards and 

Haime, 1851. Devonian; Torquay, England.

These cerioid and aphroid rugose corals have wide
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 57

corallites, a broad marginarium of partly large lonsdaleioid dissepiments and a narrow to very wide tabularium comprising closely spaced tabulae. There is no 

axial structure.

Genera provisonally assigned to the Endophyllidae 

are Endophyllum Edwards and Haime, 1851, Yassia 

Jones, 1930, and Australophyllum Stumm, 1949. 

Undescribed Late Silurian corals in the Great Basin 

and the Klamath Mountains are related to Australophyllum; the Klamath forms are a separate genus, and 

those of the Great Basin a subgenus of Australophyllum.

Some corals here regarded as Endophyllidae have 

previously been assigned to Spongophyllum, a genus 

not included in this family. It is proposed that the 

term Spongophyllum be applied to species with slender 

corallites agreeing in general structure and proportions 

with the Devonian S. sedgwicki as originally illustrated 

by Edwards and Haime (1853, pi. 56, figs. 2, 2a-c, 2e).

In addition to the wide lonsdaleioid marginarium, 

Late Silurian representatives of Yassia and a subgenus 

of Australophyllum manifest a decided tendency to 

abbreviate and lose septa in mature growth stages. 

These characteristics are not observed in Spongophyllum.

Genus AUSTRALOPHYLLUM Stumm, 1949 

1911. Spongophyllum cyathophylloides Etheridge, p. 7-8, pi.

A, fig. 3, pi. C, figs. 1-2.

1949. Australophyllum Stumm, p. 34, pi. 16, figs. 1-2. 

1956. (?) Australophyllum cyathophylloides (Etheridge). 

Hill, fig. 207-4a-b.

Type species. Spongophyllum cyathophylloides 

Etheridge; by author designation. "Lower Middle 

Devonian"; Douglas Creek, Clermont, Queensland, 

Australia.

Diagnosis. The cerloid Endophyllidae have 

medium wide to narrow, closely spaced, proximally 

sagging tabulae. The wide to very wide dissepimentarium has scattered to wholly lonsdaleioid dissepiments, some of which are large.

Remarks. Australophyllum differs from Spongophyllum in possessing multiple columns of elongate, 

lonsdaleioid dissepiments, and more closely spaced 

sagging tabulae which lack a peripheral depression. 

Stumm's diagnosis of Australophyllum includes carinate septa, although his figures do not show these 

convincingly. The wall of typical Australophyllum is 

somewhat thickened stereoplasmically; septal crests 

do not appear to be characteristic, as they are in the 

Silurian subgenus.

Australophyllum landerensis n. sp. 

Plate 25, figures 1-4

Type material Holotype, USNM 159353; paratype, USNM 159354. Lower Devonian Rabbit Hill 

Limestone, Toquima Range, locality Ml 150.

Diagnosis. This Australophyllum has long minor 

septa; all septa are thickened in the tabularium. The 

outer dissepiments are steeply inclined for the genus; 

the outer wall is thickened and beaded with wall 

crests. The tabularium is medium wide; close-set tabulae have a pronounced sag.

External features. Occurs in compact ceriod heads 

up to 15 cm or more in diameter.

Transverse sections. Septa are about 44, all of 

which are long and usually thickened within the tabularium; minor septa are more than one-half the length 

of major septa. Some major septa reach the axis where 

they are twisted. Some septa may be minutely wavy 

and, where thickened, show minute nodes and lateral 

bumps, but there are no true elbow carinae. The stereome-thickened wall has a beaded appearance because 

of the wall crests; other septal crests are sparse in the 

lonsdaleioid band.

Longitudinal sections. The tabularium, about onethird the corallite width at maturity, is very sharply 

set off from the dissepimentarium. The tabulae, 4 or 5 

per millimeter, have a pronounced sag which may be 

angulate axially. Large outer dissepiments are steep 

for this genus; the smaller ones become vertical at the 

tabularium margin. Dissepiment columns range from 

7 to 12. Thickened axial segments of major septa 

reveal minute lateral bumps.

Reproductive offsets. One mature corallite shows 

three calice wall offsets internally situated.

Fine structure. Some thickened septa may have a 

median clear trace; trabecular structure and lamination are poorly defined in the septal stereozone.

Comparison with related forms. Australophyllum 

cyathophylloides, the type species, differs in having 

fewer septa which lack thickening in the tabularium. 

Australophyllum sp. (plate 25, fig. 5) from the upper 

part of the Vaughn Gulch Limestone, Owens Valley, 

Calif, (locality M1093), resembles landerensis but 

lacks the thickened septa. The Vaughn Gulch form 

occurs in beds of Late Silurian or Early Devonian age 

that are immediately above the Late Silurian coral 

zone E which contains an Australophyllum so different 

that it is placed in a new subgenus. The form from 

Silurian coral zone E has broader mature corallites 

with a considerably smaller septal count and lacks 

septal thickening; this form has outer lonsdaleioid dissepiments inclined at a low angle, a larger number of 

very large dissepiments and, unlike landerensis, tends 

to reduce and lose septa at maturity.

Measurements. Mature corallite diameter and 

septum count:
58 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Corallites .......................... A

Diameter (in mm) ..........13

Major septa ......................22

Holotype 

USNM 159S5S

B

14

22

C

10

22

Paratype 

USNM 159SS4

A B

13 10

20 22

Occurrence. Lower Devonian Rabbit Hill Limestone, Petes Canyon, locality Ml 150 at north end of 

Toquima Range, southern Lander County, Nev. At 

locality Ml 150 this colonial coral is associated with a 

typical Rabbit Hill fauna like that of the type locality. 

Colonial corals of this kind are uncommon in the Rabbit Hill Limestone, which usually contains only the 

small solitary rugose coral Syringaxon.

Family DISPHYLLIDAE Hill, 1939

Reference form. Disphyllum caespitosum (Goldfuss), 1826. Middle Devonian, Germany.

These phaceloid, cerioid, thamnastraeoid, and aphroid rugose corals have smooth or carinate, continuous 

lamellar, and usually little-dilated septa. The tabularium is wide, combining straight and slightly domed 

tabulae with marginal tabellae. Dissepiments in several columns are normal-globose or elongate; rarely 

lonsdaleioid. There are no horseshoe dissepiments. 

Species with a wide dissepimentarium have bases of 

outer dissepiments nearly horizontal.

This large complex Devonian family includes the 

following genera and subgenera:

Disphyllum de Fromentel, 1861

Cylindrophyllum Simpson, 1900

Acinophyllum McLaren, 1959

Hexagonaria Giirich, 1896

Hexagonaria (Pinyonastraea) new subgenus

Billingsastraea Grabau, 1917

Taimyrophyllwn Chernychev, 1941

Aphroidophyllum Lenz, 1961

Cerioid Hexagonaria-like genera are common in the 

Silurian and Devonian; most of these do not fit well 

in this family. Among these are the Australian Xystriphyllum Hill with wedge-thickened smooth septa and 

a narrow dissepimentarium and Columnarza-like species with few and irregularly distributed dissepiment 

columns of the German Devonian (Glinski, 1955). 

Certain other European Devonian species assigned to 

Hexagonaria appear to be affiliates of the Family Phillipsastraeidae (Rozkowska, 1960; Moenke, 1954).

The true spongophyllids and the Endophyllidae, 

which include Endophyllum and Australophyllum, are 

distinguished by a lonsdaleioid marginarium. However, this type of dissepiment arose independently in 

several rugosan families and may be expected to 

appear sporadically in Hexagonaria and other disphyllid genera.

The Family Phillipsastraeidae as here interpreted 

does not include the disphyllid corals (Hill, 1956, p. 

F279-F282; Stumm, 1964). Phillipsastraeidae are 

characterized by highly distinctive internal features

such as columns of true horseshoe dissepiments with 

contiguous peculiar trabecular bundles or sheaves. 

Moreover, in the Great Basin stratigraphic column the 

disphyllid forms are older, occurring in coral zones 

D2 and F; the true Phillipsastraeidae are of Late 

Devonian age, present only in coral zone I.

Genus DISPHYLLUM Fromentel, 1861

1826. Cyathophyllum caespitosum Goldfuss, p. 60, pi. XIX,

fig. 2b only; not pi. XIX, figs. 2a, 2c, 2d. 

1846. Cladocora goldfussi Geinitz (in part), p. 569. New name

for Cyathophyllum caespitosum Goldfuss, 1826, p. 60,

pi. XIX, figs. 2a-d; not pi. XIII, fig. 4. 

1935. Cyathophyllum caespitosum Goldfuss 1826 (p. 60, pi.

XIX, fig. 2b only). Lang and Smith, p. 545. 

1940. Disphyllum de Fromentel. Lang, Smith, and Thomas,

p. 53.

1945. Disphyllum Fromentel. Smith, p. 20-21 (in part only). 

1949. Disphyllum de Fromentel. Stumm, p. 32, pi. 15, figs.

1-5. 

1956. Disphyllum de Fromentel. Hill, p. F280, fig. 191-10a-b.

Type species. Cyathophyllum caespitosum Goldfuss, 1826, p. 60, pi. XIX, fig. 2b only (Lang and Smith, 

1935, p. 545; Smith, 1945, p. 20-21). Middle Devonian; Eifel district, Germany. Details of the complex 

taxonomic history relating to this type fixing are 

reviewed by Lang and Smith (1935, p. 545; Lang, 

Smith, and Thomas, 1940, p. 53; and Smith, 1945, p. 

20-21).

Diagnosis. These bushy and phaceloid rugose corals have fairly long slightly thickened major septa 

which do not reach the axis; minor septa are over half 

the length of the major septa. The tabularium is 

medium to wide; most tabulae are continuous and 

fairly straight, may be complete in some specimens 

and commonly are in combinations with periaxial tabellae. There may be a minor circular sag at the periphery of the tabularium, but not a pronounced uparching 

or axial sag. The dissepimentarium has several columns of small to medium dissepiments; the peripheral 

dissepiments are nearly flat, but the axial ones steepen 

to vertical in a border zone of small dissepiments. 

Large dissepiments are uncommon; there are no true 

lonsdaleioid dissepiments. Some septa may have zigzag 

carinae, but these are uncommon; there are no yardarm carinae.

Remarks. Disphyllum is here interpreted according to illustrations of the type species of Smith (1945, 

pi. 11, figs. 8a-c) and of Stumm (1949, pi. 15, figs. 

1-5). Acinophyllum McLaren (1959, p. 2228) has a 

narrower dissepimentarium and wider, straighter tabulae without marginal tabellae. Some other phaceloid 

rugose corals which might be confused with Disphyllum have marginal horseshoe dissepiments; these 

include Phacellophyllum Giirich and Peneckiella Soshkina, possibly a synonym of Phacellophyllum (see
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 59

McLaren, 1959, p. 22-23; p. 28-29). Cylindrophyllum 

Simpson has yardarm carinae; Eridophyllum has carinate septa and an aulos. Sociophyllum, which is the 

commoner coral of this general type in the Nevada 

Formation, has a wide peripheral zone of lonsdaleioid 

dissipiments and lacks carinae.

Disphyllum nevadense (Stumm), 1937 

Plate 18, figures 7-10

1937. Spongophyllum nevadense Stumm, p. 435, pi. 53, fig. 10,

pi. 55, figs. 5a, b. 

(?) Spongophyllum expansum Stumm, p. 436, pi. 53,

fig. 11, pi. 55, figs. 6a, b. 

1940. not Spongophyllum cf. expansum Stumm. Merriam, p.

56-57. 

not Disphyllum nevadense Stumm, p. 63.

Type material. Holotype, USNM 94454; lower 

part of the Nevada Formation, Lone Mountain, 

Eureka County, Nev.

Diagnosis. This slender Disphyllum has few dissepiment columns, including both globose and narrow 

elongate dissepiments; large, elongate dissepiments 

are interseptal and not lonsdaleioid.

Transverse section. Major septa, 16 to 18, extend 

to the axis in many corallites. Minor septa vary from 

short to more than half the length of the major septa. 

Septa, thin and wavy or straight, are more inclined 

to waviness in the tabularium; they are noncarinate. 

The wall is thin and lacks a stereozone.

Longitudinal section. The dissepiment columns 

range from one to three. The dissepiments are either 

small and globose or elongate. In places a nearly flat, 

elongate dissepiment extends from the periphery to 

the tabularium. Commonly there are one or two small 

nearly vertical dissepiments at the inner margin. The 

tabularium is wide; tabulae, mostly continuous or complete, are usually uparched distally with a periaxial 

sag; they are rarely straight.

Reproductive offsets. Lateral offsets are developed 

at the calice periphery to form branches.

Comparison with related forms. Disphyllum eurekaensis n. sp. has larger corallites with zigzag carinate 

septa and many more dissepiment columns lacking the 

elongate individual dissepiments. Disphyllum goldfussi 

(Geinitz) has more small dissepiment columns and 

lacks the elongate dissepiments of D. nevadense.

Remarks. Disphyllum nevadense Stumm, (1940) 

is a Late Devonian species of coral zone I and not 

closely related to the form here reassigned to Disphyllum.

Measurements. Diameter and major septa, three 

corallites of colony USNM 159320:

Corallites........................ 123

Diameter (in mm) ........ 8.3 7.3 7

Major septa ..................18 16 16

Occurrence. Nevada Formation, unit 2. Devonian 

coral zone D2 . Lone Mountain: Eureka County, Nev. 

locality M1046; at this locality it is associated with 

Disphyllum eurekaensis n. sp.

Disphyllum eurekaensis n. sp. 

Plate 18, figures 5, 6

Type material. Holotype, USNM 159319. Nevada 

Formation, unit 2; Devonian coral zone D2 . Lone 

Mountain locality M1046.

Diagnosis. This Disphyllum has corallites of large 

diameter and multiple columns of predominantly small 

dissepiments. The tabulae are mostly wide and fairly 

straight; some specimens are complete and some have 

peripheral tabellae and a tendency for peripheral 

depression of tabulae. The septa are slightly thickened 

in the dissepimentarium; some have weak zigzag 

carinae.

External features. Longitudinal grooves are not 

well shown; in places transverse rugae are fairly pronounced. There are lateral mergings of corallites in 

positions of marginal calicinal offsets.

Transverse sections. Major septa, which number 

18 to 22 do not reach the axis; minor septa are about 

two-thirds the length of the major septa. The septa 

are minutely wavy to zigzag and have sporadic zigzag 

carinae in the middle part of the dissepimentarium. 

Outer dissepiments are rather irregular; some are 

concave peripherally. All septa are continuous to the 

wall. The wall is thin and lacks a stereozone. Some 

are thickened slightly toward the inner edge of the 

dissepimentarium.

Longitudinal sections. Dissepiments, small and 

medium size, are in as many as eight columns on each 

side; outer three or four columns are medium, globose, 

and nearly flat to low dipping axially; inner four columns are small and steep to vertical. There are some 

large elongate dissepiments in the outer zone. The 

tabulae are rather closely set and for the most part 

fairly continuous and straight; some are complete, 

with large periaxial tabellae. Peripheral parts of the 

tabulae are depressed in some specimens. Traces of 

carinae appear in local patches.

Reproductive offsets. The merging of corallites 

laterally indicates reproductive offsets from calice 

margins.

Comparison with related forms. Disphyllum eurekaensis differs from D. goldfussi (Geinitz) in having 

more wavy and partly zigzag, weakly carinate septa; 

the tabularium of D. goldfussi (see Smith, 1945, pi. 11, 

figs. 8a-c; Stumm, 1949, pi. 15, figs. 2-3) is wider with 

more tabellae at the tabularium margin. Disphyllum 

nevadense (Stumm, 1937) has narrower corallites 

lacking the zigzag carinae, fewer dissepiment columns,
60 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

and partly elongate dissepiments. Disphyllum nevadense Stumm (1940, p. 63, pi. 7, fig. 10; pi. 8, figs, llab), a much younger species from the Devils Gate Limestone, is more slender, has fewer columns of dissepiments and a wide tabularium with medial flat tabellae, 

and lacks the more nearly complete straight tabulae 

of D. eurekaensis.

Measurements. Diameter and major septa; three 

corallites of colony USNM 159319:

Corallites .............................. 123

Diameter (in mm) ............19 16.6 17

Major septa ........................20 20 22

Occurrence. Nevada Formation, unit 2. Devonian 

coral zone D2 . Lone Mountain: locality M1046, associated with D. nevadense (Stumm, 1937).

Genus HEXAGONARIA Giirich, 1896 

1900. Prismatophyllum Simpson, p. 218.

Type species. Cyathophyllum hexagonum Goldfuss; by subsequent designation (Lang and others, 

1940, p. 69). Middle Devonian; Eifel district and Bensberg, Germany.

Diagnosis. The colonies are spherical and mushroom shaped; the polygonal corallites are separated by 

a discrete, usually unthickened wall. Calices have a 

flat or axially sloping peripheral platform and a deep 

axial pit. Major septa are unbroken from the wall into 

the tabularium and commonly reaching the axis; minor 

septa end at tabularium margin. Septa are normally 

thin, modified by waviness, minor zigzag elbow carinae 

and strong yardarm carinae. Dissepimentarium, usually wide, has multiple columns in which peripheral 

dissepiments lie nearly flat. There are no horseshoe 

dissepiment columns and no associated trabecular 

bundles.

Remarks. Most species of true Hexagonaria have 

long septa, some of which reach the axis. The septa 

commonly thin abruptly within the tabularium. There 

are, however, aberrant forms with all the septa shortened and an overly widened tabularium. The patterns 

of tabulae are usually irregular and include variable 

combinations of arched tabellae against inclined and 

incomplete tabulae; straight complete tabulae are unusual. Thickened walls and septa are uncommon, although there are species in which some individuals 

have moderate radially extended dilation or sporadic 

small bumpy or knobby swellings (Rozkowska, 1960, 

figs. 9, 11, 12).

Most species presently interpreted as Hexagonaria 

have carinae; usually yardarm carinae are in some but 

not all corallites or are restricted to parts of a single 

corallite. For example, Stumm's figures of the type 

species Hexagonaria hexagona Goldfuss from the Eifel 

district show definite and numerous yardarm carinae

(Stumm, 1948a, pi. VI, figs. 1, 2). However, Soraufs 

(1967, p. 10, fig. 4) figures of a specimen assigned to 

hexagona from Frasnes, Belgium, reveal no clear indication of yardarm carinae. It is assumed that the Eifel 

district material from the Middle Devonian is more 

nearly representative of the Goldfuss type species.

Unrelated Hexagonaria-Zz&e Rugosa. Any family of 

dissepimented rugose corals may theoretically give rise 

to compact cerioid species by adaptive radiation. 

Within the Phillipsastraeidae have evolved rather 

closely related solitary, bushy, and lastly cerioid forms 

which parallel Hexagonaria. Convergent or homeomorphic cerioid corals of this sort are known in Paleozoic 

strata from Silurian to Carboniferous. All share the 

compact growth habit, by itself not a feature of overriding taxonomic importance. Five of these families 

have cerioid radicles: (1) Kyphophyllidae, (2) Endophyllidae, (3) Spongophyllidae, (4) Disphyllidae, and 

(5) Phillipsastraeidae. Within a wide range of internal 

structure combinations, each family gave rise to lineages paralleling or mimicking Hexagonaria of the 

Disphyllidae.

Silurian rugose coral genera possessing the cerioid 

growth habit of Hexagonaria have received little attention from paleontologists. Undescribed corals from the 

Klamath Mountains Gazelle Formation possess a narrow, weakly developed tabularium and are classified 

with the Kyphophyllidae. Other western Silurian 

forms are endophyllids with strong lonsdaleioid dissepiments. Late Silurian Hexagonaria-like species of 

eastern North America are members of the cerioid 

genus Entelophylloides Rukhin, as are some Russian 

species.

Stratigraphic range of Hexagonaria. Hexagonaria 

has been reported in Devonian rocks of western Europe 

as old as Emsian and Siegenian (Sorauf, 1967, p. 24). 

In general those species which are convincingly disphyllid appear in the Eifelian rocks. In Europe the 

highest known Hexagonaria is Frasnian Late Devonian. The Hexagonaria from the Great Basin appears 

as the subgenus Pinyonastraea in the Nevada Formation, unit 2 and coral zone D2 , probably in earliest Eifelian. True Hexagonaria peaks in coral zone F, which is 

the higher Eifelian of this province. No coral zone I or 

Frasnian Hexagonaria is known with assurance in the 

Great Basin; all cerioid species of zone I thus far identified are Phillipsastraeidae.

Hexagonaria systematics. Characters by which 

Hexagonaria may be subdivided taxonomically and 

distinguished from Phillipsastraeidae and other families have been reviewed by Stumm (1948a), Schouppe 

(1958), and more recently by Sorauf (1967). At present Hexagonaria is a large morphologically diverse and
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 61

complex genus of worldwide distribution in need of 

descriptive study and taxonomic appraisal. Some 

morphologically similar taxa within its scope are 

doubtless products of convergent evolution. As with 

Billingsastraea and other disphyllids, it is not unlikely 

that similar internal structures developed in not very 

closely related genetic stocks; in this light Hexagonaria may be quite polyphyletic.

Current investigations of Hexagonaria differentiation in Great Basin Devonian coral zones D and F suggest that subgenera may be used appropriately. Accordingly the earliest Hexagonaria of coral zone D is 

discribed as subgenus Pinyonastraea.

Of uncertain relationship to Hexagonaria are three 

noncarinate Devonian groups of Hexagonaria-like corals: (1) the genus Xystriphyllum; (2) a group having 

a few small sporadic lonsdaleioid dissepiments; and 

(3) aberrant species structurally intermediate between Hexagonaria and Columaria.

Xystriphyllum, unlike Hexagonaria, has a thickened 

wall, thickened gradually wedge-tapering smooth 

septa, narrow tabulae, and very steeply inclined or 

vertical dissepiments. Group 2, with its scattered small 

lonsdaleioid features, may conceivably be a Hexagonaria derivative, for dissepiments of this kind clearly 

arose in numerous genetically unrelated lineages. Australophyllum of the Endophyllidae differs in possessing 

a well-defined lonsdaleioid band.

Group 3, a heterogeneous, probably polyphyletic 

form group, from the Devonian of Western Europe, 

comprises thick-walled species with a very wide tabulae, a reduced dissepimentarium having few columns, 

and thick wedge-tapering, commonly short septa 

(Glinski, 1955). Generic or subgeneric grouping may 

eventually prove appropriate for these local differentiates.

Certain Polish species of "Hexagonaria" described 

by Moenke (1954) appear, on examination of the published figures, to be better assigned to the Phillipsastraeidae. The origin of this family is unexplained. 

Whether the distinctive horseshoe dissepiment columns and trabecular bundles arose in a Middle Devonian disphyllid line can only be inferred at present. 

In the Central Great Basin, beds of coral zone F with 

Hexagonaria also contain disphyllids of the Taimyrophyllum type surficially mimicking Late Devonian 

(coral zone I) Pachyphyllum of the Phillipsastraeidae. 

These mimics lack the horseshoe dissepiment columns 

and trabecular bundles.

Study of Hexagonaria suites from coral zone F, supplemented by review of the literature on Disphyllidae, 

suggests that the following characters are of special 

taxonomic value within this and allied genera.

1. Septal configuration: thin, smooth and 

straight; or thickened, tapering gradually 

from wall; or wavy, zigzag with elbow carinae, yardarm carinae, or combination of 

these.

2. Wall in transverse section: thin and wavy or 

straight; or thickened stereoplasmically 

with obtusely wedge-tapering septum bases.

3. Width of tabularium relative to dissepimentarium.

4. Outer dissepiments in longitudinal section: 

vertical to steeply inclined toward axis; or 

nearly horizontal.

5. Outer dissepiments in transverse section: normal concentric, or irregular with herringbone pattern.

The width of the tabularium is a feature having a good 

deal of latitude within a species and between species. 

Normally about one-third the corallite diameter in 

most species, the tabularium may broaden to exceed 

one-half the corallite diameter, and the dissepiment 

band narrows correspondingly as the septa shorten. 

At the opposite extreme is Hexagonaria with narrow 

tabularium and thin axial septum extensions which 

meet in a twisted axial plexus accompanied by median 

uparching of tabulae. A small raised median calice boss 

may result, but no discrete axial rod or columella.

Subgenus PINYONASTRAEA, new subgenus 

1937. Prismatophyllum kirki Stuirtm, p. 437; pi. 55, figs. 7a-b. 

1940. "Prismatophyllum" kirki Stumm. Merriam, pi. 16, fig. 5.

Type species. Prismatophyllum kirki Stumm, here 

designated. Nevada Formation, unit 2, Devonian coral 

zone D2 ; Lone Mountain.

Diagnosis. This Hexagonaria has a thin wall and 

thin minutely wavy septa lacking yardarm carinae. 

The dissepimentarium is wide and has numerous columns of globose dissepiments; the peripheral dissepiments are nearly flat. The calice is shallow with a broad 

brim platform and an axial pit.

Remarks. The subgenus Pinyonastraea differs 

from Hexagonaria (Hexagonaria) as typified by its 

type species hexagona Goldfuss (Stumm, 1948a, p. 11- 

15, pi. VI, flgs. 1-2) in lacking crossbar or yardarm 

carinae and other septal thickenings. Carinae, if developed in Pinyonastraea, are weak and of the zigzag 

elbow or angulation type.

Stumm, in 1937 (p. 437), called attention to this 

coral type as possibly having developed from ancestry 

different from that of other "Prismatophyllum" If it 

has, study of more adequate material might justify 

separate generic designation eventually.

Selected individuals of Pinyonastraea kirki resemble 

internally the associated Billingsastraea nevadensis,
62 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

especially subspecies arachne; the principal difference 

is lack of the outer wall. As elsewhere noted, it is conceivable the two come from common ancestry.

Pinyonastraea is a relatively uncommon coral in the 

higher part of Nevada Formation, unit 2, unlike the 

local occurrence of Hexagonaria s.s. which is abundant 

in higher Middle Devonian coral zone F. The new subgenus has not been recognized outside of the Great 

Basin Province.

Hexagonaria (Pinyonastraea) kirki (Stumm) 

Plate 23, figures 5-10

1937. Prismatophyllum kirki Stumm, p. 437, pi. 55, figs. 7a-b. 

1940. "Prismatophyllum" kirki Stumm. Merriam, pi. 16, fig. 5. 

1949. Hexagonaria kirki (Stumm), pi. 15, figs. 16,17.

Type material. Holotype USNM 94456; lower part 

of the Nevada Formation, Lone Mountain, Eureka 

County, Nev.

Diagnosis. This Pinyonastraea has large corallites, 

minutely undulant, very weakly zigzag, thin carinate 

septa and thin, even, commonly straight walls. Some 

of the outer dissepiments are sublonsdaleioid in the 

peripheral irregular zone of the dissepimentarium.

External features. This species occurs in large lenticular colonies 25 cm or more in greatest diameter. Individual corallites are considerably larger than average 

for Hexagonaria. A well-preserved distal surface (pi. 

23, fig. 7) shows shallow calicinal pits with a weak 

crateriform rim; the calicinal platform is concavely 

depressed to almost flat, and the thin outer wall stands 

sharply in relief.

Transverse sections. Major septa number 20-23; 

these are uniformly thin, and for the greater part extend without discontinuity to the axial vicinity. Minor 

septa are more than half to nearly as long as the major 

septa, which extend to the tabularium margin. Some 

minor septa are discontinuous, represented by crests, 

or lacking; minor septa commonly do not pass peripherally to the wall. Septa range from straight and nearly 

smooth to minutely undulant and zigzag carinate. 

However, the carinae are not abundant; they are 

minute, and in some specimens confined to a few septa, 

in which they occur alternately in radial succession. 

There are no yardarm carinae. The peripheral zone of 

irregular dissepiments is wide; the outer sublonsdaleioid dissepiments, if present, usually are small.

Longitudinal sections. The flat-lying dissepiments 

which make up most of the dissepimentarium show 

little size regularity; they range from small to large, 

the larger predominate. Significant features are nearly 

straight, continuous, horizontal, and somewhat thickened tabulae in parts of this zone.

Comparison with related forms. In the Roberts 

Mountains at locality M1054, a subcerioid variety with

corallite walls partly rounded transversely is associated with normal kirki; this fact suggests a phaceloid 

tendency (Merriam, 1940, pi. 16, fig. 6). An Hexagonaria-like species occurring in the Toiyabe Range at 

locality Ml 151 near Reeds Canyon (plate 25, fig. 10) 

has wide corallites and may be allied either to kirki 

of coral zone D or to Hexagonaria flexum (Stumm) of 

coral zone F.

Hexagonaria (Pinyonastraea) kirki has no described 

close relatives elsewhere. It is possible this form is 

derived from the stock which produced the corals 

assigned to Billingsastraea nevadensis.

Measurements. Holotype USNM 94456; diameter 

of a mature corallite, 17 mm; major septa, 20. Figured 

specimen USNM 159345: diameter of a mature corallite, 17 mm; major septa, 23.

Occurrence. Lower part of the Nevada Formation, 

unit 2. Devonian coral zone D2 . Lone Mountain: locality M1052. Northern Roberts Mountains: locality 

M1054. Northern Antelope Range: locality M1053. At 

Lone Mountain this coral may range into coral zone D3 

at locality M1052.

Genus BILLINGSASTRAEA Grabau, 1917

1917. Phillipsastraea (Billingsastraea) Grabau, p. 957. 

1937. Billingsastraea Grabau. Stumm, p. 437, pi. 53, fig.

13, pi. 55, figs. 8a-b. 

1949. Billingsastraea Grabau. Stumm, p. 35, pi. 16, figs.

8-11, ?fig. 7. 

1951. Billingsastraea Grabau. Ehlers and Stumm (in

part).

1953. Billingsastraea Grabau. Ehlers and Stumm, p. 1. 

1956. Billingsastraea Grabau. Hill, p. F280, fig. 191-7. 

Not 1958. Billingsastraea Grabau. Schouppe, p. 235-237. 

1964. (?) Billingsastraea Grabau. Stumm, p. 43, pi. 40,

figs. 1-3.

1964. Billingsastraea Grabau. Oliver, p. B1-B5, pis. 1-2. 

1964. Radiastraea Stumm. Pedder, p. 446-449, pis. 71-73. 

Type species. Phillipsastraea verneuili Edwards 

and Haime, 1851 (by monotypy). According to Edwards and Haime, the type occurrence is in the State 

of Wisconsin; the type is now believed to have come 

from glacial drift.

Diagnosis. Oliver's (1964) diagnosis is as follows: 

Astraeoid, thamnastraeoid, or slightly aphroid corals with calices having a central pit and a broad horizontal or reflexed 

peripheral platform. Septa are radially arranged and lightly 

to heavily carinate with zigzag or crossbar carinae. Major and 

minor septa extend from the periphery; minor septa terminate 

at the outer margin of the tabularium; major septa commonly 

extend to or almost to the axis. At their peripheral ends the 

septa abut against those in the adjacent corallite or are continuous into the next corallite. The tabularium is narrow and 

composed of closely spaced, horizontal complete tabulae. The 

dissepimentarium is composed of gently to strongly curved 

dissepiments that are horizontally arranged except next to the 

tabularia where they are inclined toward the corallite axes.

Remarks. Grabau (1917, p. 957) introduced the
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 63

name Billingsastraea in a subgeneric sense without 

definitive treatment. Stumm first defined Billingsastraea as a genus in 1937. Redefining the genus in 1949, 

Stumm (p. 35) in effect interpreted his genus Radiastraea (Stumm, 1937, p. 439) as an objective synonym 

of Billingsastraea, thus suppressing Radiastraea. As 

noted by Oliver, the species Radiastraea arachne 

Stumm (type species of Radiastraea) was regarded by 

Stumm (1949, p. 35) as a weakly carinate species of 

Billingsastraea. Oliver's diagnosis above supplements 

that of Stumm in taking into account the characters of 

R. arachne.

Because the internal features of "Phillipsastraea" 

verneuili Edwards and Haime, type species of Billingsastraea, are unknown, the status of this genus remains 

clouded. Pedder (1964, p. 447) regards certain species 

assigned to Billingsastraea by Ehlers and Stumm 

(1951; 1953) as constituting a separate but undescribed compound Heliophyllum-like genus; these are 

doubtless the strongly yardarm-carinate forms. He has 

accordingly resurrected Radiastraea for the noncarinate to not so strongly carinate species of the arachne 

type.

At present too little is known of the taxonomic significance of prominent crossbar or yardarm carinae to 

decide whether or not these features constitute a sufficient basis for separating the species bearing them as 

a genus distinct from the species presently in Billingsastraea which have noncarinate septa or zigzag carinae. The problem is similar to that in connection with 

Hexagonaria.

In accord with Stumm's 1949 review and the reevaluation of Oliver (1964), the term Billingsastraea is 

provisionally retained for the Nevada forms here dealt 

with, in the hope that eventually the type of verneuili 

may be sectioned and studied.

Billingsastraea neuadensis (Stumm) andB. arachne 

(Stumm) were both described from the lower part of 

the Nevada Formation at Lone Mountain, Eureka 

County, Nev. Both occur in the lower part of the Nevada (in unit 2) and in coral zone D2 ; it is not improbable they are within the range of variation of a 

single species. However, full confirmation of this identity will depend upon variation studies of more complete suites of specimens from the same locality and 

horizon than are at present available. Probably the 

best course at this time is to consider B. arachne 

merely a subspecies of B. nevadensis.

Billingsastraea nevadensis (Stumm) 

Plate 24, figures 1-3, 5, 6-8

1937. Billingsastraea billingsi nevadensis Stumm, p. 438, pi.

53, fig. 12, pi. 55, figs. 9a-b. 

1940. Billingsastraea nevadensis Stumm. Merriam, pi. 12, fig.

1.

1949. Billingsastraea nevadensis Stumm, pi. 16, fig. 8.

Type and figured material. Holotype USNM 

94467; lower part of Nevada Formation, Lone Mountain. Figured specimens: USNM 159346, 159347, 

localities M1055, M1035, Antelope Range; USNM 

159348-159350, locality M55, Lone Mountain. Devonian coral zone D2 .

Diagnosis. This thamnastraeoid Billingsastraea 

has uniformly thin finely wavy or zigzag septa and 

numerous weak zigzag elbow carinae. Longer major 

septa reach the axis; their waviness generally increases 

as they approach the center. Minor septa are twothirds the length of the major septa; their axial tips 

just enter the tabularium. Concentric dissepiment 

traces, close-set at the inner margin of dissepimentarium, form a false inner ring that normally has no 

significant thickening. Tabulae are mostly irregular, 

arched distally, and some of them are complete. Flat 

outer dissepiments range in size from small to large; 

inner steeply inclined dissepiments are small.

External features. Ovoidal colonies that are 15 cm 

or more in greatest diameter show polygonal outlines 

resembling those of large corallites that have raised 

margins with the external appearance of a wall. Calice 

pits commonly lack the raised crateriform margin 

characteristic of subspecies arachne.

Transverse sections. Major septa average about 

16, most of them extend to the axis; in the tabularium, 

major septa, wavy and irregular, commonly produce an 

incipient plexus, join laterally or with their opposite 

septa. Some individuals reveal a moderate thickening 

of both major and minor septa only in those parts 

where they pass from dissepimentarium to tabularium; 

others show no such thickening. No wall separates 

corallites, most septa pass into septa of adjacent 

corallites peripherally. Dissepiment traces are usually 

evenly concentric in the close-spaced periaxial area; 

they become more irregular or nearly straight in the 

more open peripheral area. Septal waviness varies considerably from those with numerous small, even bends 

without carinae to those with closely spaced, zigzag 

bends having fine sharply projecting carinae at the 

elbow angles. No crossbar (yardarm) carinae are present. Astraeoid patterns are slightly developed peripherally, but no aphroid tendencies have been recognized 

in this predominantly thamnastraeoid species.

Longitudinal sections. The diameter of the tabularium is about one-third of or slightly less than the 

corallite diameter. In a broad zone of nearly flat lying 

dissepiments, small to large dissepiments have irregular distribution; limbs of larger dissepiments subtend 

as many as five small underlying ones. The innermost 

column of small dissepiments stands near vertical. The
64 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

tabulae, rather closely spaced, commonly are arched 

distally in an irregular fashion and have a periaxial 

depression. A few tabulae nearly complete, most terminate as large tabellae. Carinal traces show in scattered patches. Horseshoe dissepiments are lacking.

Comparison with related forms. Billingsastraea 

nevadensis differs from B. verrilli (Meek) in the structure of its septa, which are thickened within the dissepimentarium of verrilli (Smith, 1945, pi. 19, figs. 

2b-c; and in cellulose peels of a Meek para type); B. 

nevadensis has uniformly thin septa in this region. B. 

verrilli, figured by Pedder (1964, pi. 71, figs. 1-5; pi. 

72, figs. 4-5), is less thickened here than is Meek's 

paratype specimen, but it has thicker than normal 

nevadensis septa in the dissepimentarium.

Among the adequately illustrated species of Billingsastraea, the one under consideration bears a significant resemblance to B. affinis (Billings) as figured by 

Oliyer (1964). The differences are a greater number of 

major septa (about 19 in affinis, 16 in nevadensis) and 

possibly in the vertical sides of calicular pits in affinis. 

Billingsastraea affinis, reported by Oliver (1964, p. B3) 

from the Grande Greve Formation of Coblenzian age, 

is possibly not a great deal older than B. nevadensis 

from the beds of coral zone Dz .

Measurements. The measurements of representative corallites of figured specimens are:

USNM No. .............................................. ...159346 159347 159349

Diameter (in mm)........................16 14 11

Major septa ..................................19 17 16

Occurrence. Nevada Formation, unit 2; Devonian 

coral zone D2 . Lone Mountain: localities 104 (Merriam, 1940), M55. Northern Roberts Mountains: locality 3 (Merriam, 1940). Northern Antelope Range: localities M1035, M1055. At localities 3 and 104 (Merriam, 1940) B. nevadensis is associated with subspecies 

arachne.

Billingsastraea nevadensis subsp. arachne (Stumm) 

Plate 24, figure 4

1937. Radiastraea arachne Stumm, p. 439, pi. 53, fig. 13; pi.

55, figs. 8a-b.

1940. Radiastraea arachne Stumm. Merriam, pi. 13, fig. 5. 

1949. Billingsastraea arachne (Stumm), p. 35, pi. 16, fig. 9,

10, 11.

1964. Radiastraea arachne Stumm. Oliver, p. B3. 

1964. Radiastraea arachne Stumm. Pedder, p. 446-447, pi.

72, figs. 1-3, pi. 73, figs. 1-5.

Type material. Holotype USNM 94458. Lower 

part of the Nevada Formation, Lone Mountain.

Diagnosis. This Billingsastraea has uniformly thin, 

moderately wavy to only slightly wavy or nearly 

straight septa, with a few zigzag carinae limited to a 

few septa. The distal surface of type has prominent 

raised crateriform rim at the edge of calicular pits.

Remarks. Stumm (1937, p. 439) defined Radiastraea to contain species having these features, together 

with an inner aulos or "tubular ring." Later Stumm 

(1949, p. 35) noticed that an aulos was not present; 

accordingly he concluded that R. arachne is a weakly 

carinate species of Billingsastraea. This appears to be a 

reasonable interpretation. Pedder (1964, p. 446-449) 

has reinterpreted Stumm's Radiastraea as a genus and 

has proposed its adoption to include arachne and verrilli (Meek).

Occurrence. Lower part of the Nevada Formation, 

unit 2. Devonian coral zone D2 . Lone Mountain: localities 58, 104, 105 (Merriam, 1940). Northern Roberts 

Mountains: localities 3, 36 (Merriam, 1940), M1073. 

Southern Roberts Mountains: locality 15 (Merriam, 

1940). At localities 3 and 104 this subspecies occurs 

with typical Billingsastraea nevadensis.

Billingsastraea? sp. T 

Plate 25, figures 6-9

This Billingsastraea? has thamnastraeoid to aphroid 

growth habit. Its thickened septa lack carinae but are 

minutely wavy. Septal dilation is rather evenly distributed radially through the dissepimentarium; there 

is a greater swelling of twisted axial segments in the 

tabularium. Closely spaced tabulae, mostly incomplete 

and irregular, have a slight proximal sag. In longitudinal section this species resembles Billingsastraea 

nevadensis (Stumm).

Billingsastraea? sp. T differs from B. nevadensis in 

lacking septal waviness and angulation carinae and in 

having thickened septa. Moreover, nevadensis normally is not aphroid.

The partly aphroid growth of this form is suggestive 

of Billingsastraea-like corals with dilated septa occurring in Nevada Formation, unit 4 (coral zone F) at 

Lone Mountain. These undescribed coral zone F forms 

are more appropriately classified as Aphroidophyllum 

Lenz (1961, p. 505) or Taimyrophyllum Chernychev 

1941, the relationships of which have recently been 

discussed by Pedder (1964).

Occurrence. Toiyabe Range, Nev.; Austin quadrangle, locality M1151, west range front 0.8 mile southsouthwest of mouth of Reeds Canyon. Lower part of 

Devonian limestones correlative with the Nevada Formation. Middle Devonian within the interval of coral 

zones D to F.

Family CYSTIPHYLLOIDAE Slumm, 1949

Reference form. Cystiphylloides aggregatum (Billings) 1859. Devonian; Ontario, Canada.

These compound and solitary Devonian cystimorphs 

lack septa beyond initial growth stages, except for 

traces or weak radiating septal ridges on mature calice
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 65

floor and wall. The transverse thin sections of some 

species do not show septal crests. The calice is deep 

and has a V-shaped profile. The tabularium is generally narrow, in keeping with the deep, conical calice. 

The dissepimentarium is not abruptly set off from the 

tabularium. Straight tabulae are uncommon; the horizontal structures consist of two or more tabellae with 

bases nearly horizontal or axially inclined at a low 

angle. All the dissepiments are concave peripherally 

in lonsdaleioid fashion.

The Devonian Cystiphylloidae differ from the Silurian Cystiphyllidae in lacking the trabecular septal 

spines that extend inward from dissepiment surfaces. 

Transitional forms probably exist between the Cystiphylloidae and the Digonophyllidae, which generally 

have septal crests and ridges.

Genus CYSTIPHYLLOIDES Chapman, 1893

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

1893. Cystiphylloides Chapman, p. 46.

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

1940. not Cystiphylloides Yoh. Lang, Smith, and Thomas, p.

48. 

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).

Type species. Cystiphyllum aggregation Billings, 

1859. Devonian; Onandaga, near Simcoe, Ontario, 

Canada. Cystiphylloides Yoh, 1937 is a homonym and 

is probably not congeneric with Cystiphylloides Chapman, although it has generally been so regarded 

(Stumm 1949, p. 39). Unfortunately the internal 

structure of Billings' type of Cystiphyllum aggregatum 

is unknown, and the specimen is lost. According to 

D. J. McLaren (written commun., Sept. 1, 1966), Billings' species is almost certainly the common Onondaga dendroid to phaceloid form, as suggested by Billings' figure of the exterior. Sections of this species are 

available; the diagnosis is based on them.

Diagnosis. These are medium and large dendroid 

to phaceloid and solitary cystiphylloid Rugosa; individual corallites range from trochoid to cylindrical. 

The mature dissepimentarium, of medium width, is 

composed mostly of small- and medium-sized, rather 

uniformly globose dissepiments. The border between 

the tabularium and the dissepimentarium is rather 

well defined but not sharp; there is a change from 

small steep dissepiments to large axially inclined tabellae. The tabularium is medium wide to wide. Septa 

and septal spines or crests are commonly absent.

Remarks. A series of transverse sections of a dendroid, a moderately compact colony from the Onondaga Limestone of Hagersville, Ontario (Royal Ontario Museum Collections No. 8950H), (photographs

of which were supplied by McLaren), show corallites 

in neanic and mature growth stages. None have septal 

crests, even at the periphery where they appear in 

some cystiphylloids. Most of the dissepiments decrease 

in size axially; they become less globose in the columns 

bordering the wide tabularium where there is a rather 

abrupt change to large tabellae.

Yoh's Cystiphylloides (1937), from the Middle Devonian of Kwangsi Province, South China, was proposed without reference to prior use of this generic 

name by Chapman. Yoh's excellent figures suggest 

that it is a solitary coral with a wider dissepimentarium and a narrower tabularium which is less abruptly 

set off from the former. Septal crests are present peripherally, a feature not noted in the Onondaga form.

Cystiphylloides, according to either Chapman's or 

Yoh's definition, rather conclusively does not belong 

in synonymy with Plasmophyllum where it has recently been placed by Birenheide (1964, p. 17).

The cystiphylloid corals from the Nevada Formation may be segregated as colonial or solitary and, on 

the basis of septal crests, into two groups; one group 

lacks septal crests entirely or has some scattered but 

poorly developed, and the other group has well-developed peripheral septal crests. Within this morphological range there is a great diversity of type with respect 

to features of the dissepimentarium and tabularium. 

The term Cystiphylloides of Chapman is applicable to 

some of these types, especially to the colonial forms. 

Other types, morphologically closer to Yoh's Cystiphylloides may require a new generic name. Generic 

names such as Mesophyllum and Digonophyllum are 

available for these Cystiphylloides with false septa or 

septal crests alined radially in the separate growth 

cones.

Cystiphylloides robertsense (Stumm) 

Plate 23, figures 1-4

1937. Mesophyllum robertsense Stumm, p. 440, pi. 53, fig. 14; 

pi. 55, figs. lOa-c.

Type material. Holotype USNM 94459, Frazier 

Creek, northern Roberts Mountains; paratype USNM 

94460, Lone Mountain, Eureka County, Nev. Lower 

part of the Nevada Formation.

Diagnosis. The mature corallites are cylindrical 

and narrow for the genus; probably colonial. A definite 

peripheral zone of small dissepiments passes rather 

abruptly into an inner band of large dissepiments that 

are not sharply set off from the tabularium.

External features. The cylindrical corallites have 

rather coarse external longitudinal ribs. The parallel 

alinement of corallites in the matrix. A bushy colonial 

growth form, although no lateral connection was 

observed.
66 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Transverse section. The peripheral zone shows 

about four columns of small dissepiments; at the axis 

there is a fairly abrupt change to large dissepiments. 

Within the outer band of small dissepiments there is 

stereoplasmic thickening but septal crests, if present, 

are indistinct. No septal crests were observed internally

Longitudinal section. The change from small dissepiments to large ones in the peripheral zone is conspicuous, but the separation between the tabularium 

and the large dissepiments is indistinct; the innermost 

roughly horizontal arched features considered to be 

tabellae occupy the axial part of the corallite that is 

usually narrow.

Reproductive offsets. Stumm (1937, p. 440) notes 

the presence of a calice offset in the holotype.

Comparison with related forms. Other Great Basin 

cystyphylloids become cylindrical when mature, but 

this specimen is the only one I have reason to think 

may be a phaceloid colonial.

Measurements. Figured specimen USNM 159343: 

mature corallite diameter 22 mm; width of peripheral 

band of small dissepiments 3 mm.

Occurrence. Lower part of Nevada Formation, 

lower part of unit 2. Devonian coral zone D. Lone 

Mountain: locality M1046. Northern Roberts Mountains, Nev.; type locality of species on Frazier Creek. 

At Lone Mountain locality M1046 this coral is associated with Disphyllum nevadense (Stumm) and D. 

eurekaensis n. sp.

Cystiphylloides lonense (Stumm) 

Plate 19, figures 1-9

1937. Mesophyllum lonense Stumm, p. 440, pi. 53, fig. 15; pi. 

55, figs, lla-b.

Type material and figured specimens. Holotype 

USNM 94461, paratype USNM 94461a. Figured specimens USNM 159322-159327: lower part of the Nevada Formation, Lone Mountain, Eureka County, 

Nev.

Diagnosis. This medium-sized, solitary, ceratoid 

Cystiphylloides has large tabellae not sharply set off 

from large interior dissepiments; few recognizable septal crests are present in transverse sections, but the 

deep calice has weak, raised, discontinuous radial striations. Both large and small dissepiments are in the 

peripheral band.

Transverse sections. About two-thirds the diameter of the mature stages consists of larger dissepiments 

and tabellae; a few large dissepiments are at the periphery. There are a few scattered septal crests in the 

peripheral band, and incomplete cones of stereoplasmic thickening occur at about the inner edge of the 

dissepimentarium.

Longitudinal sections. Most dissepiments, including those in the peripheral band, are steeply inclined. 

The tabularium is commonly occupied by two large 

tabellae, which are either nearly flat or axially inclined 

at a low angle; in the early neanic stages there may be 

a few complete and flat tabulae.

Comparison with related forms. The younger Cystiphylloides robertsense of coral zone D2 has a greater 

concentration of small dissepiments in the outer band 

of the dissepimentarium and is cylindrical and probably colonial when mature. Cystiphylloides americanum (Edwards and Haime) has a much wider dissepimentarium with numerous columns, the outer columns 

are nearly flat. Its tabularium usually shows several 

laterally contiguous tabellae in a rather wide and fairly 

discrete tabularium.

Measurements. Three figured specimens (in mm):

USNM No. ........................................................159SU 159S2S 159S2^

Length of corallum restored............39 69 70

Diameter at calice edge....................22.5 31 31

Outside diameter at

calice bottom ................................17 17 23

Depth of calice..................................l6 35 29

Occurrence. Lower part of the Nevada Formation, 

unit 1. Devonian coral zone C. In association with Papiliophyllum elegantulum, Siphonophrentis (Breviphrentis) kobehensis, and Acrospirifer kobehana. Lone 

Mountain: localities M74, M286.

Cystiphylloides sp. d 

Plate 19, figures 10, 11

1884. (?) Cystiphyllum americanum Milne-Edwards. Walcott, 

p. 106.

1937. (?) Mesophyllum flexum Stumm, p. 441, pi. 55, figs. 

12a-b.

1938. (?) Mesophyllum vesiculosum (Goldfuss). Stumm, p. 

483, pi. 59, figs. 8a-b.

These cystiphylloid rugose corals resemble Cystiphylloides americanum (Edwards and Haime). They 

occur in the lower part of the Nevada Formation, probably ranging up through coral zone D to coral zone E. 

In coral zone D these corals are represented by the 

form here referred to as Cystiphylloides sp. d. There is 

insufficient material to justify a specific name at 

present.

Like C. americanum, this form grows to a large size 

with numerous columns of medium and large dissepiments that are not sharply set off from from tabellae 

in the axial vicinity. In C. sp. d the dissepiments are inclined steeply and many are elongate, whereas in C. 

americanum the peripheral dissepiments are less 

steeply inclined. The calice of C. americanum is deep 

and bellshaped, whereas the calice of C. sp. d may be 

very deep and conical or funnel shaped. No septal 

crests are noted.
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 67

Occurrence. Nevada Formation, unit 2. Devonian 

coral zone D2 . Southern Sulphur Spring Range: locality M36.

Family DIGONOPHYLLIDAE Wedekind, 1924

Reference form. Mesophyllum defectum Schluter 

1889. Middle Devonian, Germany.

Large, solitary, pervasively dissepimented rugose 

corals have septa which tend to be discontinuous 

lamellar plates, both vertically and radially. Bilateral 

symmetry is obscure in mature growth stages; at the 

mature stage the fossula is usually not clearly defined.

The Digonophyllidae include forms ranging from a 

type with conspicuous septa to a type with greatly 

reduced mature septa, which may appropriately be 

called cystimorphs. The dissepimentarium is always 

wide; the tabularium is narrow and may be poorly differentiated. Many of these corals have a wide calice 

brim or platform.

As noted by Wedekind (1924), the septal tissue of 

the Digonophyllidae develops discontinuously in successive growth cones (septal cones of Wedekind).

The family and generic classification of these corals 

remains in a confused state. Multiplication of genera 

by Wedekind and associates reflects the taxonomic 

philosophy of the extreme "splitter," although the 

coral divisions appear to have been related to stratigraphic units. Birenheide (1964), in a painstaking and 

detailed revision, eliminates many of the Wedekind 

genera. Hill (1956, p. F314-F320) has adopted some 

Wedekind genera as subgenera in a practical and reasonably satisfactory scheme based on current understanding of this complex group.

For purposes of this study two subfamilies of the 

Digonophyllidae are adopted: Digonophyllinae and 

Zonophyllinae. In the Great Basin Province the Digonophyllidae appear in Devonian coral zone D2 and are 

not known above coral zone F. Here, as in Germany, 

they are characteristic of Middle Devonian, especially 

the Eifelian.

Subfamily DIGONOPHYLLINAE Wedekind, 1924

Reference form. Mesophyllum defectum Schliiter 

1889. Middle Devonian, Germany.

The large Digonophyllidae have septa uniformly developed or sporadic and vestigal in growth cones. The 

septa in the growth cones are either radially continuous plates or radially alined ridges and crests. Strip 

carinae are present in the marginarium of some genera. 

Normal but irregular dissepiments of the periaxial 

band change peripherally to irregularly lonsdaleioid. 

The dissepiment pattern is highly complex and nonuniform in the marginarium of species having peripherally suppressed septa.

In all Digonophyllinae the tabularium is relatively 

narrow and poorly differentiated and lacks straight 

tabulae. The calice is deep and conical, floored by two 

or more tabellae; in those forms having a wide brimplatform, the inner conical part of the calice is a 

steep-sided pit.

Hill (1956, p. F317) and Birenheide (1964, p. 6) 

note the presence of a fossula in some Digonophyllidae; however, no fossula is recognized in the Great 

Basin species, and it seems probable that symmetry 

features of this kind are not generally conspicuous in 

the Digonophyllinae.

No preparations of Great Basin material convincingly show vertical continuity of homologous septal 

lamellae from one incremental growth cone to those 

above and below, within successive nested cone 

sequences.

The radially broken septa of some species commonly 

have such closely spaced component crests that the 

septa almost look continuous. Observed in transverse 

thin section, each crest or spine is an outgrowth from 

one or more tabellae or from one or more of the dissepiments.

As noted elsewhere, it is probable that Digonophyllinae with almost wholly lonsdaleioid dissepiments intergrade with the Cystiphylloidae.

Genera and subgenera classified with the Digonophyllinae are as follows:

Digonophyllum (Digonophyllum) Wedekind, 1923 

Digonophyllum (Mochlophyllum) Wedekind, 1923 

Mesophyllum (Mesophyllum) Schliiter, 1889 

Mesophyllum (Arcophyllum) Markov, 1926

Arcophyllum is classified as a subgenus under Mesophyllum by Hill (1956, p. F318). Other possible subgenera of Mesophyllum are: Lekanophyllum Wedekind, 1923, Atelophyllum Wedekind, 1925, and Dialytophyllum Amanshauser, 1925.

The Digonophyllinae appear in Early and early Middle Devonian coral subzones D2 and D3 of the Great 

Basin with the genera Mesophyllum and Arcophyllum. 

Corals of this subfamily peak in the higher Eifelian 

interval of coral zone F.

Genus MESOPHYLLUM Schluter, 1889 

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

Type species. Mesophyllum defectum Schluter, 

1889. Middle Devonian; Eifel district, Germany.

Diagnosis. These Digonophyllinae have nonuniformly discontinuous septa which become obsolete 

peripherally in a wide band of irregularly lonsdaleioid 

dissepiments, with or without localized groups of strip 

carinae.

Remarks. The corals assigned to Mesophyllum 

range in structure from maturely cylindrical cysti-
68 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

morphs with only a few weak septal traces and scattered or no bar carinae to those with numerous discontinuous and fairly strong continuous irregularly 

thickened septa and abundant strip carinae.

At present the adoption of Mesophyllum as a genus 

with subgenera, more or less in accord with the scheme 

used by Hill (1956, p. F317-F320) seems to be a reasonably satisfactory rationalization of the taxonomic 

dilemma in this difficult and complex group. Birenheide's detailed systematics (1964) are, on the other 

hand, quite different; he classifies Mesophyllum as a 

subgenus of Plasmophyllum.

Of the seven subgenera of Mesophyllum listed by 

Hill, only two, Mesophyllum (Mesophyllum) and Mesophyllum (Arcophyllum), are pertinent in this study. 

Cystiphylloides Chapman is not here classified as a 

subgenus of Mesophyllum.

Subgenus MESOPHYLLUM sensu stricto 

1956. Mesophyllum (Mesophyllum). Hill, p. F318, fig. 219-5.

Type species. Same as genus.

Diagnosis. This Mesophyllum has a highly discontinuous, patchy, nonuniform, and unstable pattern of 

prevailingly weak septa, with or without a few rather 

poorly defined bar carinae.

Remarks. On the one hand, this subgenus includes 

cystimorphs resembling Cystiphylloides with only a 

few weak septal traces; on the other, it probably grades 

into the sugenus Arcophyllum with partly strong septal crests and numerous bar or strip carinae.

Corals of this general category occur in Nevada Formation, unit 2. Because they are known only from 

fragmentary material, they are accordingly designated 

provisionally by letter only.

Mesophyllum (Mesophyllum) sp. b 

Plate 21, figures 1, 2

Figured specimens. USNM 159334. Lower part of 

the Nevada Formation, Sulphur Spring Range, Nev.

Early ephebic transverse sections show many rather 

stout septal crests of variable length; most of these are 

concentrated in a zone halfway between wall and axis. 

Inner dissepiments and tabellae are large. Carinae 

were not recognized. In fully mature stages the septal 

crests probably become thinner and more scattered.

Occurrence. Lower part of the Nevada Formation, 

unit 2; Devonian coral zone D2 . Southern Sulphur 

Spring Range, Nevada; locality M36.

Mesophyllum (Mesophyllum) sp. c 

Plate 21, figures 3, 4

Figured specimens. USNM 159335. Lower part of 

the Nevada Formation, northern Antelope Range,

Nev.

Mature transverse sections show sporadic, mostly 

vague septal traces in groups with widely scattered 

short septal crests. Inner dissepiments and tabellae are 

relatively smaller than in sp. b. In longitudinal section 

all the rather small dissepiments in a very wide zone 

are steeply inclined, the tabularium is very narrow, 

and the septal cones and calice are acutely conical. 

Thickened trabecular septal crests are developed at 

the inner edge of the dissepimentarium. Carinae were 

not recognized.

Occurrence. Lower part of Nevada Formation, 

unit 2; Devonian coral zone D. North end of the Antelope Range, Eureka County, Nev.; locality M1035.

Subgenus ARCOPHYLLUM Markov, 1926 (as a genus)

1926. Arcophyllum Markov, p. 49-60, pi. 3.

1940. Arcophyllum Markov. Lang, Smith, and Thomas, p. 20,

41.

1949. Arcophyllum Markov. Stumm, p. 39, 43, pi. 21, figs. 1-3. 

1922. Cosmophyllum Vollbrecht (not Blanchard, 1851), p. 17. 

1925. Cosmophyllum Vollbrecht. Wedekind, p. 39. 

1931. Cosmophyllum Vollbrecht. Wedekind and Vollbrecht,

pi. 28, figs. 1-6, pi. 32, fig. 8. 

1931. (?)Pseudocosmophyllum Wedekind and Vollbrecht, pi.

19, figs. 1-4, pi. 20, figs. 1, 2, 5.

1931. (?) Hemicosmophyllum Wedekind and Vollbrecht, pi. 

31, figs. 1-12, pi. 32, figs. 1-7.

1932. Cosmophyllum Vollbrecht. Wedekind and Vollbrecht,

pi. 33, figs. 1-4. 

1932. Hemicosmophyllum Wedekind and Vollbrecht, pi. 36,

figs. 1-8. 

1932. (?) Pseudocosmophyllum Wedekind and Vollbrecht, pi.

38, figs. 1-5. 

1937. Mesophylloides kirki Stumm, p. 441, 442, pi. 55, figs.

13a-b. 

1956. Mesophyllum (Arcophyllum) Markov. Hill, p. F318,

fig. 219-1. 

1956. (?) Mesophyllum (Hemicosmophyllum) Wedekind and

Vollbrecht. Hill, p. F318, fig. 219-3a, b, c. 

Type species. Arcophyllum typus Markov, 1926 

(by original designation). Middle Devonian; Calceola 

beds, western slope of Ural Mountains, Russia.

Diagnosis. The solitary Digonophyllinae have 

large, commonly elongate subcylindrical mature 

growth stages with thick external annulations and 

rejuvenescence constrictions. The calice, deep and 

inversely bell shaped to funnel shaped, is reflected 

peripherally as a platform or brim, which is either 

inclined axially, flattened, or broadly convex distally. 

The brim surface is ornamented by numerous edges 

of strip carinae. Septa are numerous, somewhat withdrawn from axial tabellae, and range from continuous 

to discontinuous as radially alined septal crests, the 

more continuous septa predominate. Septa are thickened locally in the tabularium; septa pass peripherally into a wide band of irregular dissepiments, within 

which some septa terminate short of the outer wall. 

Concentric rows of strip carinae are vaguely defined
SYSTEMATIC AND DESCRIPTIVE PALEONTOLOGY 69

peripherally in transverse thin sections; outermost 

strip carinae are commonly detached from the septa 

with which they are alined. Minor septa range from 

about one-half to nearly full length of the major septa; 

minor septa are more prone to discontinuity as alined 

septal crests. The dissepimentarium is wide, is commonly three-fourths of corallum radius, and consists 

of mostly small steeply inclined dissepiments which 

pass axially without discontinuity into the narrow 

medial zone of the tabellae. Outer dissepiments are 

mostly lonsdaleioid; inner dissepiments form concentric irregular chevrons and forks. Flat continuous tabulae are few. In longitudinal thin section, prominent 

isolated groups of diagonally inclined strip carinae 

abut in the outer dissepimentarium against parts lacking carinae.

Remarks. Because of delay in publication, Vollbrecht's (1922) genus Cosmophyllum appeared in 

print before Markov's (1926, p. 49) Arcophyllum, with 

which it is probably congeneric (Lang and others, 

1940, p. 20). However, the generic name Cosmophyllum, being an homonym, is superseded by Arcophyllum (Stumm, 1949, p. 43; Hill, 1956, p. F318).

Cosmophyllum-like digonophyllid Rugosa similar to 

Arcophyllum are recorded by Wedekind (1924, p. 84, 

85; 1925, p. 70, 71; 1926, table, p. 201) from the middle 

part of the Middle Devonian, Eifel district, Germany. 

The species in question are variously assigned to Cosmophyllum, Hemicosmophyllum, and Pseudocosmophyllum. Further taxonomic revision of these Eifel 

district genera is called for, in the light of field stratigraphic occurrence, to establish their relationship to 

Arcophyllum. Available illustrations of Hemicosmophyllum and Pseudocosmophyllum suggest that these 

corals may be closely related to, if not synonyms of, 

Arcophyllum; detailed variation study is needed, noting such features as presence of strip carinae in fully 

mature individuals. Hill's (1956, p. F317-F319) revision of the Digonophyllinae interprets Arcophyllum 

and other genera of this subfamily as subgenera of 

Mesophyllum Schliiter. Arcophyllum-like corals from 

southeastern Alaska have continuous septa, not the 

radially broken septa of Mesophyllum, and may 

accordingly be more closely allied to the genus Digonophyllum.

Mesophyllum (Arcophyllum) kirki (Stumm) 

Plate 21, figures 5-7; plate 22, figures 1-6

1937. Mesophylloides kirki Stumm, p. 441, pi. 55, figs. 13a-b.

Type and figured material. Holotype USNM 

94463; figured specimens USNM 159336-159342. 

Lower part of Nevada Formation, Lone Mountain, 

Eureka County, Nev.

Diagnosis. This large Arcophyllum has a subcylindrical mature corallum. The wide dissepimentarium 

consists of many columns of small steeply inclined dissepiments; the relatively narrow tabularium has few 

tabellae. The septa are withdrawn from the axis, somewhat thickened toward axis, and partly discontinuous 

as septal crests. Strip carinae of the peripheral part of 

the dissepimentarium are conspicuous in longitudinal 

sections, inconspicuous or shadowy in transverse sections. There is no stereozone, fossula, or other indication of bilateral symmetry in the mature corallum. The 

funnel-shaped calice has a wide brim, which is distally 

convex and proximally inclined toward narrow axial 

pit.

External features. The epitheca is thin and is 

absent over most of mature surface, possibly due to 

wear. The complete coralla of some individuals are 

very elongate and have annular folds and rejuvenescence constrictions. Strip carinae project conspicuously from the wide calice brim.

Transverse sections. The septal count is 80; 

this count includes minor septa, some of which are 

nearly as long as the major septa; some reach the corallum periphery, but most become unrecognizable 

peripherally in the generally complex dissepiment tissue. The septa are partly continuous and partly discontinuous as septal crests. Minute to small lonsdaleioid dissepiments compose a broad peripheral band; the 

dissepiments increase in size axially and pass without 

discontinuity into the tabellae of the narrow tabularium. Inner dissepiments are composed of a complex 

tissue which includes concentric but uneven traces, 

herringbone chevrons, forks, and minute semiglobose 

dissepiments that terminate against a single septum 

at all edges. Septal thickenings are of the tapering 

variety, without nodes and bumps of the acanthophylloids. In transverse sections the strip carinae appear 

as concentric radial rows of shadowy yet translucent 

thickenings mainly within the peripheral lonsdaleioid 

part of the dissepimentarium.

Longitudinal sections. The narrow tabularium 

about one-sixth the diameter of mature growth stages, 

consists of three or four axially inclined tabellae or less 

commonly a single complete tabula. Small peripheral 

dissepiments reveal a progressive size increase axially 

and pass without break into the less steeply inclined 

tabellae. Groups of prominent strip carinae, which 

terminate abruptly against the normal dissepiments, 

reveal an earlier growth stage calice brim.

Comparison with related forms. Mesophyllum 

(Arcophyllum) kirki of this report is, with some misgivings, assigned to Stumm's species founded upon an 

incomplete specimen from Nevada Formation, unit 2 

at Lone Mountain; this unit is also the source of the
70 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

material here described. The holotype transverse thin 

section of Stumm's Mesophylloides kirki shows the 

inner part of a corallum lacking the outer dissepimentarium band which normally reveals strip carinae in 

Arcophyllum. Stumm's holotype has thinner, more 

continuous septa with fewer septal crests than the 

specimens here described.

The material referred to Mesophyllum (Arcophyllum) kirki resembles M. (A.) dachsbergi (Vollbrecht) 

and M. (A.) limbatum (Wedekind and Vollbrecht), 

the first was originally placed in Cosmophyllum and 

the second in Hemicosmophyllum. Mesophyllum 

(Arcophyllum) dachsbergi differs from M. (A.) kirki 

in having a broader tabularium and more numerous 

tabellae. These differences apply also to M. (A.) limbatum. An undescribed Middle Devonian Arcophyllumlike coral from the Alberto Islands, southeast Alaska, 

has continuous septa, a wider tabularium with more 

numerous tabellae, and a well-developed strip carinae 

in the larger individuals.

Occurrence. Lower part of the Nevada Formation, 

unit 2. Devonian coral subzone D3 . Lone Mountain; 

localities M29, M1036. The corals here assigned to 

Mesophyllum (Arcophyllum) kirki characterize the 

uppermost beds in the "Spirifer pinyonensis" zone of 

previous usage. Comparison with Arcophyllum dachsbergi and A. limbatum suggests that coral subzone D3 

is within the Middle Devonian. According to Wedekind and Vollbrecht (1931, pi. 28; 1932, pi. 36), Arcophyllum dachsbergi and A. limbatum occupy the zone 

of "Cosmophyllum dachsbergi" at Gerolstein, Eifel 

district, Germany. Wedekind (1925, p. 71 table) 

assigns the zone of "Cosmophyllum dachsbergi" to the 

upper part of his coral zone "St-Stenophyllumstufe," 

which is shown by him as well up in the middle Middle 

Devonian. Markov's type of Arcophyllum, being 

reported from Ca/ceo/a-bearing beds of the Ural Mountains, may be older. The comparable southeast Alaskan ArcophyHum-like coral from the Alberto Islands 

is associated with Acanthophyllum, Australophyllum, 

and Heliolites in a fauna which probably underlies 

that containing Stringocephalus in this Alaskan stratigraphic section.

Subfamily ZONOPHYLLINAE Wedekind. 1924

Reference form. Zonophyllum duplicatum Wedekind. Early Middle Devonian; Germany.

The cystimorph Digonophyllidae lack continuous 

septal lamellae; septal crests in growth cones that tend 

to be greatly thickened stereoplasmically, form septal 

wreaths or festoons where in contact laterally.

The Zonophyllinae range from cystimorphs with a 

few sporadic septal crests to those with radially alined, 

usually stout thickened crests and either a crescentic

festoon or a completely circular stereoplasmic septal 

wreath.

Nepionic growth stages have thick stumpy septa in 

lateral contact. The stereoplasm of septal wreaths 

passes from one growth cone to those adjacent, as 

revealed in longitudinal thin section.

In Germany and in the Great Basin, the Zonophyllinae are characteristic of the early Middle Devonian; 

none have been recognized above coral zone D2 .

Genus ZONOPHYLLUM Wedekind, 1924

1924. Zonophyllum Wedekind, p. 12-21, figs. 1-8.

1940. Zonophyllum Wedekind, Lang, Smith, and Thomas, p.

142. 

1949. Zonophyllum Wedekind, Stumm, p. 46-47, pi. 22, figs.

16-22. 

1956. Zonophyllum Wedekind, Hill, p. F314-F315 (in part),

fig. 216-la-c.

Type species. Z. duplicatum Wedekind by subsequent designation, Lang, Smith and Thomas (1940 p. 

142). Lower Middle Devonian; Nohner Schichten at 

Nohn Eifel district Germany. From Wedekind's (1924, 

p. 85) zone D (Digonophyllumstufe), which is the 

lowermost Middle Devonian as interpreted by him.

Diagnosis. Medium-size and small solitary cystiphylloids of ceratoid to cylindrical growth habit have 

thick stumpy to long septal crests in the peripheral 

zone of mature individuals; in longitudinal section the 

septal crests reveal thick trabecular spines which are 

directed distally and axially. The peripheral zone of 

small and medium-sized steeply inclined dissepiments 

passes axially into a zone of large dissepiments and 

periaxial tabellae. The axial zone has irregular, distally 

convex tabulae, some of which may be almost flat and 

wide. The ontogeny is distinctive. The late nepionic 

stage has about twelve thick stumpy septa touching 

laterally and has a suggestion of bilateral symmetry 

though no specific cardinal septum may be recognizable. In mid-neanic stage, there is a peripheral zone 

of dissepiments with thinner septal crests and an inner 

periaxial zone of thickened septal crests forming a septal wreath. In later growth stages, dissepiments and 

tabellae are introduced between the two septal zones. 

Septal crests are usually thick and numerous peripherally in earlier ephebic stages. Advanced ephebic stages 

are cystiphylloid with traces of septal crests.

Remarks. Wedekind's (1924) researches by means 

of serial sections demonstrated that cystiphylloid 

rugosa of the Zonophyllum group were distinct from 

those of the true Cystiphyllum group. In this connection ontogenetic changes seem to be highly significant. 

Cystiphyllum itself is a Silurian genus (Lang and others, 1940, p. 48). The Devonian strata include descendants and no doubt numerous homeomorphic strains, 

such as Zonophyllum of the late Early and the Middle
LOCALITY REGISTER 71

Devonian. In accordance with Wedekind's views 

(1924, p. 29), the early Middle Devonian Zonophyllum 

group was possibly the ancestral stock whence came 

the very numerous Digonophylloids of the ensuing 

Middle Devonian, all of which include cystiphylloid 

types.

Wedekind's theory of the septal cone (septalkegel) 

is set forth in connection with his description of Zonophyllum and its presumed evolutionary offshoots 

(1924, p. 21). This theory is reviewed by Hill (1956, 

p. F252, F314), who noted that in some of the rugose 

corals under consideration, septal tissue is developed 

only in successive inverse cones of thickening in which 

large trabeculae are present.

Zonophyllum haguei n. sp. 

Plate 20, figures 3, 4, 9,10

Type material. Holotype USNM 159333, paratype 

USNM 159331; Grays Canyon, Eureka district, 

Nevada. Lower part of the Nevada Formation; coral 

zone D2 .

Diagnosis. Zonophyllum has a peripheral wreath 

of thick septal crests in the dissepimentarium and 

small very steep outer dissepiments. For part of the 

septal wreath, thick crests are joined discontinuously 

in a general stereoplasmic mass. The tabularium is 

occupied by large axially inclined tabellae and rather 

widely spaced tabulae which are, for the most part, 

distally convex.

External features. The mature corallum is rather 

small and subcylindrical.

Transverse sections. All dissepiments and tabellae 

are concave peripherally. There is progressive increase 

in size from small outer dissepiments to the large tabellae but no discrete border between dissepimentarium 

and tabularium. Some thick outer septal crests originating at or near the outer wall penetrate two or more 

rows of the small peripheral dissepiments. Stubby septal crests are sparsely developed within the tabularium.

Longitudinal sections. The stereoplasmic deposits 

in the outer septal wreath partly fill the dissepimentarium. Large and elongate trabeculae extend inward 

and distally; in some instances they penetrate columns 

of smaller dissepiments. Dissepiments merge with size 

increase into large less steeply inclined periaxial tabulae. A few nearly flat tabulae may be more than onethird the corallite diameter in the mature stages.

Comparison with related forms. Zonophyllum 

haguei differs from Z. duplicatum Wedekind, the type 

species, in lacking a well-defined inner wreath of thickened septal spines; the outer wreath of spines in haguei 

shows a more extensive deposit of stereoplasm. Other 

forms assigned to Zonophyllum from the Great Basin 

may represent distinct species but, being known from

insufficient material, are not formally described.

Occurrence. Nevada Formation, unit 2; Devonian 

coral subzone D2 . Grays Canyon, Northern Fish Creek 

Range; localities M3, M51.

Zonophyllum sp. a 

Plate 20, figures 5, 6

Figured specimens. USNM 159332; lower part of 

the Devonian section, Ranger Mountains, Nevada.

Septal crests are short and scattered, not joined as 

in haguei, to form a discrete septal wreath. The tabellae are large in the wide tabularium.

Occurrence. Lower part of Devonian section; probably coral zone DL Ranger Mountains, Nev.: locality 

M1058.

Zonophyllum sp. b 

Plate 20, figures 11, 12

Figured specimens. USNM 159351, lower part of 

the Nevada Formation; northern Fish Creek Range, 

Nevada.

The few septal crests are largely confined to an inner 

zone of diffuse stereoplasmic thickening which borders 

the dissepimentarium.

Occurrence. Lower part of Nevada Formation, unit 

2; Devonian coral zone D2 . Grays Canyon, northern 

Fish Creek Range; locally M51.

LOCALITY REGISTER

(see figs. 1, 2, 3, 4)

7. Southern Great Basin

1. Desert Range, Clark County, Nev.

Locality M1057. Army Map Service, Las Vegas, Nev., 

quadrangle; southwest edge of Desert Range east of 

Braun's Playa, in the WY2 sec. 10, T. 16 S., R. 58 E., 

altitude 3,200 ft. Early Middle Devonian. Collected by 

C. R. Longwell.

2. Ranger Mountains, east of Frenchman Flat, Clark County,

Nev. 

Locality M1034. Frenchman Lake SE quadrangle,

Nevada; lat 36°46'45" N., long 115°52'00" W. Between

10 and 25 ft above base of Nevada Formation. Collected

by F. G. Poole. 

Locality M1058. Frenchman Lake quadrangle, Nevada;

southeast side of Frenchman Flat, 1.5 miles east of Nye

County-Clark County line in higher part of the southern

Ranger Mountains; 1,500 ft northwest of summit 4907.

Limestone about 20 ft above base of Nevada Formation;

fossil zone 4 (Devonian unit A) of M. S. Johnson and

D.E.Hibbard(1957).

3. Northern Panamint Mountains, Ubehebe district, Calif.

Locality M184. Marble Canyon quadrangle, California; 

south side of Andy Hills in northwest corner of quadrangle, east side of Hidden Valley, on central ridge of 

Andy Hills near summit 5612. Upper part of McAllister's 

(1952, p. 15-17) Hidden Valley Dolomite, unit 3b containing Papiliophyllum and Costispirifer arenosus.

Locality M1065. Marble Canyon quadrangle, California. 

Andy Hills, south side; measured section across Me-
72 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

Allister's unit 3b of the Hidden Valley Dolomite; in

same area as locality Ml84. Zoned collections A through

H by J. F. McAllister, April 1965. 

4. Funeral Mountains, Calif.

Locality M1059. Ryan quadrangle, 1.8 miles N. 48° W.

of Pyramid Peak, Funeral Mountains. Uppermost part

of Hidden Valley Dolomite with rugose coral fauna.

Late Early Devonian. Collected by J. F. McAllister,

June 1962. 

Locality M1060. Ryan quadrangle, 1.6 miles N. 58° W.

of Pyramid Peak, Funeral Mountains. Uppermost part

of Hidden Valley Dolomite with rugose coral fauna.

Late Early Devonian. Collected by J. F. McAllister,

April 1962. 

Locality M1061. Ryan quadrangle, northwest slope of

4298 hill, 4.5 miles S. 47° E. of Pyramid Peak, Funeral

Mountains, altitude 3,750 ft. Uppermost part of Hidden

Valley Dolomite with rugose coral fauna. Collected by

J. F. McAllister, May 1964. 

Locality M1062. Ryan quadrangle, 6.2 miles S. 71° E. of

Pyramid Peak, Funeral Mountains, altitude 3,450 ft.

Uppermost part of Hidden Valley Dolomite with rugose

coral fauna. Collected by J. F. McAllister, November

1963. 

Locality M1063. Ryan quadrangle, 4.7 miles S. 51° E. of

Pyramid Peak, Funeral Mountains, altitude 3,700 ft.

Uppermost part of Hidden Valley Dolomite with rugose

coral faunas. Collected by J. F. McAllister, April 1965. 

Locality M1064. Ryan quadrangle, 6.23 miles S. 72° E. of

Pyramid Peak, Funeral Mountains, altitude 3,450 ft.

Uppermost part of Hidden Valley Dolomite with rugose

coral faunas. Collected by J. F. McAllister, March 1965.

II. South-central Great Basin

1. Hot Creek Range, Nev., middle part

Locality M1066. Army Map Service, Tonopah, Nev., 

sheet. Probably in Hot Creek Canyon about 4 miles 

west-northwest of Hot Creek Ranch houses. Devonian 

including lower part of Nevada Formation with rugose 

corals. Collected by H. E. Cook (thesis, Univ. California, 

Berkeley, 1966).

2. Monitor Range, Nev., middle part

Locality M1067. Army Map Service, Tonopah, Nev.

sheet. Dobbin Summit, northern Nye County, Nev.; 1

mile southeast of East Dobbin Summit Spring on east

side of canyon; west edge sec. 32, T. 13 N., R. 49 E.

Lower part of Rabbit Hill Limestone. 

Locality Ml 068. Dobbin Summit, northern Nye County,

Nev.; same as Ml067, but a higher zone in Rabbit Hill

Limestone. 

Locality M1069. Dobbin Summit, northern Nye County,

Nev.; same as M1067, but upper part of Rabbit Hill

Limestone.

117. Central Great Basin

1. Monitor Range, northern part

Locality M48. Horse Heaven Mountain quadrangle,

Nevada. West side Copenhagen Canyon, southeast slope

Rabbit Hill, altitude 7,100 ft. Type section Rabbit Hill

Limestone, Lower Devonian. 

Locality M49. Horse Heaven Mountain quadrangle,

Nevada. One mile north of Locality M48, SE^SE1/^ sec.

35, T. 16 N., R. 49 E.; in Rabbit Hill Limestone. 

Locality Ml 87. Horse Heaven Mountain quadrangle,

Nevada. Same as Locality M48 on southeast side of 

Rabbit Hill. Rabbit Hill Limestone, Lower Devonian.

2. Antelope Range, northern part

Locality M1035. Bellevue Peak quadrangle, Nevada. 

North tip of Antelope Range, west side of lower slope 

east of hill 6754, altitude 6,680 ft. Nevada Formation, 

unit 2.

Locality M1035a. Same as Locality M1035, but a lower 

stratigraphic horizon.

Locality M1053. Bellevue Peak quadrangle, Nevada. 

North end of Antelope Range, NEy4 sec. 16, T. 16 N., 

R. 51 E., 2,200 ft north-northwest of summit 7602, altitude 7,000 ft. Nevada Formation, unit 2.

Locality M1055. Bellevue Peak quadrangle, Nevada. 

North end of Antelope Range; top of spur, altitude 7,100 

ft and 500 ft southwest of Ml053. Nevada Formation, 

unit 2.

3. Southern Fish Creek Range

Locality M1033. Bellevue Peak quadrangle, Nevada. 

North tip of Fenstermaker Mountain, 3 miles southwest 

of Fish Creek Springs near top of range at altitude 6,700 

ft; 1,200 ft north of southern edge of quadrangle. Nevada 

Formation, unit 2.

Locality M1070. Cockalorum Wash quadrangle, Nevada. 

First north-south ridge east of southern Fish Creek 

Range, near south boundary of Eureka County; 0.8 mile 

southeast of Eightmile Well near top of knob at altitude 

6,400 ft. Nevada Formation, unit 2.

Locality Ml 043. Cockalorum Wash quadrangle, Nevada. 

One mile south of Nyeka (BM 7352), on line between 

sees. 4 and 5, T. 14 N., R. 52 E. Limestone conglomerate.

4. Northern Fish Creek Range

Locality M3. Pinto Summit quadrangle, Nevada. Near 

mouth of Grays Canyon, 1 mile south of Pinnacle Peak 

and 800 ft east of quadrangle boundary; east side of 

canyon near bottom, altitude 7,250 ft. Nevada Formation, unit 2.

Locality M51. Same as Locality M3.

5. Mahogany Hills

Locality M27. Bellevue Peak quadrangle, Nevada. Combs 

Peak area, 1.7 miles south of Combs Peak on top of spur 

in saddle at altitude 7,520 ft. Nevada Formation, unit 2.

Locality M1071. Whistler Mountain quadrangle, Nevada. 

Mahogany Hills, east side (Mountain Boy Range) at 

Modoc Peak; near quadrangle boundary at south side of 

Modoc Peak. Nevada Formation, units 1 and 2.

Locality M1084. Bellevue Peak quadrangle, Nevada. 

West side of Mahogany Hills, 1.75 miles west-southwest 

of summit of Combs Peak. Lower part of Nevada Formation.

6. Lone Mountain (see fig. 3)

Locality Ml045. Bartine Ranch quadrangle; Lone Mountain, northwest side, 2,200 ft S. 50° W. of U. S. Mineral 

Monument No. 6828 and about 1 mile northwest of 

summit of Lone Mountain, Nevada Formation, unit 2, 

upper 250 ft.

Locality Ml052. Bartine Ranch quadrangle; Lone Mountain, northwest side. Same as locality M1045.

Locality Ml048. Bartine Ranch quadrangle; Lone Mountain, northwest side, 2,500 ft S. 33° W. of U. S. Mineral 

Monument No. 6828, altitude 6,640 ft. Nevada Formation, unit 2.

Locality M286. Bartine Ranch quadrangle; Lone Mountain, northwest side, 2,700 ft S. 39° W. of U.S. Mineral
LOCALITY REGISTER 73

Monument No. 6828, altitude 6,560 ft. Nevada Formation, unit 1.

Locality Ml036. Bartine Ranch quadrangle; Lone Mountain, northwest side, 3,500 ft N. 62° W. of Lone 

Mountain summit 7,936 at altitude 6,900 ft. Nevada Formation, unit 2 near middle.

Locality M1044. Bartine Ranch quadrangle; Lone Mountain, west side, 2,400 ft southwest of Lone Mountain 

summit 7,936 at altitude 7,280 ft. Nevada Formation, 

unit 1, upper beds.

Locality M29. Bartine Ranch quadrangle; Lone Mountain, west side; 2,000 ft south-southwest of Lone Mountain summit 7,936, above altitude of 7,200 ft. Nevada 

Formation, unit 2.

Locality M55. Bartine Ranch quadrangle; Lone Mountain, west side. Same vicinity and zone as locality M29.

Locality Ml4. Bartine Ranch quadrangle; Lone Mountain, south side, 3,800 ft S. 7° E. of Lone Mountain 

summit 7,936 at altitude 7,020 ft. Nevada Formation, 

unit 1.

Locality M1050. Bartine Ranch quadrangle; Lone Mountain, south side, 4,000 ft S. 16° E. of Lone Mountain 

summit 7,936 at altitude 6,960 ft; 1,200 ft west of quadrangle boundary. Nevada Formation, unit 2, lowermost 

beds.

Locality Ml046. Bartine Ranch quadrangle; Lone Mountain, south side, 4,500 ft S. 23° E. of Lone Mountain 

summit 7,936 at altitude 7,000 ft; 700 ft west of quadrangle boundary. Nevada Formation, unit 2.

Locality M1038. Whistler Mountain quadrangle; Lone 

Mountain, south side; at west edge of quadrangle, 2,500 

ft S. 36° W. of summit 7,360 a.t altitude 6,890 ft. Nevada 

Formation, unit 2.

Locality M1049. Whistler Mountain quadrangle; Lone 

Mountain, south side; 1,700 ft S. 34° W. of summit 7,360 

at altitude 6,920 ft; 600 ft east of quadrangle boundary. 

Nevada Formation, unit 2, upper beds.

Locality M1037. Whistler Mountain quadrangle; Lone 

Mountain, south side. Southeast pediment slope 1.5 

miles north of Highway 50; 6.500 ft S. 11° E. of summit 

7,360, altitude 6,420 ft. Nevada limestone in fault contact 

with Permian beds.

7. Southern Roberts Mountains

Locality M1042. Roberts Creek Mountain quadrangle, 

Nevada; 1.75 miles north of Roberts Creek Ranch house, 

0.75 mile east of Roberts Creek and 1.25 miles northwest 

of top of hill 7,504 (Pyramid Hill); top of knob, altitude 

6,850 ft. Nevada Formation, unit 1.

8. Northern Roberts Mountains

Locality M1072. Roberts Creek Mountain quadrangle, 

Nevada. Near top of main spur south of Niel Creek, 1.4 

miles S. 30° W. of Western Peak and 1,000 ft northeast 

of summit 8,168, altitude 7,800 ft. Same as Merriam 

(1940, pi. 1) locality 1 (R 24). Lower Devonian.

Locality M1073. Roberts Cr&ek Mountain quadrangle, 

Nevada. In saddle 1,500 ft southeast of Cooper Peak at 

altitude 9,150 ft. Nevada Formation, unit 2. Same as 

Merriam (1940, pi. 1) locality 11 (R 1).

Locality M1054. Same as locality M1073.

9. Northern Simpson Park Range

Locality M1074. Walti Hot Springs quadrangle, Nevada. 

Foothills 0.5 mile southeast of Walti Ranch and west of 

McClusky Peak at middle of north edge sec. 4, T. 23 N., 

R. 48 E. Rabbit Hill Limestone; Lower Devonian. Collections by R. J. Roberts and R. E. Lehner, 1954.

Locality M1075. Horse Creek Valley quadrangle, Nevada. 

Near mouth of Coal Canyon, east side along ridge top of 

summit 6,909, SEy4 sec. 17, T. 25 N., R. 49 E. Rabbit 

Hill Limestone, Lower Devonian.

Locality M1032. Horse Creek Valley quadrangle, Nevada. 

East side of Coal Canyon, NW% sec. 21, T. 25 N., R. 

49 E. Rabbit Hill Limestone.

Locality M1076. Horse Creek Valley quadrangle, Nevada. 

Near mouth of Coal Canyon, east side. Mostly float 

below ridge top of summit 6,909 and down slope to west. 

Rabbit Hill Limestone, Lower Devonian. 

10. Southern Sulphur Spring Range (see fig. 5)

Locality M4. Garden Valley quadrangle, Nevada. Prince 

of Wales mine area; 1,200 ft southeast of mine and 

northwest of summit, 7,530, altitude 7,300 ft near old 

charcoal pit. Nevada Formation, unit 1.

Locality M69. Garden Valley quadrangle, Nevada. 

Prince of Wales mine area. Foat material near locality 

M4.

Locality M1041. Garden Valley quadrangle, Nevada. 

Prince of Wales mine area. Measured section through 

locality M4 southeast of mine, and northwest of summit 

7,530. Nevada Formation, unit 1.

Locality M68. Garden Valley quadrangle, Nevada. 

Prince of Wales mine area. Top of ridge east of mine at 

signal station 7,530. Nevada Formation, upper part of 

unit 1; Acrospirifer kobehana fauna.

Locality M67. Garden Valley quadrangle, Nevada. 

East side of range 0.8 mile N. 65° E. of Bailey Pass at 

altitude 6,520 ft, 0.1 mile south of road on top of spur. 

Nevada Formation, unit 1.

Locality M36. Garden Valley quadrangle, Nevada. 

South of Bailey Pass 0.9 mile at altitude 7,200 ft on 

north side of knob. Nevada Formation, unit 2 with 

Spirifer pinyonensis and Chonetes macrostriata.

Locality M198. Garden Valley quadrangle, Nevada. 

Same as Locality M36.

Locality M197. Garden Valley quadrangle, Nevada. 

S. 12° E. of Bailey Pass 0.8 mile on ridge top at altitude 

7,080 ft; 1,000 ft northeast of Locality M36. Rabbit Hill 

Lower Devonian fauna.

Locality M1031. Garden Valley quadrangle; southern 

Sulphur Spring Range, 0.5 mile north-northwest of 

Romano Ranch house at east end of old lake terrace, 

altitude 6,000 ft; 500 ft west of BM 5,836. Nevada Formation, unit 2 with rugose corals.

Locality M1078. Garden Valley quadrangle; southern 

Sulphur Spring Range 0.5 mile northwest of Romano 

Ranch house, south side of old lake terrace at altitude 

6,000 ft; 1,000 ft southwest of BM 5,836 and 500 ft southwest of Locality M1031. Nevada Formation, unit 2.

Locality M1047. Garden Valley quadrangle; southern 

Sulphur Spring Range about 0.5 mile west-northwest of 

Romano Ranch house, altitude 6,000 ft; about 0.5 mile 

southwest of BM 5,836. Nevada Formation, unit 2.

Locality M1079. Garden Valley quadrangle; southern 

Sulphur Spring Range 1.5 miles N. 83° W. of Romano 

Ranch house; 2,000 ft northeast of summit 6,933 at 

altitude 6,560 ft. Nevada Formation, unit 2 with quartzite interbeds and fossils.

Locality M1080. Garden Valley quadrangle; southern 

Sulphur Spring Range; S. 86° E. of Mulligan Gap 1.2 

miles on top of northeast spur, altitude 6,885 ft; 2,500
74 LOWER AND LOWER MIDDLE DEVONIAN RUGOSE CORALS OF THE CENTRAL GREAT BASIN

ft northeast of 7,446 summit. Lower Devonian with 

rugose corals.

Locality Ml081. Garden Valley quadrangle; southern 

Sulphur Spring Range. In ravine 600 feet west of 

Locality M1080 at altitude 6,800 ft; 2,000 ft northeast of 

7,446 summit. Lower Devonian with Rabbit Hill Helderberg fauna.

Locality M1040. Garden Valley quadrangle, Nevada; 

southern Sulphur Spring Range. South-southeast of 

Bailey Pass 1.1 miles near range top and 750 ft northeast of summit 7,378, altitude 7,240 ft. Lower Devonian 

beds with Oriskany fossils.

Locality M56. Garden Valley quadrangle, Nevada. 

Southernmost Sulphur Spring Range, 1.6 miles S. 25° 

E. of Mulligan Gap, east side of ravine on top spur at 

altitude 6,690 ft. Nevada Formation, unit 1 with Oriskany fauna.

Locality M186. Garden Valley quadrangle, Nevada. 

Southernmost Sulphur Spring Range, 1.7 miles S. 22° 

E. of Mulligan Gap, east side of ravine, altitude 6,560 

ft, and 750 ft southwest of Locality M56. Nevada Formation, Beacon Peak Dolomite Member with Rabbit 

Hill Helderbergian fauna.

Locality M1039. Garden Valley quadrangle, Nevada. 

Southernmost Sulphur Spring Range. 1.5 miles southsoutheast of Mulligan Gap and 1,500 ft west of 6,927 

summit on east side of ravine. Nevada Formation, unit 1.

Locality M1051. Garden Valley quadrangle, Nevada. 

Southernmost Sulphur Spring Range. 1.25 miles southsoutheast of Mulligan Gap near top main spur at altitude 7,150 ft; 2,700 ft south of summit 7,446. Nevada 

Formation, unit 2.

Locality Ml 077. Garden Valley quadrangle, Nevada. 

Southernmost Sulphur Spring Range; 1.3 miles S. 25° 

E. of Mulligan Gap on east side of ravine at altitude 

6,720 ft. Nevada Formation, unit 1, with Lower Devonian rugose coral fauna.

Locality M1082. Garden Valley quadrangle, southernmost Sulphur Spring Range, east side. 1.2 miles due 

west of BM 5,867 on top spur at altitude 6,440 ft; 1,400 

ft east-northeast of summit 6,474. Lower Devonian 

Beacon Peak Dolomite Member with Rabbit Hill 

Helderberg fauna.

7V. North-central Great Basin

1. Northern Sulphur Spring Range

Locality M1018. Mineral Hill quadrangle, Nevada. 

Northwest of Union Summit in McColley Canyon. 

Lower beds of the McColley Canyon Member of the 

Nevada Formation of Carlisle and others (1957). Lower 

Devonian with rugose coral fauna.

Localities M1021 to M1025 inclusive. Mineral Hill quadrangle, Nevada. Same general areas as locality M1018, 

in same stratigraphic unit. Lower Devonian with rugose 

corals.

2. Cortez Mountains

Locality M1083. Cortez quadrangle, Nevada. Southeast 

side of Mt. Tenabo, east slope of the Cortez Mountains; 

northeast side of upper Horse Canyon, 1,000 ft north, 

700 ft west of SE cor. sec. 4, T. 26 N., R. 48 E., altitude 

7,900 ft. Rabbit Hill Lower Devonian fauna with 

Syringaxon and Pleurodictyum.

V. West-central Great Basin

1. Northern Toquima Range

Locality Ml 150. Wildcat Peak quadrangle, Nevada. 

SWy4 sec. 16, T. 16 N., R. 46 E., top of spur 1,600 ft 

south of summit 7,188 and 2,800 ft east of Petes Canyon 

road. Rabbit Hill Limestone on Silurian graptolitebearing shale. 

2. Toiyabe Range

Locality M1151. Austin quadrangle, Nevada. SEJ/4 sec. 

36, T. 17 N., R. 42 E.; in first main canyon at range 

front 1 mile south of Reeds Canyon. Devonian coralbearing limestone.

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INDEX

Page 

A

abditum, Endophyllum .................................. 56

Acanthophyllum ...................................... 21, 25, 70

Acinophyllum .................................................... 58

Acknowledgments ............................................ 6

acuminatum, Laccophyttum ......................... 39

Syringaxon ................................................ 39

affinis, Bittingsastraea .................................. 30, 64

Age, coral zone B ............................................ 32

coral zone C ............................................ 30, 32

coral zone D .............................................. 30

coral subzone Dj .................................... 32, 33

coral subzone D2 ...................................... 31

coral zones, distant regions .................. 29

Early Devonian ........................ 29, 32, 33

Early middle Devonian .................. 29

Late Silurian .................................... 33

Nevada Formation, unit 2 ...................... 31

aggregatum, Cystiphylloides ...................... 64, 65

Agoniatites nevadensis .................................. 30

Alaska .................................................. 29, 31, 66, 69

Alberto Islands, Alaska .................................. 70

Alleynia .............................................................. 39

(Nicholsonia) ............................................ 39

americanum, Cystiphylloides ........................ 66

Ammonoids .............................................. 18, 30, 31

Amphipora ............................................ 9, 15, 17, 22

Amplexus invaginatus .................................... 42

lonensis .................................................... 42, 43

magnus ...................................................... 56

nevadensis ................................................ 42, 56

Anastrophia verneuili .................................... 29

Andy Hills ...................................................... 32, 33

Antelope Range .............. 6, 43, 52, 56, 62, 63, 68

Antelope-Roberts Mountains facies belt .... 1,

6, 13, 15, 17, 29 

antelopensis, Bethanyphyllum ...... 21, 33, 53, 54

Aphroidophyllum ............................................ 58, 64

arachne, Billingsastraea ................................ 64

Billingsastraea nevadensis.. 34, 38, 62, 63, 64. 

Radiastraea ............................................ 63, 64

Arachnophyllidae .............................................. 37

Arcophyllum .......................... 30, 34, 36, 67, 68, 70

dachsbergi .................................................. 70

kirki ............................................................ 30

limbatum ".................................................... 70

typus ............................................................ 68

(Arcophyllum) dachsbergi, Mesophyllum.. 70 

kirki, Mesophyllum .................... 21, 30, 34, 69

limbatum, Mesophyllum .......................... 70

Mesophyllum .............................................. 67

Acrospirifer kobehana.... 13, 20, 24, 32, 33, 52, 66

fauna .......................................................... 8, 20

zone ............................................................ 18, 30

sp .......................................................... 20, 32, 50

arenosus, Costispirifer ...................... 8, 11, 17, 19

20, 23, 29, 31, 32, 33, 49 

Assemblage, brachiopod .................................. 31

coral-brachiopod-tribolite ...................... 24

astreiformis, Stauria ...................................... 45

Atelophyttum ...................................................... 67

Atrypa nevadana .............................................. 21

Atrypa Peak ...................................................... 42

Aulacophyllum .............................. 24, 32, 33, 46, 53

convergens .................................................. 47

sp. c .................................................... 20, 33, 46

[Italic page numbers indicate major references]

Page 

Austin, Nev .................................................... 17, 31

Austin quadrangle ............................................ 64

Australophyttum .............. 28, 29, 33, 34, 57, 61, 70

cyathophylloides ........................................ 57

landerensis ............................ 20, 23, 33, 34, 57

sp .................................................................. 57

B

Barrandeophyttum ............................................ 39

perplexum .................................................. 39

Bartine Ranch quadrangle ............................ 5

Bay State Dolomite Member ............ 9, 13, 15, 16,

17, 19, 21, 22, 23 

lithology ...................................................... 16

Beacon Peak Dolomite Member .......... 6, 8, 9, 11,

15, 17, 19, 23, 28, 31, 40 

lithology ...................................................... 15

type section ................................................ 11

Beeehwood Limestone ...................................... 47

Belmont mine ............................................ 19, 22, 25

Bensberg, Germany .......................................... 60

Bethanyphyllidae .................. 17, 27, 29, 30, 33, 53

Bethanyphyllum .................................. 24, 33, 53, 54

antelopensis ................................ 21, 33, 53, 54

lonense .......................................... 21, 33, 54, 55

robustum .................................................. 53, 54

sp. d .......................................................... 33, 55

Bittingsastraea ............................ 20, 23, 24, 31, 32,

34, 35, 37, 38, 58, 61, 62 

affinis ........................................................ 30, 64

arachne ........................................................ 64

billingsi nevadensis .................................. 63

nevadensis .......... 21, 30, 31, 34, 38, 61, 62, 63

arachne .......................... 34, 38, 62, 63, 64

verritti ...................................................... 31, 64

sp. T .................................................... 32, 34, 64

(Billingsastraea), Phittipsastraea ................ 62

billingsi nevadensis, Bittingsastraea ............ 63

Biofacies, Breviphrentis ................................ 23

Hexagonaria-Sociophyllum .................. 23, 25

Kobeha ...................................................... 23, 24

Papiliophyllum ........................................ 23, 24

Phittipsastraea .................................. 19, 23, 25

Syringaxon ................................................ 23

Biofacies aspects, Devonian corals .............. 22

Biofacies indicator .......................................... 5, 23

Bioherms ................................................ 9, 22, 23, 25

Biometrics ........................................................ 34, 38

Biostratigraphy, central Great Basin

Devonian .................................... 17

Birenheide, Rudolf, cited ............................ 67, 68

bowerbanki, Endophyttum .............................. 56

Brachiopods .................................... 5, 9, 13, 15, 16,

17, 18, 19, 20, 23, 24, 25, 29, 30, 50 

spiriferoid .................................................. 17

Branik Limestone, Bohemia .......................... 39

Breviphrentis ............ 24, 27, 33, 38, 41, 42, 44, 45

invaginatus .................... 30, 42, 43, 44, 46, 56

kobehensis .................................................. 29

(Breviphrentis) invaginatus,

Siphonophrentis .............. 21, 30, 32,

33, 38, 42, 45, 54, 56 

kobehensis, Siphonophrentis .............. 20, 30,

33, 42, 43, 66 

Siphonophrentis ...................................... 32, 41

Breviphyttum ............................................ 41, 43, 56

Page 

Brevip h yllum Continued

lonensis ........................................................ 42

breviseptatum, Eurekaphyttum .................. 33, 52

Bryozoans .......................................................... 23

Biicheler Schichten ........................................ 45, 46

Bush Creek ........................................................ 30

caespitosum, Cyathophyllum ........................ 58

Disphyllum ................................................ 58

Calcarenite .................................................. 8, 11, 31

Calceola beds, Russia .................................... 68, 70

Caledonian Revolution .................................... 27

Canada, western ................................ 29, 31, 56, 64

Carlin, Nev ...................................................... 20, 31

Caryophyllia gigantea .................................... 41

castanea, Leiorhynchus ................................ 13, 16

Central Peak ...................................................... 20

Cephalopods ...................................................... 18, 24

orthoceratid .............................................. 23, 24

Ceratophyllum .................................................. 53

Charcoal Gulch .................................................. 21

Chert nodules .................................................... 8, 9

Chewelah area .................................................. 31

Chonetes macrostriata ...................... 20, 21, 32, 52

Chonophyllidae ......._............. 28, 33, 37, 38, 41, 55

Conophyllum ...................................................... 55

Cladocora goldfussi .......................................... 58

Clams .................................................................. 24

Clark County, Nev .......................................... 32, 52

Clermont, Australia ........................................ 57

Coal Canyon .................................. 17, 19, 28, 33, 40

Cockalorum Wash quadrangle ...................... 5

Coeymans Limestone, New York .... 23, 30, 42, 44

coeymanensis, Gypidula .......................... 20, 29, 50

Collection procedure ........................................ 4, 5

Columnaria ................................................ 45, 58, 61

(Cyathophylloides) disjuncta ................ 45

Combs Peak .......................................... 8, 15, 30, 54

Compressiphyttum ............................................ 41

Confusion Range .............................................. 27

Conglomerate .................................................... 15

Conodonts .................................................. 17, 19, 31

Contact Gulch .................................................. 8, 20

convergens, Aulacophyllum ............................ 47

Odontophyllum .......................................... 47

Convergent genera .......................................... 34

cooperi, Trematospira .................................... 29

Copenhagen Canyon ...................................... 17, 19

Coral assemblage, Gotlandian ................ 17, 19, 28

Coral assemblages, succession ...................... 27

Coral distribution, Devonian ........................ 22

Coral evolution, biologic factors ................ 25, 27

Devonian .................................................... 28

geologic factors ...................................... 25, 27

Late Silurian ............................................ 27, 28

related to faunal migration .................. 25

related to geologic changes .................... 25

Coral fauna, Ludlovian .................................. 19

Onondaga Limestone .............................. 25

Rabbit Hill ................................................ 19

Coral Gulch ........................................................ 8

Coral Ridge ........................................................ 13

Coral subzones .................................... 19, 20, 30, 31

Coral succession, related to

faunal migration ...................... 25

related to geologic change ...................... 25

79
80 INDEX

Page 

Coral zonation, Devonian ................................ 22

Coral zone A ........ 8, 11, 17, 18. 19, 23, 28, 29, 31

B .......... 8, 11, 17, 18, 19, 20, 23. 25, 28, 29, 32

C ................ 8, 11, 13, 17, 18, 19, 20, 23, 24, 25,

28, 29, 32, 42, 44, 47, 53 

D .............. 8, 11, 13, 17, 18, 19, 20, 21, 22, 23,

24, 25, 28, SO, 32, 42, 43, 44, 45 

E ................ 3, 17, 18, 19, 21, 25, 28, 29, 33, 45

F ................ 5, 9, 13, 15, 16, 17, 18, 19, 20, 21,

22, 23, 25,28, 31 

correlation .......................................... 31

G ...................... 9, 15, 16, 17, 18, 19, 21, 22, 28

H ...................................... 17, 18, 19, 22, 25, 28

I .................................. 17, 18, 19, 22, 23, 25, 28

Corals .............................................................. 5, 9, 23

Carboniferous ............................................ 38

colonial .................................. 21, 29, 31, 37, 38

dissepiments .............................................. 36

favositid ........................................ 16, 20, 24, 25

horn ............................................................ 8, 23

internal structures .................................. 35

Paleozoic .................................................... 37

pycnostylid ................................................ 11

rugose ............ 9, 13, 15, 16, 17, 19, 21, 22, 25

septa .................................................... 55, 37, 38

septal carinae ........................................ 36, 37

septal stereozone ...................................... 37

Silurian ........................................................ 37

skeletal features ........................................ 34

solitary .................... 21, 24, 25, 35, 36, 37, 38

stereoplasmic deposits ............................ 37

tabulae ...................................................... 36, 37

tabulate .............. 18, 19, 20, 23, 24, 25, 29, 36

Cordilleran Belt .................... 25, 29, 30, 31, 37, 56

corniculum, Streptelasma .............................. 40

Correlation, coral subzone Da ........................ 31

coral zones, Cordilleran Belt .................. 31

distant regions .................................. 29

Cortez Mountains .................................... 31

Desert Range ............................................ 32

Funeral Mountains .................................. 32

Hot Creek Range ...................................... 32

Hume Formation ...................................... 31

Nahanni Formation ................................ 31

Nevada Formation .................................. 31

unit 1 .................................................. 32

unit 2 .................................................. 32

north-central Great Basin ...................... 31

northern Inyo Mountains ...................... 32

northern Panamint Mountains ............ 32

northern Sulphur Spring Range .......... 31

Oriskany ...................................................... 31

Rabbit Hill Limestone ............................ 31

Ranger Mountains .................................. 32

south-central Great Basin ...................... 32

southern Great Basin .............................. 32

Toiyabe Range .......................................... 31

Tuscarora Mountains, southern ............ 31

west-central Great Basin ...................... 31

Cortez Mountains .......................................... 31, 40

Cosmophyllum ............................................ 68, 69, 70

dachsbergi ................................................ 30, 70

Costispirifer .................................. 13, 19, 20, 23, 29

arenosus .................................. 8, 11, 17, 19, 20,

23, 29, 31, 32, 33, 49 

Crinoids .................................................. 8, 21, 23, 24

curviseptatum, Grypophyllum ...................... 53

Cyathaxonia siluriensis .................................. 39

Cyathophylloides .............................................. 45

rhenanum .................................................. 45

(Cyathophylloides) disjuncta, Columnaria.. 45 

cyathophylloides, Australophyttum .............. 57

Spongophyllum ........................................ 57

Cyathophyllum .......................................... 21, 25, 29

caespitosum ................................................ 58

hexagonum ................................................ 60

lonense ...................................................... 53, 54

(Moravophyllum) .................................... 21

Page 

Cyathophyllum Continued

robustum .................................................... 53

sp .................................................................. 54

cycloptera, Howellella .................................... 19, 29

Cylindrophyllum .............................................. 58, 59

Cyrtina ................................................................ 20

Cyrtospirifer beds, Devils Gate .................... 19, 22

Cyrtospirifer zone ............................................ 22

Cystiphyllidae .................................................... 65

Cystiphylloidae ............................ 29, 33, 34, 35, 64

Cystiphylloides ............ 6, 21, 24, 25, 33, 34, 65, 68

aggregatum .............................................. 64, 65

americanum .............................................. 66

lonense ................................................ 20, 34, 66

robertsense ........................................ 21, 34, 65

sp. d ............................................................ 34, 66

(Cystiphylloides), Mesophyllum .................. 65

Cystiphyllum ...................................................... 70

Czechoslovakia ................................................ 40, 50

D

dachsbergi, Arcophyllum ................................ 70

Cosmophyllum .......................................... 30, 70

Mesophyllum (Arcophyllum) ................ 70

Dalmanites ........................................................ 20

meeki .................................................. 21, 32, 52

defectum, Mesophyllum .................................. 67

Dendrostella ...................................................... 33, 45

disjuncta .................................................... 46

praerhenana .............................................. 46

rhenana ...................................................... 46

romanensis ........................................ 21, 33, 45

(Dendrostella) praerhenana, Favistella .... 45

rhenana, Favistella .................................. 45

Descriptive paleontology ................................ S3

Desert Range .................................................. 32, 43

Devils Gate ............................................ 9, 18, 19, 25

Devils Gate Limestone ........................ 6, 9, 15, 17,

21,22,25,28,29, 60 

lithology ...................................................... 9, 17

type area .................................................... 22

type section .............................................. 19

Devils Gate Pass .............................................. 22

Dialytophyllum .................................................. 67

Diamond Mountains .................... 5, 6, 8, 9, 13, 15,

18, 19, 21, 22, 23, 27 

Diamond Mountains facies belt ............ 1, 6, 8, 9,

13, 15, 17, 29 

Diamond Valley ................................................ 15

Digonophyllidae .......................... 17, 24, 29, 30, 33,

34, 35, 36, 38, 65, 67, 71 

Digonophyllinae ........................................ 34, 67, 68

Digonophyttum .................................... 21, 25, 65, 69

(Digonophyllum) ...................................... 67

(Mochlophyllum) .................................... 21, 67

(Digonophyllum), Digonophyllum .............. 67

Dimorphism .................................................... 35, 38

Disconformity ............................................ 8, 19, 27

disjuncta, Columnaria (Cyathophylloides).. 45 

Dendrostella .............................................. 46

Disphyllidae .............. 17, 29, 33, 34, 38, 58, 60, 61

Disphyttum .................................. 21, 22, 24, 34, 58

caespitosum ................................................ 58

eurekaensis .................................. 21, 34, 59, 66

goldfu&si ...................................................... 59

nevadense .................................... 34, 59, 60, 66

nevadensis .................................................. 21

Dissepimentarium ............................................ 36

Dissepiments ...................................................... 36

Diversophyllum ................................................ 55, 56

Dobbin Summit ........................................ 17, 19, 40

Dolomite .................. 8, 9, 13, 15, 19, 20, 27, 28, 32

diagenetic .................................................. 3, 23

recrystallization ............................ 23

saccharoidal ...................................... 16, 22, 29

Douglas Creek, Australia .............................. 57

duplicatum, Zonophyllum ..................... 30, 70, 71

Page

E

Early Devonian coral zones ............................ 19

correlation .................................................. 29

Early middle Devonian coral zones ............ 19

correlation ...................................-. ......... 29

East Ridge .......................................................... 11

Eifel district, Germany ........ 30, 58, 60, 67, 69, 70

Eightmile Well .................................................. 15

elegantulum, Ketophyttum .............................. 50

Papiliophyllum .. 20, 24, 32, 33, 46, 47, 50, 66 

Encrinite .......................................................... 13, 15

Endophyllidae .......... 28, 33, 34, 37, 56, 58, 60, 61

Endophyllum .................................................... 57, 58

abditum ...................................................... 56

bowerbanki ................................................ 56

Entelophylloides ................................................ 60

Entelophyllum ................................................ 28, 35

Erben, H. K., cited .......................................... 30

Eridophyllum .................................................... 59

Eureka, Nev ...................................................... 27

Eureka County ........ 23, 43, 44, 46, 47, 50, 51, 52,

54, 55, 56, 59, 62, 63, 65, 68 

Eureka mining district.

Nevada .... 4, 5, 29, 32, 42, 43, 56, 71

Eureka Quartzite .............................................. 15

eurekaensis, Disphyttum .................. 21, 34, 59, 66

Eurekaphyttum .................................. 33, 47, 50, 52

breviseptatum .......................................... 33, 52

Eurekaspirifer pinyonensis ........................ 18, 21

expansum, Spongophyttum ............................ 59

Fauna, brachiopod ........................................ 31, 32

coral ............................................................ 32

coral zone A ................................ 19, 20, 28, 40

B .................. 20, 28, 30, 31, 46, 47, 49, 50

C .................. 20, 30, 31, 46, 50, 52, 54, 66

D .............................. 20, 21, 27, 31, 32, 41,

46, 56, 61, 62, 64, 66, 68 

E ............................................ 21, 57, 64, 66

F .......................................... 21, 27, 30, 32,

56, 58, 60, 61, 62, 64, 67 

G ............................................................ 21

H .......................................................... 22

I ...................................... 22, 58, 59, 60, 61

Funeral Mountains .................................. 33

Onondaga .................................................... 30

Rabbit Hill .................................. 29, 31, 50, 58

subzone Di .... 20, 21, 30, 47, 52, 54, 55, 56, 71

subzone D2 ........ 21, 30, 46, 54, 55, 56, 58, 59,

60, 61, 62, 63, 64, 66, 67, 70, 71 

subzone D3 ...................... 21, 30, 55, 62, 67, 70

Trematospira ............................................ 29

trilobite ...................................................... 31

Faunal migration, related to

coral evolution .......................... 25

Favistella .......................................................... 33, 45

(Dendrostella) rhenana .......................... 45

praerhenana ...................................... 45

Favistina ............................................................ 45

Favosites .............................................. 19, 23, 24, 25

Favositid heads ................................................ 8, 24

Fenstermaker Mountain ................................ 45

Fish Creek Range .......... 5, 6, 9, 15, 43, 45, 54, 71

flexum, Hexagonaria ........................................ 62

Mesophyllum .............................................. 66

foerstei, Syringaxon .......................... 19, 31, 33, 39

Foraminifera .................................................... 34

Fossil Gulch ...................................................... 8

Fossils, silicifled .................................. 23, 24, 25, 34

Frasnes, Belgium .............................................. 60

Frazier Creek .................................................... 65

Frenchman Flat County, Nev ...................... 32

Funeral Mountains .................................. 32, 43, 52

Fusulinids .......................................................... 34
INDEX 81

Page

G

Garden Valley Formation .............................. 15

Garden Valley quadrangle ............................ 5

Gastropods ........................................................ 15, 24

Gazelle Formation ............................................ 60

Genomorph concept ........................................ 38

Geologic change, related to

coral evolution .......................... 25

Geologic structure, Lone Mountain .............. 5

Germany .................................................... 58, 67, 70

Gerolstein, Eifel district .............................. 30, 70

gigantea, Caryophyttia .................................... 41

Siphonophrentis ........................................ 41

giganteum, Grypophyllum .............................. 53

Gold Hill mining district, Utah .................... 27

goldfussi, Cladocora ........................................ 58

Disphyllum .................................................. 59

Goniatites .................................................. 24, 30, 31

Goniophyllidae .................................................. 28

Gotland ........................................................ 44, 45, 50

Gotland reef complex ...................................... 22

Grande Greve Formation ............................ 30, 64

Graptolite beds ...................................... 5, 17, 19, 32

Grays Canyon .............................. 15, 43, 54, 56, 71

Great Basin, central ................................ 39, 43, 71

coral reference sections .......................... 6

correlation ................................................ 31, 32

Devonian Facies belts .............................. 6

faunas ...................... 24, 53, 56, 57, 60, 67, 71

north-central .............................................. 74

Rugosa, classification .............................. 33

south-central .............................................. 72

southern .................................................... 43, 71

west-central .............................................. 74

Great Basin province .... 23, 25, 27, 29, 45, 62, 67

Green Springs quadrangle ............................ 22

Greifensteiner Kalk, Germany ...................... 40

Grypophyllum curviseptatum ........................ 53

giganteum .................................................. 53

nevadense .................................................... 53

Gypidula ............................................................ 29, 50

coeymanensis .................................... 20, 29, 50

loweryi ................................................ 20, 21, 52

haguei, Zonophyllum ........................ 21, 30, 34, 71

Hallia .................................................................. 46

insignia ........................................................ 46

Halliidae .............. 17, 20, 24, 29, 30, 33, 37, 46, 53

Halliinae ............................................................ 33, 46

Halysites ............................................................ 11

Halysitids ............................................................ 28

Hamilton Formation ........................................ 30

Helderbergian fauna .................................. 8, 9, 11

helderbergium, Pseudoblothrophyllum ........ 44

Heliolites .......................................................... 25, 70

Heliophyttum ............................................ 35, 53, 63

venatum ...................................................... 41

Hemicosmophyllum .................................. 68, 69, 70

(Hemicosmophyllum), Mesophyllum .......... 68

Heterophrentis .................................. 30, 41, 42, 43

nevadensis ................................................ 42, 43

hexagona, Hexagonaria ................................ 60, 61

Hexagonaria ...................................... 21, 25, 30, 32,

34, 37, 38, 56, 58, 60, 63 

flexum .......................................................... 62

hexagona .................................................. 60, 61

(Hexagonaria) .......................................... 61

kirki ............................................................ 62

(Pinyonastraea) .................................... 32, 58

kirki ...................................... 21, 34, 38, 62

(Hexagonaria), Hexagonaria ...................... 61

hexagonum, Cyathophyllum .......................... 60

Hidden Valley .................................................... 33

Hidden Valley Dolomite ................................ 32, 39

unit 3b ........................................................ 32

Hill, Dorothy, cited ........ 33, 34, 38, 67, 68, 69, 71

Page 

History of investigation .................................. 4

Homalophyllum ................................................ 41

Homeomorphie genera .................................. 34, 70

Hot Creek Canyon ............................................ 32

Hot Creek Range ...................................... 6, 32, 52

House, M. R., cited ........................................ 29, 30

Howelletta cycloptera .................................... 19, 29

Hume Formation .............................................. 31

I, J

Independence Shale, Iowa .............................. 39

insignis, Hallia .................................................. 46

Introduction ...................................................... 1

invaginatus, Amplexus .................................... 42

Breviphrentis ................ 30, 42, 43, 44, 46, 56

Siphonophrentis (Breviphrentis) ...... 21, 30

32, 33, 38, 42, 45, 54, 56 

Inyo Mountains ................................................ 33

Joana Limestone .............................................. 28

K

Keriophyllum .................................................... 21

ketophylloides, Kobeha .................... 20, 33, 48, 49

Ketophyllum .................................................... 50, 55

elegantulum ................................................ 50

Kirk, Edwin, cited .......................................... 4

kirki, Arcophyllum .......................................... 30

Hexagonaria .............................................. 62

(Pinyonastrea) .................. 21, 34, 38, 62

Mesophyllmdes .................................. 68, 69, 70

Mesophyllum (Arcophyllum) .. 21, 30, 34, 69 

Prismatophyllum .................................... 61, 62

Klamath Mountains ...................................... 57, 60

Kobeha .............................. 8, 13, 17, 20, 23, 24, 29,

31, 32, 33, 35, 36, 38, 47, 49, 50, 52, 53 

fauna .......................................................... 8, 20

ketophylloides .............................. 20, 33, 48, 49

walcotti ........ 19, 20, 23, 29, 31, 33, 48, 49, 52

zone .............................................................. 48

Kobeha Valley project .................................... 5

kobehana, Acrospirifer .. 13, 20, 24, 32, 33, 52, 66 

kobehensis, Breviphrentis .............................. 29

Siphonophrentis (Breviphrentis) ...... 20,

30, 33, 42, 43, 66 

Kodonophyllidae ........................................ 28, 33, 44

Kodonophyllinae ............................................ 33, 45

Kodonophyllum .................................. 33, 37, 43, 45

milne-edwardsi .......................................... 44

sp. a ............................................................ 33

sp. f .............................................................. 45

Kozlowskiellina ................................................ 19

Kwangsi Province, China ............................ 56, 65

Kyphophyllidae .................................................. 60

Laccophyllidae .............................. 17, 27, 33, 37, 38

Laccophyllum .................................................... 39

acuminatum .............................................. 39

Lake Manitoba .................................................. 46

Lake Winnipegosis .......................................... 46

Laketown Dolomite .......................................... 27

Lander County, Nev ........................................ 58

landerensis, Australophyllum .... 20, 23, 33, 34, 57

Late Devonian coral zones ............................ 21

Late middle Devonian coral zones ................ 21

Leiorhynchus castanea .................................. 13, 16

Lekanophyllum .................................................. 67

(Lekanophyllum), Mesophyllum .................. 21

lenticularis, Pleurodictyum ............................ 31

Leonaspis .................................................... 19, 23, 29

sp .................................................................. 46

Leptaena ........................................................... 20

Leptocoelia ...................................................... 19, 31

sp .................................................................. 50

Leptostrophia sp .............................................. 20

Levenea subcarinata ................................ 19, 29, 31

Page 

limbatum, Arcophyllum .................................. 70

Mesophyllum (Arcophyllum) ................ 70

Limestone ................ 8, 9, 15, 16, 17, 19, 22, 27, 32

argillaceous ............................ 3, 13, 22, 24, 28

calcarenite .................................................. 23

dolomitic .......................... 16, 19, 22, 23, 25, 29

stromatoporoid .......................................... 29

Lithology, Bay State Dolomite Member ...... 16

Beacon Peak Dolomite Member ............ 15

Devils Gate Limestone ............................ 9, 17

Nevada Formation, unit 1 ...................... 8

unit 2 .................................................. 8

unit 3 .................................................. 8

unit 4 .................................................. 9

unit 5 .................................................. 9

Oxyoke Canyon Sandstone Member .... 15

Rabbit Hill Limestone ............................ 17

Sentinel Mountain Dolomite Member.. 15 

Woodpecker Limestone Member .......... 16

Locality register .............................................. 71

Lone Mountain .................... 5, 8, 9, 13, 15, 16, 18,

20, 22, 23, 24, 25, 30, 39, 43, 44, 46, 47,

50, 51, 52, 54, 55, 56, 59, 60, 61, 62, 63,

64, 65, 66, 69, 70

reference section .... 6, 8, 16, 17, 19, 28, 30, 32

Lone Mountain Dolomite ............ 8, 15, 19, 27, 28

Lone Mountain Dolomite-Nevada

Formation boundary ................ 28

lonense, Bethanyphyllum ................ 21, 33, 54, 55

Cyathophyllum ........................................ 53, 54

Cystiphylloides .................................. 20, 34, 66

Mesophyllum .............................................. 66

lonensis, Amplexus ........................................ 42, 43

Breviphyllum ............................................ 42

Louisville, Kentucky ........................................ 47

Lower-Middle Devonian boundary ................ 31

loweryi, Gypidula ...................................... 20, 21, 52

Lycophyllidae .................................................. 37, 46

M

McAllister, J. F., cited .................................... 32

McColley Canyon ............................................ 31, 49

MacGeea ...................................................... 22, 25, 35

macrostriata, Chonetes .................... 20, 21, 32, 52

Maggie Creek .................................................. 20, 31

magnus, Amplexus .......................................... 56

Mahogany Hills .............................. 8, 15, 19, 27, 43

Mapping procedures ........................................ 5

Marble Canyon quadrangle, Calif ................ 32

Martinia kirki zone .......................................... 21

undifera ...................................................... 55

masoni, Nevadaphyllum ................................ 33, 44

Mazourka Canyon ............................................. 33

meeki, Dalmanites .................................... 21, 32, 52

OdontophyUum .............. 20, 29, 30, 33, 46, 47

Meristella .......................................................... 21, 33

robertsensis .............................................. 20, 33

Merriam, C. W., cited .................................... 22

MesophyUoides kirki ................................ 68, 69, 70

Mesophyllum .............. 21, 25, 30, 33, 34, 65, 67, 68

defectum .................................................... 67

flexum .......................................................... 66

lonense ........................................................ 66

robertsense ................................................ 65

vesiculosum ................................................ 66

(Arcophyllum) ........................................ 67

dachsbergi .......................................... 70

kirki ...................................... 21, 30, 34, 69

limbatum ............................................ 70

(Cystiphylloides) ...................................... 65

(Hemicosmophyllum) .............................. 68

(Lekanophyllum) .................................... 21

(Mesophyllum) ...................................... 67, 68

sp. b ............................................ 21, 34, 68

sp. c .................................................... 34, 68

(Mesophyllum), Mesophyllum .................... 67,68

Methods of investigation ................................ 5
82 INDEX

Page 

Middle middle Devonian coral zones ............ 21

Miller, A. K., cited .......................................... 30

milne-edwardsi, Kodonophyllum .................. 44

Mineral Hill district ........................................ 31

(Mochlophyllum), Digonophyllum .............. 21, 67

Modoc Peak ...................................................... 8, 15

Moenke, Maria, cited ...................................... 61

Mollusca ...................................................... 23, 24, 29

MoOusks .............................................................. 5

Monitor Range .............................. 6, 17, 19, 39, 40

Monitor-Simpson Park facies belt.... 3, 6, 17, 29

Monte Cristo .............................................. 19, 22, 25

Moravophyttum ................................................ 53

(Moravophyllum), Cyathophyttum .............. 21

Morey Peak ........................................................ 32

Mount Tenabo ................................................ 31, 40

MucophyUum .................................................... 35

Mulligan Gap .................................................... 20

Mulligan Gap fault .......................................... 15

Mutation ............................................................ 38

Mycophyllinae .................................................. 28, 45

N

Nahanni Formation ........................................ 31

Nautiloids .......................................................... 24

Nevada Formation ........................ 6, 8, 32, 40, 47,

52, 54, 56, 59, 63, 65, 68, 71 

barren zone ........................ 3, 8, 13, 15, 21, 25

Beacon Peak Dolomite Member .......... 6, 40

type section ................................................ 8

unit 1 ...................... 8, 9, 11, 15, 17, 19, 20, 23,

24, 25, 28, 31, 44, 46, 48, 49,

50, 51, 52, 53, 66

lithology .............................................. 8

unit 2 ...................... 6, 8, IS, 15, 17, 19, 20, 21,

22, 24, 27, 28, 29, 30, 31, 39, 42, 43,

44, 45, 46, 52, 54, 55, 59, 60, 61,

62, 63, 64, 66, 68, 69, 70, 71

lithology .............................................. 8

unit 3 .................... 8, 13, 15, 19, 21, 25, 27, 29

lithology .............................................. 8

unit 4 ...................................... 8, 13, 15, 16, 19,

20, 21, 22, 25, 28, 29, 31, 64 

lithology .............................................. 9

unit 5 ........................ 8, 9, 13, 15,16, 19, 21, 22

lithology .............................................. 9

nevadana, Atrypa ............................................ 21

nevadaensis, Schizophoria .............................. 21

Schuchertella .................................................. 20, 21

Nevadaphyllum ............................ 33, 37, 38, 41, 44

masoni ...................................................... 33, 44

nevadense, Disphyllum ...................... 34, 59, 60, 66

Grypophyttum ............................................ 53

Spongophyllum .......................................... 59

Tabulophyllum ........................................ 53, 54

nevadensis, Agoniatites .................................. 30

Amplexus .................................................. 42, 56

Billingsastraea .. 21, 30, 31, 34, 38, 61, 62, 63 

Billingsastraea billingsi .......................... 63

Disphyllum ................................................ 21

Heterophrentis ........................................ 42, 43

arachne, Billingsastraea, .... 34, 38, 62, 63, 64

New York ........................................................ 29, 53

Newark Mountain ............................................ 9

Niagaran reef complex .................................. 22

Nicholsonia ........................................................ 39

(Nieholsonia) Alleynia .................................. 39

Nohner Schichten .................................... 30, 46, 70

North Sand Peak .............................................. 13

Nye County, Nev .............................................. 32

O

Odontophyllum ............................ 24, 30, 33, 46, 47

convergens .................................................. 47

meeki ................................ 20, 29, 30, 33, 46, 47

patellatum .................................................. 47

Ohio .................................................................... 46

Oklahoma ............................................................ 29

Page 

Old Whalen mine area .................................. 29, 31

Oliver, W. A., Jr., cited .................. 23, 24, 25, 38

quoted .......................................................... 62

Onondaga Limestone, New York .. 24, 29, 30, 41

Hagersville, Ontario ................................ 65

Orbitoids ............................................................ 34

Oriskany Peak .................................................. 13

Orthoceras .......................................................... 24

Orthoquartzite .................................................. 15

Orthostrophia strophomenoides .................... 29

Owens Valley, Calif ........................................ 57

Oxyoke Canyon ................................................ 21

Oxyoke Canyon Sandstone Member.. 8, 13, 15, 32

lithology ...................................................... 15

Pachyphyllum .............................. 22, 25, 29, 35, 61

Paflrath Basin, Germany .............................. 45

Paleophyllum ...................................................... 45

Panamint Mountains .................. 32, 33, 39, 43, 52

Papiliophyllinae ................................ 27, 33, 37, 47

Papiliophyllum 13, 17, 20, 24, 29, 32, 33, 47, 50, 53

elegantulum ........ 20, 24, 32, 33, 46, 47, 50, 66

subsp. d .................... 20, 30, 32, 33, 52, 55

fauna ............................................................ 8

patellatum, Odontophyllum ............................ 47

PeneckieUa ........................................................ 58

Permia ................................................................ 39

perplexum, Barrendeophyllum ...................... 39

Perry County, Tenn ........................................ 39

Petes Canyon .................................................... 58

Phacellophyttum .................................. 22, 25, 35, 58

Phacops ............................................................ 19, 23

rana .............................................................. 21

Phillipsastraea .......................................... 22, 25, 35

(Billingsastraea) ...................................... 62

biofacies ...................................................... 19

verneuili .................................................... 62, 63

Phillipsastraeidae .............................. 22, 25, 29, 33,

34, 35, 37, 58, 60, 61

Phillipsburg mine ............................................ 15

Pholidostrophia ................................................ 50

sp .................................................................. 20

phrygia, Zaphrentis ........................................ 41

Pilot Shale ........................................................ 17, 22

lower ............................................................ 19

upper .......................................................... 6

Pinto Summit quadrangle .............................. 21

Pinyonastraea ........................ 24, 30, 34, 35, 60, 61

(Pinyonastraea), Hexagonaria ................ 32, 58

kirki, Hexagonaria .................. 21, 34, ,38, 62

pinyonensis, Eurekaspirifer ........................ 18, 21

Spirifer ...................................................... 46, 52

planotabulatum, Sinospongophyllum ........ 55, 56

Plasmophyllum ................................................ 65, 68

Plethorhyncha .................................................. 19

Pleurodictyum .................................... 19, 23, 29, 31

lenticularis .................................................. 31

Polycoeliidae ...................................................... 39

Polyphyletic ...................................................... 35, 61

praerhenana, Dendrostetta ............................ 46

Favistetta (Dendrostetta) ...................... 45

Prince of Wales mine .................................. 11, 20

Prismatophyllum ............................................ 60, 61

kirki .......................................................... 61, 62

Proetus ................................................................ 19

Protocorallites .................................................. 38

Pseudoblothrophyllum ............................ 43, 44, 45

helderbergium ............................................ 44

Pseudocosmophyllum ...................................... 68, 69

punctulifera, Strophonetta .............................. 20

Purpose and scope of investigation ............ S

pustulosa, Strophonetta .................................. 20

Pycnostylidae .................................................. 28, 35

Pyramid Peak .................................................... 32

Q, R

Quartz grains

Page 

Quartzite .......................................................... 32, 55

Queensland, Australia ...................................... 57

Rabbit Hill ........................................................ 19, 40

Rabbit Hill Limestone .............. 5, 6, 8, 15, 17, 19,

23, 28, 29, 33, 39, 40, 57, 58 

lithology ...................................................... 17

type section .............................................. 17, 19

Radiastraea ................................................ 38, 63, 64

arachne ...................................................... 63, 64

rana, Phacops .................................................... 21

Ranger Mountains ............................ 32, 43, 52, 71

rectum, Tabulophyllum .................................. 56

Reeds Canyon ...................................... 17, 31, 62, 64

Reef Point .......................................................... 20

Reefs, patch .................................... 9, 22, 23, 24, 25

Reference section, coral zone C .................... 20

coral zone D .............................................. 20

coral zone F .............................................. 21

coral zone G .............................................. 21

coral zone I ................................................ 22

Lone Mountain ........................................ 6, 8

Sulphur Spring Range ............................ 6

Reference sections ............................................ 5

References selected .......................................... 74

Rensselandia ...................................................... 9, 21

rhenana, Dendrostetta ...................................... 46

Favistetta (Dendrostetta) ...................... 45

rhenanum, Cyathophylloides .......................... 45

Rhine Valley ...................................................... 46

Roberts Creek Mountain ................................ 29

Roberts Creek Ranch ...................................... 20

Roberts Mountains ............................ 6, 25, 29, 30,

49, 50, 52, 54, 62, 64, 65 

Roberts Mountains Formation ............ 19, 28, 31

robertsense, Cystiphylloides .................. 21, 34, 65

Mesophyttum .............................................. 65

robertsensis, Meristetta ................................ 20, 33

robustum, Bethanyphyllum .......................... 53, 54

Cyathophyttum .......................................... 53

romanensis, Dendrostetta ........................ 21, 33, 45

Romano Ranch .................................................. 46

Rozkowska, Maria, cited .............................. 24, 38

Rugosa ............................ 1, 8, 18, 19, 20, 22, 25, SS

calice features .......................................... 35

Carboniferous ............................................ 38

classification ............................................ SS, 35

Coeymans Limestone .............................. 23

colonial .... 20, 21, 23, 24, 25, 31, 35, 36, 37, 38

Cordilleran ................................................ 3, 4

descriptive terms ...................................... $8

digonophyllid ............................................ 69

dissepimented ............................................ 36

Early Devonian ........................................ 37

Early Silurian ............................................ 37

eastern North America .......................... 3

Eureka mining district .......................... 4

evolution .................................................... 25

Devonian ............................................ 28

Late Silurian .................................... 2S

zone D ................................................ 2S

zone F .................................................. 29

zone I .................................................. 29

exterior corallum features .................... 34, 37

fine structure .......................................... 34, 35

Great Basin .......................................... 3, 4, 34

growth stages ............................................ 37

interior corallum structures ................ 35

Nevada Formation .................................. 3

Ordovician .................................................. 37

pycnostylid ................................................ 28

Rabbit Hill Limestone ............................ 3

reproductive features .............................. S7

Rhine Valley .............................................. 4

solitary ...................... 21, 24, 25, 35, 36, 37, 38

symmetry .................................................... 37

taxonomy ...................................... 34, 36, S7, 38

western North America .......................... 3

Ryan quadrangle, Calif .................................. 32
INDEX 83

Page 

S

Sandstone, quartzitic ................................ 8, 11, 15

siliceous ...................................................... 8, 13

Schigophoria nevadaensis .............................. 21

Schuchertella ...................................................... 21

nevadaensis .............................................. 20, 21

Section Ridge .................................................... 9

sedgwicki, Spongophyttum ............................ 57

Sellersburg Limestone .................................... 47

Sentinel Mountain ............................................ 21

Sentinel Mountain Dolomite

Member .......................... 8, 13, 15, 23

lithology ...................................................... 15

Septa .......................................................... 35, 37, 38

Septal stereozone .............................................. 37

Sevy Dolomite .................................................... 27

Silurian-Devonian boundary ........................ 27, 28

siluriensis, Cyathaxonia .................................. 39

Syringaxon ........................................ 38, 39, 40

Simpson Park Mountains .................. 6, 28, 33, 40

Simpson Park Range ...................................... 6, 17

Simpson Peak Range ...................................... 19

Sinospongophyllum .............. 32, 33, 38, 41, 44, 55

planotabulatum ...................................... 55, 56

sp. d .................................................... 33, 52, 56

sp. e .......................................................... 33, 56

sp. f .................................................... 34, 44, 56

spp ................................................................ 21

Siphonophrentinae .. 17, 27, 29, 30, 33, 36, 37, 45 

Siphonophrentis ........................................ 33, 41, 45

gigantea ...................................................... 41

variabilis ...................................... 30, 42, 43, 44

(Breviphrentis) ...................................... 32, 41

invaginatua ........................ 21, 30, 32, 33,

38, 42, 45, 54, 56 

kobehensis .............. 20, 30, 33, 42, 43, 66

i Siphonophrentis) .................................. 41

(Siphonophrentis), Siphonophrentis .......... 41

Sociophyllum ............................................ 21, 25, 59

Southern Sulphur Spring Range,

reference section .................... 6, 8, 9

spatiosa, Zaphrentis ........................................ 41

Spirifer kobehana zone .................................. 30

pinyonensis .............................................. 46, 52

zone .................... 18, 20, 21, 29, 30, 55, 70

Spirifers ............................................................ 32, 33

Spongophyllidae ................................ 33, 37, 58, 60

SpongophyUum ................................................ 57

cyathophylloides ........................................ 57

expansum .................................................... 59

nevadense .................................................. 59

sedgwicki .................................................... 57

Stathmoelasma ................................................ 53, 54

Stauriidae .................................................. 33, 34, 45

Stauria ................................................................ 45

astreiformis ................................................ 45

Page 

Stereoplasmic deposits .................................... 37

Stevens County, Wash .................................... 31

Stratigraphy, Central Great Basin

Devonian Rocks ........................ 6

Streptelasma corniculum ................................ 40

Streptelasmatidae .......................................... 33, 40

Streptelasmidae .............................................. 33, 40

Striatopora ...................................................... 19, 23

Stringocephalus ...................... 9, 15, 16, 21, 46, 70

zone .............................................................. 22

Stromatoporoid-AmpTiipom biofacies ........ 19

Stromatoporoids ................ 9, 15, 17, 22, 23, 24, 25

strophomenoides, Orthostrophia .................. 29

Strophonetta ...................................................... 32

punctulifera .............................................. 20

pustulosa .................................................... 20

Stumm, E. C., cited .......... 4, 22, 33, 51, 57, 66, 69

subcarinata, Levenea .............................. 19, 29, 31

Sulphur Spring Range .................... 6, 8, 9, 11, 13,

15, 17, 18, 19, 20, 23, 24, 29, 31,

40, 43, 45, 46, 48, 49, 50, 52, 54, 67, 68

SynaptophyUum ................................................ 45

Syringaxon .............................. 5, 17, 19, 20, 23, 27,

28, 29, 31, 33, 35, 36, 37, 39, 58 

acuminatum ................................................ 39

biofacies ...................................................... 19

foerstei .......................................... 19, 31, 33, 39

siluriensis .......................................... 38, 39, 40

sp .................................................................. 31

Systematic paleontology ................................ 33

Tabulae .............................................................. 36, 37

Tabularium ...................................................... 36, 37

Tabulata .............................................................. 1, 29

Tabulophyttum .......................................... 21, 55, 56

nevadense .................................................. 53, 54

rectum .......................................................... 56

Taimyrophyllum .............. 21, 25, 29, 32, 58, 61, 64

Taxonomy, Rugosa ............................ 34, 36, 37, 38

Rugosa, interior corallum ...................... 35

Teicherticeras .................................................. 30, 31

Temnophyllum .................................................. 22

Tennessee ............................................................ 29

Tentaculites ...................................................... 24, 31

Terrace Butte .................................................. 13, 15

Thrust plates .................................................... 5

Toiyabe Range .................................... 17, 31, 62, 64

Toquima Range .......................... 17, 20, 23, 57, 58

Torquay, England ............................................ 56

Tortophyllum ...................................................... 53

Trematospira cooperi ...................................... 29

fauna ............................................................ 29

Trilobites .................................. 5, 19, 23, 24, 29, 30

Tryplasmatidae .............................................. 34, 35

Turbidity ............................................................ 24

Page 

Tuscarora Mountains .............................. 20, 23, 31

Twin Hills .......................................................... 11

typus, Arcophyllum .......................................... 68

U, V

Unconformity .................................................... 27

undifera, Martinia ............................................ 55

Ural Mountains, Russia ................................ 68, 70

Utaratuia .......................................................... 21, 25

variabilis, Siphonophrentis .............. 30, 42, 43, 44

Variation ............................................................ 38

statistical analysis .................................... 34

Vaughn Gulch Limestone ............................ 33, 57

venatum, Heliophyllum .................................. 41

verneuili, Anastrophia .................................... 29

Phittipsastraea ........................................ 62, 63

verritti, Billingsastraea .................................. 31, 64

vesiculosum, Mesophyllum .............................. 66

W, X

walcotti, Kobeha ...................................... 19, 20, 23,

29, 31, 33, 48, 49, 52 

Wales Peak ...................................................... 11, 13

Walti Hot Springs area .................................. 40

Wedekind, Rudolf, cited ........ 4, 30, 38, 67, 69, 71

Well, J. W., cited .............................................. 35

West Peak .......................................................... 8, 20

West Sand Peak ................................................ 13

Whistler Mountain quadrangle .................... 5, 22

White Pine mining district,

Nevada ............................ 4, 19, 22, 25

Wisconsin ............................................................ 62

Woodpecker Limestone Member ................ 9, IS,

15, 16, 17, 19, 21 

lithology ...................................................... 16

Xystriphyllum .................................................. 58, 61

Y, Z

Yahoo Canyon .................................................. 22

Yassia .................................................................. 57

Yoh, S. S., cited ................................................ 56

Zaphrentidae ...................................... 33, 35, 41, 53

Zaphrentis ........................................................ 33, 41

phrygia ........................................................ 41

spatiosa ...................................................... 41

Zaphrentoid sp .................................................. 48

Zoantharia .......................................................... 1

Zonophyllinae ............................................ 34, 67, 70

Zonophyllum .............................................. 30, 34, 70

duplicatum ........................................ 30, 70, 71

haguei .......................................... 21, 30, 34, 71

sp. a .......................................................... 34, 71

sp. b .......................................................... 34, 71

sp .................................................................. 32
PLATES 1-25

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

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

FIGURES 1-8. Syringaxon foerstei n. sp.

1. Oblique calice view of holotype ( x ll/2 ), USNM 159243.

2. Lateral view of paratype ( x 2), USNM 159244.

3. Calice view of paratype ( x 1), USNM 159245.

4. Calice view ( x 2).

5. Early ephebic transverse thin section of paratype (x 6), USNM 

159246.

6. Longitudinal thin section of paratype ( x 6), USNM 159247. 

7,8. Mature transverse thin sections of two individuals (x 2, x 4). 

Early Devonian, Rabbit Hill Limestone; Devonian coral zone A. Locality 

M187, Rabbit Hill, Copenhagen Canyon, Eureka County, Nev. 

9-12. Syringaxon foerstei n. sp.

9,10. Mature transverse thin section and longitudinal thin section

(X 31/3), USNM 159352.

11,12. Calice views of two silicified individuals ( x 2). 

Early Devonian; Devonian coral zone A. Locality M197, south of Bailey

Pass, Sulphur Spring Range, Nev. 

13,14. Syringaxon foerstei n. sp.

Mature transverse thin section and longitudinal thin section of paratype

(x 4), USNM 159248. 

Early Devonian, Rabbit Hill Limestone; Devonian coral zone A. Locality

M1032, Coal Canyon, Simpson Park Range, Nev. 

15,16. Syringaxon foerstei n. sp.

15. Lateral view of paratype ( x 2), USNM 159249.

16. Lateral view of paratype ( x 2), USNM 159250. 

Same horizon and locality as figures 1-8.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 1

13

SYRINGAXON
PLATE 2

FIGURES 1-6. Kobeha cf. K. walcotti n. gen., n. sp.

1,2. Exterior and calice interior (x 1). Lower Devonian; Devonian

coral zone B. Locality M1021, McColley Canyon, Sulphur Springs

Range, Nev. 

3. Exterior (x 1). Lower Devonian; Devonian coral zone B. Locality

M1018, McColley Canyon, Sulphur Spring Range, Nev. 

4,5. Longitudinal and transverse thin sections (X 1%). Lower Devonian; Devonian coral zone B. Locality M1024, McColley Canyon,

Sulphur Spring Range, Nev. 

6. Exterior of two attached corallites ( x 1). Lower Devonian; Devonian

coral zone B. Locality M1025, McColley Canyon, Sulphur Spring

Range, Nev. 

7-9. Siphonophrentoid coral.

Lateral and calice views (x 1) and longitudinal thin section (x 3).

Lower Devonian; Devonian coral zone A (Helderbergian); locality

M186, southern Sulphur Spring Range, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 2

KOBEHA AND SIPHONOPHRENTOID CORAL
PLATE 3

FIGURES 1, 2, 6. Kobeha walcotti n. gen., n. sp.

1. Calice view of paratype ( x 1), USNM 159251.

2. Lateral view of paratype ( x 1), USNM 159252.

6. Lateral view of holotype ( x 1), USNM 159253.

Lower Devonian, Nevada Formation, unit 1; Devonian coral zone B.

Locality M56, southern Sulphur Spring Range, Nev. 

3-5, 8, 9. Kobeha ketophylloides n. sp.

3-5. Lateral, calice, and calice interior views of paratype (x 1), 

USNM 159254. Lower Devonian, Nevada Formation, unit 1; Devonian coral zone B. Locality M1039, southern Sulphur Spring Range, 

Nev.

8, 9. Oblique calice view (x 1) and neanic transverse thin section 

(X 2) of paratype USNM 159255. Lower Devonian, Nevada Formation, unit 1; Devonian coral zone B. Locality M69, southeast of 

Prince of Wales mine, southern Sulphur Spring Range, Nev. 

7. Kobeha cf. K. walcotti n. gen., n. sp.

Transverse thin section (x 2) of late neanic stage. Lower Devonian, 

Nevada Formation, unit 1; Devonian coral zone B. Locality M1040, 

southern Sulphur Spring Range, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 3

2 

^
PLATE 4

FIGURES 1-4. Kobeha walcotti n. gen., n. sp.

Early ephebic (x 2), late neanic (x 4), late ephebic (x 1V£), and 

longitudinal thin section (x l 1/^) of paratype USNM 159256. Early 

Devonian, Nevada Formation, unit 1, Devonian coral zone B. Locality 

M1041, southeast of Prince of Wales mine, southern Sulphur Spring 

Range, Nev. 

5, 6. Kobeha cf. K. walcotti n. gen., n. sp.

Longitudinal thin section (x 2), neanic transverse thin section ( x 2). 

Early Devonian, Nevada Formation, unit 1, Devonian coral zone B. 

Locality M67, southern Sulphur Spring Range, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 4

KOBEHA
PLATE 5

FIGURES 1-7. Kobeha walcotti n. gen., n. sp.

1,2. Transverse and longitudinal thin sections of para type (x

USNM 159257. Lower Devonian, Nevada Formation, unit 1; Devonian 

coral zone B. Locality M56, southern Sulphur Spring Range, Nev.

3-5. Transverse and longitudinal thin sections ( x 1V&) and transverse 

thin section of late neanic stage (x 2). Lower Devonian, Nevada 

Formation, unit 1; Devonian coral zone B. Locality M4, southeast of 

Prince of Wales mine, southern Sulphur Spring Range, Nev.

6. Longitudinal thin section (x 2). Lower Devonian, Nevada Formation, unit 1; Devonian coral zone B. Locality M67, northeast of Bailey 

Pass, southern Sulphur Spring Range, Nev.

7. Longitudinal thin section of holotype ( x 1%), USNM 159253. Lower 

Devonian, Nevada Formation, unit 1; Devonian coral zone B. Locality 

M56, southern Sulphur Spring Range, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 5

6 

KOBEHA
PLATE 6

FIGURES 1-7. Kobeha ketophylloides n. sp.

1-5. Longitudinal thin section (x 2), transverse thin section (x 1), 

enlargement of previous thin section (x 2), early ephebic transverse 

thin section (x 2), and neanic transverse thin section ( x 2), all of 

holotype USNM 159258.

6. Neanic transverse thin section ( X 2).

7. Early ephebic transverse thin section of paratype (x 2), USNM

159259. 

Lower Devonian, Nevada Formation, unit 1; Devonian coral zone B.

Locality M1042, 1.75 miles north of Roberts Creek Ranch, Roberts

Mountains, Nev. 

8, 9. Kobeha walcotti n. gen., n. sp.

Neanic transverse thin sections (x 2), (x 4) of paratype USNM

159260. Lower Devonian, Nevada Formation, unit 1; Devonian coral 

zone B. Locality M69, southeast of Prince of Wales mine, southern 

Sulphur Spring Range, Nev. 

10. Kobeha walcotti n. gen., n. sp.

Transverse thin section of late nepionic growth stage (x 4). Lower 

Devonian, Nevada Formation, unit 1; Devonian coral zone B. Locality 

M56, southern Sulphur Spring Range, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 6

10

KOBEHA
PLATE 7

FIGURES 1, 2. Papiliophyllum elegantulum subsp. d.

1,2. Transverse and longitudinal thin sections (x l 1/^). Late Early 

Devonian, Nevada Formation, unit 2; Devonian coral zone Dx . Locality M1037, south pediment of Lone Mountain, Eureka County, Nev.

3. Papiliophyllum cf. P. elegantulum Stumm.

Lateral exterior (XI). From limestone cobble, Locality M1043, Cobble 

Hill, Cockalorum Wash quadrangle, Fish Creek Range, Nev.

4. Papiliophyllum elegantulum Stumm.

Calice interior (x 1). Early Devonian, Nevada Formation, unit 1; 

Devonian coral zone C. Locality M286, Lone Mountain, Eureka 

County, Nev. 

5-7. Papiliophyllum elegantulum Stumm.

Lateral and calice interior views (x 1) and early ephebic transverse 

thin section (x 4). Early Devonian, Nevada Formation, unit 1; 

Devonian coral zone C. Locality M74, Lone Mountain, Eureka County, 

Nev.

8. Papiliophyllum elegantulum Stumm.

Neanic transverse thin section ( x 4). Same locality as figures 5-7.

9. Papiliophyllum elegantulum Stumm.

Neanic transverse thin section (x 4). Same locality as figures 5-7.

10. Papiliophyllum elegantulum Stumm.

Late neanic transverse thin section (x 2). Same locality as figure 4.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 7

8

PAPILIOPHYLL UM
PLATE 8

FIGURES 1 4. Papiliophyllum elegantulum Stumm.

Mature transverse thin section ( x 2), early ephebic thin section ( x 2),

longitudinal thin section (x 2), mature transverse section (x iVfc)-

Early Devonian, Nevada Formation, unit 1; Devonian coral zone C.

Locality M286, northwest side of Lone Mountain, Eureka County,

Nev. 

5, 6. Papiliophyllum elegantulum Stumm.

Longitudinal thin section (x 1), mature transverse thin section (x 1).

Early Devonian, Nevada Formation, unit 1; Devonian coral zone C.

Locality M1044, west side Lone Mountain, Eureka County, Nev.

7. Papiliophyllum elegantulum Stumm.

Longitudinal thin section (x 1%). Early Devonian, Nevada Formation, 

unit 1; Devonian coral zone C. Locality M68, east of Prince of Wales 

mine, Sulphur Spring Range, Nev.

8. Papiliophyllum elegantulum Stumm.

Early ephebic transverse thin section ( x 1%)- Early Devonian, Nevada 

Formation. Locality M1035a, north end Antelope Range, southern 

Eureka County, Nev. 

9,10. Eurekaphyllum breviseptatum Stumm.

Longitudinal and transverse thin sections (x 1) of holotype; copies of 

original figures (Stumm, 1937, pi. 54, figs. 8a, 8b). Nevada Formation, 

Lone Mountain, Eureka County, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE

PAPILIOPHYLLUM AND EUREKAPHYLLUM
PLATE 9

FIGURES 1-5. Odontophyttum meeki n. sp.

1 4. Calice view and two lateral views (x 1%). tilted calice view 

(X 2) of holotypeUSNM 159261.

5. Lateral calice view ( x 1) of paratype USNM 159262.

Early Devonian, Nevada Formation, unit 1; Devonian coral zone C.

Locality M74, Lone Mountain, Eureka County, Nev. 

6-9. Aulacophyllum sp. c.

6. 7. Calice view ( x 1), early ephebic transverse thin section ( x

USNM 159263. 

8,9. Calice view (x 1), ephebic transverse thin section (x

USNM 159264. 

Same horizon and locality as figures 1-5.

10. Aulacophyttum-^ke rugose coral.

Lateral-interior of weathered specimen (x 1) showing down-bend of 

tabulae at fossula. Same horizon and locality as figures 1-5.

11. Aulacophyllum-\\ke rugose coral.

Mature transverse thin section ( X 2). Early Middle Devonian, Nevada 

Formation, unit 1; Devonian coral zone D. Locality M36, south of 

Bailey Pass, Sulphur Spring Range, Nev.

12. ?'Aulacophyllum sp.

Calice view ( X 1). Same horizon and locality as figures 1-5. 

13,14. ?Odontophyllum sp.

Calice and lateral views ( x 1). Same horizon and locality as figures 1-5.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 9

 V ______fc' i\ , 14 

7 ^**rr 11 

ODONTOPHYLLUM, AULACOPHYLLUM, AND AULACOPHYLLUM-UKE RUGOSE CORALS
PLATE 10

FIGURES 1-7. Bethanyphyllum antelopensis n. sp.

1,2. Transverse and longitudinal thin sections of holotype (x 2), 

USNM 159265.

3. Transverse thin section of paratype (x 2), USNM 159266.

4. Longitudinal thin section of paratype ( X 2), USNM 159267.

5. Longitudinal thin section of paratype (x 2), USNM 159268.

6. 7. Late neanic thin section (X 4) and ephebic thin section (X 2)

of paratype USNM 159269. 

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral zone

D2 . Locality M27, Combs Peak, southern Mahogany Hills, Bellevue

Peak quadrangle, Eureka County, Nev. 

8-10. Bethanyphyllum sp. d.

8, 9. Neanic transverse thin section ( X 4) and longitudinal thin section

(x 2), USNM 159270. Late Early Devonian, Nevada Formation;

Devonian coral zone Da . Locality M1037. Pediment slopes, south side

of Lone Mountain, Eureka County, Nev. 

10. Neanic transverse thin section ( x 4), USNM 159271. Same horizon

and locality as figures 8,9.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 10

BETHANYPHYLL UM
PLATE 11

FIGURES 1-4. Bethanyphyllum lonense (Stumm).

1, 2. Transverse and longitudinal thin sections ( X 2), USNM 159272. 

3, 4. Transverse and longitudinal thin sections ( x 2), USNM 159273. 

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral zone

D2 . Locality M3, Grays Canyon, southern Eureka district, Nevada. 

5, 6. Bethanyphyllum antelopensis n. sp.

Transverse and longitudinal thin sections of paratype (x 4), USNM

159274. Early Middle Devonian, Nevada Formation, unit 2; Devonian

coral zone D2 . Locality M27, Combs Peak, southern Mahogany Hills,

Bellevue Peak quadrangle, Nevada. 

7-9. Bethanyphyllum cf. B. antelopensis n. sp.

Exterior (x 1), longitudinal thin section (x 2), transverse thin section

(X 2), USNM 159275. Middle Devonian, Nevada Formation, unit 2;

Devonian, coral zone D3 . Locality M1045, northwest side of Lone

Mountain, Eureka County, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 11

>y

BETHANYPHYLL UM
PLATE 12

FIGURES 1,2. BethanyphyHum ct.B. lonense (Sturam.).

Transverse and longitudinal thin sections (x 1%). USNM 159276. 

Early Middle Devonian, Nevada Formation, unit 2. Locality M1046, 

Lone Mountain, Eureka County, Nev. 

3-5. Bethanyphyllum lonense (Stumm).

Two transverse thin sections and a longitudinal thin section (x 2), 

USNM 159277. Middle Devonian, Nevada Formation, unit 2; Devonian coral zone D2. Locality M1047, east side Sulphur Spring Range, 

Nev. 

6-8. Bethanphyllum lonense (Stumm).

Longitudinal section, early ephebic thin section and neanic thin section 

( X 2), USNM 159278. Middle Devonian, Nevada Formation, unit 2; 

Devonian coral zone D2 . Locality M1048, northwest side of Lone 

Mountain, Eureka County, Nev. 

9-10. Bethanyphyllum lonense (Stumm).

Transverse and longitudinal thin sections (X 2), USNM 159279. Devonian, Nevada Formation, unit 2; Middle Devonian coral zone D2 . 

Locality M3, Grays Canyon, southern Eureka district, Nevada. 

11. Bethanyphyllum antelopensis n. sp.

Longitudinal thin section at calice (x 2). Middle Devonian, Nevada 

Formation, unit 2; Devonian coral zone D2. Locality M27, Combs 

Peak, southern Mahogany Hills, Bellevue Peak quadrangle, Nevada.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 12

8

BETHANYPHYLL UM
PLATE 13

FIGURES 1, 2. (?) Kodonophyllum sp. f.

1. Transverse smoothed surface (X 1). Photographed in water.

2. Transverse thin section (X 3).

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral

zone D. Locality M1033. north end of Fenstermaker Mountain, Fish

Creek Range, Nev. 

3,4. Nevadaphyllum masoni Stumm.

3. Transverse thin section ( x ll/2 ) of holotype, USNM 94447.

4. Longitudinal thin section ( x 3) of paratype, USNM 94447a.

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral zone

D. Lone Mountain, Eureka County, Nev. 

5. Nevadaphyllum-Uke rugose coral.

Transverse thin section ( x 2), USNM 159280. Early Middle Devonian, 

Devonian, Nevada Formation, unit 2; Devonian coral zone D. Locality 

M1049, Lone Mountain, Eureka County, Nev. 

6, 7. Bethanyphyllum sp. d.

Transverse and longitudinal thin sections (x 2), USNM 159281. Late 

Early Devonian, Nevada Formation; Devonian coral zone Dx . Locality M1037, pediment slopes, south side Lone Mountain, Eureka 

County, Nev.

8. Bethanyphyllum cf. B. lonense (Stumm).

Transverse thin section (X 1%), USNM 159282. Early Middle Devonian, Nevada Formation, unit 2; Devonian coral zone D. Locality 

M1050, Lone Mountain, Eureka County, Nev.

9. Bethanyphyllum sp. d.

Longitudinal section (x 1V£), USNM 159281a. Same horizon and locality as figures 6, 7.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 13

9 7 

(1}KODONOPHYLLUM, NEVADAPHYLLUM, NEVADAPHYLLUM-LIKE 

RUGOSE CORAL, AND BETHANYPHYLLUM
PLATE 14

FIGURES 1-15. Siphonophrentis (Breviphrentis) kobehensis n. sp.

1. Lateral view of paratype (X 1), USNM 159283.

2. Lateral view of holotype ( X 1), USNM 159284.

Early Devonian, Nevada Formation, unit 1; Devonian coral zone C.

Locality M286, northwest side of Lone Mountain, Eureka County,

Nev.

3. Lateral view of paratype ( X 1), USNM 159285.

Early Devonian, Nevada Formation, unit 1; Devonian coral zone C. 

Locality M74, west side of Lone Mountain, Eureka County, Nev.

4. Lateral view of paratype (x 1), USNM 159286. Same horizon and 

locality as figures 1, 2.

5. Lateral view of paratype (x 1), USNM 159287. Same horizon and 

locality as figures 1,2.

6. Calice view of paratype (x 1), USNM 159288. Same horizon and 

locality as figures 1, 2.

7. Calice view of paratype (x 1), USNM 159289. Same horizon and 

locality as figures 1, 2.

8. Longitudinal thin section of paratype (x 2), USNM 159290. Same

horizon and locality as figures 1, 2. 

9,10. Transverse and longitudinal thin sections of paratype (x 4),

USNM 159291. Same horizon and locality as figure 3.

11. Longitudinal thin section of paratype (x 2), USNM 159292. Same 

horizon and locality as figure 3.

12. Longitudinal thin section of paratype (x 4), USNM 159293. Same 

horizon and locality as figure 3.

13,14. Two early neanic transverse thin sections ( x 7), USNM 159294. 

Same horizon and locality as figure 3.

15. Nepionic transverse thin section ( x 7), USNM 159295. Same horizon and locality as figure 3.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 14

11

SIPHONOPHRENTIS (BREVIPHRENTIS)
PLATE 15

FIGURES 1-11. Siphonophrentis (Breviphrentis) invaginatus (Stumm).

1,2. Transverse thin section (x 2) and longitudinal thin section

(x 4), USNM 159296. 

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral

zone D. Locality M1033, north end of Fenstermaker Mountain, Fish

Creek Range, Nev. 

3,4. Transverse thin section (x 4) and longitudinal thin section

(x 2), USNM 159297. 

Early Middle Devonian, Nevada Formation, unit 2; Devonial coral

zone D. Locality M198, southern Sulphur Spring Range, Nev. 

5, 6. Transverse and longitudinal thin sections ( x 2), USNM 159298.

Devonian coral zone D. Locality M1034, Ranger Mountains, Nevada

Test Site. 

7, 8. Transverse and longitudinal thin sections ( x 2), USNM 159299.

Same horizon and locality as figures 1,2.

9. Transverse section (X 2), USNM 159300. Early Middle Devonian, 

Nevada Formation, unit 2; Devonian coral zone D2 . Locality M3, 

Grays Canyon, southern Eureka district, Nev.

10. Longitudinal thin section (x 2), USNM 159301. Devonian coral 

zone D. Locality M1035, north end Antelope Range, Eureka County, 

Nev.

11. Longitudinal thin section (X 2), USNM 159302. Same horizon and 

locality as figure 9.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 15

SIPHONOPHRENTIS (BREVIPHRENTIS)
PLATE 16

FIGURES 1-13. Siphonophrentis (Breviphrentis) invaginatus (Sturnm).

1, 2. Transverse and longitudinal thin sections (X 2), USNM 159303, 

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral 

zone D2 . Locality M55, Lone Mountain, Eureka County, Nev.

3. Transverse thin section (X 2), USNM 159304.

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral

zone D2 . Locality M3, Grays Canyon, southern Eureka district,

Nevada.

4. Longitudinal thin section ( x 2), USNM 159305. Same horizon and

locality as figure 3. 

5-7. Transverse thin section (x 2), longitudinal thin section ( x 2),

and transverse thin section of neanic stage (x 4), USNM 159306.

Same horizon and locality as figure 3.

8. Transverse thin section (x 2), USNM 159307. Same horizon and 

locality as figure 3.

9. Transverse thin section (X 2), USNM 159308. Same horizon and 

locality as figure 3.

10,11. Transverse and longitudinal thin section ( x 2), USNM 159309.

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral 

zone D2. Locality M1033, north end Fenstermaker Mountain, Fish 

Creek Range, Nev.

12,13. Transverse and longitudinal thin sections (x 2), USNM 

159310.

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral 

zone D2 . Locality M27, Combs Peak, southern Mahogany Hills, 

Eureka County, Nev. 

14. Siphonophrentis (Breviphrentis) kobehensis n. sp.

Transverse thin section of paratype (x 2), USNM 159283. Early 

Devonian, Nevada Formation, unit 1; Devonian coral zone C. Locality M286, northwest side of Lone Mountain, Eureka County, Nev. 

15,16. Siphonophrentis (Breviphrentis) invaginatus (Stumm).

Transverse and longitudinal thin sections ( X 2), USNM 159311. Early 

Middle Devonian, Nevada Formation, unit 2; Devonian coral zone 

D2 . Locality M1036, Lone Mountain, Eureka County, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 16

SIPHONOPHRENTIS (BREVIPHRENTIS)
PLATE 17

FIGURES 1, 2. Sinospongophyllum sp. d.

Longitudinal and transverse thin sections ( x 2), USNM 159312. Late 

Early Devonian, Nevada Formation, unit 2; Devonian coral zone D^ 

Locality Ml037, south pediment slope, Lone Mountain, Eureka 

County, Nev. 

3, 4. Sinospongophyllum sp. e.

Transverse and longitudinal thin sections (x ll/2 ), USNM 159313. 

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral 

zone D2. Locality M1038, Lone Mountain, Eureka County, Nev. 

5, 6. Sinospongophyllum sp. d.

Longitudinal and transverse thin sections ( x 2), USNM 159314. Early 

Middle Devonian, Nevada Formation, unit 2; Devonian coral zone 

D2. Locality M1035, north end Antelope Range, southern Eureka 

County, Nev. 

7. Sinospongophyllum sp. f.

Transverse thin section ( x 2), USNM 159315. Early Middle Devonian, 

Nevada Formation, unit 2; Devonian coral zone D2 . Locality M3, 

Grays Canyon, southern Eureka district, Nevada. 

8, 9. Sinospongophyllum sp. f.

Transverse and longitudinal thin sections (x 2), USNM 159316. Early 

Middle Devonian, Nevada Formation, unit 2; Devonian coral zone 

D2 . Locality M1035, north end Antelope Range, southern Eureka 

County, Nev. 

10,11. Sinospongophyllum sp. e.

Transverse and longitudinal thin sections (x 2), USNM 159317. Early 

Middle Devonian, Nevada Formation, unit 2; Devonian coral zone D2 . 

Locality M1035, north end Antelope Range, southern Eureka County, 

Nev. 

12,13. Siphonophrentis (Breviphrentis) invaginatus (Stumm).

Transverse and longitudinal thin sections (x 2), USNM 94445. Holotype of Amplexus lonensis Stumm which is the type species of Breviphyllum Stumm 1949. Nevada Formation, Lone Mountain, Eureka 

County, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 17

12 13

SINOSPONGOPHYLLUM AND SIPHONOPHRENTIS (BREVIPHRENTIS)
PLATE 18

FIGURES 1-4. Dendrostella romanensis n. sp.

Transverse and longitudinal thin sections of holotype ( x 2), USNM

159318. Middle Devonian, Nevada Formation. Devonian coral zone D. 

Locality M1031, southern Sulphur Spring Range, northwest of Romano 

Ranch, Eureka County, Nev. 

5, 6. Disphyllum eurekaensis n. sp.

Transverse and longitudinal thin sections of holotype ( x 4), USNM

159319. Early Middle Devonian, Nevada Formation, unit 2; Devonian 

coral zone D2 . Locality Ml046, Lone Mountain, Eureka County, Nev. 

7-10. Disphyllum nevadense (Stumm).

7-9. Transverse and longitudinal thin sections (x 4), USNM 159320.

Same horizon and locality as figures 5,6.

10. Longitudinal thin section ( x 4), USNM 159321. Same horizon and 

locality as figures 5, 6.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 18

DENDROSTELLA AND DISPHYLLUM
PLATE 19

FIGURES 1-9. Cystiphylloides lonense (Stumm).

1,2. Calice and lateral views (x 1), USNM 159322. Early Devonian,

Nevada Formation, unit 1; Devonian coral zone C. Locality M74,

Lone Mountain, Eureka County, Nev. 

3,4. Lateral and calice views (x 1), USNM 159323. Early Devonian,

Nevada Formation, unit 1; Devonian coral zone C. Locality M286,

Lone Mountain, Eureka County, Nev.

5. Lateral view (x 1), USNM 159324. Same horizon and locality as 

figures 3, 4.

6. Transverse thin section (x 2), USNM 159325. Same horizon and 

locality as figures 1, 2.

7. 8. Longitudinal and transverse thin sections ( x 2), USNM 159326.

Same horizon and locality as figures 1, 2. 

9. Longitudinal thin section ( x 2), USNM 159327. Same horizon and

locality as figures 1, 2. 

10,11. Cystiphylloides sp. d.

Transverse and longitudinal thin sections (x 2), USNM 159328. Early

Middle Devonian, Nevada Formation, unit 2; Devonian coral zone

D2. Locality M36, Sulphur Spring Range, Nev. 

12,13. Cystiphylloides sp.

Transverse section and longitudinal section at calice (x 1%). USNM

159329. Early Middle Devonian, Nevada Formation, unit 2; Devonian

coral zone D2. Locality M1048, Lone Mountain, Eureka County, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 19

CYSTIPHYLLOIDES
PLATE 20

FIGURES 1,2. Cystiphylloides sp.

Transverse thin section (x 1V&), longitudinal thin section (x 2), 

USNM M159330. Early Middle Devonian, Nevada Formation, unit 2; 

Devonian coral zone D2 . Locality M1051, south end Sulphur Spring 

Range, Nev. 

3, 4. Zonophyllum haguei n. sp.

Transverse thin section ( x 4), longitudinal thin section ( x 2) of paratype, USNM 159331. Early Middle Devonian, Nevada Formation, 

unit 2; Devonian coral zone D2 . Locality M51, Grays Canyon, southern Eureka district, Nevada. 

5, 6. Zonophyllum sp. a.

Longitudinal and transverse thin sections ( x 2), USNM 159332. Early 

Middle Devonian; Devonian coral zone D. Locality M1058, Nevada 

Test Site, Frenchman Flat, Nev. 

7,8. Zonophyllum cf. Z. haguei n. sp.

Longitudinal and transverse thin sections (X 2). Early Middle Devonian, Nevada Formation, unit 2; Devonian coral zone D2 . Locality 

M1035, north end of Antelope Range, Eureka County, Nev. 

9,10. Zonophyllum haguei n. sp.

Transverse and longitudinal thin sections (x 2) of holotype USNM 

159333. Early Middle Devonian, Nevada Formation, unit 2; Devonian 

coral zone D2 . Locality M3, Grays Canyon, southern Eureka district, 

Nevada. 

11, 12. Zonophyllum sp. b.

Transverse thin section ( x 2), longitudinal thin section ( x 4), USNM 

159351. Same horizon and locality as figures 3,4.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 20

CYSTIPHYLLOIDES AND ZONOPHYLLUM
PLATE 21

FIGURES 1, 2. Mesophyllum (Mesophyllum) sp. b.

Two transverse thin sections (x 2), USNM 159334. Early Middle 

Devonian, Nevada Formation, unit 2; Devonian coral zone D2 . Locality M36, southern Sulphur Spring Range, Nev. 

3,4. Mesophyllum (Mesophyllum) sp. c.

Transverse and longitudinal thin sections (x 2), USNM 159335. Early 

Middle Devonian, Nevada Formation, unit 2; Devonian coral zone D. 

Locality M1035, north end Antelope Range, Eureka County, Nev. 

5-7. Mesophyllum (Arcophyllum) kirki (Stumm).

Transverse and longitudinal thin sections (x 2) and enlarged transverse thin section (x 4), USNM 159336. Middle Devonian, Nevada 

Formation, unit 2; Devonian coral zone D3 . Locality M29, Lone 

Mountain, Eureka County, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 21

6 7

MESOPHYLLUM AND MESOPHYLLUM (ARCOPHYLLUM)
PLATE 22

FIGURES 1-6. Mesophyllum (Arcophyllum) kirki (Stumm).

1. Transverse thin section ( X 2), USNM 159337.

2. Transverse thin section ( X 2), USNM 159338.

3. Enlargement of transverse thin section ( X 4), USNM 159339.

4. Longitudinal thin section at calice (X 2), USNM 159340.

5. Longitudinal thin section (X 4), USNM 159341.

6. Longitudinal thin section (X 2), USNM 159342.

Middle Devonian, Nevada Formation, unit 2; Devonian coral zone D3 .

Locality M1036, northwest side of Lone Mountain, Eureka County,

Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 22

MESOPHYLLUM (ARCOPHYLLUM)
PLATE 23

FIGURES 1-4. Cystiphylloides robertsense (Stumin).

Transverse, longitudinal, longitudinal, and transverse thin sections 

(x 2), USNM 159343. Early Middle Devonian, Nevada Formation, 

unit 2; Devonian coral zone D2 . Locality M1046, Lone Mountain, 

Eureka County, Nev. 

5-10. Hexagonaria (Pinyonastraea) kirki (Stumm) n. subgen.

5,6. Transverse and longitudinal thin sections of holotype (x 2), 

USNM 94456. Middle Devonian, Nevada Formation, unit 2; 

Devonian coral zone D. Lone Mountain, Eureka County, Nev. 

7. Upper surface of large colony (x 1), USNM 159344. Middle 

Devonian, Nevada Formation, unit 2; Devonian coral zone D3 . 

Locality M1052, northwest side of Lone Mountain, Eureka 

County, Nev.

8,9. Transverse and longitudinal thin sections ( x 2), USNM 159345. 

Middle Devonian, Nevada Formation, unit 2; Devonian coral 

zone D. Locality Ml053, north end of Antelope Range, Eureka 

County, Nevada.

10. Transverse print from stained cellulose peel (x 2). Middle 

Devonian, Nevada Formation, unit 2; Devonian coral zone D. 

Locality M1054, south side of Cooper Peak, Roberts Mountains, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 23

9 10 

CYSTIPHYLLOIDES AND HEXAGONARIA (PINYONASTRAEA)
PLATE 24

FIGURES 1-3. Billingsastraea nevadensis (Stumm).

Longitudinal thin section ( x 2), surface of colony ( x 1), and transverse 

thin section (x 2), USNM 159346. Early Middle Devonian, Nevada 

Formation, unit 2; Devonian coral zone D2 . Locality M1055, north 

end Antelope Range, Eureka County, Nev.

4. Billingsastraea nevadensis subsp. arachne (Stumm).

Upper surface of colony (x 1), USNM 94458. Early Middle Devonian, 

Nevada Formation, unit 2; Devonian coral zone D2 . Lone Mountain, 

Eureka County, Nev. Copy of Stumm's (1937, pi. 53, fig. 13) holotype 

figure.

5. Billingsastraea nevadensis (Stumm).

Transverse thin section ( x3), USNM 159347. Early Middle Devonian, 

Nevada Formation, unit 2; Devonian coral zone D2 . Locality M1035, 

north end Antelope Range, Eureka County, Nev. 

6-8. Billingsastraea nevadensis (Stumm).

6. Longitudinal thin section ( X 4), USNM 159348.

7. Transverse thin section (x 4), USNM 159349.

8. Longitudinal thin section ( x 4), USNM 159350.

Early Middle Devonian, Nevada Formation, unit 2; Devonian coral 

zone D2 . Locality M55, Lone Mountain, Eureka County, Nev.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 24

BILLINGSASTRAEA
PLATE 25

FIGURES 1-4. Australophyllum landerensis n. sp.

1,2. Transverse thin section (x2%) and longitudinal thin section

( x 2i/2 ) of holotype USNM 159353. 

3, 4. Longitudinal and transverse thin sections (x 2) of paratype

USNM 159354. 

Early Devonian, Rabbit Hill Limestone. Locality Ml 150, north end of

Toquima Range. 

5. Australophyllum sp.

Transverse thin section (x 2) . Upper Silurian or lowermost Devonian 

just above Silurian coral zone E, upper unit of Vaughn Gulch Limestone. Locality M1093, Mazourka Canyon, northern Inyo Mountains, 

Calif. 

6-9. Billingsastraeal sp. T.

6, 7. Transverse thin sections ( x 2, x 3).

8, 9. Longitudinal thin sections ( x 3).

All sections cut from same colony. Middle Devonian. Locality M1151,

Reeds Canyon area, Toiyabe Range. 

10. (?)Hexagonariasp.

Transverse thin section (x 2). Same locality and horizon as figures 6-9.
GEOLOGICAL SURVEY PROFESSIONAL PAPER 805 PLATE 25

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10 7 

AUSTRALOPHYLLUM, BILLINGSAST-RAEAt, AND (^HEXAGONARIA